Furcraea Foetida (L.) Haw
Furcraea Foetida (L.) Haw
Furcraea Foetida (L.) Haw
Department
of Agriculture
Wildland Shrubs of the
Forest Service United States and Its
International Institute
of Tropical Forestry
San Juan, PR
Territories: Thamnic
Rocky Mountain
Research Station
Descriptions:
Fort Collins, CO
General Technical
Report IITF-GTR-26
Volume 1
July 2004
John K. Francis, Editor
Abstract ______________________________________
Francis, John K. ed. 2004. Wildland shrubs of the United States and its Territories:
thamnic descriptions: volume 1. Gen. Tech. Rep. IITF-GTR-26. San Juan, PR: U.S.
Department of Agriculture, Forest Service, International Institute of Tropical Forestry, and
Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain
Research Station. 830 p.
A discussion of the general characteristics of shrubs as a life form and their distribution
within the United States is followed by 311 short monographs containing general descrip-
tions, ranges, ecology, reproductive habits, growth and management, and benefits to
humans, animals, and the environment.
The U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry
operates in cooperation with the University of Puerto Rico, Río Piedras, PR, 00936-4984
——————
The use of trade or firm names in this publication is for reader information and does not
imply endorsement by the U.S. Department of Agriculture of any product or service
I believe that it is appropriate that this effort was undertaken by the USDA Forest Service
Institute of Tropical Forestry and the Rocky Mountain Station’s Shrub Sciences
Laboratory. These institutions represent foci of significant shrub diversity. Shrub
diversity is enhanced both by warm, mesic and tropical conditions and by aridity and
diverse topography. The two institutions—one in the tropics and one in the continental
Western United States desert--represent those contrasting situations.
E. Durant McArthur
Project Leader
Shrubland Biology and Restoration Work Unit
Shrub Sciences Laboratory
Rocky Mountain Research Station
i
Introduction
The evolution of the Wildland Shrub Manual begins a long time ago with the observation and
description of plants; however, its direct ancestor is the U.S. Silvics Manual: “Silvics of North America”
Volumes 1 and 2 (Burns and Honkala 1990a, 1990b) in which the editor participated as a species author.
After this, a regional silvics manual developed at the International Institute of Tropical Forestry was
published as single species’ separate publications and later translated into Spanish and compiled into a
single volume, “Bioecología de Árboles y Exóticos de Puerto Rico y las Indias Occidentales” (Francis and
others 2000). Besides the numerous botanical references cited in the following section and in the species
descriptions, other important references were important contributing sources for this effort. The “Woody-
Plant Seed Manual” (Schopmeyer 1974 and the update in progress, Bonner and Nisley 2003) is cited in
many of the species descriptions. The “Fire Effects Information System” database (Fire Sciences
Laboratory 2003) contributed a significant portion of the information used is a number of the mainland
U.S. species in this volume. The Wildland Shrub Manual differs from its predecessors by the extensive use
of Internet information sources that have not been available previously.
Latin family, genus, species, and authority of plant names were checked against the “Plants”
database (Natural Resources Conservation Service 2003). In a few cases, individual authors disagreed with
the name given in the database and have cited their reasons for using the name in the form they did.
Common names, for which there is really no central authority and which are likely to take another
generation to stabilize, were left to the discretion of the authors. Common names cited were generally
chosen from the many in existence for each species because they were more widely used or because they
were unique or especially descriptive.
Although somewhat simplified, the format of each shrub description generally follows that used in
the silvical descriptions. Sections: General Description, Range, Ecology, Reproduction, Growth and
Management, and Benefits, are followed by References cited. Each description carries a botanical drawing
or a photo to aid in recognition. Because the number of shrubs to be described is so great, it was decided to
make descriptions summaries rather than complete treatises of all information available. Also, to make the
information useful to as wide an audience as possible, authors were encouraged to use wording that is as
simple as possible without sacrificing technical accuracy. Although arranged in alphabetical order by genus
name, a table of contents is provided to make the process of turning to an individual species easier. A
glossary of technical terms and an index of common names are also provided.
These descriptions have been previously published on the Internet (Francis 2003) and the number
of shrub descriptions continues to be added to. The Internet allows revisions of existing descriptions so that
Internet versions of the descriptions may vary somewhat from descriptions contained in this volume.
Volume 1 of the Wildland Shrub Manual contains descriptions of 311 shrub species, a number determined
more by the time permitted for the project than anything else. It is anticipated that the process of compiling
information and describing shrubs will continue and other volumes will follow. The number of shrubs in
the United States and its Territories easily justifies the publication of five or more volumes of descriptions.
I sincerely hope that shrub scientists in the coming years will continue the effort of publishing thamnical
descriptions to facilitate the management of our wildland shrub natural resource.
References Cited
Bonner, F.T., tech. coord. And R.G. Nisley, manag. ed. 2003. Woody plant seed manual. U.S. Department
of Agriculture, Forest Service, Washington, DC. http://wpsm.net/ index.htm. [not paged].
Burns, R.M., and B.H. Honkala, eds. 1990a. Silvics of North America. Vol. 1, Conifers. U.S. Department of
Agriculture, Forest Service, Agriculture Handbook 654. Washington, DC. 675 p.
Burns, R.M., and B.H. Honkala, eds. 1990b. Silvics of North America. Vol. 2, Hardwoods. U.S. Department of
Agriculture, Forest Service, Agriculture Handbook 654. Washington, DC. 877 p.
ii
Fire Sciences Laboratory. 2003. Fire effects information system. U.S. Department of Agriculture, Forest
Service, Rocky Mountain Research Station, Ft. Collins, CO. http://www.fs.fed.us/database/feis/ [not
paged].
Francis, J.K., ed. 2003. Wildland shrubs of the United States and its territories. U.S. General Technical
Report IITF-WB-1. Department of Agriculture, Forest Service, International Institute of Tropical
Forestry, San Juan, PR and Rocky Mountain Experiment Station, Shrub Sciences Laboratory, Provo UT.
http://fs.fed.us/global/iitf/ wildland_shrubs.htm. [not paged].
Natural Resources Conservation Service. 2003. Plants database. U.S. Department of Agriculture,
Washington, DC. http://plants.usda.gov/ [not paged].
Schopmeyer, C.S., tech. coord. 1974. Seeds of woody plants in the United States. Agriculture Handbook
450. U.S. Department of Agriculture, Forest Service, Washington, DC. 883 p.
Acknowledgments
I especially want to thank John A. Parrotta, then Team Leader at the International Institute of
Tropical Forestry, USDA Forest Service, for his appreciation of the need to package species information
for popular benefit and his encouragement to embark on the Shrub Manual project. I must also thank Ariel
E. Lugo, IITF Director, for allowing me to dedicate almost 3 years nearly full time to shrub research and E.
Durant McArthur, Project Leader, Shrub Sciences Laboratory, Rocky Mountain Research Station, USDA
Forest Service, for graciously welcoming me to work at the Shrub Sciences Laboratory and furnishing
considerable financial support to the project. These and some 36 other authors wrote thamnical descriptions
to contribute to the book. More than 100 scientists participated in reviewing the manuscripts before they
were edited. Isaac Nordlund helped with graphics and programming. I owe as much as any other to Louise
Kingsbury, Director of Publishing Services for the Rocky Mountain Research Station, for personally and
promptly editing every one of the manuscripts.
John K. Francis
Research Forester, Retired
iii
Contents
Page Page
Forward and acknowledgements i Caesalpinia bonduc 129
Introduction ii Cajanus cajan 132
Contents iv Callicarpa americana 135
Wildland shrubs 1 Calotropis procera 137
Thamnical Descriptions Capparis indica 140
Abrus precatorius 12 Capraria biflora 142
Acacia farnesiana 14 Capsicum annuum 144
Acacia greggii 16 Capsicum frutescens 147
Acacia retusa 19 Casearia decandra 149
Acacia tortuosa 21 Casearia sylvestris 151
Acer glabrum 24 Cassine xylocarpa 154
Acer spicatum 27 Castela erecta 156
Acnistus arborescens 30 Ceanothus velutinus 158
Aegiphila martinicensis 32 Celastrus orbiculatus 161
Agave missionum 34 Celastrus scandens 164
Allamanda cathartica 36 Celtis reticulata 167
Amyris elemifera 38 Cephalanthus occidentalis 170
Aralia spinosa 40 Cercocarpus intricatus 173
Ardisia elliptica 43 Cercocarpus ledifolius 175
Argythamnia candicans 45 Cercocarpus montanus 178
Artemisia arbuscula 47 Cereus greggii 181
Artemisia bigelovii 50 Cestrum diurnum 184
Artemisia californica 52 Cestrum laurafolium 186
Artemisia cana 57 Chamaebiateria millefolium 188
Artemisia filifolia 60 Chenopodium oahuense 190
Artemisia frigida 63 Chiococca alba 192
Artemisia longiloba 66 Chromolaena geraniifolium 194
Artemisia lugoviciana 69 Chromolaena odoratum 196
Artemisia nova 72 Chrysobalanus icaco 199
Artemisia pygmaea 75 Chrysophyllum oliviforme 201
Artemisia ridgida 77 Chrysothamnus nauseosus 203
Artemisia rothrockii 79 Chrysothamnus parryi 206
Artemisia spinescens 83 Chrysothamnus viscidiflorus 209
Artemisia tridentata 85 Cissampelos pareira 212
Artemisia tripartita 88 Citrus aurantiifolia 214
Arundinaria gigantean 90 Clematis orientalis 217
Atriplex canescens 93 Clibadium erosum 219
Atriplex confertifolia 96 Clidemia cymosa 221
Atriplex corrugata 99 Clidemia hirta 223
Atriplex cuneata 101 Coccoloba krugii 225
Baccharis pilularis 103 Coccoloba microstachya 227
Barleria prionitis 105 Coccoloba uvifera 229
Batis maritima 107 Coffea arabica 232
Bidens menziesii 110 Colubrina arborescens 235
Bixa orellana 112 Comptonia peregrina 237
Bocconia frutescens 115 Conocarpus erectus 240
Bontia daphnoides 117 Corchorus hirsutus 243
Borrichia arborescens 119 Cordia lima 245
Bourreria virgata 121 Cordia polycephala 247
Brickellia californica 123 Cornus sericea 249
Bromelia penguin 125 Coursetia caribaea 252
Brunfelsia lactea 127 Crataegus douglasii 254
iv
Critonia portoricense 257 Jatropha gossypiifolia 399
Crossopetalum rhacoma 259 Juniperus communis 401
Crotalaria lotifolia 261 Juniperus osteosperma 404
Crotalaria spectabilis 263 Kalmia latifolia 407
Cuphea hyssopifolia 266 Kaonophyllon polyodon 411
Curatella americana 268 Krameria ixine 412
Cytisus scoparius 275 Lantana camara 414
Dalbergia ecastaphyllum 278 Lantana involucrata 417
Dasiphora floribunda 280 Larrea tridentata 419
Dasylirion wheeleri 283 Lasiacis divaricata 425
Datura metel 286 Leandra krugii 427
Dasmanthus virgatus 289 Ledum groenlandicum 429
Distictis lactiflora 291 Lepianthes peltata 432
Dodonaea viscosa 293 Lepidospartum burgessii 434
Dryas octopetala 295 Leucaena leucocephala 437
Duranta erecta 299 Lindera melissifolia 440
Encelia farinosa 302 Lonicera japonica 442
Ephedra nevadensis 307 Lotus scoparius 445
Ephedra viridis 310 Ludwigia octovalvis 449
Eriogonum fasciculatum 314 Lupinus arboreus 451
Erithalis fruticosa 319 Lycium pallidum 453
Ernodea littoralis 321 Macfadyena unguis-cati 456
Erythroxylum areolatum 323 Machaonia portoricensis 459
Erythroxylum brevipes 325 Mahonia aquifolium 461
Eugenia foetida 327 Mahonia repens 464
Eugenia pseudopsidium 329 Malvastrum americanum 467
Euonymus alatus 331 Mecranium latifolium 469
Euonymus americanus 333 Melochia nodiflora 471
Euonymus atropurpureus 335 Melochia tomentosa 473
Fallugia paradoxa 337 Miconia impetiolaris 475
Faramaea occidentalis 341 Miconia prasina 477
Forestiera acuminata 343 Miconia racemosa 480
Forestiera segregata 345 Mimosa aculeaticarpa 482
Fouquieria splendens 347 Mimosa arenosa 485
Furcraea foetida 350 Mimosa ceratonia 487
Gesneria pedunculosa 352 Mimosa pigra 489
Gonzalagunia hirsuta 354 Mimosa pudica 492
Gossypium hirsutum 356 Mimulus aurantiacus 495
Gouania lupuloides 359 Mitracarpus portoricensis 500
Grayia spinosa 361 Morella caroliniensis 502
Guettarda elliptica 363 Morinda citrifolia 505
Gundlachia corymbosa 365 Murraya exotica 508
Gutierreza sarothae 367 Myrica gale 511
Gynerium sagittatum 370 Neea buxifolia 514
Hamelia patens 372 Odontonema cuspidatum 516
Helicteres jamaicensis 374 Olyra latifolia 518
Hibiscus pernambucensis 376 Opuntia leptocaulis 520
Holodiscus discolor 379 Palicourea crocea 524
Holodiscus dumosus 382 Parathesis crenulata 527
Hyperbaena laurifolia 384 Parkinsonia aculeata 529
Indigofera suffruticosa 386 Parthenocissus quinquefolia 532
Isocoma tenuisecta 388 Passiflora edulis 535
Itea virginica 391 Paullinia pinnata 538
Ixora coccinea 393 Pavonia spinifex 540
Jacquinea arborea 395 Pennisetum purpureum 542
Jasminum fluminense 397 Penstemon ambiguous 545
v
Philadelphus lewisii 548 Securidaca virgata 689
Philadelphus microphyllus 551 Senecio flaccidus 691
Phoradendron quadrangulare 553 Senecio spartioides 694
Phragmites australis 555 Senna atomaria 696
Physocarpus malvaceus 558 Senna occidentalis 699
Picramnia pentandra 561 Senna polyphylla 702
Piper aduncum 564 Serenoa repens 704
Piper amalago 567 Serjania polyphylla 708
Piper swartzianum 569 Sesbania sericea 710
Piptocoma antillana 571 Sida fallax 712
Pisonia aculeata 573 Sida rhombifolia 714
Pithecellobium unguis-cati 575 Smilax domingensis 717
Pluchea carolinensis 577 Solanum drymophilum 719
Prosopis glandulosa 580 Solanum dulcamara 721
Proustia vanillosma 584 Solanum persicifolium 724
Prunus americana 586 Solanum torvum 726
Prunus emarginata 589 Sorbus scopulina 729
Prunus pumila 591 Spermacoce verticillata 732
Prunus virginiana 594 Strumpfia maritima 735
Psidium guajava 597 Styrax americanus 737
Psorothamnus scoparius 600 Suaeda suffrutescens 739
Psychotria brachiata 602 Suriana maritima 742
Psychotria deflexa 604 Symphoricarpos albus 744
Psychotria microdon 606 Tabebuia haemantha 747
Psychotria nervosa 608 Tamarix chinensis 749
Purshia tridentate 610 Tamonea boxiana 753
Quercus havardii 613 Taxus brevifolia 755
Randia aculeata 617 Taxus canadensis 758
Rauvolfia viridis 619 Tecoma stans 761
Rhododendron macrophyllum 621 Tetrapterys inaequalis 764
Rhododendron maximum 623 Thrinax morrisii 766
Rhus copallinum 625 Toxicodendron radicans 769
Rhus glabra 628 Toxicodendron rydbergii 772
Rhus microphylla 631 Trema lamarckianum 775
Rhus trilobata 634 Trichilia hirta 777
Ribes aureum 637 Trichostigma octandrum 779
Ribes cereum 641 Triumfetta semitriloba 781
Ricinus communis 643 Turbina corymbosa 783
Rondeletia inermis 645 Urena lobata 785
Rondeletia pilosa 647 Urera baccifera 788
Rosa woodsii 649 Vernonia albicaulis 790
Rubus discolor 652 Vernonia borinquensis 792
Rubus parviflorus 655 Vernonia proctori 794
Rubus rosifolius 658 Vernonia sericea 796
Salix arctica 660 Waltheria indica 798
Salix exigua 664 Wedelia reticulata 801
Salix scouleriana 668 Zisiphus obtusifolia 803
Salvia apiana 671 Zuckia brandegeei 806
Salvia mellifera 676 Conclusion
Sassafras albidum 681 Glossary of technical terms 808
Schaefferia frutescens 684 Common name index 812
Schinus terebinthifolius 686 Authors 824
vi
Wildland Shrubs
According to the dictionary (Merriam-Webster Inc 1984), the word “shrub” came
from the Old English scrybb wood, which is akin to the Norwegian skrubbebaer that means
a dwarf hardwood species. The implication seems to be that a shrub is a dwarf tree--not far
from the current usage of the word. The designation of this life form is driven by the need
for a category between trees and herbs.
If a tree is “a woody perennial plant, typically large and with a single well-defined
stem carrying a more or less definite crown” (Ford-Robertson 1971), then a shrub must
necessarily be smaller. Most definitions require that a shrub should have more than one main
stem caused by branching below or above the ground level (Allaby 1994, Viereck and Little
1972). Other frequent qualifications include the need for the plant to be perennial and to be
lignified (woody), at least in some of its parts (Allaby 1994, Ford-Robertson 1971, Orshan
1989). Shrubs are distinguished from herbs in that herbs do not develop persistent woody
tissue above ground (Ford-Robertson 1971). However, Lawrence (1955) admits that the
term shrub is “not subject to precise circumscription”.
1
There are several reasons why shrubs cannot be precisely defined. Many species are
trees in fertile habitat or favorable conditions and shrubs in difficult habitat. Certain species
may grow as shrubs in large portions of their range and become trees in limited areas. Most
large shrubs will produce an occasional individual with a clearly tree-like habit. It is not
possible to define exactly what “large” is (referring to stem size). A number of species
normally develop single stems but only reach stem sizes below that normally associated with
trees. The degree of lignification is another source of ambiguity. Many annual herbs become
quite lignified before dieing, and some perennial shrubs have little or no woody tissue.
Finally, the concept of perennial looses much of its meaning in the humid tropics where a
shrub may complete its life cycle in a year of continuous growth, and an herb may live
without dieback for many years.
There are many subtypes of shrubs including dwarf shrubs, half-shrubs (sub-shrubs),
cushion plants, woody vines (lianes and climbers), scrambling and prostrate forms, woody
parasitic plants, and various forms of succulents and semi-succulents. A description of these
forms can be found in Lawrence (1955), and a discussion of criterion for classification of
shrub forms may be found in Orshan (1989).
The concept of the shrub layer, which is populated by functional shrubs, sidesteps
the ambiguity of classifying species. In this system, the forest is divided into vertical layers
(canopies or horizons). In their simplest form, the layers are: the tree layer, consisting of
trees and tree-sized plants, the shrub layer, including all plants with any diameter at breast
height (d.b.h.) that is less than some arbitrary upper d.b.h. limit such as 5.0 cm, and the herb
layer, which does not reach breast height. The principal components of the shrub layer are
actually young trees. This concept is useful in studies of wildlife habitat, plant ecology, and
biomass and carbon distribution.
Because of their large size, influence on the environment, and considerable economic
value, trees have received most of the attention in forest research and ecological
descriptions. However, shrubs are more important than is generally realized, especially in
terms of biodiversity. Little (1979) lists 679 species of trees native and naturalized in the
continental United States. Another 60 native tree species with an additional number of
naturalized species grow in Hawaii (Little and Skolmen 1989). Puerto Rico and the U.S.
Virgin Islands support 547 native tree species (Little and others 1974) and at least 118
2
species of naturalized exotics (Francis and Liogier 1991). With allowances for species
represented in more than one area, the U.S. and its territories support about 1,300 tree
species. Shrub species in the same area are much more numerous.
Estimating the number of shrub species in the U.S. and its territories is challenging
because the life-form is subject to the perception of authors and compilers and not indicated
or consistently identified in many of the checklists and floras. However, it is possible to
count the shrub species indicated in the floras of several of the states or regions. Alaska has
about 110 species of shrubs (Viereck and Little 1972), California about 800 species
(McMinn 1951), Hawaii about 170 (Degener 1946, Degener 1957, Degener and Degener
1963), Idaho about 160 (Davis 1952), Pennsylvania about 264 (Rhoads and Klein 1993),
Puerto Rico about 540 (Liogier 1985, Liogeir 1988, Liogier 1994, Liogier 1995, Liogier
1997), and tropical Florida about 250 (Long and Lakela 1971). These totals (with
duplication eliminated) plus other miscellaneous sources (Abrams 1940, Abrams 1944,
Abrams 1951, Abrams and Ferris 1960, Benson 1969, Correll and Johnston 1970, Everitt
and Drawe 1993, Great Plains Flora Association 1986, Jones 1975, Morley 1969, Nelson
1996) exceed 2800 species and seems to account for most of the shrub species in the United
States and its territories. The Natural Resources Conservation Service (2003) “Plants”
database recently made it possible to search and list shrub species by region within the
United States and its territories (Puerto Rico and the U.S. Virgin Islands). Although sums for
growth habits (life-forms) are higher than if each species were exclusively assigned to a
single growth habit (eg. Salix exidua Nutt. appears as both tree and shrub), the figures are
instructive, especially for comparing areas, life-forms, and taxonomic divisions. Searching
all plant species that carry the growth habit designates, “shrub” or “sub-shrub,” yielded
5,281 species for the U.S. and territories. There was a great deal of variation between
political divisions (see figure below, not drawn to scale). California and Texas supported the
greatest numbers of species with 1,311 and 1,300 respectively. North Dakota and Alaska
had the lowest with 195 and 200 respectively. The U.S. Virgin Islands had the highest
average shrub biodiversity with 527 shrub species in only 349 km2 of territory.
3
Several factors influence the amount of shrub diversity found within a political unit.
Numbers of shrub species, as well as all plant species, increase as one travels from the harsh
boreal regions to the humid tropics. Diversity of habitat, such as mountain peaks, deserts,
riparian areas, and sea shore, is also very important. Southwestern deserts where shrubs are
the dominant vegetation also tend to have high shrub diversity.
Using searches of the “Plants” database mentioned above, the shrubs of the U.S. and
territories were found to be divided among 166 families. The most important families were:
Asteraceae, 618 species; Rosaceae, 510 species; Fabaceae, 342 species; Cactaceae, 193
species; Ericaceae, 189 species; Scrophulariaceae, 182 species; Rubiaceae, 165 species;
Malvaceae, 148 species; Euphorbiaceae, 128 species; Lamiaceae, 124 species;
Polygonaceae, 123 species; Companulaceae, 112 species; Boraginaceae, 106 species;
Ramnaceae, 103 species; and 152 additional families with 82 to 1 species each.
Reaching or maintaining a high biodiversity is important to ecosystem health.
Managers and the public are beginning to realize that subordinate species, as well as the tree
canopy, are important. It will not be enough, as formerly called for in management plans, to
establish a tree cover and wait for natural succession to fill in the subordinate species.
Shrubs, which are near-climax in ecosystems such as subtropical deserts, are, thus, the
principal species to be maintained or reestablished. In moister wildlands, wildlife managers
4
have long sought to achieve a certain portion of the landscape in the “brush” stage,
preferably in disbursed, irregular patches within more advanced forest.
The use of shrubs as well as all other types of native plants for reclamation and
restoration of damaged sites is becoming a very important topic (Hansen 1989). Shrubs are
planted as seedlings of various types and seeded using the same techniques employed with
trees (Alder and Ostler 1989), except that densities must often be higher. Because it costs
less, establishment by site manipulation and natural seeding and succession is preferred
whenever possible.
Shrubs yield many benefits to humankind directly and indirectly. Berries and similar
small fruits are the most important shrub-derived foods. There are many hundreds of kinds
of edible, wild berrylike fruits throughout the world. Seasonally harvested and preserved,
they were once very important to hunter-gatherer tribes and still are important in certain
rural areas. All our commercial berries descended from wild shrubs, and their wild ancestors
remain a source of genetic material for breeders. Wildland shrubs also furnish nuts, seeds,
herbs, greens, and medicinal materials to rural peoples.
Fuel is another major direct benefit from shrubs. Although wood from shrubs is not
present in quantities as great as tree wood in forests, its accessibility and ease of harvest
have made it a very important fuel source in underdeveloped areas and during recreational
camping, especially when collected by women and children. Shrubs are even harvested to
make charcoal in areas with few trees. It has also been suggested that shrub stands could be
harvested mechanically for industrial biomass fuel (Young and others 1989).
Humans benefit indirectly from shrubs through animals that eat them. Grazing
domestic goats, camels, and, to a lesser extent, sheep and cattle obtain a large part of their
forage from woody browse. Many wild ruminants preferentially browse shrubs even when
grass is available. Others supplement their diet of grasses and forbs with forage from shrubs.
A multitude of birds, mammals, amphibians, and reptiles obtain forage, mast (fruits and
seeds), and insects from shrub hosts. The cover provided by shrubs is critical for a vast
number of wild animals.
5
Finally, wild shrubs are an important esthetic component of our forests, prairies, and
deserts. As greenbelts and semi-wild urban parks become more widespread, wild shrubs will
be more appreciated and will be managed along with trees and herbaceous plants.
Shrubs also negatively impact humans. Undesirable shrubs often invade managed
and semi-managed pastures, excluding the more desirable grass, and become weeds in
croplands. They can temporarily suppress tree seedlings and slow reforestation. Shrubs,
especially the many thorny and a few poisonous species, are the bane of cross-country hikers
and woods workers. Forests with a heavy shrub layer are usually less visually pleasing than
open forests. Shrub understories often are a vehicle for explosive fire spread in seasonally
dry forest habitats.
In the early days of forestry, it seemed to be enough to protect and manage the forest
as a whole. Later, it was realized that we needed detailed information on each of the tree
species. To this end, a number of excellent references have been produced (Burns and
Honkala 1990a, Burns and Honkala 1990b, Burns and others 1998, Francis and Lowe 1999).
The time is coming when, if we are to manage the lesser species, we will need detailed
information on each or, at least, the most important shrubs. The objective of this work is to
6
provide an accessible reference of the biology, ecology, and management of individual
species of shrubs. The study of the biology, ecology, and management of trees is called
silvics, and the resulting descriptions are termed silvical descriptions. No such terms have
come to general use for the study and description of shrubs. Perhaps the terms should be
designated “thamnics” and “thamnical descriptions” from the Ancient Greek “thamnos”,
which means shrub.
References
Abrams, L. 1940. Illustrated flora of the Pacific states. Vol. 1. Stanford University Press,
Stanford, CA. 538 p.
Abrams, L. 1944. Illustrated flora of the Pacific states. Vol. 2. Stanford University Press,
Stanford, CA. 635 p.
Abrams, L. 1951. Illustrated flora of the Pacific states. Vol. 3. Stanford University Press,
Stanford, CA. 866 p.
Abrams, L., and R.S. Ferris. 1960. Illustrated flora of the Pacific states. Vol. 4. Stanford
University Press, Stanford, CA. 732 p.
Alder, G.M., and W.K. Ostler. 1989. Native shrub propagation and nursery stock
production. In: C.M. McKell, ed. The biology and utilization of shrubs. Academic Press,
Inc., San Diego, CA. p. 535-552.
Allaby, M. 1994. The concise Oxford Dictionary of ecology. Oxford University Press,
Oxford, UK. 415 p.
Benson, L. 1969. The cacti of Arizona. The University of Arizona Press, Tucson, AZ. 218 p.
7
Burns, R.M., and B.H. Honkala, eds. 1990a. Silvics of North America. Vol. 1, Conifers.
U.S. Department of Agriculture, Forest Service, Agriculture Handbook 654. Washington,
DC. 675 p.
Burns, R.M., and B.H. Honkala, eds. 1990b. Silvics of North America. Vol. 2, Hardwoods.
U.S. Department of Agriculture, Forest Service, Agriculture Handbook 654. Washington,
DC. 877 p.
Burns, R.M., M.S. Mosquera, and J.L. Whitmore, eds. 1998. Useful trees of the tropical
region of North America. Publication 3. North American Forestry Commission,
Washington, DC. 256 p.
Correll, D.S., and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas
Research Foundation, Renner, TX. 1881 p.
Davis, R.J. 1952. Flora of Idaho. W.M.C. Brown Co., Dubuque, IA. 828 p.
Degener, O. 1946. Flora Hawaiiensis. Books 1-4. Otto Degener, Waialua, Oahu, HI. [Not
paged].
Degener, O. 1957. Flora Hawaiiensis. Book 5. Otto Degener, Waialua, Oahu, HI. [Not
paged].
Degener, O. and I. Degener. 1963. Flora Hawaiiensis. Book 6. Otto Degener, Waialua,
Oahu, HI. [Not paged].
Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs and cacti of South Texas. Texas Tech
University Press. 213 p.
Ford-Robertson, F.C., ed. 1971. Terminology of forest science, technology, practice, and
products. Society of American Foresters, Washington, DC. 349 p.
8
Francis, J.K. and H.A. Liogier. 1991. Naturalized exotic tree species in Puerto Rico. U.S.
Department of Agriculture, Forest Service, General Technical Report SO-82. Southern
Forest Experiment Station, New Orleans. 12 p.
Francis, J.K. and C.A. Lowe., eds. 1999. Bioecología de especies natives y exóticas de
Puerto Rico y las Indias Occidentales. U. S. Department of Agriculture, Forest Service,
General Technical Report IITF-15. International Institute of Tropical Forestry. Río
Piedras, PR. 583 p.
Great Plains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas.
1391 p.
Hansen, D. 1989. Reclamation and erosion control using shrubs. In: McKell, C. M., ed. The
biology and utilization of shrubs. Academic Press, Inc, San Diego. 459-478.
Jones, F.B. 1975. Flora of the Texas coastal bend. Mission Press, Corpus Christi, TX. 262 p.
Liogier, H.A. 1985. Descriptive flora of Puerto Rico and adjacent islands. Vol. 1. Editorial
de la Universidad de Puerto Rico, San Juan, PR. 352 p.
Liogier, H.A. 1988. Descriptive flora of Puerto Rico and adjacent islands. Vol. 2. Editorial
de la Universidad de Puerto Rico, San Juan, PR. 481 p.
Liogier, H.A. 1994. Descriptive flora of Puerto Rico and adjacent islands. Vol. 3. Editorial
de la Universidad de Puerto Rico, San Juan, PR. 461 p.
9
Liogier, H.A. 1995. Descriptive flora of Puerto Rico and adjacent islands. Vol. 4. Editorial
de la Universidad de Puerto Rico, San Juan, PR. 617 p.
Liogier, H.A. 1997. Descriptive flora of Puerto Rico and adjacent islands. Vol. 5. Editorial
de la Universidad de Puerto Rico, San Juan, PR. 436 p.
Little, E.L., Jr. 1979. Checklist of United States trees (native and naturalized). U.S.
Department of Agriculture, Forest Service, Agriculture
Handbook 541. Washington, DC. 375 p.
Little, E.L., Jr., R.O. Woodbury, and F.H. Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. U.S. Department of Agriculture, Forest Service, Agriculture
Handbook 449. Washington, DC. 1,024 p.
Little, E.L., Jr. and R. Skolmen. 1989. Common forest trees of Hawaii (native and
introduced). U.S. Department of Agriculture, Forest Service, Agriculture Handbook 679.
Washington, DC. 321 p.
Long, R.W. and O. Lakela. 1971. A flora of tropical Florida. University of Miami Press,
Coral Gables, FL. 962 p.
10
Natural Resource Conservation Service. 2003. Plants database. http://plants.usda.gov/. [not
paged].
Nelson, Gil. 1996. The shrubs and woody vines of Florida. Pineapple Press, Inc., Sarasota,
FL. 391 p.
Orshan, G. 1989. Shrubs as a growth form. In: McKell, Cyrus M., ed. The biology and
utilization of shrubs. Academic Press, Inc., San Diego, CA. p. 249-265.
Rhoads, A.F., and W.M. Klein, Jr. 1993. The vascular flora of Pennsylvania, annotated
checklist and atlas. American Philosophical Society, Philadelphia, PA. 636 p.
Viereck, L.A., and E.L. Little Jr. 1972. Alaska trees and shrubs. Agriculture Handbook 410.
U.S. Department of Agriculture, Forest Service, Washington, DC. 265 p.
Young, J.A., J.D. Budy, and R.A. Evans. 1989. Use of shrubs for fuel. In: C.M. McKell, ed.
The biology and utilization of shrubs. Academic Press, Inc, San Diego, CA. p. 479-492.
11
Abrus precatorius L. crab’s eye
FABACEAE
John K. Francis
12
crab’s eye is not reported, but certainly it lives Southern Forest Experiment Station, New
longer than 3 years. New plants are easily grown Orleans, LA. 331 p.
in the nursery and probably can be established by
direct seeding into prepared seed spots. Because Bailey, L.H. 1941. The standard cyclopedia of
the species is exotic in all of the U.S. Territories horticulture. Vol. 1. The MacMillan Co., New
and because it becomes a weed in range and York. 1,200 p.
semimanaged pastures (Velez and von Overgeek
1950), it is difficult to see a justification for the Howard R.A. 1988. Flora of the Lesser Antilles,
establishment of plantations. Leeward and Windward Islands.
Dicotyledoneae. Vol. 4. Arnold Arboretum,
Benefits.—Crab’s eye seeds were used anciently Harvard University, Jamaica Plain, MA. 673 p.
as balance weights in the Eastern countries (Neal
1965) and as rosary beads by the Buddhists Liogier H.A. 1988. Descriptive flora of Puerto
(Bailey 1941) because they were so uniform in Rico and adjacent islands, Spermatophyta. Vol.
weight, as well as durable. Today, the seeds are 2. Editorial de la Universidad de Puerto Rico,
used to make necklaces and other jewelry and for Río Piedras, PR 481 p.
the sounding weights in maracas. The seeds
contain a protein-based poison called abrin Ndamba, J., N. Nyazema, N. Makaza, C.
(Parrotta 2001). One seed contains enough poison Anderson, and K.C. Kaondera. 1994. Traditional
to kill a human, but the seeds are very hard and herbal remedies used for the treatment of urinary
must be chewed or ground to release the poison schistosomiasis in Zimbabwe. Journal of
(Acevedo-Rodríguez 1985). However, cooking Ethnopharmacology 42(2): 125-132.
destroys the poison so that the seeds may be eaten
(Neal 1965). The leaves and roots contain Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
glycyrrhizin, the principal component of licorice. 50. Bernice P. Bishop Museum Press, Honolulu,
These tissues prepared in various ways are used to HI. 924 p.
treat coughs and a number of other ailments
(Parrotta 2001). Crab’s eye holds out promise in Parrotta, J.A. 2001. Healing plants of Peninsular
the treatment of Schistosoma haematobium India. CAB International, Wallingford, UK.
infections. Extracts of the plant were shown to be 944 p.
lethal to adult schistosomes in hamsters (Ndamba
and others 1994). Crab’s eye vines are sometimes Pokhriyal, T.C., S.P. Chaukiyal, and H.B. Naithani.
grown for vegetative screens (Bailey 1941). 1997. Nodule biomass and nitrogen fixation
studies in some leguminous nitrogen fixing
References plants. Indian Forester 123(12): 1197-1198.
Acevedo-Rodríguez, P. 1985. Los bejucos de Vélez, I. and J. von Overgeek. 1950. Plantas
Puerto Rico. Vol. 1. Gen. Tech. Rep. SO-58. indeseables en los cultivos tropicales. Editorial
U.S. Department of Agriculture, Forest Service, Universitaria, Rìo Piedras, PR. 497 p.
13
Acacia farnesiana (L.) Willd. sweet acacia
FABACEAE
John A. Parrotta
14
seed, although branch cuttings can also be rooted vegetation for food production in arid zones.
(Webb and others 1980). A collection of seeds General Technical Report RM-150. U.S.
from Puerto Rico contained 7,600 seeds/kg, began Department of Agriculture, Forest Service,
germinating in 6 days, and achieved 57 percent Rocky Mountain Forest and Range Experiment
germination after 13 days (Francis and Rodríguez Station, Fort Collins, CO. p. 90-98.
1993). Although pregermination treatment is not
necessary, cold- or hot-water soaking, chemical Francis, J.K. and A. Rodríguez. 1993. Seeds of
scarification, and, particularly, seed-coat Puerto Rican trees and shrubs: second
scarification by abrasion with sandpaper greatly installment. Research Note SO-374. U.S.
increase germination rates (Parrotta 1992). Department of Agriculture, Forest Service,
Southern Forest Experiment Station, New
Growth and Management.—Early growth is Orleans, LA. 5 p.
relatively rapid. About 1 m of height growth can
be obtained during the first year, although growth Hughes, C.E. and B.T. Styles. 1984. Exploration
rates of 30 to 50 cm during the first year under and seed collection of multi-purpose dry zone
semiarid field conditions are more typical trees in Central America. International Tree
(Foroughbakhch and others 1987). Depending on Crops Journal 3: 1-31.
the environment, maximum heights of plants
generally range from 3 to 5 m, with stem Liogier, H.A. 1990. Plantas medicinales de Puerto
diameters up to 5 cm. Because sweet acacia is Rico y del Caribe. Iberoamericana de Ediciones,
intolerant of shade and does not compete well with Inc., San Juan, PR. 566 p.
more aggressive woody vegetation such as
Prosopis L., management activities for enhancing Little, E.L., Jr. and F.H. Wadsworth. 1964.
growth and natural regeneration in natural and Common trees of Puerto Rico and the Virgin
plantation stands may include control of Islands. Agriculture Handbook 249. U.S.
competing vegetation and periodic soil Department of Agriculture, Forest Service,
disturbance. Washington, DC. 548 p.
Benefits.—Sweet acacia is planted in many parts Parrotta, J.A. 1992. Acacia farnesiana (L.) Willd.
of its natural and introduced tropical and –Aroma, huisache. Research Note SO-ITF-SM-
subtropical range for reforestation of degraded 49. U.S. Department of Agriculture, Forest
drylands, for fuelwood and small timber, livestock Service, Southern Forest Experiment Station,
fodder, and for its flowers used in the perfume New Orleans. 6 p.
industry. In some areas, it is considered a pest due
to its ability to colonize pastures and other Parrotta, J.A. 2001. Healing plants of peninsular
disturbed habitats. The tannin-rich bark is used for India. CAB International, New York. 944 p.
tanning leather, the gum obtained from the cut
bark is used as a substitute for gum arabic (from Rzendowski, J. 1981. Vegetacíon de México.
Acacia nilotica (L.) Delile), and a useful black dye Editorial Limusa, Mexico City. 432 p.
is obtained from the pods. Various parts of the
plant are used in traditional medicine (Liogier Siegler, D.S., S. Seilheimer, J. Keesy, and
1990, Parrotta 2001). In Mexico, for example, the H.F.Huang. 1986. Tannins from four common
flowers are used to treat headache and indigestion, Acacia species of Texas and northeastern
whereas a decoction of the green pods is used to Mexico. Economic Botany 40: 220-232.
treat dysentery and skin inflammations. In India,
the bark, heartwood, and leaves are all used Webb, D.B., P.J. Wood, and J. Smith. 1980. A
medicinally to treat a variety of ailments (Parrotta guide to species selection for tropical and
2001). subtropical plantations. Tropical Forestry Paper
15. Overseas Development Administration,
References Commonwealth Forestry Institute, University of
Oxford, London. 256 p.
Fourouhbakhch, R., R. Penaloza, and H. Stienen.
1987. Increasing productivity in the matorral of
northeastern Mexico: domestication of ten
native multipurpose tree species. In: Strategies
for classification and management of native
15
Acacia greggii Gray catclaw acacia
FABACEAE
Juanita A. R. Ladyman
16
pollinated (Bowers 1993). The fruit is a light Vines 1960). It is a food plant for both adult and
brown to reddish legume often becoming larval butterflies (Taylor and others 1997) and an
constricted between the seeds. The circular seeds important “bee plant” (Dayton 1931). The lac
are 5- to 7-mm in diameter (Isely 1973). The wax- insect, Tachardia lacca, feeds on the sap of
coated seeds delay germination for several years catclaw acacia and produces a resinous substance
and need to be scarified to germinate (Bowers that can be used in making varnish and shellac but
1993, Epple 1995). Regions where catclaw acacia probably not in commercial quantities (Powell
grows are prone to flash floods and the tumbling 1998). The wood has been used for fuel and tool
action of floodwaters in sandy, gravelly washes handles (Epple 1995). It is also in the ornamental
accomplish both seed scarification and seed and landscape trade and is a good hedge plant
dispersal (Bowers 1993). (Taylor and others 1997). It has been used in
projects to re-vegetate degraded land but is
Growth and Management.—Catclaw acacia is a sensitive to some pollutants such as those found in
thicket-forming shrub that has increased in copper mine wastes (Norem and others 1982).
abundance throughout the Southwestern United
States since the 1890s (Bahre 1995). This increase References
is likely influenced if not directly caused by
livestock overgrazing and fire suppression (Bahre Bahre, C.J. 1995. Human impacts on the
1995). Overgrazing has reduced native perennial Grasslands of southeastern Arizona. In: M.P.
grasses that, when healthy and dense, can reduce McClaran and T.R. Van Devender, eds. The
woody shrub seedling establishment and also Desert Grassland. The University of Arizona
provide sufficient fuel to carry a fire. Catclaw is Press, Tucson, AZ. p. 230-264.
susceptible to fire when young, but mature plants
re-sprout from the root crown following fire Blake, J.G. 1984. A seasonal analysis of bird
(Carmichael and others 1978, McPherson 1995). communities in southern Nevada. The
Catclaw is moderately resistant to phenoxy Southwestern Naturalist. 29(4): 463-474.
herbicides, and after only one application the
shrubs tend to re-foliate or re-sprout from the Bowers, J.E. 1993. Shrubs and trees of the
crown (Hibbert and others 1974). Successive Southwest Deserts. Southwest Parks and
herbicide applications kill the plants. Apparently Monuments Assoc., Tucson, AZ. 140 p.
the invasive resinbrush, Euryops multifidus
(Thunb.) DC. out-competes, or is in some other Carmichael, R.S., O.D. Knipe, C.P. Pase, and W.
way deleterious to, catclaw shrubs and may be a W. Brady. 1978. Arizona chaparral: Plant
threat in some parts of its range (McAuliffe 1995). associations and ecology. Research Paper RM-
202. U.S. Department of Agriculture, Forest
Benefits.—Catclaw acacia shrubs provide Service, Rocky Mountain Forest and Range
protective cover and shade for wildlife (Dayton Experiment Station, Fort Collins, CO. 17 p.
1931). Pods, twigs and foliage, especially new
growth, provides browse for a variety of wildlife, Dayton, W.A. 1931. Important western browse
for example jack rabbit, white-tailed and mule plants. Miscellaneous Publication 101. U.S.
deer, and the seeds are eaten by many species of Department of Agriculture, Washington, DC.
small mammals and birds, for example, ground 214 p.
squirrels, western white-winged dove, scaled and
bobwhite quail (Graham 1941, Everitt and Drawe Epple, A.O. 1995. A field guide to the plants of
1993, Taylor and others 1997). The leaves contain Arizona. LewAnn Publishing Company, Mesa,
16 to 20 percent crude protein (Everitt and Drawe AZ. 347 p.
1993). Cattle browse the shrubs especially early in
the spring before the new leaves appear, when the Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs
young green twigs provide some relatively and cacti of South Texas. Texas Tech University
succulent forage for a short period of time, and Press, Lubbock, TX. 231 p.
when other forage is limited (Graham 1941,
USDAFS 1988). The mature foliage and pods are Gledhill, D. 1992. The names of plants.
unpalatable to livestock (Graham 1941, Vines Cambridge University Press, Cambridge, UK.
1960). The fruits were ground into a flour, or meal, 202 p.
called pinole by Native Americans (Epple 1995,
17
Graham, E.H. 1941. Legumes for erosion control Norem, M.A., A.D. Day, and K.L. Ludeke. 1982.
and wildlife. Miscellaneous Publication 412. An evaluation of shrub and tree species used in
U.S. Department of Agriculture, Washington, revegetating copper mine wastes in the south-
DC. 153 p. western United States. Journal of Arid
Environments 5: 299-304.
Hibbert, A.R., E.A Davis, and D.G. Scholl. 1974.
Chaparral conversion potential in Arizona: Part Powell, A.M. 1998. Trees & shrubs of Trans-
I: water yield response and effects on other Pecos and adjacent areas. University of Texas
resources. Research Paper RM-126. U.S. University Press, Austin, TX (First ed. copyright
Department of Agriculture, Forest Service, Big Bend Natural History Assoc.) 498 p.
Rocky Mountain Forest and Range Experiment
Station, Fort Collins, CO. 36 p. Richardson, A. 1995. Plants of the Rio Grande
Delta. University of Texas Press, Austin, TX.
Isely, D. 1973. Leguminosae of the United States: 426 p.
I. Subfamily Mimosoideae. Memoirs of the New
York Botanical Garden 25(1) 1-152. Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox,
J.O. Klemmedson, J.J. Norris, and L.C. Fierro.
Kartesz, J.T. 1994. A synonymized checklist of the 1992. Desert grasslands. In: Ecosystems of the
vascular flora of the United States, Canada and World. 8A. Natural Grasslands Introduction and
Greenland. Vol. Checklist. Second Ed. Timber Western Hemisphere. R.T. Coupland, ed.
Press Portland, OR. 622 p. Elsevier, London, UK. p. 337-362.
McAuliffe, J.R. 1995. Landscape evolution, soil Taylor, R.B., J. Zrutledge, J.G. Herrera. 1997. A
formation, and Arizona’s desert grasslands. In: field guide to common south Texas shrubs.
The Desert Grassland. M.P. McClaran and T.R. Texas Parks and Wildlife Press, Austin, TX.
Van Devender, eds. The University of Arizona 106 p.
Press, Tucson, AZ. p.100-129.
Uchytil, R.J. 1990. Acacia greggii. U.S.
McClaran, M.P. 1995. Desert grasslands and Department of Agriculture, Forest Service,
grasses. In: The Desert Grassland. M.P. Rocky Mountain Research Station, Fire Sciences
McClaran and T.R. Van Devender, eds. The Laboratory. Fire Effects Information System.
University of Arizona Press, Tucson, AZ. p. 1- http://www.fs.fed.us/database/feis/ [not paged].
30.
United States Department of Agriculture Forest
McPherson, G.R. 1995. The role of fire in desert Service. 1988. Range Plant Handbook. Dover
grasslands. In: The Desert Grassland. M.P. Publications, Inc. New York. 837 p.
McClaran and T.R. Van Devender, eds. The
University of Arizona Press, Tucson, AZ. p. Vines, R.A. 1960. Trees, shrubs, and woody vines
130-151. of the Southwest. Sixth printing. University of
Texas Press. Austin, TX. 1,104 p.
18
Acacia retusa (Jacq.) Howard zarza brava
FABACEAE
John K. Francis
19
December (Acevedo-Rodriguez 1985) and ripens scrambling patches in overgrazed cattle pasture. On
fruit in January and July. Fruit production can be the other hand, the species is a great nuisance to the
very heavy, and pods in one sample in Puerto Rico cattleman, woods workers, and hikers. It is
averaged 7.97 + 0.29 seeds/pod. The seeds frequently necessary to cut a path through zarza
averaged 0.041 + 0.001 g/seed or 24,000 seeds/kg. brava thickets with a machete, in order to move
Some of the seeds had been attacked by an through vegetation in the semiarid-zones within the
unknown species of beetle and a lepidopteran zarza brava range. When one contacts the vines,
larva. When planted in commercial potting mix, the the tips of the spines break off in the skin, causing
seeds began germinating in 4 days and by 11 days pain and sometimes resulting in infection. No
had reached their final 98 percent germination. specific control recommendations have been
Germination is epigeal. In the wild, seedlings are developed. In the past, control was attempted with
produced in abundance, but few survive beyond the mixed success by repeated cutting. Control could
first year. Zarza brava sprouts vigorously and probably be improved by treating the sprouts with
grows rapidly after fires or cutting. herbicides after cutting.
20
Acacia tortuosa (L.) Willd. twisted acacia
FABACEAE
John K. Francis
do not open. The ovoid seeds are brown and hard
(Howard 1988, Liogier 1988, Little and others
1974). The chromosome number is 2n = 26 (Long
and Lakela 1971).
21
1974). Bees and other insects pollinate the flowers
(Treeguide 2002). A collection of pods from Carambi Foundation. 2002. Vegetation. http://
Puerto Rico contained an average of 12.4 www.curacao.com/carambi/veget.htm. 1 p.
seeds/pod and a range of eight to 15 seeds/pod. The
air-dried seeds separated from them averaged
0.0440 + 0.0004 g/seed or 22,700 seeds/kg. These Howard R.A. 1988. Flora of the Lesser Antilles,
were mechanically scarified (nicked) and incubated Leeward and Windward Islands.
on moist blotter paper. Ninety-nine percent Dicotyledoneae, Part 1. Vol. 4. Arnold
germinated between 5 and 21 days after sowing Arboretum, Harvard University, Jamaica Plain,
(author’s observation). Because of the hard, MA. 673 p.
impermeable seed coat, germination without
scarification is low and erratic. Any technique that Institute of Systematic Botany. 2002. Atlas of
perforates the seed coat will work. One source Florida Plants: Acacia tortuosa. University of
(Treeguide 2002) recommends placing the seeds in South Florida, Tampa, FL.
boiling water, immediately removing the heat http://plantatlas.usf.edu/main.asp?plantID=1545.
source and letting the seeds soak for 24 hours. 1 p.
Germination is epigeal. The seeds are dispersed by
herbivores, particularly ruminants that eat the pods. Instituto Humboldt. 2002. Plantas vasculares:
Seedlings often emerge in large numbers, but angiospermas y gimnospermas. http://www.
nearly all are consumed by grazing animals before humboldt.org.co/diqadeladiversidad/resultados/v
they have a chance to become woody and thorny. asculares.html. 6 p.
Growth and Management.—Although acacias Island Resource Foundation. 2002. Threatened and
can be propagated with cuttings, the use of seeds is endangered birds of the Insular Caribbean.
recommended (Bonsai Clubs International 2002). http://www.irf.org/bbarden.htm. 13 p.
No management experience has been published;
however, nursery production and field Liogier H.A. 1988. Descriptive flora of Puerto
establishment will probably be as easy as it is for Rico and adjacent islands, Spermatophyta. Vol.
other acacias. Control of twisted acacia along with 2. Editorial de la Universidad de Puerto Rico,
other secondary dry-forest species has been Río Piedras, PR. 481 p.
accomplished in several areas in Puerto Rico and
the U.S. Virgin Islands by bulldozing. The use of Little, E.L., Jr., R.O. Woodbury, and F.H.
heavy disks might be a better option where Wadsworth. 1974. Trees of Puerto Rico and the
removal of woody debris is not required. Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Benefits.—Twisted acacia helps protect the soil in Washington, DC. 1,024 p.
its disturbed habitat. It is listed as a nitrogen-fixing
species (Winrock International 2002). The seeds Long, R.W. and O. Lakela. 1971. A flora of
are eaten by the threatened yellow-shouldered tropical Florida. University of Florida Press,
amazon parrot (Amazona barbadensis) in Bonaire Coral Gables, FL. 962 p.
and probably other parts of its range (Island
Resource Foundation 2002). The species serves as Missouri Botanical Garden. 2002. Current
cover for wildlife species. Cattle and other specimen list for Acacia tortuosa.
livestock eat the pods while goats and sheep http://mobot.mobot.org/cgi-bin/search_vast. [not
browse the foliage, especially when other forage is paged].
scarce. The wood is used to a limited extent for
fuel. Twisted acacia plants are used to create Nature Conservancy. 2002. The history of the
bonsai plants (Bonsai Clubs International 2002). Plains and its biota.
The species has been grown and pruned for hedges http://www.greatplains.org/resource/biodiver/bio
(Little and others 1974). stat/ecoregio.htm. 13 p.
22
Texas A&M Herbarium. 2002. Checklist of the Species.asp?SpeciesID=108&Region=NorthAme
vascular plants of Texas. http://csdl.tamu.edu/ rica. 2 p.
FLORA/cgi/msproj/shao/query.pl?query=Acacia
+tortuosa &rform=. 1 p. Winrock International. 2002. Nitrogen fixing trees
and shrubs. http://winrock.org/forestry/factpub/
Treeguide. 2002. Huisachillo, Acacia tortuosa, nftlist.htm. 29 p.
Willd. Fabaceae. http://treeguide.com/nn/
23
Acer glabrum Torr. Rocky Mountain maple
ACERACEAE
John K. Francis
24
autumn. Good seed crops occur every 1 to 3 years. shoots may be cooked and eaten as a vegetable
There are about 30,000 cleaned seeds/kg (Zasada (Pratt and others 2003). Native Americans made
and Strong 2003). Seeds disperse by whirling infusions and decoctions of the twigs and wood to
sideways as they descend, often carried by the treat swelling and nausea, respectively (Moerman
wind. Seeds sometimes lie dormant for one or two 1986). Rocky Mountain maple is widely used in
growing seasons before germinating. Germination environmental restoration projects within its native
is epigeal. Natural seedlings can be common to range and is an important ornamental species
abundant. valued for its petite size, tolerance to shade, and
bright red, yellow, and orange autumn colors
Growth and Management.—Rocky Mountain (Hansen 2003).
maple seedlings grow about 30 cm/year (Hansen
2003). On good sites sprouts may reach 1.3 m References
within 2 years and 3 m in 10 years. Maximum
heights are reached in 30 to 40 years (Anderson Abrams, L. 1951. Illustrated flora of the Pacific
2001). Fruits can be picked by hand from low States. Vol. 3. Stanford University Press,
shrubs or clipped or shaken from tall shrubs and Stanford, CA. 866 p.
small trees. Mechanical dewinging may be
advantageous to reduce bulk and improve handling Anderson, M.D. 2001. Acer glabrum. In: U.S.
before planting. The seeds can be stored for up to Department of Agriculture, Forest Service,
3 years (Hansen 2003). Stratification of 180 days Rocky Mountain Research Station, Fire Sciences
at 20 to 30 °C followed by 180 days at 3 to 5 °C is Laboratory, Fire Effects Information System.
recommended before sowing. Germination of http://www.fs.fed.us/database/feis/plants/shrub/
about 40 percent can be expected in about 30 days acegla/all.html. 62 p.
(Zasada and Strong 2003). Alternately, seeds are
sown after collection and beds are maintained for 2 Hansen, W.W. 2003. Native plants of the
years because a substantial amount of the Northwest: Acer glabrum (Douglas maple,
germination will occur after the second winter. Rocky Mountain maple, mountain maple).
Seeds are sown 0.6 to 2.5 cm deep and mulched in http://www.nwplants.com/plants/trees/aceraceae
shaded beds. Seedlings are produced as bare-root /acer_glabrum/ 5 p.
stock or containerized plants (Zasada and Strong
2003). The species can be propagated by layering Johnson, F.D. 1995. Wild trees of Idaho.
and rooted cuttings (Hansen 2003). University of Idaho Press, Moscow, ID. 212 p.
Benefits.—Rocky Mountain maple is an important Moerman, D.E. 1986. Medicinal plants of Native
part of the forest understory and pioneer America. Technical Reports 19. The Museum of
vegetation following fires and clearcuts. It helps Anthropology, University of Michigan, Ann
protect the soil, adds beauty to the forest, and Arbor, MI. 534 p.
furnishes food and cover for wildlife. It is an
important browse species for domestic livestock, Natural Resources Conservation Service. 2003.
especially sheep, and wild ungulates. Samples of Plants profile: Acer glabrum Torr., Rocky
summer growth contained about 8.7 percent crude Mountain maple. http://plants.usda.gov/
protein, 3.1 percent fat, 34.0 percent crude fiber, cgi_bin/plant_profile.chi?symbol=ACGL. 5 p.
51.0 percent N-free extract, and 3.2 percent ash.
Samples of winter browse contained 5.9 percent Pratt, M., J. Bowns, R. Banner, and A. Rasmussen.
crude protein, 2.4 percent fat, 33.3 percent crude 2003. Range plants of Utah: Rocky Mountain
fiber, and 54.2 percent N-free extract (Anderson maple. Utah State University Extension.
2001). The seeds and vegetative parts are http://extension.usu.edu/coop/natres/range/
consumed by ruffed and blue grouse, grosbeaks, Woody/rockymtnmaple.htm. 4 p.
and small mammals (Anderson 2001). The wood is
hard, close-grained, and flexible. It was used by Wasser, C.H. 1982. Ecology and culture of
Native Americans for arrow shafts, snowshoe selected species useful in revegetating disturbed
frames, ceremonial equipment, hut frames, and lands in the West. FWS/OBS-82/56. Fish and
drying and smoking racks. The inner bark Wildlife Service, U.S. Department of Interior,
furnished material for mats and ropes. Today, the Washington, DC. 347 p.
wood is mainly used for firewood (Hansen 2003)
and smoking meat (Johnson 1995). The young
25
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Zasada, J.C. and T.F. Strong. 2003. Acer L. In:
Higgins (eds.). 1987. A Utah flora. Great Basin F.T. Bonner, and R.G. Nisley, eds. Woody Plant
Naturalist Memoirs 9. Brigham Young Seed Manual. U.S. Department of Agriculture,
University, Provo, UT. 894 p. Forest Service, Washington, DC. 24 p.
26
Acer spicatum Lam. mountain maple
ACERACEAE
Synonyms: none
Paula M. Pijut
27
damaging to mountain maple (Johnson and Lyon 1993). Propagation of mountain maple by cuttings
1991). The larvae feed on the roots, mature, and taken early in the summer produces strong plants
overwinter in the soil. Adults are brown weevils (van Gelderen and others 1994).
that defoliate mountain maple. They are found
throughout the northeast into Michigan and the Growth and Management.—Mountain maple is a
border provinces of Canada (Johnson and Lyon slow to medium growing shrub or small, short
1991). Kabatiella apocrypta (Ell. & Ev.) v. Arx (crooked) trunked tree of bushy appearance (Dirr
(anthracnose pathogen) causes minor damage to 1998). It is hardy in zones three to seven at high
mountain maple, although severe infections can elevations (USDA Plant Hardiness), and starts to
led to premature defoliation (Sinclair and others decline at 40 to 50 years of age (Sullivan 1993).
1987). Mountain maple is also susceptible to Mountain maple is girdled at the root collar by
attack by Verticillium dahilae Kleb. (verticillium low, surface fires, and can re-sprout from
wilt), Phyllactinia guttata (Wallr.:Fr.) Lev. underground stems (Sullivan 1993). A severe, hot
(powdery mildew), Nectria sp. (nectria canker), fire is necessary in order to kill the roots.
Cristulariella depraedens (Cooke) Hohn. (leaf Mountain maple can withstand repeated and heavy
spot/blight), Pseudomonas syringae pv. syringae browsing, producing new growth even when 80
van Hall (bacterial leaf spot and dieback), percent of the annual twig growth is removed each
Phyllosticta minima (Berk. and M.A.Curtis) year (Sullivan 1993). Mountain maple is an
Underw. and Earle (leaf spot), Rhytisma sp. (tar understory or subcanopy component in a number
spots), and Venturia acerina Plakidas ex Barr (leaf of northeastern forest associations (Sullivan 1993).
blotch) (Farr and others 1989, Jones and Benson Cutting mountain maple near the ground level,
2001, Sinclair and others 1987). clear-cutting, and spraying with 2,4-D in the
spring will increase the availability of new growth
Reproduction.—Mountain maple staminate for deer and moose browse (Sullivan 1993).
flowers and pistillate flowers occur on the same Burning can be used to suppress mountain maple
plant. Some individuals may flower completely in order to promote reproduction of other, more
with staminate flowers. Mountain maple is desirable tree species.
predominately duodichogamous (producing
flowers in three consecutive phases, male-female- Benefits.—Mountain maple is a nutritious browse
male, during anthesis), with a small part of the species for moose, white-tailed deer, cottontails,
population protogynous (stigma receptive before snowshoe hares, woodland caribou, beavers, and
the pollen is shed) (van Gelderen and others 1994). ruffed grouse (Sullivan 1993). Mountain maple is
Low temperatures appear to favor development of sometimes planted as an ornamental, but has little
female flowers and flowers with less reduction of landscape value excluding its brief fall color
the non-functioning parts. (van Gelderen and display and colorful samaras.
others 1994). The shrub blossoms in May to June
after the leaves are fully developed; fruit ripens in References
September to October, with seed dispersal in
October to December (Zasada and Strong 2000). De Jong, P.C. 1976. Flowering and sex expression
Mountain maple is insect pollinated (Sullivan in Acer L. A biosystematic study. Mededelingen
1983) and the seed is wind disseminated. Seeds of Landbouwhogeschool Wageningen 76 (2): 1-
mountain maple need to be stratified for 90 to 120 202.
days at 5 ºC for good germination (Dirr 1998).
Mountain maple does not build up a seed-bank and Dirr, M.A. 1998. Manual of woody landscape
seedling establishment is better on undisturbed plants, Their identification, ornamental
soils (Sullivan 1993). Seedling reproduction of characteristics, culture, propagation, and uses.
mountain maple does not result in dense, Stipes Publishing, Champaign, IL. 1,187 p.
competitive stands, as can be the case with its
vegetative reproduction. Mountain maples Farr, D.F., G.F. Bills, G.P. Chamuris, and A.Y.
frequently layer, giving them the potential to Rossman. 1989. Fungi on plants and plant
develop relatively complex clones of varying size products in the Untied States. The American
and morphology (Zasada and Strong 2000). Phytopathological Society, St. Paul, MN.
Mountain maple can reproduce by sprouting from 1,252 p.
underground lateral stems, rarely from root
suckers, and colonies usually develop following Gleason, H.A. and A. Cronquist. 1991. Manual of
disturbances by browsing or cutting (Sullivan vascular plants of northeastern United States and
28
adjacent Canada. The New York Botanical Sullivan, J. 1993. Acer spicatum. In: U.S.
Garden, Bronx, NY. 910 p. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences
Johnson, W.T. and H.H. Lyon. 1991. Insects that Laboratory. Fire Effects Information System.
feed on trees and shrubs. Cornell University http://www.fs.fed.us/
Press, Ithaca, NY. 560 p. database/feis/plants/shrub/acespi/index.html.
14 p.
Jones, R.K. and D.M. Benson. 2001. Diseases of
woody ornamentals and trees in nurseries. The USDA, NRCS. 2002. The PLANTS Database,
American Phytopathological Society, St. Paul, Version 3.5 National Plant Data Center, Baton
MN. 482 p. Rouge, LA. http://plants.usda.gov. [not paged].
Sinclair, W.A., H.H. Lyon, and W.T. Johnson. van Gelderen, D.M., P.C. de Jong, and H.J.
1987. Diseases of trees and shrubs. Cornell Oterdoom. 1994. Maples of the World. Timber
University Press, Ithaca, NY. 575 p. Press, Portland, OR. 458 p.
Sullivan, J.R. 1983. Comparative reproductive Zasada J.C. and T.F. Strong. 2000. Acer L. In:
biology of Acer pensylvanicum and A. spicatum Woody plant seed manual. F.T. Bonner, tech.
(Aceraceae). American Journal of Botany 70(6): coord., R.G. Nisley, ed. U.S. Department of
916-924. Agriculture, Forest Service, Washington, DC.
http://www.wpsm.net/ Acer.pdf. [not paged].
29
Acnistus arborescens (L.) Schlecht. gallinero
SOLANACEAE
John K. Francis
30
Rico averaged 0.2025 + 0.0084 g/fruit. Seeds Haber, W.A. 2002. Acnistus arborescens (L.)
separated from them weighed an average of Schltdl., Solanacear-Potatoe Family.
0.00074 + 0.00017 g/seed or 1,350,000 seeds/kg. http://www.cs.umb.edu/~whaber/Monte/Plant/
Sown on commercial potting mix, 88 percent Sola/Acn-arb. htm. 2 p.
germinated between 11 and 35 days after sowing.
Some of the seedlings were killed by “damping Howard, R.A. 1989. Flora of the Lesser Antilles,
off” fungi (species unknown). A fruiting tree Leeward and Windward Islands. Vol. 6. Arnold
produces an enormous number of seeds. Birds and Arboretum, Harvard University, Jamaica Plain,
bats eat the fruits and disperse the seeds (Engriser MA. 658 p.
1995). Layering (rooting) occurs whenever the
trunk or branches come in contact with the ground; Kupchan, S.M., W.K. Anderson, P. Bollinger,
sprouts from prostrate trunks and branches R.W. Doskotch, R.M. Smith, J.A. Saenz-
sometimes become independent plants. Renauld, H.K. Schnoes, A.L. Burlingaure, and
D.H. Smith. 1969. Tumor inhibitors. XXXIX.
Growth and Management.—Gallinero has a Active principles of Acnistus arborescens.
moderate growth rate and is relatively short-lived Isolation and structural and spectral studies of
(10 to 20 years). Nursery and management Withaferin A and Withacnistin. Journal of
experience has not been published. Organic Chemistry 34(12): 3,858-3,866.
31
Aegiphila martinicensis Jacq. Caribbean spiritweed
VERBENACEAE
John K. Francis
32
m/year. Stems may live 5 years or more. No Universidad de Puerto Rico, San Juan, PR.
planting or management experience has been 617 p.
published.
Little, E.L., Jr., R.O. Woodbury, and F.H.
Benefits.—Carabbean spiritweed helps protect the Wadsworth. 1974. Trees of Puerto Rico and the
soil and furnishes food and cover for wildlife. Virgin Islands. Vol. 2. Agriculture Handbook
Larva of the hawk moth, Munduca hannibal 449. U.S. Department of Agriculture,
hannibal, apparently feed exclusively on Washington, DC. 1,024 p.
Caribbean spiritweed (Oehlke 2003). Infusions of
the leaves and twigs have been used as a diuretic Missouri Botanic Garden. 2003. Flora of Panama
and as a pleasant tonic for treating asthma (Núñez- checklist: Aegiphila martinicensis Jacq. http://
Melédez 1982). In Costa Rica, the plant is mobot.mobot.org/cgi-bin/search_pick. 1 p.
considered an aphrodisiac (Duke 2003).
Núñez-Melédez, E. 1982. Plantas medicinales de
References Puerto Rico. Editorial de la Universidad de
Puerto Rico, Río Piedras, PR. 498 p.
Duke, J. 2003. Dr. Duke’s phytochemical and
ethnobotanical databases: Tico ethnobotanical Oehlke, B. 2003. Manduca hannibal hannibal.
dictionary. http://www.ars-grin.gov/duke/ http://www.silkmoths.bizland.com/mhannhan.
dictionary/tico/a.html. [not paged]. htm. 4p.
Howard, R.A. 1989. Flora of the Lesser Antilles. Stevens, R.D., C. Ulloa-U., A. Pool, and O.M.
Vol. 6. Arnold Arboretum, Harvard University, Montiel, eds. 2001. Flora de Nicaragua.
Jamaica Plain, MA. 658 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanic Garden, St. Louis, MO.
Liogier, H.A. 1995. Descriptive flora of Puerto p. 911-2,666.
Rico and adjacent islands. Vol. 4. Editorial de la
33
Agave missionum Trel. corita
AGAVACEAE
John K. Francis
34
most plants never produce seeds. After the flowers former times.
fall off, bulbils form at the point where the flowers
were attached. Corita plants produce several References
hundred to several thousand bulbils each. A
collection of 100 bulbils from Puerto Rico weighed Breckon, G.J. and R.G. García. 2001. Vascular
(fresh) an average of 3.003 + 0.252 g/bulbil and plants of Susúa Forest. http://www.uprm.
ranged from 0.18 to 13.63 g. Placed with their edu/biology/profs.breckon/herbarium/florasusua.
bases slightly buried in moist potting mix, all of htm. 52 p.
them rooted within 1 week. They grew rapidly
under partial shade and were pricked into Britton, N.L. and P. Wilson. 1923. Botany of Porto
containers in about 3 weeks from sowing. After 2 Rico and the Virgin Islands, descriptive flora-
or 3 weeks they were moved to full sun and Spermatophyta. Scientific Survey of Porto Rico
reached about 15 cm in height in 2 additional and the Virgin Islands Vol. 5, Part 1. New York
months and were dispersed to the public (author’s Academy of Sciences, New York. 626 p.
observation). Plants originating from bulbils can be
numerous under fruiting plants. It is not known International Plant Name Index. 2002.
what specialized means of dispersal exist. International Plant Name Index Query.
However, new plants are frequently seen with no http://www.ipni.org/ipni/query_ipni.html. 1 p.
remains of flowering corita plants nearby.
Normally a plant dies after completing flowering, Liogier, H.A. and L.F. Martorell. 2000. Flora of
but many plants produce basal sprouts in the year Puerto Rico and adjacent islands, a systematic
of flowering, or before, that continue for another synopsis. 2nd ed. Editorial de La Universidad de
generation. Puerto Rico. 382 p.
35
Allamanda cathartica L. allamanda
APOCYNACEAE
John K. Francis
36
extend. In addition, trimmings dumped in vacant plantastoxicas.hpg.ig.com.br/toxicas/allcat.htm.
lots and wildlands root readily and start new 2 p.
colonies. Plants coppice vigorously when cut. The
species is invasive in Queensland, Australia Ecology and Evolutionary Biology Conservatory.
(Pacific Island Ecosystems at Risk 2002). 2002. Allamanda cathartica L. University of
Connecticut. http://florawww.eeb.uconn.edu/
Growth and Management.—Established acc_num/198500393.html. 3 p.
allamanda grows rapidly, adding 1 to 3 m to
extended length per year. Pruning is necessary to Floridata. 2002. Allamanda cathartica.
keep it under control in most landscaping http://www.floridata.com/ref/a/alla_cat.cfm. 3 p.
applications. There appears to be no reason to
establish it in wildlands, but the shrub could easily Howard, R.A. 1989. Flora of the Lesser Antilles,
be planted with cuttings placed directly in the soil. Leeward and Windward Islands. Dicotyledoneae.
Getting rid of spot infestations can be difficult. Part 3. Vol. 6. Arnold Arboretum, Harvard
Cutting is ineffective. For grubbing to work, University, Jamaica Plain, MA. 658 p.
removal of the plant stems and roots from the site
must be complete. Control with herbicides may Liogier, H.A. 1990. Plantas medicinales de Puerto
overcome some of these difficulties but is untested. Rico y del Caribe. Iberoamericana de Ediciones,
Inc. San Juan, PR. 566 p.
Benefits and Detriments.—In wild stands,
allamanda protects the soil and furnishes cover for Liogier, H.A. 1995. Descriptive flora of Puerto
wildlife. Year-long production of large, beautiful Rico and adjacent islands. Vol. 4. Editorial de la
flowers endears allamanda to gardeners and Universidad de Puerto Rico, San Juan, PR.
landscapers throughout the tropics and subtropics. 617 p.
The species is particularly useful because it will
grow in most areas with relatively little care. It is Long, R.W. and O. Lakela. 1976. A Flora of
used as ground cover, for hedges and screens, and Tropical Florida. Banyan Books, Miami, FL.
as upright shrubs. Because of its rapid growth, 962 p.
pruning is often necessary, which can expose
gardeners to the toxic sap that causes dermatitis Pacific Island Ecosystems at Risk. 2002.
symptoms of rash, blisters, and itch. Although Allamanda cathartica L., Apocynaceae.
incidence is much less common, plant parts are http://hear.org/pier_v3.3/alcat.htm. 2p.
also toxic if ingested. All parts contain the toxic
iridoid lactone, allamandin (Ecology and Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Evolutionary Biology Conservatory 2002). In Montiel, eds. 2001. Flora de Nicaragua.
herbal medicine, teas prepared from leaves and Monographs in Systematic Botany Vol. 85, No.
roots are used as a strong purgative that must be 1. Missouri Botanic Garden Press, St. Louis,
used with caution (Liogier 1990). MO. 943 p.
References
Tropilab Inc. 2002. Allamanda cathartica L.
http://www.tropilab.com/allamanda.thml. 2 p.
Barcellos, D.C. 2002. Plantas ornamentais tóxicas:
Allamanda cathartica. http://www.
37
Amyris elemifera L. torchwood
RUTACEAE
John K. Francis
38
in the dry forests that torchwood inhabits grow Paper ITF-12. U.S. Department of Agriculture,
slowly and may live for 50 years or more (author’s Forest Service, Institute of Tropical Forestry,
observation). A release cut or thinning should be Río Piedras, PR. 28 p.
effective if it were necessary to improve the
position of torchwood in a stand. Hammer, R.L. 2001. Attracting birds to your
garden. Fairchield Tropical Garden.
Benefits.—The wood of torchwood is used for http://www.ftg.org/horticulture/habitat
fence posts today but was formerly used for fuel, plants.html. 4 p.
furniture, and torches--hence the name torchwood.
In a test of untreated service life of posts, 20 of 20 Howard R.A. 1988. Flora of the Lesser Antilles,
posts were still sound after 13 years in the ground Leeward and Windward Islands.
(Chudnoff and Goytia 1972). If the wood was more Dicotyledoneae, part 1. Vol. 4. Arnold
abundant and in larger pieces, it would be in Arboretum, Harvard University. Jamaica Plain,
demand for lumber because it is fragrant, fine- MA. 673 p.
grained, strong, durable, and repellent to dry-wood
termites (Little and Wadsworth 1964). Torchwood Liogier H.A. 1988. Descriptive flora of Puerto
is valued as a honey plant in the Dominican Rico and adjacent islands, Spermatophyta. Vol.
Republic (Marcano Fondeur 1973). Birds eat the 2. Editorial de la Universidad de Puerto Rico,
fruits, but there usually are many other more Río Piedras, PR. 481 p.
attractive foods present (West and Arnold 1952). In
Florida, torchwood is valued as a larval food for Liogier, H.A. 1990. Plantas medicinales de Puetro
the rare and endangered Schaus’s swallowtail Rico y del Caribe. Iberoamericana de Ediciones,
butterfly, Papilio aristodemus ponceanus (Hammer Inc., San Juan, PR. 566 p.
2001). Key deer (Odocollous virginanus clavium)
will not eat the foliage (Schaus and others 2001). Little, E.L., Jr. and F.L. Wadsworth. 1964.
However, feral goats eat at least the bark, which Common trees of Puerto Rico and the Virgin
has caused torchwood to decrease in abundance in Islands. Agriculture Handbook 249. U.S.
Mona Island, Puerto Rico (Little and Wadsworth Department of Agriculture, Washington, DC.
1964). Taxaline, an oxazole extracted from 548 p.
torchwood, had the most effective antibacterial
activity of several bioactive chemicals from West Marcano Fondeur, E. de J. 1973. La Flora Apícola
Indian plants against Mycobacterium tuberculosis de la República Dominicana. http.//marcano.
and two other Mycobacterium species (Rastogi and Freeservers.com/nature/estudios/apicola/dicotsp.
others 1998). Root and resin extracts from html. 11 p.
torchwood and others of the genus are used as an
expectorant (Liogier 1990). Rastogi, N, J. Abaul, G-K. Seng, A. Devallois, E.
Philogene, P. Bourgeois, and K.S Goh. 1998.
References Antimycobacterial activity of chemically
defined natural substances from the Caribbean
4-H Youth Development. 2001. Florida forest flora of Guadeloupe. FEMS-Immunology and
trees: torchwood (Amyris elemifera). Medical Microbiology 20(4): 267-273.
http://www.sfrc.ufl.edu/4h/Torchwood/
torchwoo.htm. 2 p. Schaus, C., S. Wade, and J. Dunan. 2001. Key
deer and plants they won’t eat.
Britton, N.L. and C.F. Millspaugh. 1962. The http://monroe.ifas.ufl.edu/key_deer_plants.htm.
Bahama flora. The New York Botanical Garden, 3 p.
New York. 695 p.
West, E. and L.E. Arnold. 1952. The native trees
Chudnoff, M. and E. Goytia. 1972. Preservative of Florida. University of Florida Press,
treatments and service life of fence posts in Gainesville, FL. 212 p.
Puerto Rico (1972 progress report). Research
39
Aralia spinosa L. devil’s walking stick
ARALIACEAE
Kristina Connor
1998). The dark green, glaucous leaflets are sessile,
5.1 to 10.2 cms long and 3.0 to 4.1 cm wide,
mostly paired, ovate, serrate, with pointed apex and
rounded base (Dirr 2002, Johnson and Hoagland
1999, Krüssmann 1986). There are prickles on the
veins and lower surfaces; they appear to be
outgrowths of epidermal and parenchyma cells
(Briand and Soros 2002). The wood is lightweight,
brown but streaked with yellow (Synor and Cowen
[no date]). It may be that the pithy nature of the
trunk of Devil’s walking stick allows for rapid
growth but ultimately restricts crown architecture
and maximum size attainable (Briand and others
1999). The species is deciduous and spreads by
underground runners.
40
white to cream colored flowers are small, perfect, However, it will send up shoots from its rhizomes
numerous, and occur in large clusters (umbels) in and can be somewhat invasive.
mid to late summer, depending on geographic area.
Panicles of flowers may be 30 to 46 cm long. Each References
flower, composed of five sepals, five reflexed
petals, and five stamens, is borne on a hairy stalk. Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
The ovary is inferior, and there are usually five A Concise Dictionary of Plants Cultivated in the
styles, united at the base (Brickell and Zuk 1996, United States and Canada. McMillan Publishing
U.S. Department of Agriculture [no date]). The Co., Inc. New York. 1,290 p.
purple-black fruits, maturing in late fall, are
produced in large quantities on pink-red stems. Briand, C.H., S.M. Campion, D.A. Dzambo, and
They are juicy, ovoid drupes up to 6.4 mm long K.A. Wilson. 1999. Biochemical properties of the
with three to five seed-like stones. The species can trunk of the Devil’s walking stick (Aralia
be propagated from seed sown outdoors in fall spinosa; Araliaceae) during the crown-building
(Russell 1997) or from suckers taken in late winter phase: Implications for tree architecture.
and root cuttings taken in late fall and overwintered American Journal of Botany 86(12): 1677-1682.
upside down in sand (Scheper 2002). Stored seeds
require 3 to 5 months stratification, followed by 1 Briand, C.H., A.D. Daniel, K.A. Wilson, and H.E.
to 4 months at 20 oC for germination. Woods. 1998. Allometry of axis length, diameter,
and taper in the Devil’s walking stick (Aralia
Growth and Management.Devil’s walking stick spinosa; Araliaceae). American Journal of Botany
can be maintained by mowing, cutting, and 85(9): 1201-1206.
burning, and will commonly form dense thickets
by resprouting after such disturbances. When the Briand, C.H. and C. Soros. 2002. Prickle
overstory becomes too dense, the species will distribution in Aralia spinosa.
decline. Defoliation by gypsy moth in http://henson1.salisbury.edu/~biology/
Pennsylvania and Maryland has increased the Briand/research/research.html. 5 p.
presence of Devil’s walking stick by thinning the
overstory (Sullivan 1992). The species can be Brickell, C. and J.D. Zuk. 1996. The American
controlled by injecting stems larger than 2 inches Horticultural Society A-Z Encyclopedia of
in diameter with herbicide (Loftis 1978). Garden Plants. DK Publishing, Inc., New York.
1,092 p.
Benefits.—The flowers provide a pollen and nectar
source to honeybees and a variety of other insects. Dirr, M.A. 2002. Aralia spinosa – Devil’s-
The berries are edible to wildlife, a favorite of walkingstick or Hercules-club. Noble Plants.
cedar waxwings and other birds, as well as other Plant information webpage,
frugivores and omnivores, including the black bear. http://www.nobleplants.com/Classnotes/
Deer browsing has been reported (Krüssmann fall/fallprofiles/aralia.htm
1986, Sullivan 1992). The bark, roots, and berries
have also been used for medicinal purposes by both Felter, H.W. and J.U. Lloyd. 1898. Aralia
Native Americans and early settlers. While various spinosa.-Prickly Elder. King’s American
plant parts, extracted in alcohol or water, have been Dispensatory. 18th ed., 3rd revision, 2 vol. Ohio
used to treat boils, fever, toothache, eye problems, Valley Co., Cincinnati, OH. 2,172 p. + index.
skin conditions, and snakebite, the raw berries can
be mildly toxic to humans if ingested, contact with Frett, J. and B. MacKenzie. 1999. Aralia spinosa.
the bark or roots can cause a brief skin irritation, The University of Delaware Botanical Garden.
and ingesting the fresh bark will cause vomiting Species descriptions. http://bluehen.ags.udel.edu.
(Felter and Lloyd 1898, Grieve 1971). The species 1 p.
is cultivated as an ornamental plant, the lacy
appearance of the foliage, along with the prolific Godfrey, R.K. 1988. Trees, shrubs, and woody
flowers and fruit making it an interesting vines of northern Florida and adjacent Georgia
landscape plant (Dirr 2002, Odenwald and others and Alabama. The University of Georgia Press,
1996). It is also an excellent barrier plant (Frett Athens, GA. 734 p.
and MacKenzie 1999, Michigan State University
Extension 1999) and is good for urban use. Grieve, M. 1971. A Modern Herbal. The
medicinal. culinary, cosmetic and economic
41
properties, collection and folk-lore of herbs, Russell, A.B. 1997. Trees of the Maritime Forest.
grasses, fungi, shrubs and trees with their modern Department of Horticultural Science, North
scientific uses. Dover Publications Inc., New Carolina State University,
York. 902 p. http://www.ces.ncsu.edu/depts/hort/
consumer/factsheets/maritime. 2 p.
Johnson, F.L. and B.W. Hoagland. 1999. Aralia
spinosa. Catalog of the Woody Plants of Scheper, J. 2002. Aralia spinosa. Floridata. http://
Oklahoma, Oklahoma Biological Survey www.floridata.com/ref/a/aral_spi.cfm. 4 p.
http://www. biosurvey.ou.edu. 1 p.
Sullivan, J. 1992. Aralia spinosa. In: U.S.
Krüssmann, G. 1986. Manual of Cultivated Broad- Department of Agriculture, Forest Service, Rocky
Leaved Trees and Shrubs. Volume I, A-D. Mountain Research Station Fire Sciences
Timber Press, Beaverton, OR. 448 p. Laboratory (2002 February). Fire Effects
Information System
Loftis, D.L. 1978. Preharvest herbicide control of http://www.fs.fed.us/database/feis. 10 p.
undesirable vegetation in southern Appalachian
hardwoods. Southern Journal of Applied Forestry Synor, T.D. and W.F. Cowen. [No date]. Ohio
2(2): 51-54. Trees. Aralia. Department of Natural Resources,
Ohio State University Extension Bulletin 700-00.
Michigan State University Extension. 1999. Aralia http://www.ag.ohio-
spinosa – Devil’s Walkingstick. Ornamental Plants state.edu/~ohioline/b700/index. html. 2 p.
plus Version 3.0.
http://www.msue.msu.edu/msue/imp/ modzz/ U.S. Department of Agriculture. [No date].
00001941.html. 1 p. Southern wetland flora: Field office guide to plant
species. USDA Soil Conservation Service, South
Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope. National Technical Center, Fort Worth, Texas.
1996. Plants for American Landscapes. Louisiana Jamestown, ND: Northern Prairie Wildlife
State University Press, Baton Rouge. 266 p. Research Center Home Page,
http://www.npwrc.usgs.gov/resource/1999
/soutflor/soutflor.htm. [not paged].
42
Ardisia elliptica Thunb. shoebutton ardisia
MYRSINACEAE
John K. Francis
43
Birds are the principal seed dispersers, although 772 p.
raccoons and opossums in Florida also eat the
fruits and disperse seeds (Miami-Dade County Dominguez, J., A. Scott, T. Scott, G. Valdes, C.
2002). Glenn, and C. Moore. 2002. Herbivore damage
on the invasive exotic Ardisia elliptica and the
Growth and Management.—Although native A. escallonioides in Southeastern Florida.
shoebutton ardisia may grow as much as 1 m the NSF Research in Ecology: Invasive Species.
first year as a sprout, seedlings and established http://fig.cox.miami.edu/ys99/gladres.html. 3 p.
shrubs grow slowly, usually 0.25 to 0.5 m/year in
height. No figures are known for longevity, but 10 Francis, J.K. 2000. Estimating biomass and carbon
to 25 years or more are likely. It is often desirable content of saplings in Puerto Rican secondary
to control shoebutton ardisia in natural areas. Small forests. Caribbean Journal of Science 36(3-4):
numbers of seedlings and shrubs can be eliminated 346-350.
by pulling and grubbing. A broadcast spray of
glyphosate is effective in killing dense stands of Howard, R. A. 1989. Flora of the Lesser Antilles,
seedlings but will also kill native plants. Larger Leeward and Windward Islands. Vol. 6. Arnold
plants can be killed by applying a basal spray of Arboretum, Harvard University, Jamaica Plain,
triclopyhr mixed with an oil diluent (Pacific Island MA. 658 p.
Ecosystems at Risk 2002).
Liogier, H.A. 1995. Descriptive flora of Puerto
Benefits and Detriments.—Shoebutton ardisia Rico and adjacent islands. Vol. 4. Editorial de la
was once used heavily as an ornamental for accent Universidad de Puerto Rico, San Juan, PR.
and background plantings. Many plantings still 617 p.
remain, but the species is no longer cultivated
commercially (Miami-Dade County 2002). The Miami-Dade County. 2002. Shoebutton ardisia-
fruits are edible and taste slightly sour with a hint Ardisia elliptica, Ardisia humilis.
of starch but otherwise lack flavor. The species is http://www.co.miami-dade.fl.us/derm/
useful for fuel and vegetable stakes. The air-dry environment/badplants/plant%20…/shoebutton_
specific gravity of stem wood in Puerto Rico was ardisia.ht. 1 p.
measured at 0.48 + 0.01 g/cm3. Biomass (dry
weight) of plants from 137 cm in height to plants Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
with a 5-cm D.B.H. may be calculated by the 50. Bernice P. Bishop Museum Press, Honolulu,
model: Total Dry Weight in grams = 22.020(stem HI. 924 p.
length in meters) (diameter at 30 cm)2 (Francis
2000). Shoebutton ardisia is extremely invasive. It Pacific Island Ecosystems at Risk. 2002. Invasive
forms dense stands under trees in wet sites in plant species: Ardisia elliptica Thunberg,
Florida, Hawaii, and Puerto Rico that suppress Myrsinaceae. http://www.hear.org/pier/arell.
native understory plants (Center for Aquatic and htm. 2 p.
Invasive Plants 2001).
Pascarella, J.B. 1997. Breeding systems of Ardisia
References Sw. (Myrsinaceae). Brittonia 49(1): 45-53.
Center for Aquatic and Invasive Plants. 2002. Tomlinson, P.B. 1986. The botany of mangroves.
Ardisia elliptica Thumb., Myrsinaceae/Myrsine Cambridge University Press, London. 413 p.
family. University of Florida, Gainesville, FL.
www.aquat1.itas.ufl.edu. 2 p. Yuen-Po, Y. 1999. An enumeration of
Myrsinaceae of Taiwan. Botanical Bulletin of
Corner, E.J.H. 1952. Wayside trees of Malaya. the Academy Sinensis 40: 39-47.
Government Printing Office, Singapore, Malaya.
44
Argythamnia candicans Sw. argythamnia
EUPHORBIACEAE
John K. Francis
45
near seed-bearing plants. No planting experience Howard, R.A. 1989. Flora of the Lesser Antilles,
has been reported. Leeward and Windward Islands. Vol. 5. Arnold
Arboretum, Harvard University, Jamaica Plain,
Benefits.—Argythamnia is eaten by livestock and MA. 604 p.
contributes to ground cover and environmental
diversity. Liogier H.A. 1988. Descriptive flora of Puerto
Rico and adjacent islands, Spermatophyta. Vol.
References 2. Editorial de la Universidad de Puerto Rico,
Río Piedras, PR. 481 p.
Britton, N.L., and C.F. Millspaugh. 1962. The
Bahama Flora. The New York Botanical
Garden. New York. 695 p.
46
Artemisia arbuscula Nutt. low sagebrush
ASTERACEAE
47
the same chromosome number (McArthur and References
Sanderson 1999). In areas where the distributions
of these two overlap, low sagebrush usually Beetle, A.A. 1960. A study of sagebrush, the
inhabits sites that have more moisture or are at section Tridentatae of Artemisia. Bulletin 368,
higher elevations than black sagebrush (A. nova University of Wyoming Agriculture Experiment
Nelson) (Tisdale and Hironaka 1981, Ward 1953, Station, Laramie, WY. 83 p.
Zamora and Tueller 1973). Low sagebrush is
usually associated with pinyon-juniper, mountain Brunner, J.R. 1972. Observations on Artemisia in
brush, and mountain big sagebrush communities. Nevada. Journal of Range Management 25: 205-
208.
Reproduction.—Flowering occurs from August to
September, depending upon strain and elevation. Dealy, J.E., DA. Leckenby, D.M. Concannon.
Seeds ripen in October and November (McArthur 1981. Wildlife habitats in managed
and others 1979). Seeds are usually wind dispersed rangelands—the Great Basin of southeastern
during late fall or winter, and seedlings emerge Oregon. General Technical Report PNW-120,
during the following spring. Seedlings emerge USDA Forest Service, Pacific Northwest Forest
rapidly and grow quickly, although root growth and Range Experiment Station, Portland, OR,
generally outpaces stem growth during early 66 p.
development. Plants can be established on adapted
sites by direct seeding, broadcasting, and drilling. International Plant Names Index. 2003.
Seeding should be conducted in the fall on a firm International plant names index.
seedbed and seeds should remain within 2 mm of http://www.ipni.org. [not paged].
the soil surface. There are approximately 1,800
cleaned seeds per gram (Meyer 2003) Kufeld, R.C., O.C. Wallmo, and C. Feddema.
1973. Foods of the Rocky Mountain mule deer.
Management.—Low sagebrush is a source of Research Paper RM-111, USDA Forest Service,
browse for livestock and big game during winter Rocky Mountain Forest and Range Experiment
months (Kufeld and others 1973, McArthur and Station, Fort Collins, CO. 31 p.
others 1979). Some forms are browsed more
heavily than others. A gray-green form in Nevada, McArthur, E.D., A.C. Blauer, A.P. Plummer, and
for example, may be heavily browsed, while the R. Stevens. 1979. Characteristics and
green form is only lightly browsed (Brunner hybridization of important Intermountain shrubs.
1972). Stand maintenance is often crucial because III. Sunflower family. Research Paper INT-220,
shrubs provide forage and cover under conditions USDA Forest Service, Intermountain Forest and
that are intolerable for most shrubs. Range Experiment Station, Ogden, UT. 82 p.
Attempts to seed low sagebrush out of its
ecological range have been unsuccessful. McArthur, E.D. and S.C. Sanderson. 1999.
Seedlings planted “offsite” usually establish but Cytogeography and chromosome evolution in
fail to reproduce. Stands recover quickly after subgenus Tridentatae of Artemisia (Asteraceae).
burning or other disturbance. Low sagebrush American Journal of Botany 86: 1,754-1,775.
spreads vigorously and can invade perennial
seeded grass stands. Perennial herb and grass Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and
understory production usually decreases as stands R.G. Nisley, eds. Woody plant seed manual.
mature. Low sagebrush is not a good candidate for http://www.wpsm.net. 12 p.
mechanical control because habitat terrain is often
extreme. Tisdale, E.W. and M. Hironaka. 1981. The
sagebrush—grass region: a review of the
Benefits.—Low sagebrush is a native shrub that ecological literature. Bulletin 33. Forest,
adds structural and biological richness to arid Wildlife, and Range Experiment Station,
communities throughout the Intermountain West. University of Idaho, Moscow, ID. 31 p.
Low sagebrush communities provide important
habitat for a variety of domestic and wild animals Ward, G.H. 1953. Artemisia, section Seriphidium,
(Dealy and others 1981). Its ability to grow on in North America, a cytotaxonomic study.
exposed sites is important for wintering animals Contributions from the Dudley Herbarium 4:
and shrub cover helps reduce on-site soil erosion. 155-205.
48
Winward, A.H. and E.D. McArthur. 1995. Zamora, B. and P.T. Tueller. 1973. Artemisia
Lahontan sagebrush (Artemisia arbuscula ssp. arbuscula, A. longiloba, and A. nova habitat
longicaulis): a new taxon. Great Basin types in northern Nevada. Great Basin Naturalist
Naturalist 55(2): 151-157. 33: 225-242.
49
Artemisa bigelovii Gray Bigelow sagebrush
ASTERACEAE
50
and especially broom snakeweed [Gutierrezia Institution of Washington, Washington, DC.
sarothrae (Pursh) Britt. & Rusby]. Bigelow 355 p.
sagebrush is usually resistant to the common rust
diseases and insect galls common to other Kearney, T.H. and R.H. Peebles. 1960. Arizona
Tridentatae taxa (Beetle 1960). It is, however, flora, 2nd edition. University of California Press,
susceptible to wilt disease (Nelson and Krebill Berkeley, CA. 1,065 p.
1981). It grows in areas that usually receive 250 to
400 mm of annual precipitation. Kornkven, A.B., L.E. Watson, and J.R. Estes.
1998. Phylogenetic analysis of Artemisia section
Reproduction.—Flowering occurs from August to Tridentatae (Asteraceae) based on sequences
October and seeds mature during the following from the internal transcribed spacers (ITS) of
months. Seeds are wind dispersed in the fall and nuclear ribosomal DNA. American Journal of
winter, and seedlings emerge in spring. Seeds are Botany 85: 1787-1795.
among the smallest of sagebrush seeds, about 5.5
million cleaned seeds per kg (Meyer 2003). McArthur, E.D., A.C. Blauer, A.P. Plummer, and
Seedlings grow rapidly, and young shrubs are R. Stevens 1979. Characteristics and
robust and vigorous. Plants appear to have good hybridization of important intermountain shrubs.
drought tolerance, even as seedlings. III. Sunflower family. Research Paper INT-220.
Direct seeding has been limited to U.S. Department of Agriculture, Forest Service
experimental plantings. Seeded shrubs grow well Intermountain Forest and Range Experiment
and natural spread normally occurs around the Station, Ogden, UT. 82 p.
parental plant, suggesting that Bigelow sagebrush
can be successfully seeded. McArthur, E.D., C.L. Pope, and D.C. Freeman.
1981. Chromosomal studies of subgenus
Management.—Bigelow sagebrush is palatable to Tridentatae of Artemisia: evidence for
livestock and game throughout its distribution. Its autopolyploidy. American Journal of Botany 68:
twigs are less woody, aromatic, and bitter than 589-605.
most species within the big sagebrush complex
(Hall and Clements 1923). In addition, Bigelow McArthur, E.D. and S.C. Sanderson. 1999.
sagebrush is more grazing tolerant than most other Cytogeography and chromosome evolution of
sagebrushes. These shrubs, however, are generally subgenus Tridentatae of Artemisia (Asteraceae).
scattered, and extensive stands are found only near American Journal of Botany 86: 1754-1775.
the Four Corners area of Utah, Arizona, New
Mexico, and Colorado. McArthur, E.D., R. Van Buren, S.C. Sanderson,
and K.T. Harper. 1998. Taxonomy of
Benefits.—Bigelow sagebrush adds biological Sphaeromeria, Artemisia, and Tanacetum
diversity to shrub communities throughout the (Compositatae, Anthemideae) based on
Southwestern United States. This species improves randomly amplified polymorphic DNA (RAPD).
soil stability and reduces erosion. It also provides Great Basin Naturalist 58: 1-11.
palatable forage for livestock and wildlife and adds
structural diversity to the landscape. Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and
References R.G. Nisley, eds. Woody plant seed manual.
http://www.wpsm.net 12 p.
Beetle, A.A. 1960. A study of sagebrush: the
section Tridentatae of Artemisia. Bulletin 368. Nelson, D.L. and R.G. Krebill. 1981. A sagebrush
Agricultural Experiment Station, University of wilt disease of unknown origin. Great Basin
Wyoming, Laramie, WY. 83 p. Naturalist 41: 184-191.
Hall, H.M. and F.E. Clements. 1923. The Ward, G.H. 1953. Artemisia, section Seriphidium,
phylogenetic method in taxonomy, the North in North America. A cytotaxonomic study.
American species of Artemisia, Chrysothamnus, Contributions from the Dudley Herbarium 4:
and Atriplex. Publication 326. Carnegie 155-205.
51
Artemisia californica Less. California sagebrush
ASTERACEAE
Synonyms: None
52
(achenes) ripen in December and January, with Fire Effects.—California sagebrush resprouts
some variation among years and habitats, and are after fire about 25 percent of the time (Keeley
primarily wind-dispersed (Eliason and Allen 1997, 1998). Resprouting appears to be lower in burns
DeSimone and Zedler 1999). Achenes are tiny through dense vegetation, where plants are older,
(about 60 micrograms) and in bulk amount to or if fire intensity is high (Malanson and O'Leary
about 14,300,000 seeds/kg. (personal 1982, Keeley 1998, Minnich and Dezzani 1998).
communication with S&S Seeds, Carpenteria, Seedling emergence after fire is variable and low
California). Seeds germinate in canopy openings (Zedler and others 1983, Keeley 1998). Seedlings
or in small grassland clearings generated by tend to appear the second year from seeds of
gophers (Thomomys spp.), but seedlings suffer resprouts or seeds blown in from adjacent areas.
high mortality from gopher activity (Eliason and Under high fire intensities or frequency, California
Allen 1997, DeSimone and Zedler 1999, 2001). sagebrush will likely be extirpated because of its
Seedlings emerge in the rainy season, and most poor resprouting ability and poor competitive
growth occurs by May. ability of seedlings (Malanson and O'Leary 1982,
Malanson and Westman 1991). Development,
Seed Germination.—Seeds will germinate when increased fire frequency, competition with exotic
fresh, but stored seeds may need cold stratification grasses, poor growth in grasslands, and air
to enhance germination (De Hart 1994). While pollution are each detrimental to this shrub’s
testing the combined effects of light and fire survival (Eliason and Allen 1997, Allen and others
components on germination, 73 percent of seeds 1998, Minnich and Dezzani 1998, Keeley 1998).
exposed to light germinated on soil, but seeds in
the dark did not germinate (Keeley 1987). These Horticulture.—California sagebrush can be
results are consistent with field data that show compact if kept pruned and subjected to occasional
germination may be limited by reduced light pinching. These aromatic plants remain attractive
(Elliason and Allen 1997, Montalvo and others in the dry season with occasional water (Schmidt
2002). Certain components of fire also influence 1980, Keator 1994). It prefers well-drained soils
germination. In light, treatment with leachate from and a low organic matter content (Wasowski and
charred wood (charrate) alone or with heat Wasowski 1995). Naturally occurring prostrate or
increases germination by 5 to 14 percent, but low-mounding varieties are in cultivation (Browse
germination is about a third lower when seeds are 1987, Perry 1992, Wasowski and Wasowski 1995)
subjected to high temperatures (70 to 120 oC). In and look good in dry borders or as a foreground to
contrast, in darkness both heat and charrate alone contrasting taller shrubs (Keator 1994).
stimulate germination, although heat and charrate
together generally decrease germination. Thus Benefits.—California sagebrush provides habitat
buried seeds may require some exposure to fire in for many plant and animal species and is an
order to germinate, but such exposure yields important component of critical habitat for the rare
inferior germination compared to light alone California gnatcatcher, Polioptila californica
(Keeley 1987). (Weaver 1998). The bird forages preferentially in
California sagebrush and the often co-dominant
Genetics and Geographic Variation.—Near the shrub Eriogonum fasciculatum Benth. (Beyers and
coast, plants vary in pubescence, color, chemistry, Wirtz 1997), in part because they house a high
and physiology (O'Brien 1980). Shrubs vary number and diversity of arthropods. Osborne
between a green form with sparse hairs on the (1998) found high diversity and abundance of
leaves to a form with dense hairs that give the arthropods in both species and concluded that A.
plants a gray hue. Populations differ in the californica, together with other sage scrub shrub
distinctness of forms and in their relative species, is important to the maintenance of high
frequency. Seedlings raised together generally arthropod species richness and abundance in
retain the grayness of their parents but can become coastal sage scrub. California sagebrush is also
grayer with age (O'Brien 1980). Differences in involved in species-specific interactions. For
pubescence also correspond to differences in water example, it is the host for Trirhabda
content, rate and timing of shoot elongation, leaf sericotrachyla, an herbivorous beetle whose
retention, and chemistry. The lower leaf-water developmental cycle is closely associated with the
content of gray plants negatively impacts larval seasonal growth of its only known host plant
growth and fecundity of the beetle, Trirhabda (O'Brien 1980).
sericotrachyla (O'Brien 1980).
53
Historical Uses.—Leaves of California sagebrush Endangered Species and Habitats. International
have been used by Native Americans for smoking, Assoc. of Wildland Fire, Fairfield, WA. p. 81-89.
in sweat-houses, and various other purposes (Bean
Bocek, B. 1984. Ethnobotany of Costanoan Indians,
and Saubel 1972). The Cahuilla used the plant to
California, based on collections by John P.
ensure proper maturation of girls into women. It is
Harrington. Economic Botany 38: 240-255.
said to stimulate the uterine mucosa, ensuring
rapid childbirth and, if regularly consumed as a Browse, P.M. 1987. Artemisia californica 'canyon
decoction prior to the onset of each menstruation, gray'. Pacific Horticulture 48: 56.
to prevent menstrual cramps and alleviate
menopausal trauma. Fresh or dried leaves were De Hart, J. 1994. Propagation Secrets for California
Native Plants. Jeanine De Hart, Encinitas, CA.
chewed to alleviate colds (Bean and Saubel 1972).
Costanoan Indians applied leaves to wounds or 28p.
aching teeth for pain reduction, and as a decoction DeSimone, S.A. and P.H. Zedler. 1999. Shrub
to bathe patients with colds, coughs, rheumatism, seedling recruitment in unburned Californian
or to be consumed and used as a poultice for coastal sage scrub and adjacent grassland.
treatment of asthma (Bocek 1984). Ecology 80: 2018-2032.
Growth and Management.—California DeSimone, S.A. and P.H. Zedler. 2001. Do shrub
sagebrush should be planted using shallow seeding colonizers of southern Californian grassland fit
methods such as hydroseeding or dry broadcasting generalities for other woody colonizers?
followed by seed imprinting. Shallow methods are Ecological Applications 11: 1,101-1,111.
superior to planting with a range drill because the Eliason, S.A. and E.B. Allen. 1997. Exotic grass
seeds need light for germination (Montalvo and competition in suppressing native shrubland re-
others 2002). Attempts to use annual legumes as establishment. Restoration Ecology 5: 245-255.
nurse plants can result in reduced seedling growth
and survival (Marquez and Allen 1996), but Gray, J.T. 1982. Community structure and
seedling survival may be unaffected when sown productivity in Ceanothus chaparral and coastal
with other native species at low density (Montalvo sage scrub of southern California. Ecological
and others 2002). Clearing weeds before planting Monographs 52: 415-435.
is critical. California sagebrush can return Gray, J.T. 1983. Nutrient use by evergreen and
vigorously to areas where all vegetation has first deciduous shrubs in southern California. Journal
been removed, even when sown with a variety of of Ecology 71: 21-41.
other species. Spring rains or artificial late fall and
spring irrigation may enhance survival of seedlings Halligan, J.P. 1973. Bare areas associated with
(Williams and Hobbs 1989, Elliason and Allen shrub stands in grassland: the case of Artemisia
1997, Padgett and others 2000). However, survival californica. BioScience 23: 429-432.
is not enhanced by late-season watering in wet Halligan, J.P. 1974. Relationship between animal
years (DeSimone and Zedler 2001). activity and bare areas associated with California
sagebrush in annual grassland. Journal of Range
References Management 27: 358-362.
Allen, E.B., P.E. Padgett, A. Bytnerowicz, and R. Halligan, J.P. 1975. Toxic terpenes from Artemisia
Minnich. 1998. Nitrogen deposition effects on californica. Ecology 56: 999-1003.
coastal sage vegetation of southern California. Halligan, J.P. 1976. Toxicity of Artemisia
General Technical Report 166, U.S. Department californica to four associated herb species. The
of Agriculture, Forest Service. p. 131-139. American Midland Naturalist 95: 406-421.
Bean, J.L. and K.S. Saubel. 1972. Temalpakh: Harrison, A.T., E. Small, and H.A. Mooney. 1971.
Cahuilla Indian knowledge and usage of plants. Drought relationships and distribution of two
Malki Museum Press, Morongo Indian Mediterranean-climate California plant
Reservation, CA. 225 p. communities. Ecology 52: 869-875.
Beyers, J.L. and W.O. Wirtz II. 1997. Vegetative Hickman, J.C, ed. 1993. The Jepson Manual:
characteristics of coastal sage scrub sites used by Higher Plants of California. University of
California gnatcatcher: implications for California Press, Ltd., Los Angeles, CA. 1,400 p.
management in a fire-prone ecosystem. In: J.
Greenlee, ed. Fire Effects on Rare and Keator, G. 1994. Complete Garden Guide to the
54
Native Shrubs of California. Chronicle Books, Dissertation. University of California, Irvine. 125
San Francisco, CA. 314 p. p.
Keeley, J.E. 1987. Role of fire in seed germination Osborne, K.H. 1998. A description of arthropod
of woody taxa in California chaparral. Ecology community structure in southern California
68: 434-443. coastal sage scrub. M.S. Thesis. University of
California, Riverside. 133 p.
Keeley, J.E. 1998. Postfire ecosystem recovery and
management: The October 1993 large fire episode Padgett, P.E., S.N. Kee, and E.B. Allen. 2000. The
in California. In: J. M. Moreno, ed. Large Forest Effects of irrigation of revegetation of semi-arid
Fires. Backbuys Publishers, Leiden, Netherlands. costal sage scrub in Southern California.
p. 69-90. Environmental Management 26: 427-435.
Kirkpatrick, J.B. and C.F. Hutchinson. 1977. The Perry, B. 1992. Landscape plants for western
community composition of California coastal sage regions: an illustrated guide to plants for water
scrub. Vegetatio 35: 21-33. conservation. Land Design Publishing, Claremont,
CA. 318 p.
Malanson, G.P. and J.F. O'Leary. 1982. Post-fire
regeneration strategies of Californian coastal sage Poole, D.K. and P.C. Miller. 1975. Water relations
shrubs. Oecologia 53: 355-358. of selected species of chaparral and coastal sage
communities. Ecology 56: 1,118-1,128.
Malanson, G.P. and J.F. O'Leary. 1995. The coastal
sage scrub-chaparral boundary and response to Schmidt, M.G. 1980. Growing California Native
global climatic change. In: J. M. Moreno and W. Plants. University of California Press, Los
C. Oechel, eds. Global change and Mediterranean- Angeles, CA. 366 p.
type ecosystems. Volume 117. Springer-Verlag
Sigüenza, C. 2000. Nitrogen deposition and soil
New York, Inc., NY. p 203-224.
microorganisms of Artemisia californica and
Malanson, G.P. and W.E. Westman. 1991. exotic grasses in southern California. Ph.D.
Modeling interactive effects of climate change, air Dissertation. University of California, Riverside.
pollution, and fire on California shrubland. 153 p.
Climate Change 18: 363-376.
Wasowski, S. and A. Wasowski. 1995. Native
Marquez, V.J. and E.B. Allen. 1996. Ineffectiveness Gardens for Dry Climates, 1st Edition. Clarkson
of two annual legumes as nurse plants for Potter/Publishers, New York. 176 p.
establishment of Artemisia californica in coastal
Weaver, K.L. 1998. Coastal sage scrub variations of
sage scrub. Restoration Ecology 4: 42-50.
San Diego county and their influence on the
Minnich, R.A. 1985. Evolutionary convergence or distribution of the California gnatcatcher. Western
phenotypic plasticity? Responses to summer rain Birds 29: 392-405.
by California chaparral. Physical Geography 6:
Westman, W.E. 1981a. Factors influencing the
272-287.
distribution of species of California coastal sage
Minnich, R.A. and R.J. Dezzani. 1998. Historical scrub. Ecology 62: 439-455.
decline of coastal sage scrub in the Riverside-
Westman, W.E. 1981b. Seasonal dimorphism of
Perris plane, California. Western Birds 29: 366-
foliage in Californian coastal sage scrub.
391.
Oecologia 51: 385-388.
Montalvo, A.M., P.A. McMillan, and E.B. Allen.
Williams, K. and R.J. Hobbs. 1989. Control of
2002. The relative importance of seeding method,
shrub establishment by springtime soil water
soil ripping, and soil variables on seeding success.
availability in an annual grassland. Oecologia 81:
Restoration Ecology 10: 52-67.
62-66.
Munz, P.A. and D.D. Keck. 1968. A California
Yoshida, L.C. and E.B. Allen. 2001. Response to
Flora with Supplement. University of California
ammonium and nitrate by a mycorrhizal annual
Press, Berkeley, CA. 1,681 + 224 p.
invasive grass and native shrub in southern
O'Brien, P.Y. 1980. Adaptive relations between a California. American Journal of Botany 88: 1,430-
stenophagous herbivore, Trirhabda sericotrachyla 1,436.
(Coleoptera: Chrysomelidae), and its host plant,
Zedler, P.H., C.R. Gautier, and G.S. McMaster.
Artemisia californica (Compositae). Ph.D.
1983. Vegetation change in response to extreme
55
events: the effect of a short interval between fires
in California chaparral and coastal scrub. Ecology
64: 809-818.
56
Artemisia cana Pursh silver sagebrush
ASTERACEAE
57
1960). Ranges for each subspecies, however, are spread by rhizomes. There are approximately
smaller. Plains silver sagebrush is found mostly 2,900 cleaned seeds per gram (Meyer 2003).
east of the Continental Divide, through Montana,
the Dakotas, Wyoming, western Nebraska, and Management.—Silver sagebrush can provide an
northern Colorado. Mountain silver sagebrush important source of browse and is used quite
inhabits mountainous regions (around 1,800 m or extensively by livestock and big game, especially
higher) of Montana, south along the Continental when other food sources are scarce (Kufeld and
Divide, and west to Arizona, Nevada, and Oregon. others 1973, Wasser 1982). In the western Great
The range of Bolander silver sagebrush is Plains area, silver sagebrush is an important
restricted, generally to enclosed basins, in central survival food for antelope. Domestic sheep and
Oregon and eastern California. mule deer found Bolander silver sagebrush among
the most preferable sagebrushes during winter and
Ecology.—Silver sagebrush grows on soils that fall feeding trials (Sheehy and Winward 1981).
are less mature, with less phosphorous, potassium, Sheep often browse silver sagebrush in the fall
nitrogen, organic matter, and lower cation after forbs and grasses are dry.
exchange capacity than soils supporting big Silver sagebrush can occupy areas at high
sagebrush (Hazlett and Hoffman 1975). It inhabits densities, and therefore is a candidate for plant
various environments, each subspecies adapting to control. It can be difficult to control because of its
different soil conditions. Plains silver sagebrush tendency to resprout from the crown and to spread
grows particularly well on well-watered, deep soils by rhizomes. It is not as susceptible to fire as other
of the northern Great Plains, especially along species of sagebrush (Wright and others 1979,
stream bottoms and drainageways (McArthur and White and Currie 1983). White and Currie (1983)
others 1979, Walton and others 1986). Mountain reported that silver sagebrush mortality after
silver sagebrush, on the other hand, grows in areas prescribed fall and spring burns was directly
with a heavy, lingering snow pack (Beetle 1960, related to fire intensity. In general, fall burns are
McArthur and others 1979, Tisdale and Hironaka more effective at reducing silver sagebrush than
1981, Winward 1980). Bolander silver sagebrush spring burns.
grows on poorly drained, alkaline soils (Beetle
1960, McArthur and others 1979, Tisdale and Benefits.—Silver sagebrush is an abundant native
Hironaka 1981, Winward 1980). Silver sagebrush shrub that provides additional diversity to
usually grows in areas with precipitation between shrubland ecosystems. It is an important winter
350 and 600 mm but it occurs in areas of water forage source for domestic sheep and game
accumulation, such as near streams or in enclosed animals and adds structural variety to the
basins. landscape. Native Americans traditionally used a
decoction from silver sagebrush to stop coughing
Reproduction.—Flowers bloom between August (Stubbendieck and others 1997). In addition, Euro-
and September and seed ripens in October and American settlers and Native Americans used it
November. Seeds are wind dispersed during the for fuel (McArthur and others 1979).
winter months. Establishment by direct seeding
has been successful (Kelsey 1986), especially References
when sown in the fall on the soil surface or at
shallow depths (Wasser 1982). Seedlings emerge Beetle, A.A. 1960. A study of sagebrush: the
in spring and grow rapidly. Seeding silver section Tridentatae of Artemisia. Bulletin 368,
sagebrush is best done in the late autumn or early Agricultural Experiment Station, University of
spring to maintain high viability and high Wyoming, Laramie, WY. 83 p.
germination of seeds (Hou and Romo 1998, Romo
and Grilz 2002, Romo and Young 2002). Harvey, S.J. 1981. Life history and reproductive
Seedlings are best established in lightly tilled soils strategies in Artemisia. M.S. thesis, Montana
so that stands of existing perennial grasses are State University, Bozeman, MT. 132 p.
maintained (Romo and Grilz 2002).
Plants respond dramatically to moist Hazlett, D.L. and G.R. Hoffman. 1975. Plant
conditions and may reach 35 to 50 cm in 1 year. species distributional patterns in Artemisia
Silver sagebrush can also spread asexually. tridentata—and Artemisia cana—dominatied
Harvey (1981) successfully propagated plants vegetation in western North Dakota. Botanical
through hardwood cuttings, and plants can also Gazette 136: 72-77.
58
Hou, J. and J.T. Romo. 1998. Cold-hardiness of Sheehy, D.P. and A.H. Winward. 1981. Relative
silver sagebrush seedlings. Journal of range palatability of seven Artemisia taxa to mule deer
Management 51: 704-708. and sheep. Journal of Range Management 34:
397-399.
Kelsey, R.G. 1986. Emergence, seedling growth,
and crude terpenoid concentration in a sagebrush Stubbendieck, J, Hatch, S.L., Butterfield, C.H.
garden. In: McArthur, E.D. and B.L Welch, 1997. North American Range Plants. University
comps. Proceedings—symposium on the biology of Nebraska Press, Lincoln, Nebraska. 229 p.
of Artemisia and Chrysothamnus; 1984 July 9-
13, Provo, UT. General Technical Report INT- Tisdale, E.W. and M. Hironaka. 1981. The
200. U.S. Department of Agriculture, Forest sagebrush—grass region: a review of the
Service, Intermountain Research Station, Ogden, ecological literature. Bulletin 33, Forest,
UT. p. 358-365. Wildlife, and Range Experiment Station,
University of Idaho, Moscow, ID. 31 p.
Kufeld, R.C., O.C. Wallmo, and C. Feddema.
1973. Foods of the Rocky Mountain mule deer. Walton, T.P., R.S. White, and C.L. Wambolt.
Research Paper RM-111. U.S. Department of 1986. Artemisia rproductive strategies: a review
Agriculture, Forest Service, Rocky Mountain with emphasis on silver sagebrush. In: E.D.
Forest and Range Experiment Station, Fort McArthur and B.L Welch, comps.
Collins, CO. 31 p. Proceedings—symposium on the biology of
Artemisia and Chrysothamnus; 1984 July 9-13,
McArthur, E.D., A.C. Blauer, A.P. Plummer, and Provo, UT. General Technical Report INT-200.
R. Stevens 1979. Characteristics and U.S. Department of Agriculture, Forest Service,
hybridization of important intermountain shrubs. Intermountain Research Station, Ogden, UT: 67-
III. Sunflower family. Research Paper INT-220. 74.
U.S. Department of Agriculture, Forest Service
USDA Forest Service, Intermountain Forest and Ward, G.H. 1953. Artemisia, section Seriphidium,
Range Experiment Station, Ogden, UT. 82 p. in North America. A cytotaxonomic study.
Contributions from the Dudly Herbarium 4: 155-
McArthur, E.D., C.L. Pope, and D.C. Freeman. 205.
1981. Chromosomal studies of subgenus
Tridentatae of Artemisia: evidence for Wasser, C.H. 1982. Ecology and culture of
autopolyploidy. American Journal of Botany 68: selected species useful in revegetating disturbed
589-605. lands in the west. FSW/OBS-82/56. U.S.
Department of the Interior Fish and Wildlife
McArthur, E.D. and S.C. Sanderson. 1999. Service. Washington, DC. 347 p.
Cytogeography and chromosome evolution of
subgenus Tridentatae of Artemisia (Asteraceae). Winward, A.H. 1980. Taxonomy and ecology of
American Journal of Botany 86: 1,754-1,775. sagebrush in Oregon. Station Bulletin 642.
Argicultural Experiment Station, Oregon State
Meyer, S.E. 2003. Artemisia. In: F.T. Bonner, and University, Corvallis, OR. 15 p.
R.G. Nisley, eds. Woody plant seed manual.
http://www.wpsm.net 12 p. White, R.S. and P.O. Currie. 1983. The effects of
prescribed burning on silver sagebrush. Journal
Romo, J.T. and R.W. Grilz. 2002. Establishment of Range Management 36: 611-613.
of silver sagebrush in the northern mixed prairie.
Journal of Range Management 55: 217-221. Wright, H.A., L.F. Neuenschwander, and C.M
Britton. 1979. The role and use of fire in
Romo, J. T. and J.A. Young. 2002. Temperature sagebrush-grass and pinyon-juniper plant
profiles and the effects of field environments on communites, a state-of-the-art review. General
germination of silver sagebrush. Native Plants Technical Report INT-158. U.S. Department of
Journal 3: 5-13. Agriculture, Forest Service, Intermountain Forest
and Range Experiment Station, Ogden, UT. 48 p.
59
Artemisia filifolia Torr. sandsage
ASTERACEAE
60
grows in areas with an annual precipitation range Berg, W.A. 1994. Sand-sagebrush-mixed prairie,
of 220 to 600 mm. SRM 722. In: Shiflet, T. N., ed. Rangeland cover
types of the United States. Society for Range
Reproduction.—Blooming occurs during August Management, Denver, CO. p. 99.
and September, and seed ripens between October
and December (McArthur and others 1979). Seeds Hall, H.M. and F.E. Clements. 1923. The
are smaller than those of most sagebrushes, but phylogenetic method in taxonomy, the North
seedlings grow well once established. There are American species of Artemisia, Chrysothamnus,
approximately 3.2 million cleaned seeds per kg and Atriplex. Publication 326. Carnegie
(Meyer 2003). Seedlings emerge in the spring and Institution of Washington, Washington, DC.
grow and mature quickly. Survival is reduced, 355 p.
however, if it is planted in habitats that extend
beyond its natural range (Nelson and Krebill 1981). Kelsey, R.G. and F. Shafizadeh. 1979.
Sandsage has been seeded alone in most plantings Sesquiterpene lactones and systematics of the
but can be seeded with herbs adapted to arid sandy genus Artemisia. Phytochemistry 18: 1,591-
soils. It is easy to grow as bareroot stock and 1,611.
survives well when field planted. Wildings can be
transplanted in the spring or fall, and plants can be Korkven, A.B., L.E. Watson, and J.R. Estes. 1998.
easily reared from seed. Phylogenetic analysis of Artemisia section
Tridentatae (Asteraceae) based on sequences
Management.—The value of sandsage as a source from the internal transcribed spacers (ITS) of
of browse varies with community type. It is seldom nuclear ribosomal DNA. American Journal of
eaten in grasslands where other forage is adequate Botany 85: 1,787-1,795.
but is consumed by cattle, domestic sheep, and big
game in arid desert regions. It can be particularly Kornkven, A.B., L.E. Watson, and J.R. Estes.
important during dry years (Hall and Clements 1999. Molecular phylogeny of Artemisia section
1923, McArthur and others 1979). It is important in Tridentatae (Asteraceae) based on chloroplast
the vegetation cover of harsh sandy sites, although DNA restriction site variation. Systematic
limited water often reduces natural seed Botany 24: 69-84.
production. It is particularly useful in blackbrush
and pinyon-juniper communities where sandy McArthur, E.D. 1979. Sagebrush systematics and
outcrops occur. evolution. In: The sagebrush ecosystem: a
Sandsage resprouts vigorously after fire; symposium; 1978 April 27-28, Logan, UT.
fire may be useful in reducing canopy height and College of Natural Resources, Utah State
volume, thus allowing associated herbaceous University, Logan, UT. p. 14-22.
species to express more vigor (Vermeire and
others 2001. McArthur, E.D., A.C. Blauer, A.P. Plummer, and
R. Stevens 1979. Characteristics and
Benefits.—Sandsage can be an important, locally hybridization of important intermountain shrubs.
abundant, native shrub that provides habitat and III. Sunflower family. Research Paper INT-220.
forage for wildlife, including some small nongame U. S. Department of Agriculture, Forest Service
birds. This species helps reduce wind erosion by Intermountain Forest and Range Experiment
stabilizing light sandy soils. People in Mexico Station, Ogden, UT. 82 p.
have used a decoction of leaves to reduce the
effects of intestinal worms and other stomach McArthur, E.D. and C.L. Pope. 1979. Karyotypes
problems (Stubbendieck and others 1997). of four Artemisia species: A. carruthii, A.
filifolia, A. frigida, and A. spinescens. Great
References Basin Naturalist 39: 419-426.
Beetle, A.A. 1979. Autecology of selected Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and
sagebrush species. In: The sagebrush ecosystem: R.G. Nisley, eds. Woody plant seed manual.
a symposium; 1978 April 27-28, Logan, UT. http://www.wpsu.net 12 p.
College of Natural Resources, Utah State
University, Logan, UT. p. 23-26. Moss, E.H. 1940. Interxylary cork in Artemisia
with a reference to its taxonomic significance.
American Journal of Botany 27: 762-768.
61
Stubbendieck, J., S.L. Hatch, and C.H. Butterfield
Nelson, D.L. and R.G. Krebill. 1981. A sagebrush 1997. North American Range Plants. Fifth
wilt disease of unknown origin. Great Basin Edition. University of Nebraska Press, Lincoln,
Naturalist 41: 184-191. NB. 231 p.
Rasmussen, L.L. and J.D. Brotherson. 1986. Vermeire, L.T., R.B. Mitchell, and S.C.
Habitat relationships of sandsage (Artemisia Fuhlendorf. 2001. Sand sagebrush response to
filifolia) in southern Utah. In: McArthur, E.D. fall and spring prescribed burns. In: McArthur,
and B.L. Welch, comps. Proceedings— E.D. and D.J. Fairbanks, comps. Shrubland
symposium on the biology of Artemisia and ecosystem genetics and biodiversity:
Chrysothamnus; 1984 July 9-13, Provo, UT. proceedings; 2000 June 13-15, Provo, UT. U.S.
U.S. Department of Agriculture, Forest Service, Department of Agriculture, Forest Service,
Intermountain Research Station, Ogden, UT. p. Rocky Mountain Research Station, Ogden, UT.
58-66. p. 233-235.
62
Artemisia frigida Willd. fringed sage
ASTERACEAE
63
may grow in association with a variety of grasses Benefits.—Fringed sage is a good pioneer shrub
and forbs as well as with various shrubs, including for stabilizing disturbed sites. It is often used in
big sagebrush (Artemisia tridentata Nutt.), seeding Western strip mines, especially coal areas.
Bigelow sagebrush (Artemisia bigelovii Gray), Its strong taproot and numerous lateral roots help
sandsage (Artemisia filifolia Torr.), and especially stabilize gullies and reduce soil erosion. These
in overgrazed areas, with broom snakeweed rooting characteristics enable the shrub to resist
[Gutierrezia sarothrae (Pursh) Britt. & Rushby]. considerable grazing and trampling. It also
It is also a common understory shrub in ponderosa provides forage for livestock and forage and
pine communities in several Western States habitat for wildlife in both North America and
(McArthur and others 1979). Asia (Li and others 2002, USDA 1937). It is an
important food for sage grouse (Wallestad and
Reproduction.—Blooming occurs from June at others 1975). Fringed sage has some value as a
high elevations and latitudes, to November at medicinal plant (Hall and Clements 1923). Native
lower elevations and latitudes. Fringed sage is a Americans traditionally used fringed sagebrush as
prolific seed producer; each 2.5 cm length of toilet paper, to eliminate unpleasant odors from
inflorescence contains approximately 1,000 seeds dried meat, as a chewed application for wounds
(Harvey 1981). There are about 10 million cleaned prior to bandaging, and to make mats and fans
seed per kg (Plummer and others 1968). Seed (Stubbendieck and others 1997).
matures between September and December
(McArthur and others 1979) and are wind References
dispersed in winter. Seedlings emerge in spring.
Fringed sage can be established by seed, Alley, H. 1972. Range weed control. In:
transplanting young plants, or from rooted Proceedings 1970 Washington State Weed
segments of mature plants. Surface seeding on Control Conference, Yakima, WA. Washington
disturbed soils is recommended (Wasser 1982). State Weed Association. p. 39-45.
Management.—The forage value of fringed sage Cooperrider, A.Y. and J.A. Bailey. 1986. Fringed
varies considerably with location and season sagebrush (Artemisia frigida)—a neglected
(Dietz 1972). On Western ranges, it is most forage species of Western ranges. In: McArthur,
valuable to big game and livestock in late fall, E.D. and B.L Welch, comps. Proceedings—
winter, and early spring (Cooperrider and Bailey symposium on the biology of Artemisia and
1986, Kufeld 1973, USDA 1937, Wasser 1982). Chrysothamnus; 1984 July 9-13, Provo, UT.
Nutritive quality is highest in the spring but General Technical Report INT-200. U.S.
remains adequate throughout much of the year Department of Agriculture, Forest Service,
(Cooperrider and Bailey 1986, Rauzi 1982). Intermountain Research Station, Ogden, UT. p.
Individual plants are susceptible to overgrazing, 46-54.
but populations tend to increase under heavy
grazing pressure (Cooperrider and Bailey 1986). Dietz, D.R. 1972. Nutritive value of shrubs. In:
In the northwestern Great Plains, fringed sage may McKell, C.M., J.P. Blaisdell, and J.R. Goodin,
be less palatable than other available forages and tech. eds. Wildland shrubs—their biology and
occasionally invades deteriorated grasslands. management; 1971 July, Logan, UT. General
Pronghorn (Antilocapra Americana Ord) utilize Technical Report INT-1. U.S. Department of
fringed sage extensively during winter, and to a Agriculture, Forest Service, Intermountain Forest
lesser extent, year round (McArthur and others and Range Experiment Station, Ogden, UT. p.
1979). 289-302.
Fringed sage is well suited to grass and
broad-leafed herb communities. Seedlings are Hall, H.M. and F.E. Clements. 1923. The
competitive and can establish in areas with phylogenetic method in taxonomy, the North
herbaceous competition. It is adapted to mine American species of Artemisia, Chrysothamnus,
spoils, perhaps better than any other species of and Atriplex. Publication 326. Carnegie
Artemisia. It can be used as a biological control to Institution of Washington, Washington, DC.
reduce rapid weed expansion on large disturbed 355 p.
sites but can be controlled when it is too abundant
(Alley 1972). Harvey, S.J. 1981. Life history and reproductive
strategies in Artemisia. M.S. thesis, Montana
State University, Bozeman, MT. 132 p.
64
International Plant Plant Names Index. 2003. Utah. Publication 68-3. Utah Division of Fish
http//www.ipni.org.) [not paged]. and Game, Salt Lake City, UT. 183 p.
Kufeld, R.C. 1973. Foods eaten by the Rocky
Mountain elk. Journal of Range Management Rauzi, F. 1982. Seasonal variations in protein and
26: 106-113. mineral content of fringed sagewort (Artemisia
Li, F.R., A.f. Zhao, H.Y. Zhou, T.H. Zhang, and frigida). Journal of Range Management 35: 679-
X. Zhao. 2002. Effects of simulated grazing on 680.
growth and persistence of Artemisia frigida in a Stubbendieck, J., S.L. Hatch, and C.H.
semiarid sandy rangeland. Grass and Forage Butterfield, 1997. North American Range
Science 57: 239-246. Plants. Fifth Edition. University of Nebraska
Press, Lincoln, NB. 233 p.
McArthur, E.D., A.C. Blauer, A.P. Plummer, and
R. Stevens 1979. Characteristics and U. S. Department of Agriculture, Forest Service.
hybridization of important intermountain shrubs. 1937. Range plant handbook. United States
III. Sunflower family. Research Paper INT-220. Government Printing Office, Washington, DC.
U.S. Department of Agriculture, Forest Service, 816 p.
Intermountain Forest and Range Experiment
Station, Ogden, UT. 82 p. Wallestad, R., J.C. Peterson, and R.L. Eng. 1975.
Foods of adult sage grouse in central Montana.
McArthur, E.D. and C.L. Pope. 1979. Karyotypes Journal of Wildlife Management 39: 628-630.
of four Artemisia species: A. carruthii, A.
filifolia, A. frigida, and A. spinescens. Great Wasser, C.H. 1982. Ecology and culture of
Basin Naturalist 39: 419-426. selected species useful in revegetation disturbed
lands in the west. FSW/OBS-82/56. U. S.
Plummer, A.P., D.R. Christensen, and S.B. Department of the Interior, Fish and Wildlife
Monsen. 1968. Restoring big-game range in Service. Washington, DC. 347 p.
65
Artemisia longiloba (Osterhout) Beetle alkali sagebrush
ASTERACEAE
66
Reproduction.—Shrubs usually bloom in mid- Dealy, J.E., D.A. Leckenby, and D.M. Concannon.
June to early August and seed ripens in August 1981. Wildlife habitats in managed rangelands:
and September. Seeds are wind dispersed during The Great Basin of southeastern Oregon. Plant
late fall and winter, and seedlings emerge during communities and their importance to wildlife.
the following spring. Seeds are about the size of General Technical Report PNW-120. U.S.
low sagebrush seeds so we expect there would be Department of Agriculture, Forest Service,
about 2 million cleaned seeds per kg (Meyer Pacific Northwest Forest and Range Experiment
2003). Natural seeding occurs rapidly following Station, Portland, OR. 66 p.
fires and other disturbances, yet mechanical tillage
can limit establishment if seeds are sown too deep McArthur, E.D., A.C. Blauer, A.P. Plummer, and
or if the seedbed is disrupted (Monsen and Shaw R. Stevens 1979. Characteristics and
1986). hybridization of important intermountain shrubs.
III. Sunflower family. Research Paper INT-220.
Management.—Alkali sagebrush, depending on U.S. Department of Agriculture, Forest Service,
location and ecotype, can be a source of palatable Intermountain Forest and Range Experiment
browse for domestic sheep and provide habitat and Station, Ogden, UT. 82 p.
forage for other animals (Dealy and others 1981,
McArthur and others 1979). Stands of alkali McArthur, E.D., C.L. Pope, and D.C. Freeman.
sagebrush in good condition can support a diverse 1981. Chromosomal studies of subgenus
understory of perennial grasses and annual and Tridentatae of Artemisia: evidence for
perennial forbs. Such stands provide forage and autopolyploidy. American Journal of Botany 68:
cover for sage grouse, antelope, other wildlife, and 589-605.
domestic livestock (Dealy and others 1981,
Monsen and Shaw 1986). Some stands, however, Meyer, S.E. 2003. Artemisia. In: Bonner, F.T., and
are much less diverse and provide little forage or R. Nisley, eds. Woody plant seed manual.
habitat. Sites that lack a satisfactory understory http://www.wpsm.net. [not paged].
remain closed unless shrubs are reduced by fire or
other disturbance. Monsen and Shaw (1986) Monsen, S.B. and N.L. Shaw. 1986. Response of
reported that decadent stands can be rejuvenated an alkali sagebrush/fescue site to restoration
by mechanical means. treatments. In: McArthur, E.D. and B.L Welch,
This shrub has the potential to be a useful plant in comps. Proceedings—symposium on the biology
rehabilitation, especially on heavy, seasonally of Artemisia and Chrysothamnus; 1984 July 9-
dry soils. It has been observed invading 13, Provo, UT. General Technical Report INT-
roadcuts in Echo Canyon, Utah, and along 200. U.S. Department of Agriculture, Forest
roadsides near Kemmerer, Wyoming (McArthur Service, Intermountain Research Station, Ogden,
and others 1979). UT. p. 126-133.
Benefits.—Alkali sagebrush is a native shrub that Passey, H.B. and V.K. Hugie. 1962. Sagebrush on
provides habitat and forage for wildlife and relic ranges in the Snake River plains and
livestock. In addition, it has characteristics that northern Great Plains. Journal of Range
could make it useful in ecological restoration, Management. 15: 273-278.
rehabilitation, or soil conservation. It spreads well
through natural seeding and tolerates disturbance. Robertson, D.R., J.L. Nielsen, and N.H. Bare.
It also grows on argillic soils that may be too 1966. Vegetation and soils of alkali sagebrush
heavily textured for most other shrubs. and adjacent big sagebrush ranges in North Park,
Colorado. Journal of Range Management. 19:
References 17-20.
Beetle, A.A. 1959. New names within the section Tisdale, E.W. and M. Hironaka. 1981. The
Tridentatae of Artemisia. Rhodora 61: 82-85. sagebrush—grass region: a review of the
ecological literature. Bulletin No. 33, Forest,
Beetle, A.A. 1960. A study of sagebrush, the Wildlife, and Range Experiment Station,
section Tridentatae of Artemisia. Bulletin 368, University of Idaho, Moscow, ID. 31 p.
University of Wyoming Agriculture Experiment
Station, Laramie, WY. 83 p.
67
U. S. Department of Agriculture, Natural University, Oregon Agricultural Experiment
Resources Conservation Service— Station, Corvallis, OR. 15 p.
http//www.plants.usda.gov. Last visited August
25, 3003. Zamora, B. and P.T. Tueller. 1973. Artemisia
arbuscula, A. longiloba, and A. nova habitat
Winward, A.H. 1980. Taxonomy and ecology of types in northern Nevada. Great Basin Naturalist.
sagebrush in Oregon. Bull. 642. Oregon State 33: 225-242.
68
Artemisia ludoviciana Nutt. Lousiana sagewort
ASTERACEAE
69
others 1993). Estes (1969) suggested that Lousiana sagewort spreads asexually through
autopolyploidy has been the driving evolutionary rhizomes.
force within the complex. Molecular genetic
studies have also shown New World and Old Management.—Lousiana sagewort is used
World members of the complex to be different differentially by livestock and wildlife throughout
from one another (Vallès and McArthur 2001). its distribution. Populations in the southern
portions of its range, such as in southern Utah,
Range.—Lousiana sagewort is widely distributed New Mexico, and southern Colorado, are more
throughout Western North America. Its range palatable than those in the north (USDA Forest
spans from Ontario and Michigan in the east, to Service 1937, Stubbendieck and others 1997). In
British Columbia in the west, and south through these southern areas, it can provide valuable
Texas, Louisiana, California, and Mexico (USDA forage, especially on spring-fall and winter ranges.
Forest Service 1937). Lousiana sagewort grows at In more northern areas, it has a pungent taste and
various elevations from 250 to 3,050 m. is generally not eaten until after the first frost (Hall
and Clements 1923). Lousiana sagewort is
Ecology.—Lousiana sagewort grows on rocky, generally considered poor forage, although it is
sandy, or gravelly loams on open ridges, slopes, utilized by domestic sheep, deer, and elk to some
and mesas (USDA Forest Service 1937). It is also degree, and may be seasonally important (Shaw
common on plains and prairies and is usually a late and Monsen 1982, Hall and Clements 1923,
seral species that grows after grasses have declined USDA Forest Service 1937). Lousiana sagewort
(Hall and Clements 1923). Lousiana sagewort has also been used to revegetate road, logging, and
grows rapidly, is long lived, and is cold and other disturbances (Shaw and Monsen 1982). In a
drought tolerant (Shaw and Monsen 1982). It revegetation experiment at Mesa Verde National
grows well in sunny sites even though it is shade Park, CO, it survived well when transplanted in
tolerant. It is can be found in grasslands, roadcut sites (Paschke and others 1999).
sagebrush, ponderosa pine, and alpine
communities (Shaw and Monsen 1982). Lousiana Benefits.—Lousiana sagewort is an extensive and
sagewort often associates with asters (Aster L.), genetically diverse native plant that is widespread
goldenrods (Solidago L.), sunflowers (Helianthus throughout Western North America. It provides
L.), fringed sagebrush (Artemisia frigida Wild.), quick and excellent cover, reduces erosion, and
big sagebrush (Artemisia tridentata Nutt.), serves as a nurse plant for other species (Shaw and
tarragon (Artemisia dracunculus L.), needlegrass Monsen 1982). It also has pharmaceutical
(Stipa L.), blue grama [Bouteloua gracilis potential. Zavala-Sanchez and others (2002)
(H.B.K.) Lag. ex Steudel], muhly grasses found that an essential oil found in Lousiana
(Muhlenbergia Schreber), wheatgrasses sagewort, nonanal, inhibited diarrhea in mice. In
(Agropyron Gaertner), mountain dandelions addition, Lousiana sagewort extracts inhibited
[Agoseris glauca (Pursh) Raf.], and western reproduction in malarial parasites (Malagon and
yarrow (Achillea millefolium L.) (USDA Forest others 1997). The ssp. mexicana (estafiate) is used
Service 1937, Hall and Clements 1923). Mean for gastrointestinal pain, as a verimfuge, and as a
annual precipitation within its range varies bitter stimulant (Heinrich 2002). It has also been
between 400 and 1200 mm. used by Native Americans in ceremonial and
purification rituals, to deodorize feet, as pillows,
Reproduction.—Blooming begins in August and saddle pads, and to treat headaches, stomach
ends in September (Stubbendieck and others ailments, and coughs (Stubbendieck and others
1997). Seed matures in the fall and is dispersed by 1997).
wind during late fall or winter. Seeds are
relatively small, numbering about 8,360,000/kg References
(personal communication with S. Stranathan,
Upper Colorado Environmental Plant Center, Estes, J.R. 1969. Evidence for autoploid
Meeker, CO). Seed viability decreases after 2 or 3 evolution in the Artemisia ludoviciana complex
years of storage (personal communication with of the Pacific Northwest. Brittonia 21: 29-43.
K.R. Jorgensen, Great Basin Experimental Area
Ephraim, UT). Seeding aerially, by drill, or Hall, H.M. and F.E. Clements. 1923. The
broadcasting are all effective methods of planting phylogenetic method in taxonomy, the North
and should be conducted during fall or winter American species of Artemisia, Chrysothamnus,
(Shaw and Monsen 1982). In addition to seeds, and Atriplex. Publication no. 326, Carnegie
70
Institution of Washington, Washington, DC. Agriculture, Forest Service, Intermountain
355 p. Research Station, Ogden, UT. p. 123-131.
Heinrich, M. 2002. Ethnobotany, phytochemistry Stubbendieck, J., S.L. Hatch, and C.H Butterfield.
and biological/pharmacological activities of 1997. North American Range Plants. Fifth
Artemisia ludoviciana ssp. mexicana (estafiate). edition. University of Nebraska Press, Lincoln,
In: Wright, C.W., ed. Artemisia, medicinal and NB. 235 p.
aromatic plants Vol. 18. Taylor and Francis,
London, UK. p. 107-117. U.S. Department of Agriculture, Forest Service.
1937. Range plant handbook. United States
Keck, D.D. 1946. A revision of the Artemisia Government Printing Office, Washington, DC.
vulgaris complex in North America. 816 p.
Proceedings of the California Academy of
Sciences 25: 421-468. Vallès, J. and E.D. McArthur. 2001. Artemisia
systematics and phylogeny: cytogenetic and
Malagon, F., J. Vasquez, and A. Ruiz. 1997. molecular insights. In: McArthur, E.D. and D.J.
Antimalaric effect of an alchoholic extract of Fairbanks, comps. Shrubland ecosystem genetics
Artemisia ludoviciana mexicana in a rodent and biodiversity: proceedings. 2000 June 13-15,
malaria model. Parassitologia 39: 3-7. Provo, UT. Proceedings RMRS-P-21. U.S.
Department of Agrigulture, Forest Service,
Paschke, M.W., C. DeLeo, and E.F. Redente 1999. Rocky Mountain Research Station, Ogden, UT.
Revegetation of roadcut slopes in Mesa Verde p. 67-74.
National Park, U.S.A Restoration Ecology 8:
263-268. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Higgins, eds. 1993. A Utah Flora. Second
Shaw, N. and S.B. Monsen. 1982. edition. Printing Services, Brigham Young
Nonleguminous forbs for rangeland sites. In: University, Provo, UT. 986 p.
Monsen, S.B. and N. Shaw, comps.
Proceedings: Managing Intermountain Zavala-Sanchez, M.A., S. Perez-Gutierrez, C.
Rangelands—Improvement of range and Perez-Conzalez, and D. Sanchez-Saldivar.
wildlife habitats; September 15-17, 1981; Twin 2002. Antidiarrhoeal activity of nonanal, an
Falls, ID, June 22-24, 1982; Elko, NV. Gen. aldehyde isolated from Artemisia ludoviciana.
Tech. Rep. INT-157. U.S. Department of Pharmaceutical Biology 40: 263-268.
71
Artemisia nova A. Nels. black sagebrush
ASTERACEAE
72
on sites normally occupied by big sagebrush. Benefits.—Black sagebrush is an important native
Mean annual precipitation throughout the natural shrub in many areas of the Western United States.
range of black sagebrush ordinarily is between 180 It provides structural diversity within native plant
and 320 mm. communities and provides an important browse
Most stands of black sagebrush do not source for native and introduced ungulates. It can
burn because populations are relatively sparse also reduce soil erosion on steep shallow slopes.
(Tisdale and Hironaka 1981). Exceptions to this ‘Gordon Creek’ is a tested germplasm from
rule, however, have been observed in central Utah, Carbon County, Utah, that has been demonstrated
where large stands of black sagebrush burned. In a to be high in nutritive quality and palatability to
comparative study, Nelson and Krebill (1981) wildlife (Welch and others 1994).
found that black sagebrush is less susceptible to a
wilt disease than other species of Artemisia grown References
under similar conditions.
Beetle, A.A. and K.L. Johnson. 1982. Sagebrush in
Reproduction.—Mature shrubs flower in late Wyoming. Bulletin 779. Agricultural
summer (usually August), and seed matures in Experiment Station, University of Wyoming,
September and October. Black sagebrush seed Laramie, WY. 68 p.
tends to be larger than those of other sagebrushes.
There are about 2,000 seeds/g (Deitschman 1974). Behan, B. and B.L. Welch. 1985. Black sagebrush:
Seeds are wind dispersed in late fall or early mule deer winter preference and monoterpenoid
winter. Seedlings emerge in the spring and often content. Journal of Range Management 38: 278-
grow rapidly. Once established, young plants can 280.
persist well even under adverse conditions. With
favorable moisture conditions, mature plants Clary, W.P. 1986. Black sagebrush response to
produce abundant seed, and natural spreading grazing in the east-central Great Basin. In: E.D.
occurs quickly. McArthur and B.L. Welch, comps. Proceedings:
symposium on the biology of Artemisia and
Management.—Black sagebrush is usually Chrysothamnus; 1984 July 9-13; Provo, UT.
considered a valuable browse species for wildlife General Technical Report INT-200. U.S.
and livestock, especially for deer, domestic sheep, Department of Agriculture, Forest Service,
and pronghorn (Clary 1986; McArthur and Intermountain Research Station, Ogden, UT:
Plummer 1978, USDA Forest Service 1937). 181-185 p.
Black sagebrush has good winter nutritive value,
but not as good as big sagebrush (Welch 1983). Deitschman, G.H. 1974. Artemisia. In: C.S.
Palatability, however, can vary significantly Schopmeyer, technical coordinator, Seeds of
between populations (Welch and others 1981). Woody Plants of the United States. Agriculture
Deer and elk preference studies for black and big Handbook 450. U.S. Department of Agriculture,
sagebrushes have yielded inconsistent results Forest Service, Washington, DC. 235-237 p.
(Scholl and others 1977, Nagy and Regelin 1977,
Smith 1950, Wambolt 1996). Welch and others Kelsey, R.G. 1984. Glandular trichomes: a helpful
have shown that some accessions are not eaten taxonomic character for Artemisia nova (black
while others are highly preferred (Behan and sagebrush). Journal of Range Management 37:
Welch 1985, Welch and others 1981), which may 370-372.
partially explain differences in preference studies.
Black sagebrush can also be a valuable McArthur, E.D. 1983. Taxonomy, origin, and
conservation species for dry, shallow, stony soils distribution of sagebrush (Artemisia tridentata)
because it produces abundant seed and seedlings and allies (subgenus Tridentatae). In: K.L.
establish readily. Due to its habitat, it is not Johnson, ed. Proceedings of the First Utah
usually a candidate for plant control. Black Shrub Ecology Workshop; 1981 September 9-
sagebrush can successfully be seeded in fall or 10; Ephraim, UT. College of Natural Resources,
early winter. Seeds should be covered with soil, Utah State University, Logan, UT. 3-13 p.
but remain within 6 mm of the surface. It grows
well with seeded herbs, but understory production McArthur, E.D., A.C. Blauer, A.P. Plummer, and
tends to decrease as the stand matures. R. Stevens 1979. Characteristics and
hybridization of important intermountain shrubs.
III. Sunflower family. Research Paper INT-220.
73
U.S. Department of Agriculture, Forest Service
USDA Forest Service, Intermountain Forest and Welch, B.L. 1983. Improving the nutritive value of
Range Experiment Station, Ogden, UT. 82 p. winter range forage. In: S. B. Monsen and N.
Shaw, comps. Managing Intermountain
McArthur, E.D. and A.P. Plummer. 1978. rangelands—improvement of range and wildlife
Biogeography and management of native habitats: proceedings of a symposium; 1981
western North American shrubs: a case study, September 15-17; Twin Falls, ID; 1982 June 22-
section Tridentatae of Artemisia. Great Basin 24, Elko, NV. General Technical Report INT-
Naturalist Memoirs 2: 229-241. 157, U.S. Department of Agriculture, Forest
Service, Intermountain Forest and Range
McArthur, E.D. and S.C. Sanderson. 1999. Experiment Station, Ogden, UT: 158-164.
Cytogeography and chromosome evolution of
subgenus Tridentatae of Artemisia (Asteraceae). Welch, B.L., E.D. McArthur, and J.N. Davis.
American Journal of Botany 86: 1,754-1,775. 1981. Differential preference of wintering mule
deer for accessions of big sagebrush and black
Nagy, J.G. and W.L. Regelin. 1977. Influence of sagebrush. Journal of Range Management 34:
plant volatile oils on food selection by animals. 409-411.
Congress of Game Biologists 13: 225-230.
Welch, B.L., E.D. Nelson, S.A. Young, A.R.
Nelson, D.L. and R.G. Krebill. 1981. A sagebrush Sands, F.J. Wagstaff, and D.L. Nelson. 1994.
wilt disease of unknown origin. Great Basin ‘Gordon Creek’—a superior, tested germplasm
Naturalist 41: 184-191. of Wyoming big sagebrush. Research Paper INT-
461, U.S. Department of Agriculture, Forest
Smith, A.D. 1950. Sagebrush as a winter feed for Service, Intermountain Research Station. Ogden,
deer. Journal of Wildlife Management 14: 285- UT. 7 p.
289.
Welsh, S.L. and S. Goodrich. 1995. Plant novelties
Scholl, J.P., R.G. Kelsey, and F. Shafizadeh. 1977. in Lepidium (Cruciferae) and Artemisia
Involvement of volatile compounds of Artemisia (Compositae) from the Uinta Basin, Utah. Great
in browse preference by mule deer. Biochemical Basin Naturalist 55: 359-362.
Systematics and Ecology 5: 291-295.
Winward, A.H. 1976. Evolutionary development
Tisdale, E.W. and M. Hironaka. 1981. The of the Artemisia tridentata taxa. In: H. C. Stutz,
sagebrush—grass region: a review of the ed. Wildland shrubs: proceedings, symposium,
ecological literature. Bulletin 33, Forest, and workshop; 1975 November 4-6, Provo, UT.
Wildlife, and Range Experiment Station, College of Biology and Agriculture, Brigham
University of Idaho, Moscow, ID. 31 p. Young University, Provo, UT: 163.
U.S. Department of Agriculture, Forest Service. Zamora, B. and P.T. Tueller. 1973. Artemisia
1937. Range plant handbook. United States arbuscula, A. longiloba, and A. nova habitat
Government Printing Office, Washington, DC. types in northern Nevada. Great Basin Naturalist
816 p. 33: 225-242.
Wambolt, C.L. 1996. Mule deer and elk foraging
preference for 4 sagebrush taxa. Journal of
Range Management 49: 499-503.
74
Artemisia pygmaea Gray pygmy sagebrush
ASTERACEAE
75
Benefits.—Although it is not abundant, pygmy III. Sunflower family. Research Paper INT-220.
sagebrush does provide important soil cover in U.S. Department of Agriculture, Forest Service
dry, alkaline habitats where little else grows. It USDA Forest Service, Intermountain Forest and
also adds to the overall biological diversity to the Range Experiment Station, Ogden, UT. 82 p.
shrub communities where it is found.
McArthur, E.D. and A.P. Plummer. 1978.
References Biogeography and management of native
western shrubs: a case study, section Tridentatae
Beetle, A.A. 1960. A study of sagebrush, the of Artemisia. Great Basin Naturalist Memoirs 2:
section Tridentatae of Artemisia. Bulletin 368, 229-243.
University of Wyoming Agriculture Experiment
Station, Laramie, WY. 83 p. McArthur, E.D. and S.C. Sanderson. 1999.
Cytogeography and chromosome evolution of
Cronquist, A. 1994. Asterales, Intermountain flora, subgenus Tridentatae of Artemisia (Asteraceae).
volume 5. The New York Botanical Garden, American Journal of Botany 86: 1,754-1,775.
Bronx, NY. 496 p.
Ward, G.H. 1953. Artemisia, section Seriphicum,
McArthur, E.D., A.C. Blauer, A.P. Plummer, and in North America. A cytotaxonomic study.
R. Stevens. 1979. Characteristics and Contributions from the Dudley Herbarium 4:
hybridization of important intermountain shrubs. 155-205.
76
Artemisia rigida (Nutt.) Gray stiff sagebrush
ASTERACEAE
77
its present range suggests (McArthur and others Hall, H.M. and F.E. Clements. 1923. The
1979). phylogenetic method in taxonomy; the North
American species of Artemisia, Chrysothamnus,
Reproduction.—Blooming begins in late August and Atriplex. Publication 326. Carnegie
and ends in early October, and seeds ripen in Institution of Washington, Washington, DC.
October and November. Seed is wind dispersed in 355 p.
late fall or winter, and seedlings emerge during the
ensuing spring. Seed production and seed quality McArthur, E.D., A.C. Blauer, A.P. Plummer, and
are low in comparison to other sagebrush species R. Stevens 1979. Characteristics and
(McArthur and others 1979). Attempts to seed and hybridization of important intermountain shrubs.
transplant this species in arid sagebrush ranges III. Sunflower family. Research Paper INT-220.
near Boise, Idaho, have been unsuccessful. In U.S. Department of Agriculture, Forest Service
such instances, plantings were made where USDA Forest Service, Intermountain Forest and
cheatgrass (Bromus tectorum L.) and other annuals Range Experiment Station, Ogden, UT. 82 p.
have invaded, and stiff sagebrush seedlings were
unable to compete. Plantings have successfully McArthur, E.D., C.L. Pope, and D.C. Freeman.
established on prepared seedbeds, however. 1981. Chromosomal studies of subgenus
Tridentatae of Artemisia: evidence for
Management.—Due to scant foliage and stiff autopolyploidy. American Journal of Botany 68:
branches, few animals except domestic sheep 589-605
browse stiff sagebrush (Hall and Clements 1923).
It provides forage on dry sites for domestic sheep McArthur, E.D. and S.C. Sanderson. 1999.
during midsummer when herbaceous plants are Cytogeography and chromosome evolution of
dry. subgenus Tridentatae of Artemisia (Asteraceae).
American Journal of Botany 86: 1754-1775.
Benefits.—Stiff sagebrush is a native shrub that
adds biological diversity to Western ecosystems. Morris, M.S., R.G. Kelsey, and D. Griggs. 1976.
It is an important plant that provides cover and The geographic and ecological distribution of big
reduces erosion on rocky sites. Its ecology sagebrush and other woody Artemisias in
suggests that it could be used in reclaiming harsh, Montana. Proceedings Montana Academy of
disturbed sites. Sciences 36: 56-79.
78
Artemisia rothrockii Gray. rothrock sagebrush
ASTERACEAE
79
snow-melt. Rothrock sagebrush does not form a Berlow, E.L., C.M. D’Antonio, and S.A.
large soil seedbank (Reynolds unpublished data). Reynolds. 2002. Shrub expansion in montane
An average sized adult plant can produce over meadows: the interaction of local-scale
5000 seeds (Reynolds unpublished data). disturbance and site aridity. Ecological
However, field germination rates are low (< 1 Applications 12: 1103-1118.
percent), and seed viability appears to decline
rapidly after 1 year (Berlow and others 2002, and Berlow, E.L., C.M. D’Antonio, and H. Swartz.
unpublished data). 2003. Response of herbs to shrub removal across
natural and experimental variation in soil
Growth and Management.—The growing season moisture: Implications for restoration. Ecological
for rothrock sagebrush in montane habitats is Applications. (in press).
relatively short, beginning in early spring and
continuing until the soil moisture is depleted in Dull, R.A. 1995. Palynological evidence for 19th
late summer. Under mesic conditions, rothrock century grazing-induced vegetation change in
sagebrush can begin flowering within 5 years of the southern Sierra Nevada, California. Journal
establishment (Berlow and others 2002). While of Biogeography 26: 899-912.
age to senescence is unknown, individuals up to 40
years old have been documented by counting Kattelmann, R. and M. Embury. 1996. Riparian
growth rings (Bauer and others 2002). Rates of areas and wetlands. In Sierra Nevada
increase in stem diameter have been observed to Ecosystem Project. University of California,
vary between 0.10 to 0.14 mm/year. Faster growth Centers for Water and Wildland Resources,
rates were found in microhabitats characterized by Davis, CA. p. 34-68.
moist surface soil and a shallow water table,
suggesting that this woody species can use similar Kirchner, J.W., L. Micheli, and J.D. Farrington.
water sources as herbaceous species (Bauer and 1998. Effects of herbaceous riparian vegetation
others 2002, Berlow and others 2003). Rothrock on streambank stability. Technical Completion
sagebrush seedlings are not palatable to livestock Report, Project #W-872, University of
or native grazers, which may contribute to their California Berkeley Water Resources Center.
encroachment into intensively grazed meadows. 43 p.
While large areas of sagebrush expansion are
associated with stream incision, rothrock Knapp, R.A. and K.R. Matthews. 1996.
sagebrush also has the demonstrated potential to Livestock grazing, Golden Trout, and streams
invade un-incised, mesic meadow habitat given the in the Golden Trout Wilderness, California:
presence of exposed soil and a nearby seed supply impacts and management implications. North
(Berlow and others 2002). Thus, on a large scale, American Journal of Fisheries Management 16:
preventing further stream incision is critical to 805-820.
slowing sagebrush invasion. Within mesic
meadow habitat, removal of scattered individual McMinn, H.E. 1951. An Illustrated Manual of
shrubs that serve as local seed sources may be the California Shrubs. University of California
most effective way to prevent rapid expansion into Press, Berkeley, CA. p. 608-609.
un-incised areas.
Odion, D.C., T.L. Dudley, and C.M. D'Antonio.
Benefits.—Rothrock sagebrush contributes to the 1988. Cattle grazing in southeastern Sierran
diversity of California’s montane plant meadows: Ecosystem change and prospects for
communities. It stabilizes sandy soil on the slopes recovery. In: C.A. Hall and V. Doyle-Jones,
of meadow fringes and provides habitat for a editors. Plant Biology of Eastern California,
variety of birds and small mammals. Mary Decker Symposium. White Mountain
Research Station, Los Angeles, CA. p. 277-292.
References
Ratliff, R.D. 1985. Meadows in the Sierra Nevada
Bauer, K., E.L. Berlow, and C.M. D’Antonio. of California: state of knowledge. General
2002. Shrub expansion in montane meadows: Technical Report PSW-84. U.S. Department of
The relationship between climate and Rothrock Agriculture, Forest Service, Pacific Southwest
sagebrush colonization patterns. Journal of Forest and Range Experiment Station.
Range Management. 55: 620-625. Berkeley, CA. 52 p.
80
Sawyer, J.O. and T. Keeler-Wolf. 1995. A Manual USDA Forest Service. 1998. Kern Plateau
of California Vegetation. California Native Plant Ecosystem Analysis. U.S. Department of
Society, Sacramento, CA. p. 188. Agriculture, Forest Service, Inyo National
Forest, Bishop, CA. 215 p.
Shultz, L.M. 1993. Artemisia. In: J.C. Hickman,
ed. The Jepson Manual: Higher Plants of USDA Forest Service. 2000. Templeton and
California. University of California Press, Whitney Grazing Allotments Environmental
Berkeley, CA. p. 202-205. Assessment. U.S. Department of Agriculture,
Forest Service, Inyo National Forest, Bishop,
CA. 111 p.
81
Artemisia spinescens Eaton budsage
ASTERACEAE
82
Reproduction.—Blooming normally begins Cronquist, A. 1994. Asterales, Intermountain flora,
during the last week in April and continues through Vol. 5. The New York Botanical Garden, Bronx,
the last week in May. Exceptions, however, have NY. 496 p.
been documented where blooming began as early
as late March or has extended as late as mid-June Hall, H.M., and F.E. Clements. 1923. The
(Wood 1966). Seeds normally mature in early phylogenetic method in taxonomy; the North
June. There are about 3 million cleaned seeds per American species of Artemisia, Chrysothamnus,
kg (Meyer 2003). Freezing temperatures in early and Atriplex. Publication 326. Carnegie
spring frequently kill developing embryos. Intact Institution of Washington, Washington, DC.
heads regularly fall from the plant without breaking 355 p.
apart to release seeds, although some seeds are
usually dispersed independently. Abundant natural Holmgren, R.C. and S.S. Hutchings. 1972. Salt
reproduction occurs in years when seed production desert shrub response to grazing use. In: McKell,
is plentiful and moisture conditions are favorable. C.M., J.P. Blaisdell, and J.R. Goodin, tech. eds.
Proceedings-symposium on wildland
Management.—Budsage is a palatable, nutritious shrubs--their biology and utilization, 1971 July;
winter forage for upland birds, small game, big Logan, UT. Gen. Tech. Rep. INT-1. U.S.
game, and domestic sheep. It is particularly Department of Agriculture, Forest Service,
preferred just after dormancy. Budsage is high in Intermountain Forest and Range Experiment
calcium, magnesium, phosphorous, and protein Station, Ogden, UT. p. 153-164.
(Wood and Brotherson 1986). It is usually more
palatable in late winter than early winter McArthur, E.D., A.C. Blauer, A.P. Plummer, and
(Holmgren and Hutchings 1972). Palatability R. Stevens 1979. Characteristics and
again decreases once twigs have elongated and hybridization of important intermountain shrubs.
volatile oil content increases (Cook and others III. Sunflower family. Research Paper INT-220.
1954). Budsage is especially important to grazing U.S Department of Agriculture, Forest Service,
animals during late winter in areas where there is Intermountain Forest and Range Experiment
an abundance of dry grass. Unfortunately, it does Station, Ogden, UT. 82 p.
not tolerate grazing well, and continual heavy
grazing may eliminate it from certain areas McArthur, E.D. and C.L. Pope. 1979. Karyotypes
(Holmgren and Hutchings 1972). In order to of four Artemisia species: A. carruthii, A.
maintain the species in plant communities, filifolia, A. frigida, and A. spinescens. Great
periodic rest from grazing is needed. This allows Basin Naturalist 39: 419-426.
seed and seedlings to develop.
Meyer, S. E. 2003. Artemisia. In: Woody plant
Benefits.—Budsage is an important native shrub seed manual. http://www.wpsu.net [not paged].
because of its forage value to various species of
wildlife and domestic livestock. Wildlife species Vallès, J. and E.D. McArthur. 2001. Artemisia
that utilize budsage include mule deer (Odocoileus systematics and phylogeny: cytogenetic and
heminonus Rafinesque), pronghorn (Antilocapra molecular insights. In: E.D. McArthur and D.J.
Americana Ord), mountain sheep (Ovis canadensis Fairbanks, comps. Shrubland ecosystem genetics
Shaw), black-tailed jackrabbits (Lepus californicus and biodiversity: proceedings, 2000 June 13-15;
Gray), and chucker partridge (Alectoris graeca Provo, UT. Proceedings RMRS-P-21. U.S
Meisner) (Wood and Brotherson 1986). In areas Department of Agriculture, Forest Service,
of local abundance, budsage can be an important Rocky Mountain Research Station, Ogden, UT.
source of protein during late winter months. p. 67-74.
83
comps. Proceedings--symposium on the biology
of Artemisia and Chrysothamnus, 1984 July
9-13; Provo, UT. General Technical Report
INT-200. U.S. Department of Agriculture, Forest
Service, Intermountain Forest and Range
Experiment Station, Ogden, UT. p. 75-92.
84
Artemisia tridentata Nutt. big sagebrush
COMPOSITAE
Bruce L. Welch
85
western Wyoming; southern-central British seed dispersal is highly variable among stands and
Columbia, south through central Washington, to somewhat dependent on elevation; some starting in
the Columbia River; from the south side of the mid-fall other not until early winter. Most seeds
Blue Mountains in northeastern Oregon, germinate immediately after snow melt or after
throughout central and southeastern Oregon, late-winter or early spring storms (Welch 2002).
southwestern and central Idaho to northern Utah, The seeds possess no special adaptations for wind
Nevada, and northwestern California; from the dispersal, but are wind dispersed to a maximum
northern great basin areas of California, Nevada, distance of 30 m from the mother plant. Hence
and Utah, extending southward at elevations from seedlings placement is heavily depended on wind
5,000 to 7,000 feet and extending into western direction (Welch 2002). Big sagebrush forms a
Colorado, northwestern New Mexico, northern weak soil seed bank (Welch 2002). A single
Arizona; at much lower elevations in southern mature big sagebrush plant can produced upward
California and northern Lower California, of 500,000 seeds, although, excessive browsing
Mexico.” It also occupies areas of the northern can reduce inflorescences production by a factor of
Great Plains in the States of North and South 20 (Welch 2002). The seeds must lie on the soil
Dakota, Montana, Wyoming, and Nebraska surface or be buried not more than 5 mm for
(Welch 2002). successful emergence (Welch 2002).
86
of bird’s beaks, 3 species of broom rapes, and a
host of large and small mammals, birds, and Krannitz, P.G. and M.M. Caldwell. 1995. Root
reptiles (Welch 2002). Soil fertility is higher growth responses of three Great Basin
under big sagebrush canopy cover than outside the perennials to intra-and inter specific contact with
canopy (Welch 2002). Roots of other plants can other roots. Flora. 190:161-167.
tap into these islands of fertility (Krannitz and
Caldwell 1995). Also, by a process known as McArthur, E.D., A.C. Blauer, A.P. Plummer, and
“hydraulic lift or hydraulic redistribution” R. Stevens. 1979. Characteristics and
associated plants receive part of their needs from hybridization of important Intermountain shrubs.
water pumped by big sagebrush to the soil surface III. Sunflower family. Research Paper INT-220.
from greater soil depths (Welch 2002). Lastly, big U.S. Department of Agriculture, Forest Service,
sagebrush provides a safe haven for grasses and Intermountain Forest and Range Experiment
forbs growing under its canopy from grazing Station, Ogden, UT. 82 p.
domestic livestock (Welch 2002).
McArthur, E.D. and S.K. Goodrich. 1986.
References Artemisia tridentate ssp. spiciformis:
distribution and taxonomic placement. In: E.D.
Bailey, W.W. 1896. The sage brush. American McArthur and B.L. Welch, comps. Proceedings:
Naturalist 3:356-360. Symposium on the biology of Artemisia and
Chrysothamnus; July 9-13, 1984; Provo, UT.
Beetle, A.A. 1960. A study of sagebrush–The Gen. Tech. Rep. INT-200. U.S. Department of
section Tridentatae of Artemisia. Bulletin 368. Agriculture, Forest Service, Intermountain
University of Wyoming, Agricultural Research Station, Ogden, UT. 55-57.
Experiment Station. Laramie. 83 p.
McArthur, E.D. and S.C. Sanderson. 1999.
Beetle, A.A. and K.L. Johnson. 1982. Cytogeography and chromosome evolution of
Sagebrush in Wyoming. B-779, University of subgenus Tridentatae of Artemisia (Asteraceae).
Wyoming, Agricultural Experiment Station, American Journal of Botany. 86:1,754-1,775.
Laramie. 68p.
Welch, B.L. [in press]. Big sagebrush: A sea
Beetle, A.A. and A. Young. 1965. A third fragmented into lakes, puddles, and ponds.
subspecies in the Artemisia tridentata complex. General Technical Report. U.S. Department of
Rhodora 67:405-406. Agriculture, Forest Service, Rocky Mountain
Research Station, Fort Collins, CO.
Diettert, R.A. 1938. The morphology of Artemisia
tridentata Nutt. Lloydia 1:3-74.
87
Artemisia tripartita Rydb. threetip sagebrush
ASTERACEAE
88
Range.—Threetip sagebrush covers approximately sagebrush generally recovers well after
34,000 km2 in the Northern Rocky Mountains and disturbance, but hot summer fires can kill large
Great Basin States. It usually grows at elevations stands, inhibit resprouting, and significantly delay
between 910 to 2,700 m. It extends from British natural seedling recovery.
Columbia south through Montana and Wyoming to
Colorado, and west to Washington, Oregon, Benefits.—Three-tip sagebrush adds structural and
northern Nevada, and northern Utah (Beetle 1960). biological diversity to the landscape and reduces
Subspecies rubicola, however, is limited to areas the effects of wind and soil erosion. Although
of central and southeastern Wyoming and southern Beetle (1960) reported it was not palatable to
Oregon (Brunner 1972). livestock or game, Kufeld (1973) reported that elk
used it, and Kufeld and others (1973) also reported
Ecology.—Threetip sagebrush grows well on that deer use it. A form near Salmon, Idaho, is
moderate to deep, well-drained, loamy and sandy palatable to deer and may have introgressed into a
loam soils (Winward 1980). In some places, nearby population of Wyoming big sagebrush
particularly in Idaho, this species grows between (McArthur and others 1979).
the lower, hot, dry sites dominated by Wyoming
big sagebrush (Artemisia tridentata Nutt. ssp. References
wyomingensis Beetle & Young) and the higher,
cooler sites dominated by mountain big sagebrush Beetle, A.A. 1959. New names within the section
[Artemisia tridentata ssp. vaseyana (Rydb) Beetle] Tridentatae of Artemisia. Rhodora 61: 82-85.
(Schlatterer 1973). It often grows intermixed, but
as separate stands, with mountain big sagebrush, Beetle, A.A. 1960. A study of sagebrush. The
Wyoming big sagebrush, or alkali sagebrush section Tridentatae of Artemisia. Bulletin 368.
[Artemisia longiloba ([Osterh.) Beetle] in areas University of Wyoming, Agricultural
with 300 to 600 mm of annual precipitation. Experiment Station, Laramie, WY. 83 p.
Wyoming threetip sagebrush grows on rocky
knolls at high elevations (2,100 to 2,700 m) Brunner, J.R. 1972. Observations on Artemisia in
adjacent to mountain big sagebrush sites. Nevada. Journal of Range Management 25: 205-
208.
Reproduction.—Flowers bloom between July and
September and seed matures in October. Seeds are Cronquist, A. 1994. Asterales, Intermountain flora,
wind dispersed and seedlings emerge during the Vol. 5. The New York Botanical Garden, Bronx,
ensuing spring. Plants grow quickly and attain NY. 496 p.
maturity in 3 to 5 years. The seeds are smaller than
many Artemisia species; there are approximately 5 Kufeld, R.C. 1973. Foods eaten by the Rocky
million cleaned seeds/kg (Meyer 2003). Mountain elk. Journal of Range Management.
26: 106-113.
Management.—This species is a vigorous seeder.
Some forms are more palatable than others. Kufeld, R.C., O.C. Wallmo, and C. Feddema.
Threetip sagebrush establishes well by direct 1973. Foods of the Rocky Mountain mule deer.
seeding, rearing, or transplanting. Seedlings are Research Paper RM-11. U.S. Department of
able to persist with both a dense perennial and Agriculture, Forest Service, Rocky Mountain
annual understory, although it is no more effective Forest and Range Experiment Station, Fort
at excluding invasive annuals than other Collins, CO. 31 p.
sagebrushes. It frequently grows with a number of
other shrubs and herbs. It is compatible with many McArthur, E.D., A.C. Blauer, A.P. Plummer, and
introduced and native grasses including mountain R. Stevens 1979. Characteristics and
brome (Bromus carinatus H. & A.), slender hybridization of important intermountain shrubs.
wheatgrass [Elymus trachycaulus (Link) Gould ex III. Sunflower family. Research Paper INT-220.
Shinn], intermediate wheatgrass [Elymus hispidus U.S. Department of Agriculture, Forest Service,
(Opiz) Meld], and orchard grass (Dactylis Intermountain Forest and Range Experiment
glomerata L.). Station, Ogden, UT. 82 p.
Attempts to establish threetip sagebrush
on arid sites dominated by Wyoming big McArthur, E.D. and S.C. Sanderson. 1999.
sagebrush have been unsuccessful. Some ecotypes, Cytogeography and chromosome evolution of
however, can extend into drier sites. Threetip
89
subgenus Tridentatae of Artemisia (Asteraceae). Schlatterer, E.F. 1973. Sagebrush species and
American Journal of Botany 86: 1,754-1,775. subspecies. U.S. Department of Agriculture,
Forest Service, R-4 Range Improvement Notes
Meyer, S.E. 2003. Artemisia. In: F.T. Bonner, and 18(2): 1-11.
R.G. Nisley, eds. Woody plant seed manual.
http://www.wpsu.net [not paged]. Winward, A.H. 1980. Taxonomy and ecology of
sagebrush in Oregon. Bullitin 642. Oregon State
Pechanec, J. F., A.P. Plummer, J.H. Robertson, University, Oregon Agricultural Experiment
and A.C. Hull, Jr. 1965. Sagebrush control on Station, Corvallis, OR. 15 p.
rangelands. Agriculture. Handbook 277. U.S.
Department of Agriculture, Washington, DC.
40 p.
90
Arundinaria gigantea (Walt.) Muhl. giant cane
POACEAE
Kristina Connor
91
Reproduction.—Giant cane primarily spreads by References
rapid vegetative reproduction from large rhizomes.
It flowers infrequently, and at irregular intervals, Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
in early spring, forming simple panicles on the A Concise Dictionary of Plants Cultivated in the
branches. The panicles consist of a few racemes of United States and Canada. McMillan Publishing
five to 15 large, stalked spikelets, which may be Co., Inc. New York. 1,290 p.
purple (Bailey and Bailey 1976, Brickell and Zuk
1996). The inflorescence may be a single axis or Brickell, C. and J.D.. Zuk. 1996. The American
have branches. Seed production from these flowers Horticultural Society A-Z Encyclopedia of
is sparse and unpredictable. Garden Plants. DK Publishing, Inc., New York.
1,092 p.
Growth and Management.—Giant cane is a
short-lived, evergreen monocot. Individual stems Grelen, H.E. and V.L. Duvall. 1966. Common
survive less than 10 years, but the species sprouts plants of longleaf pine-bluestem range. Res. Pap.
prolifically from its rhizomes. In the presence of S0-23, U.S. Department of Agriculture Forest
fire and on the best-suited sites, it forms dense Service, Southern Forest Experiment Station,
thickets called canebrakes. While it can withstand New Orleans, LA. 96 p.
flooding, drought, and intense surface fires, giant
cane is sensitive to overgrazing (Grelen and Hitchcock, A.S. 1971. Manual of Grasses of the
Duvall 1966, Hitchcock 1971). One source reports United States. Vol. 1, 2nd Ed. Revised by A.
rapid growth (Brickell and Zuk 1996), especially Chase. Dover Publications Inc., New York. 569
after a fire. Fuel accumulates rapidly in a p.
canebrake, and, to eliminate the hazard, a short
burning cycle is advised (Hughes 1966, Walkup Hughes, R.H. 1966. Fire ecology of canebrakes.
1991). In: Proceedings 5th Annual Tall Timbers Fire
Ecology Conference, Tallahassee, FL. Tall
Benefits.—The dense thickets of giant cane Timbers Research Station. p. 148-158.
provide cover for nesting birds and small
mammals. In addition, the young shoots are edible, University of Florida. 2001. Website, Center for
sometimes used as a potherb (Hitchcock 1971), Aquatic and Invasive Plants. http://aquat1.ifas.
and of good nutritional quality. The leaves are a ufl.edu/arugig.html. 1 p.
preferred food for southern pearly eye butterfly
caterpillars (University of Florida 2001). The USDA, NRCS. 2001. The PLANTS database,
leaves extend well above the ground, plants can be National Plant Data Center, Baton Rouge, LA.
completely defoliated by cattle and are also http://plants.usda.gov. [not paged].
uprooted by swine (Hitchcock 1971, Walkup
1991). Thus, while grazing capacity of cane is Walkup, C. 1991. Arundinaria gigantea. In: U.S.
high, careful management is required to prevent Department of Agriculture, Forest Service, Rocky
deterioration of the plants (Grelen and Duvall Mountain Research Station Fire Sciences
1966). Laboratory. Fire Effects Information System.
http://www.fs.fed. us/database/feis. 9 p.
92
Atriplex canescens (Pursh) Nutt. fourwing saltbush
CHENOPODIACEAE
93
distribution usually ranges between 150 and 400 and grows rapidly (Cibils 1998, McArthur and
mm. It grows on deep, usually halophytic, well- others 1983, Peterson and others 1987). Leaves,
drained sandy soils, gravelly washes, mesas, stems, and fruits provide browse throughout the
ridges, and slopes. Soil salts enhance growth of year. Winter protein content has been shown to
fourwing saltbush by increasing days to wilting, vary between accessions (Welch and Monsen
organic matter production, water use efficiency, 1981). Relative saponin concentrations may affect
and ability to extract water by means of osmotic palatability. In mixed species communities,
adjustments (Glenn and Brown 1998). It often herbage production of fourwing saltbush and forbs
grows intermixed with a wide variety of shrubs, remains quite high. Fourwing saltbush is a
grasses and forbs. facultative selenium absorber, making it mildly
poisonous to browsing animals in selenium-rich
Reproduction.—Flowering generally occurs soils (Davis 1972).
between May and September (Blauer and others When propagating fourwing saltbush, it is
1976). This period can vary, however, with important to select ecotypes that are adapted to the
genotype and location. Flowers are wind site. If possible, seed should be from a source with
pollinated, and seed matures in October. Most a similar climate unless the strain being planted is
wildland stands of fourwing saltbush produce known to have a broad ecological range. Climatic
abundant seed 3 out of every 5 years. and biological factors may affect plant persistence.
Seed fill in high quality lots may only Plants established from southern seed sources have
reach 50 percent, although rates of 40 percent or not persisted in colder northern areas. However,
lower are generally considered substandard (Meyer seed collected from colder areas have done well at
2003). Like many species of saltbush, Atriplex warmer sites (McArthur and others 1983, Plummer
canescens exhibits high levels of dormancy and others 1966).
controlled by multiple and complex mechanisms
(Meyer 2003). Woody bracts inhibit germination Benefits.—Fourwing saltbush is a valuable plant
mechanically and also contain water-soluble that provides livestock and wildlife habitat and
substances, salt and perhaps saponin, which inhibit food. It has been widely and successfully used in
germination (Clor and others 1989). Seed normally revegetation and restoration projects including soil
requires 20 to 30 days of stratification to assure stabilization (Blauer and others 1976, Plummer and
uniform germination. There are about 120,000 others 1966). Its ability to adapt and to hybridize
cleaned, dewinged seeds (utricles)/kg (Blauer and with other woody Atriplex can facilitate in situ
others 1976). selection during revegetation (Stutz 1982).
Artificial seeding should take place during
the fall or winter unless spring frosts threaten References
seedling survival. Fall and winter seeding promotes
seed stratification and overall germinability. When Barrow, J.R. 1997. Natural asexual reproduction
sown, seeds should remain within the top 1.25 cm in saltbush Atriplex canescens (Pursh) Nutt.
of soil. Broadcast seeding followed by chaining Journal of Arid Environments 36: 267-270.
has produced excellent stands. Shrubs can also be
established using drills alone or in separate rows Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
among other species. Care should be taken to keep Stevens, and B.C. Giunta. 1976. Characteristics
herbs from displacing fourwing saltbush seedlings. and hybridization of important Intermountain
Seedlings grow vigorously and quickly produce a shrubs. II. Chenopod family. Research Paper
dense canopy although interspecific competition INT-177. U.S. Department of Agriculture, Forest
can reduce stand densities. Service, Intermountain Forest and Range
Shrubs can also be grown from nursery Experiment Station, Ogden, UT. 42 p.
stock, container stock, or by stem cuttings (Shaw
and Monsen 1984). Rooted stem cuttings have Cibils, A.F., D.M. Swift, and E.D. McArthur.
been used successfully to establish shrub seed 1998. Plant-herbivore interactions in Atriplex:
orchards with desirable traits (McArthur and others current state of knowledge. General Technical
1978, 1992). Some populations reproduce Report RMRS-GTR-14. U.S. Department of
asexually through root sprouting (Barrow 1997). Agriculture, Forest Service, Rocky Mountain
Research Station, Ogden, UT. 31 p.
Management.—Fourwing saltbush is a valuable
forage shrub because it is abundant, palatable, Clor, M.A., F.M.R. Al-Charachafchi, and N.
provides large quantities of forage, is nutritious, Mahmood. 1989. Seed dormancy and
94
germination inhibition of Atriplex canescens. Plummer, A.P., S.B. Monsen, and D.R.
Journal of Agriculture and Water Resources Christensen. 1966. Fourwing saltbush, a shrub
Research Plant Production 8: 55-64. for future game ranges. Publication 66-4. Salt
Lake City, UT: Utah State Department of Fish
Davis, A.M. 1972. Selenium accumulation in a and Game. 12 p.
collection of Atriplex species. Agronomy
Journal. 64: 823-824. Sanderson, S.C. and H.C. Stutz. 1994.
Chromosome numbers in Mojave and Sonoran
Freeman, D.C., E.D. McArthur, S.C. Sanderson, Desert Atriplex canescens (Chenopodiaceae).
and A.R. Tiedemann. 1993. The influence of American Journal of Botany 81: 1045-1053.
topography on male and female fitness
components of Atriplex canescens. Oecologia Sanderson, S.C. and H.C. Stutz. 2001.
93: 538-547. Chromosome races of fourwing saltbush
(Atriplex canescens), Chenopodiaceae. In: E.D.
Glenn, E.P. and J.J. Brown. 1998. Effects of salt McArthur and D.J. Fairbanks, comps.
levels on the growth and water use efficiency of Proceedings—shrubland ecosystem genetics and
Atriplex canescens (Chenopodicaceae) varieties biodiversity; 2000 June 13-15; Provo, UT. Proc.
in drying soil. American Journal of Botany 85: RMRS-P-21. U.S. Department of Agriculture,
10-16. Forest Service, Rocky Mountain Research
Station, Ogden, UT. p. 75-88.
McArthur, E.D. 1977. Environmentally induced
changes of sex expression in Atriplex canescens. Shaw, N. and S.B. Monsen. 1984. Nursery
Heredity 38: 97-193. propagation and outplanting of bareroot
chenopod seedlings. In: A.R. Tiedemann, E.D.
McArthur, E.D., D.C. Freeman, L.S. Luckinbill, McArthur, H.C. Stutz, R Stevens, and K.L.
S.C. Sanderson, and G.L. Noller. 1992. Are Johnson, K., comps. Proceedings-symposium on
triocy and sexual morphs of Atriplex canescens the biology of Atriplex and related chenopods,
genetically based: evidence from clonal studies. 1983 May 2-6; Provo, UT. Gen. Tech. Rep.
Evolution 46: 1708-1721. INT-172. U.S. Department of Agriculture, Forest
Service, Intermountain Forest and Range
McArthur, E.D., A.P. Plummer, G.A. Van Epps, Experiment Station, Ogden, UT. p. 251-260.
D.C. Freeman, and K.R. Jorgensen. 1978.
Producing fourwing saltbush seed in seed Stutz, H.C. 1982. Broad gene pools required for
orchards. In: Hyder, D. N., ed. Proceedings of disturbed lands. In: E.F. Aldon and W. Oaks W.
the First International Rangeland Congress; eds. Reclamation of mined lands in the
1978 Aug. 14-18; Denver, CO. Denver, CO: southwest: Proceedings of the symposium; 1982
Society for Range Management: 406-410. Oct. 20-22; Albuquerque, NM. Soil
Conservation Society of America, New Mexico
McArthur, E.D., R. Stevens, and A.C. Blauer.
Chapter, Albuquerque, NM. p.113-118.
1983. Growth performance comparisons among
18 accessions of fourwing saltbush (Atriplex
Stutz, H.C. 1978. Explosive evolution of perennial
canescens) at two sites in central Utah. Journal
Atriplex in western America. Great Basin
of Range Management. 36: 78-81.
Naturalist Memoirs. 2: 161-168.
Meyer, S.E. 2003. Atriplex L. In: Woody Plant Welch, B.L. and S.B. Monsen. 1981. Winter crude
Seed Manual, http://www.wpsm.net [not paged]. protein among accessions of fourwing saltbush
grown in a uniform garden. Great Basin
Peterson, J.L., D.N. Ueckert, R.L. Potter, and J.E. Naturalist. 41: 343-346.
Huston. 1987. Ecotype variation in western
Texas fourwing saltbush. Journal of Range
Management. 40: 361-366.
95
Atriplex confertifolia (Torr. & Frém.) S. Wats. shadscale
CHENOPODIACEAE
96
typically grows in arid climates where mean fruits have fallen in autumn. Grazing pressure can
annual precipitation can range between 180 and reduce more palatable shrub species, such as
360 mm. budsage and blacksage, resulting in increased
shadscale populations (Blaisdell and Holmgren
Reproduction.—Shadscale usually blooms during 1984). Shadscale may be more effective in
April and May. In isolated locations populations restoring and stabilizing salty areas than fourwing
may bloom as early as January and as late as June. saltbush or other saltbushes because it is more salt
Utricles mature, in general about mid-October. tolerant (McArthur and others 1978). In large
Shadscale seeds are largely dormant after stands, shadscale should be managed rather than
maturation but respond to cold treatment (chilling) mechanically controlled. Managing for shadscale
through dry after-ripening (Meyer and others is easier than reducing shrub densities and seeding
1998). Chilling and leaching soluble inhibitors in new species because limited soil moisture and
from seeds has been shown to increase precipitation make it difficult to manipulate these
germination (Garvin and others 1996). plant communities (Bleak and others 1965).
Interestingly, seeds from warm desert populations Aggressive annuals, such as cheatgrass, usually
responded to chilling more and after-ripened invade such areas if they are disturbed.
quicker than those from cold deserts (Meyer and
others 1998). Ecotypic germination variation is Benefits.—Shadscale is a genetically and
common in shadscale. morphologically diverse native shrub that occupies
Shadscale spreads naturally through seed a considerable range throughout the West. It
but is difficult to establish by artificial seeding provides soil cover, habitat for birds, and forage
(McArthur and others 1978, Monsen and for domestic livestock and wildlife. Some small
Richardson 1984). Low seed fill often compounds passerine birds, such as horned lark (Eremophila
seed dormancy and germination problems. Fall alpestris), Brewer’s sparrow (Spizella breweri),
seeding is recommended because it reduces the and sage thrashers (Oreoscoptes montanus) nest in
effects of dormancy, and utricles are more likely to shadscale stands (Medin 1990). Healthy stands
break down before spring. Seeds should remain also reduce the proliferation of exotic, undesirable
between 6 and 13 mm of the soil surface to annuals. In addition, seeds were historically
improve the likelihood of germination and ground into flour by Native-Americans for bread
establishment. Soil type and seedbed preparations (Stubbendieck and others 1997).
should be considered prior to seeding. Seed should
be planted at rates of 2.2 to 4.5 kg of pure live References
seed/ha and in separate rows from other seeded
species. Seedlings grow slowly and compete Blaisdell, J.P. and RC. Holmgren. 1984. Managing
poorly with herbs and aggressive grasses, Intermountain rangelands-salt-desert shrub
particularly within the first 2 years. Diploid races ranges. General Technical Report INT-163. U.S.
from pinyon-juniper woodlands and big sagebrush Department of Agriculture, Forest Service,
communities establish more readily in biologically Intermountain Forest and Range Experiment
diverse communities than other races. Station, Ogden, UT. 52 p.
Shadscale has also been successfully
transplanted when moisture conditions are right Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
during spring or fall (Luke and Monsen 1984). Stevens, and B.C. Giunta. 1976. Characteristics
Transplants can be from container or bare-root and hybridization of important Intermountain
stock. Survival is usually poor unless care is taken shrubs. II. Chenopod family. Research Paper
during handling and planting. Young shrubs are INT-177. U.S. Deptment of Agriculture, Forest
particularly susceptible to overwatering. Service, Intermountain Forest and Range
Transplanting is most successful if plants are Experiment Station, Ogden, UT. 42 p.
dormant at the time of planting.
Bleak, A.T., N.C. Frischknecht, A.P. Plummer,
Management.—Despite its spiny nature, and R.E. Eckert, Jr. 1965. Problems in artificial
shadscale is grazed by livestock and game. Forage and natural revegetation of the arid shadscale
value is directly related to leaf persistence, which vegetation zone of Utah and Nevada. Journal of
is a characteristic that varies by habitat. Many Range Management. 18: 59-65.
species prefer the nutritious seeds (USDA 1937).
Livestock and wildlife tend to seek seed and leaf Caldwell, M.M., R.S. White, R.T. Moore, and L.B.
accumulations under parent shrubs after leaves and Camp. 1977. Carbon balance, productivity, and
97
water use by cold desert shrub communities Monsen, S.B. and B.Z. Richardson. 1984, Seeding
dominated by C3 and C4 species. Oecologia. 29: shrubs with herbs on a semiarid mine site with
275-300. and without topsoil. In: A.R. Tiedemann, E.D.
McArthur, H.C. Stutz, R. Stevens, and K.L.
Freeman, D. C. and E.D. McArthur. 1984. The Johnson, comps. Proceedings: Symposium on
relative influences of mortality nonflowering, the biology of Atriplex and related chenopods,
and sex change on the sex ratios of six Atriplex 1983 May 2-6; Provo, UT. Gen. Tech. Rep.
species. Botanical Gazette 145: 385-394. INT-172. U.S. Department of Agriculture,
Forest Service, Intermountain Forest and Range
Garvin, S.C., S. E. Meyer, and S. L. Carlson. 1996. Experiment Station, Ogden, UT. p. 298-305.
Seed germination studies in Atriplex confertifolia
(Torr. & Frem.) Wats. In: J.R. Barrow, E.D. Sanderson, S.C., H.C. Stutz, and E.D. McArthur
McArthur, R.E. Sosebee, and R.J. Tausch, 1990. Geographic differentiation in Atriplex
comps. Proceedings: shrubland ecosystem confertifolia. American Journal of Botany 77:
dynamics in a changing environment. 1995 May 490-498.
23-25; Las Cruces, NM. General Technical
Report INT-GTR-338. U.S. Department of Stubbendieck, J., S.L Hatch, and C.H. Butterfield
Agriculture, Forest Service, Intermountain 1997. North American Range Plants. Fifth
Research Station, Ogden, UT. p. 165-169. Edition. University of Nebraska Press, Lincoln,
NB. 305 p.
Luke, F. and S.B. Monsen. 1984. Methods and
costs for establishing shrubs on mined lands in Stutz, H.C. and S.C. Sanderson. 1983.
southwestern Wyoming. In: A.R. Tiedemann, Evolutionary studies of Atriplex: chromosome
E.D. McArthur, H.C. Stutz, R. Stevens, and K.L. races of A. confertifolia (shadscale). American
Johnson, comps. Proceedings-symposium on the Journal of Botany 70: 1536-1547.
biology of Atriplex and related chenopods, 1983
May 2-6; Provo, UT. Gen. Tech. Rep. INT-172. U.S. Department of Agriculture, Forest Service.
U.S. Department of Agriculture, Forest Service, 1937. Range Plant Handbook. U.S. Government
Intermountain Forest and Range Experiment Printing Office, Washington, DC. 816 p.
Station, Ogden, UT. p. 286-292.
Wallace, A. and D.L. Nelson. 1990. Wildland
McArthur, E.D., A.P. Plummer, and J.N. Davis. shrub dieoffs following excessively wet periods:
1978. Rehabilitation of game range in the salt a synthesis. In: E.D. McArthur, E.M. Romney,
desert. In: K.L Johnson, ed. Wyoming S.D. Smith, and P.T. Tueller, comps.
shrublands: proceedings 7th Wyoming shrub Proceedings: Symposium on cheatgrass
ecology workshop; 1978 May 31-June 1; Rock invasion, shrub die-off, and other aspects of
Springs, WY. University of Wyoming, Division shrub biology and management; 1989 April 5-7;
of Range Management, Laramie, WY. p. 23-50. Las Vegas NV. General Techical Report INT-
276. Ogden, UT. U.S. Department of
Medin, D.E. 1990. Birds of a shadscale (Atriplex Agriculture, Forest Service, Intermountain
confertifolia) habitat in east central Nevada. Research Station. p. 84-90.
Great Basin Naturalist 50: 295-298.
West, N.E. and J. Gasto. 1978. Phenology of the
Meyer, S.E., S.L. Carlson, and S.C. Garvin. 1998. aerial portions of shadscale and winterfat in
Seed germination regulation and field seed bank Curlew Valley, Utah. Journal of Range
carryover in shadscale (Atriplex confertifolia: Management. 31: 43-45.
Chenopodiaceae). Journal of Arid Environments
38: 255-267.
98
Atriplex corrugata S. Wats. mat saltbush
CHENOPODIACEAE
99
with a drag or harrow. Seeds should be planted References
near the soil surface and no more than 1.3 cm
deep. Seedlings emerge rapidly and grow Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
vigorously during the first year. Growth continues Stevens, and B.C. Giunta. 1976. Characteristics
until soil moisture becomes limiting (usually by and hybridization of important Intermountain
mid-summer). There are about 220,000 cleaned shrubs. II. Chenopod family. Research Paper
seed per kg (Blauer and others 1976). INT-177. U.S. Department of Agriculture, Forest
Transplants from bare root stock, Service, Intermountain Forest and Range
wildings, or containers have all established Experiment Station, Ogden, UT. 42 p.
successfully. Wildings and bare root stock tolerate
considerable distress. Transplants survive Hall, H.M. and F.E. Clements. 1923. The
surprisingly well, even when planted on seemingly phylogenetic method in taxonomy, the North
infertile soils. Young plants have been American species of Artemisia, Chrysothamnus,
successfully transplanted into former black and Atriplex. Publication 326. Carnegie
greasewood communities in the Great Basin Institution of Washington, Washington, DC. 355
(Blauer and others 1976). p.
Management.—Mat saltbush is valuable winter Hanson, C.A. 1962. Perennial Atriplex of Utah and
forage for wildlife and domestic livestock. It often the northern deserts. M.S. thesis. Brigham
grows interspersed among other plant communities Young University, Provo, UT. 133 p.
on shale outcrops at medium elevations. Shrubs
often grow on windblown slopes and provide McArthur, E.D., A.P. Plummer, and J.N. Davis.
important, accessible winter forage. Big game 1978. Rehabilitation of game range in the salt
have been known to congregate on these areas desert. In: K.L. Johnson, ed. Wyoming
during periods of deep snow accumulation. Shrubs shrublands: proceedings of the 7th Wyoming
tolerate grazing and trampling and maintain vigor shrub ecology workshop; 1978 May 31-June1;
even after serious abuse. Rock Springs, WY. University of Wyoming,
Mat saltbush has been seeded with Division of Range Management, Laramie, WY.
selected grasses including Russian wildrye p. 23-50.
(Elymus junceus Fisch.), tall wheatgrass
[Agropyron elongatum (Host) P.Beauv.], and Sanderson, S.C., and H.C. Stutz. 2003.
crested wheatgrass [Agropyron cristatum (L.) Unpublished data. U.S. Department of
Gaertn.]. Although these species may increase Agriculture, Forest Service, Rocky Mountain
forage production, they do not usually persist in Research Station, Provo, UT
dense stands. Grasses and forbs will suppress
shrub establishment, but grasses interseeded into Stutz, H.C. 1978. Explosive evolution of perennial
established shrub stands do not reduce shrub Atriplex in western America. Great Basin
density or vigor. Naturalist Memoirs. 2: 161-168.
This species is useful in revegetation and
restoration projects and can help reclaim disturbed Stutz, H.C. 1984. Atriplex hybridization in western
areas with heavy textured soils such as road North America. In: A.R. Tiedemann, E.D.
construction sites or mine disturbances. McArthur, H.C. Stutz, R. Stevens, and K.L.
Johnson, comps. Proceedings: Symposium on
Benefits.—Mat saltbush is a native shrub that the biology of Atriplex and related chenopods,
occupies a specialized niche in Western 1983 May 2-6; Provo, UT. General. Technical
ecosystems. Its ability to grow on fine textured, Report INT-172. U.S. Department of
often saline soils helps reduce soil erosion on sites Agriculture, Forest Service, Intermountain Forest
that are inhospitable to many shrubs. It also and Range Experiment Station, Ogden, UT.
provides important forage for wildlife during p. 25-27.
winter when other sources of forage may be
inaccessible. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Higgins, eds. 1993. A Utah flora. 2nd Ed.,
revised. Brigham Young University Print
Services, Provo, UT. 986 p.
100
Atriplex cuneata A. Nels. Castle Valley clover saltbush
CHENOPODIACEAE
101
about 7 weeks after blooming (Hanson 1962). nutritious forage for wildlife and livestock on
Annual seed production is often erratic and only Western shrublands.
about 50 percent of utricles contain viable seed.
Germination rates are also erratic and vary among References
ecotypes. Fruits are small and generally number
about 180,000/kg (Blauer and others 1976). Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
This species should not be seeded in areas Stevens, and B.C. Giunta. 1976. Characteristics
beyond its natural distribution. It is poorly adapted and hybridization of important Intermountain
to big sagebrush (Artemisia tridentata Nutt.) and shrubs. II. Chenopod family. Research Paper
other upland habitats but can be seeded into most INT-177. U.S. Department of Agriculture, Forest
salt desert shrublands. Young plants grow Service, Intermountain Forest and Range
vigorously and are competitive but can be Experiment Station, Ogden, UT. 42 p.
suppressed by perennial grasses and annual weeds.
It should not be seeded directly with herbaceous Freeman, D.C. and E.D. McArthur. 1984. The
species but competes well when seeded with relative influences of mortality, non-flowering,
fourwing saltbush [Atriplex canescens (Pursh) and sex change on the sex ratios of six Atriplex
Nutt.], green ephedra (Ephedra viridis Coville), species. Botanical Gazette 145: 385-394.
and spiny hopsage [Grayia spinosa (Hook.) Moq.].
Hall, H.M. and F.E. Clements. 1923. The
Management.—Castle Valley clover saltbush is phylogenetic method in taxonomy, the North
preferred by livestock throughout the year. It American species of Artemisia, Chrysothamnus,
remains green and succulent, even through winter. and Atriplex. Publication 326. Carnegie
It is an important source of protein in winter Institution of Washington, Washington, DC.
months for both livestock and game when most 355 p.
other forages are dry (McArthur and others 1978).
A stabilized hybrid between fourwing saltbush and Hanson, C.A. 1962. Perennial Atriplex of Utah and
Castle Valley clover saltbush has great potential to the northern deserts. M.S. thesis. Brigham
improve the forage quality on winter ranges. Such Young University, Provo, UT. 133 p.
a hybrid would be taller than Castle Valley clover
saltbush and retain more green foliage than McArthur, E.D., A.P. Plummer, and J.N. Davis.
fourwing saltbush. 1978. Rehabilitation of game range in the salt
This species shows particular promise in desert. In: K.L. Johnson, ed. Wyoming
winter game range restoration and on disturbed shrublands: proceedings of the 7th Wyoming
alkaline soils. Like other woody saltbushes, it shrub ecology workshop; 1978 May 31-June1;
grows on freshly disturbed soils, such as exposed Rock Springs, WY. University of Wyoming,
substrata created by mining or road construction Division of Range Management, Laramie, WY.
activities (Blauer and others 1976, McArthur and p. 23-50.
others 1978).
Sanderson, S.C. and H.C. Stutz. 2003.
Benefits.—Castle valley clover saltbush is a native Unpublished data. U.S. Department of
shrub that is adapted to fine textured soils that are Agriculture, Forest Service, Rocky Mountain
inhospitable to many other shrubs. It adds Research Station, Provo, UT
biological and structural diversity to the plant
communities on these otherwise difficult sites. It Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
helps stabilize the soil, can be useful in wildland Higgins, eds. 1993. A Utah Flora. Second
restoration projects, and provides valuable and edition, revised. Brigham Young University
Print Services, Provo, UT. 986 p.
102
Baccharis pilularis DC. coyote brush
ASTERACEAE
Synonyms: None
Christopher Ross
103
ground cover is effective in controlling height Latto, J. and C.J. Briggs. 1995. Factors affecting
without long-term detrimental effects (Hodel and distribution of the gall forming midge
Pittenger 1994). Coyote brush may be removed by Rhopalomyia californica (Diptera:
burning and then pulling out the roots. Cecidomyiidae). Environmental Entomology
24(3): 679-686.
Benefits.—Although several cultivated varieties
are often sold as a landscaping shrub or ground McBride, J. 1974. Plant Succession in the
cover, some growers caution that it is neither an Berkeley Hills, California. Madrono 22: 317-
especially attractive plant, nor as fire-proof as 380.
sometimes stated. It is of low value as livestock
forage (McBride and Heady 1968). Coyote brush McBride, J. and H.F. Heady. 1968. Invasion of
was used by Native Americans as a tea for use on Grassland by Baccharis pilularis DC. Journal
poison oak rash (Timbrook 1987). It is considered of Range Management 21: 106-108.
problematic in recreation areas due to its
impenetrability and frequent association with
Pickart, A.J. and J. O. Sawyer. 1998. Ecology and
poison oak (Rhus diversiloba T. & G.) (McBride
Restoration of Northern California Coastal Dunes.
and Heady 1968).
California Native Plant Society. Sacramento, CA.
152 p.
References
Sundberg, S. 1993. Baccharis. In J.C. Hickman,
Da Silva, P.G. and J. W. Bartolome. 1984.
ed. The Jepson Manual: higher plants of
Interaction between a shrub, Baccharis pilularis
California. University of California Press,
subsp. consanguinea (Asteraceae) and an annual
grass, Bromus mollis (Poaceae), in coastal Berkeley and Los Angeles. 1,400 p.
California. Madrono 31: 93-101.
Timbrook, J. 1987. Virtuous herbs: plants in
Hodel, D.R. and D.R. Pittenger. 1994. Responses Chumash medicine. Journal of Ethnobiology
of eight groundcover species to renovation by Winter 1987: 171-180.
mowing. Journal of Environmental Horticulture
12: 4-7. Williams, K., R.J. Hobbs, and S.P. Hamburg.
1987. Invasion of an annual grassland in
Northern California by Baccharis pilularis ssp.
consanguinea. Oecologia 72:461-465.
104
Barleria prionitis L. porcupine flower
ACANTHACEAE
Synonyms: none
John K. Francis
105
years. Because it is spiny and generally ignored by Burkill, H.M. 1985. The useful plants of West
cattle, the species is considered a weed in much of Tropical Africa. Vol. 1. Royal Botanic Garden,
its range. Mowing, followed by spraying of the Kew, UK. 960 p.
sprouts with broadleaf herbicide, is a general
control measure for low shrubs and would probably Howard, R. A. 1989. Flora of the Lesser Antilles,
be effective against porcupine flower. Leeward and Windward Islands. Vol. 6. Arnold
Arboretum, Harvard University, Jamaica Plain,
Benefits.—Porcupine flower furnishes cover for MA. 658 p.
wildlife and protects the soil against erosion. It is
widely planted as an ornamental and cultivated in Liogier, H. A. 1997. Descriptive flora of Puerto
Asia as a hedge plant (Burkill 1985). Whole-plant Rico and adjacent islands. Vol. 5. Editorial de la
extracts of porcupine flower contain iridoid Universidad de Puerto Rico, San Juan, PR.
glycosides, barlerin, and verbascoside, which have 436 p.
shown potent activity against respiratory syncytial
virus in vitro and may account for the plant’s use in Neal, M.C. 1965. In gardens of Hawaii. Special
treating fever and several respiratory diseases in Publication 50. Bernice P. Bishop Museum
herbal medicine (Balick and others 1998). Extracts Press, Honolulu, HI. 924 p.
of the plant have also been shown to effectively
suppress the fungi Trichophyton mentagrophytes in Panwar, H.S., M.M. Nauriyal, and H.C. Joshi.
vitro (Panwar and others 1979). A mouthwash 1979. In vitro screening of certain indigenous
made from root tissue is used to relieve toothache plants for their antimycotic activity. Veterinary
and treat bleeding gums (Burkill 1985). The whole Research Bulletin 2(2): 164-167.
plant, leaves, and roots are used for a variety of
purposes in traditional Indian medicine. For Parrotta, J.A. 2001. Healing plants of Peninsular
example, the leaves are used to promote healing of India. CABI Publishing. Wellington, UK & New
wounds and to relieve joint pains and toothache York. 917 p.
(Parrotta 2001). Because of its antiseptic
properties, extracts of the plant are incorporated Prakruti. 2002. Suddh Bhangra (maka) oil. http://
into herbal cosmetics and hair products to promote www.prakrutiherbals.com/hairoil.htm 2 p.
skin and scalp health (Prakruti 2002, Probiotics
New Zealand 2002, Vaipani 2002). Probiotics New Zealand. 2002. Probiotics for
life—improving your health and quality of life
References through good bacteria. http://www.probiotics.
co.nz/prdcts_efml.asp?ProductID=5. 3 p.
Balick, M., P. Blanc, M. Morgan, J.L. Chen, R.
Cooper, M.R. Kernan, W. Nanakorn, N. Vaipani. 2002. Vaipani Herbal Ayurvedic
Parkinson, E.J. Rozhon, C.A. Stoddard, and Z.J. Partisthan—A herbal ayurvedic cosmetics
Yee. 1998. New iridoids from the medicinal company. http://www.vaipani.com/skincare/
plant Barleria prionitis with potent activity skin%20Care-Page3.html. 2 p.
against respiratory syncytial virus. Journal of
Natural Products 61(10): 1,295-1,297.
106
Batis maritima L. saltwort
BATACEAE
Synonyms: none
John K. Francis
107
America were carried by currents to establish the Indies. http://www.geocities.com/barbudaturf/
species in the Galapagos Islands (Trillmich 2002). hallophites%20in%20deep.htm. 3 p.
Growth and Management.—Saltwart plants Big Island Invasive Species Committee. 2002. Pest
established in experiments from cuttings and prevention/suppression: eradication of incipient
irrigated with fresh and brackish water grew 65 cm forest pest plants by the Big Island Species
and reached plant dry weights of 120 g in 13 Committee. http://www.hear.org/
weeks. Plants receiving saltier water grew much operationmicronia/2002biiscannual.pdf. 9 p.
less but still added height and weight with water
almost twice as salty as sea water. Tissue ash Florida Cooperative Extension Service. 2002.
concentrations did not differ significantly between Butterfly gardening in Florida. University of
treatments (Miyamoto and others 1996). Wild Florida, Gainesville, FL. http://edis.ifas.ufl.edu/
plants in stands in Puerto Rico appear to extend BODY_UW057. 24 p.
their stems about 1 m/year. Leaf succulence
increases as much as 1.5 times during the dry Howard R.A. 1988. Flora of the Lesser Antilles,
season to maintain a near static solute (chiefly Na+, Leeward and Windward Islands.
Cl-, and S04--) concentration (Lüttge and others Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain,
1989). Probably all planting for landscaping and MA: Arnold Arboretum, Harvard University.
environmental restoration is by rooted cuttings or 673 p.
uprooted wild plants. Except in Hawaii, where it is
being suppressed as an exotic plant (Big Island Liogier H.A. 1985. Descriptive flora of Puerto
Invasive Species Committee 2002), there is little Rico and adjacent islands, Spermatophyta. Vol.
reason to control stands of saltwort. It grows in 1. Editorial de la Universidad de Puerto Rico,
disturbed areas where few other plants can survive. Río Piedras, PR. 352 p.
Benefits.—The principal benefit of saltwort is that Liogier, H.A. 1990. Plantas medicinales de Puerto
it grows in, covers, and protects salty low-laying Rico y del Caribe. Iberoamericana de Ediciones,
areas where few other species will grow. It is used Inc., San Juan, PR. 566 p.
as an ornamental and ground cover on similar
sites. Salt tolerance of the species allows it to be Long, R.W. and O. Lakela. 1976. A flora of
irrigated with brackish and sea water, as well as Tropical Florida. Banyon Books, Miami, FL.
fresh water (Miyamoto and others 1996). Ashes of 962 p.
the plant were once used in the manufacture of
glass and soap. The leaves are sometimes eaten as Lüttge, U., M. Popp, E. Medina, W.J. Cram, M.
a salad (Neal 1965). However, it is toxic in large Diaz, H. Griffiths, H.S.J. Lee, C. Schäfer, J.A.
quantities (Austin 1998). Saltwort serves as the Smith, and K.H. Stimmel. 1989. Ecophysiology
larval and adult hosts for the great Southern white of xerophytic and halophytic vegetation of a
and Eastern pigmy blue butterflies (Florida coastal alluvial plain in Northern Venezuela.
Cooperative Extension Service 2002) and is an New Phytologist 111(2): 283-291.
important food source for the Galapagos marine
iguana (Amblyrhynchus cristatus) (Wrege 2002). Miyamoto, S., E.P. Glenn, and M.W. Olsen. 1996.
The species has applications in herbal medicine to Growth, water use and salt uptake of four
treat eczema, psoriasis, and other skin conditions, halophytes irrigated with highly saline water.
rheumatism, gout, blood and vein disorders Journal of Arid Environments 32(2): 141-159.
(Liogier 1990).
Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
References 50. Bernice P. Bishop Museum Press, Honolulu,
HI. 924 p.
Austin, D.F. 1998. Poisonous plants of southern
Florida. http://www.fau.edu/divdept/science/ Nelson, G. 1996. The shrubs and woody vines of
envsci/poison-pl.html. 9 p. Florida. Pineapple Press, Inc., Sarasota, FL.
391 p.
Barbuda Turf Company. 2002. Halophytes in
deep. Barbuda Turf Company, Codrington Pennings, S.C. and C.L. Richards. 1998. Effects of
Village, Barbuda, Antigua-Barbuda, West wrack burial in salt-stressed habitats: Batis
108
maritima in a southwest Atlantic salt marsh. scope.org/downloadpubs/scope45/ch01-1.3.1.
Ecography 21(6): 630-638. html. [not paged].
Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. Wiggins, I.L. and D.M. Porter. 1971. Flora of the
Montiel, eds. 2001. Flora de Nicaragua. Galápagos Islands. Stanford University Press,
Monographs of Systematic Botany Vol. 85, No. Stanford, CA. 998 p.
1. Missouri Botanical Garden Press. 943 p.
Wrege, P.H. 2002. Bee-eater research: Galapagos!
Trillmich, F. 2002. El Niño in the Galapagos Department of Ecology and Evolutionary
Islands: a natural experiment. International Biology, Cornell University, Ithaca, New York.
Council of Scientific Unions. http://www.icsu- http://www.eeb.cornell.edu/wrege/pages/iggires.
html. 4 p.
109
Bidens menziesii (Gray) Sherff ko'oko'olau
ASTERACEAE
110
Krauss, B. 1998. How to plant a native Hawaiian area, Hawaii. 1999 Hawaiian Conservation
garden. http://www.hawaii.gov/health/oeqc/ Conference, Honolulu, HI.
garden/index.html. [not paged].
Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
Sherry, K., J.M Castillo, and R.B Shaw. 1999. 1990. Manual of the Flowering Plants of
Effects of wildfire on vegetation and rare plants Hawaii. University of Hawaii Press, Honolulu.
in arid montane shrublands, Pohakaloa training 1,854 p.
111
Bixa orellana L. annatto
BIXACEAE
John K. Francis
112
Under good management, plants will fruit within 1 Barbosa e Silva, K.M., F.C.G. Almeida, F.A.G.
year of planting (Nepstad and others 1991). An Almeida, P.S. Lima e Silva, J.J.L. de
Indian plantation yielded 529 kg/ha of seed at 2 Albuquerque, and J.J.L. de Albuquerque. 1993.
years old and 2,483 kg/ha of seed at 3 years old Effect of substrate on rooting of air-layered
(Kanjilal and Singh 1995). Annatto seldom shoots of annatto. Pesquisa Agropecuaria
reaches more than 5 m in height and 10 cm in stem Brasileira 28(1): 101-106.
diameter (Little and others 1974). Pruning of
ornamentals is recommended to shape and thicken Belfort, A.J.L., O.R. Kato, and M. do S.A. Kato.
the crowns (Warren 1997). 1992. Practical method of drying annatto seeds
for seedling production. Circular Tecnica 67.
Benefits.—Annatto, obtained from the oily arils of Centro de Pesquisa Agropecuaria do Tropico
the seeds is the world’s second most important Umido, EMBRAPA-CPATU, Belem, Brazil.
(after caramel) natural colorant (Mercadante and 14 p.
Pfander 1998), yielding yellow to red colors. The
colors are produced by several apocarotenoides Bruckner, C.H., S.S. Khouri, and A.V. Melgaco.
and may reach up to 7 percent of the seed’s dry 1991. Propagation of annatto (Bixa orellana L.)
mass (Katzer 1999). World production of annatto by five grafting methods. Revista Ceres 38: 218,
seed, both for commercial and home use, was 340-344.
estimated in 1990 at 10,000 tons per year (Arkcoll
1990). Brazil is the world’s largest exporter Howard, R.A. 1989. Flora of the Lesser Antilles.
(Katzer 1999). Not only was the dye used Vol. 6. Arnold Arboretum, Harvard University,
anciently to color food, but also to dye cloth and Jamaica Plain, MA. 658 p.
paint the skin (which is still done today). The
species is also planted as an ornamental, Kanjilal, P.B. and R.S. Singh. 1995. Agronomic
particularly the varieties with bright pink pods. evaluation of annatto (Bixa orellana L.). Journal
Branches with the dry pods are used in dry floral of Herbs, Spices, and Medicinal Plants 3(3): 13-
arrangements (Warren 1997). Bees collect nectar 17.
from its flowers to make honey. The wood is light-
weight (specific gravity 0.4), weak, and not Katzer, G. 1999. Annatto (Bixa orellana L.).
durable. It was used in former times to start fires http://www-ang.kfunigraz.ac.at/~katzer/engl/
by friction. Ropes and twine were made from the Bixa_ore.html. 3 p.
fibrous bark (Little and others 1974). The pulp
surrounding the seeds is widely used in herbal Liogier, H.A. 1990. Plantas medicinales de Puerto
medicine to treat burns, bleeding, dysentery, Rico y del Caribe. Iberoamericana de Ediciones,
gonorrhea, constipation, and fever (Parrotta 2001). Inc., San Juan, PR. 563 p.
Extracts of leaves, bark, and roots are reported to
be antidotes for poisoning from Manihot esculenta Liogier, H.A. 1995. Descriptive flora of Puerto
Crantz, Jatropha curcas L., and Hura crepitans L. Rico and adjacent islands. Vol. 4. Editorial
(Liogier 1990). de la Universidad de Puerto Rico, San Juan, PR.
617 p.
References
Little, E.L., Jr., R.O. Woodbury, and F.H.
Amaral, L.I.V., M. de F.A. Pereira, and A.L. Wadsworth. 1974. Trees of Puerto Rico and the
Cortelazzo. 1995. Dormancy breaking in seeds Virgin Islands. Vol. 2. Agriculture Handbook
of Bixa orellana. Revista Brasiliera de 449. U.S. Department of Agriculture,
Fisiologia Vegetal 7(2): 151-157. Washington, DC. 1,024 p.
113
Nepstad, D.C., C. Uhl, and E.A.S. Serrao. 1991. on rooting of Bixa orellana L. stem cuttings.
Recuperation of a degraded Amazonian Orissa Journal of Horticulture 25(1): 84-86.
landscape: forest recovery and agricultural
restoration. Ambio 20(6): 248-255. von Carlowitz, P.G. 1991. Multipurpose trees and
shrubs, sources of seeds and inoculants.
Parrotta, J.A. 2001. Healing plants of Peninsular International Council for Research in
India. CAB International, Wallingford, UK. Agroforestry, Nairobi, Kenya. 328 p.
944 p.
Warren, W. 1997. Tropical plants for home and
Thirunavoukkarasu, M. and H.O. Saxena. 1997. A garden. Thames and Hudson, New York. 240 p.
short note on the effect of auxins (IAA, IBA)
114
Bocconia frutescens L. pan cimarrón
PAPAVERACEAE
John K. Francis
115
Reproduction.—Pan cimarrón flowers and fruits de Publicaciones. San Salvador, El Salvador.
throughout the year (Little and others 1974, 703 p.
Stevens and others 2001). The infrutescences may
contain tens to hundreds of fruits. Fruits in the Howard R.A. 1988. Flora of the Lesser Antilles,
infrutescences mature a few at a time beginning at Leeward and Windward Islands.
the top and proceeding downward. A collection of Dicotyledoneae, Part 1. Vol. 4. Arnold
seeds from Puerto Rico weighed (air-dried) an Arboretum, Harvard University, Jamaica Plain,
average of 0.0146 + 0.0002 g/seed (author’s MA. 673 p.
observation). Germination begins in 25 to 55 days
with 40 percent germinating. Germination is Jardín Botánico de Bogotá José Mutis. 2002.
epigeal (Ricardi and others 1977). The seeds are Conservación ex situ. http://www.jbb.gov.co/
dispersed by birds (Environmental Protection conservacion/ex_situ/semillas/body_semillas.
Agency 2002). Damaged plants usually resprout. html. 4 p.
Growth and Management.—Once established, Liogier H.A. 1985. Descriptive flora of Puerto
pan cimarrón has relatively rapid growth (Jardín Rico and adjacent islands, Spermatophyta. Vol.
Botánico de Bogotá José Mutis 2002). Only in 1. Editorial de la Universidad de Puerto Rico,
Hawaii is the species mentioned as a nuisance Río Piedras, PR. 352 p.
where it displaces native vegetation and threatens
the endangered Blackburn’s sphinx moth Little, E.L., Jr., R.O. Woodbury, and F.H.
(Environmental Protection Agency 2002). In the Wadsworth. 1974. Trees of Puerto Rico and the
absence of tests of control measures, such standard Virgin Islands. Vol. 2. Agriculture Handbook
practices as grubbing and spraying with broadleaf 449. U.S. Department of Agriculture,
herbicides are recommended. Washingtion, DC. 1,024 p.
116
Bontia daphnoides L. white alling
MYOPORACEAE
John K. Francis
shelled stone (Howard 1989, Liogier 1997, Little
and others 1974).
117
Growth and Management.—A small group of Liogier, H.A. 1997. Descriptive flora of Puerto
white alling seedlings grown in the nursery in Rico and adjacent islands. Vol. 5. Editorial de la
Puerto Rico ranged in height from 21 to 86 cm, 8 Universidad de Puerto Rico, San Juan, PR.
months after being pricked into containers. Sapling 436 p.
and sprout growth rate is moderate (about 0.5
m/year) and individual plants live 10 to 30 years. Little, E.L., Jr., R.O. Woodbury, and F.H.
Although no management experience has been Wadsworth. 1974. Trees of Puerto Rico and the
published, the species can probably be established Virgin Islands. Vol. 2. Agriculture Handbook
with nursery seedlings and be managed like other 449. U.S. Department of Agriculture,
short-statured species. Washington, DC. 1,024 p.
Benefits.—White alling helps protect the soil and Mahabir, D. and M.C. Gullifor. 1997. Use of
provides food and cover for wildlife. The wood is medicinal plants for diabetes in Trinidad and
useful for fuel and stakes. Ethanol extracts of white Tobago. Revista Panamericana de Salud Pública
alling showed promising insecticidal activity 1(3): 174-179.
against Boophilus microplus, Cylas formicarius,
and Tribolium confusum (Mansingh and Williams Mansingh, A. and L.A.D. Williams. 1998.
1998, Williams and Caleb-Williams 1997, Pesticidal potential of tropical plants II.
Williams and Mansingh 1993). White alling is Acaricidal activity of crude extracts of several
frequently used to control diabetes in Trinidad Jamaican plants. Insect Science and its
(Mahabir and Gullifor 1997). Extracts of the plant Application. 18(2): 149-155.
are used to control intestinal worms, treat herpes,
treat inflammation, insect bites, scarring, ulcers, Nelson, G. 1996. The shrubs and woody vines of
and wounds (Liogier 1990). Florida. Pineapple Press, Inc., Sarasota, FL.
391 p.
References
Sánchez deL.C., J. 2001. Plantas de la flora cubana
Hawaiian Ecosystems at Risk 2001. Mystery plant cultivadas en España. http://floraguide.es/
# 2, Bontia daphnoides Linn. (Myoporaceae). arboles/plantasdecuba.htm. 8 p.
http://hear.org/mysteryplants/autogendhtml/
mysteryplant2.htm. 3 p. Williams, L.A.D. and L. Caleb-Williams. 1997.
Insecticidally active sesquiterpene furan from
Howard, R. A. 1989. Flora of the Lesser Antilles, Bontia daphnoides L. Philippine Journal of
Leeward and Windward Islands. Vol. 6. Arnold Science 126(2): 155-162.
Arboretum, Harvard University, Jamaica Plain,
MA. 658 p. Williams, L.A.D. and A. Mansingh. 1993.
Pesticidal potential of tropical plants I.
Liogier, H.A. 1990. Plantas medicinales de Puerto Insecticidal activity in leaf extracts of sixty
Rico y del Caribe. Iberoamericana de Ediciones, plants. Insect Science and its Application. 14(5):
Inc., San Juan, PR. 566 p. 697-700.
118
Borrichia arborescens (L.) DC. sea ox-eye
ASTERACEAE
John K. Francis
119
irrigate sea ox-eye plants used in landscaping Arboretum, Harvard University, Jamaica Plain,
(Gillman 1999). MA. 658 p.
Benefits.—Sea ox-eye is available commercially Knox, G.W. 2002. Drought tolerant plants for
and is widely planted as an ornamental. It is North and Central Florida. Pinellas County
recommended for foundation plantings and as a Extension. http://coop.co.pinellas.fl.us/fyn/
border shrub (Workman 1980), particularly where publications/fyn_drought_tolerant.html. 12 p.
salt and salt spray are prevalent and in xeric
(unwatered) gardens (Knox 2002). It also attracts Liogier, H.A. 1997. Descriptive flora of Puerto
butterflies (Gilman 2002). A tea is prepared from Rico and adjacent islands. Vol. 5. Editorial de la
branch tips and leaves to relieve colds, coughs, and Universidad de Puerto Rico, San Juan, PR.
fish poisoning (Garland 2002). 436 p.
120
Bourreria virgata (Sw.) G. Don roble de guayo
BORAGINACEAE
John K. Francis
outer surface deeply grooved (author’s
observations, Liogier 1995, Little and others
1974).
121
guayo lives for several decades. No management Puerto Rican trees and shrubs: second
experience has been published. The species is not installment. Research Note SO-374. U.S.
weedy in agricultural land or forest plantations and Department of Agriculture, Forest Service,
will rarely need control. Southern Forest Experiment Station, New
Orleans, LA. 5 p.
Benefits.—Roble de guayo helps protect the soil
and furnishes food and cover for wildlife. The Liogier, H.A. 1995. Descriptive flora of Puerto
stems are too small for use as lumber but should Rico and adjacent islands. Vol. 4. Editorial de
make excellent fuel. The species would probably la Universidad de Puerto Rico, San Juan, PR.
be a good ornamental for landscaping xeric 617 p.
gardens.
Little, E.L., Jr., R.O. Woodbury, and F.H.
References Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
Breckon, G.J. and R.G. García. 2001. Vascular 449. U.S. Department of Agriculture,
plants of Susúa Forest. University of Puerto Washington, DC. 1,024 p.
Rico, Mayaguez, PR. http://www.uprm.edu/
biology/profs/breckon/herbarium/florasusua. World Wildlife Fund. 2001. Cuban cactus scrub
htm. 52 p. (NT1306). http://www.worldwildlife.org/
wildworld/profiles/terrestrial/nt/nt1306_full.
Francis, J.K. and A. Rodríguez. 1993. Seeds of html. 8p.
122
Brickellia californica (Torr. & Gray) Gray California brickellbush
ASTERACEAE
John K. Francis
123
coastal and island vegetation types in California References
(Munz 1974) and therefore at low elevations.
Disturbance is probably required for Abrams, L., and R.S. Ferris. 1960. Illustrated flora
establishment. of the Pacific States. Stanford University Press,
Stanford, CA. 732 p.
Reproduction.—California brickellbush flowers
August through October (Tarweed Native Plants Dittman, L. 2003. Asteraceae: aster, daisy or
2003). The species flowers abundantly and composite family. http://www.nazflora.org/
potentially produces vast numbers of seeds. Air- asteraceae.htm. [not paged].
dried seeds collected in Utah averaged 3.525
million/kg. Hand-sorted seeds placed without any Harrington, H.D. 1964. Manual of the plants of
pretreatment on moist blotter paper germinated at Colorado. Sage Books, Denver, CO. 666 p.
71 percent beginning between 5 and 14 days after
sowing. Germination is epigeal (author’s Las Pilitas Nursery. 2003. Brickellia californica.
observation). Seed will have to be harvested by http://www.laspilitas.com/plants/123.htm. 3 p.
hand because wild plants are scattered. Stems layer
(root) when covered by soil or colluvial material. Moerman, D.E. 1986. Medicinal plants of Native
The seeds are wind-dispersed. Seedlings probably America. Technical Reports 19. University of
appear and survive only when conditions are Michigan Museum of Anthropology, Ann
nearly ideal. Arbor, MI. 534 p.
Growth and Management.—Save for a few Munz, P.A. 1974. A flora of Southern California.
centimeters of caudex, California brickellbush University of California Press, Berkeley, CA.
adds and loses its entire height (0.5 to 1.0 m) each 1,086 p.
growing season. The aerial portion of the caudex
also dies back about every 3 to 5 years while Natural Resources Conservation Service. 2003.
others are produced each year. Plant may live for Plants profile: Brickellia Ell., brickellbush.
one to several decades, renewing themselves with http://plants.usda.gov/cgi_bin/plant_profile.cgi?
suckers. The species is rarely abundant and symbol=BRICK. [not paged].
probably does not require control. Management
experience has not been published. Oklahoma Biological Survey. 2003. Brikellia
californica (Torr. & Gray) Gray.
Benefits.—California brickellbush helps protect http://www.biosurvey.ou.edu/shrub/brca3.htm.
the soil and furnishes food and cover for wildlife. 3 p.
It has been reported to be a larval food plant for
the tiger moth, Dysschema howardi Hy. Edwards Tarweed Native Plants. 2003. Current plant
(Walsh 2003). The species has been planted as an availability list. http://www.tarweednativeplants.
ornamental, particularly for the floral fragrance, in com/botanical.html. 5 p.
natural landscaping (Oklahoma Biological Survey
2003). Native Americans used infusions of the Walsh, B. 2003. Moths of southeastern Arizona:
leaves as a ceremonial emetic, as a lotion to treat Arctiidae (tiger moths). http://nitro.biosci.
infant skin sores, and for coughs and fevers Arizona.du/zeeb/butterflies/artic.html. 8 p.
(Moerman 1986).
124
Bromelia pinguin L. maya
BROMELIACEAE
Synonyms: none
John K. Francis
125
plants averaged 81.6 percent water and ranged References
from 0.3 kg to 1.6 kg dry weight. The maya
portion of total stand dry biomass would run from Crane, J.C. 1945. Living fence posts in Cuba.
about 500 to 2,000 kg/ha. For the most part, maya Agriculture in America 5(2): 34-35, 38.
is undesirable in forests and pastures because it
takes up valuable space, restricts access, inhibits Guzman, D.J. 1975. Especies útiles de la flora
grazing, and provides nesting habitat and cover for Salvadoreña. Ministerio de Educación, Dirección
rats (Vélez and van Overbeek 1950). Maya stands de Publicaciones. San Salvador, El Salvador.
may be controlled by uprooting and piling the 703 p.
plants. If management objectives so dictate, maya
can be established by transplanting top-trimmed Howard, R.A. 1979. Flora of the Lesser Antilles,
wildlings. Leeward and Windward Islands. Vol. 3. Arnold
Arboretum, Harvard University, Jamaica Plain,
Benefits.—Historically, and to some extent MA. 586 p.
today, maya was used as a hedge or living fence
to inhibit entry into fields and homesteads (Crane Mesa, A.R. and G. Lajonchere. 1996.
1945). These hedges were never more than Micropropagation of Bromelia pinguin Lindl.
marginally effective—cattle and many species of Pastos y Forrajes 19(3): 217-223.
animals pass relatively easily and people can
ignore the spines or cut a path with a machete. Neal, M.C. 1965. In gardens of Hawaii. Special
The fruit, which has a white flesh, is bitingly acid, Publication 50. Bernice P. Bishop Museum
and tastes like pineapple, is eaten occasionally out Press, Honolulu, HI. 924 p.
of hand and used to make a refreshing drink (Neal
1965). The long, parallel leaf fibers were found to Vélez, I. and J. van Overbeek. 1950. Plantas
be of good quality and proposed for commercial indeseables en los cultivos tropicales. Editorial
production (Guzman 1975). Maya offers food and Universitario, Río Piedras, PR. 497 p.
cover for wildlife, such as small mammals and
birds.
126
Brunfelsia lactea Krug & Urban jasmín del monte
SOLANACEAE
Synonyms: none
John K. Francis
4-mm brown, elliptic seeds (Liogier 1995, Little
and others 1974).
127
Growth and Management.—Jasmín del monte http://www.tfts.org/Plant_Sale_May_2000.htm.
grows slowly in its natural habitat and appears to 12 p.
be long-lived. The author knows of no wildland
plantings or intentional management. Because the Kuipplantenkwekerij Gommer. 2003. Brunfelsia
species reproduces after disturbance, soil lactea. Kuipplantenkwekerij Gommer, Dalen,
unavoidably exposed during construction and road Netherlands. http://www.kwekerijgommer.com/
work should be left in good tilled condition to fotos/kuipplanten/Brunfelsia%20Lactea.htm.
encourage seedling establishment. 1 p.
Benefits.—Jasmín del monte helps hold the soil, Liogier, H.A. 1995. Descriptive flora of Puerto
provides cover and probably food for wildlife, and Rico and adjacent islands. Vol. 4. Editorial
adds to the aesthetics of the forest. The species is de la Universidad de Puerto Rico, Río Piedras,
planted as an ornamental in gardens and PR. 617 p.
greenhouses and is available for purchase from a
number of commercial nurseries. Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
References Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Gehrke, P. 2000. Sale plants 2000. Tropical Washington, DC. 1,024 p.
Flowering Tree Society, Miami, FL.
128
Caesalpinia bonduc (L.) Roxb. gray nicker bean
FABACEAE
John K. Francis
129
water for as long as 19 years (Markland 2002). The References
scarifying action of sand, weathering, insects, or
rodents eventually allows water to enter the seeds Ajiwe, V.I.E., C.A. Okeke, H.U. Agbo, G.A.
and they germinate and establish themselves in the Ogunleye, and S.C. Ekwuozor. 1996. Extraction,
new environments. A seed collected at Macquarie characterization and industrial uses of velvet-
Island, Antarctica, a drift of not less than 1 year, tamarind, physic-nut and nicker-nut seed oils.
was scarified and grown successfully (Costin Bioresource Technology 57(3): 297-299.
1965). The stems layer (root) whenever they come
in contact with the ground. Stems sprout when cut Ali, M.S., S. Shameel, V.U. Ahmad, and K.
and lateral roots sometimes sucker. Clonal plants Usmanghanim. 1997. Chemical constituents of
tend to remain attached to one another. Caesalpinia bonduc. Pakistan Journal of
Scientific and Industrial Research 40(1-4): 20-
Growth and Management.—Growth of gray 22.
nicker beans is fairly rapid at all stages. Seedlings
reached 26 cm in 40 days after sowing. Older Amarsinghe, A.P.G., R.D. Sharma, C. Chaturvedi,
plants grow a meter or more per year. Individual and D.K. Agarwal. 1993. Anthelmintic effect of
stems live at least 4 years. The author knows of no Ayurvedic recipe Kuberakshadi yoga in
published management experience. Plantations intestinal worms among children.Journal of
undoubtedly could be established with potted Research and Education in Indian Medicine
seedlings or by direct seeding scarified seeds. 12(1): 27-31.
Control could probably be obtained by lopping
with a machete followed by herbicide treatment of Biswas, T.K., S. Bandyopadhyay, Biswapati-
the sprouts. Mukherjee, Bhaswar-Mukherjee, B.R. Sangupta,
and B. Mukherjee. 1997. Oral hypoglycemic
Benefits.—Gray nicker bean contributes to the effect of Caesalpinia bonducella. International
biodiversity of the forests where it grows, helps Journal of Pharmacognosy 35(4): 261-264.
protect the soil, and furnishes cover for wildlife.
The seeds have been used for centuries and are still Burkill, H.M. 1995. The useful plants of West
used as jewelry, prayer beads, good luck charms, Tropical Africa. Vol. 3. Royal Botanic Gardens,
and worry stones (Rancho Leona 2002, Workman Kew, UK. 857 p.
1980). They were anciently used as standards of
weight in India (Vijayanagara Coins 2002). The Costin, A.B. 1965. Long-distance seed dispersal to
ancient African game of mancala traditionally Macquarie Island. Nature 206: 317.
employs gray nicker beans as game pieces
(Driedges 1972). The species is sometimes planted Driedges, W. 1972. The game of Boa, or Mankala,
as a hedge to prevent undesired entry into property in East Africa. Mila 3(1): 7-19.
(Nelson 1996) and could be planted for dune
stabilization. Preparations of the seeds and other Howard R.A. 1988. Flora of the Lesser Antilles,
plant parts are used to treat a large range of Leeward and Windward Islands.
ailments (Burkill 1995, Parrotta 2001). Dicotyledoneae, Part 1. Vol. 4. Arnold
Triterpenoids, fatty acid triglicerides, and sterols Arboretum, Harvard University, Jamaica Plain,
isolated from seeds may possibly explain some of MA. 673 p.
the activity (Ali and others 1997, Rostogi and
others 1996). Seed extracts have been shown to Liogier, H.A. 1988. Descriptive flora of Puerto
lower blood sugar in laboratory animals (Biswas Rico and adjacent islands, Spermatophyta. Vol.
and others 1997, Sharma and others 1997) and 2. Editorial de la Universidad de Puerto Rico,
effectively suppress or cure infections of several Río Piedras, PR. 481 p.
species of round worms (Amarsinghe and others
1993, Rastogi and others 1996). The fat content of Long, R.W. and O. Lakela. 1976. A flora of
gray nicker nut seeds was measured at 34 + 0.83 Tropical Florida. Banyon Books, Miami, FL.
percent. The semidrying oil extracted would be 962 p.
useful for manufacture of high-quality alkyd resins,
polishes, and paint (Ajiwe and others 1996). Markland, J. 2002. Drift seeds. In: West Word,
Community paper for Glenfinnan, Lochailort,
Glenuig, Arisaig, Morar, Mallaig, Knoydart, and
the Small Isles. March 2002. http://road-to-the-
130
isles.org.uk/westword/march2002.html. p. 6-7. antifilarial triglyceride fraction from Caesalpinia
bonduc. Fitoterapia 67(1): 63-64.
Nelson, G. 1996. The shrubs and woody vines of
Florida. Pineapple Press, Inc., Sarasota, FL. Sharma, S.R., S.K. Dwivedi, and D. Swarup. 1997.
391 p. hypoglycaemic, antihyperglycaemic and
hypolipidemic activities of Caesalpinia
Pacific Island Ecosystems at Rick. 2002. bonducella seeds in rats. Journal of
Caesalpinia bonduc (L.) Roxb., Fabaceae. Ethnopharmacology. 58(1): 39-44.
http://www.hear.org/pier3/cabon.htm. 2 p.
Vermillion, W.G. 2001. Caesalpinia bonduc
Parrotta, J.A. 2001. Healing plants of Pininsular (Fabaceae) new to Louisiana. Sida 19(4): 1,181-
India. CABI Publishing, Wellingford, UK and 1,182.
New York. 917 p.
Vijayanagara Coins. 2002. Coinage. http://www.
Rancho Leona. 2002. Rancho Leona jewelry vijayanagaracoins.com/htm/coinage.htm. 7 p.
samples: rainforest seed chart.
http://rancholeona.com/seed.html. 2 p. Workman, R.W. 1980. Growing native. The
Sanibal-Captiva Conservation Foundation, Inc.,
Rastogi, S., A.K Shaw, and D.K. Kulshreshtha. Sanibal, FL. 137 p
1996. Characterization of fatty acids of .
131
Cajanus cajan (L.) Millsp. pigeon pea
FABACEAE
John K. Francis
132
and forbs. Growth is moderately slow during the stems, the plant is damaged by browsing,
first 2 to 3 months of life during which time especially by cattle (Department of Primary
seedlings are not competitive with grass and Industries 2002). The species is planted as a green
weeds; afterwards pigeon pea competes well with manure crop, nurse crop, cover crop, a windbreak
vegetation equal or lower in height (van der hedge, as a host for lac insects, and as food for silk
Maesen 1990). In Puerto Rico, the species persists worms. The stalks are used for fuel, thatch, and
for a generation or two after cultivation and grows basketry (Center for New Crops and Plants
along roadsides and in waste places where seeds Products 2002). Pigeon pea forms root nodules in
have fallen, but the populations are not sustained association with Rhizobium sp. bacteria and is
indefinitely. Although seldom devastating, a large capable of fixing 41 to 280 kg/ha of nitrogen (Red
number of insects and diseases affect pigeon pea de Grupos de Agricultura de Cobertura 2002).
and can reduce yield or shorten a plant’s life Preparations of the leaves are used to treat
(American Phytopathological Society 2002). jaundice, inflammation, and sores of the mouth
(Parrotta 2001).
Reproduction.—Pigeon pea flowers nearly
throughout the year (Red de Grupos de Agricultura References
de Cobertura 2002). The flowers are self-
compatible and usually self-pollinated (Smartt American Phytopathological Society. 2002.
1976). Insects visit and pollinate the flowers (5 to Common names of plant diseases: Diseases of
40 percent cross pollination), but it is not known pigeonpea [Cajanus cajan (L.) Millsp.].
whether they increase seed yield (McGregor 2002). http://www.apsnet.org/online/common/names/
Upon drying, the pods spring and coil to release the pigeon.asp. 5 p.
seeds. Pigeon pea seeds weigh about 0.1g and
germinate in about 2 weeks (Center for New Crops Center for New Crops and Plants Products. 2002.
and Plants Products 2002). Cajanus cajan (L.) Millsp. Purdue University.
http://www.hort.purdue.edu/newcrop/duke_
Growth and Management.—Pigeon pea is energy/Cajanus_cajun.html. 6 p.
normally sown directly into prepared ground.
Seeding rates for pure stands are 12 to 25 kg of Department of Primary Industries. 2002. Legumes
seed/ha (Smartt 1976). Seeding depths of 2.5 to 5 for the tropics: pigeon pea (Cajanus cajan).
cm are recommended (Center for New Crops and Queensland Government. http://www.dpi.qld.
Plants Products 2002). No pregermination gov.au/pastures/4571.html. 2 p.
treatment of the seed is needed. Although some
varieties mature seed in 5 to 6 months, longer- Liogier, H.A. 1988. Descriptive flora of Puerto
lived, tall varieties including those that are more Rico and adjacent islands, Spermatophyta. Vol.
competitive in the wild take 10 to 12 months to 2. Editorial de la Universidad de Puerto Rico,
mature seed. These plants live about 5 years Río Piedras, PR. 481 p.
(Smartt 1976). Experimental yields of 50 dry
t/ha/year have been demonstrated; yields of 3 to 8 Long, R.W. and O. Lakela. 1976. A flora of
dry t/ha/year are obtained under normal Tropical Florida. Banyan Books, Miami, FL.
management (Van Den Beldt 1988). 962 p.
Benefits.—Pigeon peas are an important food in McGregor, S.E. 2002. Insect pollination of
developing tropical countries. An excellent source cultivated crop plants. Agriculture Research
of protein, the seeds (and sometimes the pods) are Service. http://gears.tucson.ars.ag.gov/book/
eaten as a vegetable, as a flour additive to other chap4/pig.html. [not paged].
foods, in soups, and with rice (Center for New
Crops and Plants Products 2002). Although they Nene, Y.L. and V.K. Sheila. 1990. Pigeonpea:
vary slightly, typical nutritional values for seeds geography and importance. In: Y.L. Nene, S.H.
are: moisture, 10.1 percent, protein 19.2 percent, Hall, and V.K. Sheila. The pigeonpea. CAB
fat, 1.5 percent, carbohydrates, 57.3 percent, fiber International, Wellingford, UK. p. 1-14.
8.1 percent, and ash, 3.8 percent (Smartt 1976).
About 3.4 million ha were under cultivation in Parrotta, J.A. 2001. Healing plants of Peninsular
1978-1988 period, 88 percent of it in India (Nene India. CABI Publishing, Wallingford, UK and
and Sheila 1990). Although pigeon pea makes New York. 917 p.
excellent forage, because of the brittleness of its
133
Red de Grupos de Agricultura de Cobertura. 2002. Smartt, J. 1976. Tropical pulses. Longman Group
Base de información sobre especies con Limited, London. 348 p.
potential de abonos verdes y cultivos de
cobertura. Rockefeller Foundation. http://www. Van Den Beldt, R.J. 1988. Cajanus cajan: it’s
rockfound.org.mx/cajanusbiesp.html. [not more than just a pulse crop. Nitrogen Fixing
paged]. Tree association. NFT Highlights 88-06. 5 p.
Skerman, P.J., D.G. Cameron, and F. Riveros. van der Maesen, L.J.G. 1990. Pigeonpea: origin,
1988. Tropical forage legumes. FAO Plant history, evolution, and taxonomy. In: Y.L.
Production and Protection Series 2. Food and Nene, S.H. Hill, and V.K. Sheila. The
Agriculture Organization of the United Nations. pigeonpea. CAB International. Wellingford,
692 p. UK. p. 15-46.
134
Callicarpa americana L. American beautyberry
VERBENACEAE (also placed in LAMIACEAE alt. Labiatae)
Synonyms: None
Kristina Connor
135
fall that are treated with indol-3-butyric acid (IBA; Grelen, H.E. and V.L. Duvall. 1966. Common
1000 ppm) and put in a mist bed (Dirr and Heuser plants of longleaf pine-bluestem range. Research
1987). Paper SO-23. U.S. Department of Agriculture
Forest Service, Southern Forest Experiment
Growth and Management.—American Station, New Orleans, LA. 96 p.
beautyberry is a short-lived shrub that is easy to
grow and widely distributed in the Southeastern Krüssmann, G. 1976. Manual of Cultivated Broad-
United States. It can be used as a landscape border Leaved Trees and Shrubs. Volume I, A-D.
and makes a good mass planting. However, regular Timber Press, Beaverton, OR. 448 p.
pruning is required if it is to maintain its form and
produce masses of the fruits so attractive in the fall Martin, C.O. and S.P. Mott. 1997. American
of the year. The fruits occur only on new growth, Beautyberry (Callicarpa americana). Section
and plants should be heavily pruned in early spring 7.5.8, U.S. Army Corps of Engineers Wildlife
(Martin and Mott 1997). It will not do well in Resources Management Manual. Ecosystem
areas that experience flooding but can be used to Management and Restoration Research Program
restore surface-mined sites (Martin and Sick Technical Report EL- 97.15. Conservation
1995). It will resprout after fire (Martin and Mott Communications, Vicksburg, MS. 17 p.
1997) and can be easily transplanted.
Martin, H. and G. Sick. 1995. American
Benefits.—American beautyberry is a good source beautyberry for borrow pit reclamation in South
of food for deer and the fruits attract birds. It can Carolina Restoration and Management Notes
be used in reclamation work and for erosion 13(1): 90-96.
control (Brown 1945). It easily reseeds in nature.
Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope.
References 1996. Plants for American Landscapes. Louisiana
State Univ. Press, Baton Rouge, LA. 266 p.
Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
A Concise Dictionary of Plants Cultivated in the Odenwald, N.G. and J.R. Turner. 1988. Plants for
United States and Canada. McMillan Publishing the South: A guide for landscaping design.
Co., Inc., New York. 1,290 p. Claitor’s Publication Division, Baton Rouge, LA.
565 p.
Bonner, F.T. (In press). Callicarpa americana L.
American beautyberry. In: Seeds of Woody Radford, A.E., H.E. Ahles, and C.R. Bell. 1968.
Plants in the United States. Revised edition. Manual of the Vascular Flora of the Carolinas.
University of North Carolina Press, Chapel Hill.
Brickell, C. and J.D.. Zuk. 1996. The American 1,183 p.
Horticultural Society A-Z Encyclopedia of
Garden Plants. DK Publishing, Inc., New York. Roberts, E.H. 1973. Predicting the storage life of
1,092 p. seeds. Seed Science and Technology 1: 499-514.
Brown, C.A. 1945. Louisiana trees and shrubs. USDA, NRCS. 2001. The PLANTS database.
Bulletin 1, Louisiana Forestry Commission, National Plant Data Center, Baton Rouge,
Baton Rouge, LA. 262 p. http://plants.usda.gov. LA [not paged].
Dirr, M.A. and C.W. Heuser, Jr. 1987. The USDA. [no date]. Southern wetland floral field
reference manual of woody plant propagation: office guide to plant species. U.S. Department
from seed to tissue culture. Varsity Press, Athens, of Agriculture Soil Conservation Service, South
GA. 239 p. National Technical Center, Fort Worth, TX.
http://www.npwrc.usgs.gov. [not paged]
.
136
Calotropis procera (Ait.) Ait. f. giant milkweed
ASCLEPIADACEAE
John K. Francis
137
about 100,000 seeds/kg (author’s observation). leaves are chopped and mixed with other feed,
Eighty-nine percent germination took place in consumption greatly increases with no ill effects.
potting mix between 7 and 64 days after sowing. (Abbas and others 1992, Nehra and others 1987).
Half the seed weight was found in the wing (silk). Shade-dried giant milkweed leaves contained 94.6
The seeds are dispersed by wind and may fly for percent dry matter, 20.9 percent ash, 19.6 percent
several hundred yards in gentle breezes. Seedlings crude protein, 2.2 percent fat, 43.6 percent acid
may arise in abundance after rainy periods, but detergent fiber, and 19.5 percent neutral detergent
only a few survive the first season. Using the fiber (Abbas and others 1992). Although it is
reserves in its large taproot, giant milkweed can lightweight, the wood is used in impoverished
resprout year after year when burned or cut. desert areas for a cooking fuel (Varshney and Bhoi
1988).
Growth and Management.—Giant milkweed
usually reaches heights of about 2 m, but may References
occasionally reach 5 m in height and stem
diameters of 25 cm (Little and others 1974). Abbas, B., A.E. El Tayeb, and Y.R. Sulleiman.
Growth is about 1 m the first year after sprouting 1992. Calotropis procera: feed potential for arid
in Puerto Rico. Senescence of individual stems zones. Veterinary Record 131(6): 132.
takes place in about 5 years, but plants often
resprout afterwards. Giant milkweed does not form Anver, S. and M.M. Alam. 1992. Effect of latex
dense stands, normally occurring as scattered seed dressing on interacting root-knot and
individuals. It can be a serious weed in pastures, reniform nematodes. Afro-Asian Journal of
overgrazed rangelands, and poorly managed hay Nematology 2: 1-2, 17-20.
fields. Although it probably is not possible to
eliminate existing stands through management Campolucci, P. and C. Paolini. 1990.
without ceasing grazing and harvest, it may be Desertification control in the Sahel regions—
possible to prevent their establishment. Some form low-cost large-scale afforestation techniques.
of chemical control would seem to be the only Note Tecniche 10. Centro di Sperimentazione
practical option for eliminating existing stands but Agricola e Forestale di Sperimentazione per la
no specific guidelines are available. Establishing Pioppicoltura. 24 p.
giant milkweed has been advocated for
environmental protection and as a nurse crop for Charu-Jain and P.C. Trivedi. 1997. Nematicidal
more valuable species (Campolucci and Paolini activity of certain plants against root-knot
1990). This can be done easily by planting nematode, Meloidogyne incognita, infecting
containerized seedlings or rooted cuttings. chickpea, Cicer arietinum. Annals of Plant
Protection Sciences 5(2): 171-174.
Benefits.—Giant milkweed tissues, especially the
root bark, are used to treat a variety of illness Howard, R.A. 1989. Flora of the Lesser Antilles,
including leprosy, fever, menorrhagia, malaria, Leeward and Windward Islands. Dicotyledoneae.
and snake bite (Parrotta 2001). The latex is toxic Part 3. Vol. 6. Arnold Arboretum, Harvard
and can cause blisters and rash in sensitive University, Jamaica Plain, MA. 658 p.
persons. The plant is occasionally grown as an
ornamental in dry or coastal areas because it is Liogier, H.A. 1995. Descriptive flora of Puerto
handsome, of a convenient size, and is easy to Rico and adjacent islands. Vol. 4. Editorial
propagate and manage. It is recommended as a de la Universidad de Puerto Rico, San Juan, PR.
host plant for butterflies (Mikula 2001). In the 617 p.
past, the silky hairs were used to stuff pillows
(Little and others 1974). Giant milkweed was Little, E.L., Jr., R.O. Woodbury, and F.H.
tested as a host for sandalwood, Santalum album Wadsworth. 1974. Trees of Puerto Rico and the
L., a partial root parasite. It resulted in greater Virgin Islands. Vol. 2. Agriculture Handbook
growth of sandalwood than all other species tested 449. U.S. Department of Agriculture,
(Shinde and others 1993). Extracts, chopped Washingtion, DC. 1,024 p.
leaves, and latex have shown great promise as
nematicides, in vitro and in vivo (Anver and Alam Mikula, R. 2001. Butterfly plants for your garden.
1992, Charu and Trivedi 1997). Sheep, goats, and www.butterflybreeders.com/pages/bflygdning/bu
camels will eat the leaves of giant milkweed tterflyplants.html. 5 p.
during droughts, but consumption is low. If the
138
Neal, M.C. 1965. In gardens of Hawaii. Special 11(3): 301-308.
Publication 50. Bernice P. Bishop Museum
Press, Honolulu, HI. 924 p. Sharma, B.M. 1968. Root systems of some desert
plants in Churu, Rajasthan. Indian Forester
Nehra, O.P., M.C. Oswal, and A.S. Faroda. 1987. 94(3): 240-246.
Management of fodder trees in Haryana. Indian
Farming 37(3): 31, 33. Shinde, S.R., R.D. Ghatge, and S.S. Mehetre. 1993.
Comparative studies on the growth and
Parrotta, J.A. 2001. Healing plants of Peninsular development of sandalwood tree in association
India. CAB International, Wallingford, UK and with different hosts. Indian Journal of Forestry
New York. 944 p. 16(2): 165-166.
Rahman, M.A. and C.C. Wilcock. 1991. A Varshney, A.C., and K.L. Bhoi. 1988. Cloth from
taxonomic revision of Calotropis bast fibre of the Calotropis procera (aak) plant.
(Asclepiadaceae). Nordic Journal of Botany Biological wastes 26(3): 229-232.
139
Capparis indica (L.) Druce sapo prieto
CAPPARACEAE
John K. Francis
140
sapo prieto flowers and fruits intermittently. References
Flowers were observed in Costa Rico in May and
September and fruits were present from May to Debrot, A.O., J.Y. Miller, L.D. Miller, and B.T.
September (Zamora 1989). A collection of sapo Leysner. 1999. The butterfly fauna of Curaçao,
prieto fruits from Puerto Rico weighed an average West Indies: 1996 status and long-term species
of 6.948 + 0.773 g/fruit. The air-dry seeds collected turnover. Caribbean Journal of Science 35(3-4):
from these fruits weighed an average of 0.0838 + 184-194.
0.0021 g/seed or 12,000 seeds/kg. Sown on
commercial potting mix without any pre-treatment, Howard R.A. 1988. Flora of the Lesser Antilles,
the seedlings began to emerge in 7 days and Leeward and Windward Islands.
reached a maximum germination of 49 percent in Dicotyledoneae, Part 1. Vol. 4. Arnold
64 days (author’s observation). The seeds are Arboretum, Harvard University, Jamaica Plain,
covered with a thin, pasty, scarlet pulp that is MA. 673 p.
presumably eaten by birds that transport the seeds.
Liogier H.A. 1985. Descriptive flora of Puerto
Growth and Management.—Sapo prieto has been Rico and adjacent islands, Spermatophyta. Vol.
referred to as slow growing (Zoológico Virtual de 1. Editorial de la Universidad de Puerto Rico,
Barranquilla 2002). Weaver (1990) measured one Río Piedras, PR. 352 p.
tree in St. John, U.S. Virgin Islands, as a part of a
larger study and found a 0.4 mm annual diameter Little, E.L., Jr., R.O. Woodbury, and F.H.
increment over a 5-year period. Increases in sapo Wadsworth. 1974. Trees of Puerto Rico and the
prieto density can probably be encouraged by Virgin Islands. Vol. 2. Agriculture Handbook
treatments to dry and moist forests near seed 449. U.S. Department of Agriculture,
sources to eliminate dense shade and create brush Washington, DC. 1,024 p.
patches and small openings. No wildland planting
experience has been reported.
Virgin Islands Wetlands Reserve. 2002. A list of
plants tagged along the trail at the reserve and
Benefits.—Sapo prieto makes a good ornamental
some of their uses. Virgin Islands Wetlands
that is evergreen, with attractive bicolor leaves. It
Reserve, University of the Virgin Islands, St.
has showy flowers and fruits, and a crown that can
Croix, U.S. Virgin Islands.
be shaped by pruning. However, the species is not
http://rps.uvi.edu/VIMAS/plantlist.htm. 5 p.
commonly used because it grows very slowly
(Virgin Islands Wetlands Reserve 2002, Zoológico
Weaver, P.L. 1990. Tree diameter growth rates in
Virtual de Barranquilla 2002). The wood is heavy
Cinnamon Bay Watershed, St. John, U.S. Virgin
and hard and useful for fuel, stakes, and small-
Islands. Caribbean Journal of Science 26(1-2):
diameter fence posts. Sapo prieto contributes to the
1-6.
biodiversity of the forests it inhabits, helps protect
those sites from erosion, and furnishes minor
Zamora, N. 1989. Flora arborescente de Costa
amounts of food and cover for wildlife. The
Rica. Editorial Techologica de Costa Rica, San
flowers are nectar sources for butterflies (Debrot
Jose, Costa Rica. 262 p.
and others 1999).
Zoológico Virtual de Barranquilla. 2002.
Resultados de busca por: Olivo macho.
http://zoobaq.org/formas/consultaveg2.php3?co
mun=Olivo. 1 p.
141
Capraria biflora L. goat-weed
SCROPHULARIACEAE
John K. Francis
142
published. Goatweed is considered weedy in 789 p.
pastures. In the absence of tested procedures for its
control, standard techniques of grubbing or spot Guzmán, D.J. 1975. Especies útiles de la flora
spraying with broadleaf herbicides is Salvadoreña. Ministerio de Educación,
recommended. Mowing has little lasting effect on Dirección de Publicaciones, San Salvador, El
stands. Salvador. 703 p.
Benefits.—Goatweed helps revegetate disturbed Howard, R. A. 1989. Flora of the Lesser Antilles,
areas, protects the soil, and furnishes cover for Leeward and Windward Islands. Vol. 6. Arnold
wildlife. It is a nectar source for butterflies and is Arboretum, Harvard University, Jamaica Plain,
recommended in natural landscaping (University of MA. 658 p.
Florida Extension 2002). Goatweed has many uses
in herbal medicine. A tea prepared from the leaves Liogier, H.A. 1995. Descriptive flora of Puerto
is used as an eyewash (Burkill 2000), to soothe Rico and adjacent islands. Vol. 4. Editorial de la
skin itch (Secritaría de Medio Ambiente y Universidad de Puerto Rico, San Juan, PR.
Recursos Naturales 2002), and as a general tonic. 617 p.
However, overindulgence can result in stupor,
disorientation, and paralysis (Guzmán 1975). The Long, R.W. and O. Lakela. 1976. A flora of
infusion is also used to treat fever, flu, vomiting, Tropical Florida. Banyon Books, Miami, FL.
childbirth recovery, diarrhea, hemorrhoids, 962 p.
rheumatism, and swelling (Scofield 2002). The
alkaloid, biflorine, present in the leaves has Pezzatti, B., T. Irzan, and D. Cherix. 1998. Ants
antibiotic properties (Burkill 2000), and (Hymenoptera, Formicide): Lost paradise.
sesquiterpenoids, caprariolides A and B from aerial Darwin Foundation, Galapagos, Equador.
parts, have demonstrated strong insecticidal http://www.darwinfoundation.org/articles/n5900
activity against adult sweet potato weevils, Cylas 049802.html. 10 p.
formicarius elegantulus (Collins and others 2000).
Scofield, D. 2002. Medicinal usage: Capraria
References biflora L. (Scrophulariaceae). http://www.
cassiakeyensis.com/sofl_plants/med_caprariabifl
Burkill, H.M. 2000. The useful plants of West ora.html. 1 p.
Tropical Africa. Vol. 5. Royal Botanical Garden,
Kew, UK. 686 p. Secritaría de Medio Ambiente y Recursos
Naturales. 2002. Especies con usos do
Collins, D.O., W.A. Gillimore, W.R. Reynolds, maderables en bosques de encino, pino, y pino-
L.A.D. Williams, and P.B. Reese. 2000. New encino. http://www/semarnat.gob.mx/pfnm/
skeletal sesquiterpenoids, caprariolides A-D, CaprariaBiflora.html. 2 p.
from Capraria biflora and their insecticidal
activity. Journal of Natural Products 63: 1,515- Stevens, W.D., C. Ulloa-U., A. Pool, O.M.
1,518. Montiel, eds. 2001. Flora de Nicaragua.
Monographs in Systematic Botany Vol. 85, No.
Flora of Texas Consortium. 2002. Texas vascular 3. Missouri Botanical Garden, St. Louis, MO. p.
plant checklist: Scrophulariaceae. 1,911-2,666.
http://csdl/tamu.edu/FLORA/ftc/dft/ftc_scr.htm.
5 p. University of Florida Extension. 2002. Ask a
master gardener: List of native Florida
Grisebach, A.H.R. 1963. Flora of the British West specimens for butterfly gardening. http://
Indian Islands. J. Cramer-Weinheim. New York. volusia.org/extension_service/butterfly.htm. 2 p.
143
Capsicum annuum L. bird pepper
SOLANACEAE
John K. Francis
144
Twenty-eight percent of the seeds in this collection References
germinated between 17 and 118 days after being
sown in potting mix. Seedlings may be established Bailey, L.H. 1941. The standard cyclopedia of
by sowing in prepared seed spots in the field or horticulture. Vol. 1. The MacMillan Co. New
grown in the nursery and transplanted. The seeds York, NY. 1,200 p.
are dispersed by birds that are immune to the
capsaicin the fruits contain. Rodents that would
Barrett, O.W. 1925. The food plants of Puerto
otherwise chew up and destroy the seeds will not
Rico. The Journal of the Department of
eat the fruits (UniSci 2001). Seedlings and plants
Agriculture of Puerto Rico 9(2): 168.
are common but scattered in early secondary forest
in Puerto Rico. However, bird pepper may
occasionally form small thickets.
California Antilles Trading Consortium. 2001.
Growth and Management.—Because the species Capsicum peppers. www.calantilles.com/
name “annuum” means annual and because capsicum_peppers.htm. 9 p.
cultivated forms are, or are treated as, annuals, bird
pepper has been assumed to be an annual (see
Liogier 1995). In its wild state, it is, in fact, a Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs
short-lived perennial (Floridata 2001), living 3 or 4 and cacti of South Texas. Texas Tech University
years, if conditions are favorable. Plants may Press. 213 p.
occasionally reach 5 m in height (Howard 1989).
Heights in Puerto Rico are commonly 2 to 3 m. Floridata. 2001. Capsicum spp.
The stems are slender, about 1 cm in diameter, in www.floridata.com/ref/c/caps_spp.cfm. 6 p.
larger plants (author’s observation).
Hawkens, J.G. 1991. The centers of plant diversity
Benefits.—Bird pepper has been cultivated for in Latin America. Diversity 7(1/2): 7-9
thousands of years and is the wild ancestor for
hundreds of named varieties of cultivated peppers. Hongo, H., and A.K. Karel. 1986. Effects of plant
However, the wild fruits are still harvested and extracts on insect pests of common beans.
used today. Bird pepper fruits are “hot” to very Journal of Applied Entomology 102(2): 164-
“hot” with a slight musky flavor. They are used to 169.
flavor food and make sauces. Capsicum
derivatives are used in a wide variety of medicinal Howard, R. A. 1989. Flora of the Lesser Antilles,
applications, mostly related to pain relief (eg: sore Leeward and Windward Islands. Vol. 6. Arnold
muscles, toothache, phantom limb pain). Arboretum, Harvard University, Jamaica Plain,
Capsaicin causes the brain to release endorphins MA. 658 p.
that promote a sense of well-being and deadens
pain receptors (Floridata 2001). Concentrated Krenzelok, E.P., and F.J. Provost. 1995. The ten
pepper sprays are used for riot control and most common plant exposures reported to
personal defense. Gardeners also use pepper poison information centers in the United States.
extracts to protect crops from insects and animals. Journal of Natural Toxins 4(2): 195-202.
Extracts of leaves and fruits of bird pepper have
shown insecticidal and fungicidal properties in Liogier, H.A. 1995. Descriptive flora of Puerto
laboratory tests (Hongo and Karel 1986, Patil and Rico and adjacent islands. Vol. 4. Editorial de la
Joi 1992, Williams and Mansingh 1993). Universidad de Puerto Rico, San Juan, PR.
However, to sensitive individuals, exposure to C. 617 p.
annuum fruits can be dangerous. This species
ranked fourth in plant poisoning incidents reported Nelson, Gil. 1996. The shrubs and woody vines of
to American poison information centers Florida. Pineapple Press, Inc. Sarasota, FL.
(Krenzelok and Provost 1995). In Texas and 391 p.
Arizona, several species of birds including the Rio
Grande turkey and the curve-billed thrasher eat the Patil, B.H. and M.B. Joi. 1992. Inhibition of
fruits (Everitt and Drawe 1993, UniSci 2001). tomato spotted wilt virus with leaf extracts of
some plants. Journal of Maharashtra
Agricultural University 17(2): 340-341.
145
Tewksbury, J.J., G.P. Nabhan, D. Norman, H. http://unisci.com/stories/20013/0726015.htm.
Suzan, J. Tuxill, and J. Donovan. 1999. In situ 3 p.
conservation of wild chilis and their biotic
associates. Conservation Biology 13(1): 98-107. Williams, L.A.D. and A. Mansingh. 1993.
Pesticidal potential of tropical plants—I.
UniSci 2001. Zing in the flavor helps chili pepper Insecticidal activity of leaf extracts of sixty
plants survive. Daily University Science News. plants. Insect Science and its Application 14(5):
697-700.
146
Capsicum frutescens L. red pepper
SOLANACEAE
John K. Francis
147
long as they live. The flowers are insect pollinated protection, law enforcement, and defense. Acetone
(Bosland and Votava 2000). Production of fruits and petroleum ether extracts caused complete
and seeds can be abundant. Fresh fruits from mortality of rice weevil (Sitophillus oryzae) in 15
several plants in a stand in Puerto Rico averaged days. Fruit powder was much less effective (El-
0.3668 + 0.0242 g/fruit. The variability was high Lakwah and others 1997). Red pepper has many
(CV = 39.5 percent). They ranged from 2 to 35 medicinal applications. Some of the most widely
seeds/fruit and averaged 16.6 + 2.9 seeds/fruit. Air- used and reliable are as a salve to relieve muscle,
dried seeds averaged 0.0041 + 0.0001 g/seed or joint, and toothache pain, to treat cough, asthma,
244,000 seeds/kg. Placed in moist potting mix, 72 and sore throat, as a stimulant, and to treat stomach
percent of the seeds germinated between 13 and 34 ache, seasickness, and flatulence. Anciently, it was
days after sowing. Germination is epigeous. The even used as an instrument of torture (Bentley and
seeds can be safely stored under refrigeration after Trimer 1880, Bosland and Votava 2000, Center for
air-drying and need no treatment before planting. New Crops and Plant Products 2002,
The seeds are dispersed by birds that eat the fruits Gardenguides.com 2002).
and are immune to the pungent chemicals they
contain. Young plants coppice when cut or broken. References
Rooting of cuttings and tissues has been
demonstrated for Capsicum (Bosland and Votava Bailey, L.H. 1941. The standard cyclopedia of
2000). horticulture. Vol. 1. The MacMillan Co. New
York, NY. 1,200 p.
Growth and Management.—Under continually
favorable conditions, red peppers live about 2 Bentley, R. and H. Trimer. 1880. Medicinal plants.
years. They grow rapidly during the first year, then Vol. 3. J.&A. Churchill, London. p. 147-227.
much more slowly, and finally dwindle and die.
Red peppers grown commercially are managed as Bosland, P.W. and E.J. Votava. 2000. Peppers:
annuals, much as other Capsicum peppers but for a Vegetable and spice capsicums. CABI
longer season. Plants that arise spontaneously in Publishing, Oxon, UK and New York. 204 p.
gardens and vacant land are usually allowed to
grow for occasional future harvest. Center for New Crops and Plant Products. 2002.
Capsicum pepper. Purdue University. http://
Benefits.—Red pepper is used and loved the world hort.purdue.edu/newcrop/med-aro/factsheets/
over as a condiment, added to food fresh, dried, CAPSICUM_PEPPER.html. 2 p.
refined, and ground (for Cayenne pepper and
curry), and as the principal or incidental ingredient
El-Lakwah, F.A., O.M. Khaled, M.M. Khattab,
in sauces. The source of the popular biting
and T.A. Abdel-Rahman. 1997. Toxic effects of
sensation are the capsaicinoids, principally
extracts and powders of certain plants against
capsaicin, which ranges from 600 to 13,000 ppm in
the rice weevil (Sitophilus oryzae). Annals of
the fruits (Center for New Crops and Plant
Agricultural Science (Moshtohor) 35(1): 553-
Products. 2002). The fruits are an excellent source
566.
of vitamins A and C (Bosland and Botava 2000). A
few thousand hectares are grown commercially,
Gardenguides.com. 2002. Cayenne. http://www.
and probably even more are grown in gardens or
gardenguides.com/herbs/cayenne.htm. 4 p.
harvested from the wild in rural areas of the
tropics. Coatings and powders are used with
Liogier, H.A. 1995. Descriptive flora of Puerto
varying degrees of success to deter browsing
Rico and adjacent islands. Vol. 4. Editorial de la
animals and insects, and break children of thumb
Universidad de Puerto Rico, San Juan, PR.
sucking and nail biting. Pepper spray, whose active
617 p.
ingredient is capsaicin, is used widely for personal
148
Casearia decandra Jacq. tostado
FLACOURTIACEAE
John K. Francis
149
reported to flower from August to October in References
Bolivia (Killeen and others 1993). Plants 1 m or
more in height in partial or full sunlight bear fruits. Francis, J.K. 2000. Estimating biomass and carbon
Shrubs and small trees bear from hundreds to content of saplings in Puerto Rican secondary
thousands of fruits each year. A collection of fruits forests. Caribbean Journal of Science 36(3-4):
from Puerto Rico weighed an average of 0.340 + 346-350.
0.031 g/fruit. They are extremely variable (CV =
69.9 percent), even within the same plant. Seeds Howard, R.A. 1989. Flora of the Lesser Antilles,
separated from the above collection weighed (air- Leeward and Windward Islands. Vol. 5. Arnold
dried) an average of 0.0406 + 0.0005 g/seed or Arboretum, Harvard University, Jamaica Plain,
24,600 seeds/kg. Without pre-treatment, these MA. 604 p.
seeds were sown on commercial potting mix and
91 percent germinated 7 days later (author’s Killeen, T., E. García E., and S.G. Beck. 1993.
observation). Tostado sprouts when cut or burned. Guía de árboles de Bolivia. Herbario Nacional
de Bolivia and the Missouri Botanical Garden.
Growth and Management.—Although tostado is St. Louis, MO. 958 p.
usually a shrub, if it survives several decades in
fertile sites with sufficient sunlight, it may Liogier, H.A. 1994. Descriptive flora of Puerto
occasionally become a tree 12 to 18 m in height Rico and adjacent Islands. Vol. 3. Editorial de la
(Howard 1989, Liogier 1994). Weaver (1990) Universidad de Puerto Rico. Río Piedras, PR.
reported an average 5-year annual diameter 461 p.
increment of only 0.05 cm for plants 4 to 9 cm
d.b.h. in St. John, U.S. Virgin Islands. The air- Little, E.L., Jr. and F.L. Wadsworth. 1964.
dried specific gravity of stemwood was measured Common trees of Puerto Rico and the Virgin
at 0.630 + 0.031. The above-ground average Islands. Agriculture Handbook 249. U.S.
carbon content was measured at 0.508. Total Department of Agriculture, Washington, DC.
above-ground dry weight in plants up to 5 cm 548 p.
d.b.h. can be estimated by the model: Wt =
34.356(D2S), where Wt is weight in grams, D is San José, J.J., M.R. Farinas, and J. Rosales. 1991.
diameter in cm at 30 cm above the ground-line, S is Spatial patterns of trees and structuring factors
total stem length in meters, and adjusted r-squared in a Trachypogon savanna of the Orinoco
equals 0.977 (Francis 2000). Llanos. Biotropica 23(2): 114-123.
Benefits.—The stems of tostado are sometimes Silva, D.W., E. Seitz, M. Burgo, S.M. Silva, and
used as fuel and fenceposts. This is an important J.J. Soares. 2002. Levantamento de espécies
honey plant. The small fruits are edible, but almost arbóreas no parque municipal das araucarias,
tasteless according to Little and Wadsworth (1964). Guapapuava, PR. Sociedad Botanica de São
However, the author found them sweet and Paulo, Brazil. http://www.ib.usp.br/sbsp/
pleasantly flavored. The species furnishes food and congresso/ec.htm. 1 p.
cover for wildlife, contributes to biodiversity, and
helps stabilize the soil. Weaver, P.L. 1990. Tree diameter growth rates in
Cinnamon Bay Watershed, St. John, U.S. Virgin
Islands. Caribbean Journal of Science 26(1-2):
1-6.
150
Casearia sylvestris Sw. wild-coffee
FLACOURTIACEAE
John K. Francis
151
height. Although individual plants do not flower 1990, Carvalho, and others 1999, Itokawa and
continuously, members of the population may be others 1990, Silva and others 1988). A commercial
seen flowering throughout the year (Little and perfume and a weight-loss product contain extracts
Wadsworth 1964, Stevens and others 2001). In of wild-coffee (Fontovit 2002, Multinível do Brasil
Panama, the species flowers principally in August S.A. 2002).
to December with sporadic flowering during the
dry season. The fruits mature in about 1 month References
(Croat 1978). A collection of fresh fruits (seed +
aril) from Puerto Rico weighed an average of Basile, A.C., J.A.A. Sertie, S. Panizza, T.T.
0.0286 + 0.0021 g/fruit. They vary considerably in Oshiro, and C.A. Azzolini. 1990.
size (coefficient of variation = 60.5). Air-dry seeds Pharmacological assey of Casearia sylvestris. I.
from the same collection averaged 0.0011 + 0.0001 Preventive anti-ulsur activity and toxicity of the
g/seed or 909,000 seeds/kg. Sown on peat, these leaf crude extract. Journal of
seeds germinated at 31 percent between 20 and 103 Ethnopharmacology 30(2): 185-197.
days after sowing (author’s observation). In
southern Brazil, 84,000 seeds/kg were reported and Carvalho, J.C.T., V.V. Vignoli, G.H.G. de Souza,
a low rate of germination (Universidad do Extremo K. Ujikawa, J.J. Neto, and G.H.B. de Souza.
Sul Catarinense 1999). Germination is epigeal. Just 1999. Antimicrobial activity of essential oils
one or two fruits per inflorescence are ripe at any from plants used in Brazilian popular medicine.
time. The only method of seed collection known to Acta Horticulturae 501: 77-81.
the author is to hand-pick the tiny ripe fruits
(capsules split exposing the red aril) and wet sieve Croat, T.B. 1978. Flora of Barro Colorado Island.
to separate the seeds. Birds disburse the seeds. Stanford University Press, Stanford, CA. 943 p.
Natural seedlings are rare to common. Plants past
the seedling stage have a high survival rate. Fontovit. 2002. Porangaba Fontovit.
Damaged plants coppice readily. http://fontovit.com.br/produtos/porangaba.htm.
1 p.
Growth and Management.—Wild-coffee has a
moderate growth rate and lives for at least 20 Howard, R.A. 1989. Flora of the Lesser Antilles,
years. Artificial reproduction is usually by seeds, Leeward and Windward Islands. Vol. 5. Arnold
but cuttings can also be rooted. Recommended Arboretum, Harvard University, Jamaica Plain,
spacing for plantings in Brazil is 4 by 4 m MA. 604 p.
(Universidad do Extremo Sul Catarinense 1999).
Because it becomes established as part of the early Instituto de Botánica Darwinion. 2002. Catálogo
secondary forest or in disturbed openings of more de las plantas vasculares de la Argentina:
advanced forest, management to promote Flacourtiaceae. http://www.darwin.edu.ar/
regeneration of the species might include the use of Catalogo/Flacourtiaceae.pdf. 3 p.
clearcuts or group selection in closed forest and
protection of open land from fires. Itokawa, H., N. Totsuka, H. Morita, K. Takeya, Y.
Iitaka, E.P. Schenkel, and M. Motidome. 1990.
Benefits.—The wood of wild-coffee is used for or New antitumor principles, casearines A-F, for
is suitable for fuel, fence posts, stakes, small poles, Casearia sylvestris Sw. (Flacourtiaceae).
rustic carpentry, and tool handles (Little and Chemical and Pharmaceutical Bulletin 38(12):
Wadsworth 1964). The species is recommended for 3,384-3,388.
urban planting as a food source for birds (Rio
Grande Energia 2002). It is a honey plant (Little Killeen, T.J., E. García E., and S.G. Beck. 1993.
and Wadsworth 1964). Leaves of wild-coffee Guía de árboles de Bolivia. Herberio Nacional
contains 2.5 percent essential oils with a pleasant de Bolivia and Missouri Botanical Garden, St.
aroma. They also contain caprionic acid, saponins, Louis, MO. 958 p.
alkaloids, flavonoids, and a host of other
chemicals. Extracts (particularly of the leaves) are Liogier, H.A. 1994. Descriptive flora of Puerto
used in traditional herbal medicine to treat a large Rico and adjacent Islands. Vol. 3. Editorial de la
number of different ailments (Universidad do Universidad de Puerto Rico. Río Piedras, PR.
Extremo Sul Catarinense 1999). Laboratory 461 p.
experiments have confirmed antiseptic, antitumor,
antiulser, and abortive activity (Basile and others
152
Little, E.L., Jr. and F.L. Wadsworth. 1964. rs.com.br/manual_poda/esp_recomendadas.asp.
Common trees of Puerto Rico and the Virgin 5 p.
Islands. Agriculture Handbook 249. U.S.
Department of Agriculture, Washington, DC. Silva, F.A., A.L.M. Baisch, B. Oliveira, A.M.
548 p. Battastini, F. Torres, G. Rocoski, E.S. Silva,
M.F. Alam, J.C.G. Apolinario, and A.J. Lapa.
Lugo, A.E. 1992. Comparison of tropical tree 1988. Preliminary pharmacological studies on
plantations with secondary forests of similar extracts from Casearia sylvestris Swartz. Acta
age. Ecological Monographs 62: 1-41. Amazonica Suplemento 18(1-2): 219-229.
Multinível do Brasil S.A. 2002. Uma nova geração Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
de colônias. http://www.multiniveldobrasil. Montiel, eds. 2001. Flora of Nicaragua.
com.br/bio_collection.html. 2 p. Monographs in Systematic Botany Vol. 85, No.
2. Missouri Botanical Garden Press. St. Louis,
van Os, M. 2000. The Llanos de Orinoco. MO. p. 945-1,910.
Wageningen Agricultural University,
Wageningen, Netherlands. http://www.wau.nl/ Universidad do Extremo Sul Catarinense. 1999.
rpv/ond_proj/dirjongm/Llanos/teksten/objectives Jornada Catarinense de plantas medicinais:
/chapter03.html. 8 p. Guaçatonga. http://www.unesc.rct-sc.br/
plantas_medicinais/guaca.htm. 4 p.
Río Grande Energia. 2002. Manual de arborização
e poda: espécies recomendadas. http://www.rge-
153
Cassine xylocarpa Vent. marble tree
CELASTRACEAE
John K. Francis
They are entire or with a few teeth, elliptic to
obovate, 2.5 to 12.5 cm long, with a rounded to
spine-tipped apex. The inflorescences are tiny-
flowered, branched axillary cymes. The fruit is a
yellow (at maturity), rounded drupe, 1.5 to 3 cm
long with a thin, hard-fleshy exocarp and hard,
thick-walled stone with one to three cells and
elliptic seeds (Howard 1989, Liogier 1994, Little
and others 1974).
154
percent, respectively, of air layers and misted MA. 604 p.
cuttings treated with IBA rooted. The newly
formed roots are fragile and easily broken off Instituto Nacional de Biodiversidad. 2000. Leaps
during transplanting (author’s observation). The and bounds. The Cutting Edge 7(2): 1-2,
seeds are probably dispersed by fruit bats. http://inbio.ac.cr/papers/manual_plantas?apr00le
Seedlings and saplings are relatively common in a.html. 2 p.
and around Puerto Rican stands of the species.
Liogier, H.A. 1990. Plantas medicinales de Puerto
Growth and Management.—Marble tree grows Rico y del Caribe. Iberoamericana de Edicones,
slowly. A 1-m tall sapling with a basal diameter of Inc. San Juan, PR. 566 p.
1.5 cm growing in the understory of low basal area
moist forest had 37 annual rings (author’s Liogier, H.A. 1994. Descriptive flora of Puerto
observation). Individuals as large as 20 cm in Rico and adjacent Islands. Vol. 3. Editorial de la
diameter at breast height (Little and others 1974) Universidad de Puerto Rico, San Juan, PR.
must be several decades old. Management of the 461 p.
species should include protection from fire and
from commercial development of habitat. Existing Little, E.L., Jr., R.O. Woodbury, and F.H.
understory seedlings and saplings could be Wadsworth. 1974. Trees of Puerto Rico and the
released from competition by thinning or Virgin Islands. Vol. 2. Agriculture Handbook
eliminating the overstory in patches. 449. U.S. Department of Agriculture,
Washingtion, DC. 1,024 p.
Benefits.—Marble tree is pretty enough to make a
good ornamental but has rarely been used as such. Medina, E., E. Cuevas, J. Figueroa, and A.E.
It is an important component of the plant Lugo. 1994. Mineral content of leaves from
community in the harsh environment where it is trees growing on serpentine soils under
most common and furnishes food and cover for contrasting rainfall regimes in Puerto Rico. Plant
animals. The fruits of marble tree are listed as a and Soil 158(1): 13-21.
food item for the endangered Anegada iguana
(Cyclura pinguis) (Zoological Society of San Missouri Botanical Garden. 2002a. Tropicos-Flora
Diego 2002). The wood is light-brown, hard, of Panama Checklist. http://mobot.org/cgi-
heavy, fine-textured, strong, and durable (Little bin/search_pick. 1 p.
and others 1974). It is certainly useful for fuel and
fence posts and possibly carving and turnery. The Missouri Botanical Garden. 2002b. W3-Specimen
fruits are reported to be edible, and extracts of the data base. http://mobot.org/cgi-bin/search_vast.
plant are a stimulant (Liogier 1990). 1 p.
155
Castela erecta Turp. cockspur
SIMAROUBACEAE
John K. Francis
elsewhere.
156
Arboretum, Harvard University, Jamaica Plain, Locklin, C. 2002. Guajira-Barranquilla xeric scrub
MA. 673 p. (NT1308). World Wildlife Fund, Washington,
DC. http://www.worldwildlife.org/wildworld/
Liogier, H.A. 1988. Descriptive flora of Puerto profiles/terrestrial/nt/nt1308_full.html. 7 p.
Rico and adjacent islands, Spermatophyta. Vol.
2. Editorial de la Universidad de Puerto Rico, Secretaría de Medio Ambiente y Recursos
Río Piedras, PR. 481 p. Naturales. 2002. Especies forestales no
maderables y maderables no tradicionales de
Little, E.L., Jr., R.O. Woodbury, and F.H. zonas áricas y semiáridas: Castela texana
Wadsworth. 1974. Trees of Puerto Rico and the (T.&G.) Rose. http://www.semarnat.gob.mx/
Virgin Islands. Vol. 2. Agriculture Handbook pfnm3/fichas/castela_texana.htm. 2 p.
449. U.S. Department of Agriculture,
Washingtion, DC. 1,024 p.
157
Ceanothus velutinus Dougl. ex Hook. snowbush ceanothus
RHAMNACEAE
Synonyms: none
John K. Francis
158
August and September (Banner 2002). The flowers round, although it makes up only a small portion
are pollinated by at least bees (Clark 1976). There of their diet. It is important fall and winter moose
is an average of 207,000 seeds/kg. When seeds browse in some areas. It has an in vitro dry matter
mature, beginning in August, they are ejected from digestibility of about 57 percent and a crude
the pod and dispersed a short distance (Anderson protein content of 7 to 18 percent (Anderson
2001). Good seed production begins when plants 2001). Snowbush ceanothus is important resting
are about 8 years old. As many as 1,250,000 and escape cover for many types of wildlife
seeds/ha/year can be produced. However, as much (Pollock 2003). Nitrogen is fixed by actinomycetes
as 99 percent of the annual seed crop can be of the genus Frankia in root nodules (Dalton
consumed by ants, birds, and rodents. (Pollock 1997). Snowbush ceanothus makes a beautiful and
2003). Seeds still accumulate in large numbers in hardy ornamental that can be pruned and shaped
the soil seed bank and can apparently remain (Hansen 2002). Native Americans used the leaves
viable for 200 to 300 years until conditions as a deodorant, smoke of the plant to kill bedbugs,
become favorable. Germination is epigeal (Reed and decoctions to treat flu, pain, and gonorrhea
1974). Heat, as from forest fires, appears to scarify (Moerman 1986). They also used the leaves as a
the seeds. Burned or damaged plants sprout from tobacco substitute (Pollock 2003).
the root crown. Layering also occurs when
branches come in contact with the soil (Anderson References
2001).
Abrams, L. 1951. Illustrated flora of the Pacific
Growth and Management.—The natural life States. Vol. 3. Stanford University Press,
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However, stands sometimes begin to deteriorate Anderson, M.D. 2001. Ceanothus velutinus. In:
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stand in the Cascade Mountains in Oregon Rocky Mountain Research Station, Fire Sciences
contained over 4,000 stems/ha and averaged Laboratory, Fire Effects Information System.
34,000 kg/ha of above-ground biomass (Pollock http://www.fs.fed.us/database/feis/plants/shrub/
2003). Seeds may be sown in the late summer and ceavel/all.html. 73 p.
will germinate in the spring. Pretreatments are
necessary for good germination of spring-planted Banner, R. 2002. Deerbush. Utah State University,
seeds. A hot water soak (90 °C until the water Logan, UT. http://extension.usu.edu/coop/
cools) followed by cold stratification at 1 to 5 °C natres/range/Woody/deerbush.htm. 4 p.
for 30 to 90 days. Seeds should be sown in flats at
depths of twice the diameter of the seed in Brayshaw, T.C. 1996. Trees and shrubs of British
amended mineral soil. Germination may be as high Columbia. UBC Press, Vancouver, British
as 82 percent. Seedlings should be pricked into Colunbia, Canada. 373 p.
pots when they have gained several sets of true
leaves (Reed 1974). Plants can be propagated with Britton, N.L. and A. Brown. 1913. Illustrated flora
stem cuttings. Survival of transplanted seedlings is of the northern states, Canada, and the British
low (about 9 percent). Small seedlings survive possessions. Vol. 2, 2nd ed. Scribner, New York.
better than large ones (Pollock 2003). Snowbush 735 p.
ceanothus stands are difficult to control. Broadleaf
herbicides such as 2,4-D tend to top-kill rather Clark, L.J. 1976. Wild flowers of the Pacific
than eliminate it. Opening the brush canopy also Northwest. Gray’s Publishing Limited, Sidney,
allows seeds to germinate and grow. Glyphosate British Columbia, Canada. 604 p.
and hand slashing result in greater densities of the
species in the long run (Anderson 2001). Dalton, D. 1997. Nitrogen fixation: range of
organisms that can fix nitrogen. Reed College,
Benefits.—Snowbush ceanothus adds to the Portland, OR. http://academic.reed.edu/
beauty of the forest, helps protect the soil, biology/Nitrogen/Nfix1.html. 7 p.
furnishes food and cover for wildlife, and is a
useful ornamental and medicinal plant. The Hansen, W. 2002. Ceanothus. http://www.
species is little used by cattle and horses. Sheep nwplants.com/plants/shrubs/ceanothus/. 6 p.
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Deer and elk browse snowbush ceanothus year-
159
Harrington, H.D. 1964. Manual of the plants of Pollock, T. 2003. Ceanothus velutinus. University
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Welsh, S.L., N.D. Atwood, S. Gooddrich, and L.C.
Nobs, M.A. 1963. Experimental studies on species Higgins. 1987. A Utah flora. Great Basin
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160
Celastrus orbiculatus Thunb. oriental bittersweet
CELASTRACEAE
161
bittersweet are in long terminal clusters. Both (Baker 1974, Newsome and Noble 1986, Sakai
species have the same chromosome number (n = and others 2001)–that made early horticulturists
23; Bowden 1945) and they can be hybridized prefer oriental bittersweet over American
(White and Bowden 1947, Pooler and others bittersweet (C. scandens L.) are the same
2002). The presence of some intermediate material characteristics that enable oriental bittersweet to be
in the wild with both axillary and terminal an extremely successful invasive plant.
inflorescences has led to the speculation that the
two species may hybridize in the wild (Dreyer and Reproduction.—Oriental bittersweet blooms in
others 1987, Mehrhoff in Pooler and others 2002). the spring and is pollinated by hymenopterous
Some spell the specific epithet “orbiculata” instead insects, especially bees, although wind may also
of “orbiculatus” used here. This is because be involved (Brizicky 1964). The fruit ripens in
botanists disagree as to whether the genus name the fall. The seeds are dispersed by birds and small
Celastrus L. should be treated as feminine mammals (Dreyer 1994). Humans are also
(“orbiculata”) or masculine (“orbiculatus”) (Paclt important dispersal agents since the fruiting stems
1998). In classical Greek, the generic name was are commonly used in dried flower arrangements
treated as feminine; however, Linnaeus treated the and subsequently discarded in compost and brush
genus as masculine. The matter is now being piles (Dreyer 1994). Oriental bittersweet can also
reviewed by a formal nomenclatural committee reproduce asexually through root suckering.
(Brummitt 2000).
Growth and Management.—Due to the plant’s
Ecology.—Oriental bittersweet grows in a wide highly invasive nature, it is not recommended that
variety of habitats, including, dune thickets, this plant be grown. Small infestations can be
fencerows, forests, forest edges, and roadsides. It controlled by cutting the stems and removing the
seems to do particularly well in disturbed habitats roots. Larger infestations require the application of
(Dreyer 1994, Steward and others in press). This glycophosphate herbicides (Miller 2002).
species was perhaps first introduced in the New
World during the 1860s and was later popularized References
for ornamental plantings (Patterson 1973, Rehder
1940). Oriental bittersweet is an aggressive Bailey, L.H. 1922. The standard cyclopedia of
invader that can dominate all vegetation levels of horticulture. vol 2. Macmillan Co., New York.
forested and open areas (Bergmann 2003). It 1,200 p
grows over other vegetation, completely covering
it. It can kill other plants by preventing them from Baker, H.G. 1974. The evolution of weeds. Annual
photosynthesizing and through girdling and Review of Ecology and Systematics 5: 1-24.
uprooting. Based on its native range and habitat
types, Oriental bittersweet can be expected to Bergmann, C. 2003. Oriental Bittersweet Celastrus
spread to additional areas in the United States and orbiculatus Thunb. http://www.nps.
Canada (J. Ma, personal observation). Oriental gov/plants/alien/fact/ceor1.htm. [not paged].
bittersweet is a shade tolerant species, able to
acclimate to low light levels under heavy shading Bowden, W.M. 1945. A list of chromosome
and grow explosively if shading is reduced. Once numbers in higher plants I. Acanthaceae to
large clones are established, it is difficult to Myrtaceae. American Journal of Botany 32: 81-
eliminate since it has developed such an extensive 92.
root system. While oriental bittersweet is
spreading and increasing in abundance, the native Brizicky, G.K. 1964. The genera of Celastrales in
American bittersweet is concurrently declining the southeastern United States. Journal of the
(Dreyer and others 1987, Forman and Stark 2001, Arnold Arboretum 45: 206-235.
Steward and others in press). Pooler and others
(2002) have hypothesized that the spread of Brummitt, R.K. 2000. Report of the Committee for
Oriental bittersweet and its hybridization with Spermatophyta: 49. Taxon 49: 261-278.
American bittersweet may be threatening the
genetic identity of the American bittersweet. The Cheng, C.Y., and P.H. Huang. 1999. Celastraceae.
very properties–faster growth (Bailey 1922, Flora Reipublicae Popularis Sinicae 45: 1-218
Rehder 1940), greater fecundity (Clemant and (in Chinese).
others 1991, Dreyer and others 1987, Hart 1928),
greater tolerance of environmental heterogeneity
162
Clemant, C., R. Warren, G. Dreyer, and P. Barnes. Patterson D. 1973 Distribution of Oriental
1991. Photosynthesis, water relations, and bittersweet in the United States. Journal of the
fecundity in the woody vines American and Elisha Mitchell Science Society 89: 245.
Oriental bittersweet (Celastrus scandens and
Celastrus orbiculatus). American Journal of Pooler, M.R., R.L. Dix, and J. Feely. 2002.
Botany 78 (Suppl.): 134-135. Interspecific hybridization between the native
bittersweet, Celastrus scandens and the
Clemants, S.E. 2003. Celastrus orbiculatus. New introduced invasive species, C. orbiculatus.
York Metropolitan Flora Project. Brooklyn Southeastern Naturalist 1: 69-76.
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paged]. shrubs. Macmillan Publishing Co., New York.
996 p.
Dreyer, G.D. 1994. Element stewardship abstract
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Nature Conservancy, Arlington, Virginia. Lodge, J. Molofsky, K.A. With, S. Baughman,
http://tncweeds.ucdavis.edu/esadocs/documnts/ R.J. Cabin, J.E. Cohen, N.C. Ellstrand, D.E.
celaorb.html.11p. McCauley, P.O’Neil, I.M. Parker, J.N.
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comparisons of reproductive potential between a 332.
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163
Celastrus scandens L. American bittersweet
CELASTRACEAE
164
23, Bowden 1945) and they have been hybridized reported as being poisonous to cattle (Brizicky
(White and Bowden 1947, Pooler and others 1964). Fruiting stems of the species are often used
2002). The presence of some intermediate material in dried arrangements.
in the wild with both lateral and terminal
inflorescences has led to the speculation that the References
two species may hybridize in the wild (Dreyer and
others 1987, Mehrhoff in Pooler and others 2002). Bailey, L.H. 1922. The standard cyclopedia of
horticulture Vol 2. Macmillan Co., New York.
Ecology.—American bittersweet grows in a wide 1,200 p.
variety of habitats, including dune thickets,
fencerows, forests, forest edges, and roadsides. Bailey, L.H. 1949. Manual of cultivated
The native American bittersweet is declining, plants.Macmillan Co., New York. 1,116 p.
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very properties--faster growth (Bailey 1922,
Rehder 1940), greater fecundity (Clemant and Britton, N. L. and A. Brown 1897. An illustrated
others 1991, Dreyer and others 1987, Hart 1928), flora of the northern United States, Canada, and
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and others 2001)--that made early horticulturists
prefer Oriental bittersweet over American Brizicky, G.K. 1964. The genera of Celastrales in
bittersweet are the same characteristics that enable the southeastern United States. Journal of the
Oriental bittersweet to outcompete American Arnold Arboretum 45: 206-235.
bittersweet.
Chittenden, F.J. and P.M. Synge. 1956. Dictionary
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the spring and is pollinated by hymenopterous Oxford, UK. 512 p.
insects, especially bees, although wind may also
be involved (Brizicky 1964). The fruit ripens in Clemant, C., R. Warren, G. Dreyer, and P. Barnes.
the fall. The seeds are dispersed by birds and small 1991. Photosynthesis, water relations, and
mammals (Dreyer 1994). American bittersweet fecundity in the woody vines American and
can also reproduce asexually through root Oriental bittersweet (Celastrus scandens and
suckering. Celastrus orbiculatus). American Journal of
Botany 78 (Suppl.): 134-135.
Growth and Management.—American
bittersweet does well in many conditions. It is a Clemants, S.E. 2003. Celastrus orbiculatus. New
rapid grower, adding up to several meters in one York Metropolitan Flora Project. Brooklyn
growing season. Adequate sun is important for Botanic Garden http://www.bbg.org/sci/nymf/
fruit production. It can be planted in Hardiness encyclopedia/cel/cel0030b.htm#description. [not
Zones 3 to 8, in soils with pH ranges from 3.7 to paged].
6.5. It cannot tolerate saturated soils. Propagation
can be effected either through layering or seed Dillingham, F.T. 1907. The staff-tree, Celastrus
(Chittenden and Synge 1956). Seeds should be scandens, as a former food supply of starving
stratified in the dark at about 4 °C for 3 months Indians. American Naturalist 41: 391-393.
(Young and Young 1992)
Dreyer, G.D. 1994. Element stewardship abstract
Benefits.—The branches of American bittersweet for Celastrus orbiculata Asiatic bittersweet. The
were used by Native Americans, including the Nature Conservancy, Arlington, Virginia.
Chippewa, as a last resort food source (Dillingham http://tncweeds.ucdavis.edu/esadocs/documnts/c
1907, Palmer 1871). The leaves have been elaorb.html. 11p.
165
Dreyer, G.D., L.M. Baird, and C. Fickler, 1987. Rehder, A. 1940. Manual of cultivated trees and
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comparisons of reproductive potential between a 996 p.
native and an introduced woody vine. Bulletin of
the Torrey Botanical Club 114: 260-264. Saikai, A.K., F.W. Allendorf, J.S. Holt, D.M.
Lodge, J. Molofsky, K.A. With, S. Baughman,
Forman, J. and J. Stark. 2001. The introduction of R.J. Cabin, J.E. Cohen, N.C. Ellstrand, D.E.
non-native plants into Massachusetts. http:// McCauley, P.O’Neil, I.M. Parker, J.N.
omega.cc.umb.edu/~conne/jennjim/celastrus. Thompson, and S.G. Weller. 2001. The
html. [not paged]. population biology of invasive species. Annual
Review of Ecology and Systematics 32: 305-
Hart, H.T. 1928. Delayed germination in seeds of 332.
Peltandra virginica and Celastrus scandens.
publications of the Puget Sound Biological Scoggan, H.J. 1978. The Flora of Canada Part 3 –
Station of the University of Washington 6: 255- Dicotyledoneae (Saururaceae to Violaceae).
261. National Museum of Natural Sciences, National
Museums of Canada, Ottawa. 568 p.
Hou, D. 1955. A revision of the genus Celastrus.
Annals of the Missouri Botanical Garden 42: Steward, A.M., S.E. Clemants, and G. Moore
215-302. 2003. The concurrent decline of the native
Celastrus scandens and spread of the non- native
Newsome, A.E. and I.R. Noble. 1986. Ecological Celastrus scandens in the New York City
and physiological characters of invading species. Metropolitan area. Journal of the Torrey
In: R.H. Groves and J.J. Burdon, eds. Ecology of Botanical Society 130: 143-146.
biological invasions. Cambridge University
Press, Cambridge, MS. p. 1-20. USDA, NRCS. 2003. The PLANTS Database,
Baton Rouge, LA. http:plants.usda.gov. [not
Palmer, E. 1871. Food products of the North paged].
American Indians. In H. Capron, Report of the
Commissioner of Agriculture for the Year 1870, White, O.E. and W.M. Bowden 1947. Oriental and
Washinton, D.C. p. 404-428 American bittersweet hybrids. Journal of
Heredity 38: 125-127.
Pooler, M.R., R.L. Dix, and J. Feely. 2002.
Interspecific hybridization between the native Young, J.A. and C.G. Young. 1992. Seeds of the
bittersweet, Celastrus scandens and the woody plants of North America. Dioscorides
introduced invasive species, C. orbiculatus. Press, Portland, OR. 407 p.
Southeastern Naturalist 1: 69-76.
166
Celtis reticulata Torr. netleaf hackberry
ULMACEAE
167
northern latitudes stands are often associated with Bonner, F.T. 1974. Celtis L. Hackberry. In: C.S.
rock (DeBolt and McCune 1995), which provides Shopmeyer, tech. coord. Seeds of woody plants
extra moisture and some protection from wildfire. in the United States. Handbook 450. U.S.
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Reproduction.—Wind-pollinated, nonshowy Washington, DC. p. 298-300.
flowers appear in spring, just before or as the
leaves unfold, depending on latitude. Fruits ripen Brown, D.E. 1982. Great Basin montane
from September to November and persist on the scrubland. In: Biotic communities of the
trees until midwinter. They are dispersed by birds, American Southwest–United States and Mexico.
rodents, and other small mammals. Netleaf Desert Plants 4: 83-84.
hackberry exhibits low germination percentages
(37 percent) and high seed dormancy (Bonner Carmichael, R.S., O.D. Knipe, C.P. Pase, and
1974). Seeds are hard and thick-walled, probably W.W. Brady. 1978. Arizona chaparral: plant
the major impediment to rapid germination. associations and ecology. Agriculture Research
Germination is epigeal. Dormancy can be Paper RM-202. U.S. Department of Agriculture,
overcome with stratification at 5 oC in moist sand Ft. Collins, CO. 16 p.
or other suitable media for 120 days (Bonner
1974). Mechanical scarification enhances Daubenmire, R. 1970. Steppe vegetation of
germination after 45 days (DeBolt 1992). Netleaf Washington. Technical Bulletin 62. Agriculture
hackberry can reproduce vegetatively by sprouting Experiment Station, Pullman, WA. 131 p.
from the root crown, stem base, or lateral roots. It
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hackberry in Idaho. Master’s thesis, Oregon
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from animal-, gravity-, or water-dispersed
propagules in sites free of herbaceous competition, DeBolt, A.M. and B. McCune. 1995. Ecology of
often in rocky areas or in the duff of an existing Celtis reticulata in Idaho. Great Basin Naturalist
stand. Plummer (1977) found netleaf hackberry 55: 237-248.
useful in stabilizing disturbed areas in pinyon-
juniper and mountain brush types of Utah. In Dick-Peddie, W.A. and J.P. Hubbard. 1977.
Texas and Arizona, it is increasingly being used Classification of riparian vegetation. Pages 85-
for riparian restoration and wildlife plantings. It 90 In: R.R. Johnson and A. Dale, eds.
currently receives limited use in artificial plantings Importance, preservation, and management of
in the northern portion of its range, though once riparian habitat. General Technical Report RM-
established, young plants are hardy and persistent. 43. U.S. Department of Agriculture, Ft. Collins,
Netleaf hackberry is generally slow-growing, but CO.
plants may live for 300 to 400 years (DeBolt and
McCune 1995). Elias, T.S. 1980. The complete trees of North
America. Times Mirror Magazines, Inc., New
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twigs are browsed by bighorn sheep, mule deer, Monographs 18: 473-506.
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Cephalanthus occidentalis L. buttonbush
RUBIACEAE
Kristina Connor
170
Longevity of seeds in storage is unknown but they Bonner, F.T. 1974b. Chemical components of
are thought to be orthodox. Thus, if seeds are dried some southern fruits and seeds. United States
to a moisture content of 12 percent or less, they Department of Agriculture Forest Service Res.
can be stored under refrigeration for long periods Note SO-183, New Orleans. 3 p.
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forced above ground, and no pretreatment is Bramble, W.C. and M.K. Goddard. 1943. Seasonal
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Brown, C.L. and L.K. Kirkman. 1990. Trees of
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DuBerry, A.P., Jr. 1963. Germination of
growing but short-lived shrub. It can be used as a
bottomland tree seed while immersed in water.
landscape shrub border but if not planted in moist
Journal of Forestry 61: 225- 226.
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pruned to maintain a good form. In nature,
Faber-Langendoen, D. and P.F. Maycock. 1989.
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Community patterns and environmental
resprout after fire (Vogl 1973, Wade and others
gradients of buttonbush, Cephalanthus
1980). It has no reported pest problems but is
occidentalis, ponds in lowland forests of
moderately sensitive to herbicides. Thickets can be
southern Ontario. Canadian Field-Naturalist
reduced by cutting.
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Benefits.—Buttonbush seeds are an important
Holstein, G. 1984. California riparian forests:
food for water birds but can be toxic to other
deciduous islands in an evergreen sea. In: R.E.
animals (Snyder 1991). Dense buttonbush thickets
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riparian systems: Ecology, conservation, and
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productive management Proceedings of a
for butterflies, bees, and hummingbirds. A
conference; 1981 Sept. 17-19, Davis CA.
decoction of the inner bark was used by Native
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contain bitter glycosides, cephalin and
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cephalanthin (ACES 2001), are reportedly toxic to
incompatibility, and secondary pollen
some animals, especially cattle that eat them when
presentation in Cephalanthus occidentalis
other foliage is scarce. However, buttonbush
(Rubiaceae). American Journal of Botany 80:
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parts are less toxic.
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2001. Poisonous Plants of the Southeastern
United States.ANR-975 http://www.aces.edu/
McCarron, J.K., K.W. McLeod, and W.H. Conner.
dept/extcomm/ publications. 56 p.
1998. Flood and salinity stress of wetland
woody species buttonbush (Cephalanthus
Bonner, F.T. 1974a. Cephalanthus occidentalis L.,
occidentalis) and swamp tupelo (Nyssa sylvatica
common buttonbush. In: C.S Schopmeyere,
var. biflora). Wetlands 18: 165-175.
tech. coord. Seeds of woody plants in the United
States. United States Department of Agriculture
Roberts, E.H. 1973. Predicting the storage life of
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seeds. Seed Science and Technology 1: 499-514.
Washington, D.C. 883 p.
171
Snyder, S.A. 1991. Cephalanthus occidentalis. In: Vines, R.A. 1960. Trees, shrubs, and woody vines
United States Department of Agriculture, Forest of the Southwest. University of Texas Press,
Service, Rocky Mountain Research Station Fire Austin, TX, 1,104 p.
Sciences Laboratory. Fire Effects Information
System, http://www.fs.fed.us/database/feis. 10 p. Vogl, R.J. 1973. Effects of fires on the plants and
animals of a Florida wetland. American Midland
Van Dersal, W.R. 1938. Native woody plants of Naturalist 89: 334-347.
the United States: their erosion-control and
wildlife values. United States Department of Wade, D., J. Ewel, and R. Hofstetter. 1980. Fire in
Agriculture Miscellaneous Publication 393, South Florida ecosystems. United States
Washington, D.C. 362 p. Department of Agriculture Forest Service
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125 p.
172
Cercocarpus intricatus Wats. littleleaf mountain-mahogany
ROSACEAE
Stanley G. Kitchen
173
sun. Soil pH of littleleaf mountain-mahogany Resources, Utah State University, Logan, UT.
habitat is neutral to slightly basic, (Fairchild and p. 1-13.
Brotherson 1980). Annual precipitation is 200 to
500 mm. Although undocumented, it is believed Fairchild, J.A. and J.D. Brotherson. 1980. Micro-
that this species possesses nitrogen fixing root habitat relationships of six major shrubs in
nodules similar to those reported for close relatives Navajo National Monument, Arizona. Journal of
(Davis 1990). Littleleaf mountain-mahogany is Range Management 33: 150-156.
browsed heavily by wild ungulates and domestic
sheep when it can be reached. Kitchen, S.G. and S.E. Meyer. 1990. Seed
dormancy in two species of mountain-mahogany
Reproduction.—Flowering occurs from mid-May (Cercocarpus ledifolius and Cercocarpus
to late June (Walker and Turley 1999). Fruits ripen montanus). In: K.L. Johnson, ed. Proceedings of
and disperse in July and August. Years of the Fifth Utah Shrub Ecology Workshop: The
abundant seed production are rare. A high Genus Cercocarpus; 1988 July 13-14; Logan,
percentage of empty fruits is common. Plummer UT. College of Natural Resources, Utah State
and others (1968) reported 112,000 cleaned seeds University, Logan, UT. p. 27-42.
(fruits with tails removed) per kg. Seed dispersal is
by wind. NatureServe Explorer. 2002. An online
encyclopedia of life [web application]. Version
Growth and Management.—Although 1.6. NatureServe. Arlington, VA, USA.
information on seed germination for this species is http://www.natureserve.org/explorer. [not
lacking in the literature, primary dormancy at seed paged].
dispersal is probably similar to that of curlleaf
mountain-mahogany. Kitchen and Meyer (1990) Plummer, A.P., D.R. Christensen, and S.B.
observed almost no germination for this species Monsen. 1968. Restoring big-game range in
without prechill (1 oC). They reported germination Utah. Publ. No. 68-3. Utah division of Fish and
of six seed collections after 6 weeks of prechill Game, Ephraim, UT. 183 p.
ranged from 53 to 96 percent of viable. Seedling
growth rate is slow, and plants require 10 to 30 Stutz, H.C. 1990. Taxonomy and evolution of
years to reach maturity. Management alternatives Cercocarpus in the western United States. In:
for littleleaf mountain-mahogany are limited due K.L. Johnson, ed. Proceedings of the Fifth Utah
to the nature of the sites it occupies. Shrub Ecology Workshop: The Genus
Cercocarpus, 1988 July 13-14, Logan, UT.
Benefits.—Littleleaf mountain-mahogany College of Natural Resources, Utah State
provides cover and year-round forage on harsh University, Logan, UT. p. 15-25.
landscapes. It is of particular value to native
ungulates that depend on rugged escape terrain Walker, S.C. and D. Turley. 1999. Characteristics
such as mule deer (Ococolius hemionus), bighorn of mountain mahogany (Cercocarpus) species
sheep (Ovis Canadensis), and mountain goat and hybrids in Utah hybrid zone. In: E.D.
(Oreamnos americanus). McArthur, W.K. Ostler, and C.L. Wambolt,
comps. Proceedings: Shrubland Ecotones; 1998
References. August 12-14; Ephraim, UT. Proceedings
RMRS-P-11. U.S. Department of Agriculture,
Davis, J.N. 1990. General ecology, wildlife use, Forest Service, Rocky Mountain Research
and management of the mountain mahoganies in Station, Ogden, UT. p. 32-34.
the Intermountain West. In: K.L. Johnson, ed.
Proceedings of the Fifth Utah Shrub Ecology Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Workshop: The Genus Cercocarpus; 1988 July Goodrich. 1987. A Utah Flora. Brigham Young
13-14; Logan, UT. College of Natural University Print Services, Provo, UT. 894 p.
174
Cercocarpus ledifolius Nutt. curl-leaf mountain mahogany
ROSACEAE
Synonyms: None
Christopher Ross
175
very light burns that do no apparent damage to References
mature trees result in full mortality within 1 year.
Closed mature stands may have insufficient Billings, W. D. 1976. Islands of sierran plants on
understory to carry fire, so that the fire and the arid slopes of Peavine Mountain. Mentzelia
resulting mortality are confined to the edges of the 6: 32-39.
stands. Regeneration by seed may occur after fire
if the soil is not colonized by other species. In Davis, J.N. and J. D. Brotherson. 1991. Ecological
some cases, the resulting stands occupy essentially relationships of curlleaf mountain mahogany
the same area as the burned stand. Historic fire (Cercocarpus ledifolius Nutt.) communities in
lines and burns are often clearly demarcated by the Utah and implications for management. Great
sharp boundaries of living stands. The wood is Basin Naturalist 51: 153-66.
rarely consumed by fire and often persists for
decades after, offering precise information about Dealy, J.E. 1975. Ecology of curlleaf mountain
past distribution and age classes. With changes in mahogany (Cercocarpus ledifolius Nutt.) in
fire intensity and return interval, usually related to eastern Oregon and adjacent areas. PhD.
increases in exotic annuals, curl-leaf mountain dissertation. Oregon State University, Corvallis,
mahogany in many areas is decreasing and is OR.
increasingly restricted to higher elevations, steeper
slopes, and cooler aspects (Ross 1999). Furniss, M.M. and W. F. Barr. 1969. Bionomics of
Anacamprodes clivinaria profanata on mountain
Growth and Management.—Although curl-leaf mahogany in Idaho. Agricultural Experiment
mountain mahogany is very long lived and Station Research Bulletin 73. University of
eventually assumes a tree form, young specimens Idaho, Moscow, ID. 24 p.
grow quickly as multistemmed shrubs. Stands are
typically composed of a single age class, perhaps Furniss M.M., D.C. Ferguson, K.W. Boget, J.W.
with younger specimens on the edges. It may be Burkhardt, A.R. Tiedemann, and J.L.
readily grown from seed for transplanting, although Oledmeyer. 1988. Taxonomy, life history, and
young transplants are attractive to rodents, ecology of a mountain mahogany defoliator,
lagomorphs, and large game. Seed collection is Stamnodes animata Pearsall in Nevada. Fish
easily done, although the hairy styles are a strong and Wildlife Research 3, USFWS, Washington,
respiratory and skin irritant. Management has D.C.
focused on increasing the forage production of old,
browsed stands via clipping and pruning (Davis Garrison, G.A. 1953. Effects of clipping on some
and Brotherson 1991), although this may result in range shrubs. Journal of Range Management 6:
tree mortality (Garrison 1953, Thompson 1970). 309-317.
Benefits.—Curl-leaf mountain mahogany has Kraft, G.F. 1960. Insects affecting bitterbrush and
found extensive use in the past as a fuel and other range plants. Office Report. Entomology
charcoal source. The very hard, beautiful wood Dept., Oregon State University, Corvallis, OR.
has been used for lathe work and parts for musical 55 p.
instruments (Ross 1999). Living stands provide
significant blue grouse nesting habitat (Stauffer Lis, R. 1993. Cercocarpus Mountain Mahogany.
and Petersen 1986). It is one of the few species that In: J.C Hickman, ed. 1993. The Jepson Manual:
meet protein requirements for wintering deer higher plants of California. U.C. Press, Berkeley
(Welch and McArthur 1979) and is heavily favored and Los Angeles, CA.
by bighorn sheep in summer (Rominger et al.
1988). Where present in even small amounts in Martin, F.L. 1950. A revision of Cercocarpus.
sagebrush, curl-leaf mountain mahogany allows Brittonia 7: 91-111.
greater wildlife diversity and abundance than
would otherwise occur (Furniss et al. 1988). It is Rominger, E.M., A.R. Dale, and J.A. Bailey. 1988.
also used increasingly as a commercial, residential, Shrubs in the summer diet of Rocky Mountain
and highway landscaping species. bighorn sheep. Journal of Wildlife Management
52: 47-50.
176
ledifolius Nutt.) in two adjacent Sierran and LaSal National Forest, USDA Forest Service
Great Basin mountain ranges. PhD. Dissertation, Range Improvement Notes, Vol. 15, No. 3.
University of Nevada, Reno.
USDA USFS. 1937. Range Plant Handbook.
Stauffer, D.F. and S.R. Petersen. 1986. Seasonal Washington D.C. U.S. Govt. Printing Office.
microhabitat relationships of blue grouse in p. 132–133.
southeastern Idaho. Great Basin Naturalist 46:
117-22. Welch, B.L. and E.D. McArthur. 1979. Nutritive
value of big sagebrush and other shrubs. In L.H.
Thompson, R.M. 1970. Experimental top pruning Stelter and others., eds. Proceedings, Shrub
of curlleaf mountain mahogany on the South establishment on disturbed arid and semi-arid
Horn Mountain, Ferron Ranger District, Manti- lands, Wyoming Fish and Game Dept. p. 9-22.
177
Cercocarpus montanus Raf. true mountain-mahogany
ROSACEAE
Stanley G. Kitchen
178
sterile. In contrast, hybrids formed with C. possible significance of rodents as secondary
ledifolius Nutt. spp. ledifolius, are quite fertile and dispersers of mountain-mahogany seeds is
backcross with parent populations (Stutz 1990). unknown.
179
References
Moore, T.C. 1963. A germination inhibitor in
Davis, J.N. 1990. General ecology, wildlife use, achenes of Cercocarpus montanus. Ecol. 44:
and management of the mountain mahoganies in 406-409.
the Intermountain West. In: K.L. Johnson, ed.
Proceedings of the Fifth Utah Shrub Ecology NatureServe Explorer. 2002. An online
Workshop: The Genus Cercocarpus; 1988 July encyclopedia of life. Version 1.6. NatureServe,
13-14; Logan, UT. College of Natural Arlington, VA, USA. http://www.natureserve.
Resources, Utah State University, Logan, UT. org/explorer [not paged].
p. 1-13.
Plummer, A.P., D.R. Christensen, and S.B.
Deitschman, G.H., K.R. Jorgensen, and A.P. Monsen. 1968. Restoring big-game range in
Plummer. 1974. Cercocarpus. In: C.S. Utah. Publication 68-3. Utah Division of Fish
Schopmeyer, tech. coord.. Seeds of Woody and Game, Ephraim, UT. 183 p.
Plants of the United States. Agricultural
Handbook 450. U.S. Department of Agriculture, Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
Forest Service, Washington, DC. p. 309-312. Viability of seed from thirty-two shrub and forb
species through fifteen years of warehouse
Ferguson, R.B. 1983. Use of wildland shrubs for storage. Great Basin Naturalist 4: 274-277.
wildland plantings in the Intermountain West.
In: S.B. Monsen and N. Shaw, comps. Managing Stutz, H.C. 1990. Taxonomy and evolution of
Intermountain Rangelands—Improvement of Cercocarpus in the western United States. In:
Range and Wildlife Habitats; Proceedings of K.L. Johnson, ed. Proceedings of the Fifth Utah
Symposia; 1981 September 15-17; Twin Falls, Shrub Ecology Workshop: The Genus
ID and 1982 June 22-24; Elko NV. General Cercocarpus; 1988 July 13-14; Logan, UT.
Technical Report INT-157. U.S. Department of College of Natural Resources, Utah State
Agriculture, Forest Service, Intermountain University, Logan, UT. p. 15-25.
Research Station, Ogden, UT. p. 136-147.
Turley, D. 2000. Effects of browsing on true
Hoeppel R.E. and A.G. Wollum. 1971. mountain mahogany (Cercocarpus montanus
Histological studies of ectomycorrhizae and root Raf.) MS Thesis. Brigham Young University.
nodules from Cercocarpus montanus and Provo, UT. 115 p.
Cercocarpus paucidentatus. Canadian Journal of
Botany 49: 1315-1318. Walker, S.C. and D. Turley. 1999. Characteristics
of mountain mahogany (Cercocarpus) species
Kitchen, S.G. and S.E. Meyer. 1990. Seed and hybrids in Utah hybrid zone. In: E.D.
dormancy in two species of mountain-mahogany McArthur, W.K. Ostler, and C.L. Wambolt,
(Cercocarpus ledifolius and Cercarpus comps. Proceedings: Shrubland Ecotones; 1998
montanus). In: K.L. Johnson, ed. Proceedings of August 12-14; Ephraim, UT. Proceedings
the Fifth Utah Shrub Ecology Workshop: The RMRS-P-11. U.S. Department of Agriculture,
Genus Cercocarpus; 1988 July 13-14; Logan, Forest Service, Rocky Mountain Research
UT. College of Natural Resources, Utah State Station, Ogden, UT. p. 32-34.
University, Logan, UT. p. 27-42.
Waugh, W.J. 1990. Stagnation and decadence of
Kitchen, S.G., S.E. Meyer, G.R. Wilson, and R. Cercocarpus montanus in a southeastern
Stevens. 1989. Addition of Cercocarpus Wyoming big game exclosure. In: K.L. Johnson,
montanus—true mountain-mahogany to the ed. Proceedings of the Fifth Utah Shrub Ecology
Rules. Association of Official Seed Analysts Workshop: The Genus Cercocarpus; 1988 July
Newsletter 63: 28-30. 13-14; Logan, UT. College of Natural
Resources, Utah State University, Logan, UT. p.
Marshall, K.A. 1995. Cercocarpus montanus. In: 89-95.
U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Laboratory, Fire Effects Information System, Goodrich. 1987. A Utah Flora. Brigham Young
http://www.fs.fed.us/database/feis/ 17 p. University Print Services, Provo, UT. 894 p.
180
Cereus greggii Engelmann night-blooming cereus
CACTACEAE
Juanita A. R. Ladyman
typically less than 1.5 m. The gray-brown stems
are four to five ribbed and approximately 1.25 cm
in diameter. There are 11 to 13 dark spines that
tend to be swollen at the base, per areole. The
lower spines may have some white coloration. The
flower is particularly remarkable, having waxy
white, pointed “petals” with numerous white to
yellow-tipped stamens. It is fragrant and up to 5
cm in diameter and 15 cm long. The orange-red,
oblong fruit is up to 7.6 cm long and 3.8 cm in
diameter with short spines. The root is a fleshy
taproot and typically weighs 2.3 to 6.8 kg
(Weniger 1991, Earle 1963, Vines 1960). In old
plants it may be up to 0.6 m in diameter and weigh
125 lbs (Weniger 1991). In 1919 a specimen with
purplish flowers was collected in Organ, New
Mexico, but since then only plants with white
flowers have been recorded (Weniger 1969). There
are two varieties of night-blooming cereus, Cereus
greggii var. greggii and Cereus greggii var.
transmontanus. Variety transmontanus differs
from the typical variety in that the flowers are
larger, approximately 7.5 cm in diameter, and the
hypanthium is particularly spiny, and the spines
generally 1.5 to 3.0 cm long (Benson 1982, Martin
and Hutchins 1980). The hypanthium is the cup-
like enlargement of the floral axis below the calyx
Illustration credit: Tonia Masaood that surrounds the ovary. The hypanthium of
variety greggii is covered by minute,
General Description.—The common names for inconspicuous spines and the flowers are smaller.
Cereus greggii are night-blooming cereus, Texas These varieties are in need of further study
night-blooming cereus, Arizona queen-of-the- (NMNPPAC 1984). Some sources (Natural
night, reina-de-la-noche, sweet-potato cactus, and Resources Conservation Service 2003) maintain
deer horn cactus. It is a perennial, succulent shrub that the synonym, Peniocereus greggii (Engelm.)
(Vines 1960). Night-blooming cereus is a Federal Britt. & Rose is the correct name. Night-blooming
Species of Concern and is listed as endangered by cereus provides an interesting evolutionary link. It
the State of New Mexico (Sivinski and Lightfoot appears to possess anatomical characteristics that
1995, NMRPTC 2002). It is threatened by illegal are relictual from its nonsucculent, woody,
collection throughout its range. Although this plant Pereskia-like ancestor (Mauseth and others 1998).
is cryptic and may be more common than verified
documentation suggests, many occurrences of Range.—Night-blooming cereus grows in
night-blooming cereus are known to have been southern New Mexico, southeastern Arizona,
extirpated due to collection pressures, and its western Texas, eastern Chihuahua, northeastern
range-wide abundance is believed to be declining. Durango, northern Zacatecas, and Coahuila,
Night-blooming cereus has erect or sprawling Mexico (Kearney and others 1960, Powell 1998,
stems that may grow to 2.4 m in length but are
181
Vines 1960). Benson (1982) defined the range of Growth and Management.—The stems generally
variety transmontanus as the Arizona and Sonoran grow up through, and later become supported by,
Desert that possibly also includes Hildalgo County shrubs. It is likely that the shrubs provide
in New Mexico, and that of variety greggii as the protection and a suitable microclimate for
Chihuahuan Desert of New Mexico, western Texas germination and subsequent plant development.
and Mexico. Any activity that reduces shrub cover is a threat to
night-blooming cereus. Therefore, fire is a
Ecology.—Night-blooming Cereus grows in dry potential hazard, although the direct effect of fire
alluvial soils at elevations between 370 and 1,500 on night-blooming cereus is not well documented.
m. Again, the varieties are distinguished by variety The substantial tuberous root may provide the
transmontanus growing at the lower end of the plant the means to recover if the above-ground
elevation range (300 to 1,050 m) and variety stems are killed, and therefore, it is likely that
greggii growing above 1200 m (Benson 1982). individual plants can tolerate light fires that do not
Night-blooming cereus generally grows in slightly decimate the shrubs with which they are associated
broken to level terrain in desert grassland or (Thomas 1991). More severe fires that scorch and
Chihuahuan desert scrub. Typically it grows in heat the soil are likely to be more detrimental.
sandy to silty gravelly soils on upper to mid Herbivores such as white-throated woodrats
bajadas among, depending upon the desert region, (packrats) and cactus borers (Cactobrosis
creosotebush (Larrea tridentata), mesquite fernaldialis) eat the stems, but new stems soon
(Prosopsis glandulosa), paloverde (Cercidium sprout from the tuberous root (ASDM 2002).
species), and knife-leaf condalia (Condalia Many populations throughout its range have been
species). It is also commonly associated with extirpated due to private and commercial
Ocotillo (Fouquieria splendens), tarbush collection, and the time required for population re-
(Flourensia cernua) and dropseed (Sporobolus establishment is not known. However, although
species). Habitat alteration such as that caused by slow growing, it is easily propagated by short stem
the high impact trampling of heavy grazing is cuttings and from seed. Seeds of many cacti
suspected to reduce population size (NMNPPAC species, including Cereus species, are subject to
1984). physiological dormancy. For example, seeds of
Cereus griseus were dormant at maturity but
Reproduction.—As the common name suggests, germinated after 8 weeks in a dry environment
night-blooming cereus blooms only at night. Just (Baskin and Baskin 2001). Cereus griseus seeds
after dark the flowers open in a series of “jerks” have an absolute light requirement for germination
(Benson 1982). Individual flowers only bloom for (Baskin and Baskin 2001). If the plant is dug up
one night. The total flowering period of plants for transplantation it is actually difficult to keep
within a population is also brief and generally all alive (Weniger 1969). Apparently the root is
flowers bloom within a 2-day period. However, in susceptible to fungus. Damp soil has to be
unusual circumstances, flowers on different avoided, but even if root rot is prevented, the plant
individuals within a population may bloom tends to decline over a few years if kept in a pot or
periodically over a period of almost one month. in a greenhouse (Weniger 1969).
Flowering occurs in late May into July depending
on geographic area (Earle 1963, Epple 1995, Benefits.—Night-blooming Cereus plants have
Kearney and others 1960). The fragrance of the commercial value especially among cactus
flowers is pleasant, and humans can detect the collectors (Epple 1995). Not only is it an
perfume of the flowers up to 30 m away from the aesthetically desirable cactus, but it has
plant (Benson 1982, Epple 1995, Vines 1960). considerable medicinal value. The root is generally
Flowers are pollinated principally by hawk moths believed to be the organ that has medicinal
(ASDM 2002) and possibly other night feeding properties (Moerman 1998, Moore 1989, Powell
insects (Epple 1995) that search for nectar. The 1998). However, the stems also have the same
flowers are not self-fertile and must be cross- properties although “more feebly” (Moerman
pollinated. Therefore the hawk moths (Sphingidae) 1998, Moore 1989, Powell 1998). Active
must fly hundreds of yards between the sparsely ingredients include penicerol, viperidone, desoxy
distributed plants (ASDM 2002). In areas where viperidone, viperidinone, β-sitosterol and, most
pesticides are heavily used for agriculture, the likely, caffeine (Moore 1989). It is likely that
hawkmoth populations are devastated, and most of night-blooming cereus is not unique in its
the flowers on plants in adjacent natural habitat medicinal value as other species of Cereus contain
fail to fruit (ASDM 2002). similar active ingredients (Ecdybase 2002). The
182
commercial toll on this species has not been Mauseth, J.D., T. Terrazas, and S. Loza-Cornejo.
established, but wild-harvested material is 1998. Anatomy of relictual members of
available on the Internet (Pacific West Botanicals subfamily Cactoideae, IOS Group 1a
2002). It is called “pain in the heart” by the Death (Cactaceae). Bradleya 16: 31-43.
Valley Shoshones (Moore 1989). This tribe
presumably uses it in a manner similar to Native Moerman, D.E. 1998. Native American ethno-
Americans of Nevada who ingest an infusion of botany. Timber Press, Portland, OR. 927 p.
the roots as a cardiac stimulant (Moerman 1998).
Other Native Americans have used a decoction of Moore, M. 1989. Medicinal plants of the desert
the roots for diabetes, the seedpods mixed with and canyon west. Museum of New Mexico
deer fat as a salve for sores, and the cut slices of Press, Santa Fe, NM. 184 p.
root as an externally applied cure for chest colds
(Moerman 1998, Vines 1960). The fruits, flowers, Natural Resources Conservation Service. 2003.
young stalks, and roots have been eaten for food Plants profile: Peniocereus greggii (Engelm.)
(Moerman 1998). This use of the root may account Britt. & Rose. http://plants.usda.gov/ [not
for the common name “sweet potato cactus.” paged].
Chewing the raw root has been reported to quench
thirst. Birds eat the seed and contribute to seed New Mexico Native Plants Protection Advisory
dispersal (Vines 1960). Committee. 1984. A handbook of rare and
endemic plants of New Mexico. University of
References New Mexico Press, Albuquerque, NM. 291p.
Arizona Sonora Desert Museum. 2002. New Mexico Rare Plant Technical Council. 2002.
Peniocereus greggii. Internet web site: NM Rare Plants Web Page: http://nmrareplants.
http://www.desert unm.edu/reports/ pengre.htm.
museum.org/books/nhsd_peniocereus.html#top.
Pacific West Botanicals. 2002. Internet web site:
Baskin, C.C. and J.M. Baskin. 2001. Seeds, http://www.pacificwestbotanicals.com/
ecology, biogeography, and evolution of Night_Blooming_Cereus.html.
dormancy and germination. Academic Press,
New York. 666 p. Powell, A. M. 1998. Trees & shrubs of Trans-
Pecos and adjacent areas. University of Texas
Benson, L. 1982. The cacti of the United States University Press, Austin, TX. 498 p.
and Canada. Stanford University Press,
Stanford, CA. p. 588-593. Sivinski, R. and K. Lightfoot 1995. Inventory of
rare and endangered plants of New Mexico.
Earle, W.H. 1963. Cacti of the Southwest. Desert Miscellaneous Publication 4. New Mexico
Botanical Garden, Phoenix, AZ. 210 p. Forestry and Resources Conservation Division,
Energy, Mineral and Natural Resources
Ecdybase. 2002. Internet Web Page: http:// Department, Santa Fe, NM.
ecdybase.org/index.php?row=308&action=
browse. Thomas, P.A. 1991. Response of succulents to
fire: A review. International Journal of Wildland
Epple, A.O. 1995. A field guide to the plants of Fire 1(1): 11-22.
Arizona. Falcon Press Publishing Co., Helena,
MT. 347 p. Vines, R.A. 1960. Trees, shrubs, and woody vines
of the Southwest. Sixth printing 1986.
Kearney, T.H., R.H. Peebles, J.T. Howell, and E. University of Texas Press. Austin, TX. 1104 p.
McClintock. 1960. Arizona flora. 2d ed.
University of California Press. Berkeley, CA. Weniger, D. 1969. Cacti of the Southwest.
1,085 p. University of Texas Press, Austin, TX. 249 p.
Martin, W.C. and C.R. Hutchins. 1980. A Flora of Weniger, D. 1991. Cacti of Texas and neighboring
New Mexico. Strauss & Cramer, Hirschberg, states. University of Texas Press, Austin, TX.
Germany. 2,592 p. 356 p.
183
Cestrum diurnum L. day jasmine
SOLANACEAE
John K. Francis
from cultivation in at least Florida, southern Texas,
Hawaii, Puerto Rico, Guam, and American Samoa
(Florida Exotic Plants Council 2001, Institute of
Pacific Islands Forestry 2001).
184
germinated between 7 and 30 days after sowing. Calcinogenic glycosides. http://www.ansci.
edu/plants/toxicagents/calglyco.htm. 2 p.
Growth and Management.—Day jasmine rarely
reaches more than 4.5 m in height and 8 cm in Florida Exotic Plants Council. 2001. Cestrum
trunk diameter (Little and others 1974). Early diurnum L., Solanaceae/nightshade family.
growth is relatively fast and life spans are www.fleppc.org/pdf/Cestrum%20diurnum.pdf.
relatively short (5 to 20 years). Although seeds are 2 p.
still sold by regional seed houses and ornamental
seedlings are offered by nurseries, nothing is Howard, R.A. 1989. Flora of the Lesser Antilles,
published about propagation and management. Leeward and Windward Islands. Dicotyledoneae.
With a plant so aggressive, few nursery and Part 3. Vol. 6. Arnold Arboretum, Harvard
planting problems seem likely. Because the species University, Jamaica Plain, MA. 658 p.
is a weed in many situations, control may be
desirable. In the absence of control guidelines, Institute of Pacific Islands Forestry. 2001. Plant
cutting followed by herbicide treatment of the threats to Pacific Ecosystems: Cestrum diurnum
sprouts is suggested. L., Solanaceae. www.hear.org/pier/cediu.htm.
2 p.
Benefits.—Day jasmine has long been planted as
an ornamental for its pleasing appearance, Island Resource Foundation. 2001. Threatened and
moderate size, ease of establishment, and fragrant endangered birds of the insular Caribbean: plain
flowers. The plant has the drawback of being pigeon. www.irf.org/binornat.htm. 14 p.
poisonous to livestock. The leaves contain a
calcinogenic glycoside called 1,25- Liogier, H.A. 1995. Descriptive flora of Puerto
dihydroxycholecalciferol that leads to a vitamin D Rico and adjacent islands. Vol. 4. Editorial de
toxicity that results in elevated serum calcium and la Universidad de Puerto Rico, San Juan, PR.
deposition of calcium in soft tissues. Fifteen to 30 617 p.
percent of day jasmine leaves in an animal’s diet is
sufficient to cause symptoms (Animal Science at Little, E.L., Jr., R.O. Woodbury, and F.H.
Cornell University 2001). However, the fruits of Wadsworth. 1974. Trees of Puerto Rico and the
day jasmine are one of the three foods that make up Virgin Islands. Vol. 2. Agriculture Handbook
the bulk of the diet of the endangered plain pigeon 449. U.S. Department of Agriculture,
(Columba inornata) in Puerto Rico (Island Washingtion, DC. 1,024 p.
Resource Foundation 2001).
Neal, M.C. 1965. In gardens of Hawaii. Special
References Publication 50. Bernice P. Bishop Museum
Press, Honolulu, HI. 924 p.
Animal Science at Cornell University. 2001.
185
Cestrum laurifolium L’Hér. dama de noche
SOLANACEAE
John K. Francis
186
nursery, planting, or management experience has References
been published. Protection of forests from fires and
development is recommended to encourage the Grisebach, A.H.R. 1963. Flora of the British West
formation and development of natural stands. Indies. J. Cramer-Weinheim, New York. 789 p.
Benefits.—Dama de noche contributes to the Howard, R. A. 1989. Flora of the Lesser Antilles,
biodiversity of the forests where it grows, helps Leeward and Windward Islands. Vol. 6. Arnold
protect the soil, and furnishes food and cover for Arboretum, Harvard University, Jamaica Plain,
wildlife. The stems are generally too small to use MA. 658 p.
for fuel.
Liogier, H.A. 1995. Descriptive flora of Puerto
Rico and adjacent islands. Vol. 4. Editorial de la
Universidad de Puerto Rico, San Juan, PR.
617 p.
187
Chamaebatiaria millefolium (Torr.) Maxim. fern bush
ROSACEAE
Nancy L. Shaw
Hitchcock and others 1961, Welsh and others
1987). Chromosome number is x = n = 9
(McArthur and Sanderson 1985). Variability
within the species appears low and hybridization
with other species has not been reported.
188
3-month prechill (Phillips 1949, Shaw and Hurd in Cormophyta 15. New observations on
2001). The optimum temperature for germination cyanogenesis in Rosaceae. Planta Medica 41:
of southwestern populations is 18 to 26 °C 313-327.
(Phillips 1949).
Hawksworth, F.G. and R.L. Mathiasen. 1978.
Growth and Management.—Seedlings establish Hosts of juniper mistletoe at Walnut Canyon
from shallow seeding on well-drained soils where National Monument, Arizona, USA. Great Basin
vegetative competition is limited (Shaw and Hurd Naturalist 38: 89.
2001). Fall seeding or artificial prechilling is
required for seeds of northern populations. Hickman, J.C., ed. 1993. The Jepson manual of
Seedlings develop rapidly if adequate water is higher plants of California. University of
available. Bareroot seedlings can be lifted California Press, Berkeley, CA. 1,400 p.
following one growing season. Irrigated plants
begin flowering during the second year (Shaw and Hitchcock, C.L., A. Cronquist, M. Ownbey, and
Hurd 2001). Unlike its namesake genus, J.W. Thompson. 1961. Vascular plants of the
Chamaebatia Benth., fern bush is not nodulated by Pacific Northwest. Part 3: Saxifragaceae to
nitrogen-fixing actinomycetes (McArthur and Ericaceae. University of Washington Press,
Sanderson 1985). Fern bush is a rare host of Seattle, WA. 614 p.
Phoradendron juniperinum A. Gray, the juniper
mistletoe (Hawksworth and Mathiasen 1978). McArthur, E.D. and S.C. Sanderson 1985. A
cytotaxonomic contribution to the western North
Benefits.—Fern bush provides cover for many American rosaceous flora. Madroño 32: 24-28.
organisms. It is sometimes browsed by mule deer
(Odocoilus hemionus Rafinesque), sheep (Ovis Merkle, J. 1952. An analysis of a pinyon-juniper
spp.), and goats (Capra hircus), but it receives community at Grand Canyon, Arizona. Ecology
little use by cattle (Bos spp.). First cultivated in 33:385-384.
1878 (Rehder 1940), fern bush is a valued
ornamental due to its showy flowers, long Mozingo, H.N. 1987. Shrubs of the Great Basin.
flowering period, and fernlike foliage (Hitchcock University of Nevada Press, Reno, NV. 342 p.
and others 1961, Phillips 1949). Mass plantings,
xeriscapes, screens, hedges, and specimen plants Phillips, J. 1949. Southwestern landscaping with
thrive in full sun (Phillips 1949). Native native plants. Museum of New Mexico Press,
Americans used a tea made from its leaves for Santa Fe, NM. 140 p.
treatment of stomachaches (Mozingo 1987). Plants
produce cyanic acids in their tissue (Fikensher and Rehder, A. 1940. Manual of cultivated trees and
others 1981). shrubs. Macmillan Publishing Company, Inc.,
New York. 996 p.
References
Shaw, N.L. and E.G. Hurd. 2001 Chamaebatiaria
Eggler, W.A. 1941. Primary succession on millefolium (Porter) Maxim. fern bush, desert
volcanic deposits in southern Idaho. Ecological sweet. In: F.T. Bonner and R.G. Nisley, eds.
Monographs 3: 277-298. Woody plant seed manual. Agriculture
Handbook. U.S. Department of Agriculture,
Everett, P.C. 1957. A summary of the culture of Forest Service, Washington, D.C.
California plants at the Rancho Santa Ana http://wpsm.net/index.html. 6 p.
Botanic Garden. Rancho Santa Ana Botanic
Garden, Claremont, CA. 233 p. Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Goodrich, eds. 1987. A Utah flora. Great Basin
Fikenscher, L.H., R. Hegnauer, and H.W.L. Naturalist Memoirs 9. Brigham Young
Ruijgrok. 1981. Distribution of hydrocyanic acid University, Provo, UT. 894 p.
189
Chenopodium oahuense (Meyen) Aellen. 'aheahea
CHENOPODIACEAE
190
'Aheahea prefers partial to full sun with light
watering. The species may have potential in native References
habitat restoration as a nurse species or to enhance
Anonymous. 1996. Hawaiian use of native plants.
microclimatic understory conditions in favor of
http://www.k12.hi.us/~waianaeh.
woody plant establishment.
hawaiianstudies/nplants6.html
Benefits.―The leaves and plant tips are used for
Ching, F. 2002. Ulu Ka Hoi. http://kms.
greens, and may be eaten like spinach, wrapped in
kapalama.ksbe.edu/projects/yellow/default.
ti [Cordyline fruticosa (L.) A. Chev.] leaves and
html. [not paged].
cooked on hot coals. Hawaiians used the plant to
make a poultice for minor wounds during battle.
Kawaharada, D. 2002. Visiting Mokumanamana.
Leaves were pounded or crushed and applied like
http://explorers.bishopmuseum.org. [not paged].
rubbing alcohol, the healing effect due to the
presence of chlorophyll in the leaf (Ching 2002).
Kessing, J., L. Martin, and R.FL Mau. 1993.
Hawaiians also used the dark red bark pounded in
http://www.extento.hawaii.edu/kbase/crop/
conjunction with other ingredients to create a
Type/Nysius.htm
concoction that would enhance and beautify the
skin of a newborn baby when ingested by a
Lamb, S.H. 1981. Native trees and shrubs of the
pregnant or nursing mother (Anonymous 1996).
Hawaiian Islands. The Sunstone press, Santa Fe,
Two limbs of the shrub wood were connected to
New Mexico. 158 p.
create a "makua mano," or shark hook, and in this
fashion, 'aheahea was used to catch sharks (Ching
Sherry, K., J.M. Castillo, and R.B Shaw. 1999.
2002). The native shrub is also of great importance
Effects of wildfire on vegetation and rare plants
to a certain rare beetle, Rhyncogonus biformis, that
in arid montane shrublands. Pohakaloa training
lives on Necker Island. The beetle is nocturnal and
area, Hawaii. 1999 Hawaii Conservation
emerges at night where it feeds soley on the leaves
Conference, Honolulu, HI.
of the 'aheahea (Kawaharada 2002).
Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
Detrimental Effects.―The flower heads are host
1990. Manual of the Flowering Plants of
plants for Nysius nemorivagus, a white Lygaeid
Hawaii. University of Hawaii Press, Honolulu,
bug that can become a nuisance pest for certain
HI. 1,854 p.
agricultural crops such as cabbage, cucumbers,
potatoes and all types of squash (Kessing and
others 1993).
191
Chiococca alba (L.) A.S. Hitchc. West Indian snow-berry
RUBIACEAE
John K. Francis
in Florida, the Bahamas, through the Greater and
Lesser Antilles, in Mexico, Central America, and
northern South America. The species is not known
to have naturalized outside its native range.
192
Plants develop slowly in the nursery, requiring 3 Technical Report SO-58. U.S. Department of
months to grow large enough to prick into pots and Agriculture, Forest Service, Southern Forest
about an additional year to reach outplanting size. Experiment Station, New Orleans, LA. 331 p.
Seedlings and sections of older plants can be dug
up in the forest and transplanted successfully Bailey, L.H. 1941. The standard cyclopedia of
(Workman 1980). Once established, West Indian horticulture. Vol. 1. The MacMillan Company,
snow-berry needs little maintenance. New York, NY. 1,200 p.
Benefits.—West Indian snow-berry is used to a Howard, R.A. 1989. Flora of the Lesser Antilles,
limited extent as an ornamental in naturalistic Leeward and Windward Islands. Dicotyledoneae.
landscaping (Workman 1980) and is sometimes Part 3. Vol. 6. Arnold Arboretum, Harvard
cultivated as a flowering vine in greenhouses University, Jamaica Plain, MA. 658 p.
(Bailey 1941). Being very common and
widespread, the species contributes to the Liogier, H.A. 1997. Descriptive flora of Puerto
biodiversity of many Neotropical forests. It is Rico and adjacent islands. Vol. 5. Editorial de la
grazed by cattle, goats and certainly wild ruminants Universidad de Puerto Rico, San Juan, PR.
and produces fruits that are eaten by birds and 436 p.
other animals. The roots of West Indian snow-
berry are used as a purgative, diuretic, vomitive, Liogier, H.A. 1990. Plantas medicinales de Puerto
and antidiarrhetic in herbal medicine (Liogier Rico y del Caribe. Iberoamericana de Ediciones,
1990). Inc., San Juan, PR. 566 p.
References Workman, R.W. 1980. Growing native. The
Sanibel-Captiva Conservation Foundation, Inc.,
Acevedo-Rodriguez, P. and R.O. Woodbury. 1985. Sanibel, FL. 137 p.
Los bejucos de Puerto Rico. Vol. 1. General
193
Chromolaena geraniifolia (Urban) King & H.E. Robins. geranium-leaf eupatorium
ASTERACEAE
John K. Francis
194
References
195
Chromolaena odorata (L.) King & H.E. Robins. Christmas bush
ASTERACEAE
John K. Francis
196
mechanical or hand cutting followed by herbicide Eupatorium odoratum as a weed in teak
treatment. Partial control can be obtained through (Tectona grandis) forest. Lembaga Penelitian
the use of aggressive cover crops. Relatively good Hasil hutan 312. Lopora, Indonesia. 25 p.
biological control has been obtained with
Pareuchaetes pseudooinsulata Rego Barros Howard, R.A. 1989. Flora of the Lesser Antilles,
(Lepidoptera) in Guam and several other Pacific Leeward and Windward Islands. Vol. 6. Arnold
islands (Pacific Island Ecosystems at Risk 2001). Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
Detriments and Benefits.—Invasion of Christmas
bush has been disastrous by seriously suppressing Kaleta, D., S.R. Ghosh, and C.N. Saikia. 1999.
native species in disturbed forests and pastures in Medium density particle board from weeds.
the tropics outside its native range. The shrub is Journal of Scientific and Industrial Research
reported to be highly allelopathic to nearby 58(9): 705-710.
vegetation (Muniappan 1994), a fact that has been
demonstrated in controlled studies (Sahid and Liogier, H.A. 1990. Plantas medicinales de Puerto
Sugau 1993). Christmas bush reduces the diameter Rico y del Caribe. Iberoamericana de Ediciones,
growth of teak in infested plantations (Daryono and Inc., San Juan, PR. 566 p.
Hamzah 1979). It was thought to be useful in the
control of Imperata grass and for this reason was Liogier, H.A. 1997. Descriptive flora of Puerto
deliberately introduced into the Ivory Coast, but Rico and adjacent islands. Vol. 5. Editorial de la
the results were disappointing (Binggeli 1999). Universidad de Puerto Rico, San Juan, PR.
Because of the abundance of dead leaves and dry 436 p.
shoots, Christmas bush stands are a fire hazard
(Muniappan 1994). Cattle do not eat Christmas M’Boob, S.S. 1991. Preliminary results of a
bush; however, it is browsed by white-tailed deer survey and assessment of Chromolaena odorata
(Meyer and others 1984). In herbal medicine, leaf (Siam weed) in Africa. Biotropica Special Pub.
extracts with salt are used as a gargle for sore 44: 51-55.
throats and colds. It is also used to scent aromatic
baths (Liogier 1990). Extracts of Christmas bush Meyer, M.W., R.D. Brown, and M.W. Graham.
have been shown to inhibit or kill Neisseria 1984. Protein and energy content of white-tailed
gonorrhoeae (the organism that causes gonorrhoea) deer diets in the Texas coastal bend. Journal of
in vitro (Caceres and others 1995) and to accelerate Wildlife Management 48(2): 527-534.
blood clotting (Triratana and others 1991). A
satisfactory medium-density particleboard was Muniappan, R. 1994. Chromolaena odorata (L.)
prepared from Christmas bush stems (Kaleta and R.M. King and H. Robinson. In: R. Labrada,
others 1999). During fallows between cultivation, J.C. Caseley, and C. Parker, eds. Weed
Christmas bush adds copious amounts of organic management for developing countries. Plant
matter to the soil and may reduce the populations Production and Protection Paper 120. Food and
of nematodes (M’Boob, 1991). It is also useful as Agriculture Organization of the United Nations,
mulch for row crops (Swennen and Wilson 1984). Rome. 93-94.
197
Swennen, R. and G.F. Wilson. 1984. In-situ mulch Triratana, T., R. Suwannuraks, and W.
production for plantain. Banana Newsletter 7: Naengchomnong. 1991. Effect of Eupatorium
20-22. odoratum on blood coagulation. Journal of
Medical Association of Tailand 74(5): 283-287.
198
Chrysobalanus icaco L. coco-plum
CHRYSOBALANACEAE
John K. Francis
199
Seed dispersion is presumed to be by gravity, hypoglycemic effects. Leandra 6-7: 7, 63-75.
water, birds, bats, domestic animals, and humans.
Natural reproduction may be sparse to abundant. Francis, J.K. and A. Rodríguez. 1993. Seeds of
Artificial reproduction is usually by seeds with Puerto Rican trees and shrubs: second
plants grown as potted seedlings. Apical and basal installment. Research Note SO-374. U.S.
semiwoody, leafy cuttings treated with hormones Department of Agriculture, Forest Service,
will root in 6 to 8 weeks in mist bed conditions Southern Forest Experiment Station. New
(Vargas-Simon and others 1997). The best Orleans, LA. 5 p.
treatment tested was Indol Acidic Acid (IAA) at
5,000 ppm applied to apical stem cuttings. Howard, R.A. 1988. Flora of Lesser Antilles. Vol.
Adventitious roots arose from the vascular 4. Arnold Arboretum, Harvard University,
cambium, and lateral roots arose from the Jamaica Plain, MA. 673 p.
paricycle.
Liogier, H.A. 1985. Discriptive flora of Puerto
Growth and Management.—Growth is slow in Rico and adjacent islands. Vol. 1. Editorial de la
the nursery and the field, so new plantings of coco- Universidad de Puerto Rico, San Juan, PR. 352
plum must be protected from both herbaceous and p.
woody competition.
Little, E.L., Jr., R.O. Woodbury, and F.H.
Benefits.—The fruits are edible raw and can be Wadsworth. 1974. Trees of Puerto Rico and the
made into preserves. Coco-plum seeds, which have Virgin Islands. Vol. 2. Agriculture Handbook
a high oil content, are also edible. The wood is 449. U.S. Department of Agriculture,
light brown, hard, and heavy (specific gravity 0.8) Washington, DC. 1,024 p.
and is used for fuel and rustic construction (Little
and others 1974). Various parts of the plant have Paradis, G. 1987. Concerning the division of the
been used in folk medicine. The species is known genus Chrysobalanus into species and
to have hypoglycemic effects (Costa 1977). It is a subspecies in West Africa (Chrysobalanaceae).
honey plant and furnishes food for wildlife. Coco- A Sciences Naturelles 45(3,4): 246-254.
plum is used in the Eastern and Western
Hemespheres as an ornamental. Another important Vargas-Simon, G., G. Arellano-Ostoa, and E.
benefit from the species is for dune and soil García-Villanueva. 1997. Propagación por
stabilization. estacas con hojas de icaco (Chrysobalanus icaco
L.) y anatomía del enraizamiento. Proceedings of
References the International Society for Tropical
Horticulture 41: 264-269.
Costa, O. de A. 1977. Brazilian plants with
200
Chrysophyllum oliviforme L. satinleaf
SAPOTACEAE
John K. Francis
201
information is not available. Misted hardwood capitata (Wiedemann). University of Florida.
cuttings and air layers, both treated with 0.3 http://creatures.ifas.ufl.edu/fruit/Mediterranean_
percent IBA (indol-butyric-acid), were tested as fruit-fly.htm. 19 p.
means of propagating the species vegetatively.
Thirty-five of 69 cuttings rooted in 7 months and Gilman, E.F. and D.G. Watson. 1993.
one of 39 air layers rooted in 6 months (author’s Chrysophyllum oliviforme, Satinleaf. Fact Sheet
observation). Ornamental production is by seed and ST-166. Environmental Horticulture
semihardwood cuttings under mist (Gilman and Department, Florida Cooperative Extension
Watson 1993). Service, University of Florida. 2 p.
Growth and Management.—Growth is relatively Institute of Food and Agricultural Sciences. 2002.
slow at all stages, and the plants are relatively Flowering and shade trees (A-E). University of
long-lived. Natural reproduction can probably be Florida Extension http://miami-dade.ifas.ufl.
encouraged by disturbance under low-density edu/programs/fyn/publications/dtplfloweringtree
secondary forests near seed sources. s-a-e.htm. 7 p.
Benefits.—Satinleaf contributes to the aesthetics of Liogier, H.A. 1995. Descriptive flora of Puerto
the forests where it grows, helps protect the soil, Rico and adjacent islands. Vol. 4. Editorial de la
and furnishes food and cover for wildlife. It was Universidad de Puerto Rico, San Juan, PR.
listed as an important source of nectar for 617 p.
honeybees in the Dominican Republic (Marcano
Fondeur 1973). The foliage is probably unpalatable Little, E.L., Jr. and F.L. Wadsworth. 1964.
to ungulates; at least the key deer (Odocoilleus Common trees of Puerto Rico and the Virgin
virginianus ssp. clavium) will not eat it (Schaus Islands. Agric. Handb. 249. U.S. Department of
and others 2002). The fruits are edible and Agriculture, Washington, DC. 548 p.
sometimes used to make jelly (Little and
Wadsworth 1964). In areas of infestation, satinleaf Long, R.W. and O. Lakela. 1976. A flora of
is usually heavily attacked by Mediterranean fruit Tropical Florida. Banyon Books, Miami, FL.
fly (Ceratitis capitata (Wiedemann) (Department 962 p.
of Entomology and Nematology 2002). The wood,
which has a specific gravity of 0.9, is hard, heavy, Marcano Fondeur, E. de J. 1973. La flora apícola
and strong. It is used for construction in Cuba de la República Dominicana. http.//
(Little and Wadsworth 1964). Satinleaf has been marcano.freeservers.com/nature/estudios/apicola
extensively, but not intensively, planted as an /dicotsp.htm. 11 p.
ornamental in Florida, Hawaii, and elsewhere. It
makes a pretty addition to natural landscaping, a Nelson, G. 1996. The shrubs and woody vines of
good foundation plant, and an attractive lawn, Florida. Pineapple Press, Inc., Sarasota, FL.
street, and parking lot tree. Pruning may be 391 p.
necessary to maintain a single stem and to remove
drooping branches (Gilman and Watson 1993). Pacific Island Ecosystems at Risk. 2002.
Chrysophyllum oliviforme L., Sapotaceae.
References http://www.hear.org/pier3/choli.htm. 2 p.
Crowley, M. 2002. Florida’s native plants— Schaus, C., S. Wade and J. Dunan. 2002. Key deer
protected. Native Florida Heritage Web Ring. and plants they won’t eat. Monroe County
http://nsis.org/garden/pr-plants-c.html. 5 p. Extension Service, University of Florida.
http://monroe.ifas.ufl.edu/key_deer_plants.htm.
Department of Entomology and Nematology. 4 p.
2002. Mediterranean fruit fly—Ceratitis
202
Chrysothamnus nauseosus (Pallas ex Pursh) Britt. rubber rabbitbrush
ASTERACEAE
203
Ecology.—Rubber rabbitbrush establishes readily stable and disturbed plant communities. Its ability
on disturbed areas and can often become the to stabilize soil and reduce wind and water erosion
dominant vegetation on lands that have been preserves habitat for other shrubs, forbs, and
disturbed by fire, insects, vehicles, or heavy grasses. This shrub is an excellent plant for erosion
grazing. The species spans a broad elevation control because it has deep roots, produces heavy
gradient, extending from 150 to 2,700 m. with litter, and is able to establish on severe sites
annual precipitation generally in the 250 to 350 (Aldon and Pase 1981, McArthur and others 1995,
mm range. This shrub grows well in sandy, USDA 1937). Rubber rabbitbrush is browsed by
gravelly, or clay-alkaline soils, depending on the deer, pronghorn, and by domestic animals, but
subspecies or ecotype. Some subspecies have palatability varies greatly between subspecies and
broader ecological amplitudes than others, ecotypes. The white to grayish subspecies
allowing them to establish in a variety of habitats (albicaulis, hololoeucus, and salicifolius) are more
(McArthur and others 1979). Although it grows palatable than the green subspecies (gravelens and
vigorously, it does not exclude or inhibit consimilis) (Goodrich and others 1999 Hanks and
herbaceous species. On the contrary, production of others 1975). Unlike some forage shrubs, however,
herbaceous cover was greater where rubber rubber rabbitbrush is quite resilient to browsing.
rabbitbrush was present than when it was not In addition to food, C. nauseosus also provides
present (Frischknecht 1963). cover for small mammals, birds, and newly born
pronghorn (Yoakum 1986). In addition to its
Reproduction.—Rubber rabbitbrush blooms from ecological importance, people have also
August to October. Seeds mature beginning in recognized the chemical properties of C.
mid-October until late fall. In most conditions, nauseosus. Rubber rabbitbrush has been
rubber rabbitbrush is a heavy seed producer. Seed investigated as a possible source of rubber and
germination in rubber rabbitbrush is quite high, other chemicals (Hall and Goodspeed 1919;
and seed viability remains relatively high (65 Hegerhorst and others 1987). Some accessions, for
percent) for up to 3 year’s storage under ordinary example, produce up to 6 percent stem rubber and
warehouse storage conditions (Stevens and others 20 percent resin. Finally, some forms have
1981). When exposed to cool night (4.7 oC) and ornamental shrub potential (McArthur and others
warm day temperatures, seed germinates in about 1979, Weber and others 1985).
2 days (Weber and others 1985). Under
standardized single cool temperature conditions, References
however, seeds from lower elevations and latitudes
germinate more quickly (2 weeks as compared to Aldon, E.F. and C.P. Pase. 1981. Plant species
4) than those from higher elevations and latitudes adaptability on mine spoils, in the Southwest: a
(McArthur and others 1987, Meyer and others case study. Research Note RM-398. U.S.
1989). Seed (achene) size in rubber rabbitbrush Department of Agriculture, Forest Service,
appears to be correlated to habitat; subspecies Rocky Mountain Forest and Range Experiment
adapted to sandy sites have larger achenes (Meyer Station, Fort Collins, CO. 3 p.
1997). There are about 1,530 cleaned seeds per
gram (McArthur and others 1979). Anderson, L.C. 1966. Cytotaxonomic studies in
Chrysothamnus (Asterae, Compositae).
Growth and Management.—Once established, American Journal of Botany 53: 204-212.
wind-disseminated achenes allow rubber
rabbitbrush to spread easily and quickly (Meyer Anderson, L.C. 1986. Sympatric subspecies in
1997, Young and others 1984). It is fast growing, Chrysothamnus nauseosus. In: E. D. McArthur
reaching maturity in 2 to 4 years but relatively and B.L Welch, comps. Proceedings—
short-lived shrub, usually 5 to 20 years (personal symposium on the biology of Artemisia and
observation). It can be aerially seeded. Rubber Chrysothamnus; 1984 July 9-13, Provo, UT.
rabbitbrush can be troublesome on rangelands General Technical Report INT-200. U.S.
when it invades and occupies areas at high Department of Agriculture, Forest Service,
densities. It is a difficult to control because it is Intermountain Research Station, Ogden, UT: 98-
resistant to herbicides and has the ability to 103.
resprout from the crown (Cook and others 1965).
Anderson, L.C. 1995. The Chrysothamnus-
Benefits.—Rubber rabbitbrush can be an Ericameria connection (Asteraceae). Great
important structural and ecological component in Basin Naturalist 55: 84-88.
204
Cook, C.W., P.D. Leonard, and C.D. Bonham. McArthur, E.D., R. Stevens, and S.B. Monsen
1965. Rabbitbrush competition and control on 1995. Adaptation and success of big sagebrush
Utah rangelands. Bulletin 454. Utah and rubber rabbitbrush on disturbed sites. In:
Agricultural Experiment Station., Logan, UT. G.E. Schuman and G.F. Vance, eds. Decades
28 p. later: a time for reassessment, proceedings of the
12th Annual Meeting of the American Society
Frischknecht, N.C. 1963. Contrasting effects of big for Surface Mining and Reclamation; 1995 June
sagebrush and rubber rabbitbrush on production 3 -8, Gillette, WY. American Society for
of crested wheatgrass. Journal of Range Surface Mining and Reclamation, Princeton,
Management 16: 70-74. WV. p. 811-823.
Goodrich, S., E.D. McArthur, A. Huber, and J.E. Meyer, S.E. 1997. Ecological correlates of achene
Ott. 1999. Ungulate browsing of two mass variation in Chrysothamnus nauseosus
populations of rubber rabbitbrush. In: E.D. (Asteraceae). American Journal of Botany. 84:
McArthur, W.K. Ostler, and C.L. Wambolt, 471-477.
comps. Proceedings: shrubland ecotones; 1998
August 12-14; Ephraim, UT. Proc RMRS-P-11. Meyer, S.E., E.D. McArthur, and G.L. Jorgensen.
Ogden, UT: U.S. Department of Agriculture, 1989. Variation in germination response to
Forest Service, Rocky Mountain Research temperature in rubber rabbitbrush
Station: 196-199. (Chrysothamnus nauseosus:Asteraceae) and its
ecological implications. American Journal of
Hall, H.M. and T.H. Goodspeed. 1919. A rubber Botany 76: 981-991.
plant survey of Western North America. II.
Chrysil, a new rubber from Chrysothamnus Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
nauseosus. University of California Publications Viability of seed from thirty-two shrub and forb
in Botany 7: 183-264. species through fifteen years of warehouse
storage. Great Basin Naturalist 41: 274-277.
Hanks, D.L., E.D. McArthur, A.P. Plummer, B.C.
Giunta, and A.C. Blauer. 1975. U.S. Department of Agriculture, Forest Service.
Chromatographic recognition of some palatable 1937. Range plant handbook. United States
and unpalatable subspecies of rubber rabbitbrush Government Printing Office, Washington, D.C.
in and around Utah. Journal of Range 816 p.
Management 28: 144-148.
Weber, D.J., T. Davis, E.D. McArthur, and N.
Hegerhorst, D., D.J. Weber, and E.D. McArthur. Sankhla. 1985. Chrysothamnus nauseosus
1987. Resin and rubber content in (rubber rabbitbrush): mutiple-use shrub of the
Chrysothamnus. Southwestern Naturalist 32: desert. Desert Plants 7: 172-209.
475-482.
Yoakum, J. 1986. Use of Aremisia and
International Plant Names Index. 2002. Chrysothamnus by pronghorns. In: McArthur, E.
International Plant Names Index. D. and B. L Welch, comps. Proceedings—
xhttp//www.ipni.org. [not paged]. symposium on the biology of Artemisia and
Chrysothamnus; 1984 July 9-13, Provo, UT.
McArthur, E.D., A.C. Blauer, A.P. Plummer, and General Technical Report INT-200. U.S.
R. Stevens 1979. Characteristics and Department of Agriculture, Forest Service,
hybridization of important intermountain shrubs. Intermountain Research Station, Ogden, UT.
III. Sunflower family. Research Paper INT-220. p. 176-180.
U.S. Department of Agriculture, Forest Service
USDA Forest Service, Intermountain Forest and Young, J.A., R.A. Evans, and B.L. Kay. 1984.
Range Experiment Station, Ogden, UT. 82 p. Persistence and colonizing ability of rabbitbrush
collections in a common garden. Journal of
McArthur, E.D., S.E. Meyer, and D.J. Weber. Range Management 37: 373-377.
1987. Germination rate at low temperature:
rubber rabbitbrush population differences.
Journal of Range Management 40: 530-533.
205
Chrysothamnus parryi (Gray) Greene Parry rabbitbrush
ASTERACEAE
206
In subspecies monocephalus, leaf shape common species, such as rubber and low
can vary from linear-oblanceolate to spatulate. rabbitbrushes.
Like subspecies howardii, the upper leaves usually
extend beyond the inflorescence. One unique Reproduction.—Parry rabbitbrush flowers from
characteristic is that flower heads occur singly or July to September and seed matures in October
in pairs rather than forming racemes. Flower heads and November. Achenes are 5 to 6 mm long and
contain five or six disc flowers subtended by covered with long, shaggy, oppressed hairs. We
obscurely ranked involucral bracts. This presume there are about 1,600 cleaned achenes per
subspecies occurs in high mountains between 790 gram and that they have good fertility as is the
and 3,400 m in eastern California and western case for other Chrysothamnus species
Nevada. (Deitschmann and others 1974). Achenes are wind
Subspecies nevadensis is comprised of disseminated in late fall or winter. In addition to
small shrubs up to 60 cm in height. Leaves are sexual reproduction, Parry rabbitbrush has been
linear to linear-oblanceolate and covered with observed to spread through underground roots
resin or gray tomentum. The uppermost leaves (Paulsen and Miller 1968, McArthur and others
rarely extend beyond the inflorescence. Flower 1979). We are unaware of seed germination data.
heads contain four to six yellow disk flowers and
are subtended by ranked involucral bracts with Growth and Management.—Seed germinates in
slender recurved tips. Subspecies nevadensis the early spring and seedlings become established
occurs on dry mountainsides from eastern during the following spring and summer. Plants
California to eastern Nevada, southwestern Utah, generally mature within 5 years and typically live
and Northern Arizona, being most common on the 15 to 20 years. Paulsen and Miller (1968) reported
eastern slopes of the Sierra Nevada (Anderson that, when necessary, Parry rabbitbrush can be
1986, McArthur and others 1979). controlled by the herbicide Tordon®.1
Subspecies C. parryi ssp. parryi consists
of low shrubs with erect branches and can be as Benefits.—Although Parry rabbitbrush is a
tall as 76 cm. The uppermost leaves usually extend component of western uplands, it is spotty in
beyond the inflorescence. Flower heads include 10 distribution and usually sparse in population
to 20 disc flowers and are subtended by obscurely density. It has some importance as a browse source
ranked bracts with attenuate tips. This subspecies in those areas where it is locally abundant. Some
grows on dry hillsides and plains in central subspecies of Parry rabbibrush, including C. parryi
Nevada, southern Utah, south-central Wyoming, ssp. asper and C. parryi ssp. monocephalus, are
western Colorado, and northern New Mexico particularly attractive plants and have horticultural
(Anderson 1986, McArthur and others 1979). potential as ornamentals.
Anderson (1970) made a case for this taxon being
the most primitive of all Chrysothamnus. References
Parry rabbitbrush in the form of C. parryi
ssp. attenuatus has been implicated in the Anderson, L.C. 1966. Cytotaxonomic studies in
parentage of the putative stabilized hybrid species Chrysothamnus (Asterae, Compositae). American
C. nauseosus ssp. uintahensis Anderson 1984). Journal of Botany 53: 204-212.
Range.—The range of Parry rabbitbrush extends Anderson, L.C. 1970. The karyotype of
from California in the west, to western Nebraska Chrysothamnus parryi ssp. parryi and its
in the east, and from Wyoming in the north, to implication. Transactions of the Kansas
New Mexico and Arizona in the south. Subspecies, Academy of Science 72: 399-401.
however, may be more geographically restricted.
Within its range, Parry rabbitbrush may be found Anderson, L.C. 1984. Chrysothamnus nauseosus
at elevations between 790 and 3,400 m. ssp. uintahensis: a stabilized hybrid. Great Basin
Naturalist 44:416-420.
Ecology.—Parry rabbitbrush grows on dry, open
foothills and mountains. Precipitation throughout Anderson, L.C. 1986. An overview of the genus
its range generally exceeds 380 mm. Parry Chrysothamnus. In: E.D. McArthur and B.L
rabbitbrush tends to increase on overgrazed or Welch, comps. Proceedings—symposium on the
otherwise disturbed areas. Populations are usually
smaller and more scattered than those of more 1
Use of trade names does not imply endorsement
of the U. S. Department of Agriculture
207
biology of Artemisia and Chrysothamnus; 1984 U.S. Department of Agriculture, Forest Service,
July 9-13, Provo, UT. General Technical Report Intermountain Forest and Range Experiment
INT-200. U.S. Department of Agriculture, Forest Station, Ogden, UT. 82 p.
Service, Intermountain Research Station, Ogden,
UT: 29-45. McArthur, E.D. and S.E. Meyer. 1987. A review
of the taxonomy and distribution of
Anderson, L.C. 1995. The Chrysothamnus- Chrysothamnus. In: K.L. Johnson, ed.
Ericameria connection (Asteraceae). Great Basin Proceedings of the Fourth Utah Ecology
Naturalist 55: 84-88. Workshop; 1986 September 17-18, Cedar City,
UT. College of Natural Resources, Utah State
Deitschman, G.H., K.R. Jorgensen, and A.P. University, Logan, UT. p. 9-18.
Plummer. 1974. Chrysothamnus. In: C.S.
Schopmeyer, tech. coord., Seeds of Woody Nesom, G.L. and G.I. Baird. 1993. Completion of
Plants of the United States. Agriculture Ericameria (Asteraceae: Asterae), diminution of
Handbook 450. Washington, DC, U.S. Chrysothamnus. Phytologia 75: 74-93.
Department of Agriculture, Forest Service.
p. 326-328. Paulsen, H.A., Jr. and J.C. Miller. 1968. Control of
Parry rabbitbrush on mountain grasslands of
McArthur, E.D., A.C. Blauer, A.P. Plummer, and Western Colorado. Journal of Range
R. Stevens. 1979. Characteristics and Management 21: 165-177
hybridization of important intermountain shrubs. .
III. Sunflower family. Research Paper INT-220.
208
Chrysothamnus viscidiflorus Nutt. low rabbitbrush
ASTERACEAE
209
viscidiflorus with C. nauseosus and C. parryi viscidiflorus were 37 and 58 percent, respectively.
based on molecular genetic techniques (Nesom Seeds from plants adapted to higher elevations
and Baird 1993; Anderson 1995) (where seeds ripen earlier in the year) are more
dormant than those collected from lower
Range.—Low rabbitbrush is one of the most elevations and may be more chill responsive
widely distributed shrubs in Western North (Meyer 1996). There are about 1,720 cleaned seeds
America (McArthur and others 1979). One reason per gram (Deitschman and others 1974).
for its broad distribution is the species’ great
ecological amplitude. It is found growing on dry, Growth and Management.—Seedlings emerge in
open areas from British Columbia and Montana, spring and grow quickly. After a period of growth
south to New Mexico, Arizona, and eastern and establishment, shrubs often produce seed after
California. their second growing season (Meyer 1996). Low
rabbitbrush can dominate rangelands that are
Ecology.—Due to its adaptability, low rabbitbrush grazed heavily or cleared. This species has a strong
grows in a variety of habitats. Plants generally, tendency to sprout from the base (Wasser 1982),
however, inhabit drier sites, such as foothills, which may make it more difficult to remove and
mountains, dry plains, and valleys. Annual difficult to control with herbicides (Cook and
precipitation within its range varies between 250 others 1965). Fall is the preferred time to seed;
and 535 mm, with notable exceptions at higher seed can be drilled or broadcast and should remain
elevations. Anderson (1986) has shown that within the top 3 mm of the soil surface. Seeds can
polyploid races, in subspecies where polypoidy be harvested by hand and by vacuum harvesting
occurs (ssp. lanceolatus, puberulus, and techniques.
viscidiflorus), are adapted to lower and drier sites
than their diploid counterparts. Interestingly, Benefits.—Low rabbitbrush is an important native
Anderson has also observed a correlation within component of Western North American
certain subspecies between plant stature and leaf shrublands. It can provide important browse to
size with altitude and precipitation (personal both game and livestock, especially during late fall
communication). In addition to its ability to and winter months after more desirable forage has
tolerate xeric conditions, some variants of low been consumed. As a forage shrub, the palatability
rabbitbrush can grow well in areas with poor or of low rabbitbrush varies significantly among
disturbed soils. Low rabbitbrush can be found at subspecies, varieties, locality, and season. Some
elevations ranging from 790 to 3,400 m. It is subspecies, e.g., ssp. lanceolatus, may be used
commonly associated with big sagebrush heavily by domestic livestock and wildlife,
(Artemisia tridentata), broom snakeweed whereas others are consumed little, if at all (USDA
(Gutierrezia sarothrae), and other rabbitbrushes 1937, Goodrich and others 2001). In addition to its
Chrysothamnus sp.), although it can be associated value as a browse species, low rabbitbrush has
with shadscale (Atriplex confertifolia), winterfat successfully been used to revegetate depleted
(Ceratoides lanata), fourwing saltbush (Atriplex rangelands and other disturbed sites (Plummer
canescens), greasewood (Sarcobatus 1977).
vermiculatus), and occasionally pinyon (Pinus sp.)
and juniper (Juniperus sp.). Goodrich and others References
(2001) note that C. viscidiflorus does not
commonly overtop associated herbaceous Abrams, R. and R.S. Ferris. 1960. Illustrated flora
vegetation. Elevation tends to partition the of the Pacific States, Vol. 4, Bignoniaceae to
subspecies; mountain low rabbitbrush (ssp. Compositae. Stanford University Press,
lanceolatus) especially and stickyleaf low Stanford, CA. 732 p.
rabbitbrush (ssp. viscidiflorus) grow well at higher
elevations, while hairy low rabbitbrush (ssp. Anderson, L.C. 1966. Cytotaxonomic studies in
puberlus) does best on lower desert foothills. Chrysothamnus (Asterae, Compositae).
American Journal of Botany 53: 204-212.
Reproduction.—Shrubs flower from August
through October; seed matures from October until Anderson, L.C. 1971. Additional chromosome
December. Like other members of the genus, low numbers in Chrysothamnus (Asteraceae).
rabbitbrush seeds (or achenes) are wind Bulletin of the Torrey Botanical Club 98: 222-
disseminated. Meyer (1996) found that 225.
germination rates at 15 oC for ssp. lanceolatus and
210
Anderson, L.C. 1980. Identity of narrow-leaved U. S. Department of Agriculture, Forest Service,
Chrysothamnus viscidiflorus (Asteraceae). Intermountain Forest and Range Experiment
Great Basin Naturalist 40: 117-120. Station, Ogden, UT. 82 p.
Anderson, L.C. 1986. Cytogeography of McArthur, E.D., D.L. Hanks, A.P. Plummer, and
Chrysothamnus viscidiflorus. In: E.D. McArthur A.C. Blauer. 1978. Contributions to the
and B.L. Welch, comps. Proceedings— taxonomy of Chrysothamnus visicidiflorus
symposium on the biology of Artemisia and (Astereae, Compositae) and other
Chrysothamnus; 1984 July 9-13, Provo, UT. Chrysothamnus species using paper
General Technical Report INT-200. Ogden, UT: chromatography. Journal of Range Management
U.S. Department of Agriculture, Forest Service, 31: 216-223.
Intermountain Research Station. p. 93-97.
McArthur, E.D. and S.E. Meyer. 1987. A review
Anderson, L.C. 1995. The Chrysothamnus- of the taxonomy and distribution of
Ericameria connection (Asteraceae). Great Chrysothamnus. In: K.L. Johnson, ed.
Basin Naturalist 55: 84-88. Proceedings of the Fourth Utah Ecology
Workshop; 1986 September 17-18, Cedar City,
Cook, C.W., P.D. Leonard, and C.D. Bonham. UT. College of Natural Resources, Utah State
1965. Rabbitbrush competition and control on University, Logan, UT. p. 9-18.
Utah rangelands. Bulletin 454. Utah
Agricultural Experiment Station., Logan, UT. Meyer, S.E. 1996. Chrysothamnus Nutt. In:
28 p. Woody Plant Seed Manual,
http://www.wpsm.net/Chrysothamnus.pdf. 11 p.
Deitschman, G.H., K.R. Jorgensen, and A.P.
Plummer. 1974. Chrysothamnus. In: C.S. Nesom, G.L., and G.I. Baird. 1993. Completion of
Schopmeyer, tech. coord., Seeds of Woody Ericameria (Asteraceae: Asterae), diminution of
Plants of the United States. Agriculture Chrysothamnus. Phytologia 75: 74-93.
Handbook 450. U.S. Department of Agriculture,
Forest Service. p. 326-328. Plummer, A.P. 1977. Revegetation of disturbed
intermountain area sites. In: J. L. Thames,
Goodrich, S., R.M. Thompson, and A. Huber. editor, Reclamation and use of disturbed land in
2001. A yellowbrush/grass community type the Southwest. University of Arizona Press,
from the Uinta Mountains and Utah plateaus. Tucson, AZ: 302-339.
In: E.D. McArthur and D.J. Fairbanks, comps.
Shrubland ecosystem genetics and biodiversity: U.S. Department of Agriculture, Forest Service.
proceedings; 2000 June 13-15, Provo, UT. 1937. Range plant handbook. United States
Proceedings RMRS-P-21. U. S. Department of Government Printing Office, Washington, DC.
Agriculture, Forest Service, Rocky Mountain 816 p.
Research Station, Ogden, UT. p. 344-346.
Wasser, C.H. 1982. Ecology and culture of
McArthur, E.D., A.C. Blauer, A. P. Plummer, and selected species useful in revegetating disturbed
R. Stevens. 1979. Characteristics and lands in the West. U.S. Department of the
hybridization of important intermountain shrubs. Interior, Fish and Wildlife Service, Washington,
III. Sunflower family. Research Paper INT-220. DC. 347 p.
211
Cissampelos pareira L. velvet leaf
MENISPERMACEAE
John K. Francis
212
available on the growth rate of seedlings. Growth International BioPark Foundation. 2002. Abuta,
of sprouts is rapid (at least 3 m in the first year). family Menispermaceae. http://www.biopark.
Individual stems are not long-lived, but by org/peru/abuta.html. 3 p.
sprouting and layering, plants or clones may last
many years. The creation of small forest openings
Liogier H.A. 1985. Descriptive flora of Puerto
and protection against heavy grazing are probably
Rico and adjacent islands, Spermatophyta. Vol.
good strategies to promote natural establishment of
1. Editorial de la Universidad de Puerto Rico,
new plants.
Río Piedras, PR. 352 p.
Benefits.—Velvet leaf adds to biodiversity and
Long, R.W. and O. Lakela. 1976. A flora of
biomass, helps stabilize the soil, and furnishes food
tropical Florida. Banyan Books. Miami, FL.
and cover for wildlife. A major interest in the
962 p.
species arises from the natural medicinal benefits
of the plant’s chemical contents. Known as the
“midwife’s herb,” it has been used for centuries by Morita, H., K. Matsumoto, K. Takeya, and H.
native peoples of South America to treat menstrual Itokawa. 1993a. Azafluoranthene alkaloids from
cramps, prevent threatened miscarriage, control Cissampelos pareira. Chemical and
uterine hemorrhages, and ease childbirth and Pharmaceutical Bulletin 41(7): 1,307-1,308.
postpartum pain (Rain-tree 2002). The list of other
natural medicinal applications to which the herb is
applied is large: urinary infections, kidney stones, Morita, H., K. Matsumoto, K. Takeya, H. Itokawa,
and Y. Iitaka. 1993b. Structures and solid state
arthritis, snakebite, cough, dysentery, piles, ulcers,
pain, indigestion, colic, skin irritations, stings, tautomeric forms of two novel antileukemic
intestinal worms, and wounds (Parrotta 2001, Rain- tropoloisoquinoline alkaloids, pareirubrines A
and B, from Cissampelos pareira. Chemical and
tree 2002). Ground tissues and preparations of
velvet leaf are sold throughout the world in Pharmaceutical Bulletin 41(8): 1,418-1,422.
markets, shops, and mail-order companies. The
physiological effects are apparently derived from a Nelson, G. 1996. The shrubs and woody vines of
number of alkaloids found in the tissues of the Florida. Pineapple Press, Inc., Sarasota, FL.
plant (International BioPark Foundation 2002, 391 p.
Morita and others 1993a, Morita and others
1993b). Parrotta, J.A. 2001. Healing plants of Peninsular
India. CABI Publishing, Wallingford, UK and
References New York. 917 p.
213
Citrus aurantiifolia (Christm.) Swingle Key lime
RUTACEAE
John K. Francis
214
Reproduction.—Key lime flowers in late spring citrus fruits cause phytophotodermatosis in
and fruits in fall to spring in Florida (Nelson 1996). sensitive individuals (Bruneton 1999). The wood is
In Puerto Rico, it flowers in spring to early summer hard and heavy, but is used for little other than
and fruits in summer and fall (Little and fuel. It is a good honey plant. With pruning, the
Wadsworth 1964). The fruits ripen and fall from plant can be used for a living fence post (Little and
the trees 5 to 6 months after flowering (Morton Wadsworth 1964) and can be formed into a hedge
1987). Fruits in one collection in Puerto Rico (Burkill 1997).
ranged from 3.0 to 4.5 cm in diameter and weighed
from 19 to 51 g. Air-dried seeds separated from References
them averaged 0.0702 + 0.0037 g/seed or 14,000
seeds/kg. Placed on moist blotter paper, 84 percent Bruneton, J. 1999. Toxic plants dangerous to
germinated between 15 and 49 day after sowing. humans and animals. Lavoisier Publishing,
Germination is epygeal. Many of the seeds are Paris. 545 p.
polyembryonic. Seedlings begin to fruit in 3 to 6
years (Morton 1987). In the West Indies, the seeds Burkill, H.M. 1997. The useful plants of West
are mainly dispersed by gravity and humans. There Tropical Africa. Vol. 4. Royal Botanic Gardens,
are undoubtedly animal vectors in the original Kew, UK. 969 p.
native habitat. Key lime can be air-layered using
indol buteric acid (IBA) with a high degree of Ehler, J.T. 2002. Key lime (Citrus aurantifolia).
success (Morton 1987). Food Reference, Key West, FL.
http://foodreference.com/html/artkeylimes.html.
Growth and Management.—Growth of Key lime 2 p.
is slow, less than 0.5 m/year. Plants may live for 25
years or more. Although the species is not managed Katzer, G. 2002. Lime [Citrus aurantifolia
in the wild, existing trees are usually protected by (Christm. & Panz.) Swengle]. Gernet Katzer’s
local inhabitants, whenever possible. Plantations spice pages. http://www-ang.kfunigraz.
are established using potted material that is set at ac.at/~katzer/engl/Citr_aur.html. 4 p.
7.5- by 7.5-m spacing. Pruning is usually not
necessary (Morton 1987). Liogier H.A. 1988. Descriptive flora of Puerto
Rico and adjacent islands, Spermatophyta. Vol.
Benefits.—Key lime has been under cultivation or 2. Editorial de la Universidad de Puerto Rico,
semicultivation for thousands of years. Improved Río Piedras, PR. 481 p.
horticultural varieties, which have bigger, generally
seedless fruits and thornless plants, now dominate Liogier, H.A. 1990. Plantas medicinales de Puerto
the local and international fruit markets. The Rico y del Caribe. Iberoamericana de Ediciones,
unimproved type is still managed commercially Inc., San Juan, PR 566 p.
and is the basis for a juice and lime oil industry
(Morton 1987). It is also planted for a dooryard Little, E.L., Jr. and F.L. Wadsworth. 1964.
fruit tree, and fruits are harvested from the wild. Common trees of Puerto Rico and the Virgin
The principal use is still for food, refreshing drinks, Islands. Agriculture Handbook 249. U.S.
tasty disserts, and for seasoning meats, vegetables, Department of Agriculture, Washington, DC.
salads, sauces, and casseroles (Ehler 2002, Katzer 548 p.
2002). The wild type is superior in flavor to the
improved varieties because it has a stronger flavor Morton, J.F. 1987. Citrus aurantifolia Swingle.
and a higher acid content (Ehler 2002). The from: Fruits of warm climates.
pericarp (rind) contains 7 percent essential oil with http://www.hort.perdue.edu/newcrop/morton/me
principal constituants, citral, limonene, and xican_lime.html. 8 p.
fenchon as well as terpineol, bisabolene, and other
terpenoids (Katzer 2002). Key lime is used to treat Nelson, Gil. 1996. The shrubs and woody vines of
a huge number of ailments (Burkill 1997, Liogier Florida. Pineapple Press, Inc. Sarasota, FL.
1990). The author recommends a tea prepared from 391 p.
juice, fruit rind, or leaves as an expectorant and to
relieve catarrh brought on by colds and flu. The
fresh fruits and bottled juice are an excellent source
of vitamin C and were once relied upon to prevent
scurvy. Essential oils of Key lime and some other
215
Secritaría del Medio Ambiente y Recursos
Naturales. 2002. Especies con usos no
maderables en bosques tropicales y
subtropicales: Lantana involucrata L.
http://www.semarnat.gob.mx.pfnm2.fichas/
citrus_aurantifolia.htm. 2 p.
216
Clematis orientalis L. oriental virginsbower
RANUNCULACEAE
John K. Francis
217
environmental damage is not severe, because it is References
an alien, the species should not be used in
environmental restoration plantings. No specific Griffiths, M. 1994. Index of garden plants. Timber
recommendations for control are available, but the Press, Portland, OR. 1,234 p.
general procedure of cutting plants and spot
spraying the sprouts with broadleaf herbicide Natural Resources Conservation Service. 2003.
would probably be effective. The use of potted Plants profiles: Clematis orientalis L., oriental
seedlings or rooted cuttings for ornamentals is virginsbower. http://plants.gov/cgi_bin/
recommended. They should be outplanted in the plant_profile.cgi?symbol=CLOR. 5 p
spring after the frost danger has passed. Pruning
should take place in the spring (Plants for a Future Plants For a Future. 2003. Database search results:
2003). Clematis orientalis. http://www.scs.leeds.ac.uk/
cgi-bin/pfaf/arr_html?Clematis+orientalis. 7 p.
Benefits.—Oriental virginsbower is planted as an
ornamental and ground cover. Plants escaped into Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
the wild help protect the soil, furnish cover for Hinnins, eds. 1987. A Utah flora. Great Basin
wildlife, and add beauty to the landscape, Naturalist Memoirs 9. Brigham Young
especially during the fall when it produces flower University, Provo, UT. 894 p.
and seed heads. Oriental virginsbower is used in
landscaping as a flowering ground cover. Infusions Wu Z., P.H. Raven, and Hong D. (eds.). 2001.
are used as a gargle for sore and ulcerated throats Flora of China. Vol. 6. Science Press, Beijing
and to treat dog bites (Plants for a future 2003). and Missouri Botanical Garden Press, St. Louis,
Mo. 512 p.
218
Clibadium erosum (Sw.) DC. carruzo
ASTERACEAE
John K. Francis
219
Stands along roads are routinely controlled by http://www.inbio.ac.cr/bins/k03/p13/c045/o0144
mowing. /f01362/g007091/s020661.htm. 1 p.
Benefits.—Carruzo helps protect the soil in Liogier, H.A. 1997. Descriptive flora of Puerto
disturbed areas, serves as a nurse species for later Rico and adjacent islands. Vol. 5. Editorial de la
successional species, and provides cover and Universidad de Puerto Rico, Río Piedras, PR.
possibly food for wildlife. 436 p.
220
Clidemia cymosa (Wendl. ex Spreng.) Alain camasey peludo
MELASTOMATACEAE
John K. Francis
to hundreds of tiny, light brown seeds (Liogier
1995, Little and Wadsworth 1964).
221
Growth and Management.—Camasey peludo used because of the poor form and small size of the
plants are relatively short-lived, lasting about 3 to 6 plants (Little and Wadsworth 1964).
years. Resprouting and layering may enable them
to persist somewhat longer. There is no reported References
planting or management experience for this
species. Plants cloned by layering should be used Liogier, H.A. 1995. Descriptive flora of Puerto
for planting until methods of growing the plants Rico and adjacent islands. Vol. 4. Editorial de la
from seed are developed. Natural reproduction in Universidad de Puerto Rico, San Juan, PR.
areas of abundant rainfall may be encouraged by 617 p.
cultivation or scalping to expose a bare seedbed
during wet portions of the year. Little, E.L., Jr. and F.L. Wadsworth. 1964.
Common trees of Puerto Rico and the Virgin
Benefits.—Camasey peludo is an attractive Islands. Agriculture Handbook 249. U.S.
roadside plant and should be encouraged whenever Department of Agriculture, Washington, DC.
possible. Its beauty also suggests real potential as 548 p.
an ornamental, although its life is short and it has a
tendency to unsightly decline in its last year. The Martorell, L.F. 1975. Annotated food plant catalog
fruit, currently almost unknown to the public, is of the insects of Puerto Rico. Department of
very tasty and deserves attention. The fruits are a Entomology, Agricultural Experiment Station,
food source for birds. The light-brown, moderately University of Puerto Rico, Río Piedras, PR.
hard wood has a specific gravity of 0.6 but is not 303 p.
222
Clidemia hirta (L.) D. Don Koster’s curse
MELASTOMATACEAE
John K. Francis
223
birds, feral pigs, other animals, and humans. The Franca, F., E.L. Lago, and P.D. Marsden. 1996.
seeds can remain viable in the soil for up to 4 Plants used in the treatment of leishmanial
years. The stems root on contact with moist soil; ulcers due to Leishmania (Vannia) braziliensis
even detached leaves have been observed to root in an endemic area of Bahia, Brazil. Revista da
(Smith 1992). Sociedade Brasileira de Medicina Tropical
29(3): 229-232.
Growth and Management.—Koster’s curse
grows slowly in the first months and faster Howard, R.A. 1989. Flora of the Lesser Antilles,
thereafter. It often dominates at the end of the first Leeward and Windward Islands. Vol. 5. Arnold
year (Binggeli 1997). Individual stems live 2 or 3 Arboretum, Harvard University, Jamaica Plain,
years in Puerto Rico, although plants may live MA. 604 p.
longer by sprouting. There is little motivation to
encourage Koster’s curse and often a need to Liogier, H.A. 1995. Descriptive flora of Puerto
control it. Single-pass mowing is usually Rico and adjacent islands. Vol. 4. Editorial de la
ineffective. Spraying with broadleaf herbicides is Universidad de Puerto Rico, San Juan, PR.
the most widely used control method, 617 p.
supplemented by hand pulling (Rubber Research
Institute of Malaya 1973). At best, such measures Murdiati, T.B., C.S. McSweeney, R.S.F.
are temporary. Some success has been achieved Campbell, and D.S. Stoltz. 1990. Prevention of
with biocontrol by the introduction of Liothrips hydrolysable tannin toxicity in goats fed
urichi, a thrips (Binggeli 1997). A number of other Clidemia hirta by calcium hydroxide
natural controls are being studied (State of Hawaii supplementation. Journal of Applied Toxicology
Department of Agriculture 2002). 10(5): 325-331.
Benefits and Detriments.—Desired or not, Pacific Island Ecosystems at Risk. 2002. Clidemia
Koster’s curse helps revegetate disturbed areas and hirta (L.) D. Don, Melosomataceae. http://www.
provides a reliable food source for wildlife. The hear.org/pier3clhir.htm. 5 p.
species is used in Brazil to treat Leishmania
braziliensis skin infections (Franca and others Rubber Research Institute of Malaya. 1973.
1996). On the other hand, Koster’s curse is a Clidemia hirta in South Johore. Planter’s
serious threat to understory plant species in fragile Bulletin 128: 140-144.
tropical island ecosystems (Pacific Island
Ecosystems at Risk 2002). It is a serious weed in Smith, C.W. 1992. Distribution, status, phenology,
tropical plantations such as rubber (Hevea rate of spread, and management of clidemia in
brasiliensis Muell. Arg.). Sheep have been shown Hawai’i. In: C.P. Stone, C.W. Smith, and J.T.
to control most weeds in plantations but will not Tunison, eds. Alien plant invasions in native
eat Koster’s curse (Chee and Faiz 2002). Goats ecosystems of Hawaii, management and
suffer toxicity from hydrolysable tannin when fed research. University of Hawaii Cooperative
the plant (Murdiati and others 1990). National Park Resources Studies Unit,
Honolulu, HI. p. 241-253.
References
State of Hawaii Department of Agriculture. 2002.
Binggeli, P. 1997. Clidemia hirta (L.) D. Don Plant pest control, persisting biological control
(Melastomataceae). http://members.lycos.co.uk/ problems. http://www.hawaiiag.org/hdoa/
WoodyPlantEcology/docs/web-sp5.htm. 4 p. pi_ppc_bioprob.htm. 6 p.
Chee, Y.K. and A. Faiz. 2002. Sheep grazing Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
reduces chemical weed control in rubber. Montiel, eds. 2001. A Flora de Nicaragua.
Australian Centre for International Agriculture Monographs in Systematic Botany Vol. 85, No.
Research. http://www.aciar.gov.au/publications/ 2. Missouri Botanical Garden Press, St. Louis,
proceedings/32/paper26.pdf. 4 p. MO. p. 945-1,910.
224
Coccoloba krugii Lindau wild-grape
POLYGONACEAE
Synonyms: none
John K. Francis
Range.—Wild-grape is native to the Bahamas, Benefits.—Wild-grape wood is heavy and hard and
225
useful for at least fuel. The fruits are edible and International Reptile Conservation Foundation.
tasty, though small. The fruits are counted among 2002. Effects of introduced ungulates on the
the food sources of the endangered Anegada iguana iguana on Anegada. http://cyclura.com/
(Cyclura pinguis) (International Reptile article/articleview/248/1/6/. 6 p.
Conservation Foundation 2002).
Liogier H.A. 1985. Descriptive flora of Puerto
References Rico and adjacent islands, Spermatophyta. Vol.
1. Editorial de la Universidad de Puerto Rico,
Howard R.A. 1988. Flora of the Lesser Antilles, Río Piedras, PR. 352 p.
Leeward and Windward Islands.
Dicotyledoneae, Part 1. Vol. 4. Arnold Little, E.L., Jr. R.O. Woodbury, and F.H.
Arboretum, Harvard University, Jamaica Plain, Wadsworth. 1974. Trees of Puerto Rico and the
MA. 673 p. Virgin Islands. Vol. 2. Agric. Handb. 449. U.S.
Department of Agriculture, Washington, DC.
1,024 p.
226
Coccoloba microstachya Willd. pockhout
POLYGONACEAE
John K. Francis
227
uncommon.
Liogier H.A. 1985. Descriptive flora of Puerto
Growth and Management.—Pockhout is slow Rico and adjacent islands, Spermatophyta. Vol.
growing and long lived. No published planting or 1. Editorial de la Universidad de Puerto Rico,
management experience is known to the author. Río Piedras, PR. 352 p.
Benefits.—Pockhout contributes to the Little, E.L., Jr., R.O. Woodbury, and F.H.
biodiversity of secondary and remnant forests, Wadsworth. 1974. Trees of Puerto Rico and the
helps protect the soil, and furnishes food and cover Virgin Islands. Vol. 2. Agriculture Handbook
for wildlife. The wood is good for fuel. 449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
References
Weaver, P.L., J.L. Ramirez, and J.L. Coll-Rivera.
Francis, J.K. and A. Rodríguez. 1993. Seeds of 1999. Las Cabezas de San Juan Nature Reserve
Puerto Rican trees and shrubs: second (El Faro). General Technical Report IITF-5.
installment. Res. Note SO-374. Southern Forest International Institute of Tropical Forestry, U.S.
Experiment Station, U.S. Department of Department of Agriculture, Forest Service, Río
Agriculture, Forest Service, New Orleans, LA. Piedras, PR. 58 p.
5 p.
228
Coccoloba uvifera (L.) L. seagrape
POLYGONACEAE
John A. Parrotta
229
after flowering. The ripe, purple fruits contain a and dyeing (Little and Wadsworth 1964, Uphof
single, hard, elliptic seed about 1 cm long 1968). The astingent roots and bark have been
surrounded by an edible, tartly sweet pulp and a used in traditional medicine in Puerto Rico and
thin, fleshy covering. Fresh seed weights range elsewhere in the Caribbean (Liogier 1990).
from 1,000 to 1,400 seeds/kg, based on collections Seagrape’s tolerance of saline soils and sea spray
in Puerto Rico (Parrotta 1994). The fruits are has made it an excellent and popular choice for
consumed by a variety of birds that help to ornamental plantings and coastal windbreaks. It
disperse the seeds. Natural reproduction is prunes well and makes an attractive hedge.
generally good on open coastal sites, although
seedlings do not tolerate shading or competition References
from grasses and herbs. In the nursery, seagrape
can be propagated from seed in light-textured, Adams, C.D. 1972. Flowering plants of Jamaica.
well-drained soils; no pretreatment is required to University of the West Indies, Mona, Jamaica.
break seed dormancy. Seeds should be sown on or 848 p.
near the soil surface. Germination takes place
between 18 and 50 days after sowing, and Chellman, C.W. 1978. Pests and problems of
seedlings reach plantable size (15 cm tall) 4 to 5 South Florida trees and palms. Florida
months after sowing. It can also be propagated Department of Agriculture and Consumer
vegetatively by air-layering, ground-layering, and Services, Division of Forestry, Talahassee, FL.
grafting, though mature wood cuttings are most 103 p.
commonly used to produce rooted cuttings for
mass propagation of selected cultivars (FAO 1982, Craig, R.M. 1984. Plants for coastal dunes of the
Parrotta 1994). Gulf and South Atlantic coasts and Puerto Rico.
Information Bulletin 460. U.S. Department of
Growth and Management.—Growth rates in Agriculture, Soil Conservation Service,
seagrape are strongly influenced by wind exposure; Washington, DC. 41 p.
on exposed sites, heights typically increase from
the windward to leeward edges of stands. On a less FAO. 1982. Fruit-bearing forest trees: technical
exposed coastal site with sandy soils receiving an notes. FAO For. Pap. 34. Food and Agriculture
annual rainfall of approximately 1500 mm in Organization of the United Nations, Rome.
northeastern Puerto Rico, average heights of 0.9 177 p.
and 2.9 m were recorded for 1- and 3-year-old
plantings (Parrotta 1994). No information is Gooding, E.G.B. 1974. The plant communities of
available on growth rates of older trees. Under Barbados. Government Printing Offic,e
optimal conditions seagrape can attain a height of 8 Bridgetown, Barbados. 177 p.
to 18 m and a stem diameter of 45 to 60 cm
(Record and Hess 1943). Being shade-intolerant
Liogier, H.A. 1990. Plantas medicinales de Puerto
and susceptible to competition from grasses, herbs
Rico y del Caribe. Iberoamericana de Ediciones,
and other woody species, planted stands should be
Inc., San Juan, PR. 566 p.
regularly weeded until they are taller than
competing vegetation.
Little, E.L., Jr. and F.H. Wadsworth. 1964.
Benefits.—Throughout its range, the wood is used Common trees of Puerto Rico and the Virgin
as firewood and for making charcoal. The reddish- Islands. Agriculture Handbook 249. U.S.
to dark-brown heartwood, distinguished from the Department of Agriculture, Forest Service,
light-brown sapwood in larger trees, is hard, close- Washington, DC. 548 p.
grained, and used in turning and occasionally for
furniture, inlay work and cabinetry (Gooding Martorell, L.F. 1975. Annotated food plant catalog
1974). The edible fruits can be eaten raw, used in of the insects of Puerto Rico. Agricultural
preserves, or fermented like grapes to make wine. Experiment Station, University of Puerto Rico,
The flowers yield abundant nectar, and the Río Piedras, PR. 303 p.
resulting honey is of good quality, light amber in
color, with a spicy flavor (National Research
Council 1983). The astringent red sap exuded from National Research Council. 1983. Firewood crops:
the cut bark, known commercially as West Indian shrub and tree species for energy production,
or Jamaican kino, was formerly used for tanning
230
Vol. 2. BOSTID Report 40. National Academy Stoffers, A.L. 1956. The vegetation of the
of Sciences, Washington, DC. 92 p. Netherlands Antilles. 15: Studies on the flora of
Curaçao and other Caribbean islands. Vol.
1.Nartinus Nijhoff, The Hague. 142 p.
Parrotta, J.A. 1994. Coccoloba uvifera (L.) L. –
Sea grape, uva de playa. Research Note SO-ITF-
Uphof, J.T.C. 1968. Dictionary of economic
SM-74. U.S. Department of Agriculture, Forest
plants. Verlag von J. Cramer, New York.
Service, Southern Forest Experiment Station,
591 p.
New Orleans, LA. 5 p.
USDA. 1960. Index of plant diseases in the United
Record, S.J. and R.W. Hess. 1943. Timbers of the
States. Agriculture Handbook 165. U.S.
New World. Yale University Press, New Haven,
Department of Agriculture, Washington, DC.
CT. 640 p.
531 p.
231
Coffea arabica L. coffee
RUBIACEAE
John K. Francis
232
others 2001). The flowers are insect pollinated 0.04 percent, stem 0.01 percent, and central root,
(Wrigley 1988). The fruits take from 6½ to 7 0.01 percent. Other alkaloids, xanthine, guanine,
months to mature (Bailey 1941). Seeds of the and trigonelline, which have stimulant and diuretic
varieties naturalized in Puerto Rico vary from properties, are also present (Burkill 1997). Caffeine
0.139 to 0.147 g/seed (Barrett 1925). Seeds 8 protects vegetative plant parts from insect and
weeks old germinate (95 percent in 32 days) better fungal attack and inhibits the growth of plants and
than fresh seed (Wrigley 1988). However, coffee bacteria near germinating seeds (Steiman 1997).
seeds stored more than 21 weeks at ambient Purified caffeine is widely sold as a medicinal
temperature begin to rapidly lose their viability stimulant, dietary aid, and headache remedy. Leaf
(Harrer 1963). The seeds are dispersed by birds, poultices are used to treat sores in Trinidad, and
and seedlings can be abundant. Artificial root sap or root infusions are drunk to relieve
propagation is usually by seed, but the species can scorpion stings (Burkill 1997). Coffee is also
be reproduced by budding and cuttings (Center for employed in folk medicine to treat asthma, flu,
New Crops and Plants Products 1996). headache, jaundice, nephrosis, malaria, sores, and
vertigo (Center for New Crops and Plants Products
Growth and Management.—Coffee has a 1996).
moderate growth rate. Plants begin bearing in 3 to
4 years and are in full fruit production in 6 to 8 References
years. Coffee plants may live over 100 years
(Center for New Crops and Plants Products 1996). Bailey, L.H. 1941. The standard cyclopedia of
Nursery plants are maintained in containers under horticulture. MacMillan, New York. 3,639 p.
shade for 6 to 12 months before outplanting
(Wrigley 1988). Also, plantations are established Barrett, O.W. 1925. The food plants of Puerto
by direct seeding into prepared seed spots (Center Rico. Journal of the Department of Agriculture
for New Crops and Plants Products 1996). Weed of Puerto Rico 9(2): 61-208.
control is necessary in the first years after
plantation establishment. Naturalized and escaped Burkill, H.M. 1997. The useful plants of West
coffee is usually not common enough or does not Tropical Africa. Vol. 4. Royal Botanic Gardens,
grow fast enough to warrant control. Kew, UK. 969 p.
Benefits.—As an exotic species in New World Center for New Crops and Plants Products. 1996.
forests, coffee has had a gentle impact on Coffea arabica L. Perdue University. http://hort.
biodiversity and contributes to wildlife food and perdue.edu/newcrop/duke_energy/Coffea_
cover, and soil stability. The wood is used mainly arabica.html. 7 p.
for fuel in the New World but is turned into chairs
and other types of furniture in Africa (Cheney Charrier, A. and J. Berthaud. 1985. Botanical
1925). Coffee is a good honey plant and yields a classification of coffee. In: M.N. Clifford and
light-colored honey (Little and Wadsworth 1964). K.C. Willson, eds. Coffee: botany, biochemistry,
Coffee berries, edible and slightly sweet, are eaten and production of beans and beverage. The AVI
occasionally by children and field workers. The Publishing Company, Inc., Westport, CN. p. 13-
fruit pulp, which is removed during processing, is 47.
sometimes fed to livestock but more often is
composted for fertilizer and mulch (Center for New Cheney, R.H. 1925. Coffee. The New York
Crops and Plants Products 1996). Coffee seeds University Press, New York. 244 p.
have been chewed as a stimulant in East Africa
from ancient times (Center for New Crops and Harrer, A.E. 1963. Coffee growing. Oxford
Plants Products 1996). The hot drink “coffee” is University Press, London. 127 p.
brewed from the roasted and ground seeds (or
“beans”) and is one of the world’s most popular Howard, R.A. 1989. Flora of the Lesser Antilles,
beverages. It is used to flavor candies, liquors, and Leeward and Windward Islands. Dicotyledoneae.
pastries. Probably the principal reason for its Part 3. Vol. 6. Arnold Arboretum, Harvard
popularity is the addictive stimulant alkaloid, University, Jamaica Plain, MA. 658 p.
caffeine (1,3,7-trimethylxanthine), 1.1 to 1.3
percent in the beans, but varying greatly in the Liogier, H.A. 1997. Descriptive flora of Puerto
beverage due to different brewing practices. The Rico and adjacent islands. Vol. 5. Editorial de la
alkaloid is present in the leaf at 0.30 percent, twig, Universidad de Puerto Rico, San Juan, PR.
233
436 p. Steiman, S. 1997. The effects of caffeine in Coffea
arabica L. http://www.grayskies.net/honeybear/
Little, E.L., Jr. and F.L. Wadsworth. 1964. arabica.htm. 2 p.
Common trees of Puerto Rico and the Virgin
Islands. Agriculture Handbook 249. U.S. Stevens, W.D., C. Ulloa-U., A. Pool, and
Department of Agriculture, Washington, DC. O.M. Montiel, eds. 2001. Flora de Nicaragua.
548 p. Monographs in Systematic Botany Vol. 85,
No. 3. Missouri Botanic Garden Press, St.
Pacific Island Ecosystems at Risk. 2002. Coffea Louis, MO. p. 1,911-2,666.
arabica L., Rubiaceae. http://www.hear.org/
pier_v3.3/coara.htm. 3 p.
Willson, K.C. 1985. Climate and soil. In: M.N.
Clifford and K.C. Willson, eds. Coffee: botany,
Smith, R.F. 1985. History of coffee. In: M.N.
biochemistry, and production of beans and
Clifford and K.C. Willson, eds. Coffee: botany,
beverage. The AVI Publishing Company, Inc.,
biochemistry, and production of beans and
Westport, CN. p. 97-107.
beverage. The AVI Publishing Company, Inc.,
Westport, CN. p. 1-12.
Wrigley, G. 1988. Coffee. Longman Scientific
Technical and John Wiley & Sons, Inc. New
York. 639 p.
234
Colubrina arborescens (P. Mill.) Sarg. coffee colubrina
RHAMNACEAE
John K. Francis
235
fertile soils. Four experimental plantings of coffee org/viewing/inyourbackyard/nativeplants.htm.
colubrina were made in Puerto Rico. Complete 11 p.
mortality occurred in the two drier, poorer sites,
but the trees grew 1 m/year in height for the first Francis, J.K. 1998. Tree species for planting in
few years on the better sites (Francis 1998). forest, rural, and urban areas of Puerto Rico.
Mortality probably occurred on the drier sites General Technical Report IITF-3. U.S.
because they were overcome by faster-growing Department of Agriculture, Forest Service,
competition. To ensure quick establishment, International Institute of Tropical Forestry, Río
planting of containerized stock for ornamental and Piedras, PR. 82 p.
wildland plantings is recommended. Protection
from weeds, vines, and faster-growing trees for 2 Gilman, E.F. 1999. Colubrina arborescens. Fact
or more years is imperative. Ornamental plants can Sheet FPS-137. University of Florida,
be forced into tree shape by continual pruning of Cooperative Estension Service, Gainesville, FL.
the lower branches (Gillman 1999). 3 p.
Benefits.—Coffee colubrina helps protect the soil, Liogier, H.A. 1990. Plantas medicinales de Puerto
contributes to the aesthetics of the forest, and Rico y del Caribe. Iberoamericana de Ediciones,
furnishes food and cover for wildlife. The plant Inc. 566 p.
attracts abundant insects including bees, wasps,
butterflies, and diurnal moths, which in turn Liogier, H.A. 1994. Descriptive flora of Puerto
provide food for warblers (Dendroica spp.), Rico and adjacent Islands. Vol. 3. Editorial de la
gnatchtchers (Polioptila spp.), kingbirds (Tyrannus Universidad de Puerto Rico. Río Piedras, PR.
spp.), and vireos (Vireo spp.) (Florida Fish and 461 p.
Wildlife Conservation Commission 2003). The
species is considered a honey plant, furnishing Little, E.L., Jr. and F.L. Wadsworth. 1964.
both nectar and pollen to honeybees (Apis Common trees of Puerto Rico and the Virgin
mellifera L.) May through September in the Islands. Agriculture Handbook 249. U.S.
Dominican Republic (Rivas-Laureano 2003). Department of Agriculture, Washington, DC.
Coffee colubrina is planted as a shade tree and 548 p.
ornamental in Florida, Guatemala, Nicaragua, and
El Salvador (Little and Wadsworth 1964, Stevens Long, R.W. and O. Lakela. 1976. A flora of
and others 2001). It is recommended for planting Tropical Florida. Banyon Books, Miami, FL.
in parking lot islands, in large containers, and 962 p.
small lawns, and for borders, screens, and hedges,
as well as for reclamation plantings (Gilman Nelson, G. 1996. The shrubs and woody vines of
1999). A tea made from the leaves and the wood is Florida. Pineapple Press, Inc., Sarasota, FL.
used as a remedy for rheumatism. Extracts are also 391 p.
used for antiseptic baths (Liogier 1990). The shiny
black seeds are used in necklaces in Jamaica. The Rivas-Laureano, S. 2003. Flora apícola
wood is hard and heavy with a specific gravity of Dominicana. http://www.apinetla.com.ar/rdom/
0.7. The sapwood is ivory or light brown, and the editorial-rdo/editorial1.htm. 2 p.
heartwood is yellowish brown. Because it is
resistant to decay, it is used for fence posts and Sargent, C.S. 1923. Manual of the trees of North
was formerly used for marine pilings (Little and America (exclusive of Mexico). Houghton
Wadsworth 1964). Mifflin, Boston, MA. 910 p.
236
Comptonia peregrina (L.) Coult. sweet fern
MYRICACEAE
Paula M. Pijut
237
1993). The nymphs of the Saratoga spittlebug are readily formed (Louis and Torrey 1991). These
(Aphrophora saratogensis Fitch.) feed on sweet sprouts can spread extensively over large areas
fern but do not cause serious damage (Johnson and forming thickets. Juvenile stems (7.6 cm or less in
Lyon 1991). Sweet fern is susceptible to attack by length) root when treated with 3 g/l indole-3-
Botryosphaeria dothidea (Moug.:Fr.) Ces. & butyric acid and placed under mist (Dirr 1998).
DeNot. (causing dieback) and Gymnosporangium Root pieces (10 cm long x 0.2 cm wide, or 5 cm x
ellisii (Berk.) Ellis (alternate host for aecia stage) 1.3 cm) dug in late winter or early spring, placed
(Sinclair and others 1987). Sweet fern is the horizontally at a depth of 1.3 cm in a mixture of
alternate host of the fungus, Cronartium fine sand and sphagnum peat, will develop shoots
comptoniae Arth., that causes sweet fern blister and additional roots (Dirr 1998). Sweet fern can be
rust on hard pines (Farr and others 1989, Sinclair grown in containers and successfully transplanted
and others 1987). The fungus is not seriously (Dirr 1998). Micropropagation of sweet fern has
harmful to sweet fern, but it can reduce the growth been achieved by the induction of root buds in
and even cause death of susceptible pines, such as excised root culture (Louis and Torrey 1991).
jack (Pinus banksiana Lamb.), pitch (P. rigida
Mill.), shortleaf (P. echinata Mill.), and loblolly Growth and Management.—Sweet fern is a slow
(P. taeda L.) (Del Tredici 1996, Snyder 1993). to medium growing shrub that develops a broad,
flat-topped to rounded habit as it spreads and
Reproduction.—Sweet fern flowers are small, colonizes (Dirr 1998). It is hardy in zones two to
inconspicuous, wind-pollinated, and shed large six (USDA Plant Hardiness) and is sexually
amounts of pollen (Hall and others 1976). In the mature in 2 to 3 years (Snyder 1993). Burning and
United States the shrub blossoms in April or early disking following logging will stimulate the
May with fruit maturation by fall. In Canada, growth of sweet fern (Snyder 1993). Sweet fern is
sweet fern blossoms mid-May to mid-June with used for erosion control, low maintenance
fruit maturation from July through September plantings along highway embankments, and as
(Hall and others 1976). Hall and others (1976) naturalistic plantings under power company rights-
determined the mean number of staminate catkins of-way (Del Tredici 1996, Snyder 1993). Sweet
per shoot based on 42 shoots to be 6.43 + 0.48 fern can become weedy in pastures, old fields, and
(SE) and for the pistillate catkins 0.60 + 0.01 (SE). open woods, but can be controlled with herbicides,
The fruit of sweet fern is borne in a bur-like cluster such as dicamba, 2,4-D, and glyphosate (Hall and
of bracts containing on average 5.52 + 0.43 (SE) others 1976, Heyd and others 1987, Snyder 1993).
nuts per bur (Hall and others 1976). The pericarp
is composed of a thin, fleshy outer layer and a Benefits.—Sweet fern is planted as a landscape
hard, thick inner layer (Del Tredici and Torrey plant because of its fern-like, aromatic foliage and
1976). Seeds do not germinate readily but can stems. Because it fixes nitrogen, sweet fern is
remain viable (for as long as 70 years) buried in useful for rehabilitation of disturbed sites. Sweet
the soil (Del Tredici 1977). Sweet fern seeds are in fern has limited use as a food source and cover for
deep primary dormancy when they are shed from wildlife. The fruits are consumed by flickers,
the plant and become incorporated into the soil moose and deer browse sweet fern, and prairie
(Dow and Schwintzer 1999). They enter seasonal chickens and sharp-tailed grouse use it for nesting
cycles of secondary dormancy over time in which cover (Snyder 1993). The leaves can be used for
dormancy is induced during the summer. The potpourri and tea (Stokes 1981). The chemical
seeds are then released from this dormancy by betulin occurs in the leaves, root, and stem of
chilling during the winter. However, most buried sweet fern, and its biological activities have been
seeds still require exposure to strong daily described (Duke 1996).
temperature fluctuations before they germinate
(Dow and Schwintzer 1999). Strong temperature References
fluctuations indicate that shade cover has been
removed, which stimulates the seeds to germinate Callaham, D., P. Del Tredici, and J.G. Torrey.
in an open situation favorable for seedling 1978. Isolation and cultivation in vitro of the
establishment. Fresh seed will germinate (80 actinomycete causing root nodulation in
percent) after scarification and treatment with 500 Comptonia. Science 199: 899-902.
mg/l gibberellic acid (GA3) for 24 hours (Del
Tredici and Torrey 1976). Sweet fern propagates Del Tredici, P. 1977. The buried seeds of
itself vegetatively via lateral roots that form Comptonia peregrina, the sweet fern. Bulletin of
endogenous root buds from which shoot sprouts the Torrey Botanical Club 104: 270-275.
238
Del Tredici, P. 1996. Bulldozers and bacteria: the Johnson, W.T. and H.H. Lyon. 1991. Insects that
ecology of sweet fern. Arnoldia 56: 2-11. feed on trees and shrubs. Cornell University
Press, Ithaca, NY. 560 p.
Del Tredici, P. and J.G. Torrey. 1976. On the
germination of seeds of Comptonia peregrina, Krüssmann, G. 1984. Manual of cultivated broad-
the sweet fern. Botanical Gazette 137: 262-268. leaved trees and shrubs, Volume I, A-D. Timber
Press, Beaverton, OR. 448 p.
Dirr, M.A. 1998. Manual of woody landscape
plants, Their identification, ornamental Louis, I. and J.G. Torrey. 1991. In vitro clonal
characteristics, culture, propagation, and uses. multiplication of the actinorhizal plant
Stipes Publishing, Champaign, IL. 1,187 p. Comptonia peregrina. Plant Cell, Tissue and
Organ Culture 26: 89-96.
Dow, M.A. and C.R. Schwintzer. 1999. Seed
germination, seedling emergence, and seed bank Lynham, T.J., G.M. Wickware, J.A. Mason, and
ecology of sweet fern (Comptonia peregrina H. Krause. 1998. Soil chemical changes and
(L.) Coult.). Canadian Journal of Botany 77: plant succession following experimental burning
1,378-1,386. in immature jack pine. Canadian Journal of Soil
Science 78: 93-104.
Duke, J.A. 1996. Phytochemical Database, U.S.
Department of Agriculture, Agricultural Missouri Botanical Garden. 2002. w3TROPICOS,
Research Service, Beltsville, Maryland. Nomenclatural database. http://mobot.mobot.
http://www.ars-grin.gov/cgi- org/W3T/Search/vast.html. [not paged].
bin/duke/farmacy2.pl.
Schwintzer, C.R. 1989. All field-collected
Farr, D.F., G.F. Bills, G.P. Chamuris, and A.Y. actinorhizae examined on Comptonia peregrina
Rossman. 1989. Fungi on plants and plant and Myrica pensylvanica in Maine are spore
products in the United States. The American negative. Canadian Journal of Botany 67: 1,460-
Phytopathological Society, St. Paul, MN. 1,252 1,464.
p.
Sinclair, W.A., H.H. Lyon, and W.T. Johnson.
Gleason, H.A. and A. Cronquist. 1991. Manual of 1987. Diseases of trees and shrubs. Cornell
vascular plants of northeastern United States and University Press, Ithaca, NY. 575 p.
adjacent Canada. 2nd Edition. The New York
Botanical Garden, Bronx, NY. 910 p. Snyder, S.A. 1993. Comptonia peregrina. In: U.S.
Department of Agriculture, Forest Service,
Hall, I.V., L.E. Aalders, and C.F. Everett. 1976. Rocky Mountain Research Station, Fire Sciences
The biology of Canadian weeds. 16. Comptonia Laboratory. Fire Effects Information System.
peregrina (L.) Coult. Canadian Journal of Plant http://www.fs.fed.us/database/feis/plants/shrub/c
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Heyd, R.L., R.L. Murray, and L.F. Wilson. 1987. Stokes, D.W. 1981. The natural history of wild
Managing Saratoga spittlebug in pine shrubs and vines, eastern and central North
plantations by suppressing sweet fern. Northern America. Harper and Row Publishers, New
Journal of Applied Forestry 4: 16-17. York. 246 p.
Hurd, T.M. and C.R. Schwintzer. 1997. Formation Ziegler, H. and R. Huser. 1963. Fixation of
of cluster roots and mycorrhizal status of atmospheric nitrogen by root nodules of
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689.
239
Conocarpus erectus L. button mangrove
COMBRETACEAE
John K. Francis
240
throughout the year in México (Pennington and Puerto Rican trees and shrubs: second
Sarukhan 1968) and from March to September in installment. Research Note SO-374. Southern
Florida (Nelson 1996). Tomlinson (1986) Forest Experiment Station, U.S. Department of
maintains that this species is dioecious. It is a Agriculture, Forest Service, New Orleans, LA.
consistent and abundant seed producer throughout 5 p.
most of its range. The fruit heads may contain from
35 to 56 fruits each (Masís and others 1999). Seeds Gilman, E.F. and D.G. Watson. 1993. Conocarpus
collected in Puerto Rico averaged 250,000 erectus, buttonwood. Fact Sheet ST-179. U.S.
seeds/kg. Seeds from this collection began Forest Service and Southern Group of State
germinating on moist filter paper in 9 days but Foresters, Gainesville, FL. 3 p.
were only 12 percent viable (Francis and
Rodríguez 1993). The seeds are dispersed by water Howard, R.A. 1989. Flora of the Lesser Antilles,
(Pacific Island Ecosystems at Risk 2002). Button Leeward and Windward Islands. Vol. 5. Arnold
mangrove can be propagated by partially burying Arboretum, Harvard University, Jamaica Plain,
large stakes in moist ground for living fence posts MA. 604 p.
(Little and Wadsworth 1964).
Instituto Nacional de Biodiversidad. 2002. Lista de
Growth and Management.—Button mangrove especímenes de Conocarpus erectus. http://
has a medium growth rate (Gilman and Watson www.inbio.ac.cr/bims/k03/p13/c045/o0264/f016
1993) and may live for several decades. 17/g007434/s021856.htm. 3 p.
Information is not available on wildland plantation
establishment. The planting of potted or Liogier, H.A. 1990. Plantas medicinales de Puerto
containerized seedlings seems the safest approach. Rico y del Caribe. Iberoamericana de Ediciones,
The species is not aggressive or invasive and only Inc., San Juan, PR 566 p.
requires control when land is converted to other
uses. Liogier, H. A. 1994. Descriptive flora of Puerto
Rico and adjacent Islands. Vol. 3. Editorial de la
Benefits.—Button mangrove provides food and Universidad de Puerto Rico, Río Piedras, PR.
cover for wildlife (various species of crabs and 461 p.
insects). It protects the soil during storm surges and
helps “fix” dunes (Popp and others 1989). Both Little, E.L., Jr. and F.H. Wadsworth. 1964.
button mangrove and silver buttonwood (C. erectus Common trees of Puerto Rico and the Virgin
var. sericeus Griseb.) are widely planted as Islands. Agriculture Handbook 249. U.S.
ornamentals in yards, parking lots, streets and Department of Agriculture, Forest Service,
parks (Gilman and Watson 1993, Nelson 1996). It Washington, DC. 548 p.
can be trained to form hedges. Potted plants are
used to form bonsai (Gilman and Watson 1993). Masís, A, D. Pérez, F. Chavarría, R. Espinoza, A.
Button mangrove wood is reported to be ideal for Guadamuz. 1999. Species page de Conocarpus
smoking fish and meat (Gilman and Watson 1993). erectus (Combretaceae), 14 April 1999. Species
The wood is durable but susceptible to dry-wood Home Pages. Area de Conservación Guanacaste,
termites [Cryptotermes brevis (Walker)] and is Costa Rica. http://www.acguanacaste.ac.cr/
used to make railroad ties, posts, for turnery, boat paginas_especie/plantae_online/magnoliophyta/
building, fuel, and charcoal. The bark and leaves combretaceae/conocarpus_erectus/c_erectus_14’
have been used in tannery (Little and Wadsworth ... 6 p.
1964). Bark from Belize contains 18 percent tannin
(Burkill 1985). Extracts of the bark are used to Nelson, G. 1996. The shrubs and woody vines of
treat bleeding gums, vaginal bleeding, colic, and Florida. Pineapple Press, Inc., Sarasota, FL.
skin ulcers (Liogier 1990). 391 p.
241
Agricultura y Ganadaría, México D.F., México. Stevens, W.D., C. Ulloa-U., A. Pool, and O.H.
413 p. Montiel, eds. 2001. Flora de Nicaragua.
Monographs of Systematic Botany Vol. 85, No.
Popp, M., U. Lüttge, W.J. Cram, M. Díaz, H. 1. Missouri Botanical Garden Press. p. 1-943.
Griffiths, H.J.S. Lee, E. Medina, C. Schäfer, K.-
H. Stimmel, and B. Thonke 1989. Water Tomlinson, P.B. 1986. The botany of mangroves.
relations and gas exchange of mangroves. New Cambridge University Press, Cambridge, UK.
Phytologist 111: 293-307. 419 p.
242
Corchorus hirsutus L. jack-switch
TILIACEAE
Synonyms: none
John K. Francis
Ecology.—Jack-switch is intolerant of shade and
grows in open areas or openings in low forest. It
cannot endure severe competition. The species
usually grows near the coast or inland on dry, often
excessively-drained sites. These are usually coastal
sands or rocky ridges and hillsides over limestone,
igneous, and metamorphic (including ultramaphic)
rocks. Jack-switch occurs as scattered plants or as
open stands with other species of similar size.
Because cattle do not eat or rarely eat the foliage,
the species benefits from over-grazing. It also
invades eroded and physically disturbed soils.
Reproduction.—Jack-switch blooms
continuously, except during periods of drought.
The species is insect pollinated (Marcano-Fondeur
1973). Capsules collected in Puerto Rico averaged
General Description.—Jack-switch, also known 28.0 seeds/capsule. Air-dried seeds from that
as wooly corchorus, mallet, cadillo, and collection averaged 1.109 million seeds/kg. Sown
malvavisco, is an upright woody to semiprostrate without pretreatment on the surface of wet peat, 10
shrub usually about 1 m in height but sometimes percent germinated over a 4-month period
reaching 2 m and 1.5 cm in basal diameter. It (author’s observation). Dispersal undoubtedly
usually has a single main stem and a somewhat occurs by wind and water; specialized means of
sparse, open crown. Plants examined by the author seed dispersal are unknown. Seedlings in wildland
had a deep taproot and fine lateral roots at all sites in Puerto Rico vary from common to rare.
levels. All parts except older branches and stems Plants apparently do not renew themselves by
are densely pale scruffy tomentulose with stellate resprouting after stem death. Reaction to fire is
hairs. The alternate leaves are ovate to oblong- unknown.
lanceolate, 2 to 6 cm long, have crenate to serrate
margins and obtuse to acute tips, and have petioles Growth and Management.—Jack-switch plants
2 to 7 mm long. The leaves, which have a gray- have a moderate to slow growth rate and live about
green color, tend to wilt rather than defoliate 5 years. Mechanical soil disturbance near seed
during dry seasons. Yellow flowers occur in groups sources and heavy grazing would probably lead to
of two to eight on axillary peduncles opposite the natural establishment of the shrubs. Jack-switch is
leaves. The four-chambered capsules are seldom common enough to warrant control, but if
ellipsoidal, about 12 mm long, and contain a needed, mowing may be sufficient.
number of 1.5- to 2-mm black seeds (Britton and
Millspaugh 1962, Howard 1989, Liogier 1994). Benefits.—Jack-switch furnishes cover for wildlife
and helps protect the soil. The plant is attractive
Range.—Jack-switch is native throughout the enough to be used as an ornamental in natural
West Indies and has been reported from Mexico landscaping, although that use is not yet reported.
through Central and South America and in It is a honey plant (Marcano-Fondeur 1973). The
northeastern Africa (Britton and Millspaugh 1962, leaves are cooked and eaten, and made into tea in
Howard 1989, Liogier 1994). Herbarium St. Croix, U.S. Virgin Islands. Branches were
specimens exist for Bolivia and Paraguay formerly used to make brooms (Garland 2002).
(Missouri Botanical Garden 2002). The species is
present as an exotic in the southern tip of Florida
(Institute of Systematic Botany 2002).
243
References http://www.plantatlas.usf.edu/main.asp?plantID
=4142. [not paged].
Britton, N.L. and C.F. Millspaugh. 1962. The
Bahama flora. The New York Botanical Garden, Liogier, H.A. 1994. Descriptive flora of Puerto
New York. 695 p. Rico and adjacent Islands. Vol. 3. Editorial de la
Universidad de Puerto Rico. Río Piedras, PR.
Garland, K. 2002. UVI Wetland Reserve plants: 461 p.
Corchorus hirsutus (jack-switch).
http://rps/uvi/edu/VIMAS/jackswitch.htm. 1 p. Marcano-Fondeur, E.J. 1973. La flora apícola de la
Republica Dominicana. http://marcano.
Howard, R.A. 1989. Flora of the Lesser Antilles, freeservers.com/nature/studios/apicola/dicotsp.
Leeward and Windward Islands. Vol. 5. Arnold html. 11 p.
Arboretum, Harvard University, Jamaica Plain,
MA. 604 p. Missouri Botanical Garden. 2002. W3-Specimen
Data Base: Current specimen list for Corchorus
Institute of Systematic Botany. 2002. Atlas of hirsutus. http://www.mobot.mobot.org/cgi-
Florida Vascular Plants: Corchorus hirsutus. bin/search_vast. [not paged].
University of South Florida, Tampa, FL.
244
Cordia lima (Desv.) Roemer & J.A. Schultes lija
BORAGINACEAE
John K. Francis
245
References Liogier, H.A. 1995. Descriptive flora of Puerto
Rico and adjacent islands. Vol. 4. Editorial de la
Grisebach, A.H.R. 1963. Flora of the British West Universidad de Puerto Rico, San Juan, PR.
Indian Islands. J. Cramer-Weinheim, New York. 617 p.
789 p.
246
Cordia polycephala (Lam.) I.M. Johnston black sage
BORAGINACEAE
John K. Francis
247
consumed by birds that disperse the seeds. SO-58. U.S. Department of Agriculture, Forest
Seedlings are common but rarely abundant. Service, Southern Forest Experiment Station,
New Orleans, LA. 331 p.
Growth and Management.—Black sage achieves
1 m or more of stem extension per year. Stems may Centro Nordestino de Informações Sobre Plantas.
grow to 1.5 cm of diameter and reach 5 m in height 2002. Base de dados CNIP, Checklist das
or extension from the point of rooting (Acevedo- plantas do Nordeste-Nomes provisórios-Versão
Rodríguez 1985). One 1.5-cm stem was observed 12: Cordia polycephala (Lam.) Johnst.
to have five growth rings. Black sage can live at Universidade Federal de Pernambuco.
least 5 years and perhaps much longer by http:150.161.125.13/pnechk/taxa/1057.shtml.
sprouting, which it readily does when injured. No 2 p.
planting, management, or control experience has
been published. In the absence of other control Howard, R.A. 1989. Flora of the Lesser Antilles.
information, grubbing out individual plants or spot Vol. 6. Arnold Arboretum, Harvard University,
treating with broad-leaf herbicides is Jamaica Plain, MA. 658 p.
recommended.
Liogier, H.A. 1995. Descriptive flora of Puerto
Benefits.—Black sage helps protect the soil and Rico and adjacent islands. Vol. 4. Editorial de la
furnishes food and cover for wildlife. The fruits are Universidad de Puerto Rico, San Juan, PR.
edible. It is a minor weed in pastures (Vélez and 617 p.
van Overbeek 1950).
Vélez, I. and J. van Overbeek. 1950. Plantas
References indeseables en los cultivos tropicales. Editorial
Universitaria, Río Piedras, PR. 497 p.
Acevedo-Rodríguez, P. 1985. Los bejucos de
Puerto Rico. Vol. 1. General Technical Report
248
Cornus sericea L. redosier dogwood
CORNACEAE
Paula M. Pijut
Fosberg and f. stolonifera (Michx.) Fosberg
(Missouri Botanical Garden 2002).
249
are insect-pollinated (Crane 1989). In the United thick, extensive root system (Crane 1989).
States the shrub blossoms from late May to June Redosier dogwood is relatively tolerant to high
(June to August in the Northern United States) salinity tailings waters produced as a result of oil
with fruit maturation by fall. The fruit is a drupe and sand mining, making it useful for reclamation
containing a single two-celled and usually two- of sites (Renault and others 1999, 2001). The bark,
seeded stone, but only one seed is fully developed stems, leaves, roots, and fruit of redosier dogwood
(Brinkman 1974). Redosier dogwood seeds have a were used by Native Americans for a variety of
hard seed coat and dormant embryos (Crane 1989). uses including basket weaving, dyes, and tonics
Seeds do not germinate readily (can remain viable (Stevens and Dozier 2002). Redosier dogwood
in seedbanks) and require cold stratification and thickets provide cover for birds and other small
sometimes scarification in order for germination to wildlife. It is an important food source and winter
occur (Crane 1989). Fresh seed can be fall sown or browse for deer, elk, moose, bear, cottontail
stratified for 60 to 90 days at 5 ºC (Brinkman rabbits, snowshoe hares, grouse, quail, and other
1974). Acharya and others (1992) found that small birds and mammals (Crane 1989).
native (Alberta, Canada) redosier dogwood seed
germination was strongly influenced by the References
population from which seeds were collected, and
the yearly precipitation in those areas. Redosier Acharya, S.N., C.B. Chu, R. Hermesh, and G.B.
dogwood propagates itself vegetatively via stolons Schaalje. 1992. Factors affecting red-osier
(stems just on or under the ground) and by dogwood seed germination. Canadian Journal of
layering forming dense thickets. Plants are also Botany 70: 1,012-1,016.
produced from the roots, the bases of dying
branches, and injured stems (Crane 1989). Brinkman, K.A. 1974. Cornus L. dogwood. In:
Cuttings (with leaves) root readily when treated C.S. Schopmeyer, tech. coord. Seeds of woody
with 1 g/l indole-3-butyric acid (Dirr 1998). plants in the United States. Agriculture
Hardwood cuttings root (90 to 100 percent) when Handbook 450, U.S. Department of Agriculture,
placed in the field in late winter without a rooting Forest Service, Washington, D.C. p. 336-342.
treatment (Dirr 1998). Redosier dogwood can be
grown as bare-root, containerized, or balled-and- Brown, D.A., M.T. Windham, and R.N. Trigiano.
burlapped plants (Dirr 1998). 1996. Resistance to dogwood anthracnose
among Cornus species. Journal of Arboriculture
Growth and Management.—Redosier dogwood 22(2): 83-86.
is a fast (vigorous) growing shrub that develops a
loose, broad-spreading, rounded habit with Crane, M.F. 1989. Cornus sericea. In: U.S.
horizontal branches at the base (Dirr 1998). It is Department of Agriculture, Forest Service,
hardy in zones 2 to 7 (USDA Plant Hardiness) and Rocky Mountain Research Station, Fire Sciences
is sexually mature in 3 to 4 years (Crane 1989). Laboratory. Fire Effects Information System.
Redosier dogwood generally increases following http://www.fs.fed.us/database/feis/plants/shrub/c
fire (Crane 1989). Coppicing stimulates new, omper/index.html.
vigorous stem growth. Pruning and burning
redosier dogwood helps produce long straight Dirr, M.A. 1998. Manual of woody landscape
stems. Redosier dogwood is used for soil plants, their identification, ornamental
stabilization, reclamation sites, residential characteristics, culture, propagation, and uses.
landscape plantings, mass plantings along highway Stipes Publishing, Champaign, IL. 1,187 p.
banks, parks, and golf courses (Crane 1989, Dirr
1998, Renault and others 2001). Redosier Gleason, H.A. and A. Cronquist. 1991. Manual of
dogwood can be controlled by spraying with vascular plants of Northeastern United States
mixtures of 2, 4-D and dicamba (Crane 1989). and adjacent Canada. 2nd Edition. The New York
Botanical Garden, Bronx, NY. 910 p.
Benefits.—Redosier dogwood is planted as a
landscape plant because of its bright red stems, Johnson, W.T. and H.H. Lyon. 1991. Insects that
which are especially beautiful contrasting against a feed on trees and shrubs. Cornell University
snowy landscape. Because it is easy to establish Press, Ithaca, NY. 560 p.
and grows rapidly, redosier dogwood is useful for
rehabilitation of disturbed sites. It is an excellent Jones, R.K. and D.M. Benson. 2001. Diseases of
soil stabilizer along stream banks because of its woody ornamentals and trees in nurseries. The
250
American Phytopathological Society, St. Paul, plants to high salinity oil sands tailings. Journal
MN. 482 p. of Environmental Quality 28: 1957-1962.
Missouri Botanical Garden. 2002. w3TROPICOS, Sinclair, W.A., H.H. Lyon, and W.T. Johnson.
Nomenclatural database. http://mobot. 1987. Diseases of trees and shrubs. Cornell
mobot.org/W3T/Search/vast.html University Press, Ithaca, NY. 575 p.
Renault, S., C. Croser, J.A. Franklin, and J.J. Stevens, M. and I. Dozier. 2002. Redosier
Zwiazek. 2001. Effects of NaCl and Na2SO4 on dogwood. In: U.S. Department of Agriculture,
red-osier dogwood (Cornus stolonifera Michx) Natural Resources Conservation Service, The
seedlings. Plant and Soil 233: 261-268. PLANTS Database, National Plant Data Center,
Baton Rouge, LA. http://plants.usda.gov. [not
Renault, S., E. Paton, G. Nilsson, J.J. Zwiazek, and paged].
M.D. MacKinnon. 1999. Responses of boreal
251
Coursetia caribaea (Jacq.) Lavin añil falso
FABACEAE
John K. Francis
252
0.0000 g/seed. They were sown without References
pretreatment on filter paper and yielded only 10
percent germination in 206 days. After mechanical Dubey, H.D., R. Woodbury, and R.L. Rodríguez.
scarification, 98 percent of the remainder of the 1972. New records of tropical legume
seeds germinated within 12 days. Grazing animals nodulation. Botany Gazette 133(1): 35-38.
are the principal seed vectors. Seedlings are
common and well-disbursed. Large plants are Howard R.A. 1988. Flora of the Lesser Antilles,
prolific seed producers. Even small and suppressed Leeward and Windward Islands.
plants produce a few seeds. Dicotyledoneae, Part 1. Vol. 4. Arnold
Arboretum, Harvard University. Jamaica Plain,
Growth and Management.—Añil falso grows up MA. 673 p.
to about 0.5 m per year in height. Plants live from 2
to several years. Wise management of the species International Plant Index. 2001. International plant
must preclude all but light grazing. Plantations for names index query: Coursetia caribaea
range improvement or environmental restoration www.ipni.org/ipni/query-ipni.html. 17 p.
could probably be established by sowing scarified
seed into cultivated understory sites before or at the Liogier, H.A. 1988. Descriptive flora of Puerto
beginning of the rainy season. Rico and adjacent islands, Spermatophyta. Vol.
2. Editorial de la Universidad de Puerto Rico,
Benefits.—Añil falso, when available, is grazed Río Piedras, PR. 481 p.
heavily by cattle. The species improves the soil by
fixing nitrogen. Liogier, H.A. and L.F. Martorell. 2000. Flora of
Puerto Rico and adjacent islands. Second ed.,
Rev. Editorial fe la Universidad de Puerto Rico,
Río Piedras, PR. 382 p.
253
Crataegus douglasii Lindl. black hawthorn
ROSACEAE
John K. Francis
254
Growth and Management.—Growth of newly References
established seedlings is slow (Habeck 1991).
Particulars on natural stand management are Borialforest.com 2003. Shrub species of the
lacking. Fruits are ripe when they are black and World’s boreal forests: Crataegus douglasii,
lustrous (Habeck 1991) and must be picked by black hawthorn. http://www.borealforest.org/
hand or clipped with a pruning pole. Pulp should world/herbs_shrubs/black_hawthorn.htm. 2 p.
be removed from the seeds by maceration and then
seeds should be air dried before sowing or storage. Brinkman, K.A. 1974. Crataegus L., hawthorn. In:
Seeds should be scarified with H2SO4 and C.S. Schopmeyer, tech. coord. Seeds of woody
stratified at 1 °C for 84 to 112 days before sowing. plants in the United States. Agriculture
Brinkman (1974) found that germination reached Handbook 450. U.S. Department of Agriculture,
about 30 percent in 35 to 45 days. Another source Forest Service, Washington, DC. p. 356-360.
cites 50 to 80 percent germination (Habeck 1991).
Dumroese and others (1997) report germination Dumroese, R.K., K.M. Hutton, and D.L. Wenny.
from 20 to 80 percent for lots treated identically. 1997. Propagating woody riparian plants in
Seedlings of most hawthorn species develop long nurseries. In: T.D. Landis and J.R. Thompson,
taproots and should not be kept in the nursery for tech. coords. National proceedings, Forest and
more than 1 year (Habeck 1991). Pruning should Conservation Nursery Associations. Gen. Tech.
be avoided and final target height of seedlings Rep. PNW-GTR-419. Pacific Northwest
should be 25 cm (Dumroese and others 1997). Research Station, U.S. Department of
Agriculture, Forest Service, Portland, OR. p. 71-
Benefits.—Black hawthorn contributes to the 76.
beauty of the riparian vegetation, helps protect the
soil from streambank and overflow erosion, and Habeck, R.J. 1991. Crataegus douglasii. In: U.S.
offers benefits to fauna. Although not as “showy” Department of Agriculture, Forest Service,
as some other hawthorns, the species is hardy and Rocky Mountain Research Station, Fire Sciences
has been used to a limited extent as an ornamental. Laboratory, Fire Effects Information System.
It makes excellent hedges and natural barriers, and http://www.fs.fed.us/database/feis/plants/shrub/
is planted as a wildlife cover and food plant, and cradou/all.html. 15 p.
for stream bank stabilization (Hansen 2003).
Livestock readily eat black hawthorn foliage but Hansen, W.W. 2003. Native plants of the
the thorny structure of shrubs and small trees Northwest: black hawthorn, Douglas hawthorn,
makes it difficult for them to utilize it well. They western thorn apple, and Douglas thorntree
prefer shrubs less than 1 m in height which they (Crataegus douglasii). http://www.nwplants.
may hedge. Mule deer also browse it. With 3.7 com/plants/trees/rosaceae/crataegus_douglasii/
percent protein, 3.8 percent fat, 19.3 percent fiber, index. html. 5 p.
and 4.0 percent ash, the species is moderately
palatable to browsing animals. Deer, upland game Moerman, D.E. 1986. Medicinal plants of Native
birds, song birds, and rodents eat the fruits. Black America. Technical Reports 19. University of
hawthorn makes excellent escape and nesting Michigan Museum of Anthropology, Ann
cover for wildlife (Habeck 1991). The fruits are Arbor, MI. 534 p.
made into pies, jellies, and jams (Hansen 2003).
Native Americans ate the fruits, sometimes dried Natural Resources Conservation Service. 2003.
and mixed with fat and other ingredients. The Plant profile: Crataegus douglasii Lindl., black
wood is hard and heavy and was used for tools and hawthorn. http://plants.usda.gov/cgi_bin/
weapons (Washington State Department of plant_profile.cgi?symbol=CRDO2. 5 p.
Transportation 2003). Native Americans used the
thorns for piercing ears, lancing boils, and making Sargent, C.S. 1923. Manual of the trees of North
fish hooks (Hansen 2003). They prepared a America (exclusive of Mexico). Houghton
poultice of chewed leaves to relieve swelling and Mifflin, Boston, MA. 910 p.
took decoctions of sap, bark, wood, and root for
stomach problems (Moerman 1986). Soper, J.H. and M.L. Heimburger. 1982. Shrubs of
Ontario. Royal Ontario Museum, Toronto,
Ontario, Canada. 495 p.
255
Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
trees and shrubs. Agriculture Handbook 410. Welsh, S.L. 1974. Anderson’s flora of Alaska.
U.S. Department of Agriculture, Forest Service, Brigham Young University Press, Provo, Utah.
Washington, DC. 265 p. 724 p.
Washington State Department of Transportation. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
2003. Crataegus douglasii, black hawthorne. Higgins. 1987. A Utah flora. Great Basin
http://www.wsdot.wa.gov/environment/eao/ Naturalist Memoirs 9. Brigham Young
culres/ethbot/a-c/Crataegus.htm. 2 p. University Press, Provo, UT. 894 p.
256
Critonia portoricense (Urban) Britt. & Wilson guerrero
ASTERACEAE
John K. Francis
257
a topical alcoholic rub (Liogier 1990). Guerrero Liogier, H.A. 1990. Plantas medicinales de Puerto
contributes to biodiversity of the forests in which it Rico y del Caribe. Iberoamericana de Ediciones,
grows, furnishes cover and nesting sites for birds, Inc., San Juan, PR. 566 p.
and helps protect and stabilize soil in disturbed
areas. Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
References Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Liogier, H.A. 1997. Descriptive flora of Puerto Washington, DC. 1,024 p.
Rico and adjacent islands. Vol. 5. Editorial de la
Universidad de Puerto Rico, San Juan, PR. Wiedenfeld, H., R. Guerrero, and E. Roeder. 1995.
436 p. Pyrrolizidine alkaloids from Eupatorium
portoricense. Planta Medica 61(4): 380-381.
258
Crossopetalum rhacoma Crantz maidenberry
CELASTRACEAE
John K. Francis
259
are not common. Plants readily sprout when cut or Howard, R.A. 1989. Flora of the Lesser Antilles,
burned. Leeward and Windward Islands. Vol. 5. Arnold
Arboretum, Harvard University, Jamaica Plain,
Growth and Management.—Maidenberry grows MA. 604 p.
slowly and may live for several decades. No
management experience has been published.
International Reptile Conservation Foundation.
However, cattle grazing to reduce competition
2002. Effects of introduced ungulates on the
from grass, herbs, and vines will probably
iguana of Anegada. http://www.cyclura.com/
encourage reproduction and survival.
article/articleview/248/1/6/. 12 p.
Benefits.—Maidenberry furnishes food and cover
Liogier, H.A. 1990. Plantas medicinales de Puerto
for wildlife and protects the soil. It is listed as one
Rico y del Caribe. Iberoamericana de Ediciones,
of the food plants for the endangered Anegada
Inc., San Juan, PR 566 p.
iguana, Cyclura pinguis (International Reptile
Conservation Foundation 2002). The wood of
Liogier, H.A. 1994. Descriptive flora of Puerto
maidenberry is light brown and hard, but is used
Rico and adjacent Islands. Vol. 3. Editorial de la
only for fuel. All the plant tissues, especially the
Universidad de Puerto Rico. 461 p.
roots, are diuretic and are used to promote
urination and to treat kidney and bladder infections
Little, E.L., Jr., R.O. Woodbury, and F.H.
(Liogier 1990).
Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
References
449. U.S. Department of Agriculture,
Washingtion, DC. 1,024 p.
Alarcón, C. 2001. Paraguana xeric scrub
(NT1313). World Wildlife Fund.
Nelson, G. 1996. The shrubs and woody vines of
http://worldwildlife.org/wildworld/profiles/terre
Florida. Pineapple Press, Inc., Sarasota, FL.
strial/nt1313_full.html. 14 p.
391 p.
Florida Fish and Wildlife Conservation
TreeGuide Inc. 2002. Florida crossopetalum,
Commission. 1997. Florida’s endangered
Crossopetalum rhacoma Crantz, Celestraceae.
species, threatened species and species of
http://www.treeguide.com/nn/Species.asp?
special concern. http://floridaconservation.org/
SpeciesID=345&Region=NorthAmerican. 2 p.
pubs/endanger.html. [not paged].
260
Crotalaria lotifolia L. cascabelillo axilar
FABACEAE
John K. Francis
261
Growth and Management.—Once established, References
cascabelillo axilar adds about 0.5 m in height each
year and loses most of it to die-back the following Breckon, G.J. 2002. Preliminary checklist for
dry season. Individual plants probably live 1 to 3 Sierra Bermeja. http://www.uprm.edu/biology/
years or more. No management experience has profs/breckon/herbarium/FLORABERMEJA.
been published. Scarified seed sown and htm. [not paged].
incorporated into prepared ground or seed spots at
the start of the wet season may offer a good chance Howard R.A. 1988. Flora of the Lesser Antilles,
of success. Leeward and Windward Islands.
Dicotyledoneae, Part 1. Vol. 4. Arnold
Benefits.—Cascabelillo axilar contributes in a Arboretum, Harvard University. Jamaica Plain,
minor way to standing biomass, cover for wildlife, MA. 673 p.
the aesthetics of the forest understory, and it helps
protect the soil. The species is browsed by cattle Liogier, H.A. 1988. Descriptive flora of Puerto
and consequently is more common in protected Rico and adjacent islands, Spermatophyta. Vol.
areas. Cascabelillo axilar is present in moderately 2. Editorial de la Universidad de Puerto Rico,
grazed areas but absent from seriously overgrazed Río Piedras, PR. 481 p.
areas.
262
Crotalaria spectabilis Roth showy rattlebox
FABACEAE
John K. Francis
263
The flowers are visited by honey bees and other Crotalaria spectabilis. http://ppws.vt.edu/scott/
insects (McGregor 1976). A collection of seeds weed_id/cvtsp.htm. 2 p.
from Puerto Rico averaged 0.0154 + 0.0000 or
65,000 seeds/kg. Only 3 percent of unscarified seed Burkill, H.M. 1995. The useful plants of West
placed on moist blotter paper germinated in 6 Tropical Africa. Vol. 3. Royal Botanic Garden,
months. However, 99 percent of scarified seed Kew, UK. 857 p.
germinated between 4 and 7 days after sowing.
Germination is epigeal. When sufficiently dry, Damron, B.L. and J.P. Jacob. 2001. Toxicity to
pods spring open, flinging the seeds a short poultry of common weed seeds. University of
distance. Florida, Cooperative Extension Service.
http://edis.ifas.ufl.edu/BODY_PS052. 7 p.
Growth and Management.—Showy rattlebox has
a moderate to fast growth rate and may reach a Howard R.A. 1988. Flora of the Lesser Antilles,
meter of height by 1 year old. Plants appear to live Leeward and Windward Islands.
2 or 3 years in Puerto Rico. Parrotta (2001) refers Dicotyledoneae, Part 1. Vol. 4. Arnold
to the species as biennial. Plantations are Arboretum, Harvard University. Jamaica Plain,
established by sowing scarified seed into prepared MA. 673 p.
ground.
International Legume Database and Information
Benefits and Detriments.—Showy rattlebox is a Service. 2002. ILDIS Legumes of the world:
beautiful plant and contributes to the aesthetics of Crotalaria spectabilis Roth. http://ildis.org/
wildlands and gardens where it is planted. It serves LegumeWeb/6.00/taxa/4077.shtml. 4 p.
as a nurse species during early reforestation and
helps protect the soil. Showy rattlebox is used as a Jain, S.K. and R.A. De Filipps. 1991. Medical
green manure crop, particularly where rotation for plants of India. Vol. 1. Reference Publications,
control of nematodes is important (Zago and others Inc., Algonac, MI. 408 p.
2002). It has also been grown as a fodder plant.
This has largely been abandoned because of the Liogier, H.A. 1988. Descriptive flora of Puerto
toxic alkaloids (principally monocrotaline) it Rico and adjacent islands, Spermatophyta. Vol.
contains. Concentrations are reported as: leaves 2. Editorial de la Universidad de Puerto Rico,
0.008 percent, seedpods 0.366 percent, and seeds Río Piedras, PR. 481 p.
1.958 percent (Burkill 1995). Chickens, turkeys,
and quail are very sensitive. Concentrations of Long, R.W. and O. Lakela. 1976. A flora of
crotalaria alkaloids between 0.01 and 0.1 percent of Tropical Florida. Banyan Books, Miami, FL.
the diet have adverse effects and 0.3 percent is fatal 962 p.
(Damron and Jacob 2001). Horses, cattle, and
swine are also sensitive. Sheep, goats, and dogs are McGregor, S.E. 1976. Insect pollination of
more resistant. Effects can be both acute and cultivated crop plants. Internet printing by the
chronic and include bloody diarrhea, anemia, USDA Agriculture Research Service, http://
jaundice, hair loss, and unthriftiness. There is no gears.tucson.ars.ag.gov/book/chap9/crotalaria.
known treatment (Alabama Cooperative Extension html. [not paged].
Service 2002). In herbal medicine, extracts of the
whole plant are used to treat impetigo and scabies Parrotta, J.A. 2001. Healing plants of Peninsular
(Jain and De Filipps 1991), as an antiseptic for India. CABI Publishing, Oxon, UK and New
cuts, and to treat intestinal worms (Parrotta 2001). York. 917 p.
264
Zago, C., V.P. de Melo and W.R. Maluf. 2002. A
aplicação da adubação verde em hortaliças.
Boletim Técnico de Hortaliças 16.
Departamento de Agricultura, Universidade
Federal de Lavras. http://www2.ufla.br/
~wrmaluf/bth016/bth016.html. 5 p.
265
Cuphea hyssopifolia Kunth false heather
LYTHRACEAE
John K. Francis
266
older plants can be promoted by mowing at about 5 Howard, R.A. 1989. Flora of the Lesser Antilles,
cm height, which cuts away the old plant structure Leeward and Windward Islands. Vol. 5. Arnold
from the newly layered branches. Nursery plants Arboretum, Harvard University, Jamaica Plain,
should be watered during dry periods or as needed MA. 604 p.
and given frequent light fertilizer applications.
False heather is susceptible to nematodes. Use of Liogier, H.A. 1994. Descriptive flora of Puerto
nematode-free media is recommended (Watkins Rico and adjacent Islands. Vol. 3. Editorial de la
and Sheehan 1975). Universidad de Puerto Rico. Río Piedras, PR.
461 p.
Benefits.—False heather has been cultivated
widely as an ornamental. It is used as a potted Pope, W.T. 1968. Manual of wayside plants of
plant, for low borders, and as ground cover in small Hawaii. Charles E. Tuttle Co, Publishers,
beds. False heather may become a weed in warm Rutland, VT. 289 p.
climates (Turner and Wasson 1997) and has
become a serious weed in some parts of Hawaii Turner, R.G., Jr. and Wasson, E. 1997. Botanica.
(Pope 1968). Barns and Noble, Inc. New York. 1,020 p.
267
Curatella americana L. chaparro
DILLENIACEAE
Synonyms: none
268
Genetics.—Comparative flavonoid chemistry of chaparro is one of the most common woody
chaparro and other members of the Dilleniaceae species.
family resulted in reticulate distribution of the Chaparro grows in many soil types. The
substances identified and did not permit most favorable are well-drained soils. Examples of
recognition of taxa beyond the level of genus and soils are red sands, red sandy loams, yellow sands,
family (Gurni and Kubitzki 1981). Woodson and well drained latosols, hard laterite crusts, ill-
Schery (1965) reported pentagyna as a variety of drained latosols, deeply weathered quartz gravel
chaparro in Panamá. and sands, serpentine-derived soils, entisols on
well drained sites, entisols on young alluvial
Ecology.—Chaparro is a savanna plant with a surfaces to oxisols on more ancient ones, alfisols,
uniform spatial distribution (San Jose and others and vertisols (Sarmiento 1983). Vargas Ulate
1991). It is adapted to fire, high air temperature, (2001) attributed the presence of chaparro in
and harsh soil conditions (Pittier 1926, Lundell Guanacaste, Costa Rica, to acid and infertile soils.
1937). Chaparro is among the first invaders after a In Colombia, the loss of the A and B horizons of
fire (Salas Estrada 1993). Medina and Silva (1990) granitic soils in sierra Nevada have led to
describe the savannas where chaparro grows as savannization and the establishment of chaparro in
“steady state (s) regulated by water-fire lands where it normally does not grow (Cavalier
interactions on a background of low nutrient and others 1998).
availability.” However, during the early phases of Chaparro has a deep root system reaching
growth, chaparro is susceptible to fire disturbance the water table in its adult stage. The species is a
(San José and Medina 1975). A low allocation of phreatophyte, so it grows new leaves during the
N to leaves--even young ones--reduces dry season (Sarmiento and Monasterio 1983,
photosynthetic capacity, but it is adaptive to fire as Medina and Francisco 1994).
only a small amount of N is lost when leaves burn Competition between chaparro and other
(Medina and Francisco 1994). plants in the savanna occurs in two stages:
Chaparro grows up to 500 m elevation in establishment and subsequent growth. During
Bolivia (Killeen and others 1993), and Venezuela establishment, grasses may hinder the growth of
(Sarmiento 1983), and it has extended its range in saplings and constitute a risk because of the high
Colombia to up to 500 to 1,500 m due to fire frequency. Chaparro roots face water shortages
savannization of forest lands (Cavalier and others as they grow through the grass root layer, so
1998). Frost appears to limit its northern and establishment is favored by sequences of wet years
southern distribution, and flooding or soil when fire frequency diminishes and soil moisture
saturation also limits its distribution (Sarmiento increases. Eventually, chaparro will out-grow
1983). grasses and shade them out (for example in tree
High acidity, high clay content, low groves).
mineral nutrient concentration, high exchangeable The chlorophyll content of chaparro is
Al, poor drainage, and poor aeration in the deeper low and correlated with low N and high ash
soil horizons favor chaparro (Lundell 1937, concentrations (Medina and Francisco 1994). Leaf
Gottsberger and Morawetz 1986). Chaparro will photosynthesis rate reached maximum values in
grow in depressions. In some sites, chaparro mature leaves during the humid season and
dominates the community and forms dense groves. declined in younger and older leaves and during
For example, within a matrix of flooded savannas the dry season (Sobrado 1996b). Young leaves are
in Bolivia, chaparro forms groves on ant and more efficient in water use than mature leaves.
termite hills and on hummocks (Killeen and others Stomatal conductance, and water loss,
1993). Sahales is the name used to describe groves decline during the dry season. Chaparro leaves do
of chaparro in northern Petén (Lundell 1937). not show photoinhibition. Stomatal conductance
Chaparro grows in a wide range of moisture differences between dry and wet season appear to
conditions from dry to wet savannas. In the be responsible for maintaining a homeostatic
Venezuelan Llanos, San José and Medina (1975) balance of leaf water and thus maintain xylem
report a mean annual rainfall of 1335 mm and a integrity in spite of the species vulnerability to
range of 580 to 1990 mm. A “normal” year has a 4 embolisms (Sobrado 1996a). Predawn water
to 6 month dry season. Beard (1953), Sarmiento potential, minimum leaf water potential, and
and Monasterio (1975), Sarmiento (1983), Medina hydraulic resistance of chaparro, did not show
and Silva (1990), and Medina and Huber (1992) significant differences throughout the year. In spite
report details of the climatic and edaphic of the scleromorphic appearance of leaves,
conditions of tropical American savannas where chaparro appears adapted only to short-term
269
diurnal deficits (Meinzer and others 1983). The capacity is reduced due to high mesophyll
transpiration curves for chaparro during the dry resistance to CO2 transfer (Medina 1982). Growth
season are always bimodal (Medina 1977, 1982) as is seasonal and its flowering is tardy (Sarmiento
plants close their stomata at noontime, and Monasterio 1983). It flowers and produces
independently of water availability (Medina 1967, new leaves during the dry season (Medina 1982).
1983). Transpiration rates are also low. The Leaf development takes 45 days (Nascimento and
dependency of leaf dark respiration on air others 1990), and leaf longevity is about 11
temperature changes seasonally (Medina 1982). months (Montes and Medina 1977). However,
The steepness of the relationship decreases from young, mature, and old leaves overlap in an
December, to February, to March, and to individual (Medina and Francisco 1994). Fruiting
September through November, when the and seed dispersal may continue for a short time
relationship is flat. during the rainy season. Fire does not interrupt this
The fungus Pseudocercospora curatellae cycle as burned leaves fall and within days there is
was isolated from leaves of chaparro (Furlanetto a new flush.
and Dianese 1999). The earthworm Pontoscolex Two hundred randomly selected trees of
roraimensis was associated with chaparro (Guerra chaparro growing in the seasonal savannas of
Torquemada 1994). Epiphytes are not abundant on western Venezuela had a mean stem diameter at
chaparro bark, probably due to its flaky deciduous ground level of 8.3 cm. The largest recorded
nature (Gottsberger and Morawetz 1993). Leaf diameter was 39.8 cm (Sarmiento and Monasterio
herbivory on chaparro was 4.6 percent near 1983). Chaparro wood has no growth rings and its
Cuiaba, Mato Grosso, Brazil (Nascimento and sapwood is not clearly defined (Record and Mell
others 1990). 1924).
Under a regime of annual burning,
Reproduction.—The white flowers have a chaparro will experience die back of all its annual
“disagreeable smell” (Grisebach 1963), occur in growth shoots, and growth will start the next
lateral panicles on defoliated old branches, and are season from adventitious buds that develop from
rarely terminal. Flowers are actinomorphic and vascular cambium of older branches (Sarmiento
hermaphroditic. Four to five petals are white or and Monasterio 1983). As a result of this
pinkish, 5 to 6 mm, free, and imbricate (Liogier response, “the tree behaves as a mechanical
1990). Four to five green sepals are rounded, 5 structure supporting a crop of annual branches”
mm long, persistent, and externally pilose (Schnee (Sarmiento and Monasterio 1983). The vascular
1960). Stamens are numerous, free, and persistent. cambium assumes the role of apical meristem as
The gynoecium has two hirsute carpels of 6 to 7 the main replacement tissue. Chaparro can
mm (Liogier 1990) with longitudinal dehiscence maintain itself as a hemixyle--with annual shoots
(Schnee 1960). They are coned at the base and and woody underground structures--and even
covered by long and abundant hairs. Follicles are complete the reproductive process in this growth
globose, hispid, coned in the lower half, with two form (Sarmiento and Monasterio 1983). Chaparro
seeds (Killeen and others 1993). Flowers and fruits can develop new individuals from injured roots
of chaparro are fasciculate and ramiflorous (Foldats and Rutkis 1975). Usually, saplings that
(Gentry 1993). Chaparro flowers between July and develop from injured roots have higher water
November in Bolivia (Killeen and others 1993) potentials than normal saplings because they are
and December to February in Venezuela (Montes already connected to the water table. Injury to
and Medina 1977). Seeds are oblong, bright, roots will stimulate vegetative regrowth, which in
chestnut brown to black, 3 mm long, with a small turn also facilitates formation of groves (San Jose
membranous hilum (Schnee 1960, Liogier 1983). and others 1991).
Seeds are long-lived and remain buried until the The mineralization of nitrogen, soil
proper conditions for germination occur (Medina organic matter, soil exchangeable Ca, Mg, and K,
and Francisco 1994). Fire induces germination and and total soil nitrogen were higher in soils under
sprouting of stems that are too thick to sprout chaparro trees than soils under savanna grasses.
under normal conditions (Foldats and Rutkis For soil organic matter, K, and cation exchange
1969). Seeds are dispersed by birds (Salas Estrada capacity, chaparro exceeded values measured
1993), including parrots and parakeets who eat under deciduous trees in savannas (García
chaparro fruits (Pérez Arbelaez 1978). Miragaya and others 1994, Sánchez and others
1997). Although chaparro is not an aluminum
Growth and Management.—Chaparro accumulating plant, it exhibits seasonal variation
assimilates carbon all year, but its photosynthetic in its nutrient concentration (Villela and de
270
Lacerda 1992). Potassium, Fe, Cu, and Zn were anti-inflammatory and analgesic activity of
highest in August, Ca and Mg in November, and Curatella americana L. Journal of
Al and Mn in May. The Al and Mn concentration Ethnopharmacology 67:171-177.
of mature chaparro leaves is 171 and 105 parts per
million, respectively (Medina 1977). Bailey, L.H. 1941. The standard cyclopedia of
horticulture. The Macmillan Co. New York.
Benefits.—Chaparro produces good wood with a
reddish hue and specific gravity of 0.85. Record Bastos, M.D.N.D.C. 1984. Floristic survey of
and Mell (1924) and de Matos Araujo and de fields of the state of Para Brazil. 1. The field of
Matos Filho (1977) contain wood anatomy Joannes Marajo Island. Boletim do Museu
information for chaparro. Small stem sizes limit Paraense Emilio Goeldi Serie Botanica 1:67-86.
wood utilization of shrubs and trees. However, the
wood is durable and used for charcoal, fuelwood, Beard, J.S. 1953. The savanna vegetation of
posts, small cabinetwork, and for turning (Record northern tropical America. Ecological
and Mell 1924, Standley and Record 1936, Killeen Monographs 23:149-215.
and others 1993, Salas Estrada 1993). Termites do
not attack chaparro wood (Pérez Arbelaez 1978). Behling, H. 1995. A high-resolution Holocene
Wood utilization increases where trees or shrubs pollen record from Lago do Pires, SE Brazil-
become locally abundant and grow in groves vegetation, climate and fire history. Journal of
(Pittier 1926). People use dry leaves as substitutes Paleolimnology 14:253-268.
for sand paper and for cleaning kitchen utensils or
polishing metals (Killeen and others 1993). Behling, H. and H. Hooghiemstra. 1999.
Chaparro bark produces tannins that provide a Environmental history of the Colombian
gray color when used for dying pelts (Killeen and savannas of the llanos Orientales since the last
others 1993). Ground seeds mixed with chocolate glacial maximum from lake records El Pinal and
flavors the drink in Oaxaca, México (Standley Carimagua. Journal of Paleolimnology 21:461-
1920). Chaparro was one of the most abundant 476.
pollens in honey samples during the months of
January, February, and March from an apiary Cavalier, J., T.M. Aide, C. Santos, A.M. Eusse,
north of Roraima State, Brazil (da Silva and Absy and J.M. Dupy. 1998. The savannization of
2000). Siegel and Paguaga (1991) suggested using moist forests in the Sierra Nevada de Santa
chaparro twigs and pods for prospecting for gold Marta, Colombia. Journal of Biogeography
in Costa Rica. In Venezuela, Clamens and others 25:901-912.
(1999) evaluated chaparro for industrial gum
production and found the species promising. Clamens, C., Leon de Pinto G., F. Rincon, V.
Antonio, and O. Beltran. 1999. The behavior of
Pharmaceuticals.—Chaparro is a medicinal plant Samanea saman and Curatella americana as
used extensively in folk medicine throughout its gum producers. Boletín del Centro de
range. Infusions from leaves and stems are used Investigaciones Biológicas Universidad del Zulia
for arthritis, diabetes, and to lower blood pressure 32(2): 67-78.
(Killeen and others 1993, Liogier 1990). Guerrero
and others (2002) found that a dose of 20 mg/kg of da Silva, S.J.R. and M.L. Absy. 2000. Analyses of
ethanolic extracts of chaparro showed significant pollen found in honey samples of Apis mellifera
antihypertensive activity in rats. In Brazil, L. (Hymenoptera, Apidae) in a savanna area in
chaparro is used for treatment of inflammation and Roraima, Brazil. Acta Amazonica 30:579-588.
ulcer. Anti-inflammatory and analgesic tests with
hydroalcoholic extract of bark were successful in Foldats, E. and E. Rutkis. 1969. Sprouting of
mice (Alexandre Moreira and others 1999). Curatella americana as affected by fire. XI
Cooked leaves are used to mitigate skin eruptions, International Botanical Congress Abstracts,
for dressing wounds, and the water for purifying p. 61.
blood (Liogier 1990).
Foldats, E. and E. Rutkis. 1975. Ecological studies
References of chaparro (Curatella americana) and manteco
(Byrsonima crassifolia) in Venezuela. Journal of
Alexandre Moreira, M.S., M.R. Piuvezam, A.C. Biogeography 2:159-178.
Araujo, and G. Thomas. 1999. Studies on the
271
Furlanetto, C. and J.C. Dianese. 1999. Some Científica XV. San Pedro de Macorís, República
Pseudocercospora species and a new Prathigada Dominicana.
species from the Brazilian cerrado. Mycological
Research 103: 1,203-1,209. Liogier, H.A. 1990. Las Plantas medicinales de
Puerto Rico y del Caribe. Iberoamericana de
García Miragaya, J., S. Flores, and N. Chacón. Ediciones. San Juan PR.
1994. Soil chemical properties under individual
evergreen and deciduous trees in a protected Lundell, C.L. 1937. The vegetation of Petén.
Venezuelan savanna. Acta Carnegie Institute of Washington, Washington
Oecologica/International Journal of Ecology 15: D.C. 244 p.
477-484.
Macbride, J.F. 1956. Flora of Perú. Botanical
Gentry, A.H. 1993. A field guide to the families Series Field Museum of Natural History
and genera of woody plants of northwest South Chicago IL. 13(3a)(2): 677.
America (Colombia, Ecuador, Perú).
Conservation International Washington, DC. Mayle, F.E., R. Burbridge, and T.J. Killeen. 2000.
895 p. Millennial-scale dynamics of southern
Amazonian rain forests. Science 290: 2,291-
Gottsberger, G. and W. Morawetz. 1986. Floristic 2,294.
structural and phytogeographical analysis of the
savannas of Humaita, Amazonas, Brazil. Flora Medina, E. 1967. Intercambio gaseoso de árboles
Gena 178:41-71. de las sabanas de Trachypogon en Venezuela.
Boletín No. 111 Sociedad Venezolana de
Gottsberger, G. and W. Morawetz. 1993. Ciencias Naturales 27: 56-69.
Development and distribution of the epiphytic
flora in an Amazonian savanna in Brazil. Flora Medina, E. 1977. Introducción a la ecofisiología
188:145-151. vegetal. Monografía 16 Serie de biología.
Programa regional de desarrollo científico y
Grisebach, A.H.R. 1963. Flora of the British West tecnológico. Departamento de Asuntos
Indian Islands. Wheldon & Wesley LTD and Técnicos, Secretaría General de los Estados
Hafner Publishing Co. New York. Americanos. Washington, D. C. 102 p.
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Medina, E. and J.F. Silva. 1990. Savannas of San Jose, J.J., M.R. Fariñas, and J. Rosales. 1991.
northern South America: a steady state regulated Spatial patterns of trees and structuring factors in
by water-fire interactions on a background of a Trachypogon savanna of the Orinoco Llanos.
low nutrients availability. Journal of Biotropica 23: 114-123.
Biogeography 17: 403-413.
Sarmiento, G. 1983. The savannas of tropical
Meinzer, F., V. Seymour, and G. Goldstein. 1983. America. In: F. Bourlière, ed. Tropical savannas.
Water balance in developing leaves of 4 tropical Elsevier, Amsterdam, The Netherlands. p. 245-
savanna woody species. Oecologia 60: 237-243. 288
Mérida, T. and E. Medina. 1967. Anatomía y Sarmiento, G. and M. Monasterio. 1975. A critical
composición foliar de árboles de las sabanas de consideration of the environmental conditions
Trachypogon en Venezuela. Boletín 111 associated with the occurrence of savanna
Sociedad de Venezuela de Ciencias Naturales ecosystems in tropical America. In: F.B. Golley
27:45-55. and E. Medina, eds. Tropical ecological system.
Springer Verlag, NY. p. 223-250
Montes, R. and E. Medina. 1977. Seasonal
changes in nutrient content of leaves of savanna Sarmiento, G. and M. Monasterio. 1983. Life
trees with different ecological behavior. Geo forms and phenology. In: F. Bourlière, ed.
Eco Trop 4: 295-307. Tropical savannas. Elsevier, Amsterdam, The
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Nascimento, M.T., Villela, D.M., and L.D. de
Lacerda. 1990. Foliar growth longevity and Schnee, L. 1960. Plantas comunes de Venezuela.
herbivory in two cerrado species near Cuiaba Revista de la Facultad de Agronomía de la
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274
Cytisus scoparius (L.) Link Scotch broom
FABACEAE
John K. Francis
275
plants usually survive (Michigan State University stands reproduce prolifically. The application of
Extension 1998). Scotch broom is commonly seen herbicides has sometimes been effective (Prasad
along roads, in vacant lots and inactive 2003). A number of insect enemies of Scotch
construction sites, in old logging areas, in broom have arrived accidentally or have been
neglected pastures and orchards, and along fence- introduced into the United States. They may
rows. already be having a minor impact on the
competitive ability of the species (Syrett and
Reproduction.—Scotch broom flowers in May or others 1999). Research into biological control
June (New England Wild Flower Society 2003). continues, particularly in Australia and New
The flowers are pollinated by honey bees and Zealand (CSIRO 2003).
bumblebees (Meyer 2003). Seed production starts
when plants are 2 years old in Tasmania with Benefits.—Scotch broom, which produces a
around 300 seeds/year and increases to 12,000 to spectacular annual floral display and possesses a
30,000 seeds/shrub/year. The seeds are spread by unique form, has been planted throughout the
explosive release from the pods, which can throw temperate regions of the world as border and
the seeds 1 to 3 m (Alchemy-works 2003), as well accent plants. It is particularly useful in this regard
as by water, soil movement, and machinery because it succeeds on poor soils with little care. It
(National Haritage Trust 2003). Ants also disperse has also been used successfully to protect exposed
seeds. Seeds average 125 seeds/g. (Meyer 2003). soils and stabilize dunes. Scotch broom furnishes
Seeds can remain viable in the soil seed bank for food (mainly from the seeds) and cover for
as long as 80 years (National Haritage Trust 2003). wildlife. The leaves and twigs are generally
The species can also be propagated with cuttings unpalatable to domestic livestock and wild
(University of Connecticut 2003). mammals. The tissues contain the alkaloids
cytisin, sparteine, and isosparteine, and the
Growth and Management.—Scotch broom glycoside scoparin. However, large amounts must
grows from 30 to 46 cm/year (Michigan State be ingested before illness or death results (Everest
University Extension 1998) and usually lives from and others 2003). The stems and twigs have been
10 to 15 years (Prasad 2003). Fruits can be picked used from ancient times to make brooms and were
by hand after they are mature but before they burst used for ritual brooms called besoms. The species
open. They should be spread to dry, threshed, and was also used to make baskets, thatching, wattle
screened. These seeds are orthodox and can be fencing, and a green dye, and to flavor beer before
stored for many years. Because they are hard- the introduction of hops (Alchemy-works 2003).
coated, scarification is needed to ensure adequate Decoctions of the twigs and leaves were and are
and uniform germination. This can be done by still occasionally used to treat gout, jaundice,
nicking or sanding the seed coat or by applying edema, and other conditions (Grieve 2003).
dry heat for 1 minute at 130 °C or 15 minutes at 70
°C. Viability is normally about 80 percent. References
Because roots are delicate, the use of containerized
stock is recommended for planting rather than Abrams, L. 1944. Illustrated flora of the Pacific
bare-root stock (Meyer 2003). However, because States. Vol. 2. Stanford University Press,
the species is invasive, planting for conservation or Stanford, CA. 635 p.
ornamental purposes is usually discouraged. In
fact, the species has been declared a noxious weed Alchemy-works. 2003. Cytisus scoparius-Scotch
in five Western States where planting is prohibited broom. http://www.alchemy-works.com/
(Natural Resources Conservation Service 2003). cytisus_scoparius.html. 1 p.
Scotch broom is not a problem in cultivated land
or in land fully timbered, only in pastures, Britton, N.L. and A. Brown. 1913. Illustrated flora
roadsides, sites normally occupied by low of the northern states, Canada, and the British
vegetation, and in regenerating timberlands. possessions. Vol. 2, 2nd Ed. Scribner, New York.
Control of Scotch broom is difficult. Grubbing 735 p.
large plants and pulling seedlings, or power
cultivation, effectively kills the plants, but new CSIRO-European Laboratory. 2003. Our research:
seedlings continually appear from the soil seed Scotch broom-Cytisus scoparius Link, Fabaceae.
bank. Mowing only encourages branching near the http://www.csiro-europe.org/cytisus.html. 2 p.
ground. Stands of Scotch broom do not carry a fire
readily (Parker and others 1994), and burned
276
Everest, J.W., T.A. Powe, Jr., and J.D. Freeman. New England Wild Flower Society. 2003. Invasive
2003. Poisonous plants of the southwestern plant atlas of New England: Cytisus scoparius
United States. Alabama Cooperative Extension (Scotch broom). http://webapps.lib.uconn.edu/
System. http://www.aces.edu/dept/extcomm/ ipane/browsing.cfm?descriptionid=48. 6 p.
publications/anr/anr-975.pdf. 56 p.
Parker, B., G. Miller, L.C. Burrell. 1994. Weeds:
Grieve, M. 2003. Broom. http://www.botanical. Scotch broom. PNW 130. Pacific Northwest
com/botanical/mgmh/b/broom-70.html. 9 p. Extension, Oregon State University, Corvallis,
OR. 10 p.
International Legume Database and Information
Service. 2003. Cytisus scoparius (L.) Link. http: Prasad, R. 2003. Scotch broom, Cytisus scoparius
//www.ildis.org/LegumeWeb/6.00/taxa/6046. L. in British Columbia. Canadian Forest Service,
shtml. 4 p. Ottawa, Ontario, Canada. http://www.pfc.
forestry.ca/biodiversity/broom_e.html. 9 p.
Meyer, S.E. 2003. Cytisus scoparius (L.) Link. In:
F.T. Bonner and R.G. Nisley, eds. Woody plant Syrett, P., S.V. Fowler, E.M. Coombs, J.R.
seed manual. U.S. Department of Agriculture, Hosking, G.P. Markin, Q.E. Paynter, and A.W.
Forest Service, Washington, DC. Sheppard. 1999. The potential for biological
http://wpsm.net/Cytisus.pdf. 4 p. control of Scotch broom (Cytisus scoparius)
(Fabaceae) and related weedy species.
Michigan State University Extension. 1998. Biocontrol News and Information 20: 17N-34N.
Cytisus scoparius—Scotch broom. http://www.
msue.msu.edu/msue/imp/modop/00000485.html. University of Connecticut. 2003. University of
2 p. Connecticut plant database: Cytisus scoparius.
http://www.hort.uconn.edu/plants/c/cytsco/
National Heritage Trust. 2003. English broom cytsco3.html. 4 p.
(Cytisus scoparius). http://www.bushcare.
tas.gov.au/ToAdd/Data/weeds/English%20%20 Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
broom.pdf. 2 p. Higgins. 1987. A flora of Utah. Great Basin
Naturalist Memoirs 9. Brigham Young
Natural Resources Conservation Service. 2003. University, Provo, UT. 894 p.
Plant profile: Cytisus scoparius (L.) Link.
http://plants.usda.gov/chi_bin/plant_profile.cgi?
symbol=CYSC4. 7 p.
277
Dalbergia ecastaphyllum (L.) Taubert coin vine
FABACEAE
John K. Francis
278
are abundant near coin vine plants. However, few Holdridge, L.R. and L.J. Poveda A. 1975. Arboles
survive more than a few weeks. Branches layer de Costa Rica. Vol. 1. Centro Cientifico
whenever they come in contact with the ground. Tropical, San Jose, Costa Rica. 546 p.
Growth and Management.—Early height growth Howard R.A. 1988. Flora of the Lesser Antilles,
is moderate but steady. A small group of nursery Leeward and Windward Islands.
seedlings reached about 15 cm in 5 months. Older Dicotyledoneae, Part 1. Vol. 4. Arnold
plants grow about 2 m per year. By resprouting and Arboretum, Harvard University. Jamaica Plain,
layering, plants (clones) can survive almost MA. 673 p.
indefinitely.
Liogier H.A. 1988. Descriptive flora of Puerto
Benefits.—The extensive horizontal branches and Rico and adjacent islands, Spermatophyta. Vol.
their roots cover, bind, and stabilize coastal sand 2. Editorial de la Universidad de Puerto Rico,
dunes (Burkill 1995). The stems are brittle and not Río Piedras, PR 481 p.
suitable for wooden implements or basketry. The
crushed roots and bark contain a chemical that was Liogier, H.A. 1990. Plantas medicinales de Puerto
used by Native Americans to stupefy and catch fish Rico y del Caribe. Iberoamericana de Ediciones,
(Workman 1980). Coin vine is listed among the Inc. San Juan, Puerto Rico. 566 p.
honey plants of the Dominican Republic (Marcano
Fondeur 2001). In Senegal, the leaves are put into Marcano Fondeur, E. de J. 2001. La flora apícola
inhalations and baths to treat various debilities de la Republica Dominicana. http://marcano.
(Burkill 1995). Various extracts are used in herbal Freeservers.com/nature/estudios/apicola/dicotsp.
medicine as a diuretic, an emetic, and a vermicide. html. 11 p.
Care must be taken, because some of the tissues are
toxic (Liogier 1990). Nelson, G. 1996. The shrubs and woody vines of
Florida. Pineapple Press, Inc., Sarasota, FL.
References 391 p.
Burkill, H.M. 1995. The useful plants of West Workman, R.W. 1980. Growing native. The
Tropical Africa. Vol. 3. Royal Botanci Gardens, Sanibal-Captiva Conservation Foundation, Inc.,
Kew, United Kingdom. 857 p. Sanibal, FL. 137 p.
279
Dasiphora floribunda (Pursh) Kartesz, comb. nov. ined. shrubby cinquefoil
ROSACEAE
280
Ecology.—Shrubby cinquefoil grows in moist germination (Baskin and Baskin 2001). A cold,
soils in mountain regions from lower foothills to moist stratification period of 2 months has been
sub alpine. The elevation tends to depend generally recommended for Potentilla species,
somewhat upon latitude, but generally plants are including Potentilla fructicosa. (Phillips 1987).
found between approximately 1,800 m and 3,400 Shrubby cinquefoil can also be propagated from
m. It grows in open woods, wet mountain softwood cuttings in the spring or from suckers
meadows, calcareous bogs, limestone pavements, (Tykač 1990). Container grown plants, that are
and along streams (Cronquist and other 1997, very popular within the landscape business, can be
Gibbons and Davies 1994, Vines 1986, Welsh and planted in spring and fall. Even old plants are
others 1993). It has also been reported from reported to tolerate transplanting (Tykač 1990).
relatively xeric habitats such as mountain shrub
lands and rocky mountain slopes (McGregor and Benefits.—Shrubby cinquefoil leaves are bitter
others 1986, Stace 1997, Thilenius 1972). The tasting, and the forage and browse value varies
degree of permanent soil moisture required is depending upon the part of the world in which it
likely to depend on average precipitation, grows. It is generally on summer range, although at
temperature, and localized water collection sites or lower elevations it may be valuable for fall
drainage channels. It is a frequent member of browsing (Dayton 1931). It is likely that just as the
willow (Salix)-dominated communities, and in shrubs’ morphology varies, the levels of secondary
some parts of its range, for example New Mexico, plant products that impart the bitter taste differ
it is a member of the Krummholz communities between habitats and geographic range. Generally,
found at the lower tundra edge (Dick-Peddie it is considered inferior forage for livestock, but it
1993). Shrubby cinquefoil may be the dominant is used by cattle in Arizona and Montana (Epple
shrub in many situations and can also grow in 1995, Vines 1986). It is also often an important
fairly pure stands. For example, it may be the browse for sheep and goats especially in the
dominant shrub in aspen stands or other forest Southwestern United States and in southwestern
clearings where snow accumulation prevents tree Montana and contiguous portions of southeastern
establishment (Knight 1994, Thilenius 1972). In Idaho (Dayton 1931). In Montana it is rated 18th
addition, it frequently attains dominance in among the most important browse species.
limestone-derived soils (Thilenius 1972). Similarly, it is reported unpalatable to deer
(Thilenius 1972) but does provide browse for deer
Reproduction.—Shrubby cinquefoil generally and elk in some regions (Epple 1995, Vines 1986).
flowers in June through August, although flowers On some overgrazed range it is often grazed quite
may be found until the first frost (Epple 1995, closely, and the shrubs assume a hedge-like
McGregor and others 1986, USDA FS 1988). The appearance (USDA FS 1988). Such use indicates
achenes mature in summer and fall. The bright overstocking and a critical reduction, if not
yellow flowers are insect pollinated. The fruit is a elimination, of other palatable plant species. It
densely hairy achene. provides effective protection against soil erosion
(Epple 1995, Vines 1986). Native Americans have
Growth and Management.—Shrubby cinquefoil used various parts of the plant for an assortment of
increases in response to overgrazing and other purposes including as a medicine and in
disturbance. In the Western United States it does ceremonials (Moerman 1998). The Blackfoot
not appear to be an invasive species, but in New peoples have used the leaves to fill pillows and the
England it can be an aggressive invader of bark as tinder, while the Eskimos of Alaska and the
agricultural land and may be difficult to eradicate Arctic have used dried leaves for tea (Moerman
once established (USDA FS 1988). It has been 1998). Leaves were also used for tea in Russia. The
used as an indicator species of past disturbance name “potentilla” is derived from the Latin
(Dick-Peddie 1993). Shrubby cinquefoil is widely “potens,” which means powerful and may refer to
used as an ornamental and in wild land seeding. It the medicinal value of some species. A preparation
does best in full sun as it tends to flower poorly in made from the leaves was used by the Cheyenne
shade conditions. Seed apparently does not require for protection and was also considered a deadly
cold stratification for germination and will arrow poison, only to be administered by holy
germinate immediately after harvesting (Bonde people (Carter 1997, Moerman 1998). Shrubby
1965, Tykač 1990). The highest germination rate cinquefoil has commercial value in the landscape
was achieved at 18 oC (Bonde 1965). However, it and horticultural trade. It is a popular ornamental
has been proposed that cold stratification may shrub, and many cultivars exist that have been
reduce the temperature requirements for seed chosen for the degree of leaf pubescence
281
(hairiness), flower color, and growth form. A list of ITIS, the Integrated Taxonomic Information
some of the recognized cultivars available can be System. 1996. Dasiphora floribunda (Pursh)
found in Bailey and others (1976) and Tykač Kartesz comb. nov. ined. Taxonomic Serial No.:
(1990). 565123 and Plants, database (version 5.1.1)
http://www.itis.usda.gov/ [not paged].
References.
Kartesz, J.T. 1994. A synonymized checklist of the
Bailey, L.H., E.Z. Bailey, and the staff of the vascular flora of the United States, Canada and
Liberty Hyde Bailey Hortorium. 1976. Hortus Greenland. Vol. 1 - Checklist. 2nd Ed. Timber
Third – a concise dictionary of plants cultivated Press, Portland, OR. 622 p.
in the United States and Canada. Macmillian
Publishing Company, New York. 1,290 p. Knight, D.H. 1994. Mountains and plains–the
ecology of Wyoming landscapes. Yale
Baskin, C.C. and J.M. Baskin. 2001. Seeds, University Press, New Haven, CN. 338 p.
ecology, biogeography, and evolution of
dormancy and germination. Academic Press, McGregor, R.L., T.M. Barkley, R.E. Brooks, E.K.
New York. 666 p. Schofield. 1986. Flora of the Great Plains.
University Press of Kansas, Lawrence, KN.
Bonde, E.K. 1965. Further studies on the 1,402p.
germination of seeds of Colorado alpine plants.
University of Colorado Studies 18. Moerman, D.E. 1998. Native American ethno-
botany. Timber Press, Portland, OR. 927 p.
Carter, J.L. 1997. Trees and shrubs of New
Mexico. Johnson Books, Boulder, CO. 534 p. Phillips, J. 1987. Southwestern landscaping with
native plants. University of New Mexico Press,
Cronquist, A., N.H. Holmgren, and P.K. Albuquerque, NM. 140p.
Holmgren. 1997. Intermountain flora, vascular
plants of the Intermountain West, U.S.A. Vol. 3, Polunin, O. 1997. Flowers of Greece and the
Part A. New York Botanical Garden, Bronx, Balkans–a field guide. Oxford University Press,
NY. 446 p. New York. 567p.
Dayton, W.A. 1931. Important western browse Stace, C. 1997. New Flora of the British Isles. 2nd
plants. Miscellaneous Publication 101. U.S. edition. Cambridge University Press, New York.
Department of Agriculture, Washington DC. 1,130 p.
214 p
Thilenius, J.F. 1972. Classification of deer habitat
Dick-Peddie, W.A. 1993. New Mexico vegetation in the ponderosa pine forest of the Black Hills,
– past, present, and future. University of New South Dakota. Research Paper RM-91. USDA
Mexico Press, Albuquerque, NM. 244 p. Forest Service Rocky Mountain Forest and
Range Experiment Station, Fort Collins, CO. 28
Epple, A.O. 1995. A field guide to the plants of p.
Arizona. Falcon Press Publishing Co., Helena,
MT. 347 p. United States Department of Agriculture Forest
Service. 1988. Range Plant Handbook. Dover
Gibbons, B. and P. Davies. 1994. Field guide to Publications, Inc. New York. 838 p.
wild flowers of Britain and Northern Europe.
The Crowood Press, Marlborough, Wiltshire, Vines, R.A. 1986. Trees, shrubs, and woody vines
UK. 320 p. of the Southwest. University of Texas Press.
Austin, TX. 1,104 p.
Hitchcock, C.L. and A. Cronquist. 1973. Flora of
the Pacific Northwest – an illustrated manual. Welsh, S.L., N.D. Atwood, S. Goodrich, L.C.
University of Washington Press, Seattle, WA. Higgins, eds. 1993. A Utah flora. Second
730 p. Edition, revised. Brigham Young University,
Provo, UT. 986 p.
282
Dasylirion wheeleri S. Wats. Wheeler sotol
AGAVACEAE
Synonyms: None.
Juanita A. R. Ladyman
283
caudex that is the region at the base of the stem by Native Americans (Moerman 1998). Wheeler
(McPherson 1995, Thomas 1991). However, sotol plants are commercially used as ornamentals
frequent fires and stable soil moisture regimes tend and described as “one of the most striking accent
to exclude wheeler sotol (Burgess 1995). The plants available” (Starr 1999). The leaves were
response of sotol to fire tends to vary depending once sold for use in floral arrangements, but this
upon conditions. Kittams (1973) observed that practice has been discouraged due to conservation
mature sotol (in this case Dasylirion leiophyllum) considerations (Epple 1995).
were usually killed by fire. White (1969) randomly
selected 50 individuals after a fire in desert References
grassland in Arizona and classified them with
respect to the level of the damage (severe, Ahlstrand, G.M. 1982. Response of Chihuahuan
moderate, or light) experienced. All of the lightly Desert Mountain Shrub Vegetation to Burning.
or moderately damaged plants survived, but only 3 Journal of Range Management 35 (1): 62-65.
percent of the severely damaged plants survived.
Sprouting did not occur except as refoliation from Bowers, J.E. 1993. Shrubs and trees of the
the terminal bud (White 1969). Ahlstrand (1982) southwest deserts. Southwest Parks and
investigated the response of Dasylirion Monumens Association, Tucson, AZ. 140 p.
leiophyllum to fire at sites in New Mexico and
western Texas. He reported losses of 50 percent or Burgess, T.L. 1995. The dilemma of coexisting
more on burned sites, but surviving top-killed growth forms. In: The desert grassland. M.P.
plants regained much of their pre-burn coverage McClaran and T.R. Van Devender, eds. The
after three growing seasons. This was similar to University of Arizona Press, Tucson, AZ. p. 31-
the observations after a semidesert grassland fire 67.
in southern Arizona where 47 percent of the
Wheeler sotol plants burned to death (Thomas and Carter, J.L. 1997. Trees and shrubs of New
Goodson 1992). Of the 71 Wheeler sotol plants Mexico. Johnson Books, Boulder, CO. 533 p.
that survived, all had apical regrowth, but despite
the fact that sotol may occasionally produce new Dayton, W. A. 1931. Important western browse
shoots from below ground, no offsets or seedlings plants. Miscellaneous Publication 101. U.S.
were found 11 months after the fire (Thomas and Department of Agriculture, Washington, DC.
Goodson 1992). In this study no individuals 214 p.
evaded the fire in refugia.
Dick-Peddie, W.A. 1993. New Mexico vegetation:
Benefits.—The head-like trunks and leaf bases of past, present, and future. University of New
Wheeler sotol are edible and American Indians Mexico press, Albuquerque NM. 244 p.
roasted and ate the young flower stalks (Moerman
1998, Powell 1998). The basal trunks contain Epple, A.O. 1995. A field guide to the plants of
starch and are also roasted, fermented and distilled Arizona. Falcon Press Publishing Co., Helena,
to make alcohol (Powell 1998). Currently an Montana. 347 p.
alcoholic beverage, generally known as sotol, is
made in parts of Mexico (Bowers 1993). Bighorn Gledhill, D. 1992. The names of plants. 2nd ed.
sheep and javelinas browse sotols (Epple 1995, Cambridge University Press, Cambridge,
Kearney and others 1960, Starr 1999). In times of England. 202 p.
stress Wheeler sotol has been used as feed for
livestock in a variety of ways. The trunks and leaf Ivey, R.D. 1995. Flowering plants of New Mexico.
bases have been used during drought, and the 3rd ed. Rio Rancho Printing, Albuquerque, NM.
plants have also been machine-cut and shredded 504 p.
for emergency feed (Dayton 1931, Powell 1998).
In addition, ranchers have burned off the leaves Kearney, T.H., R.H. Peebles, J.T. Howell, and E.
and split the heads (trunks) for livestock (Vines McClintock. 1960. Arizona flora. 2d ed.
1960). The leaves have been used to make mats, University of California Press. Berkeley, CA.
baskets, thatch, and paper (Moerman 1998, Powell 1,085 p.
1998). The long flowering stalks have been used
extensively within its range for corral, porch and Kittams, W.H. 1972. Effect of fire on vegetation of
house roofs, and other structures (Powell 1998). the Chihuahuan Desert region. Proc. Tall
They have also been used for ceremonial purposes Timbers Fire Ecology Conference 12: 427-444
284
McPherson, G.R. 1995. The role of fire in desert Sciences Laboratory. Fire Effects Information
grasslands. In: The desert grassland. M.P. System. http://www.fs.fed.us/database/feis/. 8
McClaran and T.R. Van Devender, eds. The p.
University of Arizona Press, Tucson, AZ. p. 130-
151. Thomas, P.A. 1991. Response of succulents to
fire: A review. International Journal of Wildland
Moerman, D.E. 1998. Native American ethno- Fire 1(1): 11-22.
botany. Timber Press, Portland, OR. 927 p.
Thomas, P.A. and P. Goodson. 1992. Conservation
Powell, A. M. 1998. Trees & shrubs of Trans- of succulents in desert grasslands managed by
Pecos and adjacent areas. University of Texas fire. Biological Conservation. 60(2): 91-100.
University Press, Austin, TX . 498 p.
Vines, R.A. 1960. Trees, shrubs, and woody vines
Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L. of the Southwest. Sixth printing 1986.
Cox, J.O. Klemmedson, J.J. Norris, and L.C. University of Texas Press. Austin, TX.
Fierro. 1992. Desert grassland. In: R.T. 1,104 p.
Coupland, ed. Natural grasslands: Introduction
and Western Hemisphere. Elsevier, New York. Warnock, B.H. 1974. Wildflowers of the
p. 337-362. Guadalupe Mountains and the Sand Dune
Country, Texas. Sul Ross State University,
Starr, G. 1999. Dasylirion – The shaggy lilies. Alpine, TX. 176 p.
Desert Plants 15(1): 29-31.
White, L.D. 1969. Effects of a wildfire on several
Tesky, J.L. 1993. Dasylirion wheeleri. In: U.S. desert grassland shrub species. Journal of Range
Department of Agriculture, Forest Service, Management 22: 284-285.
Rocky Mountain Research Station, Fire
285
Datura inoxia P. Mill. devil’s trumpet
SOLANACEAE
John K. Francis
The ovoid capsule is nodding, about 3 cm in
diameter and covered with stout, soft prickles 2 to
4 mm long. The capsules remain on the plant for a
long period. The yellowish-brown seeds are flat,
kidney-shaped, about 5 mm long, and have a small
fleshy aril. Datura species normally have 12 pairs
of chromosomes (Bonde 2001, Burkill 2000,
Howard 1989, Liogier 1995, Stevens and others
2001).
286
fragrance (Annie’s Annuals 2002), which perhaps D%20-%20G/Datura_metel_’Belle_Blanche’.
helps attract night-flying moths. The seeds are htm. 1 p.
distributed by ants and some species of birds that
are resistant to the chemicals they contain (Bonde Bonde, K. 2001. The genus Datura: From research
2001). Seeds germinate in less than 2 weeks at soil subject to powerful hallucinogen.
temperatures of 27 °C (Hardy Plants 2002). Devil’s http://leda.lycaeum.org/Documents/The_Genus_
trumpet is normally propagated with seeds. Datura:_From_Research_Subject_to_Power.
9 p.
Growth and Management.—In dry environments,
devil’s trumpet reaches a maximum height of about Burkill, H.M. 2000. The useful plants of West
0.5 m in one growing season. In moist areas, it can Tropical Africa. Vol. 5. 2nd ed. Royal Botanical
reach two to four times this height although it is Gardens, Kew, UK. 686 p.
not clear whether it does so in a single season. New
stands can be established by cultivation and Erowid. 2002. Erowid experience vaults.
planting. Although control information has not http://www.erowid.org/experiences/subs/ex_Dat
been published, weedy plants can probably be ura.shtml#retrospective_/_Summary. [not
reduced or eliminated without difficulty by paged].
cultivation or spraying with broadleaf herbicides.
Gupta, M., A. Bagchi, and A.B. Ray. 1991.
Benefits and Detriments.—Devil’s trumpet is Additional withanolides of Datura metel.
grown in all but the coldest climates as a flowering Journal of Natural Products. 54(2): 599-602.
ornamental. Beautiful white, purple, and yellow
varieties with large, single and double blossoms are Hardy Plants. 2002. Specialty perennials.
available. From ancient times continuing to the http://hardyplants.com/D.htm. [not paged].
present, the taking of Datura spp. tissues,
particularly the seeds, was used in shamanistic Howard, R. A. 1989. Flora of the Lesser Antilles,
rituals as a path to enlightenment (Schultes and Leeward and Windward Islands. Vol. 6. Arnold
Hofmann 1992). Today, people frequently Arboretum, Harvard University, Jamaica Plain,
experiment with it for the hallucinogenic effect, but MA. 658 p.
the results are so unpleasant (dark visions,
disorientation, amnesia, blurred vision, dry mouth, Janthangvith, J., P. Chumsri, K. Kraisintu, and A.
and incontinence) that they seldom recommend the Pongpan. 2000. Comparison of tropane alkaloid
experience (Erowid 2002). Overdoses can result in production by Datura innoxia and Datura metel
death. The plant has been used to treat impotence, varieties of white, soil, hydroponic, in vitro plant
asthma, diarrhea, as an analgesic, to control fever, culture and tranformed root culture. Mahidol
kill parasites, and as a drug for criminal purposes University annual Research Abstracts.
(Bonde 2001, Parrotta 2001). Devil’s trumpet http://www.hahidol.ac.th/abstracts/annual1999/0
contains a host of phytoactive chemicals including 355.htm. 2 p.
atropine, hyoscyamine, hyoscine, scopolamine,
norscopolamine, meteloidine, hydroxy-6- Liogier, H.A. 1995. Descriptive flora of Puerto
hyoscyamine, tiglic esters of dihydroxytropine, and Rico and adjacent islands. Vol. 4. Editorial de la
a number of withanolides (Gupta and others 1991, Universidad de Puerto Rico, San Juan, PR.
Janthangvith and others 2000). Alkaloid 617 p.
concentration in tissues is around 0.5 percent
(Niber and others 1992) but varies according to Missouri Botanic Garden. 2003. Missouri Botanic
environmental circumstances (Burkill 2000). Garden. Missouri Botanic Garden, St. Louis,
Devil’s trumpet is a minor weed in tropical and MO. http://www.mobot.org/. [not paged].
some subtropical countries. It causes erratic
behavior and even death of livestock that have Niber, B.T., J. Helenius, and A.L. Varis. 1992.
eaten it, but it is seldom a problem for pastured Toxicity of plant extracts to three storage beetles
animals because they carefully avoid consuming it. (Coleoptera). Journal of Applied Entomology
113(2): 202-208.
References
Parrotta, J.A. 2001. Healing plants of India. CABI
Annie’s Annuals. 2002. Datura metel, ‘Belle Publishing, Wallingford, UK and New York.
Blanche.’ http://www.anniesannuals.com/signs/ 917 p.
287
Schultes, R.E. and A. Hofmann. 1992. Plants of Stevens, R.D., C. Ulloa-U., A. Pool, and O.M.
the gods. Healing Plants Press, Rochester, NY. Montiel, eds. 2001. Flora de Nicaragua.
192 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanic Garden, St. Louis, MO. pp.
911-2,666.
288
Desmanthus virgatus (L.) Willd. wild-tantan
FABACEAE
John K. Francis
short petiole, less than seven pinnae, 10 to 25 pairs
of leaflets per pinnae that are linear-oblong and 4
to 9 mm long. The flowers occur in axillary heads
with peduncles 2 to 7.5 cm long. The petals are
white or creamy yellow. The pod, a legume, is
linear, 50 to 90 mm long, 3 to 4 mm broad,
reddish-brown, and contains 20 to 30 seeds
(Howard 1988, Liogier 1988, Skerman and others
1988). There are 2n = 28 chromosomes (Long and
Lakela 1976).
289
Reproduction.—Wild-tantan blooms at the end of Government, Brisbane, Australia. http://dpi.qld.
the wet season, and fruits mature about 1 month gov.au/pastures/4490.html. 2 p.
later. Seeds collected in Puerto Rico weighed an
average of 0.0041 + 0.0005 g/seed or 244,000 Forlin, S.M., H.Y. Rey, and L.A. Mroginski. 2000.
seeds/kg. Placed to germinate on moist filter paper, Cultivo de tejidos de Desmanthus virgatus:
only 28 percent germinated in 1 year. The atención de plantas a partir de hojas.
remaining seeds were scarified (by nicking) and Universidad Nacional del Nordeste, Corrientes,
100 percent germinated starting in 2 days (author’s Argentina. Comunicaciones Científicas y
observation). In nature, scarification occurs by Tecnológicas 2000. 3 p.
weathering of the seed coat in the soil or while
passing through an herbivore. Artificial Fox, J.E.D., A.I. Doronila, D.R. Barrett, and I.K.
scarification by treatment with concentrated Surata. 1996. Desmanthus virgatus (L.) Willd.
sulfuric acid for 8 minutes is recommended An efficient intermediate host for the parasitic
(Skerman and others 1988). The seeds are species Santalum album L. in Timor, Indonesia.
dispersed by ruminant animals. Many seedlings Journal of Sustainable Forestry 3(4): 13-23.
spring up under favorable conditions in natural
stand, but few survive. The species can be Howard R.A. 1988. Flora of the Lesser Antilles,
propagated by rooting hormone-treated leaves Leeward and Windward Islands.
(Forlin and others 2000). Dicotyledoneae, Part 1. Vol. 4. Arnold
Arboretum, Harvard University, Jamaica Plain,
Growth and Management.—Wild-tantan MA. 673 p.
normally grows from 0.5 to 1.5 m/year in height.
After establishment, the root crown gradually Institute of Systematic Botany. 2002. Atlas of
increases in size until it reaches 15 cm across in 3 Florida plants: Desmanthus virgatus. University
years. Plantations are established by sowing 2 kg of of Florida, Gainesville, FL. http://www.
scarified seed per hectare into well-prepared plantatlas.usf.edu/maps.asp?plantID=1474. 1 p.
seedbeds at a depth of 1 to 1.5 cm. Plantations can
be cut for feed 4 times/year and yield 35 Labios, R.V., J.G. Montesur, and R.O. Retales.
tons/ha/year of dry matter. There is little mortality 1994. Alley cropping in sloping upland rice
up to at least the fourth year (Skerman and others areas of Cavite, Philippines. Philippine Journal
1988). of Crop Science 19(1): 33-37.
Benefits.—Wild-tantan is planted throughout the Liogier H.A. 1988. Descriptive flora of Puerto
tropics as a forage and hay crop and is appreciated Rico and adjacent islands, Spermatophyta. Vol.
as a forage in range land. The forage is nontoxic 2. Río Piedras, PR: Editorial de la Universidad
and palatable to cattle and other ruminants. It is a de Puerto Rico. 481 p.
favorite of white-tailed deer in southern Texas
(Soltero-Gardea and others 1994). Crude protein Long, R.W. and O. Lakela. 1976. A flora of
content of whole plants from plantations ranges Tropical Florida. Banyon Books, Miami, FL.
from 10.5 to 15.5 percent, the leaves having 22.4 962 p.
percent and the stems 7.1 percent (Skerman and
others 1988). Wild-tantan is planted as a hedge- Neal, M.C. 1965. In gardens of Hawaii. Special
row plant in the alley cropping system on steep Publication 50. Bernice P. Bishop Museum
land in the Philippines (Labios and others 1994). It Press, Honolulu, HI. 924 p.
has been shown to be an efficient nursery-stage
host for sandalwood (Santalum album L.) (Fox and Skerman, P.J., D.G. Cameron, and F. Riveros.
others 1996). The species can become a serious 1988. Tropical forage legumes. 2nd. Ed. Food
weed in sugar cane fields (Neal 1965). and Agriculture Organization of the United
Nations, Rome. 692 p.
References
Soltero-Gardea, S., I.M. Ortega, and F.C. Bryant.
Department of Primary Industries. 2002. Legumes 1994. Nutrient content of important deer forage
for the tropics and subtropics: Desmanthus plants in the Texas costal bend. Texas Journal of
(Desmanthus virgatus). Queensland Science 46(2): 133-142.
290
Distictis lactiflora (Vahl) DC. pega palo
BIGNONIACEAE
John K. Francis
291
observation). Vines layer (root) when they come in References
contact with the ground.
Acevedo-Rodríguez, P. 1985. Los bejucos de
Growth and Management.—Pega palo grows at Puerto Rico. Vol. 1. Gen. Tech. Rep. SO-58.
least a meter per year from sprouts. Individual U.S. Department of Agriculture, Forest Service,
vines live several years, and strong plants develop Southern Forest Experiment Station, New
new stems periodically. Grazing of dry and moist Orleans, LA. 331 p.
forest areas probably promotes the species.
Liogier, H.A. 1995. Descriptive flora of Puerto
Benefits.—Pega palo contributes to the Rico and adjacent islands. Vol. 4. Editorial de la
biodiversity of moist and dry forests and helps Universidad de Puerto Rico, San Juan, PR.
protect the soil. Its foliage and white flowers add to 617 p.
the beauty of the forest. The species has been used
as an ornamental to a limited extent.
292
Dodonaea viscosa (L.) Jacq. ‘a‘ali‘i
SAPINDACEAE
293
colored to dark reddish purple, exceedingly phytoplasmas in Hawaii. Plant Disease. 79(11):
variable, turgid to inflated bladder or compressed. 1,094-1,097.
Seeds are compressed-ovoid, about 3 mm long.
The two to four-winged, papery capsules are either Burrows, C.J. 1995. Germination behavior of the
dispersed by wind or adhere to plumage of birds seeds of six New Zealand woody plant species.
(Burrows 1995). The seed coats of most of the New Zealand Journal of Botany 33: 365-377.
‘a’ali’i seeds are impermeable to water and there
also appears to be a micropylar plug present. The Cribb, A.B. and Cribb, J.W. 1981. Wild Medicine
hard seed coat and the micropylar plug may play a in Australia. Collins, Sydney. 228 p.
role in delayed germination. Some evidence
suggests that higher temperatures, which may Jain, R.K. 1998. Fuelwood characteristics of
enhance the breakdown of the plug, as well as medium tree and shrub species of India. In:
scarification of the seed coat, may be necessary for Bioresource technology. Volume 68, Issue 3.
germination (Burrows 1995). ‘A‘ali‘i is able to Elsevier Science Limited, Essex, UK. p. 305-
reproduce through sprouting following 308.
disturbance.
Lamb, S.H. 1981. Native trees and shrubs of the
Growth and management.—‘A‘ali‘i is popular as Hawaiian Islands. The Sunstone press, Santa
an ornamental in xeriscape gardens. Seedlings Fe, NM. 160 p.
should be planted from containers after last frost in
any kind of soil. Water infrequently once Little, E.L. and R.G. Skolmen. 1989. Common
established and prune if a more formal appearance forest trees of Hawaii (native and introduced).
is desired. Plant emits strong odor during pollen Agriculture Handbook. 679. U.S. Department of
season. Plant is frost damaged at approximately – Agriculture, Washington, DC. 321 p.
6.6° C (Anonymous 1999).
Pengelly, A. Dodonaea viscosa. www.nhaa.org.au/
Benefits.—‘A‘ali‘i has enormous potential as a forum/text/dodvis2.html.
landscape species. ‘A’ali’i also possesses many
medicinal properties and has been used by native Rojas, A., S. Cruz, H. Ponce-Monter, and R. Mata.
peoples from all regions where it is found. Its 1996. Smooth muscle relaxing compounds from
leaves can be used to combat rheumatism, skin Dodonaea viscosa. Planta Medica. 62:154-159.
infections, fevers, swellings, aches, and
“gastrointestinal disorders” including diarrhea, and Sherry, K., J.M. Castillo, and R.B Shaw. 1999.
can be used as a antispasmodic agent (Rojas and Effects of wildfire on vegetation and rare plants
others 1996). The Australian aborigines used the in arid montane shrublands. Pohakaloa training
leaves and roots as a painkiller to soothe area, Hawaii. 1999 Hawaii Conservation
toothaches and headaches (Crib and Crib 1981). Conference, Honolulu, HI.
The flowers (or hops) of ‘a’ali’i are used as a
“home-brew” substitute to bestow a bitter flavor, Wagner, H., C. Ludwig, L. Grotjahn, and M.S.Y.
and also used as a tonic. A red dye is extracted Khan. 1987. Phytochemistry 26(3): 697-701.
from the fruit, and the capsules are also commonly
used in leis (Little and Skolmen 1989). In India the Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
seeds are used as fish poison (Wagner and others 1990. Manual of the Flowering Plants of
1987). The wood from ‘a’ali’i is extremely hard Hawaii. University of Hawaii Press, Honolulu,
and has been used for tools, spears and weapons HI. 1,854 p.
(Lamb 1981). In India ‘a’ali’i is an important
source of fuelwood (Jain 1998). Williams, P.A. and R.P. Buxton. 1989. Response
to reduced irradiance of 15 species of native and
References adventive shrub and tree seedlings from eastern
Canterbury. New Zealand Journal of Ecology.
Anonymous. 1999. Dodonaea viscosa: Hopbush. 12: 95-101.
www.plantadviser.com/plants/dodovisc.shtml
West, J.G. and I.R. Noble. 1984. Analyses of
Borth, W.B., J.S. Hu, B.C. Kirkpatrick, D.E. digitized leaf images of the Dodonaea viscosa
Gardner, and T.L. German. 1995. Occurrence of complex in Australia. Taxon 33(4):595-613.
294
Dryas octopetala L. white mountain avens
ROSACEAE
295
grows is associated with latitude. Plants are found when sown outdoors in the autumn while seeds did
at 3,500 m to 3,965 m in Utah whereas they may geminate the following spring, suggesting that
occur at 100 m or less in northern parts of its cold stratification lowers the temperature
range, such as Greenland (Borge Wills 1996, requirement for germination (Elkington 1971). The
Welsh and others 1993). It typically grows above stems root easily and contribute to the rapid spread
the timberline in sites with low snow cover on of plants. As part of studies on the consequences
calcareous or basic soils (Dorn 2001, Dorn 1984, of global climate change, experiments have been
Komárková 1979, Ellenberg 1988, Stace 1997, made on the response of white mountain avens to
Weber and Wittman 2001). It is not an indicator of elevated temperatures in the Colorado Rocky
calcareous sites per se as it is common on granitic Mountains (Welker and others 2001). The results
substrates in the Colorado Rocky Mountains suggested that warmer, wetter conditions would
(Weber 1960). However, on this substrate calcium initially lead to greater productivity (increased leaf
is likely available to the plants as a leachate mass and higher numbers of seed per ramet)
(Weber and Wittman 2001). Mountain avens although decomposition may be retarded, leading
grows in dry meadows, rocky tundra, fellfields, to a lower nutrient availability in subsequent years
and in relatively moist sites in fen lands. In Russia, (Welker and others 2001). Under longer term
it has been reported as a co-dominant in “frost conditions, a switch in the source-sink activity of
mound bogs” that are essentially permanently the community occurred. Under current conditions
frozen ridges, or mounds, that alternate with wet the mountain avens community is a carbon sink of
hollows (Botch and Masing 1983). In contrast, about 8 g CO2-C m-2. However, after 4 years of
mountain avens and lichens are of primary experimental warming, the community switched to
importance on the dry upland meadows in the a carbon source of almost 10 g CO2-C m-2 per
Alaskan Arctic and on the windswept fellfield summer due primarily to an increase in the rate of
slopes of the Rocky Mountains of Colorado U.S.A. respiration without a concomitant increase in
(Komárková 1979, Langenheim 1962). These photosynthesis (Welker and others 2001). This
shrubs have an trellis-like growth habit clinging switch in source sink activity has been reported for
closely to rocks and stones and anchoring its roots Dryas and other communities in the Arctic
in cracks or between stones. The radiation is so (Welker and others 2001, Oechel and others 1993).
intense at high elevations that the temperature can This finding may be significant when considering
rise to more than 65 oC where the ground is dark the repercussions of increasing industrial
colored. In these locations the prostrate seedlings “greenhouse gas” emissions
and young plants can be heat damaged (Ellenberg
1988). A mutualistic association exists between Benefits.—White mountain avens is a pioneer
mountain avens and ectomycorrhizal fungi species and is important in stabilizing the thin soils
(Monson and others 2001). The most dominant on mountain slopes (Zwinger and Willard 1996,
fungus is Cenococcum geophilum, an ascomycete. Ellenberg 1988). It spreads relatively rapidly and
is popular in landscape rock gardens (Williams
Reproduction.—White mountain avens reproduce and others 1986). Several cultivated varieties,
by seed. Flowering is from June into August. The including hybrids, are available (Kershaw and
flowers always face the sun on calm, sunny days, others 1998). One hybrid, xSuendermannii
and their parabolic shape directs the sun’s heat on Kellerer, between Dryas octopetala and D.
to the stigma, which can then become up to 3.5 oC drummondii is morphologically similar to D.
warmer than the surrounding air temperature octopetala, but the flowers are yellow in bud
(Kershaw and others 1998). The warmer becoming white and nodding upon opening
temperatures can attract insects and also speed (Bailey and Bailey 1976). White mountain avens is
pollen germination. Hybridization occurs between an important food source for ptarmigian (Nelson
sympatric Dryas species (Komarov 1941). Seed is, 1992) and pikas (Martin and others 1951).
at least in part, distributed by wind.
References.
Growth and Management.—White mountain
avens seeds have physiological dormancy but do Bailey, L.H., E.Z. Bailey, and the staff of the
not appear to require stratification for germination Liberty Hyde Bailey Hortorium. 1976. Hortus
(Baskin and Baskin 2001). Forty-eight percent of Third. Macmillian Publishing Company, New
freshly matured seeds germinated after 4 days in York, NY. 1,290 p.
the light at 25 oC (Baskin and Baskin 2001).
However, in England seeds failed to germinate
296
Baskin, C.C. and J.M. Baskin. 2001. Seeds, Komárková, V. 1979. Alpine vegetation of the
ecology, biogeography, and evolution of Indian Peaks area, Front Range, Colorado Rocky
dormancy and germination. Academic Press, Mountains. J. Cramer, Vaduz, Germany.
New York, NY. 666 p.
Komarov, V.L. 1941. Flora of the U.S.S.R.
Borge Wills, L. 1996. Sanderling review. Marine Volume 10. Izdatel’stvo Akademii Nauk SSSR.
and Coastal Species Information System, Translated by N. Landau 1970. The Smithsonian
Conservation Management Inst., Roanoke, VA. Institution, Washington, D.C. p. 272-273.
http://fwie. fw.vt.edu [not paged].
Langenheim, J.H. 1962. Vegetation and
Botch, M. and V. Masing. 1983. Mire ecosystems environmental patterns in the Crested Butte area.
in the U.S.S.R. In: A.J.P. Gore, ed. Ecosystems Ecological Monographs 32(3): 249-285.
of the World, Vol. 4B. Elsevier Company, New
York. p. 95-152. Martin, A.C., H.S. Zim, and A.L. Nelson. 1951.
American wildlife and plants. McGraw-Hill
Britton, N.L. 1901. Manual of the flora of the Book Company, New York, NY. 500 p.
northern states and Canada. Henry Holt and
Company, New York, NY. 1,080 p. Monson, R.K., R. Mullen, and W.D. Bowman.
Plant nutrient relations. 2001. In: W.D. Bowman
and TR. Seastedt, eds. Structure and function of
Carter, J.L. 1988. Trees and shrubs of Colorado.
an alpine ecosystem. Oxford University Press,
Johnson Books, Boulder, CO. 165 p.
New York, NY. 337 p.
Dorn, R. D. 1984. Vascular plants of Montana.
Nelson, R.A. 1992. Handbook of Rocky Mountain
Mountain West Publishing Cheyenne, WY. 412
plants. Robert Rinehart Publishers, Niwot, CO.
p.
444 p.
Dorn, R. D. 2001. Vascular plants of Wyoming.
Oechel, W.C., S.J. Hastings, G.L. Vourlitis, M.A.
Mountain West Publishing Cheyenne, WY. 412
Jenkins, G. Riechers, and N. Grulke. 1993.
p.
Recent changes of arctic tundra ecosystems from
a carbon sink to a source. Nature 361: 520-523.
Elkington, T.T. 1971. Biological flora of the
British Isles. Dryas octopetala. Journal of
Ecology 59: 887-905. Praprotnik, N. 1997. Alpski botanicni vrt Juliana v
Trenti. Prirodoslovni muzej Slovenije,
Ellenberg, H. 1988. Vegetation ecology of central Ljubljana, Slovenia. 126 p.
Europe. 4th edition. Cambridge University Press,
New York, NY. 731 p. Stace, C. 1997. New Flora of the British Isles. 2nd
edition. Cambridge University Press, New York,
Gledhill, D. 1992. The names of plants. NY. 1,130 p.
Cambridge University Press, Cambridge,
England. 202 p. Weber, W.A. 1960. Some features of the
distribution of Arctic relicts at their Austral
Hitchcock, C.L. and A. Cronquist. 2001. Flora of limits. Twelfth International Botanical Congress.
the Pacific Northwest. University of Washington Recent Advances in Botany. p. 912-914
Press, Seattle, WA. 730 p.
Weber, W.A., and R.C. Wittman 2001. Colorado
Kartesz, J.T. 1994. A synonymized checklist of the flora, western slope. University Press of
vascular flora of the United States, Canada and Colorado, Boulder, CO. 488 p.
Greenland. Vol. 1 - Checklist. Timber Press,
Portland, OR. 622 p. Welker, J.M., W.D. Bowman, and T.R. Seastedt.
2001. Environmental change and future
Kershaw, L., A. MacKinnon, and J. Pojar. 1998. directions in alpine research. In: W.D. Bowman
Plants of the Rocky Mountains. Lone Pine and T.R. Seastedt, eds. Structure and function of
Publishing, Edmonton, Canada. 383 p. an alpine ecosystem. Niwot Range, Colorado.
Oxford University Press, New York. 337 p.
297
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Williams, J., R. Radebaugh, D. Hall, G. Keladis, P.
Higgins, eds. 1993. A Utah flora. Second Keladis, and P. A. Pachuta. 1986. Rocky
Edition, Brigham Young University, Provo, UT. Mountain Alpines. Timber Press, Portland, OR.
986 p. 333 p.
298
Duranta erecta L. golden dewdrop
VERBENACEAE
John K. Francis
299
throughout the year (author’s observation), and in Castro, O., M. Barrios, M. Chinchilla, and O.
Florida, both flowers and fruit are present on the Guerrero. 1996. Chemical and biological
shrubs at the same time (Nelson 1996). A evaluation of the effects of plant extracts against
collection of golden dewdrop fruits from Puerto Plasmodium berghei. Revista de Biologia
Rico averaged 0.515 + .009 g/fruit. Air-dried seeds Tropical 44(2A): 361-367.
separated from them averaged 0.0346 + 0.0005
g/seed or 28,900 seeds/kg. Set to germinate on Floridata. 1999. Duranta erecta. http://
moist filter paper without any pretreatment, 29 floridata.com/ref.d.dra_ere.cfm. 3 p.
percent of the seeds germinated beginning 128
days after sowing (Francis and Rodíguez 1993). Francis, J.K. and A. Rodríguez. 1993. Seeds of
Birds disseminate the seeds (Watkins and Sheehan Puerto Rican trees and shrubs: second
1975). However, seedlings are seldom common. installment. Research Note SO-374. U.S.
After establishment, plants thicken and spread Department of Agriculture, Forest Service,
laterally a few decimeters vegetatively. Golden Southern Forest Experiment Station, New
dewdrop are produced commercially from seeds, Orleans, LA. 5 p.
cuttings, and by layering (Floridata 1999).
Hardin, J.W. and J.M. Arena. 1969. Human
Growth and Management.—Golden dewdrop poisoning from native and cultivated plants.
plants live at least 15 years. The species has a Duke University Press, Durham, NC. 167 p.
moderate growth rate, usually about 0.5 m/year for
the first few years. Ornamental plants need regular Howard, R.A. 1989. Flora of the Lesser Antilles.
thinning and pruning to keep them under control Vol. 6. Arnold Arboretum, Harvard University,
and to remove dead branches (Floridata 1999). Jamaica Plain, MA. 658 p.
Although they withstand poor soil and drought
well, plants grow best if watered and fertilized Liogier, H.A. 1995. Descriptive flora of Puerto
(Woman’s Club of Havana 1958). Rico and adjacent islands. Vol. 4. Editorial de la
Universidad de Puerto Rico, San Juan, PR.
Benefits and Detriments.—Golden dewdrop 617 p.
forms a part of the coastal scrub community and
contributes to soil and ecosystem stability. It is a Little, E.L., Jr., R.O. Woodbury, and F.H.
popular ornamental used for accent plants and Wadsworth. 1974. Trees of Puerto Rico and the
hedges in tropical and subtropical parts of the Virgin Islands. Vol. 2. Agriculture Handbook
world because of its profuse displays of flowers 449. U.S. Department of Agriculture,
and fruits (Floridata 1999). Golden dewdrop Washington, DC. 1,024 p.
flowers attract butterflies and hummingbirds
(Floridata 1999). It is sometimes grown in Nelson, G. 1996. The shrubs and woody vines of
greenhouses in areas too cold for natural plants. Florida. Pineapple Press, Inc., Sarasota, FL.
The wood is light brown and hard and useful for 391 p.
stakes and fuel. Golden dewdrop is a poisonous
plant and has caused deaths of children. Saponins Pacific Island Ecosystems at Risk. 2002. Duranta
in the fruits and foliage cause gastroenteric erecta L., Verbenaceae. http://www.hear.
irritation, drowsiness, fever, nausea, vomiting, and org/pier3/duere.htm. 2 p.
convulsions. Dermatitis sometimes occurs from
handling the plants (Hardin and Arena 1969, Watkins, J.V. and T.J. Sheehan. 1975. Florida
Westbrooks and Preacher 1986). However, birds landscaping plants, native and exotic. The
feed on the fruits without difficulty (Nelson 1996). University Presses of Florida, Gainesville, FL.
Ethyl acetate and aqueous extracts of leaves 420 p.
showed significant antimalarial activity when
administered to mice (Castro and others 1996). In Westbrooks, R.G. and J.W. Preacher. 1986.
small quantities, fruits are used to treat intestinal Poisonous plants of Eastern North America.
worms (Whistler 2000). University of South Carolina Press, Columbia,
SC. 226 p.
References
Whistler, W.A. 2000. Tropical ornamentals, a
Bruggeman, L. 1964. Tropical plants and their guide. Timber Press, Inc., Portland, OR. 542 p.
cultivation. Thames and Hudson, London. 228 p.
300
Woman’s Club of Havana. 1958. Flowering plants
from Cuban gardens. Criterion Books, New
York. 365 p.
301
Encelia farinosa Gray ex Torr. brittlebush
ASTERACEAE
Synonyms: none
302
plants in more mesic gardens to grow less- capacity to survive drought conditions (Ehleringer
pubescent leaves than sibling plants in desert 1993). In general, however, brittlebush is drought
gardens, a response which allows plants to take adapted and responds quickly to water addition
advantage of higher water availability and through rapid CO2 uptake, leaf production, and
increased photosynthetic ability through leaf-hair stem growth (Nobel and others 1998).
reduction (Ehleringer and Clark 1988). The
inverse relationship for leaf pubescence and Reproduction.—Brittlebush flowers from March
photosynthetic activity may involve pleiotropy or through May (Munz and Keck 1968) and
linkage to other genetically determined individuals must be cross-pollinated in order to
morphological differences between populations or produce seed (Ehleringer and Clark 1988).
from environments with different moisture Flowers are visited by various insects including
regimes (Housman and others 2002). butterflies, moths, flies, bees, wasps, and beetles
(Kyhos 1971, Moldenke 1976). In one study, a
There is also clinal variation in production of beetle (Malachiidae) was found to be 10 times
chemical compounds that provide defense against more common than all other insect species
herbivores. From north to south in Baja California combined (Kyhos 1971). These potential
and east to west from the Sonoran desert to coastal pollinators do not discriminate between plants with
regions of California, plants produce progressively different disk-flower color (Kyhos 1971). Achenes
less of a sequiterpene and more of a chromene are collected from May to July (Mirov and Kraebel
toxin, which may influence local resistance to 1939) with about 770,000 bulk seeds/kg (personal
herbivores (Wisdom 1985, Kunze and others communication with S&S seeds, Carpenteria, CA).
1995). Seasonal production of these compounds Seed viability varies among years, with as few as
may also influence herbivores (Wisdom and 35 percent of seeds viable (personal
Rodriquez 1983). There are higher concentrations communication with M. Wall, Rancho Santa Ana
of these chemicals and nitrogen in young tissues. Botanical Garden, Claremont, CA). Seed
In addition, populations differ in the relative production is influenced by water stress
amount of different compounds. The specialist heightened by competition. In a desert study in
beetle Trirhabda geminata Horn experienced which nearby neighbors were removed, shrubs
lowered larval growth rates when fed higher levels experienced lower water stress, had higher
of the secondary compounds. survival, grew to nearly twice the mass, and
Ecology.—In sage scrub, brittlebush occurs on produced 53 percent more flower heads per twig
flats and slopes, primarily on weathered sandstone, and 220 percent more achenes per head than
granite, diorite, and alluvial deposits; in the desert shrubs with brittlebush neighbors within 2 m
it occurs on rocky slopes, flats, and in washes (Ehleringer 1984).
(Munz and Keck 1968, Hickman 1993). Studies on Establishment and Growth.—Brittlebush
the adaptive significance of variation in leaf form becomes established rapidly following
and pubescence both within and among disturbance. Seedlings emerge and become
populations is reviewed by Housman and others established in open areas in the winter rainy
(2002) and by Sandquist and Ehleringer (1997, season. Plants become dormant and drop many
1998). Plants react to seasonal increase in water leaves during the dry season and then sprout new
stress at the end of the rainy season by replacing leaves with the onset of winter rains. Plants can
the larger, less hairy leaves produced earlier in the reach maturity within 2 years and often live for 10
growing season with more pubescent leaves that to 15 years. In sage scrub vegetation, resprouting
are smaller and thicker. This reduces water loss success of shrubs from the base is inversely related
and regulates leaf temperature, but it also to fire intensity (Westman and others 1981, Martin
decreases photosynthetic capacity. Prolonged 1984). In one study, 2 to 30 percent of brittlebush
drought leads to dormancy and leaf drop. resprouted on slopes previously dominated by the
Brittlebush shows variation among shrub, and resprouts and seedlings surpassed
individuals and populations in water-use efficiency prefire densities within 2 years (Martin 1984). For
(the ratio of photosynthesis to transpiration, or coastal sage scrub in general, both resprouting and
delta value). Individuals with a high delta have a seedling emergence from a soil seed bank are
higher growth response if water stress is decreased negatively correlated with fire intensity (Keeley
but perform poorly in response to drought stress, 1998).
while those with a low delta show lower growth
response under low water stress and a greater Seed germination.—Some authors report seed
germination without pretreatment (Mirov and
303
Kraebel 1939, Emery 1988), but pretreatment can pellets in spring, summer, and fall in trace amounts
increase otherwise low germination rates. Padgett (Perry and others 1987). The dominant herbivores
and others (1999) found that seed stored for 6 on brittlebush leaves are the larvae and adults of
months at room temperature had 2 to 4 percent the beetle, Trirhabda geminata (Wisdom 1985).
germination while seed stored at 5 to 10 oC in a The fly Neotephritis finalis Loew lays its eggs
refrigerator had 10 to 12 percent germination. In between the florets, and the larvae feed on the
addition, treatment with gibberelic acid or achenes (Goeden and others 1987).
Ca(NO3)2 increased germination of both warm-
and cold-stored seeds approximately two-fold, and Horticulture.—Brittlebush does best with full sun
leaching with water for several days increased and good drainage (Hickman 1993, Keator 1994).
germination by about 50 percent. The rounded form with striking yellow flowers is
attractive near the back of borders or rock gardens,
Growth and Management.—Maximum growth and it is especially suitable on dry slopes (Perry
of roots occurs in the winter and early spring 1992, Keator 1994). Plants can be established
(Drennan and Nobel 1996), so plants will establish quickly from seed or containers (Perry 1992).
best if sown in late fall. Planting brittlebush seeds Provision of occasional summer water allows
can yield vegetative cover and visual appeal plants to remain attractive throughout the year
relatively quickly. However, seed mixtures should (Keator 1994).
be balanced carefully because overuse can retard
establishment of other species (Went 1942, Gray References
and Bonner 1948). Ample evidence for local
adaptation in this species underscores the need to Bean, J.L. and K.S. Saubel. 1972. Temalpakh:
collect seed material for wildland restoration from Cahuilla Indian Knowledge and Usage of Plants.
within the same ecological zone and vegetation Malki Museum Press, Morongo Indian
type as the targeted planting site so as to maximize Reservation, CA. 225 p.
success of planting projects. Because of potential
competition and hybridization, it is also important Drennan, P.M. and P.S. Nobel. 1996. Temperature
that correct native species are specified and used. influences on root growth for Encelia farinosa
Mistaken plantings of E. californica instead of (Asteraceae), Pleuraphis rigida (Poaceae), and
brittlebush, or vice versa, abound (authors’ Agave deserti (Agavaceae) under current and
observation), and hybrids between species have doubled CO2 concentrations. American Journal of
been found in such locations (personal Botany 83: 133-139.
communication with A. Sanders, University of
California, Riverside). Improper seed choices can Ehleringer, J.R. 1984. Intraspecific competitive
compromise the success of restoration efforts and effects on water relations, growth and
the genetic integrity of wild populations. reproduction in Encelia farinosa. Oecologia 63:
153-158.
Benefits.—Brittlebush was used by native tribes
for medicinal and other purposes. The resinous Ehleringer, J.R. 1993. Variation in leaf carbon
gum, heated or made into a salve, was applied to isotope discrimination in Encelia farinosa:
the chest to relieve pain and loosen bronchial implications for growth, competition, and drought
mucous. A decoction of boiled blossoms, leaves, survival. Oecologia 95: 340-346.
and stems was held in the mouth to relieve gum
and tooth ache (Bean and Saubel 1972, Moore Ehleringer, J.R. and C. Clark. 1988. Evolution and
1989). In addition, tea made from the gum has a adaptation in Encelia (Asteraceae). In: L.
numbing effect and was used to relieve arthritic Gottlieb, L. Jain and S. Jain, eds. Plant
pain (Moore 1989). The resin was also burnt as Evolutionary Biology. Chapman and Hall,
incense or melted and used as a varnish (Moore London. p. 221-248.
1989, Hickman 1993).
Brittlebush feeds numerous pollinators and Emery, D.E. 1988. Seed propagation of native
herbivores. It is an important nectar and pollen California Plants. Santa Barbara Botanical
source of the bee, Calliopsis pugionis Cockerell, Garden, Santa Barbara, CA. 23 p.
which is the host of the rare cleptoparasitic bee,
Holcopasites ruthae Cooper in Riverside County, Goeden, R.D., T.D. Cadatal, and G.A. Cavender.
CA (Visscher and Danforth 1993). Mountain 1987. Life history of Neotephritis finalis (Loew)
sheep eat brittlebush, but it is only found in fecal on native Asteraceae in southern California
304
(Diptera: Tephritidae). Proceedings of the Monson, R.K., S.D. Smith, J.L. Gehring, W.D.
Entomological Society of Washington 89: 552- Bowman, and S.R. Szarek. 1992. Physiological
558. differentiation within an Encelia farinosa
population along a short topographic gradient in
Gray, R. and J. Bonner. 1948. An inhibitor of plant the Sonoran Desert. Functional Ecology 6: 751-
growth from the leaves of Encelia farinosa. 759.
American Journal of Botany 35: 52-57.
Moore, M. 1989. Medicinal Plants of the Desert
Hickman, J.C., ed. 1993. The Jepson Manual: and Canyon West. Museum of New Mexico
Higher Plants of California. University of Press, Santa Fe, NM. 184 p.
California Press, Ltd., Los Angeles, CA. 1,400 p.
Munz, P.A. and D.D. Keck. 1968. A California
Housman, D.C., M.V. Price, and R.A. Redak. Flora with Supplement. University of California
2002. Architecture of coastal and desert Encelia Press, Berkeley, CA. 1,681 + 224 p.
farinosa (Asteraceae): consequences of plastic
and heritable variation in leaf characters. Nobel, P.S., H. Zhang, R. Sharifi, M. Castañeda,
American Journal of Botany 89: 1,303-1,310. and B. Greenhouse. 1998. Leaf expansion, net
CO2 uptake, rubisco activity, and efficiency of
Keator, G. 1994. Complete garden guide to the long-term biomass gain for the common desert
native shrubs of California. Chronicle Books, San subshrub Encelia farinosa. Photosynthesis
Francisco, CA. 314 p. Research 56: 67-73.
Keeley, J.E. 1998. Postfire ecosystem recovery and Padgett, P.E., L. Vázquez, and E.B. Allen. 1999.
management: The October 1993 large fire Seed viability and germination behavior of the
episode in California. In: J.M. Moreno, ed. Large desert shrub Encelia farinosa Torrey & A. Grey
Forest Fires. Backbuys Publishers, Leiden, (Compositae). Madroño 46: 126-133.
Netherlands. p. 69-90.
Perry, B. 1992. Landscape Plants for Western
Kunze, A., C. Muller, and P. Proksch. 1995. Regions: an Illustrated Guide to Plants for Water
Chemical variation and defense of Encelia Conservation. Land Design Publishing,
farinosa. Biochemical Systematics and Ecology Claremont, CA. 318 p.
23: 355-363.
Perry, W.M., J.W. Dole, and S.A. Holl. 1987.
Kyhos, D.W. 1967. Natural hybridization between Analysis of the diets of mountain sheep from the
Encelia and Geraea (Compositae) and some San Gabriel Mountains, California. California
related experimental investigations. Madroño 19: Fish and Game 73: 156-162.
33-43.
Sandquist, D.R. and J.R. Ehleringer. 1996. Potential
Kyhos, D.W. 1971. Evidence of different adaptability and constraints of response to the
adaptations of flower color variants of Encelia changing climates for Encelia farinosa var.
farinosa (Compositae). Madroño 21: 49-61. phenicondonta from southern Baja California,
Mexico. Madroño 43: 465-478.
Martin, B.D. 1984. Influence of slope aspect on
postfire reproduction of Encelia farinosa Sandquist, D.R. and J.R. Ehleringer. 1997.
(Asteraceae). Madroño 31: 187-189. Intraspecific variation of leaf pubescence and
drought response in Encelia farinosa associated
Mirov, N.T. and C.J. Kraebel. 1939. Collecting and with contrasting desert environments. New
handling seeds of wild plants. Forestry Phytologist 135: 635-644.
Publication 5. Civilian Conservation Corps,
United States Government Printing Office, Sandquist, D.R. and J.R. Ehleringer. 1998.
Washington, D.C. Intraspecific variation of drought adaptation in
brittlebush: Leaf pubescense and timing of leaf
Moldenke, A.R. 1976. California pollination loss vary with rainfall. Oecologia 113: 162-169.
ecology and vegetation types. Phytologia 34: 305-
361. Sandquist, D.R. and J.R. Ehleringer. 2003. Carbon
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between contrasting populations of Encelia substrate on post-fire growth of California coastal
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nesting, foraging, and investment sex ratio.
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against a specialist herbivore. Journal of
Went, F.W. 1942. The dependence of certain annual Chemical Ecology 11: 1,553-1,565.
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Bulletin of the Torrey Botanical Club 69: 100- Wisdom, C.S. and E. Rodriguez. 1983. Seasonal
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1981. The effects of fire intensity, aspect and 352.
306
Ephedra nevadensis S. Wats. Nevada ephedra
EPHEDRACEAE
Stanley G. Kitchen
Taxonomy.—Ephedra is monotypic to
Ephedraceae with a worldwide distribution (except
for Australia) and is represented by approximately
60 species (Stevenson 1993). Two varieties, of
Nevada ephedra are recognized; E. nevadensis var.
nevadensis and var. aspera (Engelm. ex S. Wats)
L. Benson (Kartesz 1994). Chromosome number is
2n = 14 or 28 (Stevensen 1993).
307
recover after burning (Stanton 1974, Wright 1980, up to 5 years (Stevens and others 1981). Although
McLaughlin and Bowers 1982). A lack of fuel seedlings tolerate drought well, they are slow
continuity in many Nevada ephedra communities growing and may take 5 to 10 years to reach 60 cm
results in fire being a rare event. Plants spread in height (Plummer and others 1968). Plants
vegetatively through underground stolons in establish well from direct seeding when more
response to burial. Natural recruitment from seeds competitive species are controlled. Successes with
is often from rodent caches (Tirmenstein 1990). transplants and cuttings to revegetate minelands
Of North American ephedra, Nevada ephedra is and other degraded shrublands have also been
considered to be the most important forage species reported (Luke and Monsen 1984, Shaw and
(Tirmenstein 1990). It is used extensively by mule Monsen 1984).
deer (Ococolius hemionus), pronghorn
(Antilocapra americana), and bighorn (Ovis Benefits.—Nevada ephedra provides forage and
canadensis) and domestic sheep during winter and cover for numerous species of wildlife, especially
in periods of drought (Stanton 1974, Smith and during winter and periods of drought (Stanton
Beale 1980). Nutritive value and palatability of 1974, Smith and Beale 1980, Dittberner and Olsen
young branchlets is considered to be fair to good 1983). It is also used as forage by domestic sheep
(Tirmenstein 1990). and cattle but is considered poor forage for horses.
The species is used effectively in restoration
Reproduction.—Nevada ephedra is dioecious planting on harsh sites (Plummer and others 1968,
(occasional monoecious) with female and male Luke and Monsen 1984, Shaw and Monsen 1984).
cones occurring on separate plants. Male plants are Indigenous humans used roasted seeds in foods
more common on dry slopes and female plants and various concoctions from seeds and stems to
dominate more favorable locations, maximizing treat a variety of symptoms including coughs,
reproductive output (Freeman and others 1976). headaches, cold, fever, and kidney ailments
Plants are wind-pollinated. The timing of cone (Steward 1938, Tirmenstein 1990). Mormon
maturation, or coning, and pollination is weather- pioneers also made teas from stems. All members
dependent and generally occurs over a period of 1 of the genus produce unique secondary compounds
to 2 weeks between early March and early June with potential medicinal applications (Caveney
(Stanton 1974, Turner and Randall 1987). Seeds and others 2001). Nevada ephedra, and other
ripen between May and August. Turner and North American members of the genus, lack the
Randall (1987) observed considerable inter-annual nueroactivating ephedrine alkaloids found in some
variation in timing of phonological development. Old World species. However, a class of
Cones are not produced every year, and good seed compounds with antimicrobial potential,
crops are typically several years apart for any kynurenates, are found in Nevada ephedra
given population (Plummer and others 1968). (Caveney and others 2001). It is occasionally used
Years of good cone production are apparently as an ornamental (Cronquist and others 1972).
synchronized among most plants of a population
by an interaction between years of above-average References
precipitation and time since last major seed crop.
There are approximately 45,000 seeds (nutlets) per Caveney, S., D.A. Charlet, H. Freitag, M. Maier-
kg (Kitchen and others 1999). Stolte, and A.N. Starratt. 2001. New
observations on the secondary chemistry of
Growth and Management.—Once ripe, seeds are world Ephedra (Ephedraceae). American
easily dislodged. Ripened seeds can be collected Journal of Botany 88: 1,199-1,208.
by gently striking or shaking seed-bearing
branches over tarps or hand-held hoppers. In good Cronquist, A., A.H. Holmgren, N.H. Holmgren,
years, large quantities of seeds can be harvested in and J.L. Reveal. 1972. Intermountain flora:
relatively short time if harvests are well timed. Vascular plants of the Intermountain West,
After drying, seeds are cleaned to remove light, U.S.A. Vol. 1. Hafner Publishing Company,
unfilled seeds, stem fragments, cone scales, and Inc., New York. 270 p.
other inert material. Purity and viability levels of
commercially available seed are generally high. Dittberner, P.L. and M.R. Olson. 1983. The plant
Seed germination is rapid across a wide range of information network (PIN) data base: Colorado,
temperatures (5 to 20 oC) with little or no Montana, North Dakota, Utah, and Wyoming.
dormancy (Young and others 1977, Kitchen and FWS/OSB-83/86. U.S. Department of the
others 1999). Seed viability has remained high for Interior, Fish and Wildlife Service, Washington
308
DC. 786 p. Smith, A.D. and D.M. Beale. 1980. Pronghorn
antelope in Utah: some research and
Freeman, D.C., L.G. Klikoff, and K.T. Harper. observations. Publication 80-13. Utah Division
1976. Differential resource utilization by the of Wildlife Resources, Salt Lake City, UT. 88 p.
sexes of dioecious plants. Science 193: 597-599.
Stanton, F. 1974. Wildlife guidelines for range fire
Kartesz, J.T. 1994. A synonymized checklist of the rehabilitation. Technical Note 6712. U.S.
vascular plants of the United States, Canada, and Department of the Interior, Bureau of Land
Greenland. Vol. II—thesaurus . 2nd ed. Portland, Management, Denver, CO. 90 p.
OR: Timber Press. 816 p.
Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
Kitchen, S.G., S. Walker, and G.R. Wilson. 1999. Viability of seed from thirty-two shrub and forb
Addition of Ephedra nevadensis—Nevada species through fifteen years of warehouse
ephedra to the Rules. The Seed Technologist storage. Great Basin Naturalist 41: 274-277.
Newsletter 73: proposal #9.
Stevensen, D.W. 1993. Ephedraceae. In: Flora of
Luke, F. and S.B. Monsen. 1984. Methods and North America. http://flora.huh.harvard.edu/
costs for establishing shrubs on mined lands in [not paged].
southwestern Wyoming. In: A.R. Tiedemann,
E.D. McArthur, H.C. Stutz, R. Stevens, and K.L. Steward, J.H. 1938. Basin-Plateau Aboriginal
Johnson, comps. Proceedings—Symposium on Sociopolitical Groups. Bulletin 120.
the biology of Atriplex and related chenopods; Smithsonian Institution Bureau of American
1983 May 2-6; Provo, UT. General Technical Ethnology, Washington, DC. [1997 reprinted by
Repeport INT-172. U.S. Department of the University of Utah Press, Salt Lake City,
Agriculture, Forest Service, Intermountain UT]. 346 p.
Forest and Range Experiment Station, Ogden,
UT. p. 286-291. Tirmenstein, D. 1990. Ephedra nevadensis. In:
U.S. Department of Agriculture, Forest Service,
McLaughlin, S.P. and J.E. Bowers. 1982. Effects Fire Sciences Laboratory. Fire Effects
of wildfire on a Sonoran Desert plant Information System, http://www.fs.fed.
community. Ecology 63: 246-248. us/database/feis/ [not paged].
Plummer, A.P. 1977. Revegetation of disturbed Turner, F.B. and D.C. Randall. 1987. The
Intermountain area sites. In: J.C. Thames, ed. phenology of dessert shrubs in southern Nevada.
Reclamation and use of disturbed lands of the Journal of Arid Environments 13: 119-128.
Southwest. University of Arizona Press, Tucson,
AZ. p. 302-337. West, N.E. and M.A. Hassan. 1985. Recovery of
sagebrush-grass vegetation following wildfire.
Plummer, A.P., D.R. Christensen, and S.B. Journal of Range Management 38: 131-134.
Monsen. 1968. Restoring big-game range in
Utah. Publication 68-3. Utah Division of Fish Wright, H.A. 1980. The role and use of fire in the
and Game, Ephraim, UT. 183 p. semidesert grass-shrub type. General Technical
Report INT-85. U.S. Department of Agriculture,
Shaw, N. and S.B. Monsen. 1984. Nursery Forest Service, Intermountain Forest and Range
propagation and outplanting of bareroot Experiment Station, Ogden, UT: 24 p.
chenopod seedlings. In: A.R. Tiedemann, E.D.
McArthur, H.C. Stutz, R. Stevens, and K.L. Young, J.A. and R.A. Evans. 1978. Population
Johnson, comps. Proceedings—Symposium on dynamics after wildfires in sagebrush
the biology of Atriplex and related chenopods; grasslands. Journal of Range Management 31:
1983 May 2-6; Provo, UT. General Technical 283-289.
Report INT-172. U.S. Department of
Agriculture, Forest Service, Intermountain Young, J.A., R.A. Evans, and B.L. Kay. 1977.
Forest and Range Experiment Station, Ogden, Ephedra seed germination. Agronomy Journal
UT. p. 251-260. 69: 209-211.
309
Ephedra viridis Coville green ephedra
EPHEDRACEAE
Stanley G. Kitchen
310
bunchgrass, black sagebrush, blackbrush, desert shattering after seeds ripen. After drying, seeds are
grassland, pinyon-juniper, mountain mahogany, cleaned to high purity and viability levels using
oak woodland, mountain shrub, mixed conifer, and conventional milling equipment. Freshly-harvested
ponderosa pine (Anderson 2001). It is present in seed may be partially dormant (Meyer and others
early, mid, and late successional stages (Young 1988). Dormancy can be broken with a 4-week
and Evans 1973, Koniak 1985). Mean annual moist prechill or stratification (Meyer and others
precipitation varies from 200 to 430 mm 1988). Germination for after-ripened or prechilled
(Anderson 2001). Plants sprout from crown or seed is relatively rapid. Germination of non-
roots following top-kill by fire (USDA Forest dormant seeds occurs over a range of cool
Service 1976, Young and Evans 1978, Koniak temperatures (Young and others 1977). Seed
1985, Everett 1987). Reported survival following viability remains high for at least 5, and perhaps as
fire is variable (Anderson 2001). Sprouting long as 15 years (Stevens and others 1981).
response following fire may vary by season of Seedling growth rate is slow. Seedlings may
burn, fire severity, or among population variation require 5 to 10 years to reach 60 cm in height
in fire tolerance. Green ephedra is found in (Anderson 2001). In restoration plantings,
communities for which a wide range of fire seedlings are susceptible to competition from
frequencies is known. It also establishes on weeds and faster-growing grasses and forbs in the
disturbed sites from seed (Koniak 1985). Green seed mix (Ferguson and Frischknecht 1981).
ephedra is used as winter forage by big game and Plants establish well from direct seeding and
livestock but receives only limited use during transplants (USDA Forest Service 1976, Plummer
other seasons (Jameson and others 1962, Kufeld 1977, Everett 1980, Luke and Monsen 1984). Stem
and others 1973). Palatability and nutritive quality cutting are capable of root development in
of young stems are generally ranked as fair to greenhouse conditions (Wieland and others 1971).
good for most wildlife species and classes of Green ephedra was toxic to domestic cattle and
domestic livestock (Dittberner and Olson 1983, sheep during pregnancy in a controlled diet study
Anderson 2001). Seeds are eaten by birds and (Keeler 1989). No detrimental effects were
rodents and appear to be moderately prefered observed for developing calves and lambs.
(Everett and others 1978). Rodents probably play Conflicting reports have been published on the
an important role in seed dispersal. possible presence of ephedrine alkaloids in green
ephedra. While Anderson (2001) suggests the
Reproduction.—Green ephedra is dioecious. presence of this group of compounds in this
Spatial segregation of the sexes results from high species, Caveney and others (2001) failed to find
male plant representation on dry ridges and slopes these compounds common to Old World ephedras
and high female plant representation on microsites in any New World species. Tannin content is high.
with relatively favorable soil moisture conditions
(Freeman and others 1976, Brunt and others 1988) Benefits.—Green ephedra is valuable for restoring
providing for greater reproductive efficiency. vegetation structure and diversity and for erosion
Plants are wind-pollinated. Timing of cone control on disturbed lands, including roadsides,
maturation, or coning, and seed maturation is minelands, and degraded rangelands (USDA
dependent upon variation in weather, elevation, Forest Service 1976, Everett 1980, Ferguson and
aspect, and geography. Coning and pollination Frischknecht 1981, Ferguson and Frischknecht
occur for approximately 2 weeks, usually between 1985, Herbel 1986), in areas in which it is adapted.
April and June. Seeds mature between June and When competition is controlled, it establishes well
September. Cones are not produced each year, and from seed and persists after wildfire. It provides
good seed crops may be several years apart important winter forage for wildlife and domestic
(Plummer and others 1968). There are livestock (USDA Forest Service 1976). Green
approximately 45,000 seeds per kg (Meyer and ephedra has had limited use as an ornamental
others 1988). (Anderson 2001). Native Americans made both
medicinal and nonmedicinal teas brewed from
Growth and Management.—In years of good stems and used seeds and roasted seeds (nutlets)
seed crops, significant quantities are easily for food (Steward 1938, USDA Forest Service
harvested by hand (Plummer and others 1968). 1976). Mormon settlers were also known to use it
Ripened cones are easily dislodged by gently in the production of a beverage, a cause for the
shaking or striking cone-bearing branches over frequently used moniker, ‘mormon tea’.
tarps or hand-held hoppers. Timing of harvest is Undetermined medicinal applications for a group
critical as thunder storms can cause considerable of unique secondary compounds associated with
311
members of the genus have not been fully establishment on reconstructed soils in semiarid
determined, including possible antimicrobial areas. In: Shrub establishment on disturbed arid
applications for kynurenates (Caveney and others and semi-arid lands: Proceedings of the
2001). symposium; 1980 December 2-3; Laramie, WY.
Wyoming Game and Fish Department. p. 57-63.
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Smithsonian Institution Bureau of American grasslands. Journal of Range Management 31:
Ethnology, U.S. Government Printing Office, 283-289.
Washington, DC. [1997 reprinted by the
University of Utah Press, Salt Lake City, UT]. Young, J.A., R.A. Evans, and B.L. Kay. 1977.
346 p. Ephedra seed germination. Agronomy Journal
69: 209-211.
313
Eriogonum fasciculatum Benth. California buckwheat
POLYGONACEAE
Arlee M. Montalvo
314
Eriogonum f. var. polifolium (Benth.) Torrey well on rocky road cuts and in shallow soils that
& A. Gray (flowering shoot in photograph) has 2n are inhospitable to annual grasses.
= 40 chromosomes (Stebbins 1942). These
greyish, pubescent, rounded shrubs occur at 600 to Reproduction.—Plants flower from early spring
2300 m on dry slopes and washes. They are through summer. The four taxa have slightly
widespread across inland mountains and deserts of shifted flowering times, with var. flavoviridae
California, from the inner coast ranges of San Luis having the earliest season (Munz and Keck 1968)
Obispo and Fresno Cos., south to San Diego Co., and var. polifolium flowering before the other two.
eastward into the Mojave Desert to southern In all taxa, stamens and stigmas are exerted and
Nevada, Utah, and Arizona, and south into open to generalist pollinators, including honey
northwestern Senora, Mexico and central Baja bees and many species of small native bees, flies,
California. In inland sage scrub, plants overlap in wasps, and beetles. Achenes mature from June to
distribution with var. foliolosum, especially in September and disperse when dried inflorescences
Riverside Co., but occupy drier sites. Leaves are shatter in autumn. The dry calyx provides
oblanceolate to obovate, densely pubescent buoyancy to detached achenes and assists dispersal
beneath, and usually tomentose above with leaf by wind and water. Harvester ants also disperse
margins sometimes barely revolute. Involucres and achenes (DeSimone and Zedler 1999). Seedlings
sepals are pubescent. Heads tend to be born in emerge in the rainy season in mid to late winter
simple to once-compound umbels. Reveal (1989) within light gaps and open areas. Survival is
reports hybridization between var. polifolium and unusually high as shown by studies in a variety of
E. cinereum Benth in areas were E. cinereum has sites and years, often ranging from 80 to 90
been introduced within the range of var. percent (Wright and Howe 1987, Miriti and others
polifolium. Synonyms are E. f. ssp. p. S. Stokes 1998, Montalvo and others 2002). Most shoot
and E. p. Benth. elongation and leaf growth occurs before
Eriogonum f. var. flavoviride Munz & flowering from late winter to mid spring (Cole
Johnson has n = 20 chromosomes (Reveal 1989) 1967). Plants can reach maturity in 1 to 2 years.
and is 2 to 3 dm tall. The rounded shrubs occur on
dry rocky slopes and washes from 200 to 1300 m Seed Germination.—Light improves germination
in creosote bush scrub of the southern Mojave (Cole 1967, Keeley 1987). Atwater (1980)
Desert, San Bernardino Co., south through eastern suggests that light increases permeability of seed
Riverside Co. to far eastern San Diego Co., and coats, water adsorption, and possibly leaching of
south to central Baja California. Leaves are light inhibitors, allowing seeds to germinate. Light-
green to yellowish-green, linear, strongly revolute, induced seed coat permeability would allow some
and glabrous above. Peduncles are glabrous; buried seeds to remain viable and dormant in the
involucres and sepals are subglabrous. Synonym is seed bank, a bet-hedging strategy. Cole (1967)
E. f. ssp. flavoviride S. Stokes. found coastal populations germinated at 15 to 25
o
C while inland populations germinated over a
Ecology.—California buckwheat occurs on dry larger range of 5 to 35 oC. Keeley (1987)
slopes and often granitic substrate (Westman examined the effects of light, dark, heat, and
1981). The woody, branched roots penetrate to leachate through charred wood (charrate) on
under 1.5 m (Kummerow and others 1977). Plants germination. Seeds germinated equally well on
lose some leaves in the dry season and can shed soil vs. filter paper, and light controls had a two-
many leaves in severe drought. Plants are fold increase in germination compared to dark
vulnerable to hot fires so resprout success is low controls (83 vs. 45 percent germination). Charrate
and most regeneration is from seeds (Keeley did not affect germination. In light, seeds exposed
1998). Frequent fires deplete the seed bank, to 120 oC for 5 min suffered a large decrease in
making populations vulnerable to local extinction. germination under all conditions relative to seeds
In Riverside Co., this shrub has decreased heated to 70 or 100 oC, suggesting fire may kill
dramatically since the 1940’s, with the largest many seeds in the upper centimeter of the
decreases on alluvial soils (Minnich and Dezzani seedbank. Heat sensitivity and a light requirement
1998). Decreases are attributed to a combination of for germination encourage colonization of
competition with exotic annual grasses, too disturbed or open sites.
frequent a fire interval, that is exacerbated by
weeds which carry fire, and by atmospheric Horticulture.—Plants can be propagated from
deposition of nitrogen, which encourages growth seeds or rooted from cuttings. Seeds and container
of competing weeds. California buckwheat does plants should be planted in the fall to early winter
315
to take advantage of the moist cool season for Vertebrates also use California
growth. In one study, seeds planted by seed buckwheat. The rare orange-throated whiptail is
imprinting and hydroseeding, shallow methods associated with open vegetation with this shrub
that allow light to reach planted seeds, had at least (Brattstrom 2000). Rare California gnatcatchers
twice the emergence success as drilling that (Polioptila californica californica) forage and nest
covered seeds with 0.5 to 1 cm of soil (Montalvo in the shrub. California buckwheat and Artemisia
and others 2002). Seeds germinate within 2 weeks californica Less. are dominant shrubs used by
if soil is moist and over about 15 oC. Plants require gnatcatchers (Mock and Bolger 1992). The rare
full sun and good drainage. No irrigation is mountain sheep (Ovis canadensis nelsoni) browses
required if planted before seasonal rains. on California buckwheat in the San Gabriel
The grey-green var. polifolium provides a Mountains of southern California (Perry and others
nice contrast to the other green varieties in the 1987), and deer eat the inflorescences
garden, but all do well in dry landscaping. The (Schopmeyer 1974).
flowers last far into the summer and become an
attractive rusty-brown in fruit. Some prostrate Genetics, Geographic Variation, Hybridization,
cultivars of California exist, including “Theodore and Fitness.—Stebbins (1942) hypothesized that
Payne,” “Prostrata,” “Dana Point,” and tetraploid populations of var. foliolosum arose
“Wildwood” (Perry 1992). from hybridization between diploid var.
fasiculatum and var. polifolium (= E. f. subsp.
Greenhouse studies indicate that California foliolosum, E. f. subsp. typicum, and E. f. subsp.
buckwheat forms facultative associations with polifolium, respectively, in the paper). He
arbuscular mycorrhizal fungi (Egerton-Warburton, examined the distribution of taxa in relation to
Montalvo, and Allen, unpublished report). The climatic factors and concluded that the
plant’s facultative dependence on mycorrhizal morphology and ecological tolerances of var.
fungi may be instrumental to successful foliolosum combined the morphology and
colonization of barren sites. Montalvo and others tolerances of the two diploid taxa. The variety
(2002) found that plants grew quickly to maturity polifolium was in areas with some snow, frost, and
on a graded site with low organic matter and only 10 to 13 cm of rainfall, fasciculatum was in
nutrients, but establishment, growth, and flowering the areas with no frost, often frequent fog, and 22
all decreased significantly with decreasing soil to 37 cm of rainfall, and foliolosum often occurred
nitrogen (NO3-N mean = 7.89 mg/g, range = 0.8- in habitats intermediate to the two diploids. Cole
69 mg/g in the study plot). (1967) tested assumptions about the distribution of
morphology and ecological tolerances of vars.
Benefits.—California buckwheat provides habitat fasciculatum and foliolosum. Cole compared their
and food for numerous animals. Osborne (1998) physiologies over an ecological gradient from the
found high abundance and diversity of arthropod cool coast across the mountains to the hot inland
species on California buckwheat. Some are valleys and found that coastal var. fasciculatum
species-specific feeders such as the larvae of the were the least hairy, and populations of var.
moth, Hemileuca electra (Wright) (Rubinoff foliolosum became more hairy inland. There was a
1998). Numerous species of butterfly larvae feed clear morphological cline from the coast to the
on California buckwheat and specialize on interior among and within ecotypes that correlated
different plant parts and taxa (Howe 1975), with habitat. Photosynthesis vs respiration ratios
sometimes according to timing of growth. For and photosynthetic rates and respiration rates all
example, the larvae of the butterfly, Apodemia varied in a clinal manner that correlated with
mormo (Felder & Felder), has three biotypes in the morphological characters. The variety
Mojave Desert. The spring morph feeds on fasciculatum forms a distinctly adaptive coastal
California buckwheat while the others feed on ecotype, while var. foliolosum forms an inland
species with different flowering times. E. ecotype.
parvifolium Sm. in Rees. is the host of the rare El
Segundo blue butterfly (Euphilotes bernardino Analysis of eight allozyme loci for eight
allyni Shields). When California buckwheat was populations of var. foliolosum and four
planted instead of later flowering E. parvifolium, fasciculatum revealed ample genetic variation
competing insect species used California (Montalvo, Ellstrand, and Clegg, unpublished
buckwheat, built up populations, and then data). All loci were polymorphic and there was an
competed with the rare butterfly for E. parvifolium average of 5.7 alleles/locus. Expected
flowers (Longcore and others 2000). heterozygosity was also high at 0.42. Nei’s genetic
distances among populations ranged from 0 to
316
0.074, with the largest distance between varieties. DeSimone, S.A., and P.H. Zedler. 1999. Shrub
Overall, the proportion of total variation explained seedling recruitment in unburned Californian
by differences among populations (GST) was coastal sage scrub and adjacent grassland.
0.025. By itself, the tetraploid var. foliolosum had Ecology 80: 2018-2032.
no significant structure (GST = 0.005), while GST =
0.028 for var. fasciculatum. These values suggest Hickman, J.C., ed. 1993. The Jepson Manual:
high levels of gene flow and outbreeding. Higher Plants of California. University of
California Press, Ltd., Los Angeles, CA. 1,400 p.
Seed Collection and Processing.—Seeds are
collected after heads have turned rusty brown. The Howe, W.H., ed. 1975. The butterflies of North
chaff can be separated from seeds by pushing America. Doubleday and Company, Inc., Garden
seeds through a screen and then separating seeds City, NY. 633 p.
from chaff with an air separator or fan (author’s
observation). Commercial seed companies usually Keeley, J.E. 1987. Role of fire in seed germination
skip the screening and just break up the chaff, air of woody taxa in California chaparral. Ecology
separate the seeds from crude chaff, and sell seeds 68: 434-443.
with the persistent calyx intact. A recommended
target for minimum quality of purchased seeds is Keeley, J.E. 1998. Postfire ecosystem recovery and
15 percent purity and 65 percent germination. management: The October 1993 large fire
Here, a bulk seed pound with about 990,000 episode in California. In: J. M. Moreno (ed.).
seeds/kg, would have about 96,800 pure live seeds Large Forest Fires. Backbuys Publishers, Leiden,
(personal communication, S & S Seeds, Netherlands. p. 69-90.
Carpenteria, CA).
Kirkpatrick, J.B., and C.F. Hutchinson. 1977. The
Growth and Management.—Seeds have been community composition of California coastal
widely planted for restoration, roadside erosion sage scrub. Vegetatio 35: 21-33.
control, slope stabilization, landscaping, and
apiary (Schopmeyer 1974, Perry 1992). Despite Kummerow, J., D. Krause, and W. Jow. 1977. Root
the clear geographic pattern and ecological systems of chaparral shrubs. Oecologia 29: 163-
affinities of the four taxonomic varieties, most 177.
researchers and consultants fail to designate
taxonomic variety on publications, plant lists, or Longcore, T., R. Mattoni, G. Pratt, and C. Rich.
plant palettes. They also commonly fail to specify 2000. On the perils of ecological restoration:
appropriate ecological zone for seeds used in lessons from the El Segundo blue butterfly. In: J.
wildland plantings. This has resulted in extensive E. Keeley, M. Baer-Keeley and C. J.
plantings outside natural ranges and habitats Fotheringham, eds. 2nd Interface Between
(Reveal 1989). Projects should use seeds of Ecology and Land Development in California:
appropriate taxa to maximize project success. U.S. Geological Survey Open-File Report 00-62.
Vol. 62. Sacramento, CA. p. 281-286.
References
Minnich, R.A., and R.J. Dezzani. 1998. Historical
Atwater, B.R. 1980. Germination, dormancy and decline of coastal sage scrub in the Riverside-
morphology of the seeds of herbaceous Perris plane, California. Western Birds 29: 366-
ornamental plants. Seed Science and Technology 391.
8: 523-573.
Miriti, M.N., H.F. Howe, and S.J. Wright. 1998.
Brattstrom, B.H. 2000. The range, habitat Spatial patterns of mortality in a Colorado desert
requirements, and abundance of the orange- plant community. Plant Ecology 136: 41-51.
throated whiptail, Cnemidophorus hyperythrus
beldingi. Bulletin of the Southern California Mock, P.J., and D. Bolger. 1992. Ecology of the
Academy of Sciences 99: 1-24. California gnatcatcher at Rancho San Diego.
Ogden Environmental and Energy Services Co.,
Cole, N.H. A. 1967. Comparative physiological Inc.; Project No. 110970000. 50 p.
ecology of the genus Eriogonum in the Santa
Monica Mountains, southern California. Montalvo, A.M., P.A. McMillan, and E.B. Allen.
Ecological Monographs 37: 1-24. 2002. The relative importance of seeding method,
317
soil ripping, and soil variables on seeding success. Rubinoff, D. 1998. Field observations on mating
Restoration Ecology 10: 52-67. behavior and predation of Hemileuca electra
(Saturniidae). Journal of the Lepidopterists'
Munz, P.A., and D.D. Keck. 1968. A California Society 52: 212-214.
Flora with Supplement. University of California
Press, Berkeley, CA. 1681 + 224 p. Schopmeyer, C.S., ed. 1974. Seeds of Woody Plants
in the United States. Vol. 450 Agricultural
Osborne, K.H. 1998. A description of arthropod Handbook. USDA Forest Service, Washington,
community structure in southern California D.C. 883 p.
coastal sage scrub. M.S. Thesis. University of
California, Riverside, CA. 133 p. Stebbins, G.L. 1942. Polyploid complexes in
relation to ecology and the history of floras. The
Perry, B. 1992. Landscape plants for western American Naturalist 76: 36-45.
regions: an illustrated guide to plants for water
conservation. Land Design Publishing, Westman, W.E. 1981. Factors influencing the
Claremont, CA. 318 p. distribution of species of California coastal sage
scrub. Ecology 62: 439-455.
Perry, W.M., J.W. Dole, and S.A. Holl. 1987.
Analysis of the diets of mountain sheep from the Wright, S.J., and H.F. Howe. 1987. Pattern and
San Gabriel Mountains, California. California mortality on Colorado Desert plants. Oecologia
Fish and Game 73: 156-162. 73:543-552.
318
Erithalis fruticosa L. black torch
RUBIACEAE
John K. Francis
319
growing, especially when situated on rock Liogier, H.A. 1990. Plantas medicinales de
outcrops. Management should mainly consist of Puerto Rico y del Caribe. Iberoamericana de
protection from fires and clearing of sites for Edicones, Inc., San Juan, PR. 566 p.
development.
Liogier, H.A. 1997. Descriptive flora of Puerto
Benefits.—Black torch contributes to biodiversity Rico and adjacent islands. Vol. 5. Editorial de la
in the forests where it grows, helps protect the soil, Universidad de Puerto Rico, San Juan, PR.
and furnishes food and cover for wildlife. It would 436 p.
probably make an acceptable ornamental for green
areas and nonirrigated gardens. The wood is good Little, E.L., Jr., R.O. Woodbury, and F.H.
for turned articles but available only in small sizes. Wadsworth. 1974. Trees of Puerto Rico and the
It is resistant to rot and has been used for piles and Virgin Islands. Vol. 2. Agriculture Handbook
posts. Formerly, it was split and used for torches 449. U.S. Department of Agriculture,
(Little and others 1974) but tended to be smoky Washington, DC. 1,024 p.
(University of the Virgin Islands 2002). The bark,
fruits, and the resin have diuretic and astringent Long, R.W. and O. Lakela. 1976. A flora of
properties and are used to treat inflammation of the Tropical Florida. Banyan Books, Miami, FL.
kidney and bladder, and blennorrhoea (Liogier 962 p.
1990). The leaves are parched and ground and used
to treat skin sores (University of the Virgin Islands Nelson, G. 1996. The shrubs and woody vines of
2002). Florida. Pineapple Press, Inc., Sarasota, FL.
390 p.
References
University of the Virgin Islands. 2002. Erithalis
Howard, R. A. 1989. Flora of the Lesser Antilles, fruticoas (blacktorch). http://rps.uvi.edu/
Leeward and Windward Islands. Vol. 6. Arnold VIMAS/blacktorch.htm. 1 p.
Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
320
Ernodea littoralis Sw. golden creeper
RUBIACEAE
John K. Francis
321
Benefits.—Golden creeper contributes to the Gilman, E.F. 1999. Ernodea littoralis. Fact Sheet
biodiversity of the sites where it grows, stabilizes FPS-196. Cooperative Extension Service,
the soils in those fragile environments, and University of Florida. http://
provides food and cover for wildlife. It is planted hort.ifas.ufl.edu/shrubs/ERNLITA.pdf. 3 p.
as an ornamental for landscaping, usually as
ground cover. Golden creeper is also used for Howard, R.A. 1989. Flora of the Lesser Antilles,
beach dune stabilization. It is particularly Leeward and Windward Islands. Dicotyledoneae.
promoted for water-restricted landscaping Part 3. Vol. 6. Arnold Arboretum, Harvard
(Workman 1980). A tea is made from the leafy University, Jamaica Plain, MA. 658 p.
shoots as a treatment for coughs (Centro Ecologico
Sian Ka’an 2002). Liogier, H.A. 1997. Descriptive flora of Puerto
Rico and adjacent islands. Vol. 5. Editorial de la
References Universidad de Puerto Rico, San Juan, PR.
436 p.
Carney-Kulig, S., M. Jolly, T. Weyant, S. Buehler,
and G. Davis. 2002. Planting the right plant in Nelson, G. 1996. The shrubs and woody vines of
the right place. Monroe County Extension Florida. Pineapple Press, Inc., Sarasota, FL.
Service, University of Florida. http:// 391 p.
monroe.ifas.ufl.edu/gardeningguide_5.htm. 6 p.
Workman, R.W. 1980. Growing native. The
Centro Ecologico Sian Ka’an. 2002. Flora. Sanibel-Captiva Conservation Foundation, Inc.,
http://cesiak.org/flora1.htm. 3 p. Sanibel, FL. 137 p.
322
Erythroxylum areolatum L. false cocaine
ERYTHROXYLACEAE
John K. Francis
323
performed. Rico and adjacent islands, Spermatophyta. Vol.
2. Editorial de la Universidad de Puerto Rico,
Benefits.—False cocaine protects the soil and Río Piedras, PR. 481 p.
furnishes food and cover for wildlife. The wood is
heavy, hard, fine-grained, durable, and strong. The Little, E.L., Jr. and F.L. Wadsworth. 1964.
sapwood is light brown and the heartwood is Common trees of Puerto Rico and the Virgin
reddish or chocolate brown (Little and Wadsworth Islands. Agriculture Handbook 249. U.S.
1964). Available only in small sizes, it is useful for Department of Agriculture, Washington, DC.
stakes, fuel, carving, and turnery. The foliage of 548 p.
this species does not contain useful amounts of
cocaine, which is extracted from a sister species (E. National Trust for the Cayman Islands. 2002.
coca Lam.) from the Andean Highlands (Little and Virtual herbarium of the National Trust for the
Wadsworth 1964). Cayman Islands: Erythroxylaceae, Erythroxylum
areolatum L. http://www.virtualherbarium.org/
References vh/CAYM/403.htm. [not paged].
Britton, N.L. and C.F. Millspaugh. 1962. The Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Bahama flora. The New York Botanical Garden, Montiel, eds. 2001. Flora de Nicaragua.
New York. 695 p. Monographs in Systematic Botany, Vol. 85, No.
1. Missouri Botanical Garden, St. Louis, MO.
Francis, J.K. and A. Rodríguez. 1993. Seeds of 943 p.
Puerto Rican trees and shrubs: Second
installment. Research Note SO-374. U.S. Vásquez, O.J. and D.A. Kolterman. 1998. Floristic
Department of Agriculture, Forest Service, composition and vegetation types of the Punta
Southern Forest Experiment Station, New Guaniquilla Natural Reserve—Cabo Rojo,
Orleans, LA. 5 p. Puerto Rico. Caribbean Journal of Science
43(304): 265-279.
Liogier, H.A. 1988. Descriptive flora of Puerto
324
Erythroxylum brevipes DC. brisselet
ERYTHROXYLACEAE
John K. Francis
325
of 33 brisselet stems 4 to 14 cm in diameter in St. Liogier, H.A. 1988. Descriptive flora of Puerto
John, U.S. Virgin Islands at 0.5 mm/year. Rico and adjacent islands, Spermatophyta. Vol.
Apparently, plants live for several decades. No 2. Editorial de la Universidad de Puerto Rico,
management experience has been published. Río Piedras, PR. 481 p.
Benefits.—Brisselet is an important component of Liogier, H.A. and L.F. Martorell. 2000. Flora of
dry and moist forests, protects the soil, and Puerto Rico and adjacent islands, a systematic
furnishes cover for wildlife. Although seasonal, the synopsis. 2nd Ed. Editorial de la Universidad de
fruits are a consistent source of food for birds. Bees Puerto Rico, Río Piedras, PR. 382 p.
are attracted to the fragrant flowers (Little and
others 1974). The sapwood is light tan while the Little, E.L., Jr., R.O. Woodbury, and F.H.
heartwood is dark brown, hard, and flexible. Wadsworth. 1974. Trees of Puerto Rico and the
Because of its small diameters and lengths, the Virgin Islands. Vol. 2. Agriculture Handbook
wood is useful for little other than fuel. Brisselet 449. U.S. Department of Agriculture,
tissues do not contain cocaine, as obtained from its Washington, DC. 1,024 p.
more famous relative Erythroxylum coca Lam.
Stevens, W.D., C. Ulloa-U., A. Pool, and O.H.
References Montiel, eds. 2001. Flora de Nicaragua.
Monographs of Systematic Botany Vol. 85, No.
Howard, R.A. 1988. Flora of the Lesser Antilles, 1. Missouri Botanical Garden Press. p. 1-943.
Leeward and Windward Islands.
Dicotyledoneae, Part 1. Vol. 4. Arnold Weaver, P.L. 1990. Tree diameter growth rates in
Arboretum, Harvard University, Jamaica Plain, Cinnamon Bay Watershed, St. John, U.S. Virgin
MA. 673 p. Islands. Caribbean Journal of Science 26(1-2):
1-6.
326
Eugenia foetida Pers. boxleaf stopper
MYRTACEAE
John K. Francis
327
Reproduction.—Flowering of boxleaf stopper in ruminant herbivores. Although small, the fruits are
Puerto Rico is reported to occur in spring and edible to humans, and birds consume them when
summer (Little and others 1974). It flowers in the available. Boxleaf stopper is recommended and
summer in Florida (Gilman 1999). Fresh fruits used for screens, hedges, parking lot, yard, and
collected in Puerto Rico averaged 0.181 + 0.032. street trees in Florida (Gilman 1999).
Air-dried seeds separated from them averaged
14,900 seeds/kg. Placed in moist potting mix, 51 References
percent germinated beginning 69 days after
sowing. The new seedlings grow slowly. In the Gilman, E.F. 1999. Eugenia foetida. Fact Sheet
wild, seeds are dispersed by birds. Seedlings and FPS-200. Cooperative Extension Service,
saplings are widely scattered to relatively University of Florida, Gainesville, FL. 3 p.
common.
Howard, R.A. 1989. Flora of the Lesser Antilles,
Growth and Management.—The growth rate of Leeward and Windward Islands. Vol. 5. Arnold
boxleaf stopper is moderate to slow, depending on Arboretum, Harvard University, Jamaica Plain,
habitat. Apparently, it is relatively long lived (a MA. 604 p.
few decades). Potted stock is available from
nurseries for planting. Although expensive for Liogier, H.A. 1994. Descriptive flora of Puerto
wildland plantings, this may be the best way to Rico and adjacent Islands. Vol. 3. Editorial de la
establish them in order to get them above Universidad de Puerto Rico. Río Piedras, PR.
competing weeds and grass. Ornamental plants 461 p.
may be pruned into hedges or forced to single-
stemmed trees (Gilman 1999). Because they are Little, E.L., Jr., R.O. Woodbury, and F.H.
relatively slow growing and not aggressive Wadsworth. 1974. Trees of Puerto Rico and the
invaders, control of the species in natural stands Virgin Islands. Vol. 2. Agriculture Handbook
should not be necessary. 449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
Benefits.—Boxleaf stopper protects the soil,
furnishes cover for wildlife, and contributes to the Nelson, G. 1996. The shrubs and woody vines of
aesthetics of the forest. Aromatic compounds Florida. Pineapple Press, Inc., Sarasota, FL.
released from the leaves of this species apparently 391 p.
are responsible for the slight musky smell present
in forests in Puerto Rico where it is common. Schaus, C., S. Wade and J. Dunan. 2003. Key deer
Foliage of the species is not eaten by Key deer and plants they won’t eat. http://monroe.
(Odocolleus virginanus clavium) (Schaus and ifas.ufl.edu/key_deer_plants.htm. 4 p.
others 2003) and probably is avoided by other
328
Eugenia pseudopsidium Jacq. wild guava
MYRTACEAE
John K. Francis
329
ground level, usually with more than one shoot References
(author’s observation). Birds disperse the seeds.
Seedlings and saplings are well-scattered and Howard, R.A. 1989. Flora of the Lesser Antilles,
relatively common. Wild guava sprouts readily Leeward and Windward Islands. Vol. 5. Arnold
after cutting or damage. Arboretum, Harvard University, Jamaica Plain,
MA. 604 p.
Growth and Management.—The growth rate of
wild guava appears to be relatively good for an Liogier, H.A. 1994. Descriptive flora of Puerto
understory shrub. A 2.5-m tall sapling, 2.5 cm Rico and adjacent Islands. Vol. 3. Editorial de la
d.b.h., under partial shade in Puerto Rico, had five Universidad de Puerto Rico. Río Piedras, PR.
growth rings in its trunk near the ground. Wild 461 p.
guava germinates, survives, and transplants well in
the nursery, but growth is somewhat slower than Little, E.L., Jr., R.O. Woodbury, and F.H.
most commercial nursery plants. Unfortunately, Wadsworth. 1974. Trees of Puerto Rico and the
no plantation or ornamental use has been reported. Virgin Islands. Vol. 2. U.S. Department of
Agriculture, Agriculture Handbook 449.
Benefits.—Wild guava is used to a limited extent Washington, DC. 1,024 p.
for firewood and fence posts. With its dark green
leaves, bright red fruits, and clean form, the Smithsonian. 2001. Regional plant species lists for
species has a pleasing appearance and probably Guyana. Mason.gmu.edu/~ckelloff/region2.pdf.
would make a fine ornamental, especially in shady 6 p.
locations. It provides food and cover for wildlife
and contributes to the scenic beauty and
biodiversity of the forests where it grows.
330
Euonymus alatus (Thunb.) Sieb. winged burning bush
CELASTRACEAE
331
mm or more and in elevations ranging from 40 to References
2,700 m (Ma 2001). It was first introduced into the
United States about 1860 (Rehder 1940), as an Brizicky, G.K. 1964. The genera of Celastrales in
ornamental plant, and it eventually became the southeastern United States. Journal of the
naturalized in the Eastern United States (Ebinger Arnold Arboretum. 45: 210-215.
1996, Gleason and Cronquist 1991, J. Ma in
personal observation). Currently winged burning Chittenden, F.J. and P.M. Synge. 1956. Dictionary
bush is known from 21 Eastern and Mid-Western of gardening. Vol. 2, 2nd Ed. Clarendon Press,
States (Connecticut, Illinois, Indiana, Iowa, Oxford, UK. 583 p.
Kentucky, Maryland, Massachusetts, Michigan,
Minnesota, Missouri, North Carolina, New Ebinger, J.E. 1996. Euonymus alatus. In: J.M.
Hampshire, New Jersey, New York, Ohio, Randall, J.M. and J. Marinelli, eds. Invasive
Pennsylvania, Rhode Island, South Carolina, plants–weeds of the global garden. 21st Century
Virginia, Vermont, West Virginia), as well as Gardening Series. Brooklyn Botanic Garden,
Montana (USDA 2003). It can be expected to be Brooklyn, NY. p 55.
found elsewhere with the continued spread of
naturalized populations and additional material Gilstra, Z. 1991. Proposal to conserve the spelling
escaping from cultivation. of 4618 Euonymus (Celastraceae). Taxon 40:
137-139.
Ecology.—It is not as aggressive an invader as the
related Oriental bittersweet (Celastrus orbiculatus Gleason, H.A. and A. Cronquist 1991. Manual of
Thunb). It is found in disturbed habitats, such as Vascular Plants of northeastern United States
roadsides, forest margins and woodlands that are and adjacent Canada. New York Botanical
in early stages of succession (Ma, personal Garden, Bronx, NY. 910 p.
observation).
Ma, J.S. 2001. A revision of Euonymus
Reproduction.—Hymenopterous and dipterous (Celastraceae). Thaiszia Journal of Botany 11:
insects, especially bees and flies, appear to be the 1-264.
main pollinators, and the attractive arillate seeds
are widely dispersed by birds and small mammals Martin, T. 2002. Weed alert! Euonymus alata
(Brizicky 1964). The flowers bloom in the spring (Thunb.) Siebold. http://tncweeds.ucdavis.
and the fruit mature in the fall. The species can be edu/alert/alrteuon.html. [not paged].
asexually propagated through stem cuttings and
root separation. Paclt, J. 1998. Proposal to amend the gender of
Euonymus, nom. cons. (Celastraceae) to
Growth and Management.—Winged burning feminine. Taxon 47: 473-474.
bush grows in a variety of soil conditions. It is not
a fast growing plant, reaching perhaps up to 1 m of Rehder, A. 1940. Manual of cultivated trees and
growth in a single growing season. Propagation shrubs. Macmillan Publishing Co., New York.
can be effected by layering or seed (Chittenden 996 p.
and Synge 1956). However, there is concern about
the continued use of this plant in ornamental Smith, B. 2003. The bonsai guide
plantings, since it can be invasive in some areas http://www.thebonsaiguide.com/BPG/BS_BPBT
(Martin 2002) E.HTM. [not paged].
Benefit.—The foliage of winged burning bush USDA, NRCS. 2003. The PLANTS Database.
turns bright scarlet in autumn, giving the plant its Baton Rouge, LA. http:plants.usda.gov. [not
common name burning bush. The purplish paged].
capsules and seeds with orange arils add to the
“burning” effect of the plant late in the year. This
effect of the plant has made it a popular
ornamental and horticultural species. Some
specimens can be shaped into bonsai (Smith 2003).
332
Euonymus americanus L. strawberry bush
CELASTRACEAE
333
classical Greek, the generic name was treated as hundreds of the “bursting” red capsules. In the
feminine; however, Linnaeus treated the genus as winter, the bright green twigs are also handsome.
masculine. The spelling “Euonymus” has formally
been conserved over Linnaeus’s originally spelling References
“Evonymus,” which has also occasionally been
used (Gilstra 1991) Britton, N.L. and A. Brown. 1913. Illustrated flora
of the northern states, Canada, and the British
Range.—Strawberry bush occurs throughout possessions. Vol. 2, 2nd Ed. Scribner, New York.
much of the Eastern United States (Alabama, 735 p.
Arkansas, Delaware, District of Columbia, Florida,
Georgia, Illinois, Indiana, Kentucky, Louisiana, Brizicky, G.K. 1964. The genera of Celastralesin
Maryland, Mississippi, Missouri, New Jersey, the Southeastern United States. Journal of the
New York, North Carolina, Ohio, Oklahoma, Arnold Arboretum. 45: 206-235.
Pennsylvania, South Carolina, Tennessee, Texas,
Virginia and West Virginia), except New England. Chittenden, F.J. and P.M. Synge. 1956. Dictionary
(Shetler and Orli 2000, USDA 2003) It is also of gardening. Vol. 2, 2nd ed. Clarendon Press,
known from Ontario (Gleason and Cronquist 1991, Oxford, UK. 583 p.
Ma 2001, Scoggan 1978).
Gilstra, Z. 1991. Proposal to conserve the spelling
Ecology.—Strawberry bush grows well in many of 4618 Euonymus (Celastraceae). Taxon 40:
soils types, up to 1 m in a growing season. It is 137-139.
usually found in forests and thickets, including
lowland areas and edges of streams. It occurs from Gleason, H.A., and A. Cronquist 1991. Manual of
sea level up to 400 m in elevation (Ma 2001). vascular plants of northeastern United States and
adjacent Canada. 2nd Ed. New York Botanical
Reproduction.—Hymenopterous insects, Garden, Bronx, NY. 910 p.
especially bees, appear to be the main pollinators,
although wind may also be involved (Brizicky Ma, J.S. 2001. A Revision of Euonymus
1964). The seeds of strawberry bush are dispersed (Celastraceae). Thaiszia, Journal of Botany 11:
by birds and other small mammals (Brizicky 1-264.
1964). Strawberry bush blooms in late spring and
early summer, and the fruits mature in autumn. Paclt, J. 1998. Proposal to amend the gender of
Euonymus, nom. cons. (Celastraceae) to
Growth and Management.—Strawberry bush feminine. Taxon 47: 473-474.
prefers a humus-rich, slightly acidic soil. It does
well in shady situations, tolerating even full shade. Scoggan, H.J. 1978. The Flora of Canada Part 3 –
It can be planted in USDA Zones 5 to 9. Dicotyledoneae (Saururaceae to Violaceae).
Propagation can be effected through stem cuttings, National Museum of Natural Sciences, National
layering and seed (Chittenden and Synge 1956) Museums of Canada, Ottawa. 568 p.
Seeds require approximately 3 months of
prechilling and stratification. Shetler, S.G. and S.S. Orli. 2000. Annotated
checklist of the vascular plants of the
Benefits.—Strawberry bush is an excellent Washington-Baltimore area. Part. 1. Ferns, fern
ornamental species. The leaves turn a dark red in allies, gymnosperms, dicotyledons. Smithsonian
autumn. The red capsules and scarlet arillate seeds Institution, Washington, D.C. 186 p.
add to the “burning effect” and contributes to some
of its common names. Strawberry bush is best USDA, NRCS. 2003. The PLANTS Database.
used in naturalistic settings, in the shade of larger Baton Rouge, LA. http:plants.usda.gov. [not
shrubs and trees. Large specimens can have paged].
334
Euonymus atropurpureus Jacq. eastern wahoo
CELASTRACEAE
335
to 16 weeks cold stratification prior to sowing. The adjacent Canada. 2nd Ed. New York Botanical
plant can also be propagated with cuttings. Garden, Bronx, NY. 910 p.
Benefits.—The common name “burning bush” Ma, J.S. 2001. A Revision of Euonymus
comes from the plant’s bright red fall foliage. The (Celastraceae). Thaiszia, Journal of Botany 11:
red fruit and arillate seeds add to the “burning” 1-264.
effect. This makes the eastern wahoo an excellent
shrub for horticultural use (Rehder 1940). The Paclt, J. 1998. Proposal to amend the gender of
seeds and the vegetative organs of eastern wahoo Euonymus, nom. cons. (Celastraceae) to
are said to be poisonous (Brizicky 1964). feminine. Taxon 47: 473-474.
Gleason, H.A. and A. Cronquist 1991. Manual of USDA, NRCS. 2003. The PLANTS Database.
vascular plants of northeastern United States and Baton Rouge, LA. http:plants.usda.gov. [not
paged].
336
Fallugia paradoxa (D. Don) Endl. ex Torr. Apache-plume
ROSACEAE
Juanita A. R. Ladyman
end of each stalk during the fruiting period. The
length of the flowering period varies depending
upon geographic location and elevation. It flowers
in June through August in all parts of its range but
flowering is extended from May to December in
Texas (Powell 1998) and from April to October in
Arizona and New Mexico (Epple 1995, Martin and
Hutchins 1980). The common name for Fallugia
paradoxa, Apache-plume, was coined because the
feathery seed clusters look like Indian feather
headdresses (Elmore 1976). Other common names
include poñil, fallugie (Dayton 1931), feather rose,
and feather duster bush (Epple 1995). Fallugia is a
monotypic genus, that is, a genus of a single
species (Cronquist and others 1997). In addition,
Apache-plume shows minimal genetic variation
throughout its wide range (Jones and Johnson,
1998). Hybrids between Apache-plume and
cliffrose (Purshia mexicana (D. Don) Henrickson)
have been reported (Blauer and others 1975,
McArthur and others 1977).
337
moisture than the surrounding area. Apache-plume year, environmental conditions, and availability of
is relatively drought tolerant and requires 20 to 50 alternate browse appears to influence palatability
cm of annual precipitation (Hayward 1990). It is and use. It provides cover for a variety of small
tolerant of weakly saline and neutral to moderately mammals and birds (Haywood 1990, McWilliams
basic soils (McWilliams 2000). Endomycorrhizae 2000). It was used for arrow shafts and brooms by
were associated with Apache-plume in New many Native American peoples (Dunmire and
Mexico (Williams and Aldon 1976). Tierney 1995, Moerman 1998). Apache-plume
brooms are specifically kept inside some Sandia
Reproduction.—Apache-plume reproduces Pueblo houses because there is a belief that the
sexually by seed and vegetatively by sprouts brooms exert a positive spiritual effect on the
(suckers) from the roots. The feather-tailed seeds household (Dunmire and Tierney 1995). The long
are probably primarily wind dispersed. Apache- roots were used as cord to tie fencing and make
plume seeds apparently have no dormant period ramadas (Dunmire and Tierney 1995, Moerman
(Baskin and Baskin 2001). They are small and 1998). An infusion of leaves was used as a
average approximately 925,932 per kg (Vines shampoo and hair growth stimulant by some
1986). Native American tribes, and petals are reported to
prevent stomach gas (Dunmire and Tierney 1995,
Growth and Management.—Seed can be sown Moerman 1998). One tribe used it in witchcraft to
by broadcasting it on a prepared bed and covered cause “insanity” (Moerman 1998). Shrubs are
by approximately 5 mm of fine loam or sand. attractive as ornamentals and are popularly used
Germination rate is from 19 to 65 percent (Vines for landscaping (Hayward 1990, Phillips 1987,
1986). In pots, the germination rate is 30 to 40 Morrow 1995).
percent, but seedlings are reported to damp off
easily and require adequate air circulation References
(Hayward 1990). A 4-litre plant is fully established
and ready for sale in 18 months for the landscape Aro, R.S. 1971. Evaluation of pinyon-juniper
market (Hayward 1990). Plants can also be grown conversion to grassland. Journal of Range
fairly easily from root cuttings. It is closely grazed Management. 24(3): 188-197
on overstocked range and, although tolerant of
grazing with “excellent recuperative powers,” it is Baldwin B.G., S. Boyd, B.J. Ertter, R.W.
also reported to decrease under grazing pressure Patterson, T.J. Rosatti, and D.H. Wilken. 2002.
(Dayton 1931, McGregor and others 1986). The Jepson Desert Manual: Vascular plants of
Availability of adequate moisture may be an southeastern California. University of California
important factor in the degree of recovery. Press, Berkeley, CA. 624 p.
Apache-plume sprouts after fire (Shaw and
Monson 1983, McWilliams 2000). In a pinyon Baskin, C.C. and J.M. Baskin. 2001. Seeds.
juniper community it was reported to tolerate fire Ecology, biogeography, and evolution of
well (Aro 1971). dormancy and germination. Academic Press,
New York, NY. 666 p.
Benefits.—Apache-plume is valuable for erosion
control (Dayton 1931, Stubbendieck and others Blauer, A. C., A.P. Plummer, and E.D. McArthur.
1993, Vines 1986). Its long roots provide effective 1975. Characteristics and hybridization of
soil stabilization especially in arid situations important Intermountain shrubs. I. Rose family.
(McWilliams 2000). Apache-plume is important Research Paper INT-169. U.S. Department of
winter range for livestock (Dayton 1931, Agriculture, Forest Service, Intermountain
McGregor and others 1986, Stubbendieck and Forest and Range Experiment Station, Ogden,
others 1993, USDAFS 1988). It is reported to have UT. 36 p.
only a fair palatability for sheep, goats and cattle
but closely cropped, stunted plants have been cited Cronquist, A., N.H. Holmgren, and P.K.
as evidence of high palatability (Dayton 1931, Holmgren. 1997. Intermountain Flora – vascular
McGregor and others 1986, Stubbendieck and plants of the Intermountain west, U.S.A. Vol. 3
others 1993, Warnock 1974). It also furnishes part A. Subclass Rosidae. The New York
important browse for some species of wildlife Botanical Garden, Bronx, NY. 446 p.
(Dayton 1931, Stubbendieck and others 1993) and
makes up 1 percent to 5 percent of desert mule Dayton, W.A. 1931. Important western browse
deer diet (Krausman and others 1997). Time of plants. Miscellaneous Publication 101. U.S.
338
Department of Agriculture, Washington, DC. Moerman, D.E. 1998. Native American ethno-
214 p. botany. Timber Press, Portland, OR. 927 p.
Dick-Peddie, W.A. 1993. New Mexico Vegetation Morrow, B.H. 1995. Best Plants for New Mexico
– past, present, and future. University of New Gardens and Landscapes. University of New
Mexico Press, Albuquerque, NM. 244 p. Mexico Press, Albuquerque, NM. 267 p.
Dunmire, W.W. and G.D. Tierney. 1995. Wild Phillips, J. 1987. Southwestern landscaping with
plants of the Pueblo Province. Museum of New native plants. Museum of New Mexico Press,
Mexico Press, Santa Fe, NM. 290 p. Santa Fe, NM. 141 p.
Elmore. F.H. 1987. Shrubs and Trees of the Powell, A.M. 1998. Trees and shrubs of Trans-
Southwest Uplands. Southwest Parks and Pecos and adjacent areas. University of Texas
Monuments Assoc. Tucson, AZ. 214 p. University Press, Austin, TX . 498 p.
Epple, A.O. 1995. A field guide to the plants of Shaw, N.L. and S.B. Monson. 1983. Phenology
Arizona. Falcon Press Publishing Co., Helena, and growth habits of nine antelope bitterbrush,
MT. 347 p. desert bitterbrush, stansbury cliffrose, and
apache-plume accessions. In: A.R. Tiedemann
Hayward, P. 1990. Fallugia paradoxa. American and K.L. Johnson, comps. Proceedings:
Nurseryman. 172(7): 126 Research and Management of bitterbrush and
cliffrose in Western North America: 1982, April
Jones, T.A. and D A. Johnson. 1998. Integrating 13-15, Salt Lake City, UT. General technical
Genetic Concepts into Planning Rangeland Report: INT-152, USDA Forest Service,
Seedings. Journal of Range Management Intermountain Forest and Range Station, Ogden,
51(6): 594-606. UT.
Krausman, P.R., A.J. Kuenzi, R.C. Etchberger, Stubbendieck, J., S.L. Hatch and C.H. Butterfield.
K.R. Rautenstrauch, L.L. Ordway, and J.J. 1993. North American range plants. 4th edition.
Hervert. 1997. Diets of mule deer. Journal of University of Nebraska Press, Lincoln, NE.
Range Management. 50(5): 513-522. 493 p.
Martin, W.C. and C.R. Hutchins. 1980. A Flora of Tidestrom, I. and T. Kittell. 1941. A flora of
New Mexico. Strauss & Cramer, Hirschberg, Arizona and New Mexico. The Catholic
Germany. 2,592 p. University of America Press, Washington, DC.
897 p.
McArthur, E.D., B.C. Giunta, and A.P. Plummer.
1977. Shrubs for restoration of depleted range USDA Forest Service. 1988. Range Plant
and disturbed areas. Utah Science. 35: 28-33. Handbook. Dover Publications, Inc., New York,
NY. 842 p.
McGregor, R.L. and the Great Plains Flora
Association. 1986. Flora of the Great Plains. USDA PLANTS database. 2002. Internet site:
University Press of Kansas, Lawrence, KS. http:// plants.usda.gov/cgi_bin/plant_profile.
1,402 p. cgi? symbol=FAPA#. [not paged].
McWilliams, J. 2000. Fallugia paradoxa. In: U.S. Vines, R.A. 1986. Trees, shrubs, and woody vines
Department of Agriculture, Forest Service, of the Southwest. Sixth printing. University of
Rocky Mountain Research Station, Fire Sciences Texas Press. Austin, TX. 1,104 p.
Laboratory. Fire Effects Information System.
http://www.fs.fed.us/database/feis/ 19 p. Warnock, B.H. 1974. Wildflowers of the
Guadalupe Mountains and the sand dune
Meyer, S. 1974. Fallugia paradoxa (D. Don) Endl. country, Texas. Sul Ross university, Alpine, TX.
ex Torr. Apache-plume. In: . F.T. Bonner and 176 p.
R.G. Nisley, eds. USDA Forest Service Woody
Plant Seed Manual Internet site: http:// wpsm.net Williams, S.E. and E.F. Aldon. 1976.
4 p. Endomycorrhizal (vesicular arbuscular)
339
associations of some arid zone shrubs. The
Southwestern Naturalist. 20(4): 437-444.
340
Faramea occidentalis (L.) A. Rich. cafeíllo
RUBIACEAE
John K. Francis
341
TDW = 30.045D2L (adjusted R-squared = 0.962), Holdridge, L.R. 1967. Life zone ecology. Tropical
where TDW is total dry weight in grams, D is Science Center, San José, Costa Rica. 206 p.
diameter in cm at 30 cm above the ground, and L
is stem length in meters. Lower stem density of Howard, R.A. 1989. Flora of the Lesser Antilles,
Puerto Rican shrubs averaged 0.62 + 0.04 g/cm3 Leeward and Windward Islands. Vol. 6. Arnold
(Francis 2000). Stem density of saplings from Arboretum, Harvard University, Jamaica Plain,
Panama was 0.57 g/cm3 (King 1990). Allometric MA. 658 p.
relationships for cafeíllo in Panama may be found
in O’Brien and others (1995) and King (1990). As King, D.A. 1990. Allometry of saplings and
an understory tree, cafeíllo is slow-growing and understory trees of a Panamanian forest.
long lived. Based on recent growth rates, the Ecology 4(1): 27-32.
largest individuals in a Panamanian stand were
estimated to exceed 200 years in age. Liogier, H.A. 1990. Plantas medicinales de Puerto
There is no record of plantations or Rico y del Caribe. Iberoamericana de Ediciones,
planted ornamental cafeíllo. The species is able to Inc., San Juan, PR. 566 p.
reproduce by sprouting after injury. Thirty-six
percent of the individuals in a Panamanian stand Liogier, H.A. 1997. Descriptive flora of Puerto
were of sprout origin (Putz and Brokaw 1989). Rico and adjacent islands. Vol. 5. Editorial de la
Universidad de Puerto Rico, San Juan, PR.
Benefits.—Cafeíllo wood is heavy, hard, and 436 p.
tough. However, uses are mainly limited to fence
posts, vegetable stakes, tool handles, and Little, E.L., Jr. and F.H. Wadsworth. 1964.
fuelwood. In herbal medicine, cafeíllo is Common trees of Puerto Rico and the Virgin
reportedly used as an antiseptic, astringent, and for Islands. Agriculture Handbook 249. U.S.
diarrhea and anemia (Liogier 1990). Birds Department of Agriculture, Forest Service,
consume the fruits. The use of cafeíllo as an Washington, DC. 548 p.
ornamental or in forest restoration efforts has not
been reported. Cafeíllo is a fine looking plant and O’Brien, S.T., S.P. Hubbell, P. Spiro, R. Condit,
could probably be used as a back-ground plant or and R.B. Foster. 1995. Diameter, height, crown,
screen under shade in yards, estates, and parks. and age relationships in eight neotropical tree
species. Ecology 76(6): 1926-1939.
References
Putz, F.E. and N.V.L. Brokaw. 1989. Sprouting of
Francis, J.K. 2000. Estimating biomass and carbon broken trees on Barro Colorado Island, Panama.
content of saplings in Puerto Rican secondary Ecology 70(2): 508-512.
forests. Caribbean Journal of Science 36(3-4):
346-350. Schupp, E.W. 1990. Annual variation in seedfall,
postdispersal predation, and recruitment of a
Francis, J.K. and A. Rodríguez. 1993. Seeds of neotropical tree. Ecology 71(2): 504-515.
Puerto Rican trees and shrubs: second
installment. Research Note SO-374. U.S. Travers, S.E. 1998. The effect of rust infection on
Department of Agriculture, Forest Service, reproduction in a tropical tree (Faramea
Southern Forest Experiment Station, New occidentalis). Biotropica 30(3): 438-443.
Orleans, LA. 5 p.
342
Forestiera acuminata (Michx.) Poir. swamp privet
OLEACEAE
Kristina Connor
Drawing credit: USDA—Forest Service collection, Hunt Reproduction.—Swamp privet plants are
Institute monoecious; flowers emerge in the spring before
the leaves unfold. They are yellow, fragrant and
General Description.—Swamp privet, also called small, the males sessile or almost so, female
eastern swamp privet, is a deciduous shrub or flowers in small panicles or fascicles (Bailey and
small, open-crowned tree, occasionally reaching Bailey 1976, Botanical Institute 2002, Johnson and
10 m in height but more often averaging 1.5 to 2.5 Hoagland 1999, Krüssmann 1986). They have no
m (Krüssmann 1986, Johnson and Hoagland petals and minute or absent sepals. Male flowers
1999). It is commonly multitrunked (Mackay and have one to four stamens, female flowers have
Finical 1999, U.S. Army Corps of Engineers simple or two-lobed stigmas (Johnson and
1997). The bark ranges from gray to dark brown in Hoagland 1999, Botanical Institute 2002, USGS
color and is either smooth or ridged (Johnson and 2002). Fruits are dark purple or black oblong
Hoagland 1999, USGS 2002). Twigs are light drupes, 8 to 12 mm long with one, rarely two,
brown, glabrous, and have conspicuous lenticels seeds. They appear in summer and are promptly
(Johnson and Hoagland 1999). Leaves are simple, shed (Johnson and Hoagland 1999). Although no
opposite, and dull, yellowish green. They range in data are provided to support this, the Plants for a
size from 3 to 11 cm in length, and 2.5 to 5 cm in Future website (2001) suggests that seeds be sown
width. Leaves have long, pointed tips (acuminate) as soon as they ripen and that cuttings can be taken
and petioles that are slender and slightly winged at of half-ripe wood in summer or mature wood in
the base. While leaf margins are usually smooth, it winter.
is not uncommon for the apex half to be finely
serrated. Growth and Management.—Swamp privet,
while adaptable to many soil types, must be
Range.—Swamp privet is found in the wet located in moist areas. It is moderately tolerant of
woodlands and swamps of the Southeastern United heat and can tolerate full or partial sunlight
States, from South Carolina south to Florida and (Mackay and Finical 1999). It has been reported in
westward into Texas, Oklahoma, and Kansas. It cultivation (Krüssmann 1986).
ranges as far north as Illinois, Indiana, and
Missouri (Krüssmann 1986, USGS 2002). Benefits.—The fruits of swamp privet are
considered good food for wildlife (Johnson and
343
Hoagland 1999). A decoction of the roots and bark Mackay, W. and L. Finical. 1999. A special
has been used as a health beverage, and the wood project of the Texas Agricultural Experiment
has been used for turning (Bailey and Bailey Station. Benny Simpson’s Texas Native Trees.
1976), although reports on its durability vary Texas A&M University, College Station.
(Plants for a Future 2001). http://aggie-horticulture.tamu.edu/natives. 1 p.
344
Forestiera segregata (Jacq.) Krug & Urban Florida privet
OLEACEAE
John K. Francis
General Description.—Florida privet, also known Reproduction.—In Florida, Florida privet flowers
as Florida swamp privet, wild olive, and ink-bush, in the spring (Nelson 1996). It is insect pollinated.
is a semideciduous shrub or small tree to 7 m in A collection of fruits from Puerto Rico weighed an
height. The plant is supported by a weak taproot average of 0.1209 + 0.0035 g/fruit. Air-dried seeds
and more robust, tan-colored lateral roots. The cleaned from them weighed an average of 0.0359 +
branches are slender. The opposite leaves have 0.0006 g/seed or 27,900 seeds/kg. Sown in moist
short petioles, 1 to 6 mm long, and narrowly potting mix, they began germinating in 7 days and
elliptic to obovate blades, 1 to 7 cm long, with completed 89 percent germination in 49 days
entire margins. Tiny, greenish-yellow flowers are (author’s observation). The seeds are dispersed by
borne in clusters at the leaf axils. The fruits are birds that eat the fruits. Seedlings do not occur in
slightly curved, ovoid, black drupes, 6 to 10 mm large numbers.
long, that have a bitter, unpleasant flavor and stain
cloth and skin. Each contains one greenish-yellow Growth and Management.—Florida privet grows
seed (Howard 1989, Liogier 1995, Long and slowly in the early seedling stage but later
Lakela 1976, Nelson 1996). accelerates so that seedlings are ready to transplant
at the end of 1 year. Nursery production is
Range.—Florida privet is native to Florida, normally from seed. Wildlings of all sizes can be
Georgia, South Carolina, the Bahamas, the Greater
345
successfully transplanted (Workman 1980). Howard, R.A. 1989. Flora of the Lesser Antilles,
Beyond the seedling stage, it has a moderate Leeward and Windward Islands.
growth rate (University of Florida Cooperative Dicotyledoneae. Vol. 6. Arnold Arboretum,
Extension Service 2002). The largest Florida privet Harvard University, Jamaica Plain, MA. 658 p.
recorded measures 20.2 cm diameter at breast
height and 5.5 m in height (American Forests Liogier, H.A. 1995. Descriptive flora of Puerto
2002). Rico and adjacent islands. Vol. 4. Editorial de la
Universidad de Puerto Rico, San Juan, PR.
Benefits.—Florida privet is recommended for 617 p.
xeriscape (unwatered) landscaping (South Florida
Water Management District 2001) and is available Long, R.W. and O. Lakela. 1976. A flora of
through a number of commercial nurseries. It is tropical Florida. Banyon Books, Miami, FL.
useful for hedges and foundation and specimen 962 p.
plantings. Warblers, bluebirds, mockingbirds, and
vireos feed upon the fruits in South Florida National Trust for the Cayman Islands. 2002.
(Broschat and Verkade 2002, Cowley 2002). Virtual herberium: Oleaceae, Forestiera
Honeybees, butterflies, and other insects are segregata (Jacq.) Krug & Urban. http://www.
attracted to the flowers for the nectar (Workman virtualherbarium.org/vh/CAYM/1002.html. 2 p.
1980).
Natural Resources Conservation Service. 2002.
References Plants profile: Forestiera Poir. swampprivet.
http://plants.usda.gov/cgi_bin/plant_profile.cgi?
American Forests. 2002. National register of big symbol=FORES. 3 p.
trees: Florida-privet, Forestiera segregata.
http://www.americanforests.org/resources/bigtre Nelson, G. 1996. The shrubs and woody vines of
es/register.php?details=2000. 1 p. Florida. Pineapple Press, Inc., Sarasota, FL.
391. p.
Broschat, T.K. and S.V. Verkade. 2002.
Landscaping to attract birds in South Florida. South Florida Water Management District. 2001.
http://www.floridaplants.com/landscape/birds. Quick facts on xeriscape: how to save water
htm. 3 p. through water-smart landscaping. http://www.
sfwmd.gov/images/pdfs/splash/splxeris.pdf. 2 p.
Cowley, M. 2002. Florida native plants: olive
family. Native Florida Web Ring. University of Florida Cooperative Extension
http://nsis.org/garden/family/olive.html. 2 p. Service. 2002. Native shrubs of South Florida.
http://edis.ifas.ufl.edu/BODY_EH159. 9 p.
Everglades Mitigation Bank. 2002. Endangered
and threatened plants at the EMB. Workman, R.W. 1980. Growing native. The
http://www.fpl.com/environment/emb/contents/e Sanibel-Captiva Conservation Foundation, Inc.,
ndangered_plants.shtml. 1 p. Sanibel, FL. 137 p.
346
Fouquieria splendens Engelm. ocotillo
FOUQUIERIACEAE
Synonyms: none
347
and Young 1986). It is a slow-growing species, cultivated genotypes originally adapted to another
and estimates of its life span range from less than region. This may pose a threat although, because
60 to greater than 72 years (Mathews 1994). It is of the slow growth and low turnover of
likely that life span is related to latitude and individuals, the effects would be a long time in
substrate. The leaves are winter- and drought- becoming apparent. For example, the cultivated
deciduous (Schmutz and others 1992). The leaves genotypes may have particularly high fecundity
appear after rains and are a reliable indicator of and may be physiologically robust but do not have
recent rainfall (Powell 1998). They tend to mature the genetic composition to be resistant to
and fall within a few weeks indicating that environmental or biological (e.g. disease or insect
available photosynthate is limited and that growth infestation) events that are periodic (perhaps
is dependent upon precipitation. Ocotillo is a decades apart) in the local area. Hedges or fences
“water storer” in that the stems store relatively of living stems are common, especially in Mexico.
large amounts of water that buffer against drought It does not appear to be grown for its medicinal
stress (Burgess 1995). Wildfires may be value. Historically the Mahuna Native Americans
detrimental to ocotillo populations. It rarely used it as a “blood medicine” (Moerman 1998).
sprouts from the root crown following damage The Apache Indians used powdered roots to treat
from fire, and its sprouting ability is probably wounds and painful swellings and they also bathed
dependent on fire severity (Ahlstrand 1982, in an ocotillo root mixture to relieve fatigue
McPherson 1995, White 1969). Mortality rates of (Krochmal and others 1954, Powell 1998, Vines
40 percent to 50 percent were reported after light 1960). A beverage made from ocotillo flowers was
to moderate fire in Arizona (Thomas 1991). used for cough medicine (Vines 1960). The
Seedlings are not known to establish in recently Cahuilla, Papago and Yavapai tribes used the
burned areas (Mathews 1994), and its seeds are flowers in beverages, the fruits for a “candy,” and
unlikely to survive for long in the soil (Zedler the stalks as building material (Moerman 1998).
1981). In fact, burning has been suggested as a The thorns were used for ear piercing by the
control method for woody species, such as Papago. The resin and wax from ocotillo bark was
Fouquieria splendens, in desert grasslands used for conditioning leather (Krochmal and others
(Mathews 1994). 1954, Powell 1988, Vines 1960). It seems that all
uses are localized, personal, or historic in the
Benefits.—The endangered Lucifer humming- Southwestern United States. It is not known if
bird's habitat in New Mexico centers on slopes and ocotillo is more widely used today in Mexico.
adjacent canyons in arid montane areas dominated
by ocotillo and agave species (New Mexico References
Department of Game and Fish 1991). Ocotillo has
no recorded forage value, presumably because its Ahlstrand, G.M. 1982. Response of Chihuahuan
leaves are so short-lived (Dayton 1931). Ocotillo Desert Mountain Shrub Vegetation to Burning.
plants are popularly used for landscaping. Plants Journal of Range Management 35 (1): 62-65.
sold in reputable nurseries are obtained from
private land under permit. There are typically two Burgess, T.L. 1995. The dilemma of coexisting
“types” of product. Smaller plants are sold in 1 growth forms. In: The desert grassland. M.P.
gallon pots or larger specimens are sold “balled McClaran and T.R. Van Devender, eds. The
and burlapped.” Although the plants transplant University of Arizona Press, Tucson, AZ. p. 31-
well if there are sufficient roots attached, many 67.
that are dug are not treated well and two thirds or
more of those dug up are likely not to survive. Dayton, W.A. 1931. Important western browse
Propagation has been reported using cuttings plants. Miscellaneous Publication 101. U.S.
although conversations with horticulturalists Department of Agriculture, Washington, DC.
indicate that this may be less easy than the 214 p.
literature indicates. The plant can also be grown
from seed but is a relatively slow grower. Seeds Epple, A.O. 1995. A field guide to the plants of
apparently collected from Mexico can be Arizona. Falcon Press Publishing Co., Helena,
purchased in the United States (Tradewinds MT. 347 p.
Import-Export Company 2002). This trade
between regions where the plant is native does Flores-Mata, G.F. J.J. Lopez, X.M. Sánchez, F.M.
raise the potential for genetic modification, or Ruiz, and F.T. Takaki. 1971. Memoria del Mapa
“dilution,” of locally adapted genotypes by Tipos de Vegetación de la República Mexicana.
348
Secretaria de Recursos Hidráulicos, Dirección Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L.
de Agrología, Mexico, DF. 59 p. Cox, J.O. Klemmedson, J.J. Norris, and L.C.
Fierro. 1992. Desert grassland. In: Natural
Krochmal, A., S. Paur, and P. Duisberg. 1954. grasslands: Introduction and Western
Useful native plants in the American Hemisphere. R.T. Coupland, ed. Elsevier, New
Southwestern deserts. Economic Botany. 8: 3- York. p. 337-362.
20.
Thomas, P.A. 1991. Response of succulents to
Martin, W.C. and C.R. Hutchins. 1980. A Flora of fire: A review. International Journal of Wildland
New Mexico. Strauss & Cramer, Hirschberg, Fire 1(1): 11-22.
Germany. 2,592 p.
Tradewinds Import-Export Company. 2002. Seeds
Mason, C.T. and P.B. Mason. 1987. A handbook Department – USA. P.O. Box 218650, Houston,
of Mexican roadside flora. The University of Texas 77218 USA. http://www.abonline.
Arizona Press, Tucson, AZ. 380 p. com/products/seeds_f.htm#f. [not paged].
Matthews, R.F. 1994. Fouquieria splendens. The Van Devender, T.R. 1995. Desert grassland
Fire Effects Information System Data base. U.S. history: changing climates, evolution,
Department of Agriculture, Forest Service, biogeography, and community dynamics. In:
Intermountain Research Station, Intermountain The desert grassland. M.P. McClaran and T.R.
Fire Sciences Laboratory, Missoula, MT. http:// Van Devender, eds. The University of Arizona
www.fs.fed.us/database/feis/plants/shrub/fouspl/ Press, Tucson, AZ. p. 68-99.
all.html. 15 p.
Vines, R.A. 1986. Trees, shrubs, and woody vines
McAuliffe, J.R. 1988. Markovian dynamics of of the Southwest. Sixth printing. University of
simple and complex desert plant communities. Texas Press. Austin, TX. 1,104 p.
American Naturalist. 131(4): 459-490.
Waser, N.M. 1979. Pollinator availability as a
McPherson, G.R. 1995. The role of fire in desert determinant of flowering time in ocotillo
grasslands. In: The desert grassland. M.P. (Fouquieria splendens). Oecologia. 39(1): 107-
McClaran and T.R. Van Devender, eds. The 121.
University of Arizona Press, Tucson, AZ. p.
130-151. White, L.D. 1969. Effects of a wildfire on several
desert grassland shrub species. Journal of Range
Moerman, D.E. 1998. Native American ethno- Management 22: 284-285.
botany. Timber Press, Portland, OR. 927 p.
Yeaton, R.I., J. Travis, and E. Gilinsky. 1977.
New Mexico Department of Game and Fish. 1991. Competition and spacing in plant communities: the
Handbook of species endangered in New Arizona upland association. Journal of Ecology. 65:
Mexico. Dept. of Game and Fish., Santa Fe, 587-595
NM. 185 p.
Young, J.A. and C.G.Young. 1986. Collecting,
Powell, A.M. 1988. Trees & shrubs of Trans- processing, and germinating seeds of wildland
Pecos Texas including Big Bend and Guadalupe plants. Timber Press. Portland, OR. 236 p.
Mountains National Parks. Big Bend Natural
History Association, Big Bend National Park, Zedler, P.H. 1981. Vegetation change in chaparral
TX. 536 p. and desert communities in San Diego County,
California. In: D.C. West, H.H. Shugart, D. B.
Powell, A.M. 1998. Trees & shrubs of Trans- Botkin, eds. Forest succession: Concepts and
Pecos and adjacent areas. University of Texas application. Springer-Verlag, New York. p.
University Press, Austin, TX . 498 p. 406-430
349
Furcraea foetida (L.) Haw. Mauritius hemp
AGAVACEAE
John K. Francis
Range.—Mauritius hemp is native to the Greater
Antilles, and from Guadaloupe south through
northern South America to Brazil (Grisebach
1963, Howard 1979). The species has been widely
planted. It has naturalized in Florida, Hawaii,
Marquesas Islands, French Polynesia, Tonga, and
is present and probably naturalized in many other
places (Pacific Island Ecosystems at Risk 2003).
350
crassulacean acid metabolism by which it fixes although not heavily, used as a landscaping plant
CO2 during the night and incorporates it into for accent and curiosity. A variegated form is
carbohydrates during the day (personal available (Desert-Tropicals Nursery 2003).
communication with Ernesto Medina, plant Mauritius hemp was once widely cultivated for
physiologist, Centro de Ecología, Instituto de fiber, hence the common name. Extracts of the
Investigaciones, Caracas, Venezuela). roots are used as ingredients in tonics for purifying
the blood, and the dried leaves are used to control
Reproduction.—Flowering, which may occur at swelling and to aid in wound healing (Núñez-
any time of the year, apparently begins when Meléndez 1982).
plants attain sufficient size and vigor to support the
large flower stalk. Plants die about 1 year after the References
onset of flowering. The flowers are visited by
honey bees (Apis mellifera L.). There is no record Bailey, L.H. 1941. The standard cyclopedia of
in the literature of seed weights. Fresh bulbils (n = horticulture. Vol. 2. The MacMillan Company,
100) collected by the author in Puerto Rico New York. p. 1,201-2,422.
weighed an average of 2.72 + 0.31 g/bulbil with a
range of 0.18 to 16.27 g. Placed in moist potting Desert-Tropicals Nursery. 2003. Mauritius hemp,
mix, 100 percent of them rooted within 1 week. sisal, maguey. http://www.desert-tropicals.com/
Bulbils are formed by the thousands on single Plants/Agavaceae/Furcraea_foetida.html. 2 p.
plants and can form dense thickets. Gravity is the
only known natural means of dispersal. The Grisebach, A.H.R. 1963. Flora of the British West
presence of scattered individuals indicates other Indian Islands. J. Cramer, Weinheim, Germany.
transport, perhaps fruit bats. 789 p.
Growth and Management.—The best way to Howard, R.A. 1979. Flora of the Lesser Antilles,
artificially propagate Mauritius hemp is to place Leeward and Windward Islands. Vol. 3. Arnold
bulbils in pots of potting mix. A group of bulbils Arboretum, Harvard University, Jamaica Plain,
planted in Puerto Rico grow rapidly with no MA. 586 p.
mortality, reached an average of 17.8 + 0.4 cm in
height in 1 month, and were ready to outplant in 1 Liogier, H.A. and L.F. Martorell. 2000. Flora of
to 2 months. New plants need protection from Puerto Rico and adjacent islands: a systematic
weeds for a few months, but afterwards the plants synopsis. 2nd ed. Editorial de la Universidad de
need little care. Individual plants probably live Puerto Rico, Río Piedras, PR. 382 p.
from 5 to 20 years depending on growing
conditions. One owner reported that an ornamental Núñez-Meléndez, E. 1982. Plantas medicinales de
plant died after flowering at 15 years old (author’s Puerto Rico. Editorial de la Universidad de
observation). Mauritius hemp is rarely common Puerto Rico, Río Piedras, PR. 498 p.
enough in its native range to need control. When
control is needed, particularly in invaded areas, Pacific Island Ecosystems at Risk. 2003. Furcraea
individual plants can be killed by grubbing them foetida (L.) Haw., Agavaceae. http;//www.hear.
out or by spray or drizzle application of 2,4-D or org/pier_v3.3/fufoe.htm. 3 p.
triclopyr in oil (Pacific Island Ecosystems at Risk
2003). Schlegel, R. 2003. Plant breeding update: crop
plants II. http://www.desicca.de/plant_breeding/
Benefits.—Mauritius hemp helps hold the soil, Crop_plants/Crops_II/hauptteil_crops_ii.html.
furnishes cover for wildlife, and adds to the [not paged.].
aesthetics of wildlands. The species is widely,
351
Gesneria pedunculosa (DC.) Fritsch arbol de navidad
GESNERIACEAE
John K. Francis
352
wind. Seedlings are relatively common on bare References
ground near fruiting adult plants. Adult and sapling
plants sprout when cut or damaged. American Gloxinia and Gesneriad Society. 2002.
Registered gesneriads. American Gloxinia and
Growth and Management.—Arbol de navidad is Gesneriad Society, Salsbury, CT. http://
slow growing at all stages. It probably lives several www.aggs.org/ir_ges/. 7 p.
decades. No plantations are known, and
recommended management practices have not been Liogier, H.A. 1995. Descriptive flora of Puerto
published. Disturbance of the soil surface in steep, Rico and adjacent islands. Vol. 4. Editorial de la
rocky terrain in moist areas near seed sources Universidad de Puerto Rico, San Juan, PR.
would likely result in seedling establishment. 617 p.
Benefits.—Arbol de navidad is a pretty plant that Little, E.L., Jr., R.O. Woodbury, and F.H.
adds to the aesthetics of the forest. It helps protect Wadsworth. 1974. Trees of Puerto Rico and the
the soil in fragile areas and furnishes cover for Virgin Islands. Vol. 2. Agriculture Handbook
wildlife. Because of the small diameters and 449. U.S. Department of Agriculture,
quantities in which it is available, the wood, which Washington, DC. 1,024 p.
is light brown and hard, is useful mainly for fuel.
353
Gonzalagunia hirsuta (Jacq.) K. Schum. rabo de ratón
RUBIACEAE
John K. Francis
areas in advanced secondary forests, and along
roads. In a Puerto Rican study (Myster and Walker
1997), rabo de ratón was the fifth most common
species colonizing recent landslides. Dense
thickets sometimes form. The species grows in
areas receiving from about 1100 to 3400 mm of
annual rainfall and at elevations ranging from near
sea level to more than 600 m. Apparently, all types
of soils are colonized, if not excessively well
drained or very poorly drained. The root system is
shallow, much branched, and extensive.
354
predation (Myster 1997). The seeds are dispersed Howard, R.A. 1989. Flora of the Lesser Antilles,
by birds (Parrotta 1995). Leeward and Windward Islands. Dicotyledoneae.
Part 3. Vol. 6. Arnold Arboretum, Harvard
Growth and Management.—Rabo de ratón is University, Jamaica Plain, MA. 658 p.
fast growing and may reach 4 m in height and 3
cm in lower stem diameter. Individual plants Liogier, H.A. 1997. Descriptive flora of Puerto
usually live 1 to 2 years (Vélez and von Overbeek Rico and adjacent islands. Vol. 5. Editorial de la
1950). The species can probably be established by Universidad de Puerto Rico, San Juan, PR.
traditional methods (planted seedlings and direct 436 p.
seeding), but no plantations have been reported.
Natural reproduction may be encouraged by Myster, R.W. 1997. Seed predation, disease and
cultivation near seed sources. germination on landslides in neotropical lower
montane wet forest. Journal of Vegetation
Benefits.—Rabo de rató is beneficial to the Science 8: 55-64.
environment in that it readily invades and helps to
stabilize disturbed areas. Because the shrub forms Myster, R.W., and L.R. Walker. 1997. Plant
a thin canopy, it does not inhibit the invasion of successional pathways on Puerto Rican
later successional vegetation. While probably not landslides. Journal of Tropical Ecology 13: 165-
suitable for formal gardens, rabo de rató would 173.
contribute to wildflower gardens, forest parks, and
greenbelts. The species is an abundant producer of Parrotta, J.A. 1995. Influence of overstory
fruits that are eaten by birds. composition on understory colonization by
native species in plantations on a degraded
References tropical site. Journal of Vegetation Science 6:
627-636.
Francis, J.K. and A. Rodríguez. 1993. Seeds of
Puerto Rican trees and shrubs: second Vélez, I., and J. von Overbeek. 1950. Plantas
installment. Research Note SO-374. U.S. indeseables en los cultivos tropicales. Editorial
Department of Agriculture, Forest Service, Universitaria, Río Piedras, PR. 497 p.
Southern Forest Experiment Station, New
Orleans, LA. 5 p.
355
Gossypium hirsutum L. wild cotton
MALVACEAE
John K. Francis
356
(Center for New Crops and Plants Products 2002). It has been selected, bred, and, in some cases,
Soil disturbance is usually necessary for its hybridized to give the manageable and productive
establishment. It tolerates moderate amounts of varieties planted today. Evidence from Mexico
salts in the soil and salt spray. Wild cotton shows that wild cotton has been used by man for
competes well with weeds and grasses but does not making string and cloth since at least 200 BC
tolerate shade and cannot grow under tree cover. In (Iziko Museums of Cape Town 2000). Today,
Puerto Rico, wild cotton is most competitive at low cotton is the world’s most widely used natural
rainfalls (750 to 900 mm of annual precipitation) fiber. The seeds of cultivated cotton form the basis
but grows in well-drained areas that receive up to for an important oil and animal food industry. The
1700 mm of precipitation at from near sea level to seeds contain 7.3 g water, 23.1 g protein, 22.9 g
about 300 m elevation. Wild cotton does not fat, 43.2 g total carbohydrate (16.9 g fiber), and 3.5
tolerate frost. g ash per 100 g (Center for New Crops and Plant
Products 2002). Gossypol, a sesquiterpinoid, is
Reproduction.—Wild cotton is dormant during present in vegetative parts of wild cotton as well as
the winter dry season and becomes vegetative as the cultivated varieties. It is a natural deterrent to
the rains begin again in early summer. At the end insects but is toxic to man and nonruminant
of the summer, vegetative growth ceases and plants animals. It has been shown to have strong
begin to flower and fruit (Kohel and Lewis 1984). antifertility, antitumor, antiparasite, and anti-HIV
Three collections of wild cotton seeds of Puerto properties (Bajaj 1989). Gossypol is widely used in
Rico from three locations averaged 0.088 + 0.001, China as a male contraceptive (Center for New
0.075 + 0.001, and 0.081 + 0.001 g/seed or about Crops and Plant Products 2002). Raw cottonseed
12,000 seeds/kg. Germination is epigeal. Seeds oil (containing gossypol) used alone showed
from two of the collections were sown without promise as an insecticide (Tadas and others 1994).
pretreatment in commercial potting mix. One In herbal medicine, cottonseed and roots have been
collection germinated at 100 percent starting in 17 used to treat nasal polyps, asthma, diarrhea,
days and ending in 41 days. The second group hemorrhoids, dysentery, uterine fibroids and
germinated at 67 percent starting in 22 days and certain cancers, to induce abortions, and as a
ending in 140 days. The wind disburses the seeds diuretic. (Center for New Crops and Plant Products
over short distances by blowing the lint until it 2002). Wild cotton is occasionally planted as an
catches on vegetation. Birds also move seeds when ornamental. The species contributes to biodiversity,
they use cotton lint for nest building materials. soil stability, wildlife habitat, and scenic beauty in
Under greenhouse conditions, seedlings may grow its beach strand and lowland habitat.
more than 1 m in 6 months. In wild stands,
seedlings can be abundant but they naturally thin References
themselves quickly.
Bijaj, Y.P.S. 1998. Cotton. In: Y.P.S. Bijaj, ed.
Growth and Management.—Although wild Biotechnology in Agriculture and Forestry 42.
cotton in its cultivated form is managed as an Springer-Verlag, New York. p. 3-36.
annual, it is a true perennial, and if allowed to do
so under favorable conditions, will live for several Center for New Crops and Plant Products. 2002.
years (Kohel and Lewes 1984) and become tall and Gossypium hirsutum L. Purdue University.
woody. Wild cotton in Puerto Rico grows 1 to 1.5 http://www.hort.perdue.edu/newcrop/duke_ener
m in its first year. The wild variety requires 15 or gy/Gossypium_hirsutum.html. 6 p.
more nodes to be formed along the main axis
before it can flower. Consequently, the plant is Howard, R.A. 1989. Flora of the Lesser Antilles,
rarely able to bloom during its first year (Kohel and Leeward and Windward Islands. Vol. 5. Arnold
Lewis 1984). Cotton can be transplanted using Arboretum, Harvard University, Jamaica Plain,
containerized nursery plants. However, direct MA. 604 p.
seeding following removal of overhead
competition and cultivation of the soil is Iziko Museums of Cape Town. 2000. Gossypium
recommended. Soil temperatures must be at least (cotton). Biodiversity explorer. http://www.
18 °C before planting (Center for New Crops and museums.org.za/bio/plants/malvaceae/gossypiu
Plant Products 2002). m.htm. 2 p.
Benefits.—About 87 percent of the cotton grown Kohel, R.J. and C.F. Lewis, eds. 1984. Cotton.
commercially is G. hirsutum (Vijayakumar 2002). Agronomy 24. American Society of Agronomy,
357
Inc., Crop Science Society of America, Inc., and Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
Soil Science Society of America, Inc., Madison, 50. Bernice P. Bishop Museum Press, Honolulu,
WI. 605 p. HI. 924 p.
Liogier, H.A. 1994. Descriptive flora of Puerto Nelson, G. 1996. The shrubs and woody vines of
Rico and adjacent islands. Vol. 3. Editorial de la Florida. Pineapple Press, Sarasota, FL. 391 p.
Universidad de Puerto Rico. Río Piedras, PR.
461 p. Tadas, P.L., H.K. Kene, and S.D. Deshmukh.
1994. Effect of raw cottonseed oil against
Little, E.L., Jr., R.O. Woodbury, and F.H. sucking pests of cotton. PKV Research Journal
Wadsworth. 1974. Trees of Puerto Rico and the 18(1): 142-143.
Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture, Vijayakumar, T. 2002. Cotton. http://www.
Washington, DC. 1,024 p. geocities.com/vijayakumar777/cotton1.html.
14 p.
Long, R.W. and O. Lakela. 1976. A flora of
Tropical Florida. Banyon Books. Miami, FL. Watt, Sir G. 1907. The wild and cultivated cotton
962 p. plants of the world. Longmans, Green, and Co.,
London. 406 p.
358
Gouania lupuloides (L.) Urban chew-stick
RHAMNACEAE
John K. Francis
known to have been planted or naturalized
elsewhere.
359
chew-stick grows relatively rapidly. In the presence Kennelly, E.J., W.H. Lewis, R.E.K. Winter, S.
of seed sources, reproduction is usually adequate. Johnson, M. Elvin-Lewis, and J. Gossling. 1993.
Thinning the overstory would probably allow Triterpenoid saponins from Gouania lupuloides.
existing suppressed plants to grow to larger sizes Journal of Natural Products 56(3): 402-410.
and fruit.
Liogier, H.A. 1994. Descriptive flora of Puerto
Benefits.—Chew-stick contributes to biodiversity Rico and adjacent Islands. Vol. 3. Editorial de la
and biomass accumulation, helps protect the soil, Universidad de Puerto Rico. Río Piedras, PR.
and furnishes wildlife cover in forests where it 461 p.
grows. It is an important honey plant (Townsend
1984). In Jamaica and the West Indies, stem Nelson, G. 1996. The shrubs and woody vines of
cuttings are used to clean teeth and gums and Florida. Pineapple Press, Inc., Sarasota, FL.
extracts are used to manufacture a dentifrice 391 p.
(Secretaría de Medio Ambiente y Recursos
Naturales 2002). Triterpenoid saponins may New York Botanical Garden. 2002. Flora and
contribute to the beneficial activity (Kennelly and mycota of Acre, Brazil. http://nybg.org/
others 1993). In Colombia, the plant is used for a bsci/acre/rhamnaceae.html. [not paged].
bitter flavoring in alcoholic beverages and to treat
water retention and stomach problems (Pérez- Pérez-Arbelaez, E. 1978. Plantas útiles de
Arbelaez 1978). Colombia. Litografía Arco, Bogotá, Colombia.
831 p.
References
Secretaría de Medio Ambiente y Recursos
Acevedo-Rodríguez, P. 1985. Los bejucos de Naturales. 2002. Especies con usos no
Puerto Rico. Vol. 1. General Technical Report maderables en bosques tropicales y
SO-58. U.S. Department of Agriculture, Forest subtropicales. http://www.semarnat.gob.mx/
Service, Southern Forest Experiment Station, pfnm2/fichas/gouania_lupuloides.htm. 2 p.
New Orleans, LA. 331 p.
Stevens, W.D., C. Ulloa-Ulloa, A. Pool, O.M.
Croat, T.B. 1978. Flora of Barro Colorado Island. Montiel, eds. 2001. Flora de Nicaragua.
Stanford University Press, Stanford, CA. 943 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanic Garden Press, St. Louis,
Howard, R.A. 1989. Flora of the Lesser Antilles, MO. 2,666 p.
Leeward and Windward Islands. Vol. 5. Arnold
Arboretum, Harvard University, Jamaica Plain, Townsend, G.F. 1984. Multi-purpose trees for
MA. 604 p. honey production. Echo technical note.
http://www.echonet.org/tropicalag/technotes/Mu
ltiPur.pdf. 3 p.
360
Grayia spinosa (Hook.) Moq. spiny hopsage
CHENOPODIACEAE
361
with natural regeneration of native vegetation and 163. U.S. Department of Agriculture, Forest
result in increased fine fuel accumulation. Service, Intermountain Research Station, Ogden,
UT. 52 p.
Reproduction.—Most plants are dioecious, but
the proportion varies among populations. Flowers
Daubenmire, R. 1970. Steppe vegetation of
appear in February to May and are wind
Washington. Technical Bulletin 62. Washington
pollinated. Fruits ripen in March to June (Shaw
State University, Pullman, WA. 131 p.
and others 2001) and are dispersed by wind,
gravity, and insects. Herbage, flower, and fruit
Hitchcock, C.L., A. Cronquist, M. Ownbey, and
production are highly variable among years
J.W. Thompson. 1964. Vascular plants of the
(Rickard and Warren 1981); many plants fail to
Pacific Northwest. Part 2. Flora of the Pacific
produce flowers in dry years (McArthur and
Northwest. University of Washington Press,
Sanderson 1984). During a drought, spiny hopsage
Seattle, WA. 730 p.
in a southern Idaho wildland planting began
flowering 4 years after seeding. Incubating
debracted utricles at 15 and 5 oC (8 hrs, 16 hrs) or McArthur, E.D. and S.C. Sanderson. 1984.
at 15 oC for 14 days provides an estimate of Distribution, systematics, and evolution of the
germination. A 30 to 60 day prechill at 3 to 5 oC is Chenopodiaceae, In: Proceedings—symposium
required to enhance germination of utricles from on the biology of Atriplex and related
northern populations. Shaw and others (2001) chenopods. General Technical Report INT-172.
reported an average of 1,030,600 debracted U.S. Department of Agriculture, Forest Service,
utricles/kg. Germination is epigeal. Intermountain Research Station, Ogden, UT. p.
14-24.
Growth and Management.—Seedlings develop
rapidly if adequate water is available. Mature Rickard, W.H. and R.F. Keough. 1968. Soil-plant
plants are drought tolerant. Spiny hopsage is relationships of two steppe desert shrubs. Plant
considered one of the most palatable shrubs in salt and Soil. 19:205-212.
desert shrub communities in spring and early
summer. Its use by wildlife and livestock declines
rapidly as plants enter summer dormancy. Rickard, W.H. and J.L. Warren. 1981. Response of
steppe shrubs to the 1977 drought. Northwest
Although fairly tolerant of grazing, some
populations receiving heavy use have been Science. 55:108-112.
replaced by less palatable species (Blaisdell and
Holmgren 1984). Shaw, N.L., M.R. Haferkamp, and E.G. Hurd.
2001. Grayia Hook. & Arn. hopsage. In: F.T.
Bonner, and R.G. Nisley, eds. Woody Plant
Benefits.—Spiny hopsage contributes to diversity
on native sites that generally support few other Seed Manual. U.S. Department of Agriculture,
woody species. It provides cover for birds and Forest Service, Washington, D.C.
http://wpsm.net/ 12 p.
other small animals, spring and early summer
forage for big game and livestock, and soil
stabilization on moderate slopes. Litter rich in Shaw, N.L., E.G. Hurd, and M.R. Haferkamp.
1996. Spiny hopsage fruit and seed morphology.
potassium and other accumulated cations
enhances growth and nutrient content of plants Journal of Range Management. 49:551-553.
growing beneath its canopy (Rickard and Keough
1968). Native Americans ground parched seed of Stubbendieck, J., S.L. Hatch, and K. Hirsch. 1986.
spiny hopsage to prepare pinole flour North American range plants. University of
(Stubbendieck and others 1986). Nebraska Press, Lincoln. 465 p.
362
Guettarda elliptica Sw. velvetseed
RUBIACEAE
John K. Francis
363
difficult habitat where most occur. Individuals smooth- and rough-leaved velvetseed,
probably live from two to several decades. No Guettarda elliptica Sw. and G. scabra in the
management experience has been published. Everglades. American Journal of Botany
However, long-term protection of velvetseed sites 83(supplement): 70.
is recommended.
Liogier, H.A. 1997. Descriptive flora of Puerto
Benefits.—Velvetseed contributes to biodiversity Rico and adjacent islands. Vol. 5. Editorial de
and wildland biomass while it helps protects the la Universidad de Puerto Rico, San Juan, PR.
soil from erosion and provides food and cover for 436 p.
wildlife. The species is listed as the larval food
plant for the moths, Calidota strigosa Walker in Little, E.L., Jr., R.O. Woodbury, and F.H.
Florida and the West Indies (Barns 2002) and Wadsworth. 1974. Trees of Puerto Rico and the
Hylesia lineata Druce in Mexico (Pescador 1995). Virgin Islands. Vol. 2. Agriculture Handbook
The wood is useful for fuel and stakes. Velvetseed 449. U.S. Department of Agriculture,
is a pretty shrub and would make a fine foundation Washington, DC. 1,024 p.
plant in seminatural landscaping.
Nelson, G. 1996. The shrubs and woody vines of
References Florida. Pineapple Press, Inc., Sarasota, FL.
390 p.
Barns, M.J.C. 2002. Moths of Jamaica, Calidota
strigosa. http://members.fortunecity.co.uk/ Pescador, A.R. 1995. Distribution and abundance
jamaicamoth/images/calstr.htm. 1 p. of Hylesia lineata egg masses in a tropical dry
forest in western Mexico. Southwestern
Champion Tree Project. 2002. National Champion Entomologist 20(3): 367-375.
Trees, Florida. http://www.championtrees.org/
database/championsFL.htm. 8 p. West, E. and L.E. Arnold. 1952. The native trees
of Florida. University of Florida Press,
Koptur, S. and J.H. Richards. 1996. Comparative Gainesville, FL. 212 p.
floral biology and breeding systems of
364
Gundlachia corymbosa (Urban) Britt. ex Boldingh yambush
ASTERACEAE
John K. Francis
365
furnishes cover for wildlife. Although there is no http://www.unesco.org/csi/pub/papers/gerace.
record of it having been used for ornamental htm. 20 p.
purposes, yambush seems an excellent choice for
groundcovers and borders in seaside and xeric Howard, R.A. 1989. Flora of the Lesser Antilles,
gardens. Poultices of crushed leaves relieve and Leeward and Windward Islands. Vol. 6. Arnold
help heal allergic dermatitis (Wood 2003). Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
References
Liogier, H.A. 1997. Descriptive flora of Puerto
Britton, N.L. and C.F. Millspaugh. 1962. The Rico and adjacent islands. Vol. 5. Editorial de la
Bahama flora. The New York Botanical Garden, Universidad de Puerto Rico, Río Piedras, PR.
New York. 695 p. 436 p.
Darwin Institute. 2002. Plan for biodiversity New York Botanical Garden. 2003. Specimens
management and sustainable development detailed results: Gundlachia domingensis A.
around Turks and Caicos Ramsar site: Gray var. corymbosa Urb. (syntype).
introduction. http://www.ukotcf.org/pdf/ http://scisun.nybg.org:8890/searchdb/owa/www
TCIRamMP/Parts%20One%20and%Two.pdf. catalog.detail_list. 1 p.
48 p.
Wood, K.M. 2003. Medicinal plants for what ails
Gerace, D.T., G.K. Ostrander, and G.W. Smith. you in Paradise. http://www.timespub.tc/
2003. San Salvador, Bahamas. Coastal region Natural%20History/Archive/Winter%20200102/
and small island papers 3. UNESCO, Rome. medicinal.htm. 2 p.
366
Gutierrezia sarothrae (Pursh) Britt. & Rusby broom snakeweed
ASTERACEAE
Juanita A. R. Ladyman
367
drought (Schmutz et al. 1992). It suffers from dryness, and an absence of grazing for at least one
drought to about the same extent as black grama season prior to burning has been recommended
grass, but when insect pressure is present the (Tirmenstein 1999). In many cases, fire may
snakeweed mortality is much higher (Dick-Peddie prove an ineffectual control. For example, 5 years
1993). However, by virtue of its high seed after a wildfire snakeweed had gone from 0
production and rapid seedling establishment percent canopy cover to over 40 percent canopy
(Osman and Pieper 1988) it often increases, or cover on monitored plots in Arizona (Pase and
“comes in”, after prolonged drought and other Pond 1964). In west-central Utah broom
disturbances (Dayton 1931). snakeweed was found on eight out of the nine age-
classes of burns, being absent on the 71-year-old
Reproduction.—Snakeweed flowers from July to burns and most abundant on the 22-year-old burns
December depending upon the latitude and water (Barney and Frischknecht 1974). Broom
availability. The plant is insect pollinated snakeweed is susceptible to some herbicides
(Mayeux and Leotta 1981). It is a prodigious (Whitson et al. 1991).
producer of seed that can remain viable in the soil
for a considerable length of time (Osman and Benefits.—Broom snakeweed provides some
Pieper 1988). A single plant is capable of stability to the soil against both wind and water
producing 9,000 to 10,000 seeds annually erosion (Tirmenstein 1999, USDAFS 1988). It is
(Tirmenstein 1999). The oval fruit (seeds) are fair browse for wildlife but is poisonous to
achenes and covered with stiff hairs or nearly livestock due to the saponin content, which is
glabrous. Chaffy scales, called the pappus, are on variable and depends upon both growth phase and
one end of the achene and aid seed dispersal by environmental conditions (Hatch and Pluhar
wind (Cronquist, 1994). Seeds are dispersed by 1993). The vegetation is most toxic during leaf
animals as well as wind, although it has been formation (Whitson and others 1991). At times
reported that they also accumulate under the shrub when there has been no alternative forage,
canopy (Mayeux and Leotta 1981). Light enhances snakeweed has been used by livestock with no ill
germination, which suggests that the best position effects (USDAFS 1988). Presumably saponin
for seeds is on or near the soil surface (Mayeux levels have been low at these times. It has several
and Leotta 1981). medicinal uses among the Navajo (Mayes and
Lacy 1994). It is used to heal cuts and bites in
Growth and Management.—Snakeweed is not humans and animals. A snakeweed medicine is
palatable and can be poisonous to livestock and given in childbirth and is used to treat
increases under grazing pressure (Cable 1975; stomachaches and other internal disorders. It is
Carter 1988; Cronquist 1994; Pase and Pond reported to be a powerful diuretic (Powell 1998). It
1964). Most losses in sheep and cattle are due to is also used in various Navajo ceremonies
abortions (Whitson et al. 1991), but the plant also including as an emetic and fumigent (Mayes and
apparently affects the kidneys of both sheep and Lacy 1994). A yellow dye is made from the
horses (Dayton 1931). Data suggest that the flowers, and the stems can be used to make fire by
presence of jackrabbits may reduce cattle losses friction. Birds, for example quail, eat the seeds
from poisonous plants because, under moderate (Epple 1995). Broom snakeweed also provides
stocking rates, jackrabbits consume increased cover and resting sites for species of small
amounts of poisonous plants and shrubs such as mammals, for example jackrabbits (Daniel and
snakeweed but have minimal forage competition others 1993) and birds, for example lesser prairie
with cattle (Daniel et al. 1993). When abundant, chicken (USDAFS 1988) and Columbia basin
snakeweed is generally considered an indicator of burrowing owl (Green and Anthony 1989).
overgrazing (Dayton 1931; Hatch and Pluhar
1993; Powell, 1998). Pastures are also not References
properly utilized when broom snakeweed is
present amongst the grasses (Whitson et al. 1991). Barney, M.A. and N.C. Frischknecht. 1974.
Snakeweed is killed by certain borers and other Vegetation changes following fire in the pinyon-
insects. Species from the genera, Crossidius, juniper type of west-central Utah. Journal of
Diplotaxis, and Mecas have been proposed as Range Management 27(2): 91-96.
possible biological controls (Dayton 1931). Fire
can be used to reduce snakeweed (Humphrey Burgess, T.L. 1995. The dilemma of coexisting
1949; Tirmenstein 1999). However, the success of growth forms. In: M.P. McClaran and T.R. Van
controlled burns depends upon fuel quantity and
368
Devender, eds. The desert grassland. The Greenland. Vol. 1 – Checklist. 2nd ed. Timber
University of Arizona Press, Tucson, AZ. 346 p. Press, Portland, OR. 622 p.
Cable, D.R. 1975. Range management in the Mayes, V.O. and B.B. Lacy, 1994. Nanise’ Navajo
chaparral type and its ecological basis: the status herbal. Navajo Community College Press,
of our knowledge. Research Paper RM-155. Tsaile, AZ. 153 p.
U.S. Department of Agriculture, Forest Service,
Rocky Mountain Forest and Range Experiment Mayeux, H.S., Jr. and L. Leotta. 1981. Germination
Station. Fort Collins, CO. 30 p. of broom snakeweed (Gutierrezia sarothrae) and
threadleaf snakeweed (G. microcephalum) seed.
Carter, J. L. 1988. Trees and Shrubs of Colorado. Weed Science. 29: 530-534.
Johnson Books, Boulder, CO. 165 p.
Osman, A. and Pieper, R.D. 1988. Growth of
Cronquist, A. 1994. Intermountain Flora, Vascular Gutierrezia sarothrae seedlings in the field.
Plants of the Intermountain West, USA. Vol. 5. Journal of Range Management 41(1): 92-93.
Asterales. New York Botanical Garden, Bronx,
NY. 496 p. Pase C.P. and F.W. Pond. 1964. Vegetation
changes following the Mingus Mountain burn.
Daniel, A., J. L. Holechek, R. Valdez, A. Tem, L. Note RM-18. U.S. Department of Agriculture,
Saiwana, M Rusco, and M. Carden. 1993. Range Forest Service, Rocky Mountain Research
condition influences on Chihuahuan Desert Station, Fort Collins, CO. 8 p.
cattle and jackrabbit diets. Journal of Range
Management 46(4): 296-301. Powell, A. M. 1998. Trees & shrubs of Trans-
Pecos and adjacent areas. Big Bend Natural
Dayton, W. A. 1931. Important western browse History Association. (Reprinted by University of
plants. Miscellaneous Publication 101. U.S. Texas University Press, Austin, TX.) 498 p.
Department of Agriculture, Washington, DC.
214 p. Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox,
J.O. Klemmedson, J.J. Norris, and L.C. Fierro.
Dick-Peddie, W.A. 1993. New Mexico Vegetation 1992. Desert Grasslands. In: R.T. Coupland, ed.
– past, present, and future. University of New Ecosystems of the World, 8A; Natural
Mexico Press, Albuquerque, NM. 244 p. Grasslands Introduction and Western
Hemisphere. Elsevier, London, NY. p. 337-362
Epple, A.O. 1995. A Field Guide to the Plants of
Arizona. Falcon Press Publishing Co., Helena, Texas A&M University “Virtual Herbarium”.
Montana. 347 p. Uvalde Research and Extension Center. 2000.
Copyright Texas A&M University System.
Green, G.A. and R.G Anthony. 1989. Nesting http://uvalde.tamu.edu/herbarium/gusa.htm [not
success and habitat relationships of burrowing paged].
owls in the Columbia Basin, Oregon. The
Condor. 91: 347-354. Tirmenstein, D. 1999. USDA Forest Service,
Rocky Mountain Research Station, Fire Sciences
Hatch, S.L. and J. Pluhar. 1993. Texas range Laboratory. Fire Effects Information System,
plants. The W.L. Moody Jr Natural History http://www.fs.fed.us/database/ [not paged].
Series 123. Texas A&M University Press,
College Station, TX. 326 p. Whitson, T.D., ed., L.C. Burrill, S.A. Dewey, D.W.
Cudney, B.E. Nelson, R.D. Lee, and R. Parker.
Humphrey, R.R. 1949. Fire as a means of 1991. Weeds of the West. The Western Society
controlling velvet mesquite, burroweed, and of Weed Science, Western U.S. Land Grant
cholla on southern Arizona ranges. Journal of Universities Cooperative Extension Services and
Range Management. 2: 175-182. the University of Wyoming, Pioneer of Jackson
Hole, Jackson WY. p 132-133.
Kartesz, J.T. 1994. A synonymized checklist of the
vascular flora of the United States, Canada, and United States Department of Agriculture, Forest
Service. 1988. Range Plant Handbook. Dover
Publications, Inc. New York. 837 p.
369
Gynerium sagittatum (Aubl.) Beauv. wild cane
POACEAE
John K. Francis
370
20 m from the parent plants (Pohl 1983). Segments Croat, T.B. 1978. Flora of Barro Colorado Island.
of culm or rhizome, carried by floodwaters and Stanford University Press, Stanford, CA. 943 p.
covered with soil or debris, sprout and start new
colonies. Howard, R.A. 1979. Flora of the Lesser Antilles,
Leeward and Windward Islands. Vol. 3. Arnold
Growth and Management.—Growth of wild cane Arboretum, Harvard University, Jamaica Plain,
is rapid. Nursery seedlings reached 20, 30, and 50 MA. 586 p.
cm after 1, 2, and 4 months (Kalliola and others
1992). How long seedlings take to reach maturity Instituto Nacional de Biodiversidad. 2002.
and how rapidly suckers grow is unknown. Jerarquía taxonómica: Lista de especímenes de
Theoretically, baring catastrophes and invasion and Gynerium sagittatum. http://www.inbio.ac.cr/
shading by trees, clones can endure indefinitely. bims/k03/p13/c046/00159/f01382/g008647/s027
Culms of Amazon Basin plants produced close to 308.htm. 2 p.
200 leaves during their lifetimes, having from 19 to
28 living leaves at a time (Kalliola and others Kalliola, R., M. Puhakka, and J. Salo. 1992.
1992). Unbranched culms die after flowering, but Interspecific variation, and the distribution and
only the branches of branched culms die. If not ecology of Gynerium sagittatum (Poaceae) in the
controlled, wild cane slowly invades wet western Amazon. Flora 186(3-4): 153-167.
bottomland pastures and eliminates forage plants.
Periodic mowing appears to be adequate for control Kalliola, R., J. Salo, M. Puhakka, and M. Rajasilta.
of advancing clumps. 1991. New site formation and colonizing
vegetation in primary succession on the western
Benefits.—Wild cane provides cover for wildlife Amazon floodplains. Journal of Ecology 79(4):
and protects stream banks from erosion (Kalliola 877-901.
and others 1992). Its culms lack the strength and
toughness of hardwoods and bamboo but still are Pohl, R.W. 1983. Gynerium sagittatum (caña
used in rude construction, drying racks, vegetable brava, cane). In: D.H. Janzen, ed. Costa Rican
stakes and fruit props, and for weaving mats, natural history. University of Chicago Press,
baskets, and hats (Kalliola and others 1992). In the Chicago and London. p. 248-249.
Amazon area, arrow shafts are made from the dried
culms (Russo 2002). Plumes are used for dry floral Russo, E.B. 2002. Village life of the Machiguenga.
arrangements. http://montana.com/manu/village.html. 4 p.
371
Hamelia patens Jacq. firebush
RUBIACEAE
John K. Francis
modified dichasium with flowers that are tubular,
12 to 22 mm long, and orange to red in color. The
fruit is a berry, spherical to elliptical, 7 to 10 mm
long, turning red and then black at maturity. The
seeds are orange-brown, 0.6 to 0.9 mm long
(Howard 1989, Liogier 1997).
372
Another experiment using an auto-humid chamber Cunningham, S.A. 1994. Measuring the
obtained 100 percent rooting of softwood cuttings relationships between floral duration and fruit
treated with hormones and 40 percent without set for Hamelia patens (Rubiaceae). Biotropica
hormone treatment (Maiti 1974). 26(2): 227-229.
Growth and Management.—In its native and Desert-Tropicals. 2001. Texas firecracker bush,
other frost-free habitat, firebush grows as a typical scarlet bush. http://www.desert-
shrub, but cultivated in warm temperate areas with tropicals.com/Plants/Rubiaceae/Hamelia_patens.
hard frosts, it will freeze to the ground and sprout html. 2 p.
again in the spring. Firebush is also planted as an
annual bedding plant in temperate climates. Floridata. 2001. Hamelia patens.
Ornamental plants need frequent watering during http://www.floridata.com/ref/h/hame_pat.cfm.
the establishment phase, but afterward, they are 3 p.
drought tolerant (Floridata 2001). Pruning of
ornamental plants is recommended to keep them Howard, R. A. 1989. Flora of the Lesser Antilles,
from becoming tall and spindly (Desert-Tropical Leeward and Windward Islands. Vol. 6. Arnold
2001). However, pruning inhibits blooming, which Arboretum, Harvard University, Jamaica Plain,
occurs at the terminals. MA. 658 p.
Benefits.—Firebush is loved and planted as an Liogier, H.A. 1997. Descriptive flora of Puerto
ornamental almost worldwide in warm, moist Rico and adjacent islands. Vol. 5. Editorial de la
areas. The fruit is edible (Little and others 1974). A Universidad de Puerto Rico, San Juan, PR.
phytochemical screening found firebush tissues 436 p.
rich in phytochemicals including alkaloids and
flavonoids (Raintree Nutrition 2001). Firebush is Liogier, H.A. 1990. Plantas medicinales de Puerto
used in herbal medicine to treat athlete’s foot, skin Rico y del Caribe. Iberoamericana de Ediciones,
lesions and rash, insect bites, nervous shock, Inc. San Juan, PR. 566 p.
inflammation, rheumatism, headache, asthma, and
dysentery (Liogier 1990, Mast Arboretum 2001, Little, E.L., Jr., R.O. Woodbury, and F.H.
Raintree Nutrition 2001). Firebush is one of the Wadsworth. 1974. Trees of Puerto Rico and the
woody species foraged by sheep and goats in Virgin Islands. Vol. 2. Agriculture Handbook
brushy pastures in Costa Rica. These animals 449. U.S. Department of Agriculture,
consumed 1.3 percent of their body weight of this Washington, DC. 1,024 p.
species in a free-choice test (Benavides 2001).
Firebush contains 17.5 percent crude protein and Maiti, R.G. 1974. Regeneration of terminal
has an in vitro digestibility of 61.6 percent greenwood cuttings of some ornamental shrubs
(Benavides 2001). under auto-humid chamber. Plant Science 6:
104-105.
References
Mast Arboretum. 2001. Hamelia patens, firebush.
Benavides, J.E. 2001. Arboles y arbustos Steven F. Austin State University.
forrajeros: una alternativa agroforestal para la http://www.sfasu.edu/ag/arboretum/plants/hamel
ganadería. Conferencia electrónica de la FAO iapatens/hamelia%20patens.htm. 3 p.
sobre la producción animal en Latinoamerica.
http://lead.virtualcentre.org/es/ele/conferencia1/ Paciorek, C.J., B.R. Moyer, R.A. Levin, and S.L.
bnvdes23.htm. 22 p. Halpern. 1995. Pollen consumption by the
hummingbird flower mite, Proctolaelaps
Bhattacharjee, S.K. and M. Balakrishna. 1992. kirmsei, and possible fitness effects on Hamelia
Studies on propagation of Hamelia patens Jacq. patens. Biotropica 27(2): 258-262.
and Ixora singaporensis Hort. from stem
cuttings. Progressive Horticulture 24(3/4): 157- Raintree Nutrition. 2001. Scarlet bush. Raintree
164. Nutrition, Inc. http://www.rain-
tree.com/scarletbush.htm. 3 p.
373
Helicteres jamaicensis Jacq. cowbush
STERCULIACEAE
John K. Francis
374
The fibrous bark is strong and was once used for http://jamaicans.com/tourist/bites.htm. 3 p.
making rope (Little and others 1974). Cowbush is
reported to have several uses in herbal medicine. A Liogier, H.A. 1994. Descriptive flora of Puerto
tea made from the wood and leaves is used to treat Rico and adjacent Islands. Vol. 3. Editorial de la
bilious conditions and as an emetic. A decoction of Universidad de Puerto Rico. 461 p.
those same tissues is good for fever and cough, and
the cooked root is able to stop bleeding (Liogier Liogier, H.A. 1990. Plantas medicinales de Puerto
1990). The hairs of the species are reported to Rico y del Caribe. Iberoamericana de Ediciones,
cause serious irritation (Jamaicans 2001), but this Inc., San Juan, PR 566 p.
is not reported by Little and others (1974) and was
found to be untrue by the author, at least for Puerto Little, E.L., Jr., R.O. Woodbury, and F.H.
Rican plants. Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
References 449. U.S. Department of Agriculture,
Washingtion, DC. 1,024 p.
Griffiths, M. 1994. Index of garden plants. Timber
Press, Portland, OR. 1,234 p. Reforesta. 2001. Restauración ribereña.
http://www.reforesta.com/projects/Espanol
Jamaicans. 2001. Jamaica: Bites and Stings, etc. /riparian_restoration_esp.htm. 2 p.
375
Hibiscus pernambucensis Arruda seaside mahoe
MALVACEAE
John K. Francis
capsules develop, and upon drying, split open
along five lines to release their seeds.
376
(Aleyrodidae), and mealy bugs (Pseudococcideae) cleared planting spots with no other treatment. The
sometimes cause chlorosis and slower growth. fast-growing species that were planted overtopped
the sprouting seaside mahoe and eventually
Reproduction.—The principal means of suppressed them.
reproduction is by the layering and sprouting of
prostrate stems. The old prostrate stems eventually Benefits.—The fibrous inner bark was formerly
die and rot, leaving the sprouts as new, isolated used to make ropes, cords, fish nets, floats, cloth,
individuals. Seeds are produced in relative and mats. It is still used for emergency ties and
abundance and occasionally result in seedlings that lashings. The wood is used for fuel in
can eventually establish new clones. Streams and underdeveloped areas (National Academy of
floodwaters are the only known effective vector of Sciences 1983). The flowers make a colorful and
seed dispersal. tasty addition to salads and can be cooked as a
In wetter habitat, seaside mahoe flowers vegetable (Workman 1980). The young leaves,
continuously throughout the year (Little and bark, and roots were reportedly used for food
Wadsworth 1964). The species flowers seasonally during famines in former times (Little and
in habitat with drought stress during part of the Wadsworth 1964). Bees make honey from the
year. The flowers are pollinated by insects, and nectar. The plant is used as an ornamental and to
yield mature seeds about 1 month later. Fruits can stabilize stream banks (Workman 1980).
potentially produce at least 40 seeds per fruit. In
one collection from upland habitat in Puerto Rico, References
fruits averaged only 7.7 filled seeds per fruit. In
two other Puerto Rican collections from upland Howard, R.A. 1989. Flora of the Lesser Antilles,
habitat, the seeds averaged 0.018 + 0.006 and Leeward and Windward Islands. Vol. 5. Arnold
0.024 + 0.003 g/seed. These seed collections Arboretum, Harvard University, Jamaica Plain,
yielded 41 and 53 percent germination, which MA. 604 p.
occurred between 15 and 220 days after sowing.
Germination is epigeal. Seedlings are easily raised Liogier, H.A. 1994. Descriptive flora of Puerto
in the nursery from seed using traditional Rico and adjacent islands. Vol. 3. Editorial de la
techniques. Plants can be established from cuttings Universidad de Puerto Rico. Río Piedras, PR.
and air layers (Workman 1980). In one small 461 p.
planting in Puerto Rico, 40 of 140 untreated
cuttings (45 cm long and 2.5 cm minimum Little, E.L., Jr. and F.L. Wadsworth. 1964.
diameter) placed directly in the planting site rooted Common trees of Puerto Rico and the Virgin
and grew aggressively. Plants can also be started Islands. Agriculture Handbook 249. U.S.
from very large cuttings as living fence posts Department of Agriculture, Washington, DC.
(Little and Wadsworth 1964). 548 p.
Growth and Management.—Seaside mahoe Malavasi, U.C. and M.M. Malavasi. 1999. Tree
grows rapidly from sprouts and can extend its shade preferred by recreational users of a
leader 3 m or more per year. Seedlings grow Brazilian beach sidewalk. Journal of
slowly at first. A group of 6-month-old seedlings Arboriculture 25(2): 85-87.
averaged only 17 cm in height. Probably, 1 year is
required before nursery seedlings are ready for Martorell, L.F. 1975. Annotated food plant catalog
outplanting. Seaside mahoe may occasionally of the insects of Puerto Rico. Agricultural
reach 12 to 15 m in height (Workman 1980), but Experiment Station, University of Puerto Rico,
more typically reaches only 6 m in height and 15 Río Piedras, PR. 303 p.
cm in stem diameter. Because the species is
mainly valued in natural stands for environmental National Academy of Sciences. 1983. Firewood
protection and because it aggressively maintains crops: shrub and tree species for energy
nearly pure stands, little or no management is production. BOSTID Report 40. National
required once it is established. One unwanted Academy of Sciences. Washington, DC. 92 p.
upland stand was converted to timber species by
slashing the seaside mahoe, piling the slash West, E. and L.E. Arnold. 1952. The native trees
between the planting spots, and planting seedlings of Florida. University of Florida Press,
of Eucalyptus grandis Hill ex Maiden, Terminalia Gainsville, FL. 212 p.
ivorensis L., and Hibiscus elatus Sw. in the small
377
Workman, R.W. 1980. Growing native. The
Sanibel-Captiva Conservation Foundation, Inc.
Sanibal, FL. 137 p.
378
Holodiscus discolor (Pursh) Maxim. creambush oceanspray
ROSACEAE
Nancy L. Shaw
on rocky talus slopes with shallow sandy to clay
loam soils. Remnant stands are found on higher
peaks of Great Basin mountain ranges (Hitchcock
and others 1961, Stark 1966, USDA Forest Service
1937).
379
grown from cuttings, but rooting varies widely Ericaceae. University of Washington Press,
among clones, cutting type, and propagation Seattle, WA. 614 p.
technique (Antieau 1987, Link 1993).
King, J.E. 1947. The effect of various treatments
Growth and Management.—Seedlings develop to induce germination of seeds of some plants
slowly, producing spreading, fibrous root systems valuable for soil conservation and wildlife.
(Wright and others 1979). Growth rate is moderate Master’s thesis. University of Idaho, Moscow,
and protection from introduced grass and forb ID. 97 p.
competition is required for one or two growing
seasons. Excessive browsing is rarely a threat to Kruckeberg, A.R. 1982. Gardening with native
developing seedlings, but they may be girdled by plants of the Pacific Northwest. University of
rodents or trampled by big game or livestock. Brush Washington Press, Seattle, WA. 252 p.
fields may be rejuvenated by prescribed burning at
10 to 15 year intervals (Orme and Leege 1980). Link, E. 1993. Native plant propagation techniques
for National Parks interim guide. Rose Lake
Benefits.—Creambush oceanspray provides habitat Plant Materials Center, East Lansing, MI. 240 p.
for a wide variety of organisms. Its palatability and
forage value vary geographically but are generally Marchant, C. and J. Sherlock. 1984. A guide to
low for big game and domestic livestock. Use may selection and propagation of some native woody
be greater in winter when other forage is limited. species for land rehabilitation in British
Palatability may be improved following fire. Columbia. Forest Research Report RR84007-
Creambush oceanspray offers considerable potential HQ. Ministry of Forests, Victoria, BC. 117 p.
for revegetating disturbed areas. Populations
capable of growing on dry, rocky, well-drained sites Morgan, P. and L.F. Neuenschwander. 1988. Seed-
or exhibiting low palatability may be particularly bank contributions to regeneration of shrub
valuable. It has been recommended for use in species after clear-cutting and burning. Canadian
highway plantings, riparian areas, windbreaks, Journal of Botany 66: 169-172.
erosion control, wildlife habitat improvement,
conservation plantings, and low maintenance Orme, M.L. and T.A. Leege. 1980. Phenology of
landscaping (Antieau 1987). Native Americans ate shrubs on a north Idaho elk range. Northwest
the seeds and made digging sticks and arrow shafts Science 54: 187-198.
from the hard, straight branches.
Shaw, N.L., E.G. Hurd, and P.F. Stickney.
References Holodiscus (K. Koch) Maxim. 2001. Ocean-
spray. In: F.T. Bonner and R.G. Nisley, eds.
Antieau, C.J. 1987. Field notes: Holodiscus Woody plant seed manual. U.S. Department of
discolor. American Nurseryman 166: 110. Agriculture, Forest Service, Agriculture
Handbook. Washington, D.C.
Berch, S.M., S. Gamiet, and E. Deom. 1988. http://wpsm.net/index.html. 7 p.
Mycorrhizal status of some plants of
southwestern British Columbia. Canadian Stark. N. 1966. Review of highway planting
Journal of Botany 66: 1,924-1,928. information appropriate to Nevada. College of
Agriculture Bulletin B-7. University of Nevada,
Everett, P.C. 1957. A summary of the culture of Desert Research Institute. Reno, NV. 209 p.
California plants at the Rancho Santa Ana
Botanic Garden. The Rancho Santa Ana Botanic USDA Forest Service. 1937. Range plant
Garden, Claremont, CA. 223 p. handbook. U.S. Government Printing Office,
Washington, D.C. 512 p.
Harrington, C.A., J.M. McGrath, and J.M. Kraft.
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380
Wright, H.A., L.F. Neuenschwander, and C.M.
Britton. 1979. The role and use of fire in
sagebrush-grass and pinyon-juniper plant
communities: A state of the art review. General
Technical Report INT-58. U.S. Department of
Agriculture, Forest Service, Intermountain
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UT. 48 p.
381
Holodiscus dumosus (Nutt. ex Hook.) Heller rockspirea
ROSACEAE
Synonyms: Spiraea dumosa (Nutt.) T. & G.
Schizonotus dumosus Koehne.
Holodiscus microphyllus Rydb.
Sericotheca dumosa Rydb.
Nancy L. Shaw
382
rocky, inaccessible areas and on summer ranges range shrubs. Journal of Range Management 6:
where other species receive preferential use. Based 309-317.
on results of clipping studies in Oregon, Garrison
(1953) recommended a maximum of 50 to 60 Harrington, H.D. 1954. Manual of the plants of
percent use for sustained production. On some Colorado. Sage Books, Denver, CO. 666 p.
western juniper/big sagebrush habitat types of
eastern Oregon, the species tends to increase when Hitchcock, C.L., A. Cronquist, M. Ownbey, and
other species receive excessive use (Ferguson J.W. Thompson. 1961. Vascular plants of the
1983, Hopkins and Kovalchik 1983). Pacific Northwest. Part 3: Saxifragaceae to
Ericaceae. University of Washington Press,
Benefits.—Rockspirea is a potentially valuable Seattle. 614 p.
species for revegetation as it is drought tolerant
and adapted to sites with dry, rocky, unstable Hopkins, W.E. and B.L. Kovalchik. 1983. Plant
surface conditions in a variety of forested and associations of the Crooked River National
nonforested communities (Stark 1966). Its use has Grassland. R6 Ecology 133-1983. U.S.
been limited by a lack of high quality seed Department of Agriculture, Forest Service,
supplies. Palatability and forage value of Pacific Northwest Region, Portland, OR. 98 p.
rockspirea are low (USDA 1937), but it does
King, J.E. 1947. The effect of various treatments
provide food and cover for small animals. Kufeld
to induce germination of seeds of some plants
and others (1973) reported its use by mule deer
valuable for soil conservation and wildlife. .
(Odocoilus hemionus) was moderate in fall and
Master’s thesis. University of Idaho, Moscow,
light during the remainder of the year. Rockspirea ID. 97 p.
is used in summer by bighorn sheep (Ovis
canadensis) and rabbits (Sutton and Johnson 1974, Kufeld, R.C. 1973. Foods eaten by the Rocky
Todd 1975). It is an attractive, low water use Mountain elk. Journal of Range Management
ornamental. Native Americans made digging sticks 26: 106-113.
and arrow shafts from the hard, straight branches
(Anderson and Holmgren 1969, Daubenmire 1970, McArthur, E.D. and S.C. Sanderson. 1985. A
Hopkins and Kovalchik 1983). Native Americans of cytotaxonomic contribution to the western North
the Great Basin ate the seeds, and pioneers made American rosaceous flora. Madroño 32: 24-28.
nails from its wood.
Stark. N. 1966. Review of highway planting
References information appropriate to Nevada. College of
Agriculture Bulletin B-7. University of Nevada,
Anderson, B.A. and A.H. Holmgren. 1969. Desert Research Institute, Reno. 209 p.
Mountain plants of northeastern Utah. Circular
319. Utah State University, Logan, UT. 148 p. Sutton, R. and C.W. Johnson. 1974. Landscape
plants from Utah’s mountains. EC-368. Utah
Daubenmire, R. 1970. Steppe vegetation of State University, Logan, UT. 137 p.
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State University, Pullman, WA. 131 p. Todd, J.W. 1975. Foods of Rocky Mountain
bighorn sheep in southern Colorado. Journal of
Ferguson, R.B. 1983. Use of rosaceous shrubs for Wildlife Management 39: 108-111.
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Intermountain rangelands--Improvement of handbook. U.S. Government Printing Office,
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General Technical Report INT-157. U.S.
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Station, Ogden, UT. p. 136-139. Naturalist Memoir 9. Brigham Young
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Garrison, G.A. 1953. Effects of clipping on some
383
Hyperbaena laurifolia (Poir.) Urban limestone snakevine
MENISPERMACEAE
John K. Francis
384
growth rate, and plants known to the author appear Leeward and Windward Islands.
to be several decades old. However, no data on Dicotyledoneae, Part 1. Vol. 4. Arnold
growth rate and longevity or experience on Arboretum, Harvard University, Jamaica Plain,
management have been published. MA. 673 p.
Benefits.—The fruits of limestone snakevine are Liogier H.A. 1985. Descriptive flora of Puerto
slightly acid, bitter, and unpleasant to the taste and Rico and adjacent islands, Spermatophyta. Vol.
smell. The fruit pulp immediately stains skin and 1. Editorial de la Universidad de Puerto Rico,
cloth a lasting reddish-brown and might be useful Río Piedras, PR. 352 p.
as a natural dye in crafts. The wood is heavy and
hard and would certainly be good for small- Little, E.L., Jr., R.O. Woodbury, and F.H.
diameter fuel. The species contributes to the Wadsworth. 1974. Trees of Puerto Rico and the
biodiversity of local forests and in a minor way, Virgin Islands. Vol. 2. Agriculture Handbook
furnishes wildlife food and cover, and protects 449. U.S. Department of Agriculture,
against erosion. Washingtion, DC. 1,024 p.
385
Indigofera suffruticosa P. Mill. wild indigo
FABACEAE
John K. Francis
386
produced per hectare from which 227 kg of indigo HealthLink. 2001. Monograph: indigo naturalis.
paste can be separated. Wild indigo tested for http://www.healthlink.com.au/nat_lib/htm-
revegetation of soil-lignite overburden varied in data/htm-herb/bhp1016.htm. 3 p.
biomass yield from 162 to 2,432 kg/ha and
maintained from 8 to 30 percent cover over a 3- Howard, R.A. 1988. Flora of the Lesser Antilles,
year period (Skousen and Call 1987). Surface Leeward and Windward Islands.
sowing at the start of the rainy season on freshly Dicotyledoneae, Part 1. Vol. 4. Arnold
tilled ground is recommended. Plants usually live 2 Arboretum, Harvard University, Jamaica Plain,
or 3 years in Puerto Rico. MA. 673 p.
Benefits.—Although indigo obtained from other Liogier H.A. 1988. Descriptive flora of Puerto
species of Indigofera was used in the Old World, Rico and adjacent islands, Spermatophyta. Vol.
the use of wild indigo by pre-Columbian natives of 2. Editorial de la Universidad de Puerto Rico,
Mexico to dye cloth and paint in various shades of Río Piedras, PR. 481 p.
blue was passed down to the Spanish colonists
(Haude 1997). Indigofera species became Liogier, H.A. 1990. Plantas medicinales de Puerto
important commercial crops in various tropical and Rico y del Caribe. Iberoamericana de Ediciones,
subtropical areas. The blue dye was produced by Inc., San Juan, PR. 566 p.
fermentation of the leaves, usually with caustic
soda or sodium hydrosulfite, and the exudates Moreira-Casasola, P., J.P. Grime, and M.L.
processed into dry cake. The blue color developed Martinez. 1994. A comparative study of the
as the cake was exposed to the air (Simon and effects of fluctuations in temperature and
others 1984). Indigo was used as a bluing to moisture supply on hard coat dormancy in seeds
counter the yellowing in clothes from washing with of coastal tropical legumes in Mexico. Journal of
soap (Vélez and van Overbeek 1950). In the last Tropical Ecology 10(1): 67-86.
few decades, natural indigo has been almost wholly
replaced by synthetic dyes. Poultices and extracts Neal, M.C. 1965. In gardens of Hawaii. Special
of wild indigo leaves, alone or in combination with Publication 50. Bernice P. Bishop Museum
other ingredients, are used in herbal medicine to Press, Honolulu, HI. 924 p.
treat fever, headaches, hemorrhages, convulsions,
acute cough, skin parasites, and boils (HealthLink Opler, P.A. and V. Malikul. 1992. A field guide to
2001, Liogier 1990). Domestic and wild ruminant eastern butterflies. Peterson field guide 4.
animals browse wild indigo plants. It and another Houghton-Mifflin Co., Boston, MA. 396 p.
Indigofera species are reported to be the sole hosts
of the false dusky wing butterfly [Gesta gesta Pacific Island Ecosystems at Risk. 2001. Invasive
(Herrich-Schäffer)] (Opler and Malikul 1992). plant species: Indigofera suffruticosa Miller,
Wild indigo is useful as a natural cover plant in Fabaceae. http://www.hear.org/pier/insuf.htm.
disturbed areas. The species may become weedy, 2 p.
but is seldom aggressive or common enough to
cause difficulties in croplands. Simon, J.E., A.F. Chadwick, and L.E. Craker.
1984. Herbs: An indexed bibliography. 1971-
References 1980. Scientific literature on selected herbs, and
aromatic and medical plants of the Temperate
Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs Zone. Archon Books, Hamden, CT. 770 p.
and cacti of South Texas. Texas Tech University
Press. 213 p. Skousen, J.G. and C.A. Call. 1987. Grass and forb
species for revegetation of mixed soil-lignite
Haude, M.E. 1997. Identification and classification overburden in East Central Texas. Journal of
of colorants used during Mexico’s early colonial Soil and Water Conservation 42(6): 438-441.
period. Book and Paper Group Annual Vol. 16.
The American Institute of Conservation. Vélez, I. and J. van Overbeek. 1950. Plantas
http://aic.stanford.edu/conspec/bpg/annual/v16// indeseables en los cultivos tropicales. Editorial
bp16-05.html. 26 p. Univeresitaria, Río Piedras, PR. 497 p.
387
Isocoma tenuisecta Greene burro weed
ASTERACEAE
Juanita A. R. Ladyman
388
higher mortality rate than the Santa Rita three-awn and vegetation of the Rincon Mountains, Pima
(Blydenstein 1966, Cable 1967, McPherson 1995). County, Arizona. Desert Plants 8(2): 51-94.
Reproduction.—Burro weed flowers from August Cable, D.R. 1967. Fire effects on semidesert
into October (Martin and Hutchins 1980). The grasses and shrubs. Journal of Range
seeds (achenes) are partially covered by silky hairs Management 20: 170-176.
(Martin and Hutchins 1980). The long hairs that
comprise the pappus at one end of the seed aid Dayton, W.A. 1931. Important western browse
dispersal (Parker 1990). plants. Misc. Publ. 101. U.S. Department of
Agriculture. Washington, DC. 214 p.
Growth and Management.—Burro weed is a
resilient species that can flourish on highway Elmore, F.H. 1987. Shrubs and Trees of the
shoulders, parking lots, dirt roads, vacant lots and Southwest Uplands. Southwest Parks and
other disturbed areas. It tends to be sporadically Monuments Assoc. Tucson, AZ. 214 p.
scattered on range covered by perennial bunch
grasses. However, on disturbed ranges, with even Epple, A.O. 1995. A field guide to the plants of
a good stand of annual grasses, it can become one Arizona. LewAnn Publishing Company, Mesa,
of the principal plant species (USDA FS 1988). AZ. 347 p.
Burro weed foliage is poisonous to livestock
(AVDL 1998, Bowers 1993). Horses are most Goldberg, D.E. and R.M. Turner. 1986. Vegetation
sensitive while sheep are least sensitive. If change and plant demography in permanent
livestock eat large amounts of the vegetation they plots in the Sonoran Desert. Ecology 67(3): 695-
suffer from “trembles,” “alkali sickness,” or “milk- 712.
sickness” and humans can become ill after
drinking the milk from affected cows (Dayton Ivey, R.D. 1995. Flowering plants of New
1931, Epple 1995). The toxic substance Mexico. 3rd Ed. Rio Rancho Printing,
transmitted by the milk is called tremetol (AVDL Albuquerque, NM. 504 p.
1998, Kingsbury 1964). Many other species of
Isocoma contain the same toxic constituent Kearney, T.H., R.H. Peebles, and collaborators.
(AVDL 1998, Kingsbury 1964). As a consequence 1960. Arizona flora. 2nd Ed. University of
of its toxic properties, livestock avoid it unless California Press, Berkeley, CA. 1,085 p.
there is nothing else to eat. In overgrazed land
burro weed behaves as an invasive weed Kingsbury, J.M. 1964. Poisonous Plants of the
(Whitfield and Anderson 1938, Parker 1990). United States and Canada, Prentice-Hall, Inc.,
Englewood Cliffs, NJ. 626 p.
Benefits.—Burro weed provides cover and
stabilizes soils that are denuded by overgrazing or Lane, M.A. and R.L. Hartman. 1996.
other disturbance. It adds to the biodiversity of Reclassification of North American
communities of which it is part and may be Haplopappus (Compositae: Asteraceae)
important for pollinators and other arthropods in Completed: Rayjacksonia gen. nov. American
areas that are depauperate in native species. Journal of Botany 83(3): 356-370.
389
Parker, K.F. 1990. An illustrated guide to Arizona Taylor, R.J. 1998. Desert wildflowers of North
weeds. The University of Arizona Press, America. Mountain Press Publishing Co.,
Tucson, AZ. 338 p. Missoula, MT. 349 p.
Pima Community College. 2001. Internet site. United States Department of Agriculture, Forest
http://wc.pima.edu/Bfiero/tucsonecology/plants/ Service. 1988. Range Plant Handbook. Dover
shrubs_bw.htm [not paged]. Publications, Inc. New York. 837 p.
Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox, White, L.D. 1969. Effects of a wildfire on several
J.O. Klemmedson, J.J. Norris, and L.C. Fierro. desert grassland shrub species. Journal of Range
1992. Desert grasslands. In: R.T. Coupland, ed. Management 22: 284-285.
Ecosystems of the World, 8A; Natural
grasslands: Introduction and Western Whitfield, C.J and H.L. Anderson. 1938.
Hemisphere. Elsevier, London, UK. p. 337-362. Secondary succession in the desert plains
grassland. Ecology 19 (2): 171-180.
390
Itea virginica L. Virginia sweetspire
SAXIFRAGACEAE
Synonyms: None
Kristina Connor
391
Growth and Management.—Virginia sweetspire Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope.
is semievergreen and fairly long lived. With its 1996. Plants for American Landscapes. Louisiana
long flowering period and attractive fall foliage, State Univ. Press, Baton Rouge, LA. 266 p.
the species can do well in landscape plantings. Its
adaptability to both sun and shade, and to wet and Ohio State University. 2002. Horticulture and Crop
dry areas, gives it a versatility lacking in some Science in Virtual Perspective.
other shrub species. However, it can grow beyond http://www.hcs.ohio-state.edu/hcs/TMI/
desired boundaries because it spreads by root Plantlist/it_inica.html. 4 p.
suckers, and if not pruned it will form thickets. It
grows fairly rapidly and easily from basal shoots Seiler, J.R. and J.A. Peterson. 2001. Sweetspire.
and is relatively free from insect pests and diseases Virginia Tech. Dendrology Webpage
(The Ohio State University 2002). http://www.fw.vt.edu/
dendro/dendrology/syllabus/ivirginica.htm. 1 p.
Benefits.—The dense thickets of Virginia
sweetspire provide cover for wildlife (Silberhorn Silberhorn, G. 1996. Virginia willow (Itea
1996). Although the dry fruit capsules and small virginica). Technical Report. Wetlands Flora. No.
seeds are not a preferred food, they are eaten by 96-5. Virginia Institute of Science, School of
birds. The species can be used for erosion control Marine Science, College of William and Mary,
on wet or sloped sites. The fragrant flowers are Gloucester Point, VA. 2 p.
attractive to butterflies.
U.S. Department of Agriculture. [no date].
References Southern wetland flora: Field office guide to
plant species. U. S. Department of Agriculture,
Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: A Soil Conservation Service, South National
Concise Dictionary of Plants Cultivated in the Technical Center, Fort Worth, TX. Jamestown,
United States and Canada. McMillan Publishing ND: Northern Prairie Wildlife Research Center
Co., Inc., New York. 1,312 p. Home Page. http://www.npwrc.usgs.gov. [not
paged].
Johnson, F.L. and B.W. Hoagland. 1999. Itea
virginica. Catalog of the Woody Plants of USDA-ARS-GRIN. 2001. Germplasm Resources
Oklahoma, Oklahoma Biological Survey. Information Network, National Germplasm
http://www.biosurvey.ou.edu. 1 p. Resources Laboratory. Plant Sciences Institute,
Beltsville, Agricultural Research Center,
Krüssmann, G. 1976. Manual of Cultivated Broad- Agricultural Research Service, U.S. Department of
Leaved Trees and Shrubs. Volume I, A-D. Timber Agriculture. http://www.ars-grin.gov. [not paged].
Press, Beaverton, OR. 448 p.
USDA, NRCS. 2001. The PLANTS database,
Version 3.1. National Plant Data Center, Baton
Rouge, LA. http://plants.usda.gov. [not paged].
392
Ixora coccinea L. jungle-geranium
RUBIACEAE
John K. Francis
393
most popular tropical flowering shrubs. It is used Latha, P.G. and K.R. Panikkar. 2000. Inhibition of
for hedges and borders, for accent plants, in chemical carcinogenesis in mice by Ixora
planters and as an indoor potted plant. The species coccinea flowers. Pharmaceutical Biology 38(2):
may also be shaped into bonsais (Frommer 2001). 152-156.
In laboratory tests, extracts of jungle-geranium
have shown antibacterial (Kumer and others 1997) Liogier, H. A. 1997. Descriptive flora of Puerto
and antitumor (Latha and Panikkar 2000, Serrame Rico and adjacent islands. Vol. 5. Editorial de la
and Lim-Sylianco 1995) activity. From a methanol Universidad de Puerto Rico, San Juan, PR.
extract of the flowers, 13 chemicals were 436 p.
identified, including ursolic acid, which has known
antitumor and antiviral activity (Monteath and Michaelia, R., S. Philosoph-Hadasa, J. Riovb, and
others 2001). In the traditional medicine of India, S. Meira. 1999. Chilling-induced leaf abscission
infusions of the leaves and juice from the roots are of Ixora coccinea plants. I. Induction by
used to treat a wide variety of ailments including oxidative stress via increased sensitivity to
dysentery, ulcers, and gonorrhea (Parrotta 2001). ethylene. Physiologia Plantarum 107(2): 166-
The flowers are visited by butterflies (Collins 173.
2001), and birds eat the fruits.
Monteath, S.A.F.A., V.V. Veiga, A.C. Pinto, A.
References Echevarria, and M.A.M. Maciel. 2001.
Constituentes químicos das flores de Ixora
Bailey, L.H. 1941. The standard cyclopedia of coccinea. Universidade Federal Rural do Rio de
horticulture. The MacMillan Company, New Janeiro. http://www.sbq.org.br/ranteriores/23/
York. 3,639 p. resumos /1147. 3 p.
Collins, M. 2001. Marge Collins flying flowers: Parrotta, J.A. 2001. Healing plants of peninsular
Marge’s story. http://www.ffgc.org/horticulture India. CAB International, Wallingford, UK and
/flgardens/collins/garden_collins.htm. 4 p. New York. 944 p.
Florida Gardener. 2001. Plant of the month: Ixora Serrame, E. and C.Y. Lim-Sylianco. 1995. Anti-
coccinea. http://www.floridagardener.com tumor promoting activity of decoctions and
/ixora.htm. 2 p. expressed juices from Philippine medicinal
plants. Philippine Journal of Science 124(3):
Frommer A. 2001. Growing an indoor bonsai. 275-281.
Arthur Frommer’s Budget Travel Newsletter.
http://www.dummies.com/Lifestyle/ Tropilab. 2001. Ixora coccinea L. Tropilab Inc.
Home_and_Garden/Healthy…/0-7645-5102- http://www.tropilab.com/jungle-ger.html. 1 p.
7_0019.htm. 4 p.
Watkins, J.V. and T.J. Sheeham. 1975. Florida
Griffiths, M. 1994. Index of garden plants. Timber landscape plants, native and exotic. The
Press, Portland, OR. 1,234 p. University Presses of Florida, Gainesville, FL.
420 p.
Kumer, T.K., C.E. Rajini, V.K. Sasidharan, and T.
Krishna-Kumar. 1997. Antibacterial and Whistler, W.A. 2000. Tropical ornamentals, a
antifungal activity of secondary metabolites guide. Timber Press, Portland, OR. 542 p.
from some medicinal and other common plant
species. Journal of Life Science 2: 14-19.
394
Jacquinia arborea Vahl barbasco
THEOPHRASTACEAE
John K. Francis
from the J. armillaris group: J. keyensis Mez from
the Northern West Indies (Florida and the
Bahamas), J. arborea in the Central Caribbean, and
J. armillaris Jacq. in the Lesser Antilles, northern
South America, and eastern Brazil. Stearn (1992)
gives the range of barbasco as the Cayman Islands,
Jamaica, Hispaniola, Puerto Rico, the Virgin
Islands, the Lesser Antilles, Trinidad, and Curaçao.
He has J. arborea and J. armillaris coexisting in
the Lesser Antilles. The Missouri Botanical Garden
Herbarium lists specimens for Jamaica, Puerto
Rico, the Virgin Islands, Quintana Roo, Mexico,
and Honduras (Missouri Botanical Garden 2002).
Barbasco is reported to have naturalized on Key
Largo, Florida (Nelson 1996).
395
Growth and Management.—Growth of barbasco Liogier, H.A. 1995. Descriptive flora of Puerto
seedlings in the nursery is very slow. A group of Rico and adjacent islands. Vol. 4. Editorial de la
seedlings averaged only 6.2 cm 1 year after Universidad de Puerto Rico, San Juan, PR.
pricking into containers. However, survival was 617 p.
relatively good (82 percent). Growth of native-
grown plants of all ages is also believed to be very Little, E.L., Jr., R.O. Woodbury, and F.H.
slow. They seem to be long-lived. Probably the Wadsworth. 1974. Trees of Puerto Rico and the
best strategy for management of native stands is Virgin Islands. Vol. 2. Agriculture Handbook
strict protection from fire, harvesting, and 449. U.S. Department of Agriculture,
development. Washington, DC. 1,024 p.
Benefits.—Barbasco helps hold the soil in Missouri Botanical Garden. 2002. Current
topography where there is little soil to lose and specimen list for Jacquinia arborea.
contributes significantly to the aesthetics of island http://mobot.org/cgi-bin/search_vast. 1 p.
coastlines. The fruits were once used to poison or
stupefy fish and because of that are suspected to be Nelson, G. 1996. The shrubs and woody vines of
poisonous to humans (Little and others 1974). The Florida. Pineapple Press, Inc., Sarasota, FL.
sister species J. keyensis is used as an ornamental 391 p.
(Nelson 1996), and probably barbasco has been
used as well. It is a pretty and well-shaped plant, Stahl, B. 1992. On the identity of Jaquinia
suited for borders and low backgrounds. armillaris (Theophrastaceae) and related
species. Brittonia 44 (1): 54-60.
References
Stearn, W.T. 1992. The genus Jacquinia
Howard, R. A. 1989. Flora of the Lesser Antilles, (Theophrastaceae) in Jamaica. Nordic Journal of
Leeward and Windward Islands. Vol. 6. Arnold Botany 12(2): 231-238.
Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
396
Jasminum fluminense Vell. Brazilian jasmine
OLEACEAE
John K. Francis
397
means of cuttings (Florida Exotic Plant Council Bailey, L.H. 1941. The standard cyclopedia of
2001). horticulture. The MacMillan Company, New
York, NY. 3,639 p.
Growth and Management.—The stems of
established plants may extend as much as 2 or 3 m Florida Exotic Plant Council. 2001. Jasminum
in 1 year. Seedlings grow much more slowly. fluminense Vell. www.fleppc.
Brazilian jasmine can engulf shrubs, small trees, org/pdf/Jasminum%20fluminense.pdf. 2 p.
and fences and can ascend power poles. Control of
thickets and mats is often needed. Young plants Howard, R.A. 1989. Flora of the Lesser Antilles,
can be pulled up by hand. Older plants should be Leeward and Windward Islands. Dicotyledoneae.
cut at the ground level and the stumps treated with Vol. 6. Arnold Arboretum, Harvard University,
herbicide. Follow-up treatments will probably be Jamaica Plain, MA. 658 p.
required (Pacific Island Ecosystems at Risk 2001).
Liogier, H.A. 1995. Descriptive flora of Puerto
Benefits.—Brazilian jasmine has been widely Rico and adjacent islands. Vol. 4. Editorial de la
planted as an ornamental, both in greenhouses and Universidad de Puerto Rico, San Juan, PR. 617
outdoors (Bailey 1941, Liogier 1995). Because of p.
its invasive nature, the species should not be
planted where it might escape to the wild (Nelson Miami-Dade County. 2001. Brazilian jasmine-
1996). It is a food source for birds and mammals. Jasminum fluminense. www.co.miami-
The fruits are heavily consumed by raccoons in dade.fl.us/derm/environment/badplants/plant%2
Florida (Miami-Dade County 2001). Goats browse 0…/brazilian_jasmine.ht. 1 p.
the foliage.
Nelson, Gil. 1996. The shrubs and woody vines of
References Florida. Pineapple Press, Inc. Sarasota, FL.
391 p.
Acevedo-Rodríguez, P. 1985. Los bejucos de
Puerto Rico. Vol. 1. General Technical Report Pacific Island Ecosystems at Rick. 2001. Invasive
SO-58. U.S. Department of Agriculture, Forest plant species: Jasminum fluminense Vell.,
Service, Southern Forest Experiment Station, Oleaceae. www.hear.org/pier/jaflu.htm. 2 p.
New Orleans, LA. 331 p.
398
Jatropha gossypiifolia L. red physic nut
EUPHORBIACEAE
John K. Francis
399
considered an agricultural weed but can be Horsten, S.F., A.J. van den Berg, J.J. Kettenes-van
suppressed by common weed-control practices. den Bosch, B.R. Leeflang, and R.P. Lababie.
1996. Cyclogossine A: a novel cyclic
Detriments and Benefits.—Red physic nut heptapeptide isolated from the latex of Jatropha
contains toxins in the seeds, sap and other tissues, gossypifolia. Planta Med 62(1): 46-50.
capable of killing humans (Marcano-Fondeur
2002). Apparently, poisoning cases are rare. Howard, R.A. 1989. Flora of the Lesser Antilles,
Extracts of the plant are used as a purgative and Leeward and Windward Islands. Vol. 5. Arnold
emetic, and to treat headache, diarrhea, venereal Arboretum, Harvard University, Jamaica Plain,
disease, skin sores, mouth sores, and cancer MA. 604 p.
(Burkill 1994, Parrotta 2001). The use of the seeds
in herbal medicine is advised against because of Liogier H.A. 1988. Descriptive flora of Puerto
their high toxicity (Liogier 1990). Phytoactive Rico and adjacent islands, Spermatophyta. Vol.
chemicals found in tissues of red physic nut 2. Editorial de la Universidad de Puerto Rico,
include the protease curcain, the alkaloid Río Piedras, PR. 481 p.
jatrophine, the cyclic heptapeptide, cyclogossine A,
saponins, and numerous fatty acids (Burkill 1994, Liogier, H.A. 1990. Plantas medicinales de Puerto
Horsten and others 1996, Marcano-Fondeur 1992, Rico y del Caribe. Iberoamericana de Ediciones,
Ogbobe and Akano 1993). The seeds contain 36 Inc., San Juan, PR. 566 p.
percent oil (Ogbobe and Akano 1993) and are used
as an illuminant in Africa (Burkill 1994). Swaths of Marcano-Fondeur, E. de J. 1992. Las plantas
plants are frequently established around villages in venenosas en la medicina popular. Naturaleza
Africa as fire barriers. The species is also attributed Dominicana. http://marcano.freeservers.com/
magical powers that protect against lightning, nature/conference/venen.html. 6 p.
snakes, and violence (Burkill 1994). The wood is
soft and weak and of little use. A red-leafed Ogbobe, O. and V. Akano. 1993. The physico-
variety is used as an ornamental (Howard 1989). chemical properties of the seed and seed oil of
Jatropha gossipifolia. Plant Foods for Human
References Nutrition 43(3): 197-200.
Burkill, H.M. 1994. The useful plants of West Parrotta, J.A. 2001. Healing plants of Peninsular
Tropical Africa. Vol. 2. Royal Botanic Gardens, India. CABI Publishing, Wallingford, UK and
Kew, UK. 636 p. New York. 917 p.
400
Juniperus communis L. common juniper
CUPRESSACEAE
Juanita A. R. Ladyman
401
climates (Lanzara and Pizzetti 1977). Male and Benefits.—Common juniper has low value for
female cones are on separate plants. The dark blue short-term re-vegetation projects but moderate to
seed (female) cones are globose to ovoid in shape, high value for long-term rehabilitation projects
6 to 13 mm in diameter and are resinous to and is useful in preventing soil erosion
obscurely woody with two to three triangular- (Tirmenstein 1999). Common juniper provides
shaped seeds. From pale green the female cones important cover and browse for wildlife such as
ripen to dark blue in late summer and then tend to mule deer (Dusek 1975). The cones are eaten by
whiten with a waxy “bloom.” The staminate cones several species of birds and are important food
are 3 to 6 mm in diameter (Cronquist and others sources for wild turkeys (Decker and others 1991),
1972). Although the cones are generally terminal Cedar and Bohemian waxwings (Catling and
in other species of juniper, in J. communis the Brownell 1998), and others. Domestic livestock
cones are axillary (FNA 1993). The fruits are of rarely utilize common juniper (Tirmenstein 1999).
relatively low nutritional quality and on average The seed cones of common juniper are used to
number 80,500 seeds/kg (Stiles 1980). The seed is flavor gin, and the word for this alcoholic
dispersed by birds, at least in North America, and beverage was derived from the Latin “iuiperus”
commonly require a long maturation and through the Old French and Dutch words
germination period (Crane and Fischer 1986; Stiles “genevre” and “genever” respectively (Elmore
1980). Because seeds do not readily germinate, 1987). The seed cones are also used as a flavoring
the establishment of seedlings, for example for re- for foods. Many Native American tribes used
vegetation projects, can be a problem (Dietz and common juniper for medicinal and ceremonial
others 1980, Stiles 1980). As well as poor purposes (Moerman 1998). It has been used
germination, poor seed dispersal has been blamed medicinally for childbirth and to treat urinary
for the relative rarity of juniper in areas where the infections (Pojar and Mackinnon 1994). However,
majority of potential habitat is not occupied it can also cause miscarriages. The different
(Diotte and Bergeron 1989). Plant age also affects varieties of common juniper make excellent,
reproduction; 40 to 60 percent of old individuals vigorous landscaping shrubs, which are readily
are sterile and up to 94.8 percent of seeds of seed- propagated by cuttings in the horticultural trade
producing old plants are non-viable. In contrast, 80 (Cronquist and others 1972).
percent of seeds produced by young plants are
viable (Diotte and Bergeron 1989). References
402
turkeys. Canadian Journal of Zoology. 69 (8): Mallik, A. U. 1995. Conversion of temperate
2,128-2,132. forests into heaths: role of ecosystem
disturbance and ericaceous plants.
Dietz, R.D., D.W. Uresk, H.E. Messner, and L.C. Environmental Management. 19(5): 675-684.
McEwen. 1980. Establishment, Survival, and
Growth of Selected Browse Species in a Moerman, D.E. 1998. Native American
Ponderosa Pine Forest. General Technical Ethnobotany. Timber Press, Inc, Portland, OR.
Report RM-219. USDA Forest Service Rocky 927 p.
Mountain Forest and Range Experiment Station.
Fort Collins, CO. 12 p Pojar, J. and A. Mackinnon. 1994. Plants of the
Pacific Northwest Coast. British Columbia
Diotte, M. and Y. Bergeron. 1989. Fire and the Ministry of Forests and Lone Pine Publishing,
distribution of Juniperus communis L. in the Vancouver, Canada. 527 p.
boreal forest of Quebec, Canada. Journal of
Biogeography 16: 91-96. Stace, C. 1997. New Flora of the British Isles.
Second ed. Cambride University Press,
Dusek, L. 1975. Range relations of Mule Deer and Cambridge, England. 1,130 p.
cattle in prairie habitat. Journal of Wildlife
Management. 39(3): 605-616.
Steele, F.L. 1982. At Timberline: A Nature Guide
to the Mountains of the Northeast. Appalachian
Elmore. F.H. 1987. Shrubs and Trees of the
Mountain Club, Boston, MA. 285 p.
Southwest Uplands. Southwest Parks and
Monuments Assoc. Tucson, AZ. 214 p.
South Carolina Heritage Trust. 2000. Website.
http://www.abi.org/nhp/us/sc/speclist.htm. 1 p.
Flora of North America Editorial Committee.
1993. Flora of North America. Vol. 2. Oxford
Stiles, E.W. 1980. Patterns of fruit presentation and
University Press, New York. 475 p. seed dispersal in bird-disseminated woody plants
in the eastern deciduous forest. The American
Georgia Natural Heritage Program. 2001. Naturalist. 116(5): 670-688.
http://www.dnr.state.ga.us/dnr/wild/natural/sppl
_t.htm. 1 p Tirmenstein 1999. Juniperus communis. U.S.
Department of Agriculture, Forest Service,
Kartesz, J.T. 1994. A synonymized checklist of the Rocky Mountain Research Station, Fire Sciences
vascular flora of the United States, Canada, and Laboratory. Fire Effects Information System,
Greenland. Vol.1 – Checklist. 2nd Ed. Timber http://www.fs.fed.us/database/feis/ [not paged].
Press, Portland, OR. 622 p.
Ward, L.K. 1982. The Conservation of Juniper.
Lanzara, P. and M. Pizzetti. 1977. Simon and Journal of Applied Ecology. 19: 165-188.
Schuster’s Guide to Trees. Simon and Schuster
Inc., New York. 316 p.
403
Juniperus osteosperma (Torr.) Little Utah juniper
CYPRESSACEAE
John K. Francis
404
commonly found on rocky, gravely, and sandy Growth and Management.—Utah juniper grows
sites, usually originating from igneous (Stuart and slowly, usually only 0.13 cm/year in diameter. An
Sawyer 2001) and sedimentary (sandstone and 86-year-old stand in Utah had 17.2 percent canopy
limestone) rocks, at 900 to 2,500 m above sea cover and a basal area of 7.7 m2/ha (Zlatnik 1999).
level (Zlatnik 1999). Mean annual precipitation Poor site conditions may result in 15 cm-tall plants
varies from 300 to 510 mm, but the species can over 50 years old. Individuals of the species may
withstand severe droughts (Natural Resources live 650 years or longer. Closing Utah juniper
Conservation Service 2003) and is a fierce stands eliminate most forage plants; killing trees
competitor for the little available moisture in its can result in a doubling of herbaceous cover
habitat. It has a specific leaf area of 18 cm2/g (Zlatnik 1999). Controlling Utah juniper has been
(Grier and others 1992). Utah juniper also tolerates attempted with fire, but fires on these sites are
cold and heat well. However, the species is difficult to start under humid, still conditions and
intolerant of shade and does not survive under the hard to control under dry, windy conditions.
canopy of taller vegetation. It is alleged to exert an Controlled burns can be done with the greatest
allelopathic effect on some understory grass precision in late winter or early spring when the
species. Utah juniper is a climax species in many wood is low in water content (personal
vegetation types with species such as Artimisia communication with Stanley Kitchen, Shrub
tridentata Nutt., Pinus edulis Engelm., and Pinus Sciences Laboratory, USDA Forest Service,
monophylla Torr. & Frém. The range of Utah Provo, UT). The shrubs and trees can be killed
juniper is known to have expanded since European with chainsaws or herbicides, but the preferred
colonization, and stands have changed from open, method is “chaining,” dragging sections of anchor
savanna types to dense stands probably because of chain between two bulldozers. Many young trees
reduction of fine fuels by livestock grazing, fire are not killed, but the treatment does open stands
suppression, and changes toward warmer and drier and allows greater shrub and herbaceous growth
climates (Tausch 1999). Utah juniper is easily (Plummer and others 1968, Stevens 1999).
killed by fire, both by scorching the top or heat-
girdling through the thin bark. However, the Benefits.—Utah juniper is a major constituent of
habitat seldom has sufficient fine fuels to sustain a semidry ecosystems of the West. It protects the
fire. Stands most at risk are young stands with soil from wind erosion but in some instances may
scattered trees in brush and grass, and continuous aggravate sheet erosion from rains by suppressing
stands of large, decadent trees during high winds understory grasses, forbs, and shrubs. The species
(Zlatnik 1999). Utah juniper is sometimes heavily furnishes cover and reproductive habitat for many
infested by the mistletoes, Phoradendron species and is a major source of food for wildlife,
juniperinum Engelm. ex Gray and P. bolleanum especially during winter. Many birds, rodents, as
(Seem.) Eichl. (Zlatnik 1999). The larvae of well as jackrabbits (Lepus californicus) and
Eurytoma juniperina Marcovitch attack the seeds coyotes (Canis latrans) eat the fruits and mule
(Bonner 2003). deer (Odocoileus hemionus), and elk (Cervus
elaphus) browse the foliage when better food is
Reproduction.—Utah juniper flowers from not available. It is not eaten by cattle and is
January to April (depending on site) and is wind browsed to a limited extent by domestic sheep
pollinated. Fruits ripen in the second year and (Stuart and Sawyer 2001, Zlatnik 1999). The
germinate the following spring or the second forage is of low quality because of low
spring after dispersal (Zlatnik 1999). Fruits may be digestibility (44 percent) and low protein content
collected by hand from the shrubs or low trees or (6 to 8 percent) (Zlatnik 1999). The wood is used
picked up by hand after they have fallen. Seed for firewood, charcoal, and novelties (Secretaría de
should be extracted by maceration with water Medio Ambiente y Recursos Naturales 2003). The
before the fruits dry out. Cleaned seed from bark is used as tinder and formerly was used as a
Arizona ranged from 7,900 to 15,700 seeds/kg. “slow match” to maintain and transport fire (Plants
Seed should be dried to about 10 percent moisture For a Future 2003). Native Americans used the
and can be stored at 5 to -18 °C for several years. bark to make sandals, mats, bedding, baskets,
Moist stratification at 3 to 5 °C for 30 to 180 days thatch, and cordage, and they made meal from the
is necessary for good germination. Drilled or fruits (Peattie 1953). The wood is highly decay
broadcast in a nursery bed, seeds should be resistant, which makes it desirable for fence posts
covered with 6 mm of fine soil or sand. and rude construction despite its poor form
Germination ranges from 50 to 64 percent (Bonner (Zlatnik 1999). Tissues of Utah juniper are not
2003). high enough in cedarwood oil to be commercially
405
valuable. Heartwood contains 1.19 percent oil, Utah. Publication 68-3. Utah Division of Fish
principally thujopsene (40 percent) and cedrol and Game, Salt Lake City, UT. 183 p.
(13.2 percent) (Adams 1987). The species was
used by Native Americans to treat bladder and Secretaría de Medio Ambiente y Recursos
kidney troubles, wounds, and mouth sores (Plants Naturales. 2003. Especies con usos no
for a Future 2003). Utah juniper has been used to a maderables en bosques de Encino, pino y pino-
limited extent in revegetation projects and encino: Juniperus osteosperma (Torr.) Little.
xeriscape gardening. http://www.semarnat.gob.mx/pfnm/Juniperus
Osteosperma.html. 2 p.
References
Stevens, R. 1999. Mechanical chaining and
Abrams, L. 1940. Illustrated flora of the Pacific seeding. In: S.B. Monson and R. Stevens,
States. Vol. 1. Stanford University Press, comps. Proceedings: ecology and management
Stanford, CA. 538 p. of pinyon-juniper communities within the
Interior West; Sep. 15-18, 1997. Provo, UT.
Adams, R.P. 1987. Investigation of Juniperus PMRS-P-9. U.S. Department of Agriculture,
species of the United States for new sources of Forest Service, Rocky Mountain Research
cedarwood oil. Economic Botany 41(1): 48-54. Station, Ogden, UT. p. 281-284.
American Forests. 1998. National register of big Stuart, J.D., and J.O. Sawyer. 2001. Trees and
trees 1998-1999. American Forests 104(1): 24- shrubs of California. University of California
46. Press, Berkeley, CA. 467 p.
Bonner, F.T. 2003. Juniperus L., juniper. In: F.T. Tausch, R.J. 1999. Historic pinyon and juniper
Bonner and R.G. Nisley, eds. Woody Plant Seed woodland development. In: S.B. Monson and R.
Manual. U.S. Department of Agriculture, Forest Stevens, comps. Ecology and management of
Service, Washington, DC. 15 p. pinyon-juniper communities within the Interior
West. Proc. RMRS-P-9. U.S. Department of
Grier, C.C., K.J. Elliot, and D.G. McCullough. Agriculture, Forest Service, Rocky Mountain
1992. Biomass distribution and productivity of Research Station, Ogden, UT. p. 12-19.
Pinus edulis-Juniperus monosperma woodlands
of north-central Arizona. Forest Ecology and Terry, R.G., R.S. Nowark, and R.J. Tausch. 2000.
Management 5093-4): 331-350. Genetic variation in chloroplast and nuclear
ribosomal DNA in Utah juniper (Juniperus
Natural Resources Conservation Service. 2003. osteosperma, Cupressaceae): evidence of
Plants profile: Juniperus osteosperma (Torr.) interspecific gene flow. American Journal of
Little, Utah juniper. http://plants.usda.gov/ Botany 87: 250-258.
cgi_bin/plant_search.cgi?mode=Scientific+Nave
&keywordquery=Juniperus+osteosperma&earl= Tree Guide. 2003. Treeguide species details: Utah
plant_sea. 5 p. juniper. http://www.treeguide.com/Species.
asp?SpeciesID=544. 3 p.
Peattie, D.C. 1953. A natural history of Western
trees. Houghton Mifflin Company, Boston, MS. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
749 p. Higgins, eds. 1987. A Utah flora. Brigham
Young University, Provo, UT. 894 p.
Plants For a Future. 2003. Plants for a future:
database search results: Juniperus osteosperma. Zlatnik, E. 1999. Juniperus osteosperma. In: U.S.
Plants for a Future, Cornwall, UK. http://www. Department of Agriculture, Forest Service,
ibiblio.org/pfaf/cgi-bin/arr_html?Juniperus+ Rocky Mountain Research Station, Fire Sciences
osteosperma&CAN=LATIND. 8 p. Laboratory. Fire Effects Information System.
http://www.fs.fed.us/database/feis/plants/tree/
Plummer, A.P., D.R. Christensen, and S.B. junost/all.html. 33 p.
Monson. 1968. Restoring big-game range in
406
Kalmia latifolia L. mountain laurel
ERICACEAE
407
by reducing light and increasing competition for Benefits and Detriments.―Mountain laurel has
water. long been cultivated as an ornamental--since about
1740 when American naturalist John Bartram sent
Reproduction.―The showy, white to pink 2- to plants to England (Jaynes 1997). Cultivars of
3- cm saucer-shaped flowers form convex clusters mountain laurel are particularly valuable as
at the ends of branches and appear after the ornamentals and are propagated commercially by
foliage, from March to July, depending on latitude many nurseries. Nearly 100 horticultural varieties
and altitude (Olson and Barnes 1974). Purple spots of have been selected for variation in flower color,
mark 10 anther pockets in the petals. The method leaf variegation, and growth habit (Jaynes 1997).
of pollen dispersal in Kalmia is unusual in that the Mountain laurel burls have been used for making
anthers are under tension as the flower matures, pipes for smoking tobacco, and early European
and they spring forward when disturbed by an settlers in America used the fine-grained wood for
insect, forcefully and effectively transferring making eating utensils and weaver's shuttles, and
grains of pollen to the insect (Jaynes 1997). The produced a yellow dye from its foliage. Tinctures
flowers are pollinated by more than a dozen of fresh Kalmia leaves have been used for home
insects, primarily bumblebees, but generally not medical remedies of skin disorders. An
honeybees, likely because the flowers produce unidentified anticancer compound has been
little nectar (Jaynes 1997). Glandular hairs on the extracted from its sap (Jaynes 1997). Mountain
flower stalk, calyx, and corolla may deter laurel provides cover and stability to thin soils,
nonflying insects. Flowers are most abundant on however its presence can reduce water yields
plants in full sunlight and occur rarely on plants in (Johnson and Kovner 1956). Foliage of mountain
dense shade (Kurmes 1961). Fruit ripens in late laurel is toxic to domestic cattle and sheep but
summer and consists of a brown, erect, five-celled, provides subsistence winter food for ruffed grouse
globular capsule, 4 to 7 mm wide. Each capsule (Bonasa umbellus L.) and white-tailed deer
contains about 600 seeds, which are 0.8 to 1 mm (Odocoileus virginianus Boddaert) (Robinette
long and average about 27 million per kilogram 1974). Foliage from new sprouts of burned
(Olson and Barnes 1974). The nonwinged seeds mountain laurel provides greater potential benefit
are dispersed by wind, but only for distances of to wildlife diets compared to unburned foliage
less than 15 m (Robinette 1974). Seeds remain (Thackston and others 1982).
viable for more than a year under field conditions.
Light, cold stratification, and moisture enhance Fire.―Mountain laurel contributes to intensity of
germination (Olson and Barnes 1974). Seeds fires in mountainous terrain and is classified as
germinate on mineral soil and on moss (Robinette "extreme" in rate of spread when it occurs in dense
1974). Kalmia may also regenerate by layering, thickets beneath hardwood stands (Jemison and
sprouts, or suckers (Robinette 1974). Techniques Keetch 1942). Flame length can exceed 30 m in
for commercial production of mountain laurel fires burning in Table Mountain pine (Pinus
plants have been well developed (Jaynes 1997). pungens Lambert) stands with mountain laurel
(Waldrop and Brose 1999). Moisture of Kalmia
Growth and Management.―Growth rate of foliage is lower than other species and is little
mountain laurel is relatively slow; young plants (< affected by seasonal changes of soil moisture
15 years) add about 12 cm in height and 9 cm in (Reifsnyder 1961). The high flammability of
width annually, depending on site moisture Mountain laurel thickets results from low foliate
relations (Monk and others 1985). Older mountain moisture (Richards 1940) and a high proportion of
laurel stems may attain heights up to 9 m and total biomass in small (<5 mm) dead twigs.
diameter at ground level of 15 cm (Jaynes 1997). Caloric content of foliage is estimated at about
Mountain laurel stem density can range from 5,000 cal/g based on similar vegetation (Hough
sparse on mesic sites to over 26,000 per ha in 1969). Allometric equations for estimating dry
thickets on upper, southwest slopes; basal area at 3 biomass of 6- to 8-year old open-grown mountain
cm above ground level can exceed 25 m2/ha. laurel stems were developed by Boring and Swank
Herbicides may be used to control mountain laurel (1986), which can be used to predict fuel loadings
on sites where establishment of tree seedlings is of leaf (log10Y = -0.754 + 1.882 log10X), branch
desired (Neary and others 1984). It is sensitive to (log10Y = -1.222 + 2.359 log10X), and bole (log10Y
juglone, a substance produced by black walnut = -0.636 + 1.948 log10X) components of individual
(Juglans nigra L.) trees. Mountain laurel generally plants, where Y is dry weight (g) and X is stem
does not present a problem to timber management diameter (mm) at 3 cm from the ground. Kalmia is
activities on submesic to mesic sites. killed more readily by headfires than by backfires
408
(Hooper 1969), but almost all topkilled stems Hooper, R.M. 1969. Prescribed burning for laurel
recover by producing basal sprouts (Clinton and and rhododentrol control in the southern
others 1993, Johnson and Kovner 1956). Equations Appalachians. Research Note SE-116. U.S.
for estimating stem, foliage, and total biomass of Department of Agriculture, Forest Service,
young mountain laurel sprouts on recently burned Southeastern Forest Experiment Station,
areas were developed by Elliott and Clinton Asheville, NC. 6 p.
(1993).
Hough, W.A. 1969. Caloric value of some forest
References fuels of the southern United States. Research
Note SE-120. U.S. Department of Agriculture,
Al-Hamdani, S.H., P.B. Nichols, and G.R. Cline. Forest Service, Southeastern Forest Experiment
2002. Seasonal changes in the spectral Station, Asheville, NC. 6 p.
properties of mountain laurel (Kalmia latifolia
L., Ericaceae) in north east Alabama. Castanea. Jaynes, R.A. 1997. Kalmia--Mountain laurel and
67(1): 25-32. related species. Timber Press, Portland, OR.
295 p.
Boring, L.R. and W.T. Swank. 1986. Hardwood
biomass and net primary production following Jemison, G.M. and J.J. Keetch. 1942. Rate of
clearcutting in the Coweeta basin. In: Brooks, spread of fire and its resistance to control in the
R.T., Jr. ed. Proceedings of the 1986 southern fuel types of eastern mountain forests. Technical
forest biomass workshop. Tennessee Valley Note 2. U.S. Department of Agriculture, Forest
Authority, Knoxville, TN: 43-50. Service, Appalachian Forest Experiment Station,
Asheville, NC. 15 p.
Bryson, G.M. and A.V. Barker. 2002. Sodium
accumulation in soils and plants along Johnson, E.A. and J.L. Kovner. 1956. Effect on
Massachusetts roadsides. Communications in streamflow of cutting a forest understory. Forest
Soil Science and Plant Analysis. 33:67-78. Science. 2: 82-91.
Chapman, G.L. 1950. The influence of mountain Kurmes, E.A. 1961. The ecology of mountain
laurel undergrowth on environmental conditions laurel in southern New England. Yale
and oak reproduction. Doctoral dissertation. University, New Haven, CT. 85 p. Doctoral
Yale University, New Haven, CT. Dissertation.
Clinton, B.D., J.M Vose, and W.T. Swank. 1993. Lipscomb, M.V. and Nilsen, E.T. 1990.
Site preparation burning to improve southern Environmental and physiological factors
Appalachian pine-hardwood stands: vegetation influencing the natural distribution of evergreen
composition and diversity of 13-year-old stands. and deciduous ericaceous shrubs on northeast
Canadian Journal Forest Research. 23: 2,271- and southwest slopes of the southern
2,277. Appalachian Mountains. II. Water relations.
American Journal Botany. 77(4): 517-526.
Ebinger, J.E. 1974. A systematic study of the
genus Kalmia (Ericaceae). Rhodora. 76: 315- McNab, W.H., S.A. Browning, S.A. Simon, P.E.
398. Fouts. 1999. An unconventional approach to
ecosystem unit classification in western North
Elliott, K.J. and B.D. Clinton. 1993. Equations for Carolina, USA. Forest Ecology and
estimating biomass of herbaceous and woody Management. 114: 405-420.
vegetation in early-succession Southern
Appalachian pine-hardwood forests. Research Monk, C.D., D.T. McGinty, and F.P. Day, Jr.
Note SE-365. U.S. Department of Agriculture, 1985. The ecological importance of Kalmia
Forest Service, Southeastern Forest Experiment latifolia and Rhododendron maximum in the
Station, Asheville, NC. 7 p. deciduous forest of the Southern Appalachians.
Bulletin Torrey Botanical Club. 112(2): 187-
Flemer,, W., III 1949. The propagation of Kalmia 193.
latifolia from seed. Bulletin of the Torrey
Botanical Club. 76: 12-16. Neary, D.G., J.E. Douglass, J.L. Ruehle, and W.
Fox. 1984. Converting rhododendron-laurel
409
thickets to white pine with picloram and Thackston, R.E., P.E. Hale, A.S. Johnson, and
mycorrhizae-inoculated seedlings. Southern M.J. Harris. 1982. Chemical composition of
Journal of Applied Forestry. 8(3): 163-168. mountain-laurel leaves from burned and
unburned sites. Journal Wildlife Management.
Olson, D.F., Jr. and R.L. Barnes. 1974. Kalmia 46(2): 492-496.
latifolia L., mountain laurel. In: C.S
Schopmeyer, ed. Seeds of woody plants in the Waldrop, T.A. and P.H. Brose. 1999. A
United States. Agriculture Handbook 450. U.S. comparison of fire intensity levels for stand
Department of Agriculture, Forest Service, replacement of table mountain pine (Pinus
Washington, DC. p. 470-471. pungens Lamb.). Forest Ecology and
Management. 113: 155-166.
Robinette, S.L. 1974. Mountain laurel-Kalmia
latifolia. In: J.D. Gill and W.M. Healy, eds. Waterman, J.R., A.R. Gillespie, J.M. Vose, and
Shrubs and vines for northeastern wildlife. W.T. Swank. 1995. The influence of mountain
General Technical Report NE-9. U.S. laurel on regeneration in pitch pine canopy gaps
Department Agriculture, Forest Service, of the Coweeta Basin, North Carolina, U.S.A.
Northeastern Forest Experiment Station, Radnor, Canadian Journal Forest Research. 25: 1,756-
PA. p. 102-105. 1,762.
410
Koanophyllon polyodon (Urban) King & H.E. Robins. polyodon
ASTERACEAE
John K. Francis
411
Krameria ixine L. abrojo colorado
EUPHORBIACEAE
John K. Francis
412
References Missouri Botanical Garden. 2002. Manual de la
Flora de Costa Rica. http://www.mobot.org/
Howard, R.A. 1988. Flora of the Lesser Antilles, manual.plantas/019706/S019706.html. 1 p.
Leeward and Windward Islands.
Dicotyledoneae, Part 1. Vol. 4. Arnold New York Botanical Garden. 20023. Specimens
Arboretum, Harvard University, Jamaica Plain, search results: Krameria ixine Loefl.
MA. 673 p. http://scis.../wwwspecimen.search_list?taxon=
Krameria+ixine+Loefl.+++++&projcode=VAS.
Instituto Nacional ce Biodiversidad. 2002. Lista de 1 p.
especímenes de Krameria ixine.
http://www.inbio.ac.cr/bims/k03/p13/c045/o013 Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
2/f01646/g008002/s024181.htm. 2 p. Montiel, eds. 2001. Flora de Nicaragua.
Monographs in Systematic Botany Vol. 85, No.
Liogier H.A. 1988. Descriptive flora of Puerto 2. Missouri Botanical Garden, St. Louis, MO.
Rico and adjacent islands, Spermatophyta. Vol. p. 945-1,910.
2. Editorial de la Universidad de Puerto Rico,
Río Piedras, PR. 481 p.
413
Lantana camara L. lantana
VERBENACEAE
John K. Francis
414
when ripe and contain one seed each. They are In some areas, competition by lantana results in a
eaten by birds and are widely scattered. If not reduction of biodiversity (Kumar and Rohatgi
eaten, they dry and remain on the shrub for weeks. 1999). Despite the establishment of a number of
A sample of seed collected in Puerto Rico natural enemies of lantana into exotic populations,
contained 100,700 seeds/kg of which 75 percent control of lantana populations has been usually
germinated within 7 weeks of sowing. Early limited or a failure (Day and others 1999, Hill
growth is rapid. Also, lantana can be propagated 1999). In thick stands, the shrub increases costs in
with cuttings and air layers. forest management by inhibiting access in stands
for thinning and felling, competes with
Growth and Management.—Ornamental plants reproduction, and increases fire hazards (Graaff
are established as potted seedlings. This is 1986). Lantana leaves contain poisonous
probably also the best method to establish plants triterpines and lantadenes A and B that cause death
for environmental protection projects such as of horses, cattle, sheep, goats, and rabbits by
reclamation of mine spoils. Lantana may reach 3 failure of the liver and other organs (Morton 1994,
m in height within 3 or 4 years. In naturalized Munyua and others 1990). However, most animals
habitat, the species often forms dense thickets. carefully avoid eating this plant when given a
These brush stands may be eliminated at choice. Green fruits also contain the poisons and
considerable expense from pastures and tree have caused illness and death in children (Morton
plantations by cutting and herbicide application. 1994). Lantana leaves and their leachates exert
The preferred methods are to cut the clumps and alelopathic effects in vetro and to a lesser extent in
drench the stumps with herbicide or to later spray soil on seed germination, root elongation, and
the tender regrowth with postemergence herbicide. plant growth of many species (Casado 1995, Sahid
Once established, tree plantations with closed and Sugau 1993).
canopies will keep lantana from reestablishing
itself. References
415
Herzog, F., Z. Farah, and R. Amado. 1994. popular garden flower; some cases of poisoning
Composition and consumption of gathered wild in Florida. Economic Botany 48(3): 259-270.
fruits in the V-Baoule, Cote d’Ivoire. Ecology of
Food and Nutrition 32(3-4): 181-196. Munyua, S.J.M., M.J. Nienga, T.P. Karitu, T.P.
Kimoro, J.E. Kiptoon, and I.B.J. Buoro. 1990. A
note on clinical-pathological findings and serum
Howard, R.A. 1989. Flora of the Lesser Antilles.
enzyme activity in sheep, goats and Friesian
Vol. 6. Arnold Arboretum, Harvard University,
calves with acute Lantana camara poisoning.
Jamaica Plain, MA. 658 p.
Bulletin of Animal Health and Production in
Africa 38(3): 275-279.
Kumar, S. and N. Rohatgi. 1999. The role of
invasive weeds in changing floristic diversity.
Oyedapo, O.O., F. C. Sab, and J.A. Olagunju.
Annals of Forestry 7:(1): 147-150.
1999. Bioactivity of fresh leaves of Lantana
camara. Biomedical Letters 59: 179-183.
Lal, L. 1987. Studies on natural repellents against
potato tuber moth (Phthorimaea operculella
Sahid, I.B. and J. B. Sugau. 1993. Allelopathic
Zeller) in country stores. Potato Research 30(2):
effects of lantana (Lantana camara) and siam
329-334.
weed (Chromolaena odorata) on selected crops.
Weed Science 41(2): 303-308.
Liogier, H.A. 1990. Plantas medicinales de Puerto
Rico y del Caribe. Iberoamericana de Ediciones,
Sinha, S., B. Sinha, and A. Sharma. 1995.
Inc., San Juan, PR. 563 p.
Chromosome composition of Lantana camara
L.: karyotype, basic number and DNA diversity.
Liogier, H.A. 1995. Descriptive flora of Puerto
Nucleus Calcutta 38(1-2): 16-22.
Rico and adjacent islands. Vol. 4. Editorial de la
Universidad de Puerto Rico, San Juan, PR.
Weyerstahl, P., H. Marschall, A. Eckhardt, and C.
617 p.
Christiansen. 1999. Constituents of commercial
Brazilian lantana oil. Flavour and Fragranced
Morton, J.F. 1994. Lantana, or red sage (Lantana
Journal 14(1): 15-28.
camara L. Vergenaceae), notorious weed and
416
Lantana involucrata L. wild sage
VERBENACEAE
John K. Francis
417
Growth and Management.—The growth rate is Howard, R.A. 1989. Flora of the Lesser Antilles.
moderate and plants live for 10 years or more. Vol. 6. Arnold Arboretum, Harvard University,
Propagation may be by seed, by rooting of Jamaica Plain, MA. 658 p.
softwood cuttings, or by transplanting wildlings.
Wildlings transplant well if the roots are not badly Liogier, H.A. 1995. Descriptive flora of Puerto
damaged and about one-third of the top is pruned Rico and adjacent islands. Vol. 4. Editorial de
away. Periodic pruning of ornamental plants is la Universidad de Puerto Rico, San Juan, PR.
necessary to prevent them from becoming lank 617 p.
(Workman 1980).
Long, R.W. and O. Lakela. 1971. A flora of
Benefits.—Wild sage contributes to biodiversity, tropical Florida. University of Miami Press,
helps protect the soil, and furnishes wildlife food Coral Gables, FL. 962 p.
and cover. The wood is useful for fuel, but it is
generally too small for anything but campfires. The Malone, K. 2002. Palm Beach County
species is used as a foundation plant and to form (Southeastern Florida) top butterfly nectar
low hedges (Workman 1980). It is listed as one of flowers. North American Butterfly Association.
the best butterfly nectar plants (Malone 2002). The http://www.naba.org/ftp/pbco.pdf. 4 p.
leaves are used as a condiment in cooking, and the
essential oil is used in cosmetics and liquors McNary-Wood, K. 2002. Medicinal plants for
(Secritaría del Medio Ambiente y Recursos what ails you in paradise. http://www.
Naturales 2002). Leaves are added to baths to ease timespub.tc/Natural%History/Archive/Winter%
heat rashes and mild insect bites (McNary-Wood 20200102/medicinal.htm. 2 p.
2002). Extracts of the plant are also said to have a
sedative effect (Beattie and others 2002). Teas and Nelson, G. 1996. The shrubs and woody vines of
decoctions of leaves and twigs are used to control Florida. Pineapple Press, Inc., Sarasota, FL.
colic, vomit, cough, fever, and congestion 391 p.
(Secritaría del Medio Ambiente y Recursos
Naturales 2002). It is reportedly not as poisonous Secritaría del Medio Ambiente y Recursos
as the more widespread L. camara L. (Workman Naturales. 2002. Especies con usos no
1980). maderables en bosques tropicales y
subtropicales: Lantana involucrata L.
References http://www.semarnat.gob.mx.pfnm2.fichas/
lantana_involucrata.htm. 1 p.
Beattie, L., M. Martin, M. Shaposhnik, and K.
Vaga. 2002. Stress treatments. http://members. Workman, R.W. 1980. Growing native. The
tripod.com/~Moiraine/treatment.html. 7 p. Sanibel-Captiva Conservation Foundation, Inc.,
Sanibel, FL. 137 p.
Britton, N.L. 1918. Flora of Bermuda. Charles
Schribner’s Sons, New York. 585 p.
418
Larrea tridentata (Sesse' and Moc. ex DC.) Coville creosote bush
ZYGOPHYLLACEAE
James E. Nellessen
419
caliche layer. The shallow roots are able to utilize increases associated with spring temperatures and
small rainfall events. Creosote roots inhibited moisture availability (Mabry and others 1977). It
further elongation of the roots of both adjacent has high water use efficiency and ability to resorb
creosote bushes and the shrub Ambrosia dumosa some nutrients, especially phosphorus, 72 to 86
(A. Gray) Payne (Mahall and Callaway 1992). percent, while nitrogen resorption is within the
Stem angle facilitates water stemflow to the roots, general range for other evergreen shrubs, 47 to 57
which is maximal with stems greater than 65° and percent. Creosote bush stem and leaf orientations
almost none less than 45°. Creosote bushes in the maximize light collection during cooler and
driest part of their range, for example, Death moister early morning periods. Creosote bush has
Valley, generally have steeper angles, while there a greater ability to withstand xylem cavitation
is more variability in stem angle in the Chihuahuan during droughts than riparian species (Pockman
Desert. Shrubs with shallower stem angles take on and Sperry 2000). An Arizona population of
a more hemispherical shape as opposed to an creosote bush experiences freezing-induced xylem
inverted cone shape. Shrub canopy architecture vessel cavitation and consequent disruption of
also affects soil characteristics creating "islands of water transport at temperatures from -11 to -20 °C
fertility." Shrubs with hemispherical shapes collect (Pockman and Sperry 1997). These temperatures
much more litter underneath, hence soil nitrogen is correspond to 20+ year minimum isotherms and
greater under hemispherical shrubs than under also correspond with the northern distribution in
inverted cone shaped shrubs (Whitford 2002). both the Sonoran and Mojave Deserts. A few
Creosote bushes often require a nurse shrub such studies have examined the effects of
as the small shrub A. dumosa for seedling/sapling environmental pollutants on creosote bush. Foliage
establishment, and creosote bush itself will serve demonstrates leaf injury when exposed to acute
as a nurse plant for certain other species (Whitford levels of sulfur dioxide (SO2) (Olyszyk and others
2002). Creosote bush often forms monotypic 1987) and elevated CO2 concentrations may
stands that extend for miles, although its density ameliorate the effects of heat stress (Hamerlynck
varies greatly from site to site, with the greatest and others 2000). Creosote foliage may
densities occurring in the Chihuahuan Desert (up accumulate heavy metals from industrial pollution
to 8,000 plants/ha), and lesser densities in the as demonstrated in a 1980 to 1995 study from El
Sonoran (up to 4,000 per ha) and the Mojave (up Paso, TX, and Ciudad Juarez, Mexico (Mackay
to 1,800 per ha) (Mabry and others 1977). Above- and others 1998).
ground standing crop biomass tends to reflect
these density trends. Chihuahuan Desert Reproduction.—Creosote bush blooms
populations tend to show a greater size or age class periodically during the growing season, blooming
structural diversity than those in other deserts correlated first with seasonality, secondarily with
(Mabry and others 1977). The resinous coating on the timing of moisture availability and other local
the leaves helps retard water loss. During extreme environmental factors. Peak blooming periods
droughts creosote bush will produce smaller, occur in spring (February to May) and again in late
tougher leaves, and will also shed many of its summer into the fall (August to December). A
older leaves. It quickly produces new shoot single mature shrub may produce more than 2,000
growth after substantial summer rains. Creosote flowers (or about 50 g of flowers) during the
bush has the C3 photosynthetic pathway, with growing season. Numerous insects visit the
peak photosynthetic activity (Pmax) in the spring flowers, but bees are the most effective pollinators.
and during summer monsoon seasons especially Many species of bees are known to visit the
when pre-dawn water potentials are high flowers; some are generalists but some are
(Whitford 2002). Pmax is as high as that found in specialists and forage solely on creosote bush
C4 plants (Fitter and Hay 1987), and creosote (Minckley and others 2000). It is an outcrossing
shrubland has a similar community energy balance species but is always self-compatible (Mabry and
as the C4 desert grasslands they have invaded others 1977) with localized populations having
(Dugas and others 1996). Creosote bush has higher levels of inbreeding in the Chihuahuan
demonstrated a threshold for stomatal closure at - Desert. The flowers are borne solitary in the axils,
5.8 MPa, has maintained full turgor at -3.0 MPa, are 2 to 3 cm in diameter, are perfect, with five
with positive net photosynthesis occurring at -8 to yellow clawed petals, obovate to spatulate, 6 to 10
-16 MPa (Fitter and Hay 1987, Mabry and others mm long, and 5 mm wide. There are five
1977). It can tolerate leaf water potentials below - yellowish green, unequal, deciduous sepals, 5 to 8
50 MPa. It is able to maintain a relatively stable mm long. There are 10 stamens with winged
photosynthetic rate all year in Death Valley, with filaments, and the ovary has five locules. The
420
petals twist at a 90 degree angle after pollination. fertilized, irrigated, or unmodified ungrazed black
The fruit is a roundish capsule, about 5 to 7 mm in grama (B. eriopoda) grassland, a creosote
diameter, covered with a dense concentration of shrubland, or a heavily grazed grassland. The only
white to reddish hairs, and separates into five site in which seedlings survived was in the heavily
indehiscent one-seeded carpels at maturity (Carter grazed grassland (Whitford 2002). Prior to heavy
1997, Correll and Johnston 1970, Kearney and livestock grazing creosote bush was generally
others 1951, Martin and Hutchins 1980-81). Both restricted to well-drained gravelly or sandy soils of
fruits and flowers are commonly found on the steep slopes. Creosote bush and mesquite
plant at the same time. Flowering may begin on (Prosopis glandulosa Torrey) invasion likely
plants as young as 4 to 6 years, although heavy occurred in episodes in conjunction with both
flowering and fruiting generally occurs after 8 to reduced grass cover and drought (Grover and
13 years. Greenhouse-raised seedlings may flower Musick 1990). Localized decreases have also been
at 2 years of age. Fruit production ranges from 39 reported. In an area of Big Bend National Park,
to 278 fruits per 100 g of branches or 120 to 1,710 previously overgrazed by livestock, the dry period
per plant (Young and Young 1992). There are from 1960 to 1967 showed increased creosote
about 370 seeds/g. The seeds have mechanical bush cover, while the wetter period from 1967 to
dormancy and scarification aids germination 1981 showed a decrease in creosote bush cover
(Young and Young 1992). Germination success and increased cover of perennial grasses, forbs,
varies by population, latitudinal and local and most other shrubs (Wondzell and Ludwig
environmental conditions with reported success 1995). At a site in southern New Mexico creosote
rates of 2 to 70 percent. Salt (NaCl) at 10,000 ppm shrub size has decreased over a 10-year period,
(-0.78 MPa) resulted in 0 percent germination, and possibly due to longer term climate change (Miller
seed germinated equally well at pH's from 7 to 10 and Huenneke 1996). Certain creosote bush
(Barbour 1968), although some studies have populations in the Sonoran Desert of Mexico
shown some variation from these pH's (reviewed declined by 50 to 90 percent, perhaps due to
in Baskin and Baskin 2001). Leachates from fruit prolonged droughts from 1936 to 1964 (Turner
coats inhibit germination of some plant species 1990). Creosote bush shrublands have been
[Bouteloua eriopoda (Torr.) Torr. but not experimentally treated with a variety of herbicides
Muhlenbergia porteri Scribn. ex Beal], but not of for shrub management. Several liquid-applied
creosote bush seeds (Baskin and Baskin 2001). herbicides (2,4-D, dicamba, picloram, 2,3,6-TBA)
Seeds stored dry in paper bags at room either singly or in combination with other
temperature for as long as 8 years showed some herbicides have demonstrated greater than 50
germinability (Barbour 1968). percent root kill (Herbel and Gould 1995). But
individual shrub treatment with dry herbicides
Growth and Management.—Growth response (bromacil, fenuron) or aerially applied tebuthiruon
and productivity can be rapid after rain, but there pellets have given greater control, 80 to 90
is not a strong pattern of productivity and rainfall, percent. Perennial grass and annual forb
while nitrogen has been shown as an important production increased substantially on tebuthiuron-
factor in creosote bush productivity (Whitford treated plots.
2002). Maximum leaf longevity is 16 months.
Maximal root growth occurs in the fall. Branches Benefits and Disadvantages.—Creoste bush is
will root if covered with soil. Ring counts of stems unpalatable as livestock forage and is usually
have indicated that creosote bush is long-lived, at toxic, sometimes causing death (Gay and Dwyer
least 30 to 50 years, but ring counts may not be 1998, Kearney and others 1951, Stubbendieck and
reliable. Individual shrubs may sometimes others 1997, USDA 1937). Jackrabbits are one of
reproduce by fragmentation or separation of the the few mammals that will consume the plant to a
root crowns along with new basal sprouting, certain degree, largely based on the water content
causing the plant to spread out and away from the of a particular plant (Whitford 2002). Some
center. Such fragmentation may contribute to long- woodrat populations will also consume creosote
term survival estimated at more than 100 years to bush, demonstrating differentially developed
several thousand years. Creosote bush shrubland tolerances to the resin (Mangione and others
has been expanding over the past 100 years or 2000). Woodrats consume creosote leaves and
more due to overgrazing of desert grasslands by stems in the Mojave Desert when necessary to
livestock, although climate trends have also been maintain water balance but suffer ill effects such
implicated. One experiment to test this involved as loss of body mass (Karasov 1989). Both rabbits
transplanting seedling creosote bushes into and woodrats will peel open the stems to get water.
421
Once a particular shrub is browsed by a jackrabbit, principal compounds in creosote bush resin is the
that same shrub is much more likely to be browsed anti-oxidant nordihydroguaiaretic acid (NDGA).
again than another shrub that has never been This is a phenolic aglycone and comprises 5 to 10
browsed (Ernest 1994). Younger leaves have a percent of leaf dry weight, 80 percent of all
higher concentration of resins that affect relative phenolics in the resin, and once was used as an
leaf palatability and digestibility. Resin content anti-oxidant in foods, pharmaceuticals, and
may be as high as 26 percent in young leaves, industrial materials (Mabry and others 1977).
dropping to 11 to 16 percent in older leaves, with NDGA has demonstrated an ability to inhibit
insects showing preference for the older leaves tumors and cancers but does have toxic side effects
(Mabry and others 1977). A phenoloxidase system to both animals and humans. Its use as a
in the leaves appears to make digestibility of the chemoprevention for skin cancer and protection
resinous leaves even more difficult. Chemical against the effects of ultaviolet light has been
treatment of creosote foliage to remove the resins investigated (Gonzales and Bowden 2002). The
results in nutritious edible forage, but due to botanical dietary supplement "chapparal" has been
economics this practice is not often implemented associated with nonviral toxic hepatitis and
(Mabry and others 1977). Numerous species of contains lignans similar to estrogens (Obermeyer
arthropods set up habitation in creosote bush. In and others 1995) and may cause acute liver
particular males of the grasshopper species, damage (Sheikh and others 1997). NDGA has
Ligurotettix coquilletti McNeil, select a particular shown potential to inhibit the human
shrub as their territory and defend it against other immunodeficiency virus (HIV) (Gnabre and others
male grasshoppers (Mabry and others 1977). The 1995). NDGA (masoprocol) lowered glucose and
mounds created by kangaroo rats (Dipodomys triglyceride levels in rats and may have potential in
spp.) appear to have a positive effect on creoste Type II diabetes treatment (Reed and others 1999).
bush survival, flowering, and fruiting, while Numerous flavanoid aglycones, flavanoid
creosote bush itself has a negative effect on glycosides, wax esters, saponins, and volatile
kangaroo rat populations (Chew and Whitford constituents such as terpenes, have been identified
1992). Creosote bush exposure to 0.1 to 0.2 ppm from creosote bush.
ozone consistently reduced the toxic
nordihydroguaiaretic acid content of leaves, which References
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Guadalupe Mountains and the Sand Dune
Obermeyer, W.R., S.M. Musser, J.M. Betz, R.E. Country, Texas. Sul Ross State University,
Casey, A.E. Pohland, and S.W. Page. 1995. Alpine, TX. 176 p.
Chemical studies of phytoestrogens and related
compounds in dietary supplements: flax and Whitford, Walter G. 2002. Ecology of Desert
chaparral. Proceedings of the Society for Systems. Academic Press, San Diego, CA. 343
Experimental Biology and Medicine 208(1): 6- p.
12.
Wondzell, S. and J.A. Ludwig. 1995. Community
Olszyk, D.M, A. Bytnerowicz, C.A. Fox, G. Kats, dynamics of desert grasslands: influence of
P.J. Dawson, and J. Wolf. 1987. Injury and climate, landforms, and soils. Journal of
physiological responses of Larrea tridentata DC Vegetation Science 6(3): 377-390.
Coville exposed in-situ to sulfur dioxide.
Environmental Pollution 48(3): 197-212. Young, J. A. and C. G. Young. 1992. Seeds of
Woody Plants in North America. Dioscorides
Press, Portland, Oregon. 407 p.
424
Lasiacis divaricata (L.) A.S. Hitchc. wild bamboo
POACEAE
John K. Francis
425
percent germination and the other 11 percent. The and cattle. The species seems resistant to light and
seeds are somewhat delicate and probably damaged moderate browsing. The tough stems, which often
by scarification. The pericarp can be removed by run across low brush, make it difficult to walk
rubbing with great care with unknown effect on through the forest.
germination. It is assumed that birds and possibly
rodents are seed dispersers. Seedlings are not References
common. Culms layer (root) whenever they come
in contact with the ground and are covered with CromoPar. 2002. Recuentos cromosómicos
litter. Wild bamboo sprouts when cut or damaged foráneos de plantas presentes en Paraguay.
to renew damaged culms. Universidad de Barcelona, Spain. http://
www.ub.es/botanica.cromopar.cro-out/pdf. 24 p.
Growth and Management.—Growth of wild
bamboo from sprouts is rapid. Sprouts from mature Hitchcok, A.S. 1935. Manual of the grasses of the
plants may reach most of their eventual length United States. Miscellaneous Publication 200.
during the first year. Thereafter, branches are U.S. Department of Agriculture, Washington,
added for 1 or 2 years. After old culms die, new DC. 1,040 p.
ones grow from the root crown. It is not known
how fast seedlings grow. Wild cane seldom forms Nelson, G. 1996. The shrubs and woody vines of
thickets and mostly grows in forests and brush Florida. Pineapple Press, Inc., Sarasota, FL.
lands so that control does not seem warranted. If it 390 p.
should be needed, grubbing out the root crowns or
cutting and spot-spraying the resulting sprouts with Stevens, W.D., C. Ulloa-U., A. Pool, O.M.
glyphosate would probably be effective. Montiel, eds. 2001. Flora de Nicaragua.
Monographs in Systematic Botany Vol. 85, No.
Benefits.—Wild bamboo furnishes food and cover 3. Missouri Botanic Garden Press, St. Louis,
for wildlife and helps protect the soil. The leafy MO. p. 1,911-2,666.
and more succulent portions are browsed by horses
426
Leandra krugii (Cogn.) Judd & Skean leandra
MELASTOMATACEAE
John K. Francis
427
References W3Tropicos. 2002. Calycogonium krugii Cogn.,
flowering shrub.http://digitalis.mobot.org/mrsid/
Liogier, H.A. 1995. Descriptive flora of Puerto bin/mosid/mosid.pl?client=203&image=MOA-
Rico and adjacent islands. Vol. 4. Editorial de la 06799_001.sid&title=Calycogonium+krugii+ Cog...
Universidad de Puerto Rico, San Juan, PR. 1 p.
617 p.
428
Ledum groenlandicum Oeder Labrador tea
ERICACEAE
James E. Nellessen
429
mined peatland, despite the absence of new as a folk remedy for lung ailments, dysentery,
sphagnum moss colonization (Berube and Lavoie indigestion, and to kill lice and treat leprosy.
2000). Successful traditional use for gout treatment may
be due to the presence of phenolics and tannins
Reproduction.—Labrador tea flowers are within the plant (Owen and Johns 1999). This
relatively small, about 1 cm wide and 2 cm long, species, as well as others in the genus, has
white, with five to seven stamens, and are grouped generally been regarded as having low palatability,
in terminal clusters. It blooms from May to July, being unpalatable, or even slightly poisonous
sometimes as late as August. The fruit is a slender (USDA 1937). Since this species often grows in
capsule, oval in shape, about 5 to 8 mm long and 2 wet boggy habitats, it is generally not accessible to
to 3 mm wide, with a persistent style (Gleason and most livestock. The plant may serve as reindeer
Cronquist 1963, Britton and Brown 1913). The forage in Alaska. Its unpalatability may also be
seeds are elongated, numerous and small. For due to essential oils of the monoterpene family
germination, they should be placed onto a good (sabinene and limonene) and the sesquiterpene
moisture-supplying substrate and covered with a family (alpha- and beta-selinene and germacrone)
clear plastic film (Young and Young 1992). Seeds (Belleau and Collin 1993). Germacrone has been
germinate best after a 30-day cold stratification shown to be a feeding deterrent for snowshoe
period, with optimal germination temperatures of hares (Lepus americanus Erxleben), although they
20 to 25 °C, and light is required (Calmes and do eat the plant to some extent (Reichardt and
Zasada 1982, Baskin and Baskin 2001). The seeds others 1990, MacCracken and others 1988).
are considered to have non-deep physiological
dormancy because of the relatively shorter cold References
periods needed and that some seeds can germinate
shortly after ripening. The cold stratification Addison, P.A., S.S. Malhotra, and A.A. Khan.
period can decrease the germination temperature 1984. Effect of sulfur dioxide on woody boreal
requirement. Full sunlight and high soil moisture forest species grown on native soils and tailings.
seem to be requirements, including a low soil pH, Journal of Environmental Quality 13(3): 333-
e.g. 5.5 (Karlin and Bliss 1983, Baskin and Baskin 336.
2001). Apparently winter-time cuttings will root
well (Dirr and Heuser 1987, Young and Young Baskin, C.C. and J.M. Baskin. 2001. Seeds:
1992). ecology, biogeography, and evolution of
dormancy, and germination. Academic Press,
Growth and Management.—This shrub is San Diego, CA. 666 p.
generally considered a slow growing, late
successional, pre-climax or climax species. The Belleau, F. and G. Collin. 1993. Composition of
low shrub stratum consisting of Labrador tea and the essential oil of Ledum groenlandicum.
leatherleaf was estimated to have an aboveground Phytochemistry (Oxford) 33(1): 117-121.
productivity of 0.4 and 2.0 t/ha/yr on perched and
raised bogs, respectively, in northern Minnesota Berube, M.E. and C. Lavoie. 2000. The natural
(Grigal and others 1985). Rust fungi of the genus revegetation of a vacuum-mined peatland: eight
Chrysomyxa alternate between spruce and years of monitoring. Canadian Field Naturalist
members of the Ericaceae such as Labrador tea. 114(2): 279-286.
Chrysomyxa reticulata sp. nov. may spread from
Ledum spp. to cultivated rhododendrons (Crane Britton, N.L. and A. Brown. 1913 (1970 Dover
2001). edition). An illustrated flora of the northern U.S.
and Canada, Vol. 2. Dover Publications Inc.,
Benefits.—As the name implies, the dried leaves NY. 735 p.
have been used for tea. Native Americans, such as
the Chippewa, have used this shrub for various Brooks, R.R. 1987. Serpentine and its vegetation:
purposes. The root has been used for medicinal A multidisciplinary approach. Dioscorides
purposes to treat ulcers and the leaves for making Press, Portland, OR. 454 p.
tea (Densmore 1928). As a tea this plant has been
used for treating asthma, colds, stomach aches, Calmes, M.A. and J.C. Zasada. 1982. Some
kidney problems, scurvy, and fevers (Foster and reproductive traits of four shrub species in the
Duke 1990). Externally it has been applied as a black spruce forest type of Alaska. Canadian
wash for burns, ulcers, and stings. It has been used Field Naturalist 96: 35-40.
430
Crane, P.E. 2001. Morphology, taxonomy, and Kron, K.A. and W.S. Judd. 1990. Phylogenetic
nomenclature of the Chrysomyxa ledi complex relationships within the Rhodoreae Ericaceae
and related rust fungi on spruce and Ericaceae in with specific comments on the placement of
North America and Europe. Canadian Journal of Ledum. Systematic Botany 15(1): 57-68.
Botany 79(8): 957-982.
MacCracken, J.G., W.D. Steigers, Jr., and P.V.
Densmore, F. 1928 (1974 Dover edition). How Mayer. 1988. Winter and early spring habitat
Indians use wild plants for food, medicine and use by snowshoe hares, Lepus americanus, in
crafts. Dover Publications, New York. 397 p. south-central Alaska, USA. Canadian Field
Naturalist 102(1): 25-30.
Dirr, M.A. and C.W. Heuser. 1987. The reference
manual of woody plant propagation. Varsity McCance, R.M., Jr. and J.F. Burns, eds. 1984.
Press, Athens, GA. Ohio endangered and threatened vascular plants:
Abstracts of state listed taxa. Department of
Foster, S.A. and J.A. Duke. 1990. A field guide to Natural Resources, Columbus, OH. 635 p.
medicinal plants (eastern/central). Peterson Field
Guide Series, Houghton Mifflin Co., Boston, Owen, P.L. and T. Johns. 1999. Xanthine oxidase
MA. 366 p. inhibitory activity of northeastern North
American plant remedies used for gout. Journal
Gleason, H.A. and A.R. Cronquist. 1963. Manual of Ethnopharmacology 64(2): 149-160.
of the vascular plants of northeastern U.S. and
adjacent Canada. D. Van Nostrand Co., New Pugh, R.E., D.G. Dick, and A.L. Fredeen. 2002.
York. 810 p. Heavy metal (Pb, Zn, Cd, Fe, and Cu) contents
of plant foliage near the Anvil Range lead/zinc
Great Plains Flora Association. 1986. Flora of the mine, Faro, Yukon Territory. Ecotoxicology and
Great Plains. University Press of Kansas, KS. Environmental Safety 52(3): 273-279.
1,392 p.
Reichardt, P.B., J.P. Bryant, B.J. Anderson, D.
Grigal, D.F., C.G. Buttleman, and L.K. Kernik. Phillips, T.P. Clausen, M. Meyer, and K. Frisby.
1985. Biomass and productivity of the woody 1990. Germacrone defends Labrador tea from
strata of forested bogs in northern Minnesota. browsing by snowshoe hares. Journal of
Canadian Journal of Botany 63(12): 2416-2424. Chemical Ecology 16(6): 1961-1970.
Inderjit, A.U.M. 1996. Growth and physiological Sheard, J.W. 1986. Distribution of uranium series
responses of black spruce (Picea mariana) to radionuclides in upland vegetation of northern
sites dominated by Ledum groenlandicum. Saskatchewan, Canada I: Plant and soil
Journal of Chemical Ecology 22(3): 575-585. concentrations. Canadian Journal of Botany
64(11): 2446-2452.
Karlin, E.F. and L.C. Bliss. 1983. Germination
ecology of Ledum groenlandicum and Ledum U.S. Department of Agriculture Forest Service.
palustre ssp. decumbens. Arctic Alpine 1937 (1988 Dover edition). Range plant
Research 15: 397-404. handbook. Dover Publications Inc., New York.
816 p.
Kartesz, J.T. 1994. A synonymized checklist of
the vascular flora of the United States, Canada, Young, J. A. and C. G. Young. 1992. Seeds of
and Greenland, Vol. 1, 2nd Edition. Biota of woody plants in North America. Dioscorides
North America Program of the North Carolina Press, Portland, OR. 407 p.
Botanical Garden. Timber Press, Portland, OR.
622 p.
431
Lepianthes peltata (L.) Raf. monkey’s hand
PIPERACEAE
John K. Francis
432
abundant. Monkey’s hand plants are easy to in many of the same ways in the West Indies
propagate from cuttings. A small group of stem (Liogier 1990).
cuttings were treated with IBA and placed in a mist
bed. After about 5 weeks, 85 percent had rooted.
Because rooting takes place at the nodes, a node 1
Garlon is named here for identification purposes
should be included in the buried portion of the only. The Forest Service does not endorse any
cutting. commercial product.
433
Lepidospartum burgessii B.L. Turner gypsum scalebroom
ASTERACEAE
Synonyms: none
Juanita A. R. Ladyman
of the involucre on the flower heads clearly
distinguish it from any species in that genus.
434
winter temperature ranging from lows of –2 oC and populations are reproductively compatible. Some
highs of 14 oC. Freezing is common during this protective measures, including erecting exclosures,
time. Precipitation comes primarily during the have been made for some colonies growing in an
summer (ranging from 13.1 to 46.8 cm). However, Area of Critical Concern and managed by the
substantial spring rains have been reported, and Bureau of Land Management (BLM 1997). This
there is a high degree of year-to-year variability species appears to be facing relatively imminent
characteristic of most arid and semiarid climates extinction. From a simple model that assumed the
(Noy-Mier 1973). rate of death was linear, extinction is predicted
within 35 years (Ladyman and others 1999).
Reproduction.—The peak of flowering is during
late July through early September, but within a Benefits.—Gypsum scalebroom is visited by a
population, flowering can extend from late April to wide variety of arthropods and possibly provides
early October. The number of flowers per shrub is cover and food for small wildlife species. It is
highly variable; from one flower to literally unlikely to be palatable to livestock, but they do
hundreds of flower heads on any given shrub. appear to brush against it even if they do not use it
There does not appear to be a close relationship as food (anecdotal information from ranchers in
between the size of the plant and the number of area and author’s personal observation). This is
flowers because the largest shrubs are often in a likely due to the absence of other large shrubs in
decadent condition, that is, with more than 50 the region. It is a beautiful silvery ornamental
percent dead wood. In 1991/1992 and 1997 plants shrub. It has been suggested that one way to slow
were counted in seven colonies in New Mexico its eventual extinction is to introduce it into the
(Huenneke 1991, Ladyman and others 1999). horticultural and landscape trade.
Approximately 15 percent of the individuals
counted within those colonies in 1991/1992 were References
dead in 1997. The number of juvenile plants had
also declined. Recruitment is only by clonal Baldwin, B.G., S. Boyd, B.J. Ertter, R.W.
propagation. No seeds have been observed despite Patterson, T.J. Rosatti, and D.H. Wilken. 2002.
abundant yellow flowers and apparently adequate The Jepson Desert Manual, vascular plants of
pollinators (Ladyman and others 1999). The results southeastern California. University of California
of studies on pollen germination on stigmas Press, Berkeley, CA. 624 p.
indicate an incompatibility system is operating
(Ladyman and others 1999). The prematurely
stunted pollen tube growth observed by florescent Bureau of Land Management. 1997. Areas of
microscopy is indicative of sporophytic Critical Environmental Concern Resource
incompatibility, which is typical of members of the Management Plan Amendment, August 1997.
Asteraceae and other species with trinucleate Bureau of Land Management, Las Cruces
pollen (Ladyman and others 1999, de Nettancourt District, Caballo Resource Area, Otero County,
1977). Las Cruces, NM [not paged].
435
Humphrey, R.R. 1958. The desert grassland, a Noy-Mier, I. 1973. Desert ecosystems:
history of vegetational change and an analysis of environment and producers. In: Annual Reviews
causes. Botanical Review 24:193-252. of Ecology and Systematics IV: 25-51.
Ladyman, J.A.R., L. DeLay, P. Gegick, M. Bogan. Powell, A. Michael. 1998. Trees & shrubs of
1999. Status and Reproductive Biology of Trans-Pecos and adjacent areas. University of
Lepidospartum burgessii (Burgess broomshrub Texas University Press, Austin, TX (First Ed.
or gypsum broomscale). Unpublished report copyright Big Bend Natural History Assoc.) 498
submitted to the USGS Biological Survey, p.
Albuquerque, NM. 108 p.
Rotem, J. 1994. The Genus Alternaria – Biology,
Ladyman, J.A.R. and Patricia Gegick. 2001. Epidemiology, and Pathogenicity. APS, St. Paul,
Status of Lepidospartum Burgessii (Burgess MN. 325 p.
broomshrub or gypsum broomscale). In:
Southwestern Rare and Endangered Plants. Turner, B.L. 1977. Lepidospartum burgessii
Proceedings of Third Conference. RMRS-P-23. (Asteraceae, Senecioneae), a remarkable new
USDA Forest Service, Rocky Mountain gypsophilic species from Trans-Pecos Texas.
Research Station, Fort Collins, CO. p. 116-127. Wrightia 5: 354-355.
436
Leucaena leucocephala (Lam.) de Wit leadtree
FABACEAE
John K. Francis
437
disturbed, eroded, and partially compacted soils. Individual shrubs live 10 to 20 years or more.
However, the species does not grow well in poorly Plantations are relatively easy to establish with
drained soils or periodically flooded soils, in soils containerized seedlings or by direct seeding
with high salinity or those below pH 5.0 or above scarified seed into prepared seedspots at the
pH 8.0, at temperatures below 10 °C, and in areas beginning of the wet season. Because of its
with mean annual rainfall below 750 mm or above aggressive nature, it is not advisable to introduce
2500 mm (Skerman and others 1988). Leadtree is leadtree into areas where it does not already occur.
intolerant of shade. It does best in the open, but Within the native or naturalized range, disturbed
competes aggressively in low stands of weeds and wildlands are usually quickly colonized by leadtree
grass and will grow in low forests with low basal from the soil seedbank or from nearby seed trees.
areas. It often forms thickets that briefly eliminate However, where seed is not present, planting the
nearly everything underneath but soon self-thin to species may be an excellent way of rehabilitating
fairly open stands with normal understory damaged sites. Grass swards should be cultivated
vegetation. Leadtree is drought tolerant. It folds its or burned or sprayed followed by cultivation before
leaves under water stress preventing water loss planting. About 5,000 to 10,000 plants/ha should
and, under severe conditions, will defoliate until be established for erosion control and sward
the rains return (National Academy of Sciences conversion. Dense stands of leadtree needing
1977). control may be slashed and grazed or sprayed with
broadleaf herbicide.
Reproduction.—Leadtree blooms throughout the
year (Pennington and Sarukhan 1968) and may Benefits.—Leadtree quickly invades disturbed
begin blooming as young as 4 to 6 months of age areas and prevents further erosion while allowing
(National Academy of Sciences 1977). The flowers succession to secondary forest cover. It is one of
are largely self-compatible and self-fertilized. It only a few species that can invade and rehabilitate
takes about 4 months after flowering for pods to dense tall grass swards such as Panicum maximum
mature and liberate seeds (Binggeli 1997). From Jacq. and Imperata cylindrica (L.) Raeusch.
10,000 to 21,000 seeds/kg are reported although it (author’s observation, National Academy of
is sometimes unclear from what variety they were Sciences 1977). Leadtree furnishes food and cover
sampled (Brewbaker 1997, Parrotta 2000). A for wildlife, especially browsing animals. The
sample of seeds from shrubby plants from the north species has been planted widely in agroforestry
coast of Puerto Rico averaged 0.0392 ± 0.0006 applications. It is useful for forming erosion
g/seed or 25,500 seeds/kg (author’s observation). barriers, windbreaks, for shade and support, green
Leadtree seeds have a hard coat that requires manure, and cut fodder. It is known to fix useful
scarification for reliable germination. This can be quantities of nitrogen (Parrotta 2000). Plantations
done by abrasion, or treatments with hot water or are established for pasture and cut for hay. The
concentrated acid. Germination of scarified seed is foliage is highly digestible (60 to 70 percent)
usually between 50 and 98 percent in 6 to 10 days (Brewbaker 1997). Young leaves are reported to
after sowing. Seed can be stored for up to 5 years contain 68 calories/100 g and 80 percent moisture
at temperatures of 2 to 6 °C (Parrotta 2000). Pods (Duke 1983). By dry weight, leaves contain 21
open while still attached to liberate their seeds and percent crude protein, 18 percent crude fiber, 8
are dispersed by granivorous and herbivorous percent ash, 6 percent fat, and 46 percent total
animals (Pacific Island Ecosystems at Risk 2002) digestible nutrients (FAO 2002). Cattle readily eat
and machinery. Detached pods are blown some the forage but suffer symptoms of mimosine
distance by wind and occasionally retain attached toxicity if given a pure diet of the species. Pigs and
seeds. Reproduction is common in disturbed sites chickens must not be given more than 5 to 10
near seed sources. Plants aggressively coppice percent in their diet, but goats have no difficulty
when burned, cut, or broken off. The plant can be with the forage, and sheep can become accustomed
propagated vegetatively from cuttings (Parrotta to diets rich in leadtree foliage (National Academy
2000). of Sciences 1977). The wood is a preferred fuel
that burns slowly with little smoke or ash and
Growth and Management.—Leadtree grows makes an excellent charcoal (Brewbaker 1997).
moderately rapidly (not as fast as the giant variety The stems are used for stakes and tool handles. The
for which most of the data are available). Seedlings seeds are widely employed for making jewelry and
may add about 1 m of height in the first year and placemats (Howard 1988). The young pods and
0.5 to 0.75 m for 2 or 3 years thereafter. Diameter green seeds are cooked as a vegetable, and the
growth may vary from 2 to 10 mm per year. mature seeds are used as a coffee substitute and
438
parched for snacks (Duke 1983). The flowers are a Little, E.L., Jr. and F.H. Wadsworth. 1964.
pollen source for honeybees (Little and Wadsworth Common trees of Puerto Rico and the Virgin
1964). Extracts of the roots and bark are a powerful Islands. Agriculture Handbook 249. U.S.
emetic and have been used as an abortifacient Department of Agriculture, Forest Service,
(Liogier 1990). Washington, DC. 548 p.
439
Lindera melissifolia (Walt.) Blume pondberry
LAURACEAE
440
numerous insect species. Flower and fruit Kral, R. 1983. A report on some rare, threatened,
production are highly variable (Morgan 1983, or endangered forest-related vascular plants of
Tucker 1984). Some years the flowers are the South: Volume 1, Isoetaceae through
damaged by late frosts, while other years flower Euphorbiaceae. Technical Publication R8-TP 2.
and fruit production may be heavy, with up to 100 U.S. Department of Agriculture, Forest Service,
fruits per plant. Southern Region, Atlanta, GA. 718 p.
Growth and Management.—Seedlings are rarely Morgan, S. 1983. Lindera melissifolium: A rare
observed (Devall and others 2001). Stems usually southeastern shrub. Natural Areas Journal 3: 62-
live 6 or 7 years, and when a stem dies it is almost 67.
always replaced by a new stem that grows from
the base of the plant. Clones expand vegetatively, Radford, A.E., H.E. Ahles, and C.R. Bell. 1968.
and a mature colony often includes numerous leafy Manual of the Vascular Flora of the Carolinas.
stems along with some dead stems (U.S. Fish and University of North Carolina, Chapel Hill, NC.
Wildlife Service 1993). Most pondberry colonies 1,183 p.
occur in light shade, but a few grow in almost full
sun. In unshaded conditions, competition may be a Smith, C.G. III, P.B. Hamel, M.S. Devall and
problem. More knowledge is needed about N.M. Schiff. 2003. Hermit thrush is the first
management, and studies are currently being observed dispersal agent for pondberry.
conducted on the Delta National Forest in Castanea, in press.
Mississippi and at other locations. The U.S. Fish
and Wildlife Service (1993) states that pondberry Steyermark, J.A. 1949. Lindera melissifolia.
should be protected from forestry and agricultural Rhodora 51: 153-162.
management actions and protected from grazing
and browsing animals. Tucker, G.E. 1984. Status report on Lindera
melissifolia (Walt) Blume. Provided under
Benefits.—The fruits are eaten by hermit thrushes contract to the U.S. Fish and Wildlife Service,
(Catharus guttatus), cardinals (Cardinalis Southeast Region, Atlanta, GA. 41 p.
cardinalis) and other birds (Smith and others
2003). Steyermark (1949) reported that the fruits U.S. Fish and Wildlife Service. 1986. Endangered
are used as ammunition in pop guns, tubular and threatened wildlife and plants;
contrivances constructed from elderberry determination of endangered status for Lindera
[Sambucus nigra L. ssp. canadensis (L.) R. Bolli] melissifolia. Federal Register 51: 27,495-27,500.
twigs.
U.S. Fish and Wildlife Service. 1993. Recovery
References plan for pondberry (Lindera melissifolia). U.S.
Fish and Wildlife Service. Atlanta, GA. 56 p.
Devall, M., N. Schiff, and D. Boyette. 2001.
Ecology and reproductive biology of the Wright, R.D. 1989. Reproduction of Lindera
endangered pondberry, (Lindera melissifolia melissifolia in Arkansas. Proceedings of the
(Walt) Blume. Natural Areas Journal 21: 250- Arkansas Academy of Science 43: 69-70.
258.
Wright, R.D. 1990. Photosynthetic competence of
Klomps, V.L. 1980. The status of Lindera an endangered shrub, Lindera melissifolia.
melissifolia (Walt.) Blume, pondberry, in Proceedings of the Arkansas Academy of
Missouri. Transactions of the Missouri Academy Science 44: 118-120.
of Science 14: 61-66.
441
Lonicera japonica Thunb. Japanese honeysuckle
CAPRIFOLIACEAE
John K. Francis
442
precipitation limit (except where exogenous water mentioned herbicides to the cut surfaces (Bravo
is available), roughly corresponding with native 2003). Mowing is ineffective (Nuzzo 2003).
hardwood forests. Japanese honeysuckle survives
fire by sprouting from stem tissue buried under Benefits and Detriments.—In spite of the
litter and mineral soil. It generally disappears from invasive and weedy tendencies of the species,
communities (such as longleaf pine) where fires Japanese honeysuckle is still widely used as an
are frequent (Munger 2002). ornamental. It is especially desirable in arid areas
where it has little tendency to spread. Managed for
Reproduction.—Japanese honeysuckle in the U.S. screens and ground cover, Japanese honeysuckle is
flowers from April through July (sometimes beautiful in and out of flower and especially
through October) varying by local climate (Nuzzo captivating because of the strong and pleasant
2003). The flowers open in late afternoon and aroma of its blooms. Some gardeners favor it
remain open for about 3 days (Nuzzo 2003). The because it attracts hummingbirds. The fruits are
flowers are pollinated by hummingbirds and toxic to humans if large quantities are eaten and
insects (nocturnal hawkmoths and diurnal bees). result in vomiting, diarrhea, rapid heartbeat,
The species requires pollen from genetically respiratory failure, and coma. However, children
distinct individuals for successful pollination and suck the nectar from flowers without ill effects
as a result is pollinator limited. Naturally- (Russell 1997). The species was formerly planted
pollinated flowers produced seeds in 17.4 percent for erosion control and wildlife benefits. Planting
of the resulting fruits while hand-pollinated is currently discouraged or prohibited because the
flowers produced seeds in 78.7 percent of the species has dominated large areas and threatens
fruits (Larson 2001). The seeds are dispersed sensitive native species (Nuzzo 2003). Japanese
primarily by birds. Germination of most seeds honeysuckle is important wildlife cover and an
appears to occur in the spring following dispersal important, sometimes the most important, browse
and occurs at similar rates in mineral soil and species for white tailed deer in Eastern and
under leaf litter (Munger 2002). Japanese Southern United States. It is also eaten by
honeysuckle regenerates vegetatively by sprouting cottontail rabbits, cattle, sheep, and goats. The
from underground stems and by the layering of leaves yield 8 to 12 percent crude protein and have
above-ground stems that come in contact with the an in vivo dry matter digestibility of 35 to 70
soil. Discarded prunings will also take root. percent. Wild turkeys, northern bobwhite, and a
number of songbirds eat the fruits (Munger 2002).
Growth and Management.—Seedlings grow Japanese honeysuckle tissues contain anti-
slowly for the first 2 years but afterwards may complementary polysaccharides and polyphenolic
extend their stems 1.5 m/year (Nuzzo 2003). The compounds. Used in herbal medicine, extracts are
species begins bearing fruit at age 3 in full sun and reported to protect against cellular injury, help
at age 5 when growing in the shade. Fruit maintain vascular health, and prevent infections
production generally peaks between ages 4 and 6 (Nuzzo 2003)
(Munger 2002). By fertilization, researchers were
able to increase browse produced by Japanese References
honeysuckle by over 2,000 kg/ha and increase
crude protein as well (Dyess and others 1994). The Bravo, M.A. 2003. Japanese honeysuckle,
species is easy to propagate with cuttings and most Lonicera japonica Thunb. Alien Plant Working
ornamental plants are multiplied in this fashion. Group, National Park Service, Washington, DC.
The greater challenge is to eliminate infestations http://www.nps.gov/plants/alien/fact/loja1.htm.
of Japanese honeysuckle. Pulling and grubbing 6 p.
individual plants and small infestations can be
effective but will require repeat visits to eliminate Dyess, J.G., M.K. Causey, and H.L. Stribling.
missed sprouts and new seedlings. Tethered goats 1994. Effects of fertilization on production and
can provide effective control for spot infestations quality of Japanese honeysuckle. Southern
(Bravo 2003). Over-the-top spraying with Journal of Applied Forestry 18(2): 68-71.
glyphosate, Dichlorprep + 2,4-D, picloram + 2,4-
D, tebuthiuron, sulfometuron, or triclopyr will kill Larson, K.C. 2001. Lack of pollinators limits fruit
it (Nuzzo 2003) but also tend to eliminate most set in the exotic Lonicera japonica. Botanical
other plant species as well. A more selective Society of America, Botany 2001 Abstracts.
method is to cut stems and apply the above p. 39.
443
Liogier, H.A. and L.F. Martorell. 1982. Flora of Nuzzo, V. 2003. Element stewardship abstract for
Puerto Rico and adjacent islands: a systematic Lonicera japonica, Japanese honeysuckle. The
synopsis. Editorial de la Universidad de Puerto Nature Conservancy, Arlington, VA.
Rico, Río Piedras, PR. 342 p. http://tncweeds.ucdavis.edu/esadocs/documnts/
lonijap.html. 23 p.
Miller, J.H. 1999. Japanese honeysuckle, Lonicera
japonica. Protection Report RX-PR 036. U.S. Pacific Island Ecosystems at Risk. 2003. Lonicera
Department of Agriculture, Forest Service, japonica Thunb., Caprifoliaceae. http://www.
Region 8, Atlanta, GA. 2 p. hear.org/pier/lojap.htm. 3 p.
Munger, G.T. 2002. Lonicera japonica. In: U.S. Russell, A.B. 1997. Poisonous plants of North
Department of Agriculture, Forest Service, Carolina, scientific name: Lonicera japonica.
Rocky Mountain Research Station, Fire Sciences North Carolina State University, Charlotte, NC.
Laboratory. Fire Effects Information System. http://www.ces.ncsu.edu/depts/hort/consumer/
http://www.fs.fed.us/database/feis/plants/vine/ poison/Lonicja.htm. 2 p.
lonjap/all.html. 43 p.
Schierenbeck, K.A., J.L. Hamricks, and R.N.
Natural Resources Conservation Service. 2003. Mark. 1995. Comparison of allozyme variability
Plants profile: Lonicera japonica Thunb. in a native and an introduced species of
http://plants.usda.gov/cgi_bin/plant_search.cgi? Lonicera. Heredity 75(1): 1-9.
mode=Scientific+Name&keywordquery+Lonice
ra+japonica&earl=plant_search.cgi. 6 p. Sharma, A., A. Murayama, T. Osaki, K. Ooi, T.
Yahara, M. Ikegami, Silva-O., M. 1997. Plant
conservation in the Juan Fernandez archipelago,
Chile. Aliso 16(2): 89-101.
444
Lotus scoparius (Nutt.) Ottley California broom
FABACEAE
Arlee M. Montalvo
445
observation). Variety brevialatus occurs from Los Jones and Cruzan 1999) primarily by native bees
Angeles Co., south into Baja California primarily in the genera Bombus, Hoplitus, Anthophora,
in the hotter and drier interior regions of Riverside, Habropoda, Osmia, and Anthidium, but flowers
Los Angeles, western San Bernardino, and eastern are also visited by butterflies and non-native
San Diego Counties. Although the two varieties honeybees (author’s observation, Jones and
have somewhat different geographic distributions Cruzan 1999). The indehiscent pods ripen in about
and are easily distinguished by floral characters 4 to 6 weeks and are primarily passively dispersed
(Isely 1981, Steppan 1991), they overlap and short distances. The hard seeds require heat or
hybridize in some areas of Los Angeles, Riverside, mechanical scarification to break dormancy. Of
and San Diego Counties. Putative hybrids have several treatments including control, soil heated to
been observed in contact zones between the two 100 oC for 1 hr, ash /chemical fertilizer, and heat
varieties (Isely 1981, Steppan 1991, Montalvo and plus fertilizer, Christensen and Muller (1975)
Ellstrand 2001). Flowers of synthetic F1 hybrids found that heat treatment yielded the highest
are somewhat intermediate, but have wings as long germination. In addition, Keeley (1987) found that
as the keel. Many areas of overlap may be due to heating seeds in their pods to 120 oC for 5 minutes
natural secondary contact, but some are clearly due increased germination over that of unheated
to seeding projects along highways and utility controls.
corridors. The various floral forms are maintained
when plants are grown from seed in a common Genetics.—There are significant genetic
environment (Montalvo and Ellstrand 2000, 2001). differences among populations of the two varieties
of California broom (Montalvo and Ellstrand
Ecology.—In nature, seeds typically germinate 2000, 2001). An analysis of genetic marker data
after scarification by fire but some germinate in (13 allozyme loci) from three populations of var.
open, disturbed sites (Munz and Keck 1968, brevialatus and nine populations of var. scoparius
Keeley and Keeley 1984). Between 2 to 3 years showed significant population substructure due
after fire in sage scrub vegetation, following a primarily to differences among populations of the
burst of herbaceous species, California broom can two varieties. In an analysis of all populations, 18
become the dominant canopy species, eventually percent of the variation was due to differences
becoming replaced by long-lived shrub species. Its among populations, while analysis of just var.
abundance gradually decreases in 5 to 10 years brevialatus or var. scoparius populations showed
after fire. The branched tap root is relatively only 1 and 8 percent of the variation due to
shallow (< 1 m), and roots form symbiotic differences among populations. Thus, populations
associations with nitrogen fixing bacteria and within a variety are substantially more genetically
arbuscular mycorrhizal fungi (author’s similar to each other than to populations of the
observation). Plants are facultatively drought- other variety. Inbreeding coefficients were low for
deciduous, a trait commonly associated with all populations, a pattern consistent with
shallow-rooted shrubs of coastal sage scrub substantial cross pollination, severe loss of inbred
vegetation in California. Seasonality of leaf progeny, or both (mean f = 0.09, range = 0.00 –
production, nutrient accumulation, and leaf drop in 0.18) (Montalvo, Clegg, and Ellstrand, manuscript
response to summer drought has been studied in preparation).
extensively in L. scoparius var. scoparius (Nilsen
There is also genetically based geographic
and Muller 1980, 1981a,b, 1982, Nilsen and
variation in floral form in this species. In San
Schlesinger 1981, Nilsen 1982). Other researchers
Diego Co., Steppan (1991) detected distinct
have examined response to photoperiod
discontinuities in floral morphology between
(Comstock and Ehleringer 1986).
varieties and moderate correlations among
environmental variables and floral traits of wild
Reproduction.—Seeds of California broom
populations. Montalvo and Weaver (unpublished
germinate in mid to late winter during the rainy
data) measured floral traits on 12 populations of
season. Plants establish quickly with normal
plants raised in a common environment and made
rainfall and typically reach flowering size the
pairwise comparisons of genetic, floral,
second year. Flowering occurs primarily from
geographic, and environmental distances of source
March to June but may start as early as January in
populations. Floral morphological distance
warm, wet winters, and last much longer in more
correlated with environmental distance, genetic
moist, coastal areas and in years with long-lasting
distance correlated with floral distance, but genetic
soil moisture. Flowers are self-compatible and
distance did not correlate with geographic distance
insect-pollinated (Moldenke 1976, Hickman 1993,
446
until var. brevialatus was removed from the July depending on location. Pods or cleaned seeds
analysis. Results were consistent with floral form can be covered with boiling water and left to soak
having a genetic basis and with environmental to break dormancy (Atwater 1980, Young and
factors playing a role in the evolution of floral Young 1986, Emery 1988, Montalvo and Ellstrand
divergence. 2000, 2001), or exposed to dry heat (Keeley 1987).
Whole pods can be dry or wet broadcast in the fall
The substantial genetic differentiation of
for revegetation and restoration. Seedlings plugs
populations has implications for translocation of
can also be used for small projects. In greenhouse
California broom for restoration. Common garden
plantings, seeds germinate best when planted less
experiments testing plants originating from seed
than 2 mm deep (author’s observation). Despite
collected from 12 source populations (both
the strong geographic and morphological
varieties represented) demonstrated a significant
differences, the two varieties have been used
home site advantage (Montalvo and Ellstrand
indiscriminately in many seeding projects. Due to
2000). Furthermore, Montalvo and Ellstrand
the adaptive differences and the observed
(2001) directly tested the potential for
outbreeding depression following hybridization,
“outbreeding depression,” a loss of fitness upon
land managers should use the variety native to the
crossing genetically differentiated populations, by
planting location.
crossing individuals from six populations of the
two varieties in every combination and testing the
Benefits.—California broom is an important, fast
progeny in two common gardens at wild sites.
growing, early successional species used
Seeds per flower and seedling emergence
extensively in erosion control, post-fire mitigation,
decreased significantly with an increase in genetic
and habitat restoration in California, especially in
distance of the crossed parental populations.
the rapidly disappearing coastal sage scrub. The
Among variety crosses were only 70 percent as fit
flowers and seeds are an important food resource
as within variety crosses by the time seedlings
for a variety of insects and seed foraging rodents
emerged, and further fitness differences
and birds (Duncan 1968). The plants provide
accumulated after seedlings were outplanted into
valuable forage for deer, especially in drought
field plots. In these common gardens, success of
years when growth of herbaceous vegetation is
progeny decreased with increasing differences
sparse (Dale 2000).
between parental environments and the transplant
location. Results indicate that careful attention
References
should be given to the similarity of taxonomic
position and environment of source populations Atwater, B.R. 1980. Germination, dormancy and
relative to planting location, especially when morphology of the seeds of herbaceous
choosing source populations for restoration, ornamental plants. Seed Science and Technology
mitigation, and roadside landscaping. 8: 523-573.
Christensen, N.L. and C.H. Muller. 1975. Relative
Interspecific Hybridization.—Putative hybrids
importance of factors controlling germination and
have been reported between California broom and
seedling survival in Adenostoma chaparral. The
L. junceus (Benth.) Greene and L. benthamii Green
American Midland Naturalist 93: 71-78.
in central and northern California, as well as
between other members of the species complex Comstock, J. and J.R. Ehleringer. 1986.
(Isely 1981), but there have not been any genetic Photoperiod and photosynthetic capacity in Lotus
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(1990), however, did genetic studies on San 612.
Clemente Is. and documented hybridization
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447
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448
Ludwigia octovalvis (Jacq.) Raven primrose willow
ONAGRACEAE
John K. Francis
449
and Lakela 1976). In Texas, it is reported to bloom Correll, D.S. and M.C. Johnston. 1970. Manual of
from July through October (Correll and Johnston the vascular plants of Texas. Texas Research
1970). Seeds collected in Puerto Rico averaged 54 Foundation, Renner, TX. 1,881 p.
million per kg. Placed on moist filter paper, 28
percent of them germinated within 14 days, most Fish and Wildlife Information Exchange. 2002.
on the seventh day (author’s observation). Local Species moorhen, common, Hawaiian.
dispersal is by wind and water. Long-distance http://fwie.vt.edu/WWW/esis/lists/e101014.htm.
dispersal is probably accidental on birds, 17 p.
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Hodgdon Herbarium. 2002. Dicotyledons
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grows up to 1 m/year. Under favorable conditions, http://unh.edu/herbarium/bolivia/dicots.htm. 5 p.
plants live at least 3 years and possibly more.
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Control may be necessary in drainage ditches to Leeward and Windward Islands. Vol. 6. Arnold
maintain flow and in rice paddies. Control can be Arboretum, Harvard University, Jamaica Plain,
achieved by cultivation, hand pulling, and by MA. 658 p.
applying commercial weed killers.
Jones, F.B. 1975. Flora of the Texas Costal Bend.
Benefits and Detrements.—Primrose willow Mission Press, Corpus Christi, TX. 262 p.
grows in wet areas where few other plants will
grow and as such it aids in the successional Liogier, H.A. 1995. Descriptive flora of Puerto
process. It furnishes cover for wildlife. Its yellow Rico and adjacent islands. Vol. 4. Editorial de
flowers add a splash of color to areas often devoid la Universidad de Puerto Rico, San Juan, PR.
of colorfully flowering plants. In herbal medicines, 617 p.
preparations of the leaves are used to treat diarrhea,
dysentery, as a laxative and a vermifuge, and to Liogier, H.A. 1990. Plantas mediciales de Puerto
relieve headache, chest pains, and rheumatoid pain Rico y del Caribe. Iberoamericana de Ediciones,
(Burkill 1997, Liogier 1990, Parrotta 2001). A test Inc., San Juan, PR. 566 p.
of a local crude drug from Taiwan made from
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cariogenic bacteria Streptococcua mutans (Chen Tropical Florida. Banyon Books, Miami, FL.
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450
Lupinus arboreus Sims yellow bush lupine
FABACEAE
Synonyms: None
Christopher Ross
451
enrichment by lupine potentially alter fire regimes, Gadgil, R.L. 1971. The nutritional role of Lupinus
increasing frequency and intensity of burns. arboreus in coastal sand dune forestry. Plant
and Soil 35:113-126.
Growth and Management.—Yellow bush lupine
is fast growing and short lived (maximum 7 years) Hitchcock, C.L. and A. Cronquist. 1973. Flora of
(Davidson and Barbour 1977). Stands may grow the Pacific Northwest: an illustrated manual.
rapidly and die off in as little as 3 years (Strong et University of Washington Press. Seattle, WA.
al. 1995). Restoration of lupine-invaded sites 730 p.
should include litter and duff removal in addition
to direct removal of lupine, because it enriches soil Kittelson, P.M. and J.L. Maron. 2000.
to the detriment of native competitors (Pickart and Outcrossing rate and inbreeding depression in
others 1998). In New Zealand, where it is used as a the perennial yellow bush lupine, Lupinus
soil developer, it is controlled prior to tree planting arboreus (Fabaceae). American Journal of
by crushing and herbicides (Gadgil 1971). Coastal Botany 87: 652-660.
dune restoration in northern California has focused
heavily on removal of yellow bush lupine. Manual Maron, J.L. and P.G. Connors. 1996. A native
removal is accomplished by pulling up small nitrogen-fixing shrub facilitates weed invasion.
plants and chopping larger ones, which rarely Oecologia 105:302-312.
crown sprout. Heavy equipment has also been used
successfully, but all mechanical methods are labor Maron, J.L. and R.L. Jefferies. 1999. Bush lupine
intensive and expensive (Pickart and Sawyer mortality: Altered resource availability and
1998). Although others have noted a long-lived alternative vegetation states. Ecology 80: 443-
seed bank (see Maron and Connors 1996 and 454.
above), Pickart and Sawyer (1998) found no
recruitment by 3 years after removal of established Pickart, A.J., L.M. Miller, and T.E. Duebendorfer.
stands. 1998. Yellow bush lupine invasion in northern
California coastal dunes. Ecological impacts
Benefits.—Yellow bush lupine has been widely and manual restoration techniques. Restoration
cultivated as a sand stabilizer and ornamental in Ecology 6(1): 59-68.
coastal California. It is a strikingly beautiful plant,
both in and out of bloom, although its short life Pickart, A.J. and J.O. Sawyer. 1998. Ecology and
and messy litter probably limit its value as an Restoration of Northern California Coastal
ornamental. Elsewhere it has been used to prepare Dunes. California Native Plant Society.
coastal and dune soils for planting of Sacramento, California. 152 p.
commercially useful trees (Gadgil 1971). It
provides habitat for birds, reptiles, and rodents. Sholars, T. 1993. Lupinus Lupine in Hickman,
J.C., ed. 1993. The Jepson Manual: higher
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452
Lycium pallidum Miers pale wolfberry
SOLANACEAE
Synonyms: None
453
Growth and Management.—Pale wolfberry References
sprouts readily when cut or broken (Van Dersal
1938). Once established, shrub longevity may be Baldwin, B.G., S. Boyd, B.J. Ertter, R.W.
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Benefits.— Pale wolfberry is an important source
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Matthews, R. F. 1994. Lycium pallidum. U.S.D.A.
Forest Service, Rocky Mountain Research
Vines, R.A. 1986. Trees, shrubs, and woody vines
Station, Fire Sciences Laboratory. Fire Effects
of the Southwest. University of Texas Press.
Information System. http://www.fs.fed.
Austin, TX. 1,104 p.
us/databse/feis/plants.
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
McGregor, R. L., T.M. Barkley, RE Brooks, and Higgins. 1993. A Utah Flora. Second Edition,
EK Schofield. 1986. Flora of the Great Plains. revised. Brigham Young University, Provo,
Utah. 986 p.
455
Macfadyena unguis-cati (L.) A.H. Gentry cat’s claw
BIGNONIACEAE
John K. Francis
456
moderately shade tolerant as a young plant and still a handsome ornamental. It is often used to
grows in both full sun and under forest canopies. It cover fences or screen unsightly buildings. Cat’s
may be less shade tolerant as an adult. The species claw is recommended for areas where there is little
is common in savannas, secondary forest, and chance of invasion and especially in desert areas
remnant high forest. It can survive grazing and fire because it requires relatively little water for
but is eliminated by deep grass swards. maintenance (Desert-Tropicals 2001). It was
reported that a preparation made from cat’s claw
Reproduction.—In Hawaii, flowering takes place was used to treat dermatitis from Hippomane
during the spring (Rauch and Weissich 2000) mancinella L. (Michell and Rook 2001). A
whereas there are actually two periods of number of other uses in herbal medicine from
flowering in Puerto Rico, both during dry seasons diverse locations are cited by Liogier (1990).
(Acevedo-Rodríguez 1985). In Costa Rica, nearly
all the plants flower at once and flowering lasts References
just a short time (Gentry 1983). The capsules
mature about 6 months after flowering. The Acevedo-Rodríguez, P. 1985. Los bejucos de
winged seeds are dispersed by the wind. Seedlings Puerto Rico. Vol. 1. General Technical Report
are common and widespread in suitable habitat. SO-58. U.S. Department of Agriculture, Forest
Eleven capsules collected in Puerto Rico had a Service, Southern Forest Experiment Station,
maximum of 212 seeds and a minimum of 106 New Orleans, LA. 331 p.
seeds. These seeds (air dry) averaged 0.0224 ±
0.0005 g/seed or 45,000 seeds/kg. One hundred Aquatic Plant Control Operations Support Center.
percent of the seeds tested germinated between 49 2001. Macfadyena unguis-cati—catclaw vine.
and 95 days after sowing in commercial potting www.saj.usace.army.mil/conops/apc/newtt/cat1
mix. Young plants sprout when damaged and layer maps/macfadyenaunguis-cati.htm. 1 p.
(root) whenever stems touch the ground. Cat’s
claw, for ornamental uses, have been propagated Desert-Tropicals. 2001. Philippe and Sura’s
by seed, cuttings, and layering (Turner and Phoenix Tropical Gardens. http://www.desert-
Wasson 1999). tropicals.com. 3 p.
Growth and Management.—Stem extension is Florida Exotic Plant Council. 2001. Macfadyena
moderately rapid, especially in sprouts. Diameter unguis-cati (L.) A. Gentry. http://www.
growth of cat’s claw stems is slow. The vines are fleppc.org/pdf/Macfadyena%20unguis-cati.pdf.
long-lived, nearly as long as the trees that they 2 p.
claim for support. There are relatively few
problems in establishing cat’s claw. Insect and Gentry, A.H. 1983. Macfadyena unguis-cati (uña
disease problems are rare. The biggest problem is de gato, cat-claw, bignone). In: D.H. Janzen, ed.
controlling ornamental plants. Yearly pruning Costa Rican natural history. University of
after the flowering season is essential to Chicago Press, Chicago and London. p. 272-
maintaining them within the bounds desired. In its 273.
native range cat’s claw vines have only a minor
effect on the trees they parasitize. However, the Howard, R.A. 1989. Flora of the Lesser Antilles,
species is beginning to seriously suppress native Leeward and Windward Islands.
vegetation in parts of Florida (Florida Exotic Plant Dicotyledoneae. Vol. 6. Arnold Arboretum,
Council 2001) and Australia (Stockard 2001). The Harvard University, Jamaica Plain, MA. 658 p.
only means of control recommended at present is
to cut the vines and paint the cut ends with Liogier, H.A. 1990. Plantas medicinales de Puerto
glyphosate herbicide (Stockard 2001). However, Rico y del Caribe. Iberoamericana de Edicones,
Vélez and van Overbeek (1950) report that older Inc. San Juan, PR. 566 p.
plants are killed by simply cutting the stems. A
more practical long-term solution is hoped for in Liogier, H.A. 1995. Descriptive flora of Puerto
the ongoing biological control introductions in Rico and adjacent islands. Vol. 4. Editorial de la
several countries (Pest Cabweb 2001). Universidad de Puerto Rico, San Juan, PR.
617 p.
Benefits.—Despite the propensity to naturalize
and compete with native vegetation, cat’s claw is Michell, J. and A. Rook. 2001. Botanical
457
Dermatology Database. http://bodd.cf.ac.UK/ Stockard, J. 2001. The regeneration of Wingham
BotDermFolder/BotDermB/BIGN.html. 3 p. Brush, NSW. AABR lecture series. http://
www.zip.com.au/~aabr/info/seminars/seminar0.
Pacific Island Ecosystems at Risk. 2001. 4.html. 5 p.
Macfadyena unguis-cati (L.) Gentry,
Bigniniaceae. http://www.hear.org/pier/maung. Turner, R.J. and E. Wasson. 1999. Botanica.
htm. 2 p. Barnes and Noble, Inc., New York. 1020 p.
Pest Cabweb. 2001. Spotlight on biological control Vélez, I. and J. van Overbeek. 1950. Plantas
of weeds. http://pest.cabweb.org/Features/ indeseables en los cultivos tropicales. Editorial
Spotlight/spot00-2.htm. 21 p. Universitaria, Río Piedras, PR. 497 p.
Rauch, F.D. and P.R. Weissich. 2000. Plants for Watkins, J.V. and T.J. Sheehan. 1975. Florida
tropical landscapes, a gardener’s guide. University landscape plants, native and exotic. The
of Hawaii Press, Honolulu, HI. 139 p. University Presses of Florida, Gainesville, FL.
420 p.
458
Machaonia portoricensis Baill. alfilerillo
RUBIACEAE
Synonyms: none
John K. Francis
459
cover for wildlife. The wood, which is light- New York Botanical Garden. 2002. West Indian
brown, hard, and brittle, is good for fuel. vascular plant types catalog. http://
scisun.nybg.org:8890/searchdb/owa/wwwcatalo
References g.detail_list. [not paged].
Liogier, H.A. 1997. Descriptive flora of Puerto University of the Virgin Islands. 2002. Endangered
Rico and adjacent islands. Vol. 5. Editorial de la and threatened plants and animals in the U.S.
Universidad de Puerto Rico, San Juan, PR. Virgin Islands. http://rps.uvi.edu/CES/
436 p. endangered.html. 4 p.
Little, E.L., Jr., R.O. Woodbury, and F.H. Vázquez, O.J. and D.A. Kolterman. 1998. Floristic
Wadsworth. 1974. Trees of Puerto Rico and the composition and vegetation types of the Punta
Virgin Islands. Vol. 2. Agriculture Handbook Guaniquilla Natural Reserve—Cabo Rojo,
449. U.S. Department of Agriculture, Puerto Rico. Caribbean Journal of Science 34(3-
Washington, DC. 1,024 p. 4): 265-279.
460
Mahonia aquifolium (Pursh) Nutt. Oregon grape
BERBERIDACEAE
John K. Francis
General Description.—Oregon grape, also called Ecology.—Oregon grape is tolerant of shade and
tall mahonia, hollyleaved barberry, mountain normally grows in the forest understory. Natural
grape, Oregon grape-holly, and Oregon holly- stands contain scattered individual plants arising
grape, is an upright evergreen shrub from 0.6 to 3 from seeds and clumps of the species arising both
m in height. The stems are slender, stiff, usually from seed and suckers. The species grows well in
upright, and usually without branches. The wood soils with a wide variety of properties. However,
is whitish and brittle and has a significant pith. ornamentals do not do well in dry, wind-swept
Bark of young stems is green; older bark is rough sites, or soils with high pH’s (above 8.0). The tops
and gray-brown with yellow inner bark. The root are severely injured at –29 °C (Dirr 1983). It
systems produce horizontal rhizomes up to 1.8 m grows in areas with natural precipitation ranging
long from which suckers arise. The foliage from 950 to 5450 mm annually (Las Pilitas
attaches directly to the stems. Alternate, stiff and Nursery 2003). A number of diseases and insects
leathery, pinnately compound leaves have five to attack the leaves and fruits but usually do not
13 leaflets with a form like holly (coarse dentate cause serious damage.
with prickles). Leaves are dark green above,
except in winter, when exposed leaves take on a Reproduction.—Oregon grape blooms from April
purplish color. The species is monoecious with to June (Clark and Trelawny 1976). Flowers are
perfect, bright-yellow flowers in terminal self-fertile and pollinated by insects (Plants for a
(sometimes lateral) racemes. Flowers have a Future 2003). The fruits mature in late July
honey-like fragrance. Fruits (berries), 6 to 10 mm through September and remain on the plants into
long, ripen from green to blue or blue-black in late December (Center for Applied Nursery Research
summer (Abrams 1950, Center for Applied 2003). Fruits collected by the author from
Nursery Research. 2003, Dirr 1983). Oregon grape ornamental plants in Utah averaged 0.312 ± 0.013
is the state flower of Oregon (Paghat.com 2003). g/fruit. Air-dried seeds separated from them
averaged 0.0067 + 0.0002 g/seed or 149,000
Range.—Oregon grape is native to British seeds/kg. The number of seeds/fruit varied from
Colombia, Washington, Oregon, northern zero to nine. Fruits can be harvested in quantity by
hand and can be cleaned by gentile maceration,
461
wet sieving, and screening. Seeds should not be The fruit serves as a safe and gentle laxative
allowed to dry out completely and should be (Plants for a Future 2003).
stratified at 5 °C before sowing. Taken in
November and treated with 0.8 to 1.0 percent of References
IBA (Indolbuteric acid), cuttings root well.
Division of established plants is an alternate Abrams, L. 1950. Illustrated flora of the Pacific
method of vegetative propagation (Dirr 1983). In States. Vol. 2. Stanford University Press,
nature, mostly birds disperse the seeds Stanford, CA. 635 p.
(Paghat.com 2003) and both sexual and asexual
reproduction are effective (Auge and Brandl Auge, H. and R. Brandl. 1997. Seedling
1997). recruitment in the invasive clonal shrub,
Mahonia aquifolium (Pursh) Nutt. Oecologia
Growth and Management.—Oregon grape is Berlin 110(2): 205-211.
moderately slow growing. Individual stems reach a
60- to 90-cm height in 3 to 4 years (Dirr 1983). Center for Applied Nursery Research. 2003.
While individual stems probably have a life of not Mahonia aquifolium—Oregongrapeholly,
more than 10 years, clones can last much longer. Oregon Grapeholly, Oregon Hollygrape, Oregon
Plants are grown for the nursery trade in containers grape. http://www.nobleplants.com/classnotes/
and are usually 1 year old when sold. The species spring/springprofiles/evergreen/mahoniaaquifoli
is planted both in sunny and shady locations. um.htm. 2 p.
Ornamental plants tolerate pruning well (Plants for
a Future 2003). Data on management of the Clark, L.J. and J.G. Trelawny. 1976. Wild flowers
species in natural stands are lacking. of the Pacific Northwest. Gray’s Publishing
Limited, Sidney, B.C., Canada. 604 p.
Benefits.—Oregon grape shrubs contribute to the
diversity and beauty of understory vegetation in Dirr, M.A. 1983. Manual of woody landscape
native habitats, help protect the soil, and furnish plants: their identification, ornamental
food and cover for wildlife. The species is planted characteristics, culture, propagation and uses.
in temperate areas throughout the world as an Stipes Publishing Company, Champaign, IL.
ornamental, for hedges, and foliage, and leafy 826 p.
branches with their fall colors are sometimes used
for Christmas decorations. The fruits, edible but Felter, H.W. 1922. The eclectic material medica,
sour, are made into jellies and juice drinks (Clark pharmacology and therapeutics: Berberis
and Trelawny 1976). Flowers are eaten in salads, (Mahonia) aquifolium. published as a database
cooked in tempuras, and used to make a lemonade- in: http://www.ibiblio.org/herbmed/eclectic/
like drink (Paghat.com 2003). Yellow, green, felter/hahonia-aqui.html. 2 p.
violet, and purplish-blue dyes are made from
various tissues for arts and crafts (Plants For a Gieler, U., A. Von der Weth, and M. Heger. 1995.
Future 2003). Mahonia aquifolium—a new type of topical
treatment for psoriasis. Journal of
Herbal Medicine.—Infusions of Oregon grape Dermatological Treatment 6(1): 31-34.
root and doses of isolated alkaloid have been used
effectively in the past to relieve many of the Las Pilitas Nursery. 2003. Mahonia aquifolium.
symptoms of syphilis (Felter 1922). Ointments Las pilitas Nursery, Escondido, CA.
made from the plant and isolated alkaloids are http://www.laspilitas.com/plants/420.htm. 2 p.
currently enjoying tremendous popularity in
treating psoriasis and similar skin conditions. In Muller, K., K. Ziereis, and I. Gawlik. 1995. The
clinical trials, symptoms improved or disappeared anipsoriatic Mahonia aquifolium and its active
in 81 percent of 443 patients suffering from constituents: II. Antiproliferative activity against
subacute and chronic psoriasis (Gieler and others cell growth of human keratinocytes. Planta
1995). Of the constituent compounds tested, the Medica 61(1): 74-75.
benzylisoquinoline alkaloids berbamine and
oxyacanthine were principally responsible for Natural Resources Conservation Service. 2003.
inhibition of abnormal cell growth, with berberine Plants profile: Mahonia aquifolium (Pursh) Nutt.
showing lesser activity (Muller and others 1995). http://plants.usda.gov/cgi_bin/topics.cgi. 4 p.
462
Plants For a Future. 2003. Plants for a future:
Paghat.com. 2003. Paghat’s garden: Mahonia database search results: Mahonia aquifolium.
aquifolium. http://www.paghat.com/ http://www.ibiblio.org/pfaf/cgi-bin/arr_html
oregongrape.html. 4 p. ?Mahonia+aquifolium. 12 p.
463
Mahonia repens (Lindl.) G. Don creeping barberry
BARBERIDACEAE
John K. Francis
464
1991). It tolerates acidic soils but grows poorly State University Research and Extension 1996). It
when concentrations (less than 2.5 mg/L) of boron is included in reclamation plantings (Welsh and
are low (Harrison 2003). Creeping barberry grows others 1987) and has been used especially for
at elevations of 1,125 to 3,300 m in the inland revegetation of mine spoils (Harrison 2003). Mule,
West (Harrison 2003, Walkup 1991). Although it white-tailed deer, and sometimes elk browse
is most common on north-facing slopes, it grows creeping barberry during fall and winter. The
on south-facing slopes and ridge tops at higher protein content ranges from 4.7 to 5.5 percent, and
elevations. the species contains high contents of carotene in
July. Creeping barberry foliage is mildly toxic and
Reproduction.—Creeping barberry flowers in domestic animals normally make little use of it for
May through June and matures fruit in June forage except when other food is scarce (Walkup
through July (Rudolf 1974). The flowers are 1991). In one case, browsing of creeping barberry
pollinated by bees and butterflies (Paghat.com killed cattle in Arizona (Harrison 2003). Many
2003). Flowers not cross pollinated with other mammals (including black bears) and birds eat the
plants will self pollinate, but produce much fewer fruits (Walkup 1991). The fruits become edible to
seeds (Walkup 1991). Ratio weight of fresh fruits humans after they have passed through at least one
to air-dried seeds is 12.5:1 (Auger and others frost. They are used to make jelly, wine, and a
2002). There are 119,000 to 157,000 seeds/kg juice drink (Harrison 2003). Native Americans
(Rudolf 1974). Cleaned seed can be sealed and used the berries and the roots for lavender and
stored for 5 years (Harrison 2003). Rudolf (1974) yellow dyes, respectively, and made teas from the
recommends stratification of 30 days at 1 °C roots to treat a range of afflictions (Lesica 2002,
followed by 60 days at 21 °C, and then 196 days at Paghat.com 2003). Creeping barberry contains the
1 °C. Treated thus, 74 percent of test seeds alkaloids berberine and oxyacanthin (Harrison
germinated. Alternately, a hot water treatment 2003). The species is an alternate host for black
followed by cold stratification for 60 days is stem rust of cereal grains (Walkup 1991).
recommended (Kjelgren 2003). Seeds can be sown
in fall without pretreatment. In nature, seeds are References
dispersed by birds and to a lesser extent by
mammals including black bears and rodents Auger, J., S.E. Meyer, and H.L. Black. 2002. Are
(Auger and others 2002). Once established, plants American black bears (Ursus americanus)
spread through sprouts from rhizomes (Walkup legitimate seed dispersers for fleshy-fruited
1991). shrubs? American Midland Naturalist 147: 352-
367.
Growth and Management.—Growth is slow.
Individual stems may live 10 years or more. It is Harrison, S. 2003. Mahonia repens. Crop
difficult to imagine the need to control this Development Centre, University of
unobtrusive species in its native habitat. On the Saskatchewan, Saskatoon, Saskatchewan,
other hand, it should frequently be planted as part Canada. http://www.usask.ca/agriculture/
of environmental restoration projects. The species plantsci/classes/range/berberis.html. 4 p.
can be artificially propagated by seed, divisions,
cuttings, and layers (Walkup 1991). Direct seeding Kansas State University Research and Extension.
in the fall is the most practical method to introduce 1996. Low water use plants for Kansas
it to disturbed sites. Repellants or poisons may be landscapes. MF2067. Kansas State University,
necessary because the seeds are readily consumed Manhattan, KS. 4 p.
by mice (Kjelgren 2003).
Kjelgren, R. 2003. Mahonia repens. Utah State
Benefits.—Creeping barberry adds beauty to the University, Logan, UT. http://www.hort.usu.
forest, helps protect the soil, provides food for edu/natives/shrubs/repens.htm. 1 p.
wildlife, and makes a useful ornamental. It is
planted as ground cover, especially under light tree Lesica, P. 2002. A flora of Glacier National Park,
canopy cover or on poor, gravelly ground. In the Montana. Oregon State University Press,
summer, the leaves are dark green; in winter they Corvallis, OR. 512 p.
are green, mottled with bright red, or a bonze-
purple. The species is particularly recommended Natural Resources Conservation Service. 2003.
because it requires little or no irrigation (Kansas Plants profile: Mahonia repens (Lindl.) G. Don,
465
creeping barberry. http://plants.usda.gov/ woody plants of the United States. Agric.
cgi_bin/plant_search.chi?mode=Scientific+Nam Handb. 450. U.S. Department of Agriculture,
e&keywordquerry=Mahonia+repens&earl=plant Forest Service, Washington, DC. p. 247-251.
_search.cgi. 5 p.
Walkup, C.J. 1991. Mahonia repens. In: U.S.
Paghat.com. 2003. Creeping mahonia, aka Department of Agriculture, Forest Service,
creeping barberry, aka creeping hollygrape, aka Rocky Mountain Research Station, Fire Sciences
ash barbarry, aka small or dwarf Oregon grape. Laboratory. Fire Effects Information System.
http://www.paghat.com/oregongrape2.html. 4 p. http://www.fs.fed.us/database/feis/plants/shrub/
mahrep/all.html. 10 p.
Rehder, A. 1951. Manual of cultivated trees and
shrubs. The MacMillan Company, New York. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
996 p. Higgins. 1987. A Utah flora. Great Basin
Naturalist Memoirs 9. Brigham Young
Rudolf, P.O. 1974. Berbers L., barberry, mahonia. University Press, Provo, UT. 894 p.
In: C.S. Schopmeyer. tech. coord. Seeds of
466
Malvastrum americanum (L.) Torr. Indian Valley false mallow
MALVACEAE
John K. Francis
467
level to 1000 m in elevation in Nicaragua (Stevens Howard, R.A. 1989. Flora of the Lesser Antilles,
and others 2001). Leeward and Windward Islands. Vol. 5. Arnold
Arboretum, Harvard University, Jamaica Plain,
Reproduction.—Indian Valley false mallow is MA. 604 p.
reported to flower and fruit throughout the year in
Nicaragua (Stevens and others 2001). In Puerto Institute of Systematic Botany. 2003. Atlas of
Rico, plants begin flowering at about 15 cm in Florida vascular plants: Malvastrum
height and continue through life except for pauses americanum. http://www.plantatlas.usf.edu/
during dry seasons. Air-dried seeds collected in main.asp?plantID=3662. 2 p.
Puerto Rico averaged 1.19 million seeds/kg.
Placed on moist filter paper without pretreatment, Lavia, G., G. Seijo, A. Fernández, and A.
15 percent germinated over a period of 6 months. Krapovickas. 2000. Los cariotipos de algunas
No specialized means of dispersal is apparent; the especies de Malvastrum A. Gray (Malvaceae).
seeds are small and undoubtedly are moved to Universidad Nacional del Nordeste, Corrientes,
some extent by wind, water, and animals. Argentina. http://www.unne.edu.ar/cyt/2000/
Seedlings are relatively common in favorable sites. 6_biologicas/6_pdf/6_034.pdf. 3 p.
The dry fruits can easily be collected in quantity
by hand but cleaning the tiny seeds is difficult. Liogier, H. A. 1994. Descriptive flora of Puerto
Rico and adjacent Islands. Vol. 3. Editorial de la
Growth and Management.—Growth rates are Universidad de Puerto Rico, Río Piedras, PR.
moderate (0.3 to 0.5 m/year) and plants live 1 to 5 461 p.
years, depending on competition and growing
conditions. Although control is occasionally Mensah, R., M. Dillon, M. Kahn, C. Tann, and L.
needed in rangelands and disturbed areas, no Wilson. 2003. Cotton insect pests and their weed
specific recommended treatments are known to the hosts. Australian Cotton Cooperative Research
author. Centre. http://www.cotton.pi.csiro.au/Assets/
PDFFiles/IPMGL99/IPMSO4.pdf. 2 p.
Benefits and Detriments.—Indian Valley false
mallow helps protect the soil, furnishes cover for Natural Resources Conservation Service. 2003.
wildlife, and because of its blooms contributes Plants profile: Malvastrum americanum (L.)
moderately to the aesthetics of wildlands where it Torr., Indian Valley false mallow. http://
grows. It is a host for the cotton bollworm, plants.usda.gov/cgi_bin/plant_search.cgi?mode=
Helicoverpa armigera (Hubner) (Mensah and Scientific+Name&keywordquery=Malvastrum+
others 2003). The species also is the larval food- americanum&earl=plant. 3 p.
plant for the Laviana white-skipper butterfly,
Heliopetes laviana (Hewitson) of Texas (Quinn Quinn, M. 2003. Caterpillar food plants for the
2003) and a Cuban butterfly, Strymon columella lower Rio Grande Valley. http://www.naba.
cybira (Hewitson) (Fernández 2001). org/ftp/plants_info.pdf. 8 p.
468
Mecranium latifolium (Cogn.) Skean camasey almendro
MELASTOMATACEAE
John K. Francis
469
averaged 0.138 ± 0.005 g/fruit. The seeds are tiny; ornamental in natural landscaping if it proves
an air-dried sample averaged 25.4 million/kg. The adaptable at lower elevations where the island
seeds can be cleaned by crushing, wet sieving with urban centers are located.
a fine mesh screen, filtering and drying on the
paper filter, and rubbing the seed off the paper. References
Without pretreatment seeds took 4 months to begin
germinating and completed 11 percent germination Liogier, H.A. 1995. Descriptive flora of Puerto
in 11 months. Germination is epigeal, and Rico and adjacent islands. Vol. 4. Editorial de la
seedlings are about 1 mm across the two cotyledon Universidad de Puerto Rico, San Juan, PR.
leaves. It is assumed that birds eat the fruits and 617 p.
disperse the seeds. Seedlings are scattered to
relatively common in suitable, disturbed habitat. Little, E.L., Jr., R.O. Woodbury, and F.H.
Layering (rooting) occurs whenever horizontal Wadsworth. 1974. Trees of Puerto Rico and the
stems come in contact with the ground. Camasey Virgin Islands. Vol. 2. Agriculture Handbook
almendro sprouts when stems are broken or cut. 449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
Growth and Management.—Camasey almendro
grows moderately slowly and probably lives at McKenny, S. 2002. DNA evidence for the origin
least 2 or 3 decades. No planting or management of Mecranium latifolium (Melastomataceae), a
experience has been published. Removal of the Puerto Rican endemic. Abstract in: Elkin Isaac
forest canopy and soil disturbance near seed Research Symposium, Apr. 18, 2002, Albion
sources would likely result in reproduction. College, Albion, MI. 1 p.
Benefits.—Camasey almendro helps protect the Weaver, P.L. 2000. Elfin woodland recovery 30
soil and furnishes food and cover for wildlife. It is years after a plane wreck in Puerto Rico’s
an attractive plant and adds to the aesthetics of the Luquillo Mountains. Caribbean Journal of
forest. The species might be suitable as an Science 36(1-2): 1-9.
470
Melochia nodiflora Sw. malva colorada
STERCULIACEAE
John K. Francis
471
and Uranga 1993). The species is controlled by Howard, R.A. 1989. Flora of the Lesser Antilles,
mowing, cutting with machete, and by spraying Leeward and Windward Islands. Vol. 5. Arnold
with broadleaf weed killers. Arboretum, Harvard University, Jamaica Plain,
MA. 604 p.
Benefits.—Malva colorada is browsed by goats
and to a limited extent by cattle. It protects against Liogier, H.A. 1994. Descriptive flora of Puerto
erosion in recently disturbed areas. Rico and adjacent islands. Vol. 3. Editorial de la
Universidad de Puerto Rico. Río Piedras, PR.
References 461 p.
Grisebach, A.H.R. 1864. Flora of the British West Sánchez, P. and H. Uranga. 1993. Plantas
Indian Islands. J. Cramer-Weinheim, New York. indeseables de importancia economica en los
789 p. cultivos tropicales. Editorial Científico-Técnisa.
La Habana, Cuba. 166 p.
472
Melochia tomentosa L. pyramid bush
STERCULIACEAE
John K. Francis
473
Sown without pretreatment on moist filter paper, Agropecuaria do Tropico Semi-Arido. 28 p.
79 percent germinated within a short period 6
months later. The species can be propagated with Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs,
cuttings (San Antonio Botanical Gardens 2002). and cacti of South Texas. Texas Tech University
There appear to be no specialized means of seed Press. Lubbock, TX. 213 p.
transport; the small seeds are probably disbursed
incidentally by wind, water, machinery, and cattle. Howard, R.A. 1989. Flora of the Lesser Antilles,
Leeward and Windward Islands. Vol. 5. Arnold
Growth and Management.—Nursery-grown Arboretum, Harvard University, Jamaica Plain,
pyramid bush seedlings in Texas reached 60 cm in MA. 604 p.
just 2 months after outplanting (San Antonio
Botanical Gardens 2002). However, adult plants in Instituto Nacional de Biodiversidad. 2002. The
Puerto Rico add only about 30 cm to their height cutting edge. Vol. 7, No. 3. http://www.
each year. Based on ring counts in Puerto Rico, inbio.ac.cr/papers/manual_plantas/jul00/jul00lea
plants appear to live about 10 years. Overgrazing .html. 1 p.
to reduce competition may be the best way of
promoting this species. Pyramid bush is often Liogier, H.A. 1994. Descriptive flora of Puerto
considered a weed in rangeland (Liogier 1990). Rico and adjacent islands. Vol. 3. Editorial de la
Mowing may be sufficient to control it. Universidad de Puerto Rico. Río Piedras, PR.
461 p.
Benefits.—Pyramid bush appears to have great
potential as a flowering ornamental for xeric Liogier, H.A. 1990. Plantas medicinales de Puerto
landscaping (San Antonio Botanical Gardens Rico y del Caribe. Iberoamericana de Ediciones,
2002). In natural habitat, it contributes to Inc., San Juan, PR. 566 p.
biodiversity, soil stability, and wildlife cover.
Although cattle avoid it, sheep and goats in Brazil Long, R.W. and O. Lakela. 1976. A flora of
browse it in mixture with other native species Tropical Florida. Banyon Books, Miami, FL.
(Charles and others 1983). A tea is made from the 962 p.
foliage to treat colds and as an eye wash (Liogier
1990). Recursos Hidricos. 2002. Plano diretor de recursos
hídricos das bacias hidrográficas dos tributaries
References estaduaia dos rios vazas barris e real. Vol. 3, part
4.1. http://hidricosw.mg.gov.br/vazareal/vol3/
Charles, T.N.P., A.M. Maia, C. Guimaraes Filho, 41.htm. 4 p.
L.M.C. Salviano, and E.A.P. de Figueiredo.
1983. Effect of roughage supplementation and San Antonio Botanical Gardens. 2002. Friendly
mineral mixture plus deworming on sheep and natives for Texas. http://www.sabot.org/
goat production. II. Lamb and kid growth rates. natives/native.html. 3 p.
Boletim de Pesquisa No. 20. Centro de Pesquisa
474
Miconia impetiolaris (Sw.) D. Don ex DC. camasey de costilla
MELASOMATACEAE
John K. Francis
475
2.3 million seeds/kg. Sown on wet peat, 56 References
percent of the seeds germinated between 13 and
23 days after sowing. The new seedlings are tiny Croat, T.B. 1978. Flora of Barro Colorado Island.
and develop slowly (author’s observation). Stanford University Press, Stanford, CA. 943 p.
Growth and Management.—Camasey de costilla Howard, R.A. 1989. Flora of the Lesser Antilles,
is relatively slow growing. About 0.25 to 0.5 m of Leeward and Windward Islands. Vol. 5. Arnold
height is added annually. Plants live at least 10 Arboretum, Harvard University, Jamaica Plain,
years and probably much longer. No known MA. 604 p.
attempts have been made to plant or manage
camasey de costilla. Probably management Liogier, H.A. 1995. Descriptive flora of Puerto
practices that maintain forests interspersed with Rico and adjacent islands. Vol. 4. Editorial de
pastures, brushlands, and roadsides will ensure at la Universidad de Puerto Rico, San Juan, PR.
least a presence of camasey de costilla. 617 p.
476
Miconia prasina (Sw.) DC. camasey blanco
MELASTOMATACEAE
John K. Francis
477
and persists as small individuals until the pastures goats. The fruits are eaten by the Puerto Rican
are abandoned, after which they grow rapidly and birds, Mimocichla ardosiaceae portoricensis
dominate for 10 to 20 years (Pascarella 2002). It Bryant, Vircosylva calidris calidris L., Spindalis
usually grows as dispersed individual shrubs or portoricensis Bryant (Whetmore 1916) and
small clumps. Camasey blanco disappears in a few certainly many other species across the range.
seasons after being overtopped by a forest canopy. Camasey blanco serves as a transitional
(successional) species between pasture, farmland,
Reproduction.—Camasey blanco flowers from or disturbed forest and secondary forest.
February to June and fruits from April to
September in Nicaragua (Stevens and others References
2001). The species is reported to flower and fruit
twice each year in Panama (Croat 1978). In Puerto Croat, T.B. 1978. Flora of Barro Colorado Island.
Rico, it is reported to flower and fruit almost Stanford University Press, Stanford, CA. 943 p.
throughout the year (Little and Wadsworth 1964).
Fruits collected in Puerto Rico averaged 0.137 ± Englerth, G.H. and E. Goytía-Olmedo. 1960.
0.003 g/fruit. Seeds separated from them averaged Preservation of Puerto Rican fence posts treated
10.6 million seeds/kg. Sown on wet peat, 57 by cold soaking and hot-and-cold bath method.
percent of the seeds germinated between 20 and Tropical Forest Notes 2. U.S. Department of
63 days after sowing. The seedlings are tiny and Agriculture, Forest Service, Tropical Forest
difficult to manage. In nature, birds are the Recearch Center, Río Piedras, PR. 4 p.
principal dispersers of seeds. Camasey blanco
sprouts readily when disturbed.
Francis, J.K. 2000. Estimating biomass and carbon
content of saplings in Puerto Rican secondary
Growth and Management.—The largest
forest. Caribbean Journal of Science 36(3-4):
camasey blanco recorded in Puerto Rico had a
346-350.
diameter at breast height of 16.3 cm and a height
of 11 m (Puerto Rico Champion Tree List on file
Granzow-de la Cerdal, I., N. Zamora, J.
at the International Institute of Tropical Forestry,
Vandermeer, and D. Boucher. 2002. Diversidad
Río Piedras, PR). The species has a moderate
de especies arbóreas en el bosque tropical
growth rate and is relatively short lived. None of a
húmedo del Caribe nicaragüense siete años
group of 24 camasey blanco stems in a survey in
después del huracán Juana. http://www.rbt.
Puerto Rico were present 24 years later at a
ac.cr/revistas/45-4/granzow.htm. 15 p.
second visit (Weaver 1979). However, by
suckering and ground layering, plants can
Howard, R. A. 1989. Flora of the Lesser Antilles,
perpetuate themselves somewhat longer than
Leeward and Windward Islands. Vol. 5. Arnold
individual stems. Nursery propagation of the
Arboretum, Harvard University, Jamaica Plain,
species has not been reported.
MA. 604 p.
Benefits.—Camasey blanco wood is not durable
in the ground and therefore not suitable as fence Killeen, T.J., E. García-E., and D.G. Beck. 1993.
posts (Little and Wadsworth 1964), unless treated. Guía de árboles de Bolivia. Herbario Nacional
Penetration of preservative was about average for de Bolivia and Missouri Botanical Garden, La
52 Puerto Rican species tested (Englerth and Paz and St. Louis, MO. 958 p.
Goytía-Olmedo 1960). The wood is used for fuel
and occasionally as tool handles. Wood specific Liogier, H.A. 1995. Descriptive flora of Puerto
gravity of 30 plants sampled in Puerto Rico was Rico and adjacent islands. Vol. 4. Editorial de la
measured at 0.630 + 0.021. A weighted average of Universidad de Puerto Rico, San Juan, PR. 617
carbon content for above-ground parts was 50.0 p.
percent. Total above-ground biomass in grams
may be predicted by multiplying the stem Little, E.L., Jr. and F.H. Wadsworth. 1964.
diameter in cm squared at 30 cm above the soil Common trees of Puerto Rico and the Virgin
times stem length in meters by 38.344 (R2 = Islands. Agriculture Handbook 249. U.S.
0.981) (Francis 2000). The fruits are edible but Department of Agriculture, Forest Service,
seldom eaten by people. Camasey blanco is Washington, DC. 548 p.
browsed sparingly by cattle and more heavily by
478
Lugo, A.E. 1992. Comparison of tropical tree Monographs in Systematic Botany, Vol. 85, No.
plantations with secondary forests of similar 2. Missouri Botanical Garden, St. Louis, MO.
age. Ecological Monographs 62: 1-41. pp. 945-1,910.
Missouri Botanical Garden. 2002. Flora of Costa Weaver, P.L. 1979. Tree growth in several tropical
Rica: Miconia prasina. http://www.mobot.org/ forests of Puerto Rico. Research Paper SO-152.
manual.plantas/023023/S024494.html. 2 p. U.S. Department of Agriculture, Forest Service,
Southern Forest Experiment Station, New
Pascarella, J.B. 2002. Tropical forest succession in Orleans, LA. 15 p.
Puerto Rico. http://chiron.valdosta.edu/
jbpascar/Research/Prico/puerto_rico.htm. 6 p.
Whetmore, A. 1916. Birds of Porto Rico. Bulletin
326. U.S. Department of Agriculture.
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Washington, DC. 140 p.
Montiel, eds. 2001. Flora de Nicaragua.
479
Miconia racemosa (Aubl.) DC. camasey de felpa
MELASTOMATACEAE
John K. Francis
480
abundant, the species is common at least in Puerto regeneration in openings and beneath closed
Rico, growing in small clumps or as scattered canopy in sub-tropical wet forest. Ph.D.
individuals. Camasey de felpa coppices readily dissertation, Yale University, New Haven, CN.
when cut. 307 p.
Growth and Management.—Camasey de felpa Fundaçao Andre Tosello. 2002. Base de dados
has a moderate growth rate and probably lives 10 tropicles: Espécies arbóreas da Mata Atlântica.
to 20 years. No management experience has been http://www.bdt.fat.org.br/mata.atlantica/flora/
published. Natural reproduction could be local?SC. 5 p.
encouraged by disturbance of moist and wet forest
and unwanted plants eliminated by grubbing or
Howard, R.A. 1989. Flora of the Lesser Antilles,
spraying with broadleaf herbicides.
Leeward and Windward Islands. Vol. 5. Arnold
Arboretum, Harvard University, Jamaica Plain,
Benefits.—Camasey de felpa contributes to the
MA. 604 p.
aesthetics of forests where it grows, helps
revegetate disturbed sites and protect the soil, and
Liogier, H.A. 1995. Descriptive flora of Puerto
provides food and cover for birds and other
Rico and adjacent islands. Vol. 4. Editorial de la
wildlife. The fruits are juicy and edible but small
Universidad de Puerto Rico, San Juan, PR.
and almost tasteless (Little and others 1974). The
617 p.
wood is useful for fuel, small fence posts, and
stakes.
Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
References
Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Centro Nordestino de Informaçõas Sobre Plantas.
Washington, DC. 1,024 p.
2002. Checklist das plantas do Nordeste.
Universidade Federal de Pernambuco.
Myster, R.W. 1997. Seed predation, disease and
http://umbuzeiro.cnip.org.br/db/pnechk/fam/
germination on landslides in neotropical lower
f104.shtml. 6 p.
montane wet forest. Journal of Vegetation
Science 8(1): 55-64.
Devoe, N.N. 1989. Differential seeding and
481
Mimosa aculeaticarpa Ortega wait-a-bit
FABACEAE
Juanita A. R. Ladyman
482
are pollinated by insects. Only a few flowers in References
each inflorescence produce pods (Bowers 1993).
The fruit is an elongated flattened legume (pod), Ahlstrand, G.M. 1982. Response of Chihuahuan
the margins separate from the valves at Desert Mountain Shrub Vegetation to Burning.
dehiscence. The valves often break transversely Journal of Range Management. 35 (1): 62-65.
into one-seeded sections. The seeds exhibit high
germination rates over a relatively wide range of Barneby, R.C. 1991. Sensitivae Censitae. Memoirs
temperatures (Jordan and Haferkamp 1989). of the New York Botanical Garden Vol. 65.
Livestock and wildlife are likely to disperse the The New York Botanical Garden, Bronx, NY.
seeds (Uchytil 1990).
Bowers, J.E. 1993. Shrubs and Trees of the
Growth and Management.—Catclaw mimosa
Southwest Deserts. Southwest Parks and
forms true brier patches that may become almost
Monumens Association, Tucson, AZ. 140 p.
impenetrable to humans (author’s personal
observation). Plants can profusely re-sprout from
Buechner, H.K. 1950. Life history, ecology, and
protected buds following fire-induced mortality of
range use of the pronghorn antelope in Trans-
aboveground tissue (Cable 1975, Carmichael and
Pecos Texas. American Midland Naturalist.
others 1978, McPherson 1995). However, in
43(2): 257-354.
general, plants must be several years old before
they have the ability to re-sprout. Re-sprouting
Cable, D.R. 1975. Range management in the
decreases as soil-moisture decreases and is less
chaparral type and its ecological basis: the status
common after fires that occur during the growing
of our knowledge. Research Paper RM-155.
season compared with dormant-season fires
U.S. Department of Agriculture, Forest Service,
(McPherson 1995). Stands can recover from
Rocky Mountain Forest and Range Experiment
burning in 5 years (Ahlstrand 1982, Kittams
Station, Fort Collins, CO. 30 p.
1972). However, at least in Arizona, wildfire does
not appear to stimulate sprouting (Pase and Pond
Carmichael, R.S., Knipe, O.D., Pase, C.P. and
1964). Catclaw mimosa is moderately resistant to
Brady, W.W. 1978. Arizona chaparral: plant
phenoxy herbicides and after only one application
associations and ecology. Research Paper RM-
tend to re-foliate or re-sprout from the crown
202. U.S. Department of Agriculture, Forest
(Hibbert and others 1974). Successive applications
Service, Rocky Mountain Forest and Range
kill the plants.
Experiment Station, Fort Collins, CO. 16 p.
Benefits.—Catclaw mimosa thickets provide
Carr, M.E., Mason, C.T., Jr., and Bagby, M.O.
cover and food for a variety of small wildlife
1986. Renewable resources from Arizona trees
species, especially quail (Graham 1941, Powell
and shrubs. Forest Ecology and Management.16:
1998). The flowers are a good source of nectar for
155-167.
honey bees (Powell 1998, Vines 1986). Deer and
pronghorn browse Catclaw mimosa and its
Carter, J.L. 1997. Trees and Shrubs of New
palatability is rated good (Nichol 1938, Buechner
Mexico. Johnson Books, Boulder, CO. 533 p.
1950). Its livestock palatability is low (Dayton
1931). However, livestock use it when other
Dayton, W.A. 1931. Important western browse
forage is scarce, and by virtue of its abundance in
plants. Miscellaneous Publication 101. U.S.
New Mexico and Arizona it has been described as
Department of Agriculture, Washington, DC.
important cattle browse (Dayton 1931, Vines
214 p.
1986). Both livestock and wildlife eat the pods
(Dayton 1931, Cable 1975, Uchytil 1990).
Dick-Peddie, W.A. 1993. New Mexico
Because plants have a tendency to form thickets
Vegetation--past, present, and future.
that effectively bind soil, Catclaw mimosa shows
University of New Mexico Press, Albuquerque,
potential for erosion control (Kearney and others
NM. 244 p.
1960, Vines 1986). In a comparison with 100 other
plant species studied for the production of fuels
Dick-Peddie, W.A. and W.H. Moir. 1970.
and chemicals, it yielded substantial amounts of
Vegetation of the Organ Mountains, New
oils, polyphenols, and hydrocarbons (Carr and
Mexico. Range Science Department Science
others 1986).
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Escarpment, New Mexico-Texas. Ecology 48: climatic types in Nuevo Leon, Mexico.
793-806. American Midland Naturalist 21: 687-729.
Gledhill, D. 1992. The names of plants. Second Muller, C.H. 1947. Vegetation and Climate of
Ed. Cambridge University Press, Cambridge Coahuila, Mexico. Madrono. 9: 33-57.
England. 202 p.
Nichol, A.A. 1938. Experimental feeding of deer.
Graham, E.H. 1941. Legumes for erosion control Technical Bulletin 75. Arizona Agriculture
and wildlife. Miscellaneous Publication 412. Experiment Station. 39 p.
U.S. Department of Agriculture, Washington,
DC. 153 p. Pase C.P. and F.W. Pond. 1964. Vegetation
changes following the Mingus Mountain burn.
Hibbert, A.R., Davis, E.A., and Scholl, D.G. 1974. Note RM-18. U.S. Department of Agriculture,
Chaparral conversion potential in Arizona: Part Forest Service, Rocky Mountain Station, Fort
I: water yield response and effects on other Collins, CO. 8 p.
resources. Research Paper RM-126. U.S.
Department of Agriculture, Forest Service, Pase, C.P. and D.E. Brown. 1982. Interior
Rocky Mountain Forest and Range Experiment chaparral. In: D.E. Brown, ed. Biotic
Station, Fort Collins, CO. 36 p. Communities of the American Southwest—
United States and Mexico. Desert Plants. 4 (1-
Jordan, G.L. and M.R Haferkamp. 1989. 4): 95-99.
Temperature responses and calculated heat units
for germination of several range grasses and Powell, A.M. 1998. Trees & shrubs of Trans-
shrubs. Journal of Range Management. 42(1): Pecos and adjacent areas. (First Ed. copyright
41-45. Big Bend Natural History Assoc.) University of
Texas University Press, Austin, TX 498 p.
Kartesz, J.T. 1994. A synonymized checklist of the
vascular flora of the United States, Canada and Shreve, F. 1915. The vegetation of a desert
Greenland. Vol.1-Checklist. Second ed. Timber mountain range as conditioned by climatic
Press Portland, OR. 622 p. factors. Publication 529. Carnegie Institute,
Washington, DC. p. 1-45.
Kearney, T.H., Peebles, R.H. and collaborators.
1960. Arizona flora. 2nd ed. University of Uchytil, R.J. 1990. Mimosa biuncifera. U.S.
California Press, Berkeley, CA. 1,085 p. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences
Kittams, W.H. 1972. Effect of fire on vegetation of Laboratory. Fire Effects Information System.
the Chihuahuan Desert region. Proceedings: http://www.fs.fed.us/database/feis/.
Tall Timbers Fire Ecology Conf. A Quest for
Ecological Understanding 12: 427-444. Vines, R.A. 1986. Trees, shrubs, and woody vines
of the Southwest. University of Texas Press,
Martin, W. C. and C.R. Hutchins. 1981. A Flora of Austin, TX. 1,104 p.
New Mexico. Volume 2. J. Cramer, Vaduz,
Germany. p. 1,277-2,591. Whittaker, R.H. and W.A. Niering. 1965.
Vegetation of the Santa Catalina Mountains,
McPherson, G.R. 1995. The role of fire in desert AZ: a gradient analysis of the south slope.
grasslands. In: M. P. McClaran and T. R. Van Ecology 46: 429-452.
484
Mimosa arenosa (Willd.) Poir. tepehuiste
FABACEAE
John K. Francis
485
same period. Insects pollinate the flowers. Fruits References
collected in Puerto Rico averaged 7.0 + 0.2
seeds/pod. Seeds separated from them weighed an Liogier H.A. 1988. Descriptive flora of Puerto
average of 0.0054 ± 0.0006 g/seed or 186,000 Rico and adjacent islands, Spermatophyta. Vol.
seeds/kg. One percent of unscarified seed 2. Editorial de la Universidad de Puerto Rico,
germinated in 6 months. but a lot of mechanically Río Piedras, PR. 481 p.
scarified seed germinated at 99 percent between 3
and 5 days after sowing on moist filter paper Missouri Botanical Garden. 2002. VAST
(author’s observation). Germination is epigeal nomenclatural database. http://mobot.mobot.
(Parra 1984). Plants sprout when cut or burned. org/cgi-bin/search_vast?name=Mimosa
+arenosa. [not paged].
Growth and Management.—Tepehuiste grows
rapidly. Because the species is thorny, forms New York Botanical Garden. 2002. Neotropical
almost impenetrable thickets, and aggressively flora mycota catalog. http://scisun.nybg.org:
invades pastures and rangeland, it is a highly 8890/searchdb/owa/wwwspecimin.search_list?
undesirable species in exotic habitats. Every effort taxon=Mimosa+arenosa&projcode=NETR. [not
should be made to prevent it from invading new paged].
areas. Control techniques have not been published.
However, the same measures used against Mimosa Nunez de Medeiros, A. 2002. Caprinocultura de
pigra L. probably would be effective. Corte no Nordeste Brasileiro. Departamento de
Agropecuária, Bananeiras, PB, Brazil.
Benefits.—Tepehuiste helps sites move through http://www.capritio.com.br/art18.htm. [not
the grass stage to secondary forest, protects the paged].
soil, and provides cover for wildlife. It is an
important forage plant for goats in northeastern Parra-G., P. 1984. Estudio de la morfologia
Brazil (Nunez de Medeiros 2002). The wood is externa de plantulas de Calliandra gracilis,
useful for fuel. Mimosa albida, Mimosa arenosa, Mimosa
camporum y Mimosa tenuiflora. Revista de la
Facultad de Agronomía (Maracay) 8(1-4): 311-
350.
486
Mimosa ceratonia L. zarsa
FABACEAE
John K. Francis
487
when cut and lateral roots sometimes sucker when Barneby, R.C. 1985. The identity and synonymy
damaged. The seeds are disbursed by lateral of Acacia guiladinae, Mimosa obovata, M.
extension of the vines and by the pods clinging to pseudo-obovata and M. laticifera (Mimosaceae).
clothing or to the fur of animals. Seedlings are Brittonia 37(1): 85-87.
common, but only a few survive for more than a
few weeks. Howard R.A. 1988. Flora of the Lesser Antilles,
Leeward and Windward Islands.
Growth and Management.—Both seedlings and Dicotyledoneae, Part 1. Vol. 4. Arnold
sprouts grow rapidly, up to 2 or 3 m per year. Arboretum, Harvard University, Jamaica Plain,
Zarza plants may endure for many years. Because MA. 673 p.
of the thorny nature of this species, property
owners often choose to eliminate it from pastures Liogier H.A. 1988. Descriptive flora of Puerto
and plantations. Historically, farmers controlled it Rico and adjacent islands, Spermatophyta. Vol.
by repeated cutting. This method could probably 2. Editorial de la Universidad de Puerto Rico,
be improved by treating the resulting sprouts with Río Piedras, PR. 481 p.
broadleaf weed killers.
Marcano F., E. de J. 1973. La flora apícola de la
Detriments and Benefits.—Thickets and tangles República Dominicana. http://naturalista.
of zarza are almost impenetrable to humans. Paths virtualave.net/estudios/apicola/dicotse.html. 8 p.
may be cut to allow passage, but they soon grow
over. Because of the thorns, cattle will not eat Paganucci de Queiroz, L. 2001. Leguminosas da
zarza (Vélez and van Overbeek 1950), but it is caatinga da Bahia com potencial forrageiro.
browsed by goats. The species is a host for the http://umbuzeiro.cnip.org.br/db/forrag/fam/
diaprepes root weevil (Diaprepes abbreviatus), f19.html. 6 p.
which is of grave concern for growers of citrus
and other agricultural crops (Simpson and others Simpson, S.E., H.N. Nigg, and J.L. Knapp. 2001.
2001). Zarza is a valuable honey plant (Marcano Host plants of diaprepes root weevil and their
1973), protects the soil, and serves as wildlife implications to the regulatory process. Florida
cover. Department of Agriculture and Consumer
Services. http://www.fcprac.ifas.ufl.edu/
References citrustopics/pest%20control/Diaprepes/…/simps
on.hostplant.ht. 16 p.
Acevedo-Rodríguez, P. 1985. Los bejucos de
Puerto Rico. Vol. 1. General Technical Report Vélez, I. and J. van Overbeek. 1950. Plantas
SO-58. U.S. Department of Agriculture, Forest indeseables en los cultivos tropicales. Editorial
Service, Southern Forest Experiment Station, Universitaria. Río Piedras, PR. 497 p.
New Orleans, LA. 331 p.
488
Mimosa pigra L. black mimosa
FABACEAE
John K. Francis
489
continuously in moist habitat, intermittently in environments helps them move on to the brush and
seasonally dry habitat. Bees pollinate the flowers secondary forest stages. It also protects the soil in
and pods mature about 1 month after pollination disturbed sites. Although the cover and diversity it
(Binggeli 1997). Black mimosa grows rapidly and furnishes is probably beneficial for wildlife in
can begin producing seed in as little as 3 months artificially maintained exotic grass swards, the
after emergence (Department of Natural Resources species can reduce wildlife populations in natural
and Mines 2002). One average plant can produce swards such as river floodplains in Australia
more than 9,000 seeds annually (Marko 2002). (Marko 2001). The plant has been used as a green
The pod segments disperse by floating on water manure, a cover crop, for fuel wood, and as
and clinging to clothing and possibly animal fur beanpoles. Although cattle and horses generally do
and feathers. A sample of black mimosa pods in not eat black mimosa because of the thorns
Puerto Rico contained an average of 15.0 (Binggeli 1997), with 22 percent foliar protein, it
seeds/pod. The seeds weighed an average of was found to be perfectly acceptable to rabbits
0.0166 g/seed. Sown on commercial potting mix (Lebas and others 1986). Black mimosa can be
between 6 and 20 days, 81 percent of the seed weedy in row crops and plantations, and a serious
germinated. Germination is epigeal (author’s problem in pastures and hay fields. The thorny
observation). Scarification is required for quick branches make it difficult for people to walk
and uniform germination. In an experiment with through fields and along trails where it grows. In
several treatments of temperature and storage, Africa, it is planted as a barrier around fields and
germination percentages of 75 to 94 percent were cattle enclosures to discourage human intruders
observed (Creager 1992). Seeds can remain viable and animal predators. The species is used in a
in dry soil for more than 2 years. Plants sprout number of herbal remedies and magic rites in
freely following fires (Binggeli 1997). Layering Africa (Burkill 1995).
occurs when stems become prostrate and covered
with litter. References
Marko, M. 2002. Controlling invasion of the Swarbrick, J.T. and B.L. Mercado. 1987. Weed
exotic shrub (Mimosa pigra) in tropical science and weed control in Southeast Asia.
Australia wetlands. University of Minnesota FAO Plant Production and Protection Paper 81.
Department of Horticultural Science. Food and Agriculture Organization of the United
Nations, Rome. 203 p.
491
Mimosa pudica L. sensitive plant
FABACEAE
John K. Francis
492
soaking. Germination is epigeal. Seeds are Limited, London. 358 p.
transported by means of the bristles on the edges
of their pods that cling to clothing or to the fur of Bui, L-D. 2001. Mimosa pudica. http://bio.
mammals. Most nursery and home propagation is maimi.edu/mimosa/mimosa.html. 7 p.
done using seeds, but summer cuttings may also
be used (Bui 2001). Chieng, H-T. and T.C. Huang. 1998. Aeropollen
of the Pingtung area, South Taiwan. Taiwania
Growth and Management.—In Puerto Rico, 43(2): 73-100.
sensitive plants live 1 to 2 years. Seedlings grow
slowly for 2 or 3 months and then accelerate, Coimbra, A.F. and A. Magnanini. 1953.
reaching 0.5 to 2 m of extension at the end of the Considerations sobre Mimosa pudica no
first year. Growth of plants that survive into the combate a arosao superficial. Anucio Brazilero
second year is much slower. Potted and field- Economico da Floresta 6. Instituto Nacional,
grown individuals are sensitive to overwatering Pinho, Brazil. 131-136.
(Bui 2001). This species has been successfully
tested and recommended for erosion control Feng, Z., P.G. Hartel, R.W. Roncadori, S.J.S.
plantings using potted material at a spacing of 60 Sung, and J.E. Box. 1998. Inhibition of fungal
x 60 cm (Coimbra and Magnanini 1953). colonization on the rhizoplane of the CS2-
production plant, Mimosa pudica L. In: Plant
Benefits and Detriments.—Sensitive plant has and Soil Sciences 82. Kluwer Academic
become a serious weed in fields of corn, soybeans, Publishers, Dordrecht, Netherlands. 115-126.
tomatoes, upland rice, cotton, bananas, sugarcane,
coffee, oil palms, papayas, coconuts, and rubber in Guzmán, D.J. 1975. Especies utiles de la flora
many tropical areas. It is particularly troublesome salvadoreña. Ministerio de Educación, Dirección
where hand pulling of weeds is practiced. The de Publicaciones. San Salvador, El Salvador.
species may be controlled by a number of 703 p.
commercial broad-leaf herbicides (Bui 2001). On
the other hand, it is tolerated or valued as a forage Holm, L.G., D.L. Plucknett, J.V. Paucho, and J.P.
plant in pastures (Holm and others 1977, Turbet Herberger. 1977. The world’s worst weeds.
and Thuraisingham 1948). In fact, sheep grazing East-West Center, University of Hawaii,
is reported to control sensitive plant in pastures Honolulu, HI. 609 p.
and plantations (Simonnet 1990). The root nodules
have been shown to fix nitrogen (Pokhriyal and Howard R.A. 1988. Flora of the Lesser Antilles,
others 1990). Thickets of sensitive plant may be a Leeward and Windward Islands.
fire hazard when dry (Pacific Island Ecosystems at Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain,
Risk 2001). The seeds and other plant parts of MA: Arnold Arboretum, Harvard University.
sensitive plant contain mimonsine, an amino acid 673 p.
that is known to cause hair loss and depressed
growth in mammals (Arora 1983). An unlikely Liogier H.A. 1988. Descriptive flora of Puerto
large dose is necessary to cause problems, Rico and adjacent islands, Spermatophyta. Vol.
however. The pollen is important to honeybees in 2. Editorial de la Universidad de Puerto Rico,
the Philippines (Payawal and others 1991). Río Piedras, PR. 481 p.
Extracts of the plant have been shown in scientific
trials to be a moderate diuretic, depress duodenal Martínez R., G., F.F. Rodríguez L., C.M.
contractions similar to atropine sulphone, promote Contreras, and M. Molina H. 1996. Estudio
regeneration of nerves, and reduce menorrhagia preliminary de las possibles acciones
(Modern-natural 2001). Anitdepressant activity antidepresivas de Mimosa pudica L. In:
has been demonstrated in humans (Martínez and Resumen de ponencias del Primer Congreso
others 1996). Root extracts are reported to be a Nacional de Plantas Medicinales de Mexico.
strong emetic (Guzmán 1975). p. 69. July 24-30, 1996. Tlaxcala, Tlaxcala,
Mexico.
References
Modern-natural. 2001. Mimosa pudica Linn.
Arora, S.K., ed. 1983. Chemistry and biochemistry http://modern-natural.com/mimosa_pudica.htm.
of legumes. Edward Arnold (Publishers) 4 p.
493
Pacific Island Ecosystems at Risk. 2001. Invasive Simonnet, P. 1990. Sheep flock management in a
plant species: Mimosa pudica Mart. ex Colla, tropical environment under coconut.
Fabaceae. http://www.hear.org/pier/mipud.htm. Oleagineux Paris 45(10): 451-456.
2 p.
Siregar, M.E., B. Haryanto, and S. Tjitrosemito.
Payawal, P.C., A.C. Tilde, and A.L. Manimtim. 1990. A review of weed management in
1991. Year round pollen sources of Italian honey Indonesian pastures. BIOTROP Special
bees (Apis mellifera L.) in the Philippines. III. Publication 38. A symposium on weed
Selected areas. Philippine Agriculturist 74(4): mangement. 7-9 June 1989. Bogor, Indonesia. p.
503-509. 229-235.
Pokhriyal, T.C., H.C.S. Bhandari, D.S. Negi, S.P. Turbet, C.R. and K. Thuraisingham. 1948. Feeding
Chaukiyal, and B.B. Gupta. 1990. Identification trials with the sensitive plant Mimosa pudica.
of some fast growing leguminous tree species Tropical Agriculturist, Ceylon 104(2): 81-86.
for nitrogen fixation studies. Indian Forester
116(6): 504-507.
494
Mimulus aurantiacus W. Curtis bush monkeyflower
PHRYMACEAE (formally in SCROPHULARIACEAE)
495
orange corolla with larger, deeply-notched lobes, a hybrid populations that were intermediate between
somewhat spreading habit to 1 m tall, and occurs parental forms. For example, in San Diego County,
from coastal San Luis Obispo to Monterey County putative hybrid populations were intermediate
and along the western base of the Sierra Nevada between M. puniceus (coastal form) and M.
from Placer into Butte and Plumas Counties. M. aurantiacus ssp. australis (inland form), consistent
flemingii Munz has red corollas with limbs about with analysis of Waayers (1996). In the region
half as long as other forms, is low growing from 1 between the San Gabriel and San Bernardino
to 6 dm tall, and is restricted to the Channel Mountains, putative hybrids were intermediate
Islands. M. longiflorus (Nutt.) A. L. Grant, the between M. longiflorus ssp. calycinus (lemon
southern bush monkeyflower (see figure), has an yellow corolla) and M. longiflorus (salmon
orange-yellow to salmon corolla, a long and corolla).
pubescent calyx (to 3.5 cm), is 3 to 12 dm tall, and
occurs from central San Diego County through Ecology.—Bush monkeyflowers occur in areas
Riverside to San Bernardino County. A shorter, with cool moist winters and warm to hot, dry
pale-yellow form [M. l. ssp. calycinus (Eastw.) summers. They grow on rocky hillsides and cliffs,
Munz] occurs primarily farther inland in Riverside usually on the border of chaparral or sage scrub, or
and western San Bernardino Counties. A red- in open foothill-woodland forest. The roots are
corolla form (M. l. var. rutilus Grant) occurs fibrous, less than 2 m deep, with feeder roots
mostly in interior Los Angeles County. Finally, M. concentrated within the first 8 cm of soil (Hellmers
puniceus (Nutt.) Steudel, the red bush and others 1955). There is evidence for
monkeyflower, has a red corolla, short calyx (to differentiation and local adaptation in
2.5 cm), and erect habit from 5 to 15 dm tall. It morphological and physiological characters
occurs on Catalina Island, coastal Orange County, relating to water use. For example, differences in
and inland through the Santa Ana Mountains to wood anatomy among forms and among
southwestern Riverside County south into Baja populations are correlated with habitat (Michener
California. 1983). Plants from mesic habitats have relatively
few, large vessel-elements, while plants from xeric
Hybridization.—All bush monkeyflowers have n habitats have many, small vessel-elements. Under
= 10 chromosomes and crosses among them humid, low-stress conditions, both mesic-adapted
produce viable progeny. Natural, putative hybrids and xeric-adapted populations had high water-use
between forms have been hypothesized based on efficiency and maintained a high rate of
morphology (Munz and Keck 1968, Wells 1980, photosynthesis relative to transpiration (Mooney
Grant 1993, Tulig 2000). Preliminary phylogenetic and Chu 1983). Plants from xeric populations had
work suggests widespread introgression among high water-use efficiency under low relative
forms (personal communication with Paul humidity, while the efficiency of coastal plants
Beardsley, Colorado College, CO). Grant (1993) diminished rapidly with decreasing relative
examined pollinators in relation to several floral humidity.
forms (equals species of Diplacus in his paper) and The primary herbivore of bush monkeyflower
postulated that the populations with different floral is a specialist butterfly Euphydryas chalcedona
forms would be partially reproductively isolated. Doubleday and Hewitson. The timing of growth of
He hypothesized that the red-colored D. puniceus both plants and the butterfly is linked to water
Nutt. and salmon D. longiflorus Nutt. were availability. Emergence of larvae from diapause is
primarily hummingbird-pollinated, and that D. synchronized with the initiation of leaf growth.
calycinus Eastw. (M. l. var. calycinus) with pale The leaves produce a resin that inhibits the growth
yellow flowers and long tube) were hawkmoth- of the larvae. During the growing season the
pollinated. He found that habitats interdigitated youngest leaves have the highest nitrogen (N)
and resulted in D. puniceus and D. calycinus content, the highest carbon gain, and the highest
populations overlapping in distribution, with D. resin content. Growth rate of the larvae increases
longiflorus habitat forming a connecting link with increasing N but decreases with increasing
between the different forms. He hypothesized that resin content (Lincoln and others 1982). Larvae
the intermediate nature of D. longiflorus suggested feed initially on young leaves with high nutritional
past hybridization of D. puniceus and D. calycinus. value, then switch to older leaves with lower resin
Tulig (2000) analyzed floral and vegetative content. During flowering, N is translocated from
variation throughout the geographic range of bush the leaves, and larvae stop feeding (Mooney and
monkeyflowers and found statistical evidence for others 1981).
496
Such studies demonstrate physiological longiflorus was “generally but not exclusively”
trade-offs in resource allocation to growth, pollinated by Black-chinned and Anna's
reproduction, or defense. Han and Lincoln (1994) hummingbirds. Six populations from the Santa
found significant heritability and maternal effects Monica Mountains varied in floral form, sucrose
for resin content within one population, and they content of nectar, and anther-stigma separation.
found negative genetic and phenotypic correlations Populations with high pollinator diversity but low
between resin production and growth rate. This visitation rates had shorter corolla tubes and lower
indicates potential trade-offs between traits during sucrose:hexose ratios in nectar than populations
selection. Leaf resin content varies among with less diverse pollinators. In a different
populations, but little is known about geographic population of M. longiflorus, Eckert (1970) found
differentiation in such trade-offs. that almost all pollination was done by solitary
Hare (2002) found that nearly 30 percent of bees and that Anna's hummingbirds were only
the dry weight of leaves is resin. In a common occasional visitors. Most pollination was effected
garden experiment, he examined the chemistry and by bees in the genus Osmia and carpenter bees in
relative quantities of important resin components Ceratina. Both visited multiple flowers per plant,
for six populations of bush monkeyflower from a and self-pollination appeared common. Eckert also
range of environments and that differed in attack found that selfed flowers produced fewer capsules
by E. chalcedona. The populations differed in per plant and had more aborted ovules than
chemical components of the resins but not in the outcrossed flowers.
pattern of insect attack. This study followed
Thompson’s (1993) classification and did not Horticulture.—The bush monkeyflowers and
mention if there were differences in floral their horticultural hybrids have striking flower
morphology among the populations. The collection colors, produce masses of flowers, and are popular
locations could have represented at least three ornamental plants (Schmidt 1980, Perry 1992,
species or subspecific taxa following Munz and Keator 1994). Plants can be grown from seeds or
Keck (1969) or Tulig (2000). cuttings, but plants grown from seed tend to live
longer (personal communication with Steve
Emergence and Growth.—Seedlings emerge in Morgan, University of California Riverside
the cool rainy season, and most growth is in winter Botanic Garden, CA). The shrubs grow best in full
and spring. Plants can reach reproductive maturity sun to light shade. Flowering and leaf production
in a single growing season, with flowering may be extended by summer watering in more arid
primarily from mid spring to early summer. The regions. However, Atkinson and others (1988)
capsules mature in summer, and the many, tiny found that watering may increase susceptibility to
seeds (about 1 mm long, half as wide and nearly atmospheric pollution in the dry season. In
flat) disperse by gravity and wind. Plants become summer, unwatered plants drop most leaves, are
dormant in the summer dry season and shed many not photosynthetically active, and take up little
leaves by fall. Leaf production begins after the SO2, while watered plants retain their leaves, are
first fall rain and ends near the beginning of July photosynthetically active, and take up SO2.
(Mooney and others 1981). A large and relatively
constant proportion of carbon and nitrogen are Growth and Management.—Bush
allocated to reproduction, and reproductive monkeyflowers have been planted along highways
structures supply some of their own carbon (Alpert corridors and are used in habitat restoration. There
and others 1985). Both growth and reproduction are about 145 million seeds/kg (Mirov and Kraebel
are primarily water limited. 1939). Seeds will germinate without pretreatment
(Mirov and Kraebel 1939, Schmidt 1980), but
Reproduction.—Bush monkeyflowers are seeds need to be exposed to light and leachate
pollinated by diverse flower visitors, and the from charred wood may improve germination
dominant pollinators vary with color form and slightly (Keeley 1987). For pot culture, the tiny
location. This may reflect local differentiation, or seeds should be sown on the surface of the soil and
the relative abundances of pollinators. Fetscher kept moist until seedlings emerge. Outdoors, they
and Kohn (1999) reported Anna’s hummingbird as should be planted with shallow broadcasting
the primary pollinator of a red flower form in San methods. The phylogenetic relationships among
Diego County, California, with black-chinned, populations and the fitness effects of translocations
Costa’s, and rufous hummingbirds as occasional and hybridization are not yet known, and there are
visitors. Bromer and others (1990) reported that M. no published studies on population genetic
497
patterns. However, there is a complex pattern of Fetscher, A.E. and J.R. Kohn. 1999. Stigma
geographic variation, genetic differentiation of behavior in Mimulus aurantiacus
floral form and resin chemistry, local adaptation to (Scrophulariaceae). American Journal of
moisture environments, and extensive ability to Botany 86: 1,130-1,135.
hybridize. Consequently, attention should be made
of flower color and habitat matching when Grant, V. 1993. Origin of floral isolation between
obtaining seeds for large planting projects. Use of ornithophilous and sphingophilous plant species.
regionally local seed sources for planting projects Proceedings of the National Academy of
can mitigate unknown fitness consequences of out Sciences of the United States of America 90:
of range plantings. A careful analysis of hybrid 7729-7733.
zones and the fitness of hybrids is needed to
understand if hybridization following Han, K. and D.E. Lincoln. 1994. The evolution of
translocations will cause decreases in fitness of carbon allocation to plant secondary
adjacent wild populations. metabolites: a genetic analysis of cost in
Diplacus aurantiacus. Evolution 48: 1,550-
Benefits.—Bush monkeyflowers are important 1,563.
plants for water-wise landscaping and revegetation
projects. They provide nectar to hummingbirds Hare, J.D. 2002. Geographic and genetic variation
and large bees and are an important larval host of in the leaf surface resin components of Mimulus
the checkerspot butterfly. aurantiacus from southern California.
Biochemical Systematics and Ecology 30: 281-
References 296.
Alpert, P., E.A. Newell, C. Chu, J. Glyphis, S.L. Hellmers, H., J.S. Horton, G. Juhren, and J.
Gulmon, D.Y. Hollinger, N.D. Johnson, H.A. O'Keefe. 1955. Root systems of some chaparral
Mooney, and G. Puttick. 1985. Allocation to plants in southern California. Ecology 36: 667-
reproduction in the chaparral shrub, Diplacus 678.
aurantiacus. Oecologia 66: 309-316.
Keator, G. 1994. Complete Garden Guide to the
Atkinson, C.J., W.E. Winner, and H.A. Mooney. Native Shrubs of California. Chronicle Books,
1988. Gas exchange and SO2 fumigation studies San Francisco, CA. 314 p.
with irrigated and unirrigated field grown
Diplacus aurantiacus and Heteromeles Keeley, J.E. 1987. Role of fire in seed germination
arbutifolia. Oecologia 75: 386-393. of woody taxa in California chaparral. Ecology
68: 434-443.
Beardsley, P.M. and R.G. Olmstead. 2002.
Redefining Phrymaceae: the placement of Lincoln, D.E., T.S. Newton, P.R. Ehrlich, and K.S.
Mimulus, tribe Mimuleae, and Phryma. Williams. 1982. Coevolution of the checkerspot
American Journal of Botany 89: 1093-1102. butterfly Euphydryas chalcedona and its larval
food plant Diplacus aurantiacus: larval response
Beardsley, P.M., S.E. Schoenig, J.B. Whittall, and to protein and leaf resin. Oecologia 52: 216-223.
R.G. Olmstead. 2003. The radiation of Mimulus
(Phrymaceae) in western North America. Michener, D.C. 1983. Systematic and ecological
American Journal of Botany. In Press. wood anatomy of Californian Scrophulariaceae.
I. Antirrhinum, Castilleja, Galvezia, and
Bromer, W., J. Barnette, J. Lee, D. Green, and V. Mimulus sect. Diplacus. Aliso 10: 471-487.
Ervin. 1990. Genetic variation within and among
populations of Mimulus longiflorus in the Santa Mirov, N.T. and C.J. Kraebel. 1939. Collecting
Monica Mountains, CA: consequences of and handling seeds of wild plants. Civilian
breeding system and pollination. Bulletin of the Conservation Corps, Forestry Publication 5.
Ecological Society of America 71: 103. United States Government Printing Office,
Washington, D.C.
Eckert, J.R. 1970. Pollination studies in Mimulus
longiflorus (Nutt.) Grant. Master’s thesis. Mooney, H.A., and C. Chu. 1983. Stomatal
California State University, Los Angeles. 88 p. responses to humidity of coastal and interior
498
populations of a Californian shrub. Oecologia Thompson, D.M. 1993. Mimulus. In: J.C.
57: 148-150. Hickman, ed. The Jepson Manual: Higher Plants
of California. University of California Press,
Mooney, H.A., K.S. Williams, D.E. Lincoln, and Ltd., Los Angeles, CA. p. 1,037-1,046.
P.R. Ehrlich. 1981. Temporal and spatial
variability in the interaction between the Tulig, M. 2000. Morphological variation in
checkerspot butterfly, Euphydryas chalcedona Mimulus section Diplacus (Scrophulariaceae).
and its principal food source, the Californian Masters Thesis, Department of Biological
shrub, Diplacus aurantiacus. Oecologia 50: 195- Sciences, California State Polytechnic
198. University, Pomona, CA. 82 p.
Munz, P.A. and D.D. Keck. 1968. A California Waayers. 1996. Hybridization, introgression, and
Flora with Supplement. University of California selection in Mimulus aurantiacus ssp. australis
Press, Berkeley, CA. 1,681 + 224 p. and M. puniceus. Master’s thesis. San Diego
State University, San Diego, CA. 77 p.
Perry, B. 1992. Landscape Plants for Western
Regions: an Illustrated Guide to Plants for Water Wells, H. 1980. A distance coefficient as a
Conservation. Land Design Publishing, hybridization index: an example using Mimulus
Claremont, CA. 318 p. longiflorus and M. flemingii (Scrophulariaceae)
from Santa Cruz Island, California. Taxon 29:
Schmidt, M.G. 1980. Growing California Native 53-65.
Plants. University of California Press, Los
Angeles.
499
Mitracarpus portoricensis (Urban) Urban Puerto Rico girdlepod
RUBIACEAE
John K. Francis
500
References Liogier, H.A. and L.F. Martorell. 1982. Flora of
Puerto Rico and adjascent islands: a systematic
Liogier, H.A. 1997. Descriptive flora of Puerto synopsis. Editorial de la Universidad de Puerto
Rico and adjacent islands. Vol. 5. Editorial de la Rico, Río Piedras, PR. 342 p.
Universidad de Puerto Rico, Río Piedras, PR.
436 p.
501
Morella caroliniensis (P. Mill.) Small bayberry
MYRICACEAE
Gerry Moore
502
his original description. The recent work of Wilbur of North America. Vol. 3. Oxford University
(1994, 2002) is followed here; therefore only one Press, New York. p. 430-434.
species is recognized under the name M.
carolininesis. Britton, N.L. and A. Brown. 1913. Illustrated flora
of the northern states, Canada, and the British
Ecology.—Bayberry can tolerate full sun to dense possessions. Vol. 2, 2nd Ed. Scribner, New York.
shade and can grow in dry to wet conditions. 735 p.
Habitats where it can be found include bogs,
swamps, dunes, fields, heathlands, oak forests, Chittenden, F.J. and P.M. Synge. 1956. Dictionary
pine forests, and the margins of streams and lakes. of Gardening. 2nd Ed. Clarendon Press, Oxford,
The roots of bayberry contain nitrogen fixing UK. 623 p.
nodules, and this allows it to be particularly
tolerant of nitrogen-poor, acidic soils (Morris and Collins, B.S. and J.A. Quinn. 1982. Displacement
others 1974). Results are mixed as to whether of Andropogon scoparius on the New Jersey
bayberry’s nitrogen-fixing properties may assist Piedmont by the successional shrub Myrica
the growth of other plant species growing nearby pensylvanica. American Journal of Botany 69:
(Tiffney and Barrera 1979, Dudley and others 680-689.
1996). Leachate from the leaf litter of bayberry
may have alleopathic properties and affect some Dudley, J.L., B. Michener, and K. Lajtha. 1996.
other plants (Collins and Quinn 1982). The contributions of nitrogen-fixing symbioses
to coastal heathland succession. American
Reproduction.—Bayberry is wind pollinated and Midland Naturalist 135: 334-342.
blooms in the spring. The pollen is allergenic
(Lewis 1977). The seeds of bayberry are dispersed Everett, T.H. 1960. New illustrated encyclopedia
by birds that eat the fruits (McClanahan and Wolfe of gardening. Vol. 3. Greystone Press, New
1993, Place and Stiles, 1992, Ridley 1930). York. 479 p.
Growth and Management.—Bayberry is often Fernald, M.L. 1950. Gray’s manual of botany.
cultivated on dry, sandy, sterile soils where few American Book Co., New York. 1,632 p.
other species can grow (Bailey 1922, Rehder
1940). It thrives best in acid peaty soils; it can not Gleason, H.A. and A. Cronquist. 1991. Manual of
tolerate liming. Propagation can be by seed or vascular plants of the northeastern United States
layering (Chittenden and Synge 1956). and adjacent Canada. 2nd Ed. New York
Propagation can also be done with cuttings taken Botanical Garden, Bronx. 910 p.
in late summer or fall. The cuttings should be
placed in a bed of peat moss and sand (Everett Lewis, W.H. 1977. Medical botany. John Wiley &
1960). Sons, New York. 515 p.
Benefits.—Settlers used the bark of bayberry in McClanahan, T.R. and R.W. Wolfe. 1993.
dentifrice mixtures (Lewis 1977) and the fruits in Accelerating forest succession in a fragmented
candle making (Bornstein 1997). Bayberry candles landscape: the role of birds and perches.
remain popular today; it can take up to 8 kg of Conservation Biology 7: 279-288.
bayberry fruit to yield 1 kg of wax. The fruits are
also often used in decorative arrangements. Native Miller, P. 1768. The gardeners dictionary. 8th Ed.
Americans used the plant for various medicinal Printed for the author, London. 1,329 p.
purposes (Moerman 1986).
Moerman, D.E. 1986. Medicinal plants of Native
References America. 2 Vols. Technical Report 19.
University of Michigan Museum of
Bailey, L.H. 1922. Standard cyclopedia of Anthropology, Ann Arbor, MI. 910 p.
horticulture. Vol. 4. MacMillan Co., London.
660 p. Morris, M., D.E. Eveleigh, S.C. Riggs, and W.N.
Tiffney, Jr. 1974. Nitrogen fixing in the
Bornstein, A.J. 1997. Myricaceae. In: Flora of bayberry (Myrica pensylvanica) and its role in
North America Editorial Committee, eds. Flora
503
coastal succession. American Journal of Botany Tiffney, W.N., Jr. and J.F. Barrera. 1979.
61: 867-870. Comparative growth of pitch and Japanese black
pine in clumps of the N2 –fixing shrub, bayberry.
Place, A.R. and E.W. Stiles. 1992. Living off the Botanical Gazette 140 (Suppl.): S108-S109.
wax of the land: bayberries and yellow-rumped
warblers. Auk 190: 334-345. Wilbur, R.L. 1994. The Myricaceae of the United
States and Canada: genera, subgenera, and
Rehder, R. 1940. Manual of cultivated trees and series. Sida 16: 93-107.
shrubs, Macmillan Publishing Co, New York.
996 p. Wilbur, R.L. 2002. The identity and history of
Myrica caroliniensis (Myricaceae). Rhodora
Ridley, H.N. 1930. The dispersal of plants 104: 31-41.
throughout the world. L. Reeve & Co., Ashford,
Kent, UK. 744 p.
504
Morinda citrifolia L. noni
RUBIACEAE
Synonyms: none
John K. Francis
505
information is lacking on growth and Cambie, R.C. and J. Ash. 1994. Fijian medicinal
management. plants. CSIRO, Canberra, Australia. 365 p.
Benefits.—Although the fruits of noni are Francis, J.K. and A. Rodríguez. 1993. Seeds of
somewhat tasteless and have an unpleasant smell, Puerto Rican trees and shrubs: second
they are eaten as famine food in Indonesia, installment. Research Note SO-374. U.S.
Australia, and the Pacific Islands. The young Department of Agriculture, Forest Service,
leaves are also eaten as a vegetable and contain 4 Southern Forest Experiment Station, New
to 6 percent protein. Noni is planted as a Orleans, LA. 5 p.
windbreak, for support of pepper vines, and shade
for coffee bushes (Tan 2001). The species is Hirazumi, A. and E. Furusawa. 1999. An
useful for coastal erosion control (Association of immunomodulatory polysaccharide-rich
Societies for Growing Australian Plants 2000), substance from the fruit juice of Morinda
and it is planted as an ornamental in some areas. citrifolia (noni) with antitumour activity.
The bark has been the source of a red dye (Little Phytotherapy Research 13(5): 380-387.
and Wadsworth 1964). Roots are used to produce
an orange dye (Nelson 1996). Harvested from both Howard, R.A. 1989. Flora of the Lesser Antilles,
plantations and from the wild, noni is one of the Leeward and Windward Islands.
most important botanical remedies and food Dicotyledoneae. Part 3. Vol. 6. Arnold
supplements traded on the international market. It Arboretum, Harvard University, Jamaica Plain,
is usually the fruit juice that is sold, fresh or dried. MA. 658 p.
Leaves, bark, and roots are also used for a great
many maladies. Pain, arthritis, diabetes, high Legal, L. and M. Plawecki. 1995. Comparative
blood pressure, skin and stomach ulcers, sensitivity of various insects to toxic compounds
depression, senility, diarrhea, arteriosclerosis, from Morinda citrifolia L. Entomological
cancer, AIDS, skin parasites, and bad breath are Problems 26(2): 155-159.
all treated (Association of Societies for Growing
Australian Plants 2000, Cambie and Ash 1994, Liogier, H.A. 1997. Descriptive flora of Puerto
Tan 2001). A number of physiologically active Rico and adjacent islands. Vol. 5. Editorial de la
chemicals, anthraquinones, alkaloids, Universidad de Puerto Rico, San Juan, PR.
asperuloside, caproic, caprylic, and ursolic acids, 436 p.
β-sitosterol, and asperuloside may account for
some of the effects (Cambie and Ash 1994). Little, E.L., Jr. and F.H. Wadsworth. 1964.
Antitumor activity expressed in enhanced survival Common trees of Puerto Rico and the Virgin
of tumor-bearing mice has been demonstrated Islands. Agriculture Handbook 249. U.S.
after treating with juice extracts (Hirazumi and Department of Agriculture, Forest Service,
Furusawa 1999). Aqueous extracts of roots were Washington, DC. 548 p.
shown to have an analgesic effect on mice without
any sign of toxicity, and a sedative effect at high Nelson, G. 1996. The shrubs and woody vines of
doses (Younos and others 1990). Octanoic acid, Florida. Pineapple Press, Sarasota, FL. 391 p.
which is present in the ripe fruits, effectively
poisons a fruit fly (Drosophila melanogaster), Smith, R.S. 2002. History and tradition of
honey bee (Apis mellifera), an ant (Lasius sp.), Morinda citrifolia. http://rsscomp.freeyellow.
and cockroaches (Periplaneta americana and com/morindacitrifoliastory.htm. 2 p.
Blattella germanica) (Legal and Plawecki 1995).
Noni is the principal larval host of the hawk moth, Stampsfiji.com. 2002. The Sphingid (hawk) moths
Macroglossum hirundo vitiensis in Fiji of Fiji stamp issue. http://www.stampsfiji.
(Stampsfiji.com 2002). com/stamps/moths/ 4 p.
506
Tan, R. 2001. Mangrove and wetland wildlife at Younos, C., A. Rolland, J. Fleurentin, M.C.
Sungei Buloh Nature Park: Great morinda. Lanhers, R. Misslin, and F. Mortier. 1990.
http://www.naturia.per.sg/buloh/plants/morinda. Analgesic and behavioral effects of Morinda
htm. 3 p. citrifolia. Planta Medica 56(5): 430-434.
507
Murraya exotica L. orange jasmine
RUTACEAE
John K. Francis
508
on commercial potting mix, 65 percent germinated http://www.desert-tropicals.com/Plants/
between 25 and 60 days of sowing. Germination is Rutaceae/Murraya_paniculata.html. 2 p.
hypogenous. Seedlings quickly develop deep root
systems and grow at a moderate rate. Plants Gonzalez, G. 2002. Bonsai gallery photos of
coppice vigorously after disturbance. Nursery Murraya paniculata bonsi. Tropical
production is by seed, cuttings (Little and others Bonsai.com. http://www.Tropicalbonzai.com/
1974), and air-layers (Woman’s Club of Havana MurrayaPaniculata.htm. 1 p.
1952).
Harris, E.J. and C.Y.L. Lee. 1986. Seasonal and
Growth and Management.—Orange jasmine annual occurrence of Mediterranean fruit flies
plants can live at least 15 years (author’s (Diptera: Tephritidae) in Makaha and Waianae
observation). Once established as an ornamental, valleys, Oahu, Hawaii. Environmental
they need little care. The species has not yet been Entomology 15(3): 507-512.
reported to be a weed in any area. Orange jasmine
can be killed (with moderate success) with Howard R.A. 1988. Flora of the Lesser Antilles,
herbicides recommended for broad-leaf weeds, by Leeward and Windward Islands.
girdling (Negreros-Castillo and Hall 1994), or by Dicotyledoneae, Part 1. Vol. 4. Arnold
grubbing out the plants. The species is the Arboretum, Harvard University. Jamaica Plain,
preferred host of the citrus psyllid, Diaphorina MA. 673 p.
citri, the vector for “citrus greening” disease
(Pacific Island Ecosystems at Risk 2002), and is a Kinoshita, T. and K. Firman. 1996. Highly
host of Mediterranean fruit flies (Diptera: oxygenated flavonoids from Murraya
Tephritidae) (Harris and Lee 1986). paniculata. Phytochemistry 42(4): 1207-1210.
Benefits.—Orange jasmine is a popular hedge Liogier H.A. 1988. Descriptive flora of Puerto
plant in the tropics. It can be used as a background Rico and adjacent islands, Spermatophyta. Vol.
plant or an accent plant. It can be pruned into 2. Editorial de la Universidad de Puerto Rico,
animal and other shapes (topiary), pruned into tree Río Piedras, PR. 481 p.
form, grown as a potted plant (Whistler 2000), or
cultivated as a bonsai (Gonzalez 2002). The Little, E.L., Jr., R.O. Woodbury, and F.H.
sapwood is light yellow, and the heartwood is Wadsworth. 1974. Trees of Puerto Rico and the
light brown and heavy, hard, fine-textured and Virgin Islands. Vol. 2. Agriculture Handbook
good for small turned articles (Little and others 449. U.S. Department of Agriculture,
1974). Branches or stems 2 to 5 cm in diameter in Washington, DC. 1,024 p.
the Indian Himalayan region were evaluated for
fuelwood suitability. Results were as follows: Neal, M.C. 1965. In gardens of Hawaii. Special
oven-dry density 0.72, dry-weight caloric value Publication 50. Bernice P. Bishop Museum
20.2 kj/g, ash 0.6 percent, moisture 54.7 percent, Press, Honolulu, HI. 924 p.
and nitrogen 0.42 percent (Negi and Todaria
1993). In the wild, orange jasmine contributes to Negi, A.K. and N.P. Todaria. 1993. Fuelwood
the biodiversity, protects the soil, and furnishes evaluation of some Himalayan trees and shrubs.
food and cover for wildlife. The leaves and other Energy 18(8): 799-801.
tissues have both stimulant and astringent
properties and are used to treat diarrhea, Negreros-Castillo, P. and R.B. Hall. 1994. Four
dysentery, cuts, joint pain, body aches (Parrotta methods for partial overstory removal in tropical
2001), venereal disease (Kinoshita and Firman forests of Mexico. Journal of Environmental
1996), and as an abortive (Xiao and Wang 1991). Management 41(3): 237-243.
In addition to essential oils, tissues of orange
jasmine contain the indole alkaloid yuehchukene Pacific Island Ecosystems at Risk. 2002. Invasive
(Xiao and Wang 1991) and at least eight highly plant species: Murraya paniculata (L.) Jack,
oxygenated flavones (Kinoshita and Firman 1996). Rutaceae. http://www.hear.org/pier3/mupan.
htm. 2 p.
References
Parrotta, J.A. 2001. Healing plants of Peninsular
Desert-Tropicals. 2002. Orange jasamine. India. CABI Publishing, Wallingford, UK and
509
New York. 917 p. guide. Timber Press, Portland, OR. 542 p.
Pathak, R.K., Madhulika-Mahajan, and S.N. Woman’s Club of Havana. 1952. Flowering plants
Sachan. 1992. Powdery mildew and cultural from Cuban gardens. Criterion Books, New
practices. Indian Journal of Forestry 15(1): 73. York. 365 p.
Sheehan, M.R. 1975. Florida landscape plants, Xiao, P.G. and N.G. Wang. 1991. Can
native and exotic. The University Presses of ethnopharmacology contribute to the
Florida, Gainesville, FL. 420 p. development of anti-fertility drugs? Journal of
Ethnopharmacology 32(1-3): 167-177.
Whistler, W.A. 2000. Tropical ornamentals, a
510
Myrica gale L. sweet gale
MYRICACEAE
Gerry Moore
appearing on different plants. Individual plants
have been known to change sex from year to year
(Burges 1993). The catkins are stalkless and borne
on the upper portions of the preceding year’s
branchlets. These branchlets die after flowering.
The staminate catkins range from 6 to 10 mm long,
are crowded on the branchlet, with the individual
flowers overtopped by the broad, shining,
subtending, brown bracts. The pistillate catkins
range from 7 to 10 mm long, each flower
subtended by two wing-like bracts that remain
fused to the fruit. The fruit of the sweet gale is a
three-pointed, compressed, ovoid nutlet that at
maturity is dotted with shining red to yellow resin
glands.
511
can be readily distinguished from the waxy fruited Baker, A. and R. Parsons. 1997. Rapid
species (bayberry, wax myrtle) on the bases of assimilation of recently fixed N2 in root nodules
flowers and fruits. If sweet gale is to be recognized of Myrica gale. Physiologia Plantarum 99: 640-
as generically distinct from bayberry and wax 647.
myrtle, as is done here, the generic name Myrica is
to be retained for sweet gale, with the bayberry Britton, N.L. and A. Brown. 1913. Illustrated flora
and wax myrtle being placed in the genus Morella of the northern states, Canada, and the British
(Wilbur 1994, 2002). Various varieties and possessions. Vol. 2, 2nd Ed. Scribner, New York.
subspecies [e.g., M. gale var. subglabra Fernald, 735 p.
M. gale var. tomentosa C. DC., M. gale ssp.
tomentosa (C. DC) E. Murray] have been Bornstein, A.J. 1997. Myricaceae. In: Flora of
recognized within sweet gale based on the degree North America Editorial Committee, eds. Flora
of pubescence present on the leaves. of North America Vol. 3. Oxford University
Press, New York. p. 430-434.
Ecology.—Sweet gale can tolerate full sun to
dense shade and is usually found in moist to wet Burges, N.A. 1993. Myrica. In: T.G. Tutin, N.A.
areas. Habitats where it can be found include bogs, Burges, A.O. Chater, J.R. Edmondson, V.H.
fens, swamps, and the margins of streams and Heywood, D.M. Moore, D.H. Valentine, S.M.
lakes. The roots of sweet gale contain nitrogen- Walters, and D.A. Webb, eds. Flora Europaea.
fixing nodules, which allow it to be particularly 2nd Ed.. Vol. 1. Cambridge University Press,
tolerant of nitrogen-poor, acidic soils (Baker and Cambridge, UK. p. 66-67.
Parsons 1997). Nitrogen fixation is accomplished
by the actinomycetous fungal genus, Frankia. Chittenden, F.J. and P.M. Synge. 1956. Dictionary
of Gardening 2nd Ed. Vol. 1. Clarendon Press,
Reproduction.—Sweet gale is wind pollinated Oxford, UK. 623 p.
and blooms in the spring. The pollen is allergenic
(Lewis 1977). The fruits of sweet gale mature in Everett, T.H. 1960. New illustrated encyclopedia
the fall and are dispersed by water (Ridley 1930, of gardening. Vol. 3. Greystone Press, New
Wilbur 1994). York. 479 p.
Growth and Management.—Sweet gale is often Fernald, M.L. 1950. Gray’s manual of botany.
cultivated on moist acidic soils where few other American Book Co., New York. 1,632 p.
species can be grown (Bailey 1922, Rehder 1940).
It thrives best in peaty soils and cannot tolerate Gleason, H.A. and A. Cronquist. 1991. Manual of
liming. Propagation can be by seed, layering, or vascular plants of the northeastern United States
root suckers (Chittenden and Synge 1956). and adjacent Canada 2nd Ed. New York
Propagation can also be effected through cuttings Botanical Garden, Bronx, NY. 910 p.
taken in late summer or fall. The cuttings should
be placed in a bed of peat moss and sand (Everett Lewis, W.H. 1977. Medical botany. John Wiley &
1960). Sons, New York. 515 p.
Benefits.—The branches and bark have been used Moerman, D.E. 1986. Medicinal plants of Native
in making a gale beer and in tanning; a decoction America. 2 Vols. Technical Report 19.
has been used for an insecticide and to kill vermin University of Michigan Museum of
(Rook 1998). The branches of sweet gale were Anthropology, Ann Arbor, MI. 910 p.
used by the Native Americans, Bella Coola, to
prepare decoctions to be used as a diuretic or as a Rehder, A. 1940. Manual of cultivated trees and
treatment for gonorrhea (Moerman 1986). shrubs, Macmillan Publishing Co, New York.
996 p.
References
Ridley, H.N. 1930. The dispersal of plants
Bailey, L.H. 1922. Standard cyclopedia of throughout the world. L. Reeve & Co., Ashford,
horticulture Vol. 4. MacMillan Co., London. Kent, UK. 744 p.
660 p.
512
Rook, J.S. 1998. A boundary waters compendium. Wilbur, R.L. 1994. The Myricaceae of the United
http://www.rook.org/earl/bwca/nature/shrubs/ States and Canada: genera, subgenera, and
myricagale.html [notpaged]. series. Sida 16: 93-107.
USDA, NRCS. 2003. The PLANTS Database, Wilbur, R.L. 2002. The identity and history of
Version 3.5 Baton Rouge, LA. Myrica caroliniensis (Myricaceae). Rhodora
http:plants.usda.gov. [not paged]. 104: 31-41.
513
Neea buxifolia (Hook. f.) Heimerl nia
NYCTAGINACEAE
John K. Francis
514
(room temperature) bonsai species available and 1. Editorial de la Universidad de Puerto Rico,
offers promise as an ornamental, especially for Río Piedras, PR. 352 p.
hedges and other shaped ornamentals.
Little, E.L., Jr., R.O. Woodbury, and F.H.
References Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
Clubbe, C. 2000. British Virgin Islands: 449. U.S. Department of Agriculture,
conservation and training. Kew Scientist 17:5. Washington, DC. 1,024 p.
Liogier H.A. 1985. Descriptive flora of Puerto Zane, T.L. 2002. Neea buxifolia—Neea buxifolia.
Rico and adjacent islands, Spermatophyta. Vol. http://www.bonsai-bci.com/species/neea.html.
2 p.
515
Odontonema cuspidatum (Nees) Kuntze cardinal’s guard
ACANTHACEAE
John K. Francis
moist. It is moderately tolerant of shade and will
bloom in full sun, broken sun, and moderate shade
(author’s observation, Blomber 2000). In Central
America, it grows from near sea level to 1,400 m
in elevation (Stevens and others 2001). Cardinal’s
guard is not salt tolerant (Watkins 1975) and dies
to the ground if subjected to frost. In areas subject
to frost, it is grown as an annual or herbaceous
perennial (Blomber 2000, Watkins 1975). The
plants are mostly free of pests and disease but may
occasionally be attacked by mealy bugs
(Pseudococcidae) (Woman’s Club of Havana
1952).
Ecology.—Cardinal’s guard prospers on fertile Benefits.—In natural stands, cardinal’s guard adds
and moderately fertile soils that are continually to biodiversity and protects against soil erosion.
516
The species is widely cultivated as an ornamental Universidad de Puerto Rico, San Juan, PR.
in tropical and subtropical areas for its striking red 436 p.
flowers. It is used as an accent and background
plant and sometimes employed to form hedges Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
(Watkins 1975, Whistler 2000, Woman’s Club of Montiel, eds. 2001. Flora de Nicaragua.
Havana 1952). It is also popular in gardens Monographs in systematic botany Vol. 85, No. 1.
because it attracts butterflies and hummingbirds Missouri Botanical Garden Press. 943 p.
that feed on the nectar (Blomber 2000, Watkins
1975). Watkins, J.V. 1975. Florida landscape plants,
native and exotic. The University Presses of
References Florida, Gainesville, FL. 420 p.
Blomber, M. 2000. In the garden: tropical plants Whistler, W.A. 2000. Tropical ornamentals, a
can add diversity to gardens. guide. Timber Press, Inc., Portland, OR. 542 p.
http://gainesvillesun.com/news/marina/02-26-
00marina.shtml. 4 p. Woman’s Club of Havana. 1952. Flowering plants
from Cuban gardens. Criterion Books, New
Liogier, H.A. 1997. Descriptive flora of Puerto York. 365 p.
Rico and adjacent islands. Vol. 5. Editorial de la
517
Olyra latifolia L. carricillo
POACEAE
John K. Francis
518
failed to stimulate germination (author’s Croat, T.B. 1978. Flora of Barro Colorado Island.
observation). Birds disperse the seeds (Burkill Stanford University Press, Stanford, CA. 943 p.
1994, Judziewicz and others 1999). New plants are
uncommon. Attempts to cultivate the species in Howard, R.A. 1979. Flora of the Lesser Antilles,
temperate greenhouses have failed (Judziewicz and Leeward and Windward Islands. Vol. 3. Arnold
others 1999). Arboretum, Harvard University, Jamaica Plain,
MA. 586 p.
Growth and Management.—Carricillo is a
relatively fast growing plant. Individual culms Judziewicz, E.J., J.G. Clark, X. Londoño, and M.J.
grow 2 m or more during their first year and 1 m Stern. 1999. American bamboos. Smithsonian
or so thereafter and live 2 to 4 years. By continual Institution Press, Washington, DC.
sprouting, plants may live for many years. http://email.uwsp.edu/publicanon/Course%20Inf
Artificial vegetative propagation has not been ormation/200120%-%20Spring%202002/
explored. Until vegetative methods are developed Biology/PF-BIOL345_xF8FF_545.1_200120/
or the extended dormancy problem of seeds is American%20Bamboos%20book.doc?Cmd=
solved, planting is not advised. Eradication can be open. 175 p.
done by grubbing out individual plants or probably
by cutting and spraying the sprouts with Missouri Botanical Garden. 2002. Manual de la
glyphosate or other grass herbicides. flora de Costa Rica. Missouri Botanical Garden,
St. Louis, MO. http://www.mobot.org/manual.
Benefits.—Carricillo contributes to the diversity plantas/050531/S050880.html. 7 p.
of the forest, helps protect the soil, and furnishes
food and cover for wildlife. Cattle eat the leaves North Carolina Botanical Garden. 2002. The biota
and fine twigs. Birds eat the fertile florets and of North America program: Olyra. North
fruits, although there is speculation that they may Carolina Botanical Garden, Chapel Hill, NC.
not be able to digest the hardened seeds (Burkill http://www.funet.fi/pub/sci/bio/life/plants/magno
1994). The species is the food plant for larva of the liophyta/magnoliophytina/liliopsida/poaceae/
moth Eryphanis reevesii (Doubleday) (North olyra/ 2p.
Carolina Botanical Garden 2002). The hollow
culms are used for flutes (Judziewicz and others Shaka, J.M., W. Kabushemera, and A. Msangi.
1999) and drinking straws. Leaves, culms, roots, 1997. Soils and vegetation of Semdoe proposed
and seeds are used in a number of applications in forest reserve Bombwera Division, Muheza
herbal medicine (Burkill 1994). Blades made from District, Tanga. Tech. Paper 36. Ministry of
the culms are used by the Cuiba tribe in Colombia Agriculture, National Soil Service, Agricultural
to cut the umbilical cords of newborns (Judziewicz Research Institute, Mlingano, Tanganyika. 14 p.
and others 1999).
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
References Montiel, eds. 2001. Flora de Nicaragua.
Monographs in Systematic Botany Vol. 85, No.
Burkill, H.M. 1994. The useful plants of West 3. Missouri Botanic Garden Press, St. Louis,
Tropical Africa. Vol. 2. Royal Botanic Gardens, MO. p. 1,911-2,666.
Kew, UK. 636 p.
519
Opuntia leptocaulis DC. desert Christmas cactus
CACTACEAE
Juanita A. R. Ladyman
that is usually less than 1.5 m tall but can grow up
to 2.8 m in shady, protected environments such as
under trees (Earle 1990). Although relatively
compact, it is extensively branched from a main
trunk that is covered by scaly bark in old age. It
has pencil-like woody joints that are 12 to 40 cm
long and 5 mm in diameter and can be easily
detached from the plant. The tubercules, usually
less than 1 cm long, may be indistinct and the
joints smooth. The areoles, which are the small,
clearly defined raised areas on the surface, are oval
or almost diamond-shaped with short white wool.
The areoles may be spineless or have from one to
three spines that are from 19 to 51 mm long. There
are few, usually short, yellowish-brown glochids,
or minute bristles, in one to three small clusters in
the upper part of the areole. The leaves are green,
less than 1.3 cm long, and are early deciduous. The
greenish-yellow flowers are approximately 1.3 to
1.9 cm in diameter and up to 2.5 cm tall. The fruits
are up to 2.5 cm long, and smooth with brown
glochids in the areoles. There are usually less than
12 seeds per fruit (Earle 1990, Weniger 1991,
Vines 1986). Desert Christmas cactus is a diploid
species where n = 11 (McGregor and others 1986).
520
(DC) Colville, with which it has a cyclical the fourth year (Bunting and others 1980). Desert
relationship in the Chihuahuan desert (Yeaton Christmas cactus recovery appeared particularly
1978). Creosotebush colonizes sites and modifies sensitive to the amount of precipitation.
microhabitat and edaphic conditions for desert Precipitation was required before plants would re-
Christmas cactus establishment and is eventually sprout. Plants tolerate minimum temperatures of
replaced by it. However, a combination of 7.3 oC in cultivation (Innes and Glass 1991). In
burrowing rodents and soil erosion increases the cultivation, desert Christmas cactus is susceptible
mortality rate of desert Christmas cactus while to the cochineal bug and periodic oil spray is
having little effect on creosotebush. The openings recommended (Earle 1990).
left by the dead cactus are then re-colonized by
creosotebush. Benefits.—Weniger (1991) describes it as a major
pest and “probably one of the most hated cacti in
Reproduction.—Desert Christmas cactus our area [Texas].” This is an unfortunate sentiment
reproduces both sexually by seed and vegetatively. because, although the spines can impale people
Flowers generally appear in May to July and livestock that venture too close, it provides
depending upon regional and environmental significant food, protected nesting sites, and cover
conditions. In response to wet weather, they can for a variety of wildlife (Earle 1990, Dayton
flower as early as April and again in August. The 1931). Most birds, including the bobwhite quail
flowers open wide in late evening (Taylor and and wild turkey, as well as small mammals, favor
others 1997). The fruit ripens in December and the fruit (Powell 1998, Taylor and others 1997).
persists on the plant until approximately February. Some animals, such as white-tailed dear, consume
Seeds may sprout while in the fruit and have a the joints as well as the fruit (Everitt and Drawe
shoot of up to 7 cm long while still on the plant 1993). There is apparently no seasonal variation
(Weniger 1991). Detached joints root and grow in the crude protein content of the stems, which is
into plants. The terminal stem segments detach similar to the fruits; both are 8 percent (Taylor and
particularly easily and readily take root near the others 1997). The larger shrubs provide cover and
parent plant so that natural populations that appear nesting sites, especially for the cactus wren. In
to be dense monocultures of clonal individuals addition, there is now a commercial value to
develop (Rebman and Pinkava 2001). Cuttings maintaining these shrubs. Land management for
taken from woody stems do not root easily (Earle wildlife has become increasingly important to
1990). Hybrids between cholla species are livestock producers in the Western U.S.A. because
common in nature (Rondeau and other 1996). An of fluctuations in livestock prices. In some parts of
example herbarium specimen of such a hybrid, O. Texas, income generated from consumptive (e.g.,
leptocaulis x O. spinosior (cane cholla), can be hunting) and non-consumptive (e.g., bird
seen on the Internet (Pinkava 2001). watching) wildlife activities already exceed
revenue obtained through livestock operations
Growth and Management.—Desert Christmas (Hernandez and others 1999). Under these
cactus is frequently abundant on overgrazed range circumstances upland gamebirds, such as northern
(Powell 1998). It tends to form thickets that can bobwhites and Rio Grande wild turkeys, have
become almost impenetrable to humans and become particularly important. Recommendations
livestock. Individual plants can be long-lived and have been made that low, thorny brush composed
may be more than 53 years old (Goldberg and of species such as lotebush [Ziziphus obtusifolia
Turner 1986). In a study over a 72-year period, 40 (Hook. ex Torr. & Gray) Gray] and desert
percent of plants were determined to survive at Christmas cactus should be protected in order to
least 7 years (Goldberg and Turner 1986). enhance nesting cover (Hernandez and others
Survival after fire is dependent upon the intensity 1999) and provide a food source. Desert Christmas
and frequency of the fire (Thomas 1991, Cave and cactus also provides refugia to native plant species.
Patten 1994). Plants can re-sprout after the fire A clear example of this on a landscape scale is in
kills the above ground parts (Bunting and others Arizona where clumps of tobosa grass are almost
1980). However, desert Christmas cactus appears completely restricted to refugia that are most
more susceptible to direct damage from fire than commonly under the desert Christmas cactus
some other Opuntia species (Bunting and others canopy where it is inaccessible to livestock
1980). After a fire in a mixed-grass prairie in (McAuliffe 1995). Native Americans gathered the
Texas, mortality was 65 percent in the first year fruits and ate them raw or cooked them into a jam
and reached a cumulative value of 80 percent by (Earle 1990). The Apache tribes mixed the
521
crushed fruits with a beverage to produce narcotic semiarid environments. Proceedings from
effects (Moerman 1998). Northern Texas and South Texas Wildlife
Conservation and Management Workshop,
References Saltillo, Mexico. August 19-20. Internet site:
http://cnrit.tamu.edu.
Bowers, J.E. 1993. Shrubs and trees of the
Southwest Deserts. Southwest Parks and Innes, C. and C. Glass. 1991. Cacti. Portland
Monuments Assoc. Tucson, AZ. 140 p. House, New York. 320 p.
Bowers, J.E. and S.P. McLaughlin. 1987. Flora Kearney, T.H., R.H. Peebles, and collaborators.
and vegetation of the Rincon Mountains, Pima 1960. Arizona flora. 2nd Ed. University of
County, Arizona. Desert Plants 8(2): 51-94 California Press, Berkeley, CA. 1,085 p.
Bunting, S.C., H.A. Wright, and L.F. Neuen- McAuliffe, J.R. 1995. Landscape evolution, soil
schwander. 1980. Long-term effects of fire on formation, and Arizon’s desert grassland. In:
cactus in southern mixed prairie of Texas. M.P. McClaran and T.R. Van Devender, eds.
Journal of Range Management 33(2): 85-88 The desert grassland. The University of Arizona
Press, Tucson, AZ. p. 100-129.
Burgess, T.L. 1995. The dilemma of coexisting
growth forms. In: M.P. McClaran and T.R. Van McGregor, R.L., T.M. Barkley, RE Brooks, EK
Devender, eds. The desert grassland. University Schofield. 1986. Flora of the Great Plains.
of Arizona Press, Tucson, AZ. p. 31-67 University Press of Kansas, Lawrence, KN.
1,402p.
Cave, G.H. and D.T. Patten. 1994. Short-term
vegetation responses to fire in the upper Sonoran Moerman, D.E. 1998. Native American ethno-
desert. Journal of Range Management 37(6): botany. Timber Press, Portland, OR. 927 p.
491-496
Nobel, P.S. 1988. Environmental biology of
Crook, R. and R. Mottram, 1999. Opuntia Index – agaves and cacti. Cambridge University Press,
Part 5: Nomenclatural note and I-L. Bradleya New York. 288 p.
17: 109-131
Pinkava, D. 2001. Biologist profile, specimens at
Dayton, W.A. 1931. Important western browse Arizona State University herbarium.
plants. Misc. Publication 101. U.S. Department http://lsvl.la.asu.edu/askabiologist/profiles/
of Agriculture, Washington, DC. 214 p. pinkava/pink.html.
Earle, W.H. 1990. Cacti of the Southwest. Desert Powell, A.M. 1998. Trees & shrubs of Trans-
Botanical Garden, Phoenix, AZ. 210 p. Pecos and adjacent areas. University of Texas
University Press, Austin, TX . 498 p.
Epple, A.O. 1995. A field guide to the plants of
Arizona. Falcon Press Publishing Co., Helena, Rebman, J.P., and D. J. Pinkava. 2001. Opuntia
MT. 347 p. cacti of North America–An overview. Florida
Entomologist 84 (4): 474-483
Everitt, J.H. and D.L. Drawe. 1993. Trees, Shrubs
and Cacti of South Texas. Texas Tech Rondeau, R., T.R. Van Devender, C.D. Bertelsen,
University Press, Lubbock, TX. 213 p. P. Jenkins, R.K. Wilson, and M.A. Dimmitt.
1996. Annotated flora and vegetation of the
Goldberg, D.E. and R.M. Turner. 1986. Tucson Mountains, Pima County, Arizona.
Vegetation change and plant demography in Desert Plants 12(2): 3-46
permanent plots in the Sonoran Desert. Ecology
67(3): 695-712 Taylor, R.B., J. Zrutledge, and J.G. Herrera. 1997.
A field guide to common south Texas shrubs.
Hernandez, F., C. Kleberg, and A. Radomski. Texas Parks and Wildlife Press, Austin, TX.
1999. Northern bobwhite and Rio Grande turkey p. 106.
management in relation to livestock grazing in
522
Thomas, P.A. 1991. Response of succulents to Weniger, D. 1991. Cacti of Texas and neighboring
fire: A review. International Journal of Wildland states. University of Texas Press, Austin, TX.
Fire 1(1): 11-22. 356 p.
Vines, R.A. 1986. Trees, shrubs, and woody vines Yeaton, R.I. 1978. A cyclical relationship between
of the Southwest. Sixth printing. University of Larrea tridentata and Opuntia leptocaulis in the
Texas Press. Austin, TX. 1,104 p. northern Chihuahuan desert. Journal of Ecology.
66: 651-656.
523
Palicourea crocea (Sw.) J.A. Schultes cachimbo
RUBIACEAE
John K. Francis
524
Reproduction.—Cachimbo flowers and fruits References
irregularly throughout the year (Little and others
1974). The fruits in one Puerto Rican sample Devoe, N.N. 1989. Differential seeding and
weighed an average of 0.1488 + 0.0055 g/fruit. regeneration in openings and beneath closed
Seeds cleaned from the sample averaged 0.0314 + canopy in sub-tropical wet forest. Ph.D.
0.0048 g/seed or 32,000 seeds/kg. Thirty-five dissertation. Yale University, New Haven, CN.
percent of these seeds germinated between 39 and 307 p.
84 days after sowing in commercial potting mix.
Palicourea flowers are pollinated by humming Fetcher, N., B.L. Haines, R.A. Cordero, D.J.
birds and the fruits are dispersed by frugiverous Lodge, L.R. Walker, D.S. Fernandez, and W.T.
birds (Taylor 1996). Bees were also observed to Lawrence. 1996. Responses of tropical plants to
visit the flowers (Labrón 1977). Cachimbo suffers nutrients and light on a landslide in Puerto Rico.
relatively little seed loss to pathogens and Journal of Ecology 84(3): 331-341.
predators and germinates at a higher percent after
natural dispersal than without it (Myster 1997). Howard, R. A. 1989. Flora of the Lesser Antilles,
Seedlings are common and widespread in suitable Leeward and Windward Islands. Vol. 6. Arnold
habitat. There is no significant difference between Arboretum, Harvard University, Jamaica Plain,
seasons in seedling appearance, but regardless of MA. 658 p.
season, seed germination is always higher in the
open than in closed canopy areas (Labrón 1977). Labrón, M. L. 1977. An autoecological study of
Transplanted wildlings survive and grow poorly Palicourea riparia Bentham (RUBIACEAE): an
(personal communication with F.N. Scatena, IITF, ecologically important species in the recovery of
Río Piedras, PR) a disturbed tropical rain forest in Puerto Rico.
Ph.D. Dissertation. University of North
Growth and Management.—Cachimbo grows Carolina, Chapel Hill, NC. 238 p.
slowly (a few cm) the first year and rapidly (1 m
or more per year) for about 2 years in openings Liogier, H. A. 1997. Descriptive flora of Puerto
(Labrón 1977). After about 3 years, seedlings Rico and adjacent islands. Vol. 5. Editorial de la
sown under an open canopy had accumulated Universidad de Puerto Rico, San Juan, PR.
about 20 times more biomass than seedlings sown 436 p.
under closed canopy conditions. The above-
ground to below-ground ratio of the open-canopy Little, E.L., Jr., R.O. Woodbury, and F.H.
plants was 4.8 (Labrón 1977). Cachimbo may Wadsworth. 1974. Trees of Puerto Rico and the
reach 6 m in height and 7.5 cm in stem diameter Virgin Islands. Vol. 2. Agriculture Handbook
(Howard 1989, Little and others 1974). No annual 449. U.S. Department of Agriculture,
rings are visible. However, single stems last about Washington, DC. 1,024 p.
10 years, and plants can continue to regenerate
with new sprouts. Plants cut during plantation Myster, R.W. 1997. Seed predation, disease and
cleaning and trail clearing quickly regain their germination on landslides in neotropical lower
former height (author’s observation). Scatena and montane wet forest. Journal of Vegetation
others (1993) present an equation to predict oven- Science 8(1): 55-64.
dry above-ground biomass (To) from diameter at
breast height (D) and height (H): To = exp(0.752 Scatena, F.N., W. Silver, T. Siccama, A. Johnson,
ln (D2H) – 2.362) where R2 = 0.861. and M.J. Sánchez. 1993. Biomass and nutrient
content of the Bisley Experimental Watershed,
Benefits.—Cachimbo is a food source for several Luquillo Experimental Forest, Puerto Rico,
species of song birds (Devoe 1989). It is higher in before and after Hurricane Hugo, 1989.
the major nutrients in its tissues than most of the Biotropica 25(1): 15-27.
trees and shrubs with which it is associated
(Scatena and others 1993) and is probably a good Taylor, C.M. 1996. Palicourea (Rubiaceae).
recycler of nutrients and a good forage plant. www.mobot.org/MOBOT/Staff/Research/taylor
People in some areas of Puerto Rico use crushed //palihome.html. 9 p.
cachimbo leaves to stop bleeding (Labrón 1977).
525
Zimmerman, J.K., T.M. Aide, M. Rosario, M.
Serrano, and L. Herrera. 1995. Effects of land
management and a recent hurricane on forest
structure and composition in the Luquillo
Experimental Forest, Puerto Rico. Forest
Ecology and Management 77(1-3): 65-76.
526
Parathesis crenulata (Vent.) Hook. f. rascagarganta
MYRSINACEAE
John K. Francis
annual precipitation. In the dry end of the
precipitation range, it only occurs near streams
and extra moist hollows. The species is found
mainly on slightly- to moderately-acid soils with
loamy to clayey texture. Although areas with both
igneous and sedimentary rocks are colonized, it is
most abundant in valleys in limestone formations.
Rascagarganta grows from a little above sea level
to 760 m in elevation (Little and others 1974). The
species is shade tolerant and prefers to grow under
closed forests with moderate to light understory
vegetation. It sometimes forms its own understory
canopy under which little vegetation grows. There
appear to be few insect or disease problems in
natural stands.
Reproduction.—Rascagarganta is reported to
flower from May to July and fruits from July to
December (Little and others 1974). Fruits are
often present in quantity and are easy to pick.
General Description.—Rascagarganta, also They may be cleaned by maceration and wet
known as secagarganta, jalapón, and raisin sieving. A collection of fruits from Puerto Rico
marron, is a shrub or small tree 1 to 3 m weighed an average of 0.407 g/fruit. Air-dried
(occasionally reaching 6 m) in height and 1 to 4 seeds cleaned from that collection weighed an
cm (occasionally to 7.5 cm) in trunk diameter. The average of 0.054 g/seed or 18,000 seeds/kg. Sown
common names in Spanish imply that it will make in peat, 45 percent germinated between 48 and
one’s throat raspy or dry. The species, unless 104 days after sowing. Many seedlings died from
disturbed, has a single stem and is supported by a damping-off; therefore, a looser germination
taproot and lateral roots that are orange-brown in medium than peat is recommended. Seedlings
color. Most of the branches arise from the main grew slowly in the nursery, reaching less than 10
stem, often in whorls of three’s or four’s. The cm in 6 months (author’s observation). Birds
alternate, dark-green, 7- to 23-cm leaves are disperse the seeds. Seedlings are relatively
narrowly elliptic, pointed at both ends, and often common and widespread near fruit-bearing stands.
have wavy-toothed edges. The small pink flowers
are borne in large numbers in terminal panicles. Growth and Management.—Rascagarganta
The 6- to 8-mm fleshy fruits progress from green grows slowly and appears to live several decades.
to pink to red to black as they ripen. Each fruit No management experience has been published.
contains one seed (Howard 1989, Liogier 1995, However, the best strategy for establishment of
Little and others 1974). natural stands may be to maintain plantations or
natural forest stands in moist areas for long
Range.—Rascagarganta is native to Hispaniola, periods with closed canopies and relatively clear
Puerto Rico, and Martinique (Howard 1989, understories to allow rascagarganta to build up.
Liogier 1995, Little and others 1974). Some
taxonomists suggest that P. serrulata (Sw.) Mez, Benefits.—Rascagarganta contributes to
of Cuba and Hispaniola is synonymous with P. biodiversity, cover for wildlife, and protects the
crenulata (Howard 1989). soil from erosion in the forests where it grows.
Although not usually heavily used by wildlife, the
Ecology.—Rascagarganta grows in moist forests fruits are produced in large quantity and are
that receive from about 1500 mm to 2800 mm of available after hurricanes when overstory fruits are
527
scarce. The wood is hard, but due to its small size Liogier, H.A. 1995. Descriptive flora of Puerto
is useful mainly for stakes and fuel. Rico and adjacent islands. Vol. 4. Editorial de la
Universidad de Puerto Rico, San Juan, PR.
References 617 p.
Howard, R. A. 1989. Flora of the Lesser Antilles, Little, E.L., Jr., R.O. Woodbury, and F.H.
Leeward and Windward Islands. Vol. 6. Arnold Wadsworth. 1974. Trees of Puerto Rico and the
Arboretum, Harvard University, Jamaica Plain, Virgin Islands. Vol. 2. Agriculture Handbook
MA. 658 p. 449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
528
Parkinsonia aculeata L. Jerusalem-thorn
FABACEAE
Synonyms: none
John K. Francis
529
were scarified and germinated at 59 percent, References
beginning 2 days after sowing (Francis and
Rodríguez 1993). Jerusalem-thorn produces two Correll, D.S., and M.C. Johnston. 1970. Manuel of
kinds of seeds. About 25 percent of them have thin the vascular plants of Texas. Texas Research
testae and will germinate readily without Foundation, Renner, TX. 1,881 p.
pretreatment; the rest have hard seed coats and
must be scarified before they will germinate Everitt, J.H. 1983. Seed germination
(Floridata 2001). Mechanical scarification was characteristics of two woody legumes (retama
used by the author; soaking for 45 minutes in and twisted Acacia) from South Texas. Journal
concentrated sulfuric acid worked equally well in of Range Management 36(4): 411-414.
another test (Everitt 1983). In nature, seeds are
transported by water, birds, and animals (Pacific Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs
Island Ecosystems at Risk 2001). Artificial and cacti of South Texas. Texas Tech University
propagation is routinely done with seeds followed Press. 213 p.
by ordinary nursery culture in containers. In vitro
propagation using nodal explants has been Floridata. 2001. Plant profile: Parkinsonia
demonstrated (Jaideep and others 1992). aculeata. http://www.floridata.com/ref/p/
Jerusalem-thorn will continue to resprout after park_acu.cfm. 3 p.
many cycles of annual disturbance.
Francis, J.K. and A. Rodríguez. 1993. Seeds of
Growth and Management.—The growth rate is Puerto Rican trees and shrubs: second
moderate, from 0.5 to 1 m per year in early years. installment. Research Note SO-374. U.S.
Jerusalem-thorn plants live about 30 years (Pima Department of Agriculture, Forest Service,
Community College 2001). As an exotic, Southern Forest Experiment Station. New
Jerusalem-thorn can be problematic in rangeland, Orleans, LA. 5 p.
as it has become in Australia. Biological control
attempts are ongoing, but have not yet been Harris, S.C. 1982. Nitrogen fixation by tropical
successful. Jerusalem-thorn can be controlled with woody legumes: potential source of soil
tractors by pulling or dozing the large plants out, if enrichment. In: J.P. Roskoski, J. Monano, C. van
the seedlings are also controlled. Good control has Kessel, G. Castilleja, and P.H. Graham, eds.
been achieved with several herbicides as well Biological nitrogen fixation technology for
(Pacific Island Ecosystems at Risk 2001). tropical agriculture. Centro International de
Agricultura Tropical. Cali, Colomabia. p. 447-
Benefits.—The principal use of Jerusalem-thorn 454.
today is for landscaping. Its abundant yellow floral
display alone justifies its use. Perhaps its best use Hoekstra, D.A., P.J. Wood, and F. Anap Sang.
is in xeric gardens, where it contrasts well with 2001. Proposed second phase dryland
succulents and other desert plants (Floridata agroforestry research project.
2001). It is also useful in hedges and for living http://www.idrc.ca/library/document/031091/
fenceposts (Hoekstra and others 2001). Jerusalem- 31 p.
thorn is also planted for environmental restoration
in desert areas. Besides being hardy, it has the Holdridge, L.R. and L.J. Poveda A. 1975. Arboles
added benefit of fixing nitrogen (Harris 1982). de Costa Rica. Vol. 1. Centro Científico
The wood is moderately hard, heavy (specific Tropical, San José, Costa Rica. 546 p.
gravity 0.6), and brittle. It is used principally for
fuel. Native Americans harvested the seeds, which Howard, R.A. 1988. Flora of the Lesser Antilles,
they parched before eating (Floridata 2001). Leeward and Windward Islands.
Various extracts of leaves, flowers, fruits, and Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain,
bark are used in herbal medicine to treat arthritis MA: Arnold Arboretum, Harvard University.
and fever, and as a nerve stimulant and abortive 673 p.
(Liogier 1990). Livestock eat the foliage and fruits
(Little and Wadsworth 1964). Jerusalem-thorn is Jaideep. M., S. Mukunthakumar, and J. Mathur.
occasionally browsed by white-tailed deer and the 1992. Micropropagation of Bauhinia variegata
seeds are eaten by bobwhite quail (Everitt and and Parkinsonia aculeata from nodal explants of
Drawe 1993). mature trees. Plant Cell, Tissue and Organ
530
Culture 28(1): 119-121. Department of Agriculture, Forest Service,
Washington, DC. 548 p.
Jones, C.E. and S.L. Buchmann. 1974. Ultraviolet
floral patterns as functional orientation cues in Pacific Island Ecosystems at Risk. 2001. Invasive
hymonopterous pollination systems. Animal plant species: Parkinsonia aculeata L. Fabaceae.
Behavior 22(2): 481-485. http://www.hear.org/pier/paacu.htm. 2 p.
Liogier H.A. 1988. Descriptive flora of Puerto Pantulu, J.V. 1942. Chromosome numbers of some
Rico and adjacent islands, Spermatophyta. Vol. Caesalpiniaceae. Current Science 11: 152-153.
2. Editorial de la Universidad de Puerto Rico,
Río Piedras, PR. 481 p. Piaggio, M. 2001. Plantas natives del Uruguay.
Red Academica de Uruguay.
Liogier, H.A. 1990. Plantas medicinales de Puerto http://www.rau.edu.uy/uruguay/flora/Uy.flora3.
Rico y del Caribe. Iberoamricana de Ediciones, htm. 8 p.
Inc., San Juan, PR. 566 p.
Pima Community Collage. 2001. Desert ecology
Little, E.L., Jr., and F.H. Wadsworth. 1964. of Tucson: mexican paloverde (Parkinsonia
Common trees of Puerto Rico and the Virgin aculeata). http://wc.pima.edu/Bfiero/
Islands. Agriculture Handbook 249. U.S. tucsonecology/plants/trees_mpv.htm. 1 p.
531
Parthenocissus quinquefolia (L.) Planch. Virginia creeper
VITACEAE
John K. Francis
532
percent are sound (Gill and Pogge 1974). The diarrhea and difficult urination, infusions for
seeds are dispersed by birds (Dirr 2003). Seeds jaundice, and applied decoctions and poultices
germinate the first or second spring after dispersal; topically for swelling and lockjaw (Moerman
germination is epigeal (Gill and Pogge 1974). It 1986). The fruits are reported to be poisonous by
layers readily at the nodes when stems come in reason of the oxalic acid they contain (Russell and
contact with the soil (Crandall and Crandall 1995). others 1997). However, consuming more than a
small amount is unlikely due to their bad flavor
Growth and Management.—Once well (author, personal observation).
established, Virginia creeper grows rapidly. It
often must be pruned to prevent it from getting out References
of control in ornamental settings. It can seed into
nearby flower beds and wildlands. Virginia Coladoanto, M. 1991. Parthenocissus
creeper should not be planted to climb up quinquefolia. In: U.S. Department of
buildings with wooden siding because it is difficult Agriculture, Forest Service, Rocky Mountain
to remove and because it increases the humidity Research Station, Fire Sciences Laboratory, Fire
and hastens rot (Gilman 1999). The species Effects Information System. http://www.fs.fed.
occasionally interferes with reproduction of pines us/database/feis/plants/vine/parqui/all.html.
and hardwoods and may be controlled with 10 p.
herbicides. Fire can also be an effective control
agent (Coladoanto 1991). Fruits can usually be Compton, G. 2003. Cambridgeshire flora records
picked in quantity by hand. Seeds are separated since 1538. http://mnlg.com/gc/species2pz/
from fruit tissue by maceration followed by p/par_qui.html. [not paged].
floating off the pulp and empty seeds with water.
However, maceration must be gentle because seed Crandall, C. and B. Crandall. 1995. Flowering,
coats are soft and easily damaged. There are 16 to fruiting, and foliage vines. Sterling Publishing
27 g of seeds/100 g of fruits. The seeds may be Co., Inc., New York. 192 p.
fall-sown 1 cm deep in soil and mulched.
Alternatively, seeds may be stratified in moist peat Dirr, M.A. 2003. Parthenocissus quinquefolia—
or sand at 5 or 6 °C for 60 days and sown in the Virginia creeper, also called woodbine.
spring. Forty-one to 94 percent of the seeds will http://www.nobleplants.com/classnotes/spring/
germinate after 30 days. Seedlings are ready for springprofiles/vines/parthenoquinque.htm. 1 p.
outplanting when they reach 15 cm in height after
1 or 2 years in the nursery. The species can also be Gill, J.D. and F.L. Pogge. 1974. Parthenocissus
propagated with cuttings and layers (Gill and Planch., creeper. In: C.S. Schopmeyer, tech.
Pogge 1974). coord. Seeds of woody plants in the United
States. U.S. Department of Agriculture, Forest
Benefits.—Virginia creeper is important to the Service, Washington, DC. p. 568-571.
aesthetics of forests, helps protect the soil, and
furnishes food and cover for wildlife. Cattle and Gilman, E.F. 1999. Parthenocissus quinquefolia.
white-tailed deer sometimes browse the foliage. Fact Sheet FPS-454. Cooperative Extension
Songbirds are the principal consumers of the fruits. Service, University of Florida, Gainesville, FL.
The foliage, both as a ground cover and climber, 3 p.
provides cover for small mammals and birds
(Coladoanto 1991). It has been planted for Moerman, D.E. 1986. Medicinal plants of Native
watershed protection and erosion control (Gill and America. Technical Reports. 19. University of
Pogge 1974). The principal attraction of Virginia Michigan Museum of Anthropology, Ann
creeper is its fall color. The leaves turn fiery Arbor, MI. 534 p.
shades of purple, red, and scarlet after the first
frost (Dirr 2003). The blue-black fruits and their Natural Resources Conservation Service. 2003.
stems (which turn red also) add color and interest Plants profile: Parthenocissus quinquefolia (L.)
after the leaves have fallen. The species is planted Planch. http://plants.udsa.gov/cgi_bin/
to cover building walls, run up tree trunks, and plant_search.cgi?mode=Common+Name&keyw
form trellises and arbors. It also makes a fine ordquery=Vi.... 4 p.
ground cover (Crandall and Crandall 1995). Native
Americans took decoctions of the roots for
533
Nelson, G. 1996. The shrubs and woody vines of hort/consumer/poison/Parthqu.htm. 2 p.
Florida. Pineapple Press, Inc., Sarasota, FL.
391 p. Services Culture Editions Ressources. 2003.
Parthenocissus quinquefolia (Linné) Planchon.
Rehder, A. 1951. Manual of cultivated trees and http://crdp.ac-besancon.fr/ressourc/flore/flore/
shrubs hardy in North America. The MacMillan especes/parth_quinq.htm. 1 p.
Company, New York. 996 p.
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Russell, A.B., J.W. Hardin, L. Grand, and A. Higgins, eds. 1987. A Utah flora. Great Basin
Fraser. 1997. Poisonous plants: Parthenocissus Naturalist Memoirs 9. Brigham Young
quinquefolia. North Carolina State University, University, Provo, UT. 894 p.
Raleigh, NC. http://www.ces.ncsu.edu/depts/
534
Passiflora edulis Sims passionfruit
PASSIFLORACEAE
Synonyms: none
John K. Francis
535
Yellow passionfruit in Puerto Rico flower from inoculation with superior strains of fungi
April to September and yield fruits from June to (Cavalcante and others 2001). Wild plants are
October (Acevedo-Rodríguez 1985). In some usually scattered and attempts at management
areas, plants fruit twice each year (Popenoe 1920). have not been reported.
Plants usually begin blooming and fruiting in their
second year. Yellow passionfruit flowers have Benefits.—By far the greatest benefit of
both male and female parts but are self-sterile. passionfruit to humankind is its fruit and the
They rely mainly on carpenter bees (Xylocopa delicious juice made from it. In addition to being
spp.) for pollination. Other insects and collected by local people in the forests, the fruit is
hummingbirds also visit the flowers. The flowers now grown in vineyards in dozens of countries. It
of purple passionfruit can self-pollinate (Morton is condensed, frozen, and shipped worldwide. The
1987). Fruits of the naturalized yellow-fruited fruit pulp contains 2.2 percent protein, 0.7 percent
form range from about 45 to 120 g in Puerto Rico. fat, and 21.2 percent carbohydrates. In addition,
There is a large variation between plants in size the seeds contain 23 percent oil similar to
and shape of fruits. Small fruits are sometimes sunflower or soybean oil, and the rind residue is
completely devoid of seeds, and large fruits may used for cattle feed (Morton 1987). The fruits of
have over 200 seeds. A collection of seeds from native and naturalized stands furnish food for
naturalized plants in Puerto Rico averaged 0.0251 numerous species of wild mammals and birds. The
+ 0.0004 g/seed or 40,000 seeds/kg. Passionfruit is whole plant, especially the leaves, contains
usually propagated from seeds but can be started alkaloids and a number of other phytoactive
from cuttings, layers, and grafts (Morton 1987). chemicals. Among these is passiflorine, a known
Seeds germinate best if allowed to ferment for a sedative and tranquilizer (Morton 1987). Extracts
few days in the fruit pulp before cleaning and are of the leaves have been used for centuries as
lightly scarified by clipping or sandpapering sedatives by native Brazilians. They prepare a
(Morton 1987). A group of seeds in Puerto Rico drink from the flower to treat asthma, bronchitis,
were sown without pretreatment in commercial and whooping cough. The plant is also used as a
potting mix and began germinating in 14 days and diuretic to treat urinary infections (Rain-tree
completed germination in 24 days with 61 percent 2002).
germinated. Plants are grown in beds or pots and
transplanted when they reach about 25 cm in References
height. The seedlings are heavily watered after
outplanting (Morton 1987). Seeds are disbursed in Acevedo-Rodríguez, P. 1985. Los bejucos de
the wild by humans, animals especially pigs, and Puerto Rico. Vol. 1. General Technical Report
birds, and by vine extension. SO-58. U.S. Department of Agriculture, Forest
Service, Southern Forest Experiment Station,
Growth and Management.—Plants in fertile soil New Orleans, LA. 331 p.
extend their stems about 3 m per year. Each year
during the annual dry season, the leaves fall off Anonymous. 2001. Exotic ornamental. The Hindu.
and the twigs die, leaving the main stem and a few http://www.hinduonnet.com/thehindu/2001/04/
important branches alive to rebuild the crown after 22/stories/1322045j.htm. 3 p.
the rains begin. Because fruiting takes place on
new wood, light pruning does not reduce yield. Bailey, L.H. 1941. The standard cyclopedia of
Plants live from 3 to 8 years and do not resprout. horticulture. Vol. 3. The MacMillan Co. New
Commercial stands are managed in vineyards, York. p. 2,423-3,639.
somewhat like grapes. They are planted in trellised
rows 4.5 m apart and spaced 4.5 m apart within Cavalcante, U.M.T., L.C. Maia, C.M.C. Costa, and
rows. The orchards are replanted every 4 to 6 V.F. Santos. 2001. Mycorrhizal dependency of
years (Bailey 1941). Alternately, vineyards may passion fruit (Passiflora edulis f. flavicarpa).
be established using small trees or bamboo as Fruits 56: 317-324.
standards (Heenkenda and Punchikumarihami
1991). Fruiting plants can even be grown in pots Heenkenda, H.M.S. and S.M.A.
under glass (Anonymous. 2001). Fruits fall after Punchikumarihami. 1991. Alternate trellising
ripening on the vines and are picked up from the materials for passion fruit (Passiflora edulis
ground at least twice per week. The roots normally Sims) vines. Tropical Agriculturist 147: 11-18.
form mycorrhizal associations and benefit from
536
Liogier, H.A. 1994. Descriptive flora of Puerto Popenoe, W. 1920. Manual of tropical and
Rico and adjacent Islands. Vol. 3. Editorial de la subtropical fruits. The MacMillan Company.
Universidad de Puerto Rico, Río Piedras, PR. New York. 474 p.
461 p.
Rain-tree. 2002. Maracuja, “passion fruit”.
Morton, J.F. 1987. Fruits of warm climates. http://rain-tree.com/passionf.htm. 3 p.
Creative Resources Systems, Inc. Winterville,
NC. 505 p. Sentelhas, P.C., C.T.P. Junior, J.M.M Sigristi, R.
Kavati, and M.T. Parodi. 1996. Freezing points
Pacific Island Ecosystems at Risk. 2002. Invasive of various tropical fruits. Bragantina 55(2): 231-
plant species: Passiflora edulis Sims, 235.
Passifloraceae. http://hear.org/pier/paedu/htm.
2 p.
537
Paullinia pinnata L. bejuco de costilla
SAPINDACEAE
Synonyms: none
John K. Francis
538
intolerant, die or are seriously weakened and do Howard, R.A. 1989. Flora of the Lesser Antilles,
not pose a serious threat to the reproduction stand. Leeward and Windward Islands. Vol. 5. Arnold
Herbicides can also be applied to the sprouts to Arboretum, Harvard University, Jamaica Plain,
make the method more effective, especially in MA. 604 p.
more open stands. Bejuco de costilla is one of the
alternate hosts of the tsetse flies (Glossina palpalis Liogier, H.A. 1990. Plantas medicinales de Puerto
and G. tachnioides) and has become the target of Rico y del Caribe. Iberoamericana de Ediciones,
eradication measures in some parts of tropical Inc., San Juan, PR. 566 p.
Africa (Morris 1944).
Liogier, H.A. 1994. Descriptive flora of Puerto
Benefits.—The stems of bejuco de costilla are Rico and adjacent Islands. Vol. 3. Editorial de la
used to make baskets. Ground leaves and seeds are Universidad de Puerto Rico, Río Piedras, PR.
used to stupefy fish and in preparing herbal 461 p.
remedies (Acevedo-Rodriguez 1985). The seeds
and leaves of this species are poisonous to humans Morris, K.R.S. 1944. A large-scale experiment in
and are reported to have been used in arrow the eradication of tsetse (Glossina palpalis and
poisons (Liogier 1990). Aqueous extracts of G. tachninoides). Farm and Forest 5: 149-156.
bejuco de costilla from Africa demonstrated
inhibitory effects against several important Souza, C. de, K.K. Amegavi, K. Koumaglo, M.
infectious organisms (Souza and others 1993). The Gbeassor, and C. De Souza. 1993. Study of the
white aril associated with the seed is reported to be antimicrobial activity of the total aqueous
edible (Vélez and Overbeek 1950). extracts of ten medicinal plants. Revue de
Medecines et Pharmacopees Africaines 7: 109-
References 115.
Acevedo-Rodriguez, P. 1985. Los bejucos de Vélez, I. and J. van Overbeek. 1950. Plantas
Puerto Rico. Vol. 1. General Technical Report indeseables en los cultivos tropicales. Editorial
SO-58. U.S. Department of Agriculture, Forest Universitaria, Río Piedras, PR. 497 p.
Service, Southern Forest Experiment Station,
New Orleans, LA. 331 p.
539
Pavonia spinifex (L.) Cav. cadillo espinoso
MALVACEAE
John K. Francis
540
rapid (about 1 m) from sprouts arising from References
established suffruticose plants. Growth begins
during the early summer rains and continues Griffiths, M. 1994. Index of garden plants. Timber
through the fall rainy season. The leaves and Press, Portland, OR. 1,234 p.
branches dry out and die during the late winter-
early spring dry season (in Puerto Rico). Howard, R.A. 1989. Flora of the Lesser Antilles,
Individual plants probably live at least a decade. Leeward and Windward Islands. Vol. 5. Arnold
Except for the spiny fruits that cling to clothing, Arboretum, Harvard University, Jamaica Plain,
there seems to be little need to control cadillo MA. 604 p.
espinosa in range or forest land. When necessary,
the species probably could be controlled by spot Liogier, H.A. 1994. Descriptive flora of Puerto
spraying with broadleaf weed killer to eliminate Rico and adjacent Islands. Vol. 3. Editorial de la
individual plants. The development of a closed Universidad de Puerto Rico, Río Piedras, PR.
canopy forest eventually eliminates the shrub. 461 p.
Benefits.—Cadillo espinoso is planted to a limited Liogier, H.A. 1990. Plantas medicinales de Puerto
extent as an ornamental for its handsome yellow Rico y del Caribe. Iberoamericana de Ediciones,
flowers. It contributes to the recovery of disturbed Inc., San Juan, PR. 566 p.
areas, furnishes cover for wildlife, and is eaten by
grazing animals. In herbal medicine, infusions of Nelson, G. 1996. The shrub and woody vines of
the flowers are used to treat sore throats and skin Florida. Pineapple Press, Inc., Sarasota, FL.
problems, and infusions of leaves and twigs are 391 p.
used to treat stomach problems, gall stones, and
liver pain (Liogier 1990). New York Botanical Garden. 2002. NYBG species
search results. http://s…/wwwspecimen.
search_list?taxon=Pavonia+spinifex+(L.)+Cav.+
+++&projcode=SEB. 1 p.
541
Pennisetum purpureum Schumacher elephant grass
POACEAE
Synonyms: none
John K. Francis
542
Colonization of new habitats is slow (Skerman young shoots are eaten in soups and stews (Burkill
and Riveros 1990). 1994). Elephant grass is used for mulch in East
Africa where a 25-cm depth of mulch is needed
Growth and Management.—The growth of for good weed control (Nishimoto 1994). Extracts
culms is vertical continually, but their weight of the plant are strongly diuretic and are used for
bends them in the middle and lower part, causing a that purpose in Africa. It is also used in a number
j-shaped habit. As the lower stem makes contact of other herbal remedies (Burkill 1994). The seeds
with moist soil, it roots at every node. Individual are eaten by many bird species. However, because
culms live more than 1 year, and by layering, they of the aggressive spread of the species, it is a
can continue until disturbed or a barrier is reached. menace to native vegetation in the Galapagos
New plantations are established by planting stem Islands (Mauchamp 1997) and at the margins of
pieces with at least three nodes or with root swamps and streams in Florida (Miami-Dade
cuttings (Skerman and Riveros 1990). In an County 2002).
agricultural research plot in Puerto Rico, elephant
grass reached the height of 4 m in 3 months References
(Barrett 1925). The record yield for heavily
fertilized elephant grass is 84,800 kg/ha/yr Agus, F., D.K. Cassel, and D.P. Garrity. 1996.
(Skerman and Riveros 1990). Not-withstanding its Soil-water and soil physical properties under
value as forage, elephant grass has become one of contour hedgerow systems on sloping Oxisols.
the worst weeds in the tropics because of the Soil and Tillage Research 40(3-4): 185-199.
difficulty of controlling it in croplands and fallow
areas. Cultivation alone is usually insufficient to Barrett, O.W. 1925. The food plants of Puerto
control it in croplands (Skerman and Riveros Rico. Journal of the Department of Agriculture
1990). Frequent mowing will cause it to be of Puerto Rico 9(2): 193.
replaced by other grasses. The herbicide
glyphosate provides acceptable control in at least Burkill, H.M. 1994. The useful plants of West
aquatic sites (McCann and others 1996). Tropical Africa. Royal Botanic Gardens. Kew,
UK. 636 p.
Benefits.—Elephant grass makes up the bulk of
the diet of forest elephants in West Cameroon Center for New Crops and Plant Products. 2002.
(Tchamba and Seme 1993). The species is an Pennisetum purpureum K. Schumach. Perdue
important forage and pasture grass in its native University. http://hort.purdue.edu/newcrop/
Africa and throughout the Tropics, especially for duke_energy/Pennisetum_purpurium.html. 4 p.
cattle. It is also cut for hay and fermented for
silage. A number of forage samples of different Holm, L.G., D.L. Plucknett, J.V. Pancho, and J.P.
ages of grass from several countries varied from 4 Herberger. 1977. The World’s worst weeds.
to 15 percent in crude protein, from 28 to 40 East-West Center. Honolulu, HI. 609 p.
percent in crude fiber, from 10 to 16 percent in
ash, from 0.9 to 3.8 percent in fat, and from 39 to Karschon, R. and D. Heth. 1958. Wind speed,
49 percent in nitrogen-free extract (Skerman and wind-borne salt and agricultural crops as
Riveros 1990). The species has been employed affected by windbreaks. La Yaaran 8(3/4): 8-13,
successfully using sown cuttings to replace 32-38.
Imperata swards in the Phillippines (Agus and Long, R.W. and O. Lakela. 1971. A flora of
others 1996, Skerman and Riveros 1990). It is Tropical Florida. University of Miami Press,
planted as hedgerows for erosion protection and Coral Gables, FL. 962 p.
forage production in the alley cropping system of
agroforestry (Magcale-Macandog and others 1998, Magcale-Macandog, D.B., C.D. Predo, K.M.
Menz and others 1999). The plant is also effective Menz, and A.D. Calub. 1998. Napier grass strips
as a windbreak for agricultural crops (Karschon and livestock: a bioeconomic analysis.
and Heth 1958). Lines of plants are used to mark Agroforestry Systems 40(1): 41-58.
boundaries between plots and properties. Elephant
grass is planted on riverbanks to prevent erosion. Mauchamp, A. 1997. Threats from alien plant
In Africa, the plant is used for thatch, and the thick species in the Galapagos Islands. Conservation
culms are made into fences, screens, and Biology 11(1): 260-263.
reinforcement for mud huts. The young leaves and
543
McCann, J.A., L.N. Arkin, and J.D. Williams. Natural Resources Conservation Service. 2002.
1996. Noniddigenous aquatic and semi-aquatic Plants profile: Pennisetum purpureum
plants in freshwater systems. University of Schumacher. Washington, DC. http://plants.
Florida, Center for Aquatic Plants. usda.gov/cgi_bin/plant_profile.cgi?symbol=PEP
http://aquat1.ifas.ufl.edu/mctitle.html. [Not U2. 3 p.
paged].
Nishimoto, R.K. 1994. Weed control in coffee
Menz, K.M., D. Magcale-Macandog, and I.W. plantations. In: R. Labrada, J.C. Caseley, and C.
Rusastra, eds. 1999. Improving smallholder Parker, eds. Weed management for developing
farming systems in Imperata areas of Southeast countries. FAO Plant Production and Protection
Asia: alternatives to shifting cultivation. Paper 120. Food and Agriculture Organization
Australian Centre for International Agricultural of the United Nations. Rome. p. 354-359.
Research. Canberra, Australia. 280 p.
Pacific Island Ecosystems at Risk. 2002. Invasive
Miami-Dade County. 2002. Napier grass- plant species: Pennisetum purpureum
Pennisetum purpureum. http://co.miami- Schumacher, Poaceae. http://www.hear.org/
dade.fl.us/derm/environment/badplants/plant%2 pier3/pepur.htm. 2 p.
0desc…/napier_grass.ht. 1 p.
Skerman, P.J. and F. Riveros. 1990. Tropical
grasses. FAO Plant Production and Protection
Series 23. Food and Agriculture Organization of
the United Nations. Rome. 832 p.
544
Penstemon ambiguus Torr. sand penstemon
SCROPHULARIACEAE
James E. Nellessen
545
1997). The sterile stamen (staminode) is beardless Baskin, C.C. and J.M. Baskin. 2001. Seeds:
(hairless), a feature different from most ecology, biogeography, and evolution of
penstemons (beardtongues), hence the alternate dormancy, and germination. Academic Press,
genus name Leiostemon, leio meaning smooth and San Diego, CA. 666 p.
stemon for stamen (Kirkpatrick 1992, Weber and
Wittman 2001). It blooms from May to as late as Carter, J.L. 1997. Trees and shrubs of New
October, depending on local environmental Mexico. Johnson Books, Boulder, CO. 534 p.
conditions, although mass blooming generally
occurs from June through July. The hybrid, P. Dick-Peddie, W.A. 1993. New Mexico
ambiguus x thurberi has smaller corollas, 12 to 15 Vegetation: Past, Present and Future. University
mm long, corolla lobes white to blue-purple, the of New Mexico Press, Albuquerque, NM. 244 p.
throat and tube pale pink to deep reddish purple.
The fruit is a capsule from 5 to 10 mm long that Epple, A.O. 1995. A field guide to the plants of
contains many very small (1.2 to 2 mm) seeds. Arizona. Falcon Publishing Inc., Helena, MT.
Seed germination characteristics (for example, the 347 p.
amount of cold stratification) vary as a function of
the elevation of the source population (Meyer and Great Plains Flora Association. 1986. Flora of the
others 1995). Sand penstemon seeds have a Great Plains. University Press of Kansas, KS.
physiological dormancy requiring 8 to 15 weeks of 1,392 p.
cold stratification before germination will occur
(Kitchen and Meyer 1991, Baskin and Baskin Harborne, J. B. 1988. Introduction to ecological
2001). biochemistry, 3rd Ed. Academic Press, London.
356 p.
Growth and Management.—Seedlings take a
few years becoming established before this long- Haflin, J. 1997. Penstemons: the beautiful
lived plant becomes larger and more conspicuous breadtongues of New Mexico. Jackrabbit Press,
in the landscape. Only a few flowers may be Albuquerque, NM. 50 p.
produced the first year of blooming, before the
plant gains enough mass and root reserves for the Kartesz, J.T. 1994. A synonymized checklist of
large floral displays that make this plant so the vascular flora of the United States, Canada,
characteristic. and Greenland. Vol. 1, 2nd Edition. Biota of
North America Program of the North Carolina
Benefits.—Since this species’ typical habitat is in Botanical Garden. Timber Press, Portland, OR.
sandy, unstable type soils, it acts as a soil 622 p.
stabilizer. Because of its conspicuous and colorful
floral displays, it is a welcome addition to habitats Kearney, T.H., R. Peebles, and Collaborators.
that might otherwise appear drab. This is an 1951 with 1960 supplement. Arizona flora.
attractive plant for native plant landscaping University of California Press, Berkeley.
(Morrow 1995, Warnock 1974), and in full bloom 1,085 p.
with the numerous stems and flowers per stem, it
makes for an attractive display of pink bouquets. Kirkpatrick, Z.M. 1992. Wildflowers of the
Western Plains. University of Texas Press.
References Austin, TX. 240 p.
Allred, K.W. 2002. A working index of New Kitchen, S.G. and S.E. Meyer. 1991. Seed
Mexico vascular plant names. Available on a germination of intermountain Penstemons as
New Mexico State University web page. influenced by stratification and GA3 treatments.
http://web.nmsu.edu/~kallred/herbweb. [not Journal of Environmental Horticulture 9: 51-56.
paged].
Martin, W.C. and C.E. Hutchins. 1981. (reprinted
Arslanian, R.L., T. Anderson, and F.R. Stermitz. 2001). A flora of New Mexico. Vol. 2. Bishen
1990. Iridoid glucosides of Penstemon Singh Mahendra Pal Singh, India and Koeltz
ambiguus. Journal of Natural Products--Lloydia Scientific Books, Germany) p. 1,277-2,591.
53(6): 1,485-1,489.
546
Meyer, S.E., S.G. Kitchen, and S.L. Carlson. 1995. Warnock, B.H. 1974. Wildflowers of the
Seed germination timing patterns in Guadalupe Mountains and the Sand Dune
intermountain Penstemon (Scrophulariaceae). Country, Texas. Sul Ross State University,
American Journal of Botany 82: 377-389. Alpine, TX. 176 p.
Morrow, B.H. 1995. Best plants for New Mexico Weber, W.A. and R.C. Wittman. 2001. Colorado
gardens and landscapes. University of New flora: Eastern Slope. University Press of
Mexico Press, Albuquerque, NM. 267 p. Colorado, Boulder CO. 521 p.
547
Philadelphus lewisii Pursh Lewis mock orange
HYDRANGEACEAE
Nancy L. Shaw
It is most common at mid-elevations but grows
from near sea level to 2,100 m (USDA Forest
Service 1937). Lewis mock orange exhibits wide
ecological amplitude, growing in communities
ranging from Pseudotsuga menziesii (Mirbel)
Franco and Sequoia sempervirens (Lamb. ex D.
Don) Endl. to Artemisia L., chaparral, and Pinus
contorta Dougl. ex Loud and Pinus ponderosa P.
& C. Lawson.
548
broadcast seeded on a rough seedbed and covered shafts from the long woody shoots (USDA Forest
using a Brillion seeder or similar device. They Service 1937).
may also be spot seeded in selected, prepared areas
that are well-drained and free of herbaceous References
competition. Seeds may be mixed with other shrub
seeds that require shallow or surface planting. Everett, P.C. 1957. A summary of the culture of
Bareroot stock may be produced by fall seeding or California plants at the Rancho Santa Ana
by seeding moist, prechilled seeds in spring. Botanic Garden. The Rancho Santa Ana
Seedlings develop rapidly and can be transplanted Botanic Garden, Claremont, CA. 223 p.
as 1-year-old stock (Stickney and others 2001).
Container stock is grown from seed or propagated Hitchcock, C.L., A. Cronquist, M. Ownbey, and
from hardwood or softwood cuttings (Marchant J.W. Thompson. 1961. Vascular plants of the
and Sherlock 1984). Rooted suckers and crown Pacific Northwest. Part 3: Saxifragaceae to
divisions are also used. Ericaceae. University of Washington Press,
Seattle, WA. 614 p.
Growth and Management.—Lewis mock orange
seedlings establish well on a wide variety of soils Hopkins, W.E. and B.L. Kovalchik. 1983. Plant
but should be protected from competition and associations of the Crooked River National
browsing. Plants grow at a moderate rate, flower Grassland. R6 Ecol. 133-1983. U.S.
reliably, and are generally free of insect and Department of Agriculture, Forest Service,
disease problems. Seedlings, however, are Pacific Northwest Region, Portland, OR. 98 p.
sometimes susceptible to damping off. Plants
grown from seed may begin flowering in the Kramer, N.B. and F.D. Johnson. 1987. Mature
second or third year (Everett 1957). Flowers are forest seed banks of three habitat types in
produced on twigs of the previous year; hence, central Idaho. Canadian Journal of Botany 65:
landscape plants should be pruned after flowering. 1961-1966.
Benefits.—Seedlings or larger planting stock of Kufeld, R.C. 1973. Foods eaten by the Rocky
Lewis mock orange are transplanted into steep, Mountain elk. Journal of Range Management
rocky, unstable slopes where they provide soil 26: 106-113.
stabilization and vegetative cover. The species is
also useful in transitional areas of degraded Leege, T.A. 1968. Prescribed burning for elk in
riparian zones. Lewis mock orange is usually not northern Idaho. Proceedings: Annual tall
grazed heavily by livestock, but in some areas it timbers fire ecology conference. 8: 235-253.
does receive fair amounts of use by cattle (Bos
spp. L.) and sheep (Ovis spp. L.) (Leege 1968, Marchant, C. and J. Sherlock. 1984. A guide to
USDA Forest Service 1937). It frequently occurs selection and propagation of some native
with other species that are more palatable to big woody species for land rehabilitation in British
game, and consequently, it may receive little use, Columbia. Forest Research Report RR84007-
except under severe conditions. However, it can HQ. Ministry of Forests, Victoria, BC. 117 p.
provide good browse for deer (Odocoileus spp.
Rafinesque) and elk (Cervus canadensis Stickney, P.F., N.L. Shaw, and E.G. Hurd. 2001.
Erxleben), especially on winter ranges (Kufeld Philadelphus L. Mockorange. In: F.T. Bonner
1973, Leege 1968, Marchant and Sherlock 1984, and R.G. Nisley, eds. Woody plant seed
USDA Forest Service 1937). New growth is manual. Agriculture Handbook. U.S.
generally highly palatable to big game (Leege Department of Agriculture, Forest Service,
1968). The species provides food and cover for Washington, D.C. http://wpsm.net/index.html.
birds and other small animals. Lewis mock orange 5 p.
is a valued landscape species because of its showy,
white flowers. It is used in borders, screens, Taylor, S. 1972. Philadelphus lewisii Pursh ‘mock
hedges, and as specimen plants (Marchant and orange’ or ‘syringa’. Davidsonia 3: 4-7.
Sherlock 1984). Several commercial cultivars of
Lewis mock orange have been developed. The USDA Forest Service. 1937. Range plant
flowers are used in preparing perfumes and teas handbook. U.S. Government Printing Office,
(Taylor 1972). Native Americans made arrow Washington, D.C. 512 p.
549
Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Goodrich. 1987. A Utah flora. Great Basin
Naturalist 9. Brigham Young University, Provo,
UT. 894 p.
550
Philadelphus microphyllus Gray littleleaf mock orange
HYDRANGEACEAE
Nancy L. Shaw
551
prechill at 3 to 5 oC when seeds were subsequently Hickman, J.C. 1993. The Jepson manual: higher
incubated at 20/10 oC, but not at 15 oC. plants of California. University of California
Germination is epigeal. Press, Berkeley, CA. 1,400 p.
Growth and Management.—Little data are Hitchcock, C.L. 1943. The xerophyllous species of
available on the propagation of littleleaf mock Philadelphus in southwestern North America.
orange, but species of the genus Philadelphus are Madroño 17: 35-56.
generally easily propagated from softwood or
hardwood cuttings, rooted suckers, divisions, or Macdonald, B. 1986. Practical woody plant
layers (Hartmann and others 1990, Macdonald propagation for nursery growers. Vol. I. Timber
1986). Littleleaf mock orange is easily Press, Portland, OR. 669 p.
transplanted (Sutton and Johnson 1974). Seeds of
Philadelphus spp. are tiny and may be broadcast Munz, P.A. and D.D. Keck. 1973. A California
seeded on a rough seedbed and covered lightly or flora and supplement. University of California
spot seeded on prepared seedbeds (Stevens and Press, Berkeley, CA. 1,681 p.
others, in press). Seeds may also be surface-
planted using a Brillion or similar seeder. Best
results are obtained if seeds are planted in well- Patton, D.R. and M.G. Ertl. 1982. Run wild
drained sites free of herbaceous competition. wildlife/habitat relationships. Wildlife Unit
Seeds may be mixed with other shrub seeds that Technical Series. U.S. Government Printing
require surface or shallow planting. Growth of Office, Washington, D.C. 49 p.
littleleaf mock orange is moderate to rapid (Sutton
and Johnson 1974). Rehder, A. 1940. Manual of cultivated trees and
shrubs. 2nd Ed. The MacMillan Co., New York.
Benefits.—Littleleaf mock orange provides good 996 p.
cover for small animals. It is used to some extent
by mule deer (Odocoileus hemionus Rafinesque) Stevens, R., S.B. Monsen, and N.L. Shaw. [in
(Patton and Ertl 1982). Although the species press]. Shrubs of other families. In: S.B. Monsen
receives little use in revegetation, it has potential and R. Stevens, comps. Restoring western
for planting on disturbances on steep, rocky, ranges and wildlands. INT-GTR. U.S.
unstable slopes within its native range (Stevens Department of Agriculture, Forest Service,
and others, in press). Seedlings or larger stock are Rocky Mountain Research Station, Ogden, UT.
recommended for such use. The species might
also be used to advantage in drier areas of Stickney, P.F., N.L. Shaw, and E.G. Hurd. 2001.
degraded riparian zones. Littleleaf mock orange is Philadelphus L. Mockorange. In: F.T. Bonner
an attractive ornamental because of its showy and R.G. Nisley, eds. Woody plant seed manual.
flowers and fall coloration. It was first cultivated U.S. Department of Agriculture, Forest Service,
in 1883 (Rehder 1940). It can be used in borders, Agriculture Handbook. Washington, D.C.
screens, hedges, or as isolated specimens in sunny http://wpsm.net/ index.html. 5 p.
areas. It can also be used for low maintenance
landscaping and in recreational area plantings. The Sutton, R. and C.W. Johnson. 1974. Landscape
small seeds were eaten by Native Americans plants from Utah’s mountains. EC-368. Utah
(Cronquist and others 1997). State University, Logan, UT. 137 p.
John K. Francis
553
within the crown of the host tree. It does not References
spread vegetatively.
Howard, R.A. 1988. Flora of the Lesser Antilles,
Growth and Management.—The growth rate of Leeward and Windward Islands.
guacimilla de canario is not known. Although it is Dicotyledoneae, Part 1. Vol. 4. Arnold
a parasite and does harm its hosts, the plant takes Arboretum, Harvard University, Jamaica Plain,
many years to become sufficiently abundant to MA. 673 p.
seriously weaken a host tree. Guazuma ulmifolia
and most of the other hosts are not used as Jorgensen, P.M., and S. León-Yánez. 1999.
ornamentals and are not commercially valuable. If Catalogue of the vascular plants of Ecuador.
control is desired, probably the best method is to Missouri Botanical Garden, St. Louis, MO.
clip the guacimilla de canario plants from the 1,181 p.
limbs each year or until they die. This will prevent
it from bearing fruit and spreading within the tree. Liogier, H.A. 1985. Descriptive flora of Puerto
Alternately, one may prune out the infected limbs Rico and adjacent islands, Spermatophyta. Vol.
of the parasitized tree. 1. Editorial de la Universidad de Puerto Rico,
Río Piedras, PR. 352 p.
Benefits and Detriments.—Guacimilla de
canario contributes in a minor way to biodiversity, Missouri Botanical Garden. 2002. Manual de la
to tree canopy density, and to biomass production. flora de Costa Rica. http://www.mobot.
It furnishes fruits for wildlife food. On the other org/manual.plantas/040567/S040679.html. 4 p.
hand, a few trees/km2 are attacked, some are
weakened, and an occasional tree dies as a result.
Parasitized trees are often unsightly.
554
Phragmites australis (Cav.) Trin. ex Steud. common reed
POACEAE
John K. Francis
555
resistance. A large number of fungal species and at sooth lung pain, especially during pneumonia, and
least one nematode have been isolated from the to treat stomach problems and diarrhea (Moerman
species, none of which cause extensive damage 1986). There are numerous other applications in
(Duke 1998). herbal medicine throughout the world. The stems
were used as arrow shafts by Native Americans
Reproduction.—Common reed blooms mostly (Duke 1998). Native people of Tasmania used the
from July through November (Diggs and others hollow stems to make rafts, jewelry, baskets, and
1999). It apparently blooms and fruits throughout light spears (Australian National Botanic Gardens
the year in frost-free areas (Gould 1975). Under 2003). Common reeds are harvested today in
favorable artificial conditions, germination begins Britain for thatching (Tyler-Walters 2002). They
1 day after sowing and may reach nearly 100 were once harvested for fuel where better sources
percent (Ekstam and Forseby 1999). Although were unavailable and may hold promise as an
common reed is a relatively good seed producer energy crop in the future. Stands yield 415
and seeds are added annually to the soil seed bank, calories/100g (Duke 1998). Early in the growing
successful seedling establishment is rare (Uchytil season, common reed is high-quality forage for
1992). New stands can also be started by flood- cattle and horses and may be cut for hay. It
borne pieces of rhizome. Once a new plant is contains 11.4 percent protein, 2.3 percent fat, 42.1
established, it spreads by vegetative means. percent carbohydrates, 31.1 percent crude fiber,
and 10.8 percent ash (Duke 1998). It becomes
Growth and Management.—Because stems are woody and unpalatable later in the season. The
renewed each growing season, annual height seeds are eaten by waterfowl and the roots are
growth is from 2 to 4 m per year. Growth can eaten by muskrats. The edges are occasionally
reach 4 cm/day. In southern Manitoba, stems reach used for nesting by ducks and the interior of the
their full height by the end of July (Uchytil 1992). stands are nesting habitat for snowy egrets, black-
During periods of falling water, colonies can crowned night herons, and yellow-headed
increase in width by as much as 15 m in a single blackbirds. A few species such as deer use the
season (Fewless 2003). Annual productivity ranges stands for escape cover (Uchytl 1992). It is
from 7.5 to 63 metric tons/ha (Duke 1998). If important habitat for wildlife in Britain and
harvested during the dormant season, productivity supports such species as the bittern, the reed
is not affected, except through compaction. New bunting, and the marsh harrier (Tyler-Walters
stands are established by planting rhizome 2002). Throughout the world the stems have been
segments (Duke 1998). This is normally not used for building materials, lattice work, baskets,
recommended because of the invasive nature of the and mats. The young shoots and rhizomes are
species. Eliminating stands is difficult. The best sometimes eaten, and the rhizomes processed for
approaches are heavy grazing by cattle, and starch. The stems are processed into pulp, paper,
spraying with amatrol, dalapon, or glyphosate with and fiberboards. Stalks contain over 50 percent
follow-up treatments. Stands can be degraded by cellulose and have fibers 0.8 to 3.0 mm long by
cultivation or summer mowing (Uchytil 1992). 5.0 to 30.5 µm in diameter. The dried plumes are
used for decorations and a variegated variety is
Benefits and Detriments.—Common reed employed as an ornamental (Duke 1998).
aggressively colonizes large areas of shoreline and
shallow marsh, and with its thick sod and heavy References
stand of stems effectively prevents wave- and
current-caused erosion (Uchytil 1992). It also traps Australian National Botanic Gardens. 2003.
sediment and pollutants. In fact the species has Aboriginal plant use in South-eastern Australia:
been used to aid in settling and drying sewage Phragmites australis. http://www.anbg.gov.
sludge (Graduate College of Marine Studies 2003). au/aborig.s.e.aust/phragmites-australis.html. 2 p.
The dense single-species stands completely
displace other native marsh communities (Fewless Blossey, B. 2002. Native to North America or
2003) and many of the fauna they support introduced (or both)? Ag Web, Cornell
(Weinstein and Balletto 1999). On the other hand, University, Ithaca, NY. http://www.
strong declines in populations in many European invasiveplants.net/phragmites/phrag/natint.htm.
countries have occurred during the last two 3 p.
decades (Clevering 1998). Native Americans used
the sap from green stems to loosen phlegm and
556
Britton, N.L. and A. Brown. 1913. Illustrated flora Harrington, H.D. 1964. Manual of the plants of
of the northern states, Canada, and the British Colorado. Sage Books, Denver, CO. 666 p.
possessions. Vol. 2, 2nd Ed. Scribner, New York.
735 p. Moerman, D.E. 1986. Medicinal plants of Native
America. Technical Reports. 19. University of
Clevering, O.A. 1998. An investigation into the Michigan Museum of Anthropology, Ann
effects of nitrogen on growth and morphology of Arbor, MI. 534 p.
stable and die-back populations of Phragmites
australes. Aquatic Botany 60(1): 11-25. Natural Resources Conservation Service. 2003.
Plants profile: Phragmetes australis (Cav.) Trin.
Diggs, G.M., Jr., B.L. Lipscomb, and R.J. ex Steud. http://plants.usda.gov/cgo_bom/
O’Kennon. 1999. Thinner’s and Mahler’s plant_search.cgi?mode=Scientific+Nave&keyw
Illustrated flora of North Central Texas. ordquery=Phragmites+australis&earl=plant_s…
Botanical Research Institute of Texas, Fort 6 p.
Worth, TX. 1,622 p.
Serag, M.S. 1996. Ecology and biomass of
Duke, J.A. 1998. Handbook of energy crops: Phragmites australes (Cav.) Trin. ex Steud. In
Phragmites australis (Cav.) Trin. ex Steud. the north-eastern region of the Nile Delta,
Center for New Crops and Plants Products, Egypt. Ecoscience 3(4): 473-482.
Purdue University, West Lafayette, IN. http://
www.hort.purdue.edu/newcrop/duke_energy/ Tyler-Walters, H. 2002. Phragmites australis,
phragmites_australis.html. 5 p. common reed. Marine Life Information
Network, Plymouth, UK. http://www.marlin.
Ekstam, B. and A. Forseby. 1999. Germination ac.uk/species/Phraus.htm. 4 p.
response of Phragmites australis and Typha
latifolia to diurnal fluctions in temperature. Seed Uchytil, R.J. 1992. Phragmites australis. In: U.S.
Science Research 9(2): 157-163. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences
Fewless, G. 2003. Invasive plants of Wisconsin: Laboratory, Fire Effects Information System.
Phragmites australis: common reed. http:// http://www.fs.fed.us/database/feis/plants/gramin
www.uwgb.edu/biodiversity/herbarium/invasive oid/phraus/all.html. 19 p.
_species/phraus01.htm. 2 p.
Weinstein, M.P. and J.H. Balletto. 1999. Does the
Gould, F.W. 1975. The grasses of Texas. Texas common reed, Phragmites australis, affect
A&M University Press, College Station, Texas. essential fish habitat? Estuaries 22(3B): 793-
653 p. 802.
Graduate College of Marine Studies. 2003. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Optimized reed (Phragmites australis) function Higgins, eds. 1987. A Utah flora. Great Basin
in sludge drying beds: producing a more Naturalist Memoirs 9. Brigham Young
effective genotype. University of Delaware, University, Provo, UT. 894 p.
Newark, DL. http://www.ocean.udel.edu/
level1/facultystaff/faculty/dseliskar/sludgebeds.
html. 3 p.
557
Physocarpus malvaceus (Greene) Kuntze mallow ninebark
ROSACEAE
John K. Francis
558
stands often owe their existence to logging or fires. sometimes appear later and will have to be sprayed
The species is an obligate pioneer that increases again (Habeck 2003).
and grows rapidly following disturbance, but as
competition, particularly overhead shade, Benefits.—Mallow ninebark helps protect the soil
increased, it decreases in mean height and during secondary succession until the shrub is
gradually disappears (Habeck 2003). The species replaced by taller vegetation. It adds beauty to
is generally free of insect and disease problems, forest openings and shrublands when its leaves
although it is occasionally attacked by foliage- turn russet-red in the fall (Clark 1976). The species
eating and seed-eating insects (Youngblood and provides cover for both large and small animals. It
others 2003). is one of the principal nesting sites of the dusky
flycatcher (Empidonax oberholseri) (Sedgwick
Reproduction.—Flowering takes place between 1993). The shrub is not browsed by cattle and is
late May and late July, depending on altitude, and utilized only to a limited extent by deer, bighorn
fruits ripen between late August and late sheep, and domestic sheep. The upper third of the
September (Habeck 2003). Seed weights are annual growth of plants from a burned area
reported (geographic source unspecified) at contained 1.3 percent N, 0.3 percent P, and 1.2
165,000 seeds/kg (Youngblood and others 2003). percent Ca. Plants in an unburned site contained
Seeds collected by the author in Utah averaged even lower levels (Habeck 2003). Early summer
158,000 seeds/kg. Viability is usually less than 50 sheep grazing increases the nutritional quality of
percent (Youngblood and others 2003). The fruits mallow ninebark for fall and winter browse (Alpe
split open at maturity but most of the seeds do not and others 1999).
immediately fall. Dispersal is by gravity, wind,
and probably by browsing animals. Seeds in the References
soil seedbank are reported to be 11 percent viable
and will germinate when the surface is scarified Abrams, L. 1950. Illustrated flora of the Pacific
(Habeck 2003). Ripe fruits can be picked by hand States. Vol. 2. Stanford University Press,
or beaten off shrubs onto a tarp. Fruits may be Stanford, CA. 635 p.
threshed by rubbing for small quantities or passed
through a hammer mill for larger quantities and Alpe M.J., J.L. Kingery, and J.C. Mosley. 1999.
screened and blown to clean them. They may be Effects of summer sheep grazing on browse
stored cool and dry for at least 5 years. Seeds may nutritive quality in autumn and winter. Journal
be sown in the fall, or in the spring after 30 days of of Wildlife Management 63(1): 346-354.
prechilling. Seeds should be sown to a depth of 3
mm and mulched with another 6 mm of sawdust Clark, L.J. 1976. Wild flowers of the Pacific
(Youngblood and others 2003). Mallow ninebark Northwest. Gray’s Publishing Limited, Sidney,
can easily be vegetatively reproduced with misted BC, Canada. 604 p.
softwood cuttings or by planting hardwood
cuttings directly in the field (Youngblood and Habeck, R.J. 2003. Physocarpus malvaceus. In:
others 2003). U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences
Growth and Management.—Mallow ninebark Laboratory. Fire Effects Information System.
shrubs in northern Idaho reached their maximum http://www.fs.fed.us/database/feis/plants/shrub/
height by the fourth growing season after a fire phymal/all.html. 17 p.
(Habeck 2003). Although stems are added and lost
continuously, shrubs do not grow taller. Individual Natural Resources Conservation Service. 2003.
plants may live for many years if growing Plant profile: Physocarpus malvaceus (Greene)
conditions permit. Mallow ninebark is sometimes Kuntze. Plants database. http://plants.usda.gov/
planted for soil stabilization and other cgi_bin/plant_profile.chi?symbol=PHMA5. 3 p.
conservation objectives. Killing the shrubs to
make room for more desirable species is Palouse Prairie Foundation. 2003. Palouse Prairie
sometimes attempted. Several herbicides can be Foundation native species search results:
effective if sprayed early in the season from late Physocarpus malvaceus. http://www.
foliar development to flowering or in early fall palouseprairie.org/readppf.pl?entry=PHMA5.
before leaves change color. Sprouts will 3 p.
559
Sedgwick, J.A. 1993. Reproductive ecology of Youngblood, A.Y., J.D. Gill, and F.L. Pogge.
dusky flycatchers in western Montana. Wilson 2003. Physocarpus (Camb.) Raf. U.S.
Bulletin 105(1): 84-92. Department of Agriculture, Forest Service,
Washington, DC. http://wpsm.net/Physocarpus.
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. pdf. 5 p.
Higgins, eds. 1987. A Utah flora. Great Basin
Naturalist Memoirs 9. Brigham Young
University, Provo, UT. 894 p.
560
Picramnia pentandra Sw. Florida bitterbush
SIMAROUBACEAE
John K. Francis
561
ripening occurs in November and December and fungi were treated with ethanol extracts of
(Pascarella 1996). Fruit and seed production is various tissues. Growth reductions of Bacillus
moderately abundant and consistent from year to cereus, Bacillus subtilis, and Pseudomonas
year. Fresh fruits collected in Puerto Rico weighed aeruginosa were observed which was enhanced by
an average of 0.231 + 0.043 g/fruit. Air-dried exposure to long-wave ultraviolet radiation
seeds separated from them averaged 11,900 (O’Neal and others 2002).
seeds/kg. Placed on moist filter paper without
pretreatment, 100 percent germinated, beginning References
14 days after sowing (Francis and Rodríguez
1993). Pascarella (1996) noted fruits collected in Francis, J.K. and A. Rodríguez. 1993. Seeds of
Florida weighed an average of 0.483 + 0.034 g and Puerto Rican trees and shrubs: second
that 85 to 90 percent of the fruits contained just installment. Research Note SO-374. U.S.
one seed. Most seeds fall under the parent trees but Department of Agriculture, Forest Service,
some are dispersed by birds. Seedlings are Southern Forest Experiment Station, New
relatively common in forests where the species Orleans, LA. 5 p.
grows.
Howard R.A. 1988. Flora of the Lesser Antilles,
Growth and Management.—Growth rate of Leeward and Windward Islands.
Florida bitterbush is slow to moderate. Weaver Dicotyledoneae, Part 1. Vol. 4. Arnold
(1979) followed the growth of two plants in a Arboretum, Harvard University, Jamaica Plain,
forest in Puerto Rico with characteristic slow MA. 673 p.
growth and found diameter growth rates over the
24-year period of about 0.2 cm/year. Florida Institute of Food and Agricultural Sciences. 2003.
bitterbush in Puerto Rico live about 30 to 60 years Native landscape plants for South Florida:
(author’s observation). Plants begin flowering and shrubs and small trees (P-Z). http://miami-
fruiting when they are about 1.5 to 2 m in height if dade.ifas.ufl.edu/programs/fyn/publications/nati
sufficient light is available. Fruits are easily veplantlist/npl-shrubs-small-trees-p-z.htm. 5 p.
collected in quantity by hand and with short
pruning poles. They may be cleaned by macerating Liogier H.A. 1988. Descriptive flora of Puerto
or by working them against a screen under running Rico and adjacent islands, Spermatophyta. Vol.
water. Production of seedlings, at least to the 2. Editorial de la Universidad de Puerto Rico,
pricking-out stage, is straightforward. Where a Río Piedras, PR. 481 p.
seed source is available in favorable habitat,
natural reproduction is usually adequate. Florida Liogier, H.A. 1990. Plantas medicinales de Puerto
bitterbush has not been reported to cause any Rico y del Caribe. Iberoamericana de Ediciones,
problems in forests and therefore control measures Inc., San Juan, PR. 566 p.
are probably not needed.
Little, E.L., Jr. and F.L. Wadsworth. 1964.
Benefits.—Florida bitterbush helps protect the soil Common trees of Puerto Rico and the Virgin
and contributes to the biodiversity and aesthetics Islands. Agriculture Handbook 249. U.S.
of forest stands. A number of bird and animal Department of Agriculture, Washington, DC.
species eat the fruits. It furnishes larval food for 548 p.
the bush sulphur butterfly, Urema dina helios
(Institute of Food and Agricultural Sciences 2003). Long, R.W. and O. Lakela. 1976. A flora of
Florida bitterbush is a honey plant (Little and Tropical Florida. University of Miami Press,
Wadsworth 1964). The wood is little used, except Coral Gables, FL. 962 p.
occasionally for fuel. Florida bitterbush is planted
in Florida and Cuba as an ornamental (Little and O’Neal, C.B., J. Salazar, and E. Rodríguez. 2002.
Wadsworth 1964). A bitter tonic is made from A preliminary analysis of the anti-microbial and
tissues of the species in the Caribbean to treat phytochemical properties of Picramnia
fevers, anorexia, and stomach problems and as an pentandra (Simaroubaceae) and its potential
enema to treat for worms (Liogier 1990). relationship to gastrointestinal health.
Qualitative tests indicated the presence of Emanations 4: 57.
antioxidants, phenols, terpenoids, and alkaloids in
ethanol extracts of tissues. A number of bacteria
562
Pascarella, J.B. 1996. Reproductive ecology of Weaver, P.L. 1979. Tree growth in several tropical
Picramnia pentandra (Picramniaceae) in South forests of Puerto Rico. Research Paper SO-152.
Florida. Caribbean Journal of Science 32(1): 99- U.S. Department of Agriculture, Forest Service,
104 Southern Forest Experiment Station, New
Orleans, LA. 15 p.
563
Piper aduncum L. bamboo piper
PIPERACEAE
John K. Francis
564
paper (author’s observation). The seeds are References
dispersed by bats, birds, and possibly arboreal
rodents (Plant Protection Service 2001). Seedlings Bourke, R.M. 1997. Management of fallow species
are fairly common on disturbed ground. composition with tree planting in Papua New
Established plants thicken into clumps by suckers Guinea. http://coombs.anu.edu.au/Depts/
arising from the root crown (Pacific Island RSPAS/RMAP/bourke.htm. 8 p.
Ecosystems at Risk 2002). Plants are established
for agroforestry purposes in Papua New Guinea by Gómez, P., D. Cubillo, G.A. Mora, and L. Hilje.
shoving cuttings into moist soil (Bourke 1997). 1997. Evaluation of possible repellents for
Bemisi tabaci. II. Botanical substances. Manejo
Growth and Management.—The growth of Integrado de Plagas 46: 17-25.
bamboo piper is moderately rapid after the early
seedling stage. Sprouts and suckers grow more Howard R.A. 1988. Flora of the Lesser Antilles,
than a meter in their first year. Producing Leeward and Windward Islands.
seedlings by means of seed is difficult. Dicotyledoneae, Part 1. Vol. 4. Arnold
Propagating plants with cuttings, which need no Arboretum, Harvard University, Jamaica Plain,
hormonal treatment, in the nursery or directly in MA. 673 p.
the field, is recommended. Individual stems live 2
to several years; by sprouting, plants live much Ibrahim, J. A. Abu-Said, A. Abdul-Rashih, M.A.
longer. Infestations of bamboo piper can be Nor-Azah, M.Z. Zaridah, A.K. Azizol, Norhara-
controlled by uprooting young plants and spraying Hussein, P.S. Bacon, and K.C. Khoo. 1996.
older plants with broadleaf weed killers such as Essential oils of selected Malaysian plants and
2,4-D. Treated areas should be checked in a few their potential uses. Forestry and Forest Products
weeks and treated again if regrowth has occurred Research, Proceedings of the Third Conference,
(Plant Protection Service 2001). Oct. 3-4, 1995; Kuala Lumpur, Malaysia. p. 97-
103.
Benefits.—Bamboo piper helps revegetate
disturbed areas and contributes to the biodiversity Instituto de Botánica Darwinion. 2002. Catálogo
and biomass of forests. It also is a source of food de las plantas vasculares de la Argentina.
and cover for wildlife. Bamboo piper established http://www.darwin.edu.ar/Catalogo/indicevascul
in contour rows for soil erosion brakes helps ares.htm. [n.p.].
facilitate agroforestry on steep land in Papua New
Guinea (Bourke 1997). The wood is useful for Liogier H.A. 1985. Descriptive flora of Puerto
fuel, stakes, fences, and rude construction Rico and adjacent islands, Spermatophyta. Vol.
(Philippine Council of Agriculture 2002). 1. Editorial de la Universidad de Puerto Rico,
However, exposed wood rots quickly (Vélez and Río Piedras, PR. 352 p.
van Overbeek 1950). The species is planted as an
ornamental. The peppery fruits have been used to Liogier, H.A. 1990. Plantas medicinales de Puerto
season food (Little and Wadsworth 1964). Rico y del Caribe. Iberoamericana de Ediciones,
Essential oil content of bamboo piper tissue Inc., San Juan, PR. 566 p.
(leaves and twigs), of which dillapiole is the major
component, ranged from 1.2 to 3.4 percent (Maia Little, E.L., Jr. and F.H. Wadsworth. 1964.
and others 1998). Teas and other extracts of the Common trees of Puerto Rico and the Virgin
leaves and roots of bamboo piper are used in Islands. Agriculture Handbook 249. U.S.
herbal medicine as a tonic to ease diarrhea, Department of Agriculture, Forest Service,
dysentery, vomiting, ulcers, and to control Washington, DC. 548 p.
bleeding (Liogier 1990). The chemical 2’, 6’-
dihydroxy-4’-methoxychalcone isolated from Maia, J.G.S., M.G.B. Zohhbi, E.H.A. Andrade,
bamboo piper inhibited 98 percent of the growth A.S. Santos, M.H.L. da Silva, A.I.R. Luz, C.N.
of Leishmania amazonensis parasites in vitro with Bastos, and M.H.L. da Silva. 1998. Constituents
low host cell toxicity (Torres-Santos and others of the essential oil of Piper aduncum L. growing
1996). In addition, the essential oils have shown wild in the Amazon region. Flavour and
strong insecticidal, molluscicidal, and antibacterial Fragrance Journal 13(4): 269-272.
effects (Gómez and others 1997, Ibrahim and
others 1996, Orjala and others 1992). Orjala, J., A.D. Wright, T. Rali, and O. Sticher.
565
1992. Three new prenylated benzoic acid Stevens, W.D., C. Uloa-U., A. Pool, O.M.
derivatives and molluscicidal sesquiterpenoids Montiel, eds. 2001. Flora de Nicaragua.
from Piper aduncum leaves. Medica 58(7): 714. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanical Garden Press. p. 1911-
Pacific Island Ecosystems at Risk. 2002. Invasive 2666.
plant specie, Piper aduncum L. Rutaceae.
http://www.hear.org/pier3/ piadu.htm. 2 p. Torres-Santos, E.C., V.C. Moura, G.M. Sperandio,
D.L. Moreira, M.A. Kaplan, and B. Rossi-
Philippine Council of Agriculture. 2002. Research Bergmann. 1996. Anti-leishmanial effect of a
highlights 1997. http://www.pcarrd.dost.gov.ph/ pure chalcone isolated from Piper aduncum
division/FERD/new/highlights97.html. 18 p. (Piperaceae). Memórias Vol. 91 (Supplement).
Instituto Oswaldo Cruz, Rio de Janeiro, RJ,
Plant Protection Service. 2001. Pest alert: false Brazil. 18 p.
kava. Pest Alert No. 19. http://www.spc.org.nc/
pps/PestAlerts/PestAlertNo19-False_kave.pdf. Vélez, I. and J. van Overbeek. 1950. Plantas
2 p. indeseables en los cultivos tropicales. Editorial
Universitaria, Río Piedras, PR. 497 p.
566
Piper amalago L. higuillo de limón
PIPERACEAE
John K. Francis
thickened in older plants. The alternate, dark
green, hairless leaves are 4 to 14 cm long and 2 to
7 cm broad. They are pointed at the tip, ovate to
elliptic, and have palmate venation with five main
veins. Tiny flowers are borne in compact, cord-
like, gray-green spikes 6 to 12 cm long. At
maturity, the small (1.5 mm) fruits (drupes) are
closely packed along the spike (Liogier 1985,
Little and others 1974).
567
Growth and Management.—In Puerto Rico, Interactions between crown structure and light
higuillo de limón reaches 5.5 m in height and 8 cm environment in five rain forest Piper species.
in diameter (Little and others 1974). The species American Journal of Botany 75(10): 1,459-
may become a tree to 15 m in high forests in 1,471.
Mexico, although it remains a shrub or small tree
in successional forests and gaps (Chazdon and Elizondo C., L.H. 2000. Artibeus toltecus
others 1988). Its stems have no visible annual Saussure. http://www.Inbio.ac.cr/bims/ubi/
rings and are therefore difficult to age. Judging mamiferos/ubiespejo/ubiid=1557&-find.html.
age by stem internodal segments, individual stems 5 p.
in Puerto Rico appear to reach 5 to 10 years old.
Because resprouting occurs from the roots, Fleming, T.H., E.R. Heithaus, and W.B. Sawyer.
individual plants may live much longer. Natural 1977. An experimental analysis of the food
regeneration of higuillo de limón occurs readily in location behavior of frugivorous bats. Ecology
early secondary forest and in open stands of older 58(3): 619-627.
forest when seed sources are present. Stands can
be easily manipulated to encourage it. Small Grieve, Mrs. M. 2001. A modern herbal: pepper.
quantities of seed have been separated by hand http;//www.botanical.com/botanical/mgmh/p/
with tweezers after mashing the fruit. Larger pepper24.html. 5 p.
quantities probably can be obtained by gentle
maceration and wet sieving. Germination on the Howard R.A. 1988. Flora of the Lesser Antilles,
surface of wet soil or peat is recommended. Leeward and Windward Islands.
Plantations could be established using Dicotyledoneae, Part 1. Vol. 4. Arnold
containerized seedlings, although no plantings Arboretum, Harvard University, Jamaica Plain,
have been reported. MA. 673 p.
Benefits.—Higuillo de limón is eaten by cattle Liogier H.A. 1985. Descriptive flora of Puerto
after the more desirable grasses and forbs are Rico and adjacent islands, Spermatophyta. Vol.
gone. Infusions of the leaves are said to alleviate 1. Editorial de la Universidad de Puerto Rico,
colic and intestinal gas. The roots are used as a Río Piedras, PR. 352 p.
diuretic and to treat water retention (Liogier
1990). The fully ripe fruits and their seeds have Liogier, H.A. 1990. Plantas medicinales de Puerto
the same taste as black pepper, Piper nigrum L., Rico y del Caribe. Iberoamericana de Ediciones,
and are sometimes used as a substitute for it, Inc., Río Piedras, PR. 566 p.
especially in Jamaica. The fruits are usually dried
whole and ground as needed (Grieve 2001). Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
References Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Chazdon, R.L., K. Williams, and C.B. Field. 1988. Washingtion, DC. 1,024 p.
568
Piper swartzianum (Miq.) C. DC. higuillo
PIPERACEAE
John K. Francis
569
management experience has been published. References
Natural reproduction could probably be
encouraged by disturbance that exposes mineral Liogier H.A. 1985. Descriptive flora of Puerto
soil under a moderately shady tree canopy. Rico and adjacent islands, Spermatophyta. Vol.
1. Editorial de la Universidad de Puerto Rico,
Benefits.—Higuillo contributes to the Río Piedras, PR. 352 p.
biodiversity, scenic beauty, and stability of the soil
of the forests where it grows. Like other pipers, its Little, E.L., Jr., R.O. Woodbury, and F.H.
fruits are food for fruit bats. Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
570
Piptocoma antillana Urban Antilles velvetshrub
ASTERACEAE
John K. Francis
571
Growth and Management.—Antilles velvetshrub for background and accent in naturalistic
grows 0.5 to 1.0 m per year. The stems do not live landscaping.
long--perhaps 2 to 5 years--but plants can renew
themselves and live longer by sprouting from the References
root crown. Planting and management experience
have not been published. Because the species is Liogier, H.A. 1997. Descriptive flora of Puerto
not abundant or aggressive, control should not be Rico and adjacent islands. Vol. 5. Editorial de la
necessary. Universidad de Puerto Rico, Río Piedras, PR.
436 p.
Benefits.—Antilles velvetshrub helps protect the
soil, furnishes cover for wildlife, and adds to the Liogier, H.A. and L.F. Martorell. 1982. Flora of
aesthetics of the forest. If means of propagation Puerto Rico and adjacent islands, a systematic
can be developed, the species would be valuable synopsis. Editorial de la Universidad de Puerto
Rico, Río Piedras, PR. 342 p.
572
Pisonia aculeata L. pull-back-and-hold
NYCTAGINACEAE
John K. Francis
species may be evergreen or deciduous depending
on the climate. The leaves are variable, ovate to
elliptical and 2.5 to 15 cm long by 1.5 to 6 cm
broad. They are light to dark green, leathery to
somewhat fleshy. The flowers are small axillary
cymes that are succeeded a few months later by 11
to 15 mm long fruits (anthocarps) with five single
or double length-wise rows of sticky glands
(Acevedo-Rodríguez 1985, Howard 1988, Liogier
1985).
573
g/fruit. Seeds extracted by hand, which have the References
appearance of grains of wild rice, averaged 0.041
+ 0.011 g/seed. It is not necessary to extract seed Acevedo-Rodríguez, P. 1985. Los bejucos de
because they germinate satisfactorily within the Puerto Rico. Vol. 1. General Technical Report
fruit. Seventy-five percent of seeds from the above SO-58. U.S. Department of Agriculture, Forest
Puerto Rican collection, sown as fruits in Service, Southern Forest Experiment Station,
commercial potting mix, germinated between 21 New Orleans, LA. 331 p.
and 49 days after sowing. Pull-back-and-hold is
susceptible to damping-off as seedlings in Enquist, B.J. and J.J. Sullivan. 2001. Vegetative
germination beds. Seedlings are ready to prick out key and descriptions of tree species of the
into containers about 1 month after germination. tropical dry forests of upland Sector Santa Rosa,
The sticky seeds can cling to birds while on the Area de Conservación, Guanacaste, Costa Rica.
trees; then, after they fall, still attached to the http://www.acguanacaste.ac.cr/paginas_ especie/
infrutescences, stick to the clothing or fur of plantae_online/EnquistSullivanTreeKey.pdf.
passing humans and animals. 68 p.
Growth and Management.—Although pull-back- Florida 4-H Foundation. 2001. Florida forest
and-hold has only a moderate growth rate, it can plants: Devil’s claws (Pisonia aculeata).
live for several decades and become quite large. http://www.sfrc.ufl.edu/4h/Devils_claw/
Plants may reach 20 m of extension (Acevedo- deviclaw.htm. 2 p.
Rodríguez 1985) and 20 cm of stem diameter in
Puerto Rico. It is sometimes desirable to remove Friend, W.H. 2001. Plants of ornamental value for
pull-back-and-hold from forest stands. This the Rio Grande Valley of Texas. http://
increases safety during timber harvest and helps www.aggiehorticulture.tamu.edu/plantanswers/
ensure that the regenerating stand will not be publications/riograndeornamentals.htm. 16 p.
suffocated by resurgent vines. A time-honored
method is to cut the vines hanging from trees 1 Howard R.A. 1988. Flora of the Lesser Antilles,
year before harvest and then to cut or spray the Leeward and Windward Islands.
few surviving sprouts just before harvest. Dicotyledoneae, Part 1. Vol. 4. Arnold
Arboretum, Harvard University, Jamaica Plain,
Benefits.—Pull-back-and-hold may be used as a MA. 673 p.
hedge or a foundation plant in landscaping, but it
requires regular pruning to keep it under control Liogier, H.A. 1985. Discriptive flora of Puerto
(Friend 2001) An occasional plant occurs entirely Rico and adjacent islands. Vol. 1. Editorial de la
free of spines. Perhaps these could be propagated Universidad de Puerto Rico, San Juan, PR.
vegetatively for ornamental use. The wood of wild 352 p.
pull-back-and-hold is sometimes used for fuel. It
is hard, heavy, and often available in good Múlgura, M.E. 2000. Catálogo de las plantas
diameters and considerable lengths. Root extracts vasculares de la República Argentina.
have been used in herbal medicine as a purgative, http://www.Darwin.edu.ar/Catologo/
and extracts from leaves and bark are used to treat nyctaginaceae.pdf. 4 p.
arthritis (Acevedo-Rodríguez 1985).
574
Pithecellobium unguis-cati (L.) Benth. bread-and-cheese
FABACEAE
John K. Francis
575
Growth and Management.—Plants started from installment. Research Note SO-374. Southern
seed grow relatively slowly at first and somewhat Forest Experiment Station, U.S. Department of
faster after deep rooting. Supplemental watering Agriculture, Forest Service, New Orleans, LA.
and fertilization are unnecessary. As a landscaping 5 p.
plant, it can be easily pruned and shaped
(Workman 1980). In the poor habitat where the Howard R.A. 1988. Flora of the Lesser Antilles,
species usually occurs in the wild, growth is very Leeward and Windward Islands.
slow and plants live for several decades. Dicotyledoneae, Part 1. Vol. 4. Arnold
Arboretum, Harvard University, Jamaica Plain,
Benefits.—Although it is difficult to harvest MA. 673 p.
because of the spines, the wood of bread-and-
cheese is used for fuel. It is heavy and hard and Island Resource Foundation. 2001. Threatened and
has an oven-dry heat of combustion of 19.05 endangered birds of the insular Caribbean:
megajoules/kg (Timyan 1996). The fleshy aril is Yellow-shouldered amazon, Amazona
edible, although not highly desirable because of barbadensis. http://www.irf.org/bbarden.htm.
the lingering flavor. The seeds of bread-and- 13 p.
cheese are used to make necklaces. The species is
planted to form impenetrable hedges (UVI Liogier H.A. 1988. Descriptive flora of Puerto
Wetlands Reserve 2001). Bread and cheese is Rico and adjacent islands, Spermatophyta. Vol.
listed as a nitrogen fixing species (Winrock 2. Editorial de la Universidad de Puerto Rico,
International 2001). The fruits are one of the food Río Piedras, PR. 481 p.
plants of the endangered yellow-shouldered
amazon parrot (Amazona barbadensis) in Little, E.L., Jr., R.O. Woodbury, and F.H.
Venezuela, and the islands of Margarita, La Wadsworth. 1974. Trees of Puerto Rico and the
Blanquilla, and Bonaire (Island Resource Virgin Islands. Vol. 2. Agriculture Handbook
Foundation 2001). The large orange sulfur 449. U.S. Department of Agriculture,
(Phoebis agarithe Boisduval) and the Miami blue Washington, DC. 1024 p.
(Hemiargus thomasi Clench) butterflies both use
bread-and-cheese as rearing plants for their larva Timyan, J. 1996. Bwa yo: important trees of Haiti.
(Association of Florida Native Nurseries 2001). A South-East Consortium for International
small treehopper insect (Homoptera) has Development, Washington, DC. 418 p.
developed an extended thorax that mimics a
bread-and-cheese thorn and serves as protective UVI Wetlands Reserve. 2001. A list of plants
camouflage (Workman 1980). tagged along the trail at the Reserve, and some
of their uses. http://rps.uvi.edu/
References VIMAS/plantlist.htm. 5 p.
Association of Florida Native Nurseries. 2001. West, E. and L.E. Arnold. 1952. The native trees
Florida gardening with native plants—Legume. of Florida. University of Florida Press,
http://www.nsis.org/garden/family/legume.html. Gainesville, FL. 212 p.
4 p.
Winrock International. 2001. Nitrogen fixing trees
The Champion Tree Project. 2001. National and shrubs. http.//www.winrock.org
champion trees, Florida. /forestry/factpub/nftlist /htm. 25 p.
http://www.championtrees.org/database/champi
onsFL.htm. 8 p. Workman, R.W. 1980. Growing Native. The
Sanibel-Captiva Conservation Foundation, Inc.
Francis, J.K. and A. Rodríguez. 1993. Seeds of 137 p.
Puerto Rican trees and shrubs: second
576
Pluchea carolinensis (Jacq.) G. Don cure-for-all
ASTERACEAE
John K. Francis
577
and summer in Florida (Long and Lakela 1976) P. Lohr, and D. Atha. 2000. Medical plants used
and produces seeds prolifically. A collection of by Latino healers for woman’s health conditions
seeds from Puerto Rico weighed an average of in New York City. Economic Botany 54(3):
0.000025 g/seed or 40 million seeds/kg. Because 344-357.
the seeds failed to germinate on moist filter paper,
it is not known whether this represents a Gavilán-Yodú, R. and P.N. Hechavarría. 2002.
reasonable estimate for the species (author’s Efecto analgésico y antiinflamatorio de la tintura
observation). The seeds are wind-dispersed and de salvia (Pluchea carolinensis) al 30 % en ratas
probably require wet, bare soil to germinate and Wistar. Instituto Superior de Ciencias Médicas,
establish themselves. La Habana, Cuba. http://fcmfajardo.sld.cu/
jornada/trabajos/salvia/resumen/htm. 5 p.
Growth and Management.—In Puerto Rico,
most cure-for-all live for 2 to 4 years before dying Howard, R. A. 1989. Flora of the Lesser Antilles,
or dying back to the root and resprouting. Sprouts Leeward and Windward Islands. Vol. 6. Arnold
grow about 1.5 m in the first year. Establishment Arboretum, Harvard University, Jamaica Plain,
of new plants can probably be assured by MA. 658 p.
scarifying the soil before the wet season in the
presence of a seed source. Where it is necessary to Liogier, H.A. 1990. Plantas medicinales de Puerto
eliminate cure-for-all, grubbing out the plants or Rico y del Caribe. Iberoamericana de Ediciones,
spraying with broadleaf herbicides is Inc., San Juan, PR. 566 p.
recommended until tested treatments are available.
A seed insect, Acinia picturata (Diptera: Liogier, H.A. 1997. Descriptive flora of Puerto
Tephritidae), was introduced in Hawaii and Rico and adjacent islands. Vol. 5. Editorial de la
although now well established, has had no Universidad de Puerto Rico, Río Piedras, PR.
significant effect on the shrub (Alyokhin and 436 p.
others 2001).
Long, R.W. and O. Lakela. 1976. A flora of
Benefits.—Cure-for-all has an excellent ability to Tropical Florida. Banyan Books, Miami, FL.
colonize and stabilize disturbed areas and act as a 962 p.
nurse crop for later-successional species. Although
not specifically reported, as other members of the Natural Resources Conservation Service. 2002.
family, it probably provides a source of nectar and Plant profile: Pluchea carolinensis (Jacq.) G.
pollen for honeybees and other insects. It has Don. http://plants.usda.gov/cgi_bin/plant_profile
several herbal applications including aromatic .cgi?symbol=PLCA10. 3 p.
baths, control of fever, treatment of uterine
fibroids, relief of sore throat and stomach pain, Nelson, G. 1996. The shrubs and woody vines of
poultices for wounds and skin ulcers, as an Florida. Pineapple Press, Inc., Sarasota, FL.
analgesic, and for the treatment of malaria (Balick 391 p.
and others 2000, Liogier 1990, Vélez and van
Overbeek 1950). Analgesic and antiinflammatory Pacific Island Ecosystems at Risk. 2002. Invasive
effects have been demonstrated in laboratory trials plant species: Pluchea carolinensis (Jacq.) G.
with rats (Gavilán-Yodú and Hechavarría 2002). Don, Asteraceae. http://hear.org/pier3/plcar.htm.
2 p.
References
Peng, C.-I., C.-H. Chen, W.-P. Leu, and H.-F.
Alyokhin, A.V., R.H. Messing, and J.J. Duan. Yen. 1998. Pluchea Cass. (Asteraceae: Inuleae)
2001. Utilization of the exotic weed Pluchea in Taiwan. Botanical Bulletin of Academia
odorata (Asteraceae) and related plants by the Sinica 39(4): 287-297.
introduced biological control agent Acinia
picturata (Diptera: Tephritidae) in Hawaii. Smith, C.W. and J.T. Tunison. 1992. Fire and alien
Biocontrol Science and Technology 11: 711- plants in Hawai’i: Research and management for
718. native ecosystems. In: C.P. Stone, C.W. Smith,
and J.T. Tunison, eds. Alien plant invasions in
Balick, M.J., F. Kronenberg, A.L. Ososki, M. native ecosystems of Hawai’i: Management and
Reiff, A. Fugh-Berman, B. O’Connor, M. Roble, Research. University of Hawaii Cooperative
578
National Park Resources Studies Unit, University of Hawaii Botany. 2002. Alien plants
Honolulu, HI. p. 394-409. of Hawaii: Pluchea symphytifolia (Mill.) Gillis.
http://www.botany.hawaii.edu/faculty/cw_smith
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. /plu_sym.htm. 1 p.
Montiel, eds. 2001. Flora de Nicaragua.
Monographs in Systematic Botany. Vol. 85, No. Vélez, I. and J. van Overbeek. 1950. Plantas
1. Missouri Botanical Garden Press, St. Louis, indeseables en los cultivos tropicales. Editorial
MO. 943 p. Universitaria, Río Piedras, PR. 497 p.
579
Prosopis glandulosa Torr. honey mesquite
FABACEAE
580
Scarified seeds germinate readily and can remain as Peniocereus greggii Britt. and Rose (Tull 1987,
undigested and viable as they pass through animal author’s personal observation). It is not clear
digestive tracts (Wright 1982). The germination whether this is due to a favorable microclimate or
rate typically ranges from 75 to 90 percent (Vines a response to grazing pressures. Seeds of mesquite
1986). Seeds frequently contain weevil larvae are nutritionally rich and are important food for a
(Vines 1986). The pollen is reported responsible large number of wildlife species (Graham 1941,
for some hay fever (Kearney and others 1960) Tull 1987). Honey mesquite seeds may constitute
implying that it is also disseminated by wind. 5 to 37 percent of scaled quails' diets year round
(Davis and others 1975). Deer, javelina, and
Growth and Management.—Since livestock smaller wildlife species such as jackrabbits, feed
grazing was introduced in the Southwestern United on both pods and vegetation. It is important
States and Northern Mexico, honey mesquite has livestock feed, especially in times of drought.
increased to such an extent as to be considered a However, if livestock eat too large amounts,
pest. However, mesquite is a valuable soil binder rumen stasis, impaction, and death may result
and slows erosion of otherwise denuded land. The (Stubbendieck and others 1993). It is an important
increase in honey mesquite has been attributed to a “honey plant” and bees that forage its flowers
variety of factors. The two most often cited are a produce excellent quality honey (Dayton 1931). It
reduction in perennial grasses, which when healthy provides a good source of nectar and food for
and dense reduces mesquite seedling butterfly adults and larvae (Taylor and others
establishment, and the reduced incidence of fire 1997). The wood is resistant to termites and decay,
due to livestock over-grazing and fire suppression and is used for building. It is also prized as bar-b-
(Bahre 1995, Humphrey 1958, Fisher 1977). Other cue fuel as it is a hard, close-grained wood that
reasons include: reduced effectiveness of burns hot and slowly (Barneby 1989, CWAR
invertebrate seed predators, increased 2002). Honey mesquite was essential to Native
dissemination of seed by livestock and/or Americans who made breads from the pods, fuel
kangaroo rats, land clearing and cultivation, soil and utensils from the wood, medicines from the
compaction by livestock that hinders grass stems and leaves, rope from the roots, and dye
establishment, decreased activity of jackrabbits from the black pitch of the trunks (Moerman 1998,
and wood rats, and climate change (Bahre 1995, Tull 1987). The gum exudates of the bark may be
Glendening 1952). Depending upon the situation, a substitute for gum arabic and have been used to
herbicide and mechanical treatments are often both mend pottery (Powell 1998). An intoxicating
economical and effective in controlling scattered beverage is made from fermented meal (Bowers
and dense stands of mesquite (Schmutz and others 1993).
1992). Chaining and burning the trees destroy
above-round tissue, but old stumps of honey References
mesquite readily resprout. Burning is effective
only on small mesquite plants and where fuel is Bahre, C.J. 1995. Human impacts on the
adequate to carry a fire (Schmutz and others 1992). grasslands of southeastern Arizona In: M.P.
McClaran and T.R. Van Devender, eds. The
Benefits.—Honey mesquite plants bind soils, and Desert Grassland. The University of Arizona
plantings are recommended for erosion control Press, Tucson, AZ. p. 230-264.
(Graham 1941, Rorabaugh 1995). Mesquite,
particularly var. torreyana, is tolerant of saline Bainbridge, D.A., R.A. Virginia, and W.M. Jarrell.
soils and is a particularly useful cover in degraded 1990. Honey Mesquite. Forest, Farm, and
areas (Rorabaugh 1995, Schmutz and others 1992). Community Tree Network (FACT Net)
It has been used, with limited success, in http://www.winrock.org/forestry/factnet.htm.
revegetating copper mine wastes (Norem and
others 1982). Mesquite has been used in riparian Barneby, R.C. 1989. Fabales. Intermountain Flora,
restoration projects and provides excellent habitat Vascular Plants of the Intermountain West,
for many species of wildlife and birds (Rorabaugh U.S.A. Vol. 3, Part B. New York Botanical
1995). In one study, the number of quail using an Garden, Bronx, NY. 279 p.
area decreased as a consequence of mesquite
removal (Goodwin and Hungerford 1977). Honey Bowers, J.E. 1993. Shrubs and trees of the
mesquite also provides shelter for the germination Southwest Deserts. Southwest Parks and
and development of forbs, grasses, and cacti such Monuments Assoc. Tucson, AZ. 140 p.
581
Center for Wood Anatomy Research. 2002. U.S. Kearney, T.H., R.H. Peebles, and collaborators.
Department of Agriculture. Forest Service. 1960. Arizona flora. 2nd ed. University of
Tech. Fact Sheet. Prosopis. http://www2.fpl. California Press, Berkeley, CA. 1085 p.
fs.fed.us/TechSheets/HardwoodNA/htmlDocs/
prosop1.html. Moerman, D.E. 1998. Native American Ethno-
botany. Timber Press, Portland, OR. 927 p.
Davis, C.A., R.C. Barkley, and W.C. Haussamen.
1975. Scaled quail foods in southeastern New Norem, M.A., A.D. Day, and KL. Ludeke. 1982.
Mexico. Journal of Wildlife Management An evaluation of shrub and tree species used in
39(3): 496-502. revegetating copper mine wastes in the United
States. Journal of Arid Environments 5: 299-
Dayton, W.A. 1931. Important western browse 304.
plants. U.S. Department of Agriculture. Misc.
Publ. 101. Washington, DC. 214 p. Powell, A.M. 1998. Trees and shrubs of Trans-
Pecos and adjacent areas. University of Texas
Dick-Peddie, W.A. 1993. New Mexico Vegetation Press, Austin, TX. 498 p.
– past, present, and future. University of New
Mexico Press, Albuquerque, NM. 244 p. Rorabaugh, J.C. 1995. A superior accession of
western honey mesquite (var. torreyana) for
Everitt, J.H. and D.L. Drawe. 1993. Trees, Shrubs riparian restoration projects. Desert Plants.
and Cacti of South Texas. Texas Tech 11(4) 32-40
University Press, Lubbock, TX. 213 p.
Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L.
Fisher, CE. 1977. Mesquite and modern man in Cox, J.O. Klemmedson, J.J. Norris, and L.C.
southwestern North America. In: B.B. Simpson, Fierro. 1992. Desert Grassland. In: R.T.
ed. Mesquite, its biology in two desert shrub Coupland ed. Ecosystems of the World. Vol.
ecosystems. Downden, Hutchinson, and Ross, 8A. Elsevier, London and New York. 469 p.
Stroudsburg, PA. p 17-88.
Smiens, F.E., D.D. Diamond, and C.W. Hanselka.
Glendening, G.E. 1952. Some quantitative data on 1992. Coastal Prairie. In: R.T. Coupland, ed.
the increase of mesquite and cactus on a desert Ecosystems of the World. Vol. 8AElsevier,
grassland range in southern Arizona. Ecology. London and New York. 469 p.
33 (3): 319-328.
Sohmer, S.H. and R. Gustafson. 1987. Plants and
Goodwin, J.G. and C.R. Hungerford. 1977. Habitat flowers of Hawaii. University of Hawaii Press,
use by native Gambel’s and scaled quail and Honolulu, HI. 160 p.
released masked bobwhite quail in southern
Arizona. Research Paper RM-197. USDA Stubbendieck, J., S.L. Hatch, and C.H. Butterfield.
Forest Service, Rocky Mountain Research 1993. North American Range Plants. 4th ed.
Station, Fort Collins, CO. 8 p. Univ. of Nebraska Press, Lincoln, NB. 493 p.
Graham, E.H. 1941. Legumes for erosion control Taylor, R.B., J. Zrutledge, and J.G. Herrera. 1997.
and wildlife. U.S. Department of Agriculture A field guide to common south Texas shrubs.
Misc. Pub. 412. Washington D.C. 153 p. Texas Parks and Wildlife Press, Austin, TX.
p. 106.
Humphrey, R.R. 1958. The desert grassland: a
history of vegetational changes and an analysis Tull, D. 1987. Edible and Useful Plants of Texas
of causes. Botanical Review 24: 193-252. and the Southwest. University of Texas Press.
Austin, TX. 518 p.
Kartesz, J.T. 1994. A synonymized checklist of the
vascular flora of the United States, Canada and Uchytil, R.J. 1990. Prosopis glandulosa var.
Greenland. Vol.1 - Checklist. 2nd ed. Timber glandulosa. U.S.D.A. Forest Service, Rocky
Press, Portland, OR. 622 p. Mountain Research Station, Fire Sciences
Laboratory Fire Effects Information System,
http:// www.fs.fed. us/databse/feis/.
582
United States Department of Agriculture Forest Wright, R.A. 1982. Aspects of desertification in
Service. 1988. Range Plant Handbook. Dover Prosopis dunelands of southern New Mexico.
Publications, Inc. New York. 837 p. Journal of Arid Environments 5: 277-284.
583
Proustia vanillosma C. Wright sweet yellowcrown
ASTERACEAE
John K. Francis
584
Universidad de Puerto Rico, San Juan, PR. cgi_bin/plant_profile.cgi?symbol=PRVA. [not
436 p. paged].
Natural Resources Conservation Service. 2003. Turner, B.L. 1993. Berylsimpsonia (Compositae:
Plants profile: Proustia vanillosma C. Wright, Mutiseae), a new genus for the Greater Antilles.
sweet yellowcrown. http://plants.usda.gov/ Phytologia 74: 349-355.
585
Prunus americana Marsh. American plum
ROSACEAE
John K. Francis
586
reported 1,918 cleaned seeds/kg, and that 60 of the plants to domestic ruminants and wild game
percent germinated after cold stratification. animals varies from good to poor depending on the
Germination is hypogeal (Young and Young animal species and the location (Forest Service
1992). Seeds are dispersed by mammals and birds. 2003). The cover provided by American plum
Animals that ingest the seeds, such as black bear thickets is important to many wild animals. The
(Ursus americanus), are more effective dispersers fruits are eaten by a number of species including
than those that do not (Forest Service 2003). Seeds bluejays (Cyanocitta cristata), brown thrashers
ingested by coyotes (Canis latrans) had a (Toxostoma rufum), mockingbirds (Mimus
significantly lower germination rate than uneaten polyglottos), red-headed woodpeckers
seeds (Cypher and Cypher 1999). Seeds may (Melanerpes erythrocephalus), bobwhite quail
remain in the soil seed bank for many years until (Colinus virginianus), white-tailed deer
disturbance creates conditions for growth (Odocoileus virginianus), raccoons (Procyon
(Treeguide 2003). After dispersed plants become lotor), squirrels (Sciurus spp.), and coyotes (Canis
well established, they begin forming clonal latrans) (Cowley 2003, Cypher and Cypher 1999,
thickets. Suckers may appear as much as 3 m away Kaiser 2001). The flowers furnish nectar food for
from the parent plants (Colorado Springs Utilities great purple hairstreak [Atlides halesus (Cramer)]
2003). and Sweadner’s Jupiter hairstreak [Callophrys
gryreus (Hübner)] butterflies (Cowley 2003).
Growth and Management.—American plum is Fruits are eaten raw or cooked, made into jams and
only capable of a moderate growth rate, usually jellies, and were dried for winter food in former
less than 30 cm in height per year (Michigan State times by Native Americans and pioneers. Several
University Extension 2003). Minimum seed- horticultural varieties of plum have been derived
bearing age is 4 years (Grisez and others 2003). from American plum. It has also been used in
Individual stems rarely live longer than 20 years hybrid crosses with other species of plums
(Treeguide 2003). However, clones may last much (Bircher and Bircher 2000). The wood, which has
longer. American plum fruits should be collected a specific gravity of 0.73 (Britton and Shafer
when fully mature for best quality seed. This can 1908), makes excellent firewood except for being
be done by hand stripping or shaking or beating crooked and limby. American plum is used in
onto a tarp spread under the shrub or tree. The amenity planting for wildlife and in revegetation
seeds should be cleaned of all pulp by macerating projects.
and washing. The seeds are usually 96 to 100
percent filled. Seeds of this species can be stored References
at room temperature for up to 30 months without
loss of viability. Seeds to be stored for longer Bircher, A.B. and W.H. Bircher. 2000.
should be surface dried and placed in a sealed Encyclopedia of fruit trees and edible flowering
container at 1 to 5 °C. Stratification of 90 to 150 plants in Egypt and the Subtropics. The
days at 2 to 5 °C before planting is recommended American University in Cairo Press, Cairo,
for after-ripening of the seed. Alternately, seed Egypt. 568 p.
may be sown in nursery beds 2.5 to 5 cm deep in
the fall. Seedlings are ready for lifting (bare-root) Britton, N.L. and J.A. Shafer. 1908. North
at the end of one growing season (Grisez and American trees. Henry Holt and Company, New
others 2003). Plantings may be made with bare- York. 964 p.
root stock and potted nursery seedlings. Direct
seeding in the fall with unstratified seed or spring Colorado Springs Utilities. 2003. Plant details.
with stratified seed in prepared seedspots will http://www.csu.org.cgi-bin/xeri/Xeriinclude?
normally yield new seedlings. The species can be Xeridetail?PIS-pra. 2 p.
grafted or used as root stock, and it has been
successfully propagated from stem cuttings (Forest Cowley, M. 2003. Rose family (Rosaceae).
Service 2003). http://www.nsis.org/garden/family/rose.html.
4 p.
Benefits.—American plum is an early spring
bloomer and with its white flowers beautifies the Cypher, B.L. and E.A. Cypher. 1999. Germination
forests and prairies for a week or more each year. rates of tree seeds ingested by coyotes and
The species also helps protect the soil and provides raccoons. American Midland Naturalist 142(1):
benefits to wildlife and humans. The browse value 71-76.
587
Forest Service. 2003. Fire effects information Natural Resources Conservation Service. 2003.
system: species: Prunus americana. http://fs. Prunus americana Marsh. http://plants.usda.
fed.us/database/feis/plants/tree/pruame/all.html. gov/cgi_bin/plant_profile.chi?symbol=PRAM.
14 p. 4 p.
Grisez, T.J. 1974. Prunus L. cherry, peach, and Oklahoma Biological Survey. 1999. Prunus
plum. In: C.S. Schopmeyer, tech. coord. Seeds americana Marsh. http://www.biosurvey.ou.
of woody plants in the United States. U.S. edu/shrub/prun-ame.htm. 2 p.
Department of Agriculture, Forest Service,
Washington, DC. p. 658-673. Sargent, C.S. 1923. Manual of the trees of North
America (exclusive of Mexico). Houghton
Grisez, T.J., J.R. Barbour, and R.P. Karrfalt. 2003. Mifflin, Boston, MA. 910 p.
Prunus L. cherry, peach, and plum. In: F.T.
Bonner and R.G. Nisley, eds. Woody plant seed Treeguide. 2003. American plum, Prunus
manual. http://wpsm.net/Prunus.pdf. [not americana Marsh. http://www.treeguide.com/
paged]. Species.asp?SpeciesID=770. 3 p.
Kaiser, J. 2001. 2001 featured plant, American Young, J.A. and C.G. Young. 1992. Seeds of
plum, Prunus americana. In: Plants for woody plants in North America. Dioscorides
conservation, Vol. 4, No. 1. Elsberry Plant Press, Portland, OR. 407 p.
Materials Center, Elsberry, MO. p. 1.
588
Prunus emarginata (Dougl. ex Hook.) D. Dietr. bitter cherry
ROSACEAE
Synonyms: None
Christopher Ross
General Description.—Bitter cherry ranges from Growth and Management.—Bitter cherry has
a shrub to a tree of 10 m. It often forms dense been effectively controlled on conifer planting
thickets. Clustered, deciduous elliptic to obovate, sites by application of the herbicide 2,4,5-T (now
crenate-serrate leaves 20 to 65 mm in length are on banned in the United States) (Bock and others
3 to 12 mm petioles (Wilken 1993, Mozingo 1978). It may be heavily attacked by tent
1987). The leaves, stems, and flowers all have a caterpillars and other insects at times.
very strong cyanide odor (bitter almond) when
crushed. Benefits.—Blue grouse and other birds are fond of
the fruits of bitter cherry. They are also a preferred
Range.—Bitter cherry is found in cooler parts of summer and fall food of black bears (Unsworth
Arizona, California, far western Nevada, Idaho, and others 1989). Leaves are browsed by sheep
Utah, New Mexico and Montana and north to and cattle, despite their cyanogenic properties,
British Columbia (Mozingo 1987, Esser 1995). which have caused reported stock poisonings
(Mozingo 1987). In the Pacific Northwest, bitter
Ecology.—Bitter cherry site dominance may not cherry is eaten by deer and elk (Crouch 1968),
peak until three decades after disturbance, although it is not a preferred forage (Leege 1979,
suggesting that it is not an obligate seed bank Klebenow 1965). Native Americans used long
species (Oakley and Franklin 2001). However, in fibrous strips of bark for twine and basketry, and
some cases following fire it may quickly dominate used the roots for a variety of medicinal purposes
a site (personal. observation.). However, others but made little use of the bitter fruit (Turner and
(Morgan and Neuenschwander 1988) have Bell 1973). Bitter cherry essence is marked by
considered it an obligate seed bank species. It may herbalists, who claim that it aids patience and
achieve major dominance of disturbed sites, which short attention spans. Bitter cherry has also been
589
used for mine and highway reclamation (Everett Morgan, P. and L.F. Neuenschwander. 1988.
and others 1980 and personal observation). Seed-bank contributions to regeneration of shrub
species after clear-cutting and burning.
References Canadian Journal of Botany 66:169-72.
Bock, J.H., M. Raphael, and C.E. Bock. 1978. A Mozingo, H. N. 1987. Shrubs of the Great Basin.
comparison of planting and natural succession University of Nevada Press, Reno Nevada. 342
after a forest fire in the northern Sierra Nevada. p.
Journal of Applied Ecology 15:597-602.
Oakley, B.B. and J. F. Franklin. 2001. Bitter
Crouch, G.L. 1968. Forage availability in relation cherry (Prunus emarginata) distribution,
to browsing of Douglas-fir seedlings by black- successional dynamics, and implications for the
tailed deer. Journal of Wildlife Management 32: role of the seed bank.
542-553. http://www.cisti.nrc.ca/cisti/journals/cjb/b98-
162.html. p. 1725-1732.
Esser, L.L. 1995. U.S. Department of Agriculture,
Forest Service, Rocky Mountain Research Turner, N.C. and M.A.M. Bell. 1973. The
Station, Fire Sciences Laboratory (2001, Ethnobotany of the Southern Kwakiutl Indians
October). Fire Effects Information System, of British Columbia. Economic Botany 27: 257-
http://www.fs.fed.ss/database/feis/. [not paged]. 310.
Everett, R.L., R.O. Meeuwig, and R. I. Butterfield. Unsworth, J.W., J.J. Beecham, and L.R. Irby.
1980. Revegetation of untreated acid spoils at 1989. Female black bear habitat use in West-
Leviathan mine, Alpine County, California. central Idaho. Journal of Wildlife Management
California Geology 32: 8-10. 53: 668-673.
590
Prunus pumila L. sand cherry
ROSACEAE
Don C. Bragg
592
References Gysel, L.W. 1966. Ecology of a red pine (Pinus
resinosa) plantation in Michigan. Ecology 47:
Arkansas Department of Planning. 1974. Arkansas 465-472.
natural areas plan. Arkansas Department of
Planning, Little Rock, AR. 248 p. Lamson-Scribner, F. 1891. A sketch of the flora of
Orono, ME. Botanical Gazette 16: 228-234.
Billington, C. 1943. Shrubs of Michigan. Bulletin
20. Cranbrook Institute of Science, Bloomfield McAtee, W.L. 1920. Notes on the jack pine plains
Hills, MI. 249 p. of Michigan. Bulletin of the Torrey Botanical
Club 47: 187-190.
BONAP. 1999. Synthesis of the North American
flora, version 1.0 (CD). North Carolina Olson, J.S. 1958. Rates of succession and soil
Botanical Garden, University of North changes on southern Lake Michigan sand
Carolina, Chapel Hill, NC. dunes. Botanical Gazette 119: 125-170.
Cowles, H.C. 1899. The ecological relations of the Rehder, A. 1958. Manual of cultivated trees and
vegetation on the sand dunes of Lake shrubs, 2nd ed. The Macmillan Co., New York.
Michigan. Botanical Gazette 27: 167-202. 996 p.
Cusick, A.W. 1985. Prunus pumila var. pumila L. Robertson, G.P. and P.M. Vitousek. 1981.
(Great Lakes Sand Cherry). Abstract. Ohio Nitrification potentials in primary and
Dept. of Natural Resources Division of Natural secondary succession. Ecology 62: 376-386.
Areas and Preserves. http://www.dnr.
state.oh.us/ODNR/dnap/Abstracts/P/prupumi. Smith, E.B. 1988. An atlas and annotated list of
htm. 2 p. the vascular plants of Arkansas, 2nd ed.
Department of Botany and Microbiology,
Drayton, B. and R.B. Primack. 1996. Plant species University of Arkansas, Fayetteville, AR.
lost in an isolated conservation area in 489 p.
metropolitan Boston from 1894 to 1993.
Conservation Biology 10: 30-39. Stevens, O.A. 1961. Plants of Fargo, North
Dakota. American Midland Naturalist 66: 171-
Emmitt, D.P. and A.W. Cusick. 1983. Prunus 177.
pumila var. cuneata Willd. (Sand Cherry).
Abstract. Ohio Dept. of Natural Resources Stewart, M.M. and J. Rossi. 1981. The Albany
Division of Natural Areas and Preserves. Pine Bush: a northern outpost for southern
http://www.dnr.state.oh.us/ODNR/dnap/ species of amphibians and reptiles in New
Abstracts/P/prupumi2.htm. 2 p. York. American Midland Naturalist 106: 282-
292.
Fernald, M.L. 1923. The identities of the sand
cherries of eastern America. Rhodora 25: 69- Talbot, M. 1934. Distribution of ant species in the
74. Chicago region with reference to ecological
factors and physiological toleration. Ecology
Fernald, M.L. 1950. Gray’s manual of botany, 8th 15: 416-439.
ed. American Book Co., New York. 1,632 p.
Voss, E.G. 1954. The butterflies of Emmet and
Gleason, H.A. 1952. Illustrated flora of the Cheboygan Counties, Michigan with other
Northeastern United States and adjacent notes on northern Michigan butterflies.
Canada, Vol. 2. Lancaster Press, Inc., American Midland Naturalist 51: 87-104.
Lancaster, PA. 655 p.
Walp, R.L. 1935. Shrubs of Cheboygan and
Emmet Counties, Michigan. American
Midland Naturalist 16: 230-247.
593
Prunus virginiana L. chokecherry
ROSACEAE
Bruce L. Welch
594
Ecology.—Usually chokecherry can be found excessive livestock grazing has damaged some
growing in canyon bottoms, sheltered slopes, populations in many areas of the northern Great
along streams and roads. It grows in a wide range Plains. Hydrogen glycoside prunasin is a toxin
of soils, ranging from Entisols to Mollisols that produced by chokecherry that could be poisonous
have textures ranging from sandy loams to clays. It to grazing livestock. Highest levels (5 percent) are
can be found from 177 (Michigan) to 3,100 (Utah) found in new stems and leaves. These levels
m in elevation where the combinations of soil and diminish over the growing season to 1.2 to 2.2
topography permit greater than average percent.
accumulation of moisture (Pacific Southwest
Experiment Station. 2002). Weakly salty soils are Benefits.—Chokecherry adds to the biodiversity
tolerated by chokecherry, but it cannot tolerant of a multitude of ecosystems. It provides habitat
soils that are poorly drained or suffer from and food for a number of wildlife species, and
prolonged flooding. It grows in soil pH ranges watershed protection. Fruits, leaves, or twigs are
from 3.5 to 7.6. It is found in numerous habitat and eaten by bears, moose, bighorn sheep, pronghorn,
plant associations that range from post disturbance elk and deer. A number of small of mammals also
invaders to early successional to climax or stable consume chokecherry including coyotes,
(Pacific Southwest Experiment Station 2002). snowshoe hares, red foxes, bobcats, raccoons, and
Chokecherry is intolerant to intermediate in porcupines. Chokecherry fruits are eaten by many
tolerance of shade and resprouts from root crowns birds including robins, western, eastern, and
and rhizomes readily, thus giving it the ability to mountain bluebirds, European starlings, and
persists under open or forest canopies of moderate Columbian sharp-tailed grouse. Livestock also eat
densities. It is well adapted to disturbance by fire. chokecherry. Winter twigs contain 38.9 percent
Chokecherry is a primary host of the eastern tent total digestible nutrients, which is lower than a
caterpillar and a fungus Plowrightia stansburiana, number of winter shrubs (Dietz 1972, Welch
which causes black knot-like tumors on stems 1981.) Winter protein, calcium, and phosphorus
(Pacific Southwest Experiment Station 2002). levels are above average. Fruits are harvested to
Western X virus can kill entire stands of make wines, syrups, jellies, and jams. Chokecherry
chokecherry. plants are planted as ornamentals, for enhancing
backyard wildlife habitats, and as windbreaks.
Reproduction.—Chokecherry is among the first
of the deciduous woody shrubs to leaf out and to References
flower in the spring. It can reproduce sexually
from seeds and asexually from root crowns and Auger, J., S.E. Meyer, and H.L. Black. 2002. Are
rhizomes. Seeds are encased in a stony endocarp American black bears (Ursus americanus)
and have an after-ripening requirement for legitimate seed dispersers of fleshy-fruited
germination. Heat treatment improves shrubs?American Midland Naturalist. 147:352-
germination. Seeds weigh about 0.095 grams 367.
(Grisez 1974). Passing through digestive systems
of mammals and birds may enhance germination Cronquist, A., N.H. Holmgren, and P.K. Holmgren
(Auger and others 2002). Also a number of birds 1997. Intermountain flora: vascular plants of the
and mammals may be responsible for long Intermountain west, U.S.A. Vol. 3. Part A;
distance dispersal (Auger and others 2002). Subclass Rosidae (except Fabales). The New
Chokecherry seeds persists in the soil seedbank . York Botanical Garden, New York. 446 p.
596
Psidium guajava L. guava
MYRTACEAE
John K. Francis
597
Mediterranean fruit flies (Ceratitis capitata Wied.) The fruit rinds are candied or stewed in syrup.
(Popenoe 1948). This thin-barked species is easily Nearly all the commercial production comes from
top-killed by fire and is sensitive to frost (von improved varieties with large fruits and few seeds.
Carlowitz 1991). The ripe fruits contain the following components:
water 84 percent, ash 0.7 percent, protein 0.8
Reproduction.—Guava flowers and fruits nearly percent, fiber 5.6 percent, total sugars 5.4 percent,
throughout the year (Little and Wadsworth 1964). starch 2.5 percent, and fat 1.0 percent (Popenoe
Many individual plants bear just once per year but 1948). The fruit pulp is rich in vitamins A and C
not necessarily synchronized with other guava (Morton 1987). Guava wood is used for tool
plants in the area. Honey bees (Apis mellifera) are handles, carving, and fuel (Advisory Committee
the chief pollinators (Morton 1987). Time from on Technology Innovation 1983). In folk
flowers to ripe fruits ranges from 102 to 124 days medicine, extracts of roots, bark, and leaves are
(Samson 1986). Thirty-one ripe fruits collected used to treat gastroenteritis, vomiting, diarrhea,
from wild plants in Puerto Rico averaged 32.8 g dysentery, wounds, ulcers, toothache, coughs, sore
and ranged from 13.5 to 61.8 g. Air-dried seeds throat, inflamed gums, and a number of other
averaged 0.0079 + 0.0001 g/seed or 127,000 conditions (Morton 1987). Guava leaf tea is
seeds/kg. Sown on moist peat, 63 percent widely used to control blood sugar of diabetics in
germinated between 15 and 60 days of sowing. Japan and elsewhere. It has been shown to be
Germination was epigeal (author’s observation). effective in vitro, in mice, and in human
Birds and mammals disperse the seeds (Invasive volunteers (Deguchi and others 1998). The basis
Species Specialist Group 2002). Root cuttings are for herbal treatment of diarrhea was established by
often used for commercial propagation of demonstrating inhibition of eight bacteria species
improved varieties (Morton 1987). Branches and and amoebas, and antispasmodic activity (Tona
stems layer (root) when they come in contact with and others 1999).
moist soil (author’s observation), and suckers also
arise from roots near the trunk. Established plants References
coppice readily and withstand repeated cutting
(Advisory Committee on Technology Innovation Advisory Committee on Technology Innovation
1983). 1983). Firewood crops. Vol. 2. National
Academy of Sciences, National Academy Press,
Growth and Management.—Guava seedlings Washington, D.C. 92 p.
grow at a moderate rate, but older plants grow
more slowly. Under good conditions, guava begins Deguchi, Y., K. Osada, K. Uchida, H. Kimura, M.
bearing fruits in 3 or 4 years. Plants live 40 years Yoshikawa, T. Kudo, H. Yasui, and M.
or more (Popenoe 1948). It is probably unwise and Watanuki. 1998. Effects of extract of guava
unnecessary to plant wild guava because the leaves on the development of diabetes in the
species is tough and aggressive and often invades db/db mouse and on the postprandial blood
agricultural (especially cattle pasture) and forest glucose of human subjects. Nippon
lands. Control is sometimes attempted, especially Nogeikagaku Kaishi 72(8): 923-931.
in cattle pastures and plantations. Sheep and goats
graze the leaves and strip the bark and have been Howard, R.A. 1989. Flora of the Lesser Antilles,
used to control it. Several herbicides are effective Leeward and Windward Islands. Vol. 5. Arnold
in controlling infestations (Pacific Island Arboretum, Harvard University, Jamaica Plain,
Ecosystems at Risk 2002). Repeated heavy MA. 604 p.
plowing or repeated burning are also effective
measures (Mune and Parham 1956). Invasive Species Specialist Group. 2002. Notes
from Jim Space on his survey of invasive plant
Benefits.—Guava helps protect the soil and can be species in Tonga. The World Conservation
a major participant in reforestation of disturbed Union. http://www.issg.org/features/
areas and abandoned pastures. It furnishes food invasives_on_tonga.html. 4 p.
and cover for wildlife. Domestic animals eagerly
consume the fruits. Guava is one of the most Liogier, H.A. 1994. Descriptive flora of Puerto
important fruits in the tropics and is exported to Rico and adjacent Islands. Vol. 3. Editorial de la
temperate areas. It is consumed fresh and made Universidad de Puerto Rico. Río Piedras, PR.
into juice, jams and jellies, and paste or “cheese.” 461 p.
598
Little, E.L., Jr. and F.L. Wadsworth. 1964. Rain-tree. 2002. Guava. Raintree Nutrition, Inc.,
Common trees of Puerto Rico and the Virgin Austin, Texas. http://www.rain-tree.com/guava.
Islands. Agriculture Handbook 249. U.S. htm. 5 p.
Department of Agriculture, Washington, DC.
548 p. Samarao, S.S. and M.A. Martins. 1999. Influence
of arbuscular mycorrhizal fungi, associated with
Long, R.W. and O. Lakela. 1976. A flora of addition of rutin, on the growth of guava
Tropical Florida. Banyon Books, Miami, FL. (Psidium guajava L.). Revista Brasileira de
962 p. Fruticultura 21(2): 196-199.
Morton, J. 1987. Guava. In: J.F. Morton. Fruits of Samson, J.A. 1986. Tropical fruits. 2nd ed.
warm climates. Julia F. Morton, Maimi, FL. Longman Scientific & Technical, Harlow, UK.
p. 356-363. 336 p.
Mune, T.L. and J.W. Parham. 1956. Weed control: Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Guava and its control in Fiji. Agricultural Montiel, eds. 2001. Flora de Nicaragua.
Journal, Fiji 27(3/4): 103-108. Monographs in Systematic Botany Vol. 85, No.
2. Missouri Botanical Garden, St. Louis, MO.
Pacific Island Ecosystems at Risk. 2002. Psidium p. 945-1,910.
guajava L., Myrtaceae. http://hear.org/pier_v3.3/
psgua.htm. 4 p. Tona, L., K. Kambu, K. Mesia, K. Cimanga, S.
Apers, T. de Bruyne, L. Pieters, J. Totte, A.J.
Pennington, T.D. and J. Sarukhan. 1968. Arboles Vlietinck, and T. de Bruyne. 1999. Biological
tropicales de México. Instituto Nacional de screening of traditional preparations from some
Investigación Forestales, Secretaría de medicinal plants used as antidiarrhoeal in
Agricultura y Ganadaría. Ciudad de México, Kinshasa, Congo. Phytomedicine 6(1); 59-66.
México. 413 p.
von Carlowitz, P.G. 1991. Multipurpose trees and
Popenoe, W. 1948. Manual of tropical and shrubs: Sources of seeds and inoculants.
subtropical fruits. Hafner Press, New York. International Council for Research in
474 p. Agroforestry, Nairobi, Kenya. 328 p.
599
Psorothamnus scoparius (Gray) Rydb. broom dalea
FABACEAE
James E. Nellessen
600
Growth and Management.—This species grows Dick-Peddie, W.A. 1993. New Mexico
relatively quickly and can reach 1 m in height or Vegetation: Past, Present and Future. University
diameter within 4 to 7 years depending on local of New Mexico Press, Albuquerque, NM. 244 p.
environmental conditions. The sandy Plains-Mesa
Sand Scrub habitats in which this shrub may be a Ivey, R.D. 1995. Flowering Plants of New Mexico,
dominant are by nature poor rangelands for 3rd Ed. Published by the author, NM. 504 p.
livestock.
Kearney, T.H., R. Peebles, and Collaborators.
Benefits.—Broom dalea is an important natural 1951 with 1960 supplement. Arizona Flora.
stabilizer of dune and other sandy soils. Its University of California Press, Berkeley, CA.
relatively rapid growth rate allows for reasonably 1,085 p.
quick establishment in situations of shifting and
disturbed soils. Like many members of the legume Lucero, M.E., R.E. Estell, and E.L. Frederickson.
family, it is a nitrogen fixer and forms rhizobial 2002. The essential oil composition of
root nodules (Allen and Allen 1981). In fact, this Psorothamnus scoparius (A. Gray) Rydb.
species has been documented to transmit the United States Department of Agriculture,
rhizobial bacteria to another genus within the Agricultural Research Service, Jornada
legume family, Crotalaria (Wilson 1939). Broom Experimental Range http://www.nal.
dalea is not an important range or browse plant for usda.gov/ttic/tektran. In press, Journal of
livestock and may increase under grazing. It is not Essential Oils.
substantially browsed, likely due to oils contained
within the glands (Warnock 1974). Very little, Martin, W.C. and C.E. Hutchins. 1980-1981
until recently, has been known about the volatile (reprinted 2001). A Flora of New Mexico. Vol.
oil composition of this species. Sixty-four volatile 1. Bishen Singh Mahendra Pal Singh, India and
compounds have been identified (Lucero and Koeltz Scientific Books, Germany. p. 1-1,276.
others 2002). The three most abundant compounds
were gamma-terpinene (22.3 percent), p-cymene Rozen, Jr., J.G. and Rozen B.L. 1986. Bionomics
(14.0 percent), and alpha-pinene (9.0 percent). of crepuscular bees associated with the plant
Some of these are known deterrents to herbivore Psorothamnus scoparius: Hymenoptera,
browsing in other plant species and may serve the Apoidea. Journal of the New York
same function in broom dalea. Entomological Society. 94(4): 472-479.
601
Psychotria brachiata Sw. palo de cachimbo
RUBIACEAE
John K. Francis
others 2001).
602
0.5 to 0.8 m/year for 2 or 3 years from sprouts. Liogier, H.A. 1997. Descriptive flora of Puerto
Individual stems probably live from 10 to 20 Rico and adjacent islands. Vol. 5. Editorial de la
years, but plants may persist longer by sprouting. Universidad de Puerto Rico, Río Piedras, PR.
Natural regeneration probably can be encouraged 436 p.
by disturbance designed to create forest openings.
Planting and management experience have not Lugo, A.E. 1992. Comparison of tropical tree
been published. plantations with secondary forests of similar
age. Ecological Monographs 62: 1-41.
Benefits.—Palo de cachimbo contributes to the
biodiversity of forests, helps protect the soil, and Missouri Botancial Garden. 2002. W3Tropicos:
furnishes food and cover for wildlife. nomenclature data base. http://mobot.mobot.
org/cgi-bin/search_vast#samer. [not paged].
References
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Croat, T.B. 1978. Flora of Barro Colorado Island. Montiel, eds. 2001. Flora de Nicaragua.
Stanford University Press, Stanford, CA. 943 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanical Garden, St. Louis, MO.
p. 1,911-2,666.
603
Psychotria deflexa DC. garricillo
RUBIACEAE
John K. Francis
604
stems of young plants layer (root) readily when in Allentown, PA. http://botany2002.org.section3/
contact with the soil. abstracts/21.shtml. 1 p.
605
Psychotria microdon (DC.) Urban thicket wild coffee
RUBIACEAE
John K. Francis
606
desirable. The species is a larval host for the Olmos, F. 1997. Tapirs as seed dispersers and
Xylophanes pluto (Fabricius) moth in Puerto Rico predators. IUCN/SSC Tapir Specialist Group.
(Torres-Bauzá 2000). http://www.tapirback.com/tapirgal/iucn-ssc/tsg/
action97/ap97-05.htm. 12 p.
References
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Howard, R.A. 1989. Flora of the Lesser Antilles, Montiel, eds. 2001. Flora de Nicaragua,
Leeward and Windward Islands. Angiospermas. Monographs in Systematic
Dicotyledoneae. Part 3. Vol. 6. Arnold Botany Vol. 85, No. 3. Missouri Botanical
Arboretum, Harvard University, Jamaica Plain, Garden Press, St. Louis, MO. p. 1,911-2,666.
MA. 658 p.
Torres-Bauzá, J.A. 2000. Ciclo de vida y aspectos
Liogier, H.A. 1997. Descriptive flora of Puerto de la biología de Xylophanes pluto (Fabricius)
Rico and adjacent islands. Vol. 5. Editorial de la en Puerto Rico (Lepidoptera: Sphingidae).
Universidad de Puerto Rico, Río Piedras, PR. Caribbean Journal of Science 36(3-4): 227-232.
436 p.
607
Psychotria nervosa Sw. shiny-leafed wild coffee
RUBIACEAE
John K. Francis
lanceolate, with sunken veins and elongated tips.
A large, flattened bud is almost always visible
between the youngest pair of leaves. The flower
clusters, which are terminal or axillary, are nearly
globose and contain a number of small, white
flowers. The clusters become more diffuse as the
fruits mature. The fruits (drupes) are bright red,
elliptic or globose at maturity and 6 to 7 mm long
(Howard 1989, Liogier 1997, Nelson 1996,
Stevens and other 2001). They are fleshy and juicy
with little flavor. Each contains two seeds that
have a rounded side with shallow striations and a
flattened side with a division down the middle like
a coffee bean (author’s observation).
608
Reproduction.—Shiny-leafed wild coffee blooms References
in spring and summer in Florida (Nelson 1996)
and irregularly through the year in Nicaragua Dave’s Garden Network. 2002. Wild coffee
(Stevens and others 2001). The flowers are visited (Psychotria nervosa). The Plants Database.
by honeybees and butterflies (Dave’s Garden http://plantsdatabase.com/go/2326.html. 2 p.
Network 2002) that presumably pollinate them.
A collection of fruits from Puerto Rico weighed an Dolan, S. 2002. Native shrubs.
average of 0.256 + 0.009 g/fruit. Air-dried seeds http://www.indialantic.com/contents/ecology/
from those fruits weighed an average of 0.0175 + shurbs.htm. 3 p.
0.0004 g/seed or 57,000 seeds/kg. About 85
percent of fruits from the collection contained two Howard, R.A. 1989. Flora of the Lesser Antilles,
seeds; the reminder contained one filled seed and Leeward and Windward Islands.
one aborted seed. Sown in commercial potting Dicotyledoneae. Part 3. Vol. 6. Arnold
mix, 95 percent of the filled seeds germinated Arboretum, Harvard University, Jamaica Plain,
between 64 and 164 days after sowing (author’s MA. 658 p.
observation).
Liogier, H.A. 1997. Descriptive flora of Puerto
Growth and Management.—Shiny-leafed wild Rico and adjacent islands. Vol. 5. Editorial de la
coffee has a moderate growth rate. Small wildlings Universidad de Puerto Rico, Río Piedras, PR.
and pruned larger plants transplant well. They also 436 p.
can be grown easily from seed. Once established
in a garden or natural area, they will reproduce Nelson, G. 1996. The shrubs and woody vines of
and eventually fill the area (Workman 1980). Florida. Pineapple Press, Inc., Sarasota, FL.
Where a seed source is available, probably the 391 p.
best strategy to promoting natural establishment is
to manipulate the stand for an open understory. Olmos, F. 1997. Tapirs as seed dispersers and
predators. IUCN/SSC Tapir Specialist Group.
Benefits.—Shiny-leafed wild coffee is a http://www.tapirback.com/tapirgal/iucn-ssc/tsg/
widespread and important understory plant in dry action97/ap97-05.htm. 12 p.
and moist forest and as such contributes to the
biodiversity and biomass accumulation, and School of Forest Resources and Conservation.
protects against soil erosion. It is frequently 2002. Florida forest plants: wild coffee
planted in Florida as a native landscape plant in (Psychotria nervosa). University of Florida,
both formal and natural gardens (Workman 1980). Gainesville, FL. http://www.plantatlas.usf.edu/
It is appreciated for its attractive foliage and bright maps.asp?plantID=2811. 2 p.
red berries and because it attracts birds and
butterflies (Dave’s Garden Network 2002). The Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
fruits are food for tapirs (Olmos 1997), a number Montiel, eds. 2001. Flora de Nicaragua,
of bird species (School of Natural Resources and Angiospermas. Monographs in Systematic
Conservation 2002), and probably many other Botany Vol. 85, No. 3. Missouri Botanical
animals. Shiny-leafed wild coffee contains the Garden Press, St. Louis, MO. p. 1,911-2,666.
alkaloid dimethyltryptamine, which is
hallucinogenic (Ted 2002) and may eventually Ted, M. 2002. Smokable dimethyltryptamine from
prove to have medicinal applications. The seeds organic sources. http://www.deoxy.org/
were once used as a coffee substitute (Dave’s smokedmt.htm. 11 p.
Garden Network 2002).
Workman, R.W. 1980. Growing native. The
Sanibel-Captiva Conservation Foundation, Inc.,
Sanibel, FL. 137 p.
609
Purshia tridentata (Pursh) DC. antelope bitterbrush
ROSACEAE
610
1999, Meyer 1989). Site-adapted seed sources 1993. Rules for testing seeds. Journal of Seed
should be selected for plantings. Antelope Technology 16: 1–113.
bitterbrush should be drill seeded in late fall or
winter to provide overwinter prechilling. It may be Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
seeded with other native species that are not highly Stevens, and B.C. Giunta. 1975. Characteristics
competitive. Seeding failures resulting from rodent and hybridization of important Intermountain
predation of seed can be avoided by seeding in late shrubs. I. Rose family. U.S. Department of
fall on fairly large disturbances that do not provide Agriculture, Forest Service, Intermountain
cover for the rodents (Evans and others 1983). Forest and Range Research Station, Ogden, UT.
Containerized or 1-0 bareroot seedlings are easily 35 p.
grown, but fungicide treatments may be required
to reduce losses to damping-off organisms. Booth, D.T. 1999. Imbibition temperatures affect
Nursery stock can be planted on steep, rocky bitterbrush seed dormancy and seedling vigor.
slopes that are inaccessible to seeding equipment. Journal of Arid Environments 48: 35-39.
It establishes best if planted in early spring when
antelope bitterbrush in the surrounding area is yet Cronquist, A., N.H. Holmgren, and P.K.
dormant. Herbaceous vegetation must be cleared Holmgren. 1997. Intermountain flora: vascular
around each seedling to reduce competition. plants of the Intermountain West, U.S.A. Vol. 3,
part 1. Subclass Rosidae: (except Fabales). The
Growth and Management.—Antelope New York Botanical Garden, The Bronx, NY.
bitterbrush seedings, plantations, and wild stands 446 p.
are vulnerable to heavy use by wild or domestic
ungulates and predation by gophers, mice, birds, Evans, R.A., J.A. Young, G.J. Cluff, and J.K.
and insects. Seedlings are not competitive with McAdoo. 1983. Dynamics of antelope
weedy annuals or most introduced perennial bitterbrush seed caches. In: A.R. Tiedemann and
grasses. Many mature stands are damaged by K.L. Johnson, comps. Proceedings: Research
continuous and close browsing by deer and and management of bitterbrush and cliffrose in
livestock. western North America. GTR-INT-152. U.S.
Department of Agriculture, Forest Service,
Benefits.—Next to sagebrush, antelope bitterbrush Intermountain Forest and Range Experiment
is probably one of the most widespread shrubs in Station, Ogden, UT. p. 195–202.
the Great Basin region, although its prevalence has
been reduced by range fires and exotic plant Jabbes, M. 2000. Hybridization and its
introductions (Mozingo 1987). It is important for a evolutionary consequences in Purshia and
host of wildlife and provides nutritious browse and Cowania. Ph.D. dissertation. University of
essential cover on fall, winter, and summer mule Idaho, Moscow, ID. 185 p.
deer ranges (Plummer and others 1968). Protein
content varies among populations depending upon McArthur, E.D., H.C. Stutz, and S.C. Sanderson.
over-winter leaf retention. New growth may also 1983. Taxonomy, distribution, and cytogenetics
be heavily used in spring and summer. Because of of Purshia, Cowania, and Fallugia (Rosoideae,
the varied growth forms, attractive foliage, and Rosaceae). In: A.R. Tiedemann and K.L.
showy flowers, antelope bitterbrush has potential Johnson, comps. Proceedings, Research and
as an ornamental in low-maintenance landscapes. management of bitterbrush and cliffrose in
Populations of antelope bitterbrush with distinctive western North America. GTR-INT-152. U.S.
attributes have been recognized and are Department of Agriculture, Intermountain Forest
commercially harvested and sold. Native and Range Experiment Station. p. 4–24.
Americans reportedly used bitterbrush as a remedy
for smallpox, measles, tuberculosis, and Meyer, S.E. 1989. Warm pretreatment effects on
pneumonia, and as an antiseptic for rashes and antelope bitterbrush (Purshia tridentata)
insect bites (Mozingo 1987). germination response to chilling. Northwest
Science 63: 146–153.
References
Mozingo, H.N. 1987. Shrubs of the Great Basin: a
AOSA [Association of Official Seed Analysts]. natural history. University of Nevada Press,
Reno, NV. 342 p.
611
Nord, E.C. 1965. Autecology of bitterbrush in Research and management of bitterbrush and
California. Ecological Monographs 35: 307– cliffrose in western North America. GTR-INT-
334. 152. U.S. Department of Agriculture, Forest
Service, Intermountain Forest and Range
Plummer, A.P., D.R. Christensen, and S.B. Experiment Station, Ogden, UT. p. 55–69.
Monsen. 1968. Restoring big game range in
Utah. Publication 68-3. Utah Division of Fish Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
and Game, Salt Lake City, UT. 183 p. Viability of seed from thirty-two shrub and forb
species through fifteen years of warehouse
Righetti, T.L., C.H. Chard, and D.N. Munns. 1983. storage. Great Basin Naturalist 41: 274–277.
Opportunities and approaches for enhancing
nitrogen fixation in Purshia, Cowania and USDA Forest Service. 1937. Range plant
Fallugia. In: A.R. Tiedemann and K.L. Johnson, handbook. U.S. Government Printing
comps. Proceedings: Research and management
of bitterbrush and cliffrose in western North
Office, Washington, D.C. 512 p.
America. GTR-INT-152. U.S. Department of
Agriculture, Forest Service, Intermountain Van der Wall, S.B. 1994. Seed fate pathways of
Forest and Range Experiment Station, Ogden, antelope bitterbrush: dispersal by seed-caching
UT. p. 214–224. yellow pine chipmunks. Ecology 75: 1911-1926.
Shaw, N.L. and S.B. Monsen. 1983. Phenology Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
and growth habits of nine antelope bitterbrush, Goodrich. 1987. A Utah flora. Great Basin
desert bitterbrush, Stansbury cliffrose, and Nturalist Memoirs 9. Brigham Young University
Apache plume accessions. In: A.R. Tiedemann Press. 894 p.
and K.L. Johnson, comps. Proceedings:
612
Quercus havardii Rydb. Havard shin oak
FAGACEAE
Synonyms: None
James E. Nellessen
613
Atriplex canescens (Pursh) Nutt. (four-wing considerable amounts of water during wet weather
saltbush), Sporobolus airoides (Torr.) Torr. (alkali with up to 50 percent of their mass being water.
sacaton), and Eriogonum annuum Nutt. (annual This gives the plant a great ability to survive
buckwheat). But there are a wide variety of other drought. During spring drought it may not leaf out
shrubs, forbs and grasses comprising numerous or may defoliate and leaf out later once water
other vegetation series within the greater shinnery becomes available. It generates more negative leaf
community. The tall grasses such as sand water potentials, -25 to -32 bars, than many other
bluestem, big bluestem (Andropogon gerardii species in the surrounding community. Soils under
Vitmann), and Indiangrass [Sorghastrum nutans shin oak, in addition to being sandy, are low in
(L.) Nash] have been reduced in abundance nitrogen (0.017 percent), low in organic matter
through overgrazing of Havard shin oak (0.36 percent) neutral to slightly alkaline in pH,
communities. Havard shin oak itself may remain and low in clay and carbonates (a caliche layer is
stable or also decrease with overgrazing, the cover inhibitory). Havard shin oak has been reported to
of grass and Havard shin oak often being have allelopathic effects on other plant species.
positively correlated. Stands of Havard shin oak on
sites with shallow water tables, such as in Chavez Reproduction.—Shin oak flowers in April to
County New Mexico, may contain cottonwoods. May. As with other oaks, male flowers are born in
Hybrids with other oak species, such as Q. stellata pendulous catkins, the female flowers solitary or a
Wang (post oak), occur and may be found within few clustered within leaf axils. The male catkins
typical Havard shin oak stands, the hybrids often are 2 to 3 cm long and densely flowered. The
attaining greater height (2 to 3 m), and stand out as female catkins are 3 to 7 mm long with one to five
readily visible clumps from the surrounding flowers. For a small shrubby oak, the acorns are
shorter plants. These hybrids tend to have quite large, 12 to 25 mm long, up to about 20 mm
intermediate characteristics such as larger leaves wide, brown, ovoid to oblong in shape. They are
and smaller acorns. These hybrids tend to be more born solitary or in pairs and the cup, 10 to 12 mm
common within the eastern portion of Havard shin deep, covers about a third to a half of the fruit. The
oak’s range (Correll and Johnston 1970, Flora of fruits mature from mid-July into September during
North America 1997, Peterson and Boyd 1998). the first year of development on the current
Havard shin oak also hybridizes with Q. mohriana season’s branches (Carter 1997, Correll and
Buckley ex Rydb. (Mohr shin oak) that occurs Johnston 1970, Flora of North America 1997,
within the rocky “breaks” area of the Texas Great Plains Flora Association 1986, Martin and
panhandle. Hybrids are intermediate in Hutchins 1980). Acorns of the white oak group,
characteristics and habitat location (Muller 1952). show little or no dormancy, and will germinate
immediately (Young and Young 1992). Acorn
Autoecology.—The plant primarily spreads by its moisture content for this group cannot drop below
underground rhizomes to form large clonal clumps 30 to 50 percent otherwise seed viability will be
that may range in size from as small as 2 to 15 m lost. Within a particular clone, acorn crops
in diameter to as large as 0.8 ha. Isozyme generally only occur every 3 to 10 years. Within a
electrophoresis identified 37 clones/ha, larger larger landscape scale area containing many
clones being free of interclonal growth, and the different clones, at least one or a few clones will
sample population was outcrossing based on have acorns. Most acorns are attacked by weevils
Hardy-Weinberg expected genotype ratios (Mayes and will decompose quickly. There is little
and others 1998). There are no data on horizontal reproduction by seed because of the short lifespan
growth rates but they are believed to be slow. The of the acorn, the need for good environmental
plants are long-lived, estimated at hundreds to conditions for germination and establishment at
thousands of years (Peterson and Boyd 1998). The the time of acorn drop, attack by weevils, and
rootstocks are massive, comprising the bulk of the decomposers.
plant mass (73 percent); the above-ground shoots
are merely small side shoots to the below ground Growth and Management.—For purposes of
parts. Above-ground biomass has been measured agriculture and livestock, shin oak has generally
at 1,821 kg/ha (Sears and others 1986) and annual been managed as an undesirable species. Havard
production has been estimated to be anywhere shin oak has sometimes been classified as an
from 550 to 4,500 kg/ha depending on the site increaser under livestock grazing, but reports do
(reviewed in Peterson and Boyd 1998). The roots not substantiate this (reviewed in Peterson and
(which are mycorrhizal) and rhizomes absorb Boyd 1998). Its poor reproduction by seed restricts
614
its spread primarily to the slow growing rhizome catkins and buds were higher in livestock grazed
system. An estimated 100,000 acres of land in plots than ungrazed plots. Crude protein and other
New Mexico have been treated with the herbicide nutritional qualities did not differ between
tebuthiuron for brush control. Tebuthiuron treatments (Boyd and others 2001). The acorns are
gradually kills the plant over a 3 to 4 year period. eaten by a variety of wildlife and are a staple food
Other chemicals that have historically been used for lesser prairie chickens (Tympanuchus
on Havard shin oak are 2,4-D, benzoic acids, pallidicinctus Ridgeway) in the fall and winter
2,4,5-T, and picloram. Tebuthiruron should not be seasons. Havard shin oak communities are
used in drought years, and the treated land should important breeding and nesting habitats for the
not be summer grazed for 2 years. Various lesser prairie chicken. Prescribed fire in shin oak
techniques of rotational and continuous grazing in habitats can improve grass and forb cover and
shin oak communities have been tested with mixed associated insect abundance for prairie chickens
results for livestock productivity. But rotational but at least a 2 to 3-year recovery period, no
grazing is best for overall ecosystem management burning in spring, and burning in patches are
including maintenance of proper native grass necessary (Boyd and Bidwell 2001). The rare sand
covers, and management of the lesser prairie dune lizard (Sceloporus arenicolus Degenhardt
chicken and perhaps other wildlife. Goats have and Jones), a threatened species in New Mexico, is
been used to control shin oak with reasonable restricted to Havard shin oak communities. Both
success, although goats eating it continuously will Native Americans and early Spanish colonists
lose weight. In addition to livestock use and direct would eat the acorns raw or baked. A flour can be
agricultural conversion of shin oak habitat, another made from the large acorns by first boiling them,
reason for shin oak management concerns the boll then roasting, chopping into pieces, drying for 30
weevil (Anthonomus grandis Boheman). This minutes, then passing them through a grinder a
cotton pest is known to overwinter in oak leaf litter couple of times (Warnock 1974).
and to reinfest cotton fields for up to a half mile
away (reviewed in Peterson and Boyd 1998). References
Above-ground shoots may live 11 to 15 years.
Shoot regeneration is rapid, 30 to 60 cm within the Boyd, C.S. and T.G. Bidwell. 2002. Effects of
first year, when above ground tissues are removed prescribed fire on shinnery oak (Quercus
mechanically or through fire. The response to fire havardii) plant communities in western
is variable and dependent on timing of the fire. Oklahoma. Restoration Ecology 10(2): 324-
Burning reduced shin oak cover by at least 25 333.
percent, with the greatest reduction, 50 percent,
following spring burns (Boyd and Bidwell 2002). Boyd, C.S. and T.G. Bidwell. 2001. Influence of
Winter and annual fires increased rhizomatous prescribed fire on lesser prairie chicken habitat
grass cover, and although oak cover was reduced, in shinnery oak communities in western
regrowth resulted in greater stem densities (e.g. Oklahoma. Wildlife Society Bulletin 29(3):
increasing from 20 stems/ha to 40 stems/ha). 938-947.
Benefits.—The sandy soils that Havard shin oak Boyd, C.S., L.T. Vermeire, T.G. Bidwell, and R.L.
grows in are susceptible to wind erosion. The Lochmiller. 2001. Nutritional quality of
extensive underground rootstock system and dense shinnery oak buds and catkins in response to
above ground shoots make it an important soil burning or herbivory. Southwestern Naturalist
stabilizer. Oak leaves can be good forage for sheep 46(3): 295-301.
and goats, but are less utilized by cattle.
Consumption of too much, especially young spring Carter, J.L. 1997. Trees and Shrubs of New
shoots, can be toxic and result in illness or even Mexico. Johnson Books, Boulder, CO. 534 p.
death for cattle and lambs (Gay and Dwyer 1998,
Warnock 1974). As with many oak species, Correll, D.S. and M.C. Johnston. 1970. Manual
toxicity is due to phenolics and tannins. Later in of the Vascular Plants of Texas. Texas Research
the growing season cattle will utilize it in small Foundation, Renner, TX. 1,881 p.
and varying degrees. The crude protein content of
foliage is about 9 percent. The plants are browsed Dick-Peddie, W.A. 1993. New Mexico
by deer, pronghorn antelope, and southern plains Vegetation: Past, Present and Future.
woodrats. Phenolic content (15 to 20 percent) of
615
University of New Mexico Press, Albuquerque, Mayes, S.G., M.A. McGinley, and C.R. Werth.
NM. 244 p. 1998. Clonal population structure and genetic
variation in sand-shinnery oak, Quercus
Flora of North America Editorial Committee. havardii (Fagaceae). American Journal of
1997. Flora of North America, Vol 3. Botany 85(11): 1609-1617.
Magnoliophyta: Magnoliidae and Hamamelidae.
Flora of North America Editorial Committee, McCoy, D. 1981. Roadside Trees and Shrubs of
Oxford University Press, New York. 590 p. Oklahoma. University of Oklahoma Press,
Norman, OK. 116 p.
Freeman, C.C. 2000. Vascular plants new to three
states in the central United States. Transactions Muller, C.H. 1952. Ecological control of
of the Kansas Academy of Science 103(1-2): 51- hybridization in Quercus; A factor in the
54. mechanism of evolution. Evolution 6: 147-161.
Gay, C.W., Jr. and D.D. Dwyer. 1998 Reprint. Peterson, R.S. and C.S. Boyd. 1998. Ecology and
New Mexico Range Plants. Cooperative Management of Sand Shinnery Communities: A
Extension Service Circular 374. Revisions by: Literature Review. General Technical Report
C. Allison, S. Hatch, and J. Schickedanz. New RMRS-GTR-16. United States Department of
Mexico State University, Las Cruces, NM. Agriculture Forest Service, Rocky Mountain
84 p. Research Station, Fort Collins, CO. 44 p.
Great Plains Flora Association. 1986. Flora of the Sears, W.E., C.M. Britton, D.B. Wester, and R.D.
Great Plains. University Press of Kansas, KS. Pettit. 1986. Herbicide conversion of a sand
1,392 p. shinnery oak (Quercus havardii) community:
effects on biomass. Journal of Range
Kartesz, J.T. 1994. A Synonymized Checklist of Management 39: 399-403.
the Vascular Flora of the United States, Canada,
and Greenland, Vol. 1, 2nd Edition. Biota of Warnock, B.H. 1974. Wildflowers of the
North America Program of the North Carolina Guadalupe Mountains and the Sand Dune
Botanical Garden. Timber Press, Portland, OR. Country, Texas. Sul Ross State University,
622 p. Alpine, TX. 176 p.
Martin, W.C. and C.E. Hutchins. 1980 (reprinted Young, J.A. and C.G. Young. 1992. Seeds of
2001). A Flora of New Mexico, Vol. 1. Bishen Woody Plants in North America. Dioscorides
Singh Mahendra Pal Singh, India and Koeltz Press, Portland, OR. 407 p.
Scientific Books, Germany. 1,276 p.
616
Randia aculeata L. white indigo berry
RUBIACEAE
John K. Francis
617
tenaciously after disturbance and is prominent in References
secondary forests arising after logging or partial
clearing for pasture. Plants are usually well Dade County. 2001. Some trees and shrubs native
disbursed and seldom, if ever, form thickets. to South Florida. http://www.fnps.org/dade/
pubs/SomeTrees.html. 6 p.
Reproduction.—White indigo berry blooms and
fruits irregularly throughout the year. Fruits (n = Howard, R.A. 1989. Flora of the Lesser Antilles,
73) collected in Puerto Rico weighed an average Leeward and Windward Islands.
of 0.472 + 0.014 g/fruit. Fourteen of the fruits Dicotyledoneae. Part 3. Vol. 6. Arnold
contained from three to 14 seeds and averaged 8.2 Arboretum, Harvard University, Jamaica Plain,
seeds. The air dry seeds from these fruits averaged MA. 658 p.
0.0264 + 0.0003 g/seed or 38,000 seeds/kg. Sown
in commercial potting mix, 25 percent germinated Liogier, H.A. 1990. Plantas medicinales de Puerto
between 46 and 168 days after sowing. Rico y del Caribe. Iberoamericana de Ediciones,
Inc., San Juan, PR. 566 p.
Growth and Management.—Weaver (1990)
measured 12 plants over a 5-year period in closed Liogier, H.A. 1997. Descriptive flora of Puerto
low forest in St. John, U.S. Virgin Islands and Rico and adjacent islands. Vol. 5. Editorial de la
found the vary slow diameter growth rate of 0.06 Universidad de Puerto Rico, Río Piedras, PR.
+ 0.02 cm/yr. A 1-m tall sapling in a moist area of 436 p.
Puerto Rico with a basal diameter of 2 cm had 10
growth rings (author’s observation). Ornamental Little, E.L., Jr., R.O. Woodbury, and F.H.
plants also grow slowly (Workman 1980). Seeds Wadsworth. 1974. Trees of Puerto Rico and the
for nursery production should be cleaned by Virgin Islands. Vol. 2. U.S. Department of
maceration and wet sieving before sowing. Agriculture, Agriculture Handbook 449.
Production of seedlings in containers is Washington, DC. 1,024 p.
recommended. Because growth is slow, a year or
more in the nursery is required followed by Nelson, G. 1996. The shrubs and woody vines of
protection from weeds in the field. Florida. Pineapple Press, Inc. Sarasota, FL.
391 p.
Benefits.—White indigo berry has been used as an
ornamental to a limited extent in Florida and, Smithsonian Institution. 2001. Centers of plant
because it is native and resists drought and diversity: Columbian Central Massif.
hurricanes, it is currently being recommended for http://www.nmnh.si.edu/botany/projects/cpd/sa/
planting (Dade County 2001). It is available as sa29.htm. 14 p.
nursery-grown plants, and plants from the wild
can be successfully dug up and transplanted. The Vélez, I. and J. van Overbeek. 1950. Plantas
fruits are edible although of poor flavor indeseables en los cultivos tropicales. Editorial
(Workman 1980). White indigo berry provides Universitaria, Río Piedras, PR. 497 p.
food and nesting sites for birds (Vélez and van
Overbeek 1950) and nectar for butterflies (Dade Weaver, P.L. 1990. Tree diameter growth rates in
County 2001). The wood is hard and heavy and Cinnamon Bay Watershed, St. John, U.S. Virgin
useful for fuel. The shrub is sometimes decorated Islands. Caribbean Journal of Science 26(1-2):
for a Christmas tree, hence the Spanish name, 1-6.
árbol de navidad. Other names (tintillo and
inkberry) arose from the former use of the berries Workman, R.W. 1980. Growing native. The
for dye and for ink (Little and others 1974). The Sanibel-Captiva Conservation Foundation, Inc.,
plant is also frequently formed into bonsai plants. Sanibel, FL. 137 p.
The fruits are used in herbal medicine to control
dysentery. An unspecified part of the plant is used
to control fever, and the latex is reported to
effectively stop bleeding (Liogier 1990).
618
Rauvolfia viridis Willd. ex Roemer & J.A. Schultes bitter bush
APOCYNACEAE
John K. Francis
General Description.—Bitter bush, also known Reproduction.—A sample of bitter bush fruits
as bellyache bush, balsam, sassafras, milk tree, collected in Puerto Rico weighed an average of
and ti bois-lait, is a branchy shrub to 2.5 m in 0.124 + 0.005 g/fruit. Air-dried seeds separated
height (Howard 1989, Liogier 1995). The species from these fruits averaged 0.0247 + 0.0004 g/seed
rarely becomes a small tree to 4 m in height or 40,500 seeds/kg. Sown in commercial potting
(Liogier 1995) and 7 cm in diameter at breast mix without any pretreatment, 5 percent
height (Weaver 1990). Usually, there are multiple germinated after 11 months. Birds that eat the
stems, as suckers arising from lateral roots, fruits disburse the seeds.
especially after fire or other disturbance. The root
system is dominated by lateral roots, which Growth and Management.—Growth of bitter
support sinker roots and a moderate amount of bush is slow. The author estimated height growth
fine roots. The ovate or ovate-elliptic leaves are 5 of young shrubs from sprouts at 0.5 m/year.
to 15 cm long by 3 to 5 cm broad, and grow in Weaver (1990) measured the diameter growth rate
whorls of four or rarely three leaves. The several- of two old small trees and found an average of
to many-flowered inflorescences are usually only 0.01 cm/yr. No planting or management
terminal. The tiny flowers are tubular and white. experience has been reported.
The fruits are spherical or slightly flattened and 5
to 8 mm in diameter. Each contains two seeds Benefits.—The roots of bitter bush were analyzed
(Howard 1989, Liogier 1995). for alkaloids. Ajmalidine, a-yohimina, reserpinine,
quebrachidine, and yohimbina were isolated.
Range.—According to Howard (1989) and Reserpine, a sedative present in other Rauvolfias,
Liogier (1995) bitter bush is native to Hispaniola, was not found (Martínez Pérez and others 1997).
Puerto Rico, the Virgin Islands, the Lesser Bitter bush furnishes minor food and cover for
Antilles, Trinidad, Margarita, and Venezuela. birds and other wildlife. With other dry forest
There are also records for Cuba (Martínez-Pérez vegetation, it protects the soil from erosion.
and others 1997), Navassa Island (New York
Botanical Garden 2001), Costa Rica
619
References Tápanes, and V. Fuente Fiallo. 1997. Bases
débiles minoritarias de la raíz de Rauvolfia
Departamento de Botánica. 1999. Especies de viridis Roem. et Schult. Revista Cubana Farm
plantas nuevas para Costa Rica. Departamento 31(3): 188-191.
de Botánica, Inventario Biótico.
http://www.minae.go.cr/estrategia/estudio/planta Missouri Botanical Garden. 2001. Current
s1.htm. 4 p. specimen list for Rauvolfia viridis.
http://mobot.mobbot.org/cgi-bin/search_vast.
Howard, R.A. 1989. Flora of the Lesser Antilles, 1 p.
Leeward and Windward Islands.
Dicotyledoneae. Part 3. Vol. 6. Arnold The New York Botanical Garden. 2001. Flora of
Arboretum, Harvard University, Jamaica Plain, Navassa Island: vascular plant checklist.
MA. 658 p. http://www.nybg.org/bsci/hcol/navassa/checklist
vasc.html. 9 p.
Liogier, H.A. 1995. Descriptive flora of Puerto
Rico and adjacent islands. Vol. 4. Editorial de la Weaver, P.L. 1990. Tree diameter growth rates in
Universidad de Puerto Rico, Río Piedras, PR. Cinnamon Bay Watershed, St. John, U.S. Virgin
617 p. Islands. Caribbean Journal of Science 26(1-2):
1-6.
Martínez Pérez, J., A. Bello Alarcío, O. Montero
620
Rhododendron macrophyllum D. Don ex G. Don Pacific rhododendron
ERICACEAE
Christopher Ross
621
State Reserve, Pacific rhododendron is considered for the western hemlock zone: Mt. Hood
to be an early seral stage after fire or logging, and National Forest. R6-ECOL-232A. U.S.
before tanoak, which is being thinned in order to Department of Agriculture, Forest Service,
set back the succession process and maintain the Pacific Northwest Region, Portland, OR.
rhododendrons (California State Parks 2002). 111 p.
Alberta Government. 2002. Rhododendron Kingsbury, J.M. 1964. Poisonous plants of the
toxicity. http://www.Agric.gov.ab.ca/crops. United States and Canada. Prentice Hall, Inc.
[not paged]. Englewood, NJ. 626 p.
Burrill, L.C., W.S. Braunworth Jr., and R.D. Olson, D.F., Jr. 1974. Rhododendron L.
William, comps. 1989. Pacific Northwest weed rhododendron. In: C.S. Schopmeyer, ed. Seeds
control handbook. Oregon State University, of woody plants in the United States.
Extension Service, Agricultural Agricultural Handbook 450. U.S. Department
Communications. Corvallis, OR. 276 p. of Agriculture, Forest Service, Washington,
D.C. p. 709-712.
CalFlora. 2000. Information on California plants
for education, research and conservation. University of California Cooperative Extension.
Berkeley, California: The CalFlora Database. 2002. Sudden oak death. http://cemarin.
http://www. Calflora.org/ [not paged]. ucdavis.edu. 2 p.
California State Parks–Kruse Rhododendron State USDA NRCS. 2002. Plants Source and Reference.
Reserve. 2002. http://cal-parks.ca.gov. http://plants.usda.gov. [not paged].
Dyrness, C.T. 1973. Early stages of plant U.S. Department of Agriculture, Forest Service,
succession following logging and burning in the Rocky Mountain Research Station, Fire
western Cascades of Oregon. Ecology 54: 57- Sciences Laboratory. 2002. Fire Effects
69. Information System, http://www.fs.fed.us/ [not
paged].
Foss, C. and A. Antonelli. 1999. Hortsense.
http://pep.wsu.edu/hortsense. [not paged]. Wallace, G. 1993. Rhododendron. In: J.C.
Hickman, ed. 1993. The Jepson Manual:
Halpern, C.B. 1989. Early successional patterns of higher plants of California. U.C. Press,
forest species: Interactions of life history traits Berkeley and Los Angeles, CA. 1,400 p.
and disturbance. Ecology 70: 704-720.
Young, J.A. and C.G. Young. 1992. Seeds of
Halverson, N.M., C. Topik, and R. Van Vickle. woody plants of North America. Disceroides
1986. Plant association and management guide Press, Portland, OR. 407 p.
622
Rhododendron maximum L. rosebay rhododendron
ERICACEAE
Synonyms: none
Barton D. Clinton
optimum carbon gain, and has a tremendous
capacity for avoiding cavitation during freeze-thaw
cycles (Lipp and Nilsen 1997). Where extensive
overstory mortality has eliminated most of the
overstory, this species forms a thick and
continuous subcanopy known locally as ‘laurel
slicks’ or ‘laurel hells’. Rosebay rhododendron is
an important structural and functional component
of southern Appalachian forest ecosystems. What
isn’t clear is whether or not we are in a period of
advancement or retreat for this species. For
example, on poorly drained sites on ridge or upper
slope positions, large areas of rosebay
rhododendron, particularly at the high elevations,
have recently died out presumably due to the
General Description.—Rosebay rhododendron, phytophthora fungus, or due to recent prolonged
sometimes known as ‘great laurel’, is an periods of below-average precipitation. Yet,
understory evergreen shrub. It ranges in height rosebay rhododendron now occupies sites that
from 2 to 8 m, and can occasionally reach historically were free of evergreen understory.
diameters of 20 to 25 cm. Leaves are There are still important questions to be answered
schlerophyllous, simple, alternate, and oblong (10 regarding this species to completely understand its
to 30 cm long, 5 to 8 cm wide). It retains its waxy, role in forest understories.
deep-green leaves for up to 8 years, but once shed
are slow to decompose. It produces large, showy, Reproduction.—Rosebay rhododendron is clonal.
white to purple flowers each June. It is capable, however, of reproducing both
vegetatively and sexually. It reproduces
Range.—Worldwide, there are over 800 species in vegetatively through a process called ‘layering’
the genus Rhododendron. Most of the 26 North where it produces roots from aboveground woody
American species occur in the East along the parts when in contact with the forest floor. The
mountain chain stretching from Nova Scotia in the fruit is produced from showy flowers from March
north to Georgia and Alabama in the south, but are to August. The fruit is an oblong capsule that
concentrated in the southern Appalachians (Dobbs ripens in the fall, and splits along the sides soon
1998). Rosebay rhododendron is the most after ripening to release large numbers of minute
frequently occurring and dominant species of this seed (approx. 400 per capsule) (Schopmeyer
genus in the southern Appalachian region 1974). Microsite requirements for seed
(Swanson 1994), and occurs occasionally on mesic germination are relatively specific (e.g., high in
hill-slopes throughout the upper Piedmont organic matter such as rotting logs); hence, the
Crescent of the Southeastern United States. majority of reproduction is vegetative resulting in
a clonal distribution.
Ecology.—Approximately 1.2 million ha in the
southern Appalachians are occupied by this Growth and Management.—Seeds from rosebay
species (Dobbs 1995) where it dominates the rhododendron are minute and it is estimated that
understory. This species has historically been approximately 11 million are contained in 1 kg.
confined to riparian areas and other mesic sites but Commercial seed production is generally from
takes advantage of disturbed areas where it is cultivated hybrids. Seeds from wild sources are not
present to advance onto sub-mesic sites. It prefers commonly sold commercially. Rosebay
deep well-drained acid soils high in organic matter rhododendron is a slow-growing shrub and has a
where it produces a thick, peat-like humus. It very high sprout potential. If mechanical removal
prefers low to medium light conditions for is attempted in the case of forest management,
623
extremely high densities are attained by this Clinton. B.D. and J.M. Vose. 1996. Effects of
species in a manner of a few years. Prescribed fire Rhododendron maximum L. on Acer rubrum L.
has also been used to control this species but with seedling establishment. Castanea 61(1):38-45.
limited success (Clinton and Vose 2000).
Clinton, B.D. and J.M. Vose. 2000. Plant
Benefits.—Rosebay rhododendron is a striking succession and community restoration
and aesthetically pleasing feature of mesic following felling and burning in the southern
southern Appalachian forests. It is one of the Appalachian Mountains. In: W. K. Moser and
largest and hardiest rhododendrons grown C.F. Moser, eds. Fire and forest ecology:
commercially. Several cultivars with white to innovative silviculture and vegetation
purple flowers have been selected for the management. Tall Timbers Fire Ecology
horticultural trade (Brown and Kirkman 1990). Conference Proceedings, No. 21. Tall Timbers
Where it occurs naturally, it produces a showy, Research Station, Tallahassee, FL. p . 22-29.
white, pink, or light purple flower primarily in
June, but occurs from March into August. Rosebay Dobbs, M.M. 1995. Spatial and temporal
rhododendron maintains deep-green foliage year distribution of the evergreen understory in the
round. This species affords protection to steep southern Appalachians. Master’s Thesis,
watersheds and shelter for wildlife. The wood is University of Georgia, Athens, GA. 100 p.
very hard and is occasionally used for specialty
wood products. Dobbs, M.M. 1998. Dynamics of the evergreen
understory at Coweeta Hydrologic Laboratory,
Detrimental Effects.—For all its prized qualities North Carolina. PhD Dissertation, University
as a naturally occurring component of the of Georgia, Athens GA. 179 p.
landscape or as plantings in residential and
commercial landscaping, rosebay rhododendron Lipp, C.C. and E.T. Nilsen. 1997. The impact of
can have an inhibitory effect on regeneration of subcanopy light environment on the hydraulic
other plant species. There is some evidence to vulnerability of Rhododendron maximum to
suggest that due to fire suppression and the freeze-thaw cycles and drought. Plant, Cell
absence of other cultural activities (i.e., mountain- and Environment 20:1,264-1,272.
land grazing), this species has advanced beyond
the mesic forest sites into sub-mesic understories Nilsen, E.T., B.D. Clinton, T.T. Lei, O.K. Miller,
(Dobbs 1998). The significance of this movement S.W. Semones, and J.F. Walker. 2001. Does
onto previously unoccupied sites centers around Rhododendron maximum L. (Ericaceae) reduce
the impacts of rosebay rhododendron on plant the availability of resources above and
succession (Clinton and Vose 1996) and resource belowground for canopy tree seedlings?
availability (Nilsen and others 2001). Rosebay American Midland Naturalist 145:325-343.
rhododendron is associated with reduced woody
and herbaceous seedling abundance throughout its Nilsen, E.T., J.F. Walker, O.K. Miller, S.W.
range, and hence poses a serious impediment to Semones, T.T. Lei, and B.D. Clinton. 1999.
the production of wood products. The Inhibition of seedling survival under
mechanism(s) by which rosebay rhododendron Rhododendron maximum (Ericaceae): could
reduces seedling survival has been the subject of allelopathy be a cause? American Journal of
much debate. Possible sources of inhibition Botany 86(11):1,597-1,605.
include allelopathy, competition for resources
including light, physical and chemical attributes of Schopmeyer, C.S. 1974. Seeds of woody plants in
the forest floor and soil, and interactions between the United States. Agricultural Handbook 450.
some or all sources (Nilsen and others 1999, U.S. Department of Agriculture, Forest
Nilsen and others 2001). Service, Washington, DC. 883 p.
624
Rhus copallinum L. shining sumac
ANACARDIACEAE
Synonyms: None
Kristina Connor
625
panicles are pubescent, about 12 to 15 cm long,
and can be either terminal or axillary (Brickell and Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
Zuk 1996, Johnson and Hoagland 1999, A Concise Dictionary of Plants Cultivated in the
Krussmann 1976). Brown and Kirkman (1990) United States and Canada. McMillan Publishing
report the presence of polygamous flowers but Co., Inc., New York. 1,312 p.
state that shining sumac is functionally dioecious
since the pistils of the polygamous flowers abort. Brickell, C. and J.D.. Zuk. 1996. The American
Thus, both male and female plants are needed for Horticultural Society A-Z Encyclopedia of
good fruit set to occur (Arborquest 2001) The 3- to Garden Plants. DK Publishing, Inc., New York.
5- mm pubescent fruits are small drupes that 1,092 p.
contain a single nutlet (Coladonato 1992). Fruits
ripen from August to October. They turn dark red Brinkman, K.A. 1974. Rhus L. Sumac. In: C.S.
when ripe and often remain on the plant Schopmeyere, tech. coord. Seeds of woody
throughout the winter. The hard-coated seeds plants in the United States. Agriculture
germinate poorly without pretreatment, but a 1 hr Handbkook 450. U.S. Department of
soak in sulfuric acid at room temperature can Agriculture, Forest Service Washington, D.C.
result in a 75 percent germinative capacity 883 p.
(Brinkman 1974).
Brown, C.L. and L.K. Kirkman. 1990. Trees of
Growth and Management.—Shining sumac is a Georgia and adjacent states. Timber Press,
fast growing but short lived clonal shrub, first Portland, OR. 292 p.
cultivated in 1688 (Brinkman 1974). It is planted
as an ornamental and used in mass plantings Coladonato, M. 1992. Rhus copallinum. In: U.S.
because of its brilliant red fall foliage but it can Department of Agriculture, Forest Service,
spread well beyond desired borders because of its Rocky Mountain Research Station Fire Sciences
ability to sprout from the roots. Laboratory, Fire Effects Information System
http://www.fs.fed.us/database/feis. 9 p.
Benefits.—Shining sumac is nonpoisonous to
humans (USDA-ARS [no date], Seiler and Grelen, H.E. and V.L. Duvall. 1966. Common
Peterson 2001, Synor and Cowen [no date]), and plants of longleaf pine-bluestem range. Research
the fresh fruits can be used to make a lemon- Paper SO-23. U.S. Department of Agriculture,
tasting beverage. It was also used by Native Forest Service, Southern Forest Experiment
Americans to treat dysentery and mouth sores Station, New Orleans, LA. 96 p.
(Tenaglia 2002). Dense thickets of shining sumac
serve as cover for birds and mammals. The seeds Johnson, F.L. and B.W. Hoagland. 1999. Rhus
are eaten by a variety of birds, while the flowers copallinum L. Catalog of the Woody Plants of
attract butterflies. Deer and rabbits commonly Oklahoma, Oklahoma Biological Survey.
browse the twigs in winter; rabbits also eat the http://www.biosurvey.ou.edu. 1 p.
bark (Grelen and Duvall 1966, Oplin 2001,
Arborquest 2001, Coladonato 1992). It is not, Krüssmann, G. 1976. Manual of Cultivated Broad-
however, a preferred food and is considered a poor Leaved Trees and Shrubs. Volume I, A-D.
to moderately important browse (Coladonato Timber Press, Beaverton, OR. 448 p.
1992). However, the seeds, bark and leaves, while
low in nutritive value, have a high tannin content Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope.
and have been used by the leather industry (Bailey 1996. Plants for American Landscapes.
and Bailey 1976, Brown and Kirkman 1990, Louisiana State University Press, Baton Rouge,
Coladonato 1992). Its ability to sprout from the LA. 266 p.
roots, coupled with its rapid growth rate, make
shining sumac a good species to plant for erosion Oplin. 2001. Dwarf Sumac--Rhus copallina. Ohio
control (Arborquest 2001). Public Library Information Network (OPLIN)
and the Ohio Historical Society (OHS).
References http://www. oplin.lib.oh.us/products/tree. 2 p.
Arborquest. 2001. Rhus copallina. Seiler, J.R. and J.A. Peterson. 2001.Shining
http://www.arborquest.com. 2 p. sumac. Virginia Tech. Dendrology Webpage
626
http://www.fw.vt.edu/dendro/dendrology/syllab USDA, ARS National Genetic Resources Program.
us/ivirginica.htm. 1 p. Germplasm Resources Information Network-
(GRIN). [no date]. Online Database. National
Synor, T.D. and W.F. Cowen. [no date]. Ohio Germplasm Resources Laboratory, Beltsville,
trees. Rhus--Sumac. The Ohio State University MD. http://www.ars-grin.gov/npgs/
College of Food, Agricultural, and searchgrin.html.
Environmental Sciences, Extension Bulletin
700-00. http://www.ag.ohio- USDA, NRCS. 2001. The PLANTS database.
state.edu/~ohioline/b700. 3 p. National Plant Data Center, Baton Rouge, LA.
http://plants.usda.gov. [not paged].
Tenaglia, D. 2002. Rhus copallinum. The Missouri
Flora Website. http://www.missouriplants.com.
4 p.
627
Rhus glabra L. smooth sumac
ANACARDIACEAE
John K. Francis
gray-brown stem but often occurs in clonal
thickets formed by suckers from horizontal roots.
The relatively few branches and twigs are thick,
have a soft pith, and large, horseshoe-shaped leaf
scars. Stems and branches ooze a milky-resinous
sap when cut. The alternate leaves are uneven,
pinnately compound, up to 46 cm long, and
support seven to 29 leaflets. Leaflets are almost
stalkless and have serrate edges. (Clark 1973). The
foliage turns a brilliant red or scarlet during the
fall. Small, greenish-yellow flowers are grouped in
tight, 15- to 25-cm terminal panicles. The fruits are
dry, hairy, red to scarlet drupes that contain one
hard ovoid, dark-gray nutlet. The fruits have a
tangy, sour taste (Johnson 1995). Smooth sumac is
dioecious (male and female plants) (Wyman
1969). Total above-ground dry biomass may be
predicted by the equation: dry weight = 1.5130 +
0.6292 D2H, where D = diameter at 1 cm above
groundline, H = total height and r2 = 0.974
(Reeves and Lenhart 1988).
628
In Western areas it grows in stable communities on requires more than ordinary maintenance to keep
mountains, hillsides, and canyons. The species is smooth sumac from spreading vegetatively. One
used as a climax indicator in a number of shrub- method of keeping plants in check is to establish
grassland communities (Fire Sciences Laboratory them in soil space limited by rock or concrete
2003). Smooth sumac usually grows with full sun barriers (Clark 1973). The tops are killed by fires,
but tolerates partial shade. It grows from low but underground rhizomes (7.5 to 30 cm deep) are
elevations in many areas and up to 2,100 m in not damaged and sprout vigorously, increasing the
Arizona (Kearney and Peebles 1951). density and extent of smooth sumac stands. Seeds
from the soil seed bank are also stimulated to
Reproduction.—Smooth sumac flowers from germinate. Spring fires are more beneficial than
May to late July, depending on location. Fruits late summer fires (Fire Sciences Laboratory 2003).
ripen in September and October and persist When it is necessary to control the plant, it is
through the fall and winter (Fire Sciences recommended that they be lopped during or
Laboratory 2003). The flowers are pollinated by shortly after flowering and the sprouts spot-
bees (Rowe and Blazich 2003). Smooth sumac sprayed with glyphosate (Hutchison 2003).
produces at least some fruits and seeds nearly
every year (Fire Sciences Laboratory 2003). There Benefits and Detriments.—Smooth sumac helps
are about 2,000 seeds per well-formed panicle protect the soil in disturbed areas, adds to the
(Browse-Shrub and Forb Committee 1985), much aesthetics of wildlands, and furnishes food and
less in some areas and during dry years. Rowe and cover for wildlife. The fruits are eaten by a wide
Blazich (2003) report 50,600 to 105,600 fruits/kg variety of birds (Johnson 1995). Smooth sumac
and 52,800 to 277,200 seeds/kg. Plummer and foriage is rated poor in both energy and protein
others (1968) found 137,700 seeds/kg in samples value (Fire Sciences Laboratory 2003). White tail
from Utah. Air-dried fruits collected by the author deer (Odocoileus virginianus) browse it during fall
in Utah averaged 0.0202 + 0.0004 g/fruit. Seeds and winter but do not weaken the plants (Strauss
separated from them averaged 0.0103 + 0.0002 1991). It is also browsed by mule deer (Odocoileus
g/seed or 97,000 seeds/kg. The seeds are dispersed hemionus) (Fire Sciences Laboratory 2003).
principally by birds and form a soil seed bank Smooth sumac is valued as an ornamental for its
(Fire Sciences Laboratory 2003). Once individual brilliant red and scarlet fall colors and because it
plants are established, horizontal roots sucker to can be planted almost anywhere in the Continental
form clonal thickets. United States. It is particularly useful for
Nursery plants are usually produced from vegetating sandy banks and for low screens (Clark
seed, although production from cuttings is also 1973) and has been recommended for establishing
done to preserve favorable traits of select strains “living snow fences.” The species has been planted
(such as assuring only female plants). Rootstalks to revegetate roadsides and cuts, landfills, and strip
pulled up from wild stands in the early spring mines (Fire Sciences Laboratory 2003, Plummer
when the ground is wet will survive almost 100 and others 1968). In addition to using dried leaves
percent if planted properly (Plummer and others as an adulterant for tobacco and making drinks
1968). Seed can be collected in quantity by hand in from the fruits, Native Americans employed
fall or early winter. Seeds may be separated from smooth sumac to treat a large number of ailments,
the fruits by rubbing or beating followed by particularly mouth and throat sores, burns, to
screening (Rowe and Blazich 2003) and can be control diarrhea, and to promote urination (Fire
stored for as long as 5 years without loss of Sciences Laboratory 2003, Giese 1996, Moerman
viability (Fire Sciences Laboratory 2003). 1986). Basis for medicinal uses may be found in
Scarification to break physical (endocarp) the fact that methanol extracts of ground branches
dormancy may be done by a 3 to 4 hr soak in were effective (although not as effective as
concentrated sulfuric acid (Rowe and Blazich commercial antibiotics) against total of 11 species
2003) or emersion for less than 1 min. in boiling of bacteria. The antibacterial compounds were
water (Li and others 1999). identified as methyl gallic acid, gallic acid, and the
methyl ester of 3,4,5-trihydroxybenzoic acid
Growth and Management.—Smooth sumac has a (Saxena and others 1994). Some people develop
moderate growth rate of 30 to 46 cm in height per allergic dermatitis after contacting the sap, but the
year (Michigan State University Extension 1999). species is not as dangerous as many other Rhus
Although clones may last much longer, individual species (Clark 1973). Thickets of smooth sumac
stems are not long-lived, perhaps 2 to 10 years. It can prevent the development of tree seedlings
629
through shade and root competition and arrest in seeds of two North American Rhus species
succession (Putz and Canham 1992). (Anacardiaceae). American Journal of Botany
86(11): 1,505-1,511.
References
Michigan State University Extension. 1999. Rhus
Abrams, L. 1951. Illustrated flora of the Pacific glabra—smooth sumac. http://www.msue.msu.
States. Vol. 3. Stanford University Press, edu/msue/imp/modzz/00001245.html. 1 p.
Stanford, CA. 866 p.
Moerman, D.E. 1986. Medicinal plants of Native
Browse-Shrub and Forb Committee. 1985. America. Research Reports in Ethnobotany Vol.
Handbook on seeds of browse-shrubs and forbs. 1. University of Michigan Museum of
Tech. Pub. R8-TP8. Association of Official Seed Anthropology, Ann Arbor, MI. 534 p.
Analysts and U.S. Department of Agriculture,
Forest Service, Southern Region, Atlanta, GA. Natural Resources Conservation Service. 2003.
246 p. Plants profile: Rhus glabra L., smooth sumac.
http://plants.usda.gov/cgi_bin/plant_profile.cgi?
Clark, L.J. 1973. Wild flowers of British symbol=RHGL. 5 p.
Columbia. Gray’s Publishing Limited, Sidney,
BC, Canada. 591 p. Plummer, A.P., D.R. Christensen, and S.B.
Monsen. 1968. Restoring big-game range in
Fire Sciences Laboratory. 2003. Fire effects Utah. Pub. 68-3. Utah Division of Fish and
information system: Rhus glabra. U.S. Game, Salt Lake City, UT. 183 p.
Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences Putz, F.E. and C.D. Canham. 1992. Mechanism of
Laboratory, Missoula, MT. http://www.fs. arrested succession in shrublands: root and shoot
fed.us/database/feis/plants/shrub/rhugla/all.html. competition between shrubs and tree seedlings.
28 p. Forest Ecology and Management 49(3-4): 267-
275.
Giese, P. 1996. Red sumac: multi-tribal use.
http://www.kstrom.net/isk/food/rhususes.html. Reeves, H.C. and J.D. Lenhart. 1988. Fuel weight
4 p. prediction equations for understory woody
plants in eastern Texas. Texas Journal of
Hutchison, M. 2003. Vegetation management Science 40(1): 49-53.
guideline: smooth sumac (Rhus glabra). Illinois
nature Preserves Commission, Belknap IL. Rowe, D.B. and F.A. Blazich. 2003. Rhus L.,
http://www.inhs.uiuc.edu/VMG/smsumac.html. sumac. In: F.T. Bonner and R.G. Nisley, eds.
5 p. Woody plant seed manual. U.S. Department of
Agriculture, Forest Service, Washington, DC.
Institute of Systematic Botany. 2003. Atlas of http://wpsm.net.Rhus.pdf. 16 p.
Florida plants: Rhus glabra. http://www.
plantatlas.usf.edu/main.asp?plantID=3941. 2 p. Saxena, G., A.R. McCutcheon, S. Farmer, G.H.N.
Towers, and R.E.W. Hancock. 1994.
Johnson, F.D. 1995. Wild trees of Idaho. Antimicrobial constituents of Rhus glabra.
University of Idaho Press, Moscow, ID. 212 p. Journal of Ethnopharmacology 42(2): 95-99.
Kearney, T.H. and R.H. Peebles. 1951. Arizona Strauss, S.Y. 1991. Direct, indirect, and
flora. University of California Press, Berkeley cumulative effects of three native herbivores on
and Los Angeles, CA. 1,032 P. a shared host plant. Ecology 72(2): 543-558.
Li, X., J.M. Baskin, and C.C. Baskin. 1999. Wyman, D. 1969. Shrubs and vines for American
Anatomy of two mechanisms of breaking gardens. The Macmillan Company, London.
physical dormancy by experimental treatments 613 p.
630
Rhus microphylla Engelm. ex Gray little-leaf sumac
ANACARDIACEAE
James E. Nellessen
631
also have physiological dormancy (Baskin and University of New Mexico Press, Albuquerque.
Baskin 2001). There is a high proportion of 244 p.
species with physical dormancy in savannas, hot
deserts, and similar climates with distinct dry and Emmerich, W.E., J.D. Helmer, K.G. Renard, and
wet seasons, the wet season being the season L.J. Lane. 1984. Fate and effectiveness of
favorable for germination. The impermeable tebuthiuron applied to a rangeland watershed. J.
endocarp in the genus Rhus may have evolved as of Environmental Quality 13(3): 382-386
long ago as the Middle Eocene epoch (Baskin and
Baskin 2001). Epple, A.O. 1995. A Field Guide to the Plants of
Arizona. Falcon Publishing Inc., Helena, MT.
Growth and Management.—Little-leaf sumac is 347 p.
not one of the faster growing shrubs. It produces
numerous stems and resprouts readily. Great Plains Flora Association. 1986. Flora of the
Tebuthiuron, an herbicide often used for brush Great Plains. University Press of Kansas.
control on rangelands, proved effective in little- 1,392 p.
leaf sumac control (Emmerich and others 1984).
Tebuthiuron was applied at 0.84 kg/ha to Kearney, T.H., R. Peebles, and Collaborators.
rangeland in both pelleted and dissolved forms. 1951 with 1960 supplement. Arizona Flora.
After 21 months there was a 100 percent kill of University of California Press, Berkeley, CA.
little-leaf sumac, tarbush (Flourensia cernua DC.), 1,085 p.
and four-wing saltbush [Atriplex canescens
(Pursh) Nutt.], all dominant species on the test site. Killingbeck, K.T. and W.G. Whitford. 2001.
Tebuthiuron moved to a soil depth of 15 cm within Nutrient resorption in shrubs growing by design,
8 months, and after 21 months 38 percent of the and by default in Chihuahuan Desert arroyos.
application was remaining, while only 0.47 percent Oecologia, Berlin 128(3): 351-359.
was detected in runoff water.
Kozma, J.M. and N.E. Mathews 1997. Breeding
Benefits.—The fruits are readily eaten by many bird communities and nest plant selection in
birds. Little-leaf sumacs are also important nesting Chihuahuan desert habitats in south-central New
sites for birds. The fruits may be steeped in warm Mexico. Wilson Bulletin 109(3): 424-436.
or cold water to make soothing tea-like drinks that
can sooth sore throats (Bowers and Wignall 1993, Li, Xiaojie, J.M. Baskin, and C.C. Baskin. 1999.
Warnock 1974). Seed morphology and physical dormancy of
several North American Rhus species
References (Anacardiaceae). Seed Science Research 9(3):
247-258.
Baskin, C.C. and J.M. Baskin. 2001. Seeds:
Ecology, Biogeography, and Evolution of Martin, W.C. and C.E. Hutchins. 1980-1981
Dormancy, and Germination. Academic Press, (reprinted 2001). A Flora of New Mexico. 2
San Diego. 666 p. volumes. Bishen Singh Mahendra Pal Singh,
India and Koeltz Scientific Books, Germany.
Bowers, J.E. and B. Wignall. 1993. Shrubs and 2,591 p.
Trees of the Southwest Deserts. Southwest
Parks and Monuments Association, Tucson, AZ. McPherson, G.R., H.A. Wright, and D.B. Wester.
140 p. 1988. Patterns of shrub invasion in semiarid
Texas USA grasslands. American Midland
Carter, J.L. 1997. Trees and Shrubs of New Naturalist 120(2): 391-397.
Mexico. Johnson Books, Boulder, CO. 534 p.
Warnock, B.H. 1974. Wildflowers of the
Correll, D.S. and M.C. Johnston. 1970. Manual of Guadalupe Mountains and the Sand Dune
the Vascular Plants of Texas. Texas Research Country, Texas. Sul Ross State University,
Foundation, Renner, TX. 1,881 p. Alpine, TX. 176 p.
633
Rhus trilobata Nutt. oakleaf sumac
ANACARDIACEAE
634
populations is improved by a 0.3 to 2 hour in the United States. Agriculture Handbook 450.
sulphuric acid scarification (Babb 1959, Brinkman U.S. Department of Agriculture, Forest Service,
1974, Glazebrook 1941, Heit 1968), while 30 to Washington, DC. p. 715-719.
120 days of moist prechilling is required to release
embryo dormancy (Babb 1959, Swingle 1939). Clark, J.W. and J.E. Depuit. 1981. Analysis of
Mechanically rupturing the seedcoat improves direct seeding methods for establishment of
germination (McKeever 1938), but techniques selected native shrub species on minesoils in
have not been developed for treating large seed southeastern Montana. In: L.H Stelter, E.J.
lots. Following fire, oakleaf sumac generally DePuit, and S.A. Mikol, tech. coords. Shrub
sprouts vigorously from the root crown and establishment on disturbed arid and semi-arid
rhizomes (Wasser 1982). lands. Wyoming Game and Fish Department,
Cheyenne, WY. p. 89-103.
Growth and Management.—Local seed sources
should be selected for range seedings or Glazebrook, T.B. 1941. Overcoming delayed
transplanting projects as the range of adaptability germination in the seed of plants valuable for
for most populations is not known (Swenson erosion control and wildlife utilization. Master’s
1957). Scarified seed may be fall planted to thesis, University of Idaho, Moscow, ID. 97 p.
provide overwinter moist prechilling. Seed should
be planted approximately 1.3 cm deep or slightly Goodrich, S. and E. Neese. 1986. Uinta Basin
deeper in dry, coarse textured soils. Seedlings are flora. U.S. Department of Agriculture, Forest
considered to be drought and cold hardy but may Service, Intermountain Region, Ogden, UT. 319
succumb to competition from herbaceous species p.
when young (Clark and Depuit 1981, Monsen
1987). Range or wildland seedings normally Harrington, H.D. 1964. Manual of the plants of
require 2 to 5 years for establishment (Plummer Colorado. The Swallow Press, Inc., Chicago, IL.
and others 1968). 666 p.
Benefits.—Oakleaf sumac can be used to stabilize Heit, C.E. 1968. Thirty-five years’ testing of tree
disturbed riparian areas. It provides excellent and shrub seed. Journal of Forestry 66: 632-634.
ground cover, and the spreading root system
reduces soil erosion. It is particularly useful for Hitchcock, C.L., A. Cronquist, M. Ownbey, and
restoration of meadows and other riparian sites J.W. Thomson. 1961. Vascular plants of the
where extensive erosion and gully cutting has Pacific Northwest. Part 3: Saxifragaceae to
lowered water tables. Oakleaf sumac thickets Ericaceae. University of Washington Press,
provide excellent cover for many small birds and Seattle, WA. 614 p.
mammals, and winter browse where shrubs are
limited. It is increasingly being used by LHBH [Liberty Hyde Bailey Hortorium]. 1976.
homeowners seeking drought-tolerant native Hortus third: a concise dictionary of plants
species since it is commercially available, easy to cultivated in the United States and Canada.
grow, long-lived, responds well to pruning, and Third Edition, Macmillan Publishing, New
has attractive red, orange to yellow fall color. York. 1,290 p.
Native Americans have used oakleaf sumac for
food, tobacco substitute, basketry, and as a McKeever, D.G. 1938. The effects of various
mordant in dyeing (LHBH 1976). methods of treatment on germination of seeds of
some plants valuable for game and erosion
References purposes. Master’s thesis, University of Idaho,
Moscow, ID. 128 p.
Babb, M.F. 1959. Propagation of woody plants by
seed. Bulletin 26. Alaska Agricultural Monsen, S.B. 1987. Shrub selections for pinyon-
Experiment Station. [Place of publication juniper plantings. In: R.L. Everett (comp.).
unknown]. 12 p. Pinyon-juniper conference. INT-GTR-215. U.S.
Department of Agriculture, Forest Service,
Brinkman, K.A. 1974. Rhus L. Sumac. In: C.S. Intermountain Research Station, Ogden, UT. p.
Schopmeyer, tech. coord. Seeds of woody plants 316-334.
635
Plummer, A.P., D.R. Christensen, and S.B. USDA Forest Service. 1937. Range plant
Monsen. 1968. Restoring big game range in Handbook. U.S. Department of Agriculture,
Utah. Publication 68-3. Utah Division of Fish Forest Service, Washington, DC. Republished
and Game, Salt Lake City, UT. 183 p. by Dover Publications, London. 816 p.
Shaw, N. 1984. Producing bareroot seedlings of van Dersal, W.R. 1938. Native woody plants of the
native shrubs. In: P.M. Murphy, comp. The United States: Their erosion-control and wildlife
challenge of producing native plants for the values. Miscellaneous Publication 303. U.S.
Intermountain area. INT-GTR-168. U.S. Department of Agriculture, Washington, DC.
Department of Agriculture, Forest Service, 302 p.
Intermountain Forest and Range Experiment
Station, Ogden, UT. p. 6-15. Vories, K.C. 1981. Growing Colorado plants from
seed: a state of the art. Vol. 1: Shrubs. INT-
Swenson, W.S. 1957. Squawbush in windbreaks in GTR-103. U.S. Department of Agriculture,
eastern Colorado. Journal of Soil and Water Forest Service, Intermountain Forest and Range
Conservation 12: 184-185. Experiment Station, Ogden, UT. 80 p.
Swingle, C.F. 1939. Seed propagation of trees, Wasser, C.H. 1982. Ecology and culture of
shrubs, and forbs for conservation planting. selected species useful in revegetating disturbed
SCS-TP-12. U.S. Department of Agriculture, lands in the West. FWS/OBS-82/56. U.S.
Soil Conservation Service, Washington, DC. Department of the Interior, Fish and Wildlife
198 p. Service, Office of Biological Services, Western
Energy and Land Use Team, Washington, DC.
Thornburg, A.A. 1982. Plant materials for use on 347 p.
surface mined land in arid regions. T.P. 157,
EPA 600-7-79-133. U.S. Department of Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Agriculture, Soil Conservation Service, Higgins, eds. 1987. A Utah flora. Great Basin
Washington, DC. 88 p. Naturalist Memoir 9. Brigham Young
University, Provo, UT. 894 p.
636
Ribes aureum Pursh golden currant
GROSSULARIACEAE
637
approximately 400 to 2,800 m (Cronquist and and plants can sprout after fire (Tykač 1990,
others 1997, Carter 1997, Powell 1998). Golden Marshal 1995). Tolerance to fire in the dormant
currant grows in full sunlight but also does well in state is good (Wasser 1982). However, the
partial shade (Tykač 1990). It is tolerant to a range severity, or heat, of the fire is likely to be
of weakly acid (pH 6) to weakly basic soils important to survival, and very hot fires can kill
(Wasser 1982). Bacterial diseases have been noted shrubs completely (Marshall 1995).
to infect canes of some genotypes, but resistant
varieties are available (Wasser 1982). Ribes Benefits.—Golden currant rates from “poor” and
species are intermittent hosts of the white pine “fair” to “medium” palatability for cattle (Dayton
blister rust (Cronartium ribicola), a fungal 1931, USDA FS 1988). It has good palatability for
pathogen that can be ruinous to most native white sheep (Dayton 1931). In Montana, twigs and
pine species. Because of this, Ribes species have foliage can make up to 2 to 5 percent of a sheep’s
been subject to specific eradication programs in diet (Martin and others 1951). The fruits are an
many white pine (Pinus monticola Dougl. ex D. important food source for a variety of wildlife
Don) regions (Martin and others 1951, Van such as songbirds, chipmunks, mice, and ground
Arsdel and others 1998). However, the squirrels (Martin and others 1951, Marshall 1995,
significance of specifically golden currant in the Tykač 1990). The fruits may contribute up to 10
life cycle and pathology of this rust is uncertain. percent of a chipmunk and ground squirrels’ diet
Throughout the Sacramento Mountains of New (Martin and others 1951). Both fruit and foliage
Mexico, R. pinetorum Greene was a major are used by a range of species that include grouse,
contributor to the spread of rust to nearby pines, coyote, and beaver (Martin and others 1951). Elk
but blister rust was not found on golden currant have been reported to eat the foliage in Colorado,
that generally grew below the elevation of the and one report indicated that mule deer browsed it
pines (Van Arsdel and others 1998). infrequently, but cattle not at all, during a 1-year
study in Montana (Dusek 1975). Golden currant
Reproduction.—Golden currant generally flowers has been planted for erosion control and in
in June through August, although flowers may be restoration projects (Vines 1986, Wasser 1982). It
found from March until the first frost (Epple 1995, is particularly used for restoration of game habitat
McGregor and others 1986, USDA FS 1988, Vines (Wasser 1982). When golden currant is used in
1986). The fragrant flowers are insect pollinated. vegetation restoration projects it is usually a small
Some insects have adapted to the long narrow component of the seed mix and used at
flower tube by boring a hole at its base to “steal” approximately 461 to 922 grams per hectare
the nectar (Derig and Fuller 2001). Golden currant (Wasser 1982). The berries, both fresh, dried, and
readily crosses with other Ribes species (Wasser made into jams and cakes, have been used for food
1982). The fruits mature in summer and fall. The by many Native American tribes as well as by
seed is a densely hairy achene. The cleaned seed European settlers (Elmore 1976, Moerman 1998,
averages approximately 478,398 to 513,672 Welsh and others 1993). It has also been used as a
individuals per kg with a germination rate of 60 to pharmaceutical by some of the tribes of the Great
98 percent (Vines 1986, Wasser 1982). The seeds Plains. The dried, pulverized inner bark was
undergo a physiological dormancy that is broken sprinkled on sores, used with other ingredients in a
by 60 days of cold stratification (Baskin and poultice, and a decoction was taken for leg
Baskin 2001). Germination is epigeal. The plant swelling (Moerman 1998). The flowers are edible,
also spreads by growing shoots from rhizomes apparently flavorful, and eaten by gourmets
(Wasser 1982). (Elmore 1976). Golden currant is a popular
ornamental plant in North America, Europe, and
Growth and Management.—Golden currant is parts of the former Soviet Union (Bailey and
easily propagated from hardwood cuttings in June others 1976, Komarov 1939, Tykač 1990).
or September and bare rooted plants are best
planted when leafless in the spring or fall (Tykač References
1990). When starting plants from seed, fall
seeding is preferred, but stratified seed can be Bailey, L.H., E.Z. Bailey, and the staff of the
planted in the spring (Wasser 1982). Golden Liberty Hyde Bailey Hortorium. 1976. Hortus
currant is used as a rootstock for grafting on other Third–a concise dictionary of plants cultivated
species of currant that root poorly (Tykač 1990). in the United States and Canada. Macmillian
Cutting back old shoots stimulates new growth, Publishing Company, New York. 1,290 p.
638
Baskin, C.C. and J.M. Baskin. 2001. Seeds, Kartesz, J.T. 1994. A synonymized checklist of the
ecology, biogeography, and evolution of vascular flora of the United States, Canada and
dormancy and germination. Academic Press, Greenland. Vol.1 - Checklist. 2nd Ed. Timber
New York. 666 p. Press, Portland, OR. 622 p.
Carter, J.L. 1997. Trees and shrubs of New Kershaw, L., A. MacKinnon, and J. Pojar. 1998.
Mexico. Johnson Books, Boulder, CO. 534 p. Plants of the Rocky Mountains. Lone Pine
Publishing, Edmonton, Canada. 383 p.
Cronquist, A., N.H. Holmgren, and P.K.
Holmgren. 1997. Intermountain flora, vascular Marshall, K.A. 1995. Ribes aureum. In:
plants of the Intermountain West, U.S.A. Vol. Department of Agriculture, Forest Service,
3, Part A. New York Botanical Garden, Bronx, Rocky Mountain Research Station, Fire
NY. 446 p. Sciences Laboratory. Fire Effects Information
System, http://www.fs.fed.us/ database/feis/.
Dayton, W. A. 1931. Important western browse 13 p.
plants. Miscellaneous Publication 101. U.S.
Department of Agriculture, Washington DC. Martin, A C., Zim, H. S., and Nelson, A.L. 1951.
214 p American wildlife and plants. 2nd printing.
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371 p.
639
Van Arsdel, E.P., D.A. Conklin, J.B. Popp, and Wasser, C.H. 1982. Ecology and culture of
B.W. Geils. 1998. The distribution of white selected species useful in revegetating disturbed
pine blister rust in the Sacramento Mountains of lands in the West. FWS?OBS-82/56. Fish and
New Mexico. Finnish Forest Research Institute Wildlife Service, U.S. Department of the
Research Papers 712: 275-283. Interior, Washington, DC. 347 p.
Vines, R.A. 1986. Trees, shrubs, and woody vines Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
of the Southwest. University of Texas Press. Higgins. 1993. A Utah flora. 2nd Edition,
Austin, TX. 1,104 p. Revised. Brigham Young University, Provo,
UT. 986 p.
640
Ribes cereum Dougl. wax current
GROSSULARIACEAE
John K. Francis
641
Growth and Management.—Fruits must Brayshaw, T.C. 1996. Trees and shrubs of British
generally be picked by hand. Seeds are cleaned of Columbia. UBC Press, Vancouver, B.C.,
fruit by maceration, and air dried. They can be Canada. 373 p.
stored in sealed containers at ambient temperatures
or refrigerated (Pfister 1974). Seeds require 4 to 5 Britton, N.L., and A. Brown. 1913. Illustrated flora
months of cold stratification (at temperatures near of the northern states, Canada, and the British
freezing) after which they can be planted in the possessions. Vol. 2, 2nd Ed. Scribner, New York.
spring (Marshall 1995). Alternately, they can be 735 p.
sown in the fall and allowed to stratify naturally.
Plants can also be started from cuttings of semi- Marshall, K.A. 1995. Ribes cereum. In: U.S.
hardened wood (Plants For a Future 2003). Adult Department of Agriculture, Forest Service,
plants can be eliminated from stands by fire, Rocky Mountain Research Station, Fire Sciences
grubbing, and probably herbicides. However, in at Laboratory, Fire Effects Information System.
least one study, because many new seedlings http://www.fs.fed.us/database/feis/plants/shurb/
appeared, the number of plants increased in sites ribcer/all.html. 14 p.
disturbed by treatments (Marshall 1995). Because
wax current does poorly in shade, tree planting is Mesler, M.R. and J.O. Sawyer, Jr. 2003.
suggested as a long-term solution. Grossulariaceae, gooseberry family. http://
ucjeps.berkeley.edu/chi-bin/get_JM_treatment.
Benefits.—Wax current contributes to the beauty pl?4451,4452,4463. 3 p.
of forest lands where it grows and helps protect the
soil from erosion. Shade from their crowns shelters Natural Resources Conservation Service. 2003.
conifer seedlings and thus aids forest succession Plants profile: Ribes cereum Doul.
(Marshall 1995). The species is used to a limited http://plants.usda.gov/cgi_bin/plant_profile.chi?
extent as a foundation plant in formal landscaping symbol=RICE. 3 p.
(Wildland Nursery 2003). Wax current is fair to
poor browse for domestic livestock and wild Oehlke, B. 2003. Agapema homogena moth, Dyar,
ungulates and is mainly eaten when little else is 1908. http://www3.islandtelecom.com/
available. New annual growth contains about 5.6 ~oehlkew/sahomoge.htm. 4 p.
percent protein. Several species of birds and
rodents consume the fruits (Marshall 1995). The Pfister, R.D. 1974. Ribes L., current, gooseberry.
larva of the Rocky Mountain Agepema moth In: C.S. Schopmeyer, tech. coord. Seeds of
(Agapema homogena Dyar) feed on the foliage woody plants in the United States. Agriculture
(Oehlke 2003). Wax current berries are edible but Handbook 450. U.S. Department of Agriculture,
have little flavor. They are picked for making Forest Service, Washington, DC. p. 720-727.
jams, jellies, and pies and were formerly used by
Native Americans for making pemmican (Marshall Plants For a Future. 2003. Database search results:
1995). People of the Secwepemc tribe of British Ribes cereum. http://www.ibiblio.org/pfaf/cgi-
Columbia ate them to relieve diarrhea (Secwepemc bin/arr_html?Ribes+cereum&CAN=LATIND.
Nation 2003). An infusion of inner bark was used 5 p.
as a wash for sore eyes (Plants For a Future 2003).
Wax current and other Ribes species are alternate Secwepemc Nation. 2003. Secwepemc
hosts for white pine blister rust (Cronartium ethnobotanical garden: other plants. Sccwepemc
ribicola), which can be lethal to five-needle pines. Nation, Kamloops, BC, Canada. 4 p.
Efforts to eradicate Ribes from several forest areas
failed and did not decrease the incidence of blister Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
rust (Marshall 1995). Higgins. 1987. A flora of Utah. Great Basin
Naturalist Memoirs 9. Brigham Young
References University Press, Provo, UT. 894 p.
Abrams, L. 1944. Illustrated flora of the Pacific Wildland Nursery. 2003. Wildland nursery:
States. Vol. 2. Stanford University Press, shrubs. Wildland Nursery, Joseph, UT.
Stanford, CA. 635 p. http://www.wildlandnursery.com/shrubs/shrubs
M-Z.html. 6 p.
642
Ricinus communis L. castor bean
EUPHORBIACEAE
Synonyms: none
John K. Francis
found growing naturally and in cultivation as an
ornamental throughout the tropical and subtropical
areas of the United States and its territories.
643
and may reach 6 m in height (Kadambi and Dabral and horses are especially vulnerable. Fatal doses
1955). Annual varieties reach 1 to 2 m in height. are from 2.5 to 6 seeds for humans and about 6
Seed yields under cultivation vary from 200 to seeds for horses (CISR 1972). Symptoms are
1,700 kg/ha, depending on variety and site quality stomach irritation, diarrhea, abdominal pain,
(CISR 1972). increased heart rate, profuse sweating, collapse,
and convulsions. Broken seeds can cause skin
Benefits.—Castor beans contain from 31 to 61 irritation. The foliage is only slightly toxic
percent oil (CISR 1972). After decorticating, they (Anonymous 2000). It is advisable to completely
are subject to a series of hot or cold presses eliminate castor bean from pastures, especially
followed by solvent extraction, each step yielding a horse pastures, and pinch off flowers of ornamental
different grade of oil (Kirschenbauer 1960). The plants to prevent possible poisoning of children.
largest producers are Brazil and India. The
principal consumer is the United States References
(Encyclopedia Brittanica 2000), but an annual
demand for 100,000 tons of castor oil is reported Anonymous. 2000. Castorbean, castor oil plant.
for Europe (NF-2000 Database 2000). The seed http://www.vet.perdue.edu/depts/addl/toxic/
cake remaining after extracting the oil is used as plant11. htm. 1 p.
fertilizer or cooked to destroy the toxin and
incorporated into animal feeds. Not as popular as it CSIR. 1972. The wealth of India. Raw materials.
once was, castor oil is still widely used in Vol. 9. Publications & Information Directorate,
traditional and herbal medicine, especially in less Council for Scientific and Industrial Research,
developed countries. Its principal use in medicine New Delhi. 472 p.
is as a purgative and laxative. Castor oil is also
used as a lubricant, lamp fuel, a component of Encyclopedia Britanica. 2000. Castor oil.
cosmetics, and in the manufacture of soaps, http://www.Britanica.com/bcom/eb/article/4/0,57
printer’s ink, plastics, fibers, hydraulic fluid, break 16,2105+1+20724,00htm?query=castoroil%20
fluid, varnishes, paints, embalming fluid, textile oil. [not paged].
dyes, leather finishes, adhesives, waxes, and
fungicides (Encyclopedia Brittanica 2000, CISR Kadambi, K. and S.N. Dabral. 1955. The
1972). It is gradually being replaced as a raw silviculture of Ricinus communis Linn. Indian
material for some of the uses by petroleum-based Forester 81(1): 53-58.
products. In India, the leaves are used as food for
eri silk worms (Kadambi and Dabral 1955). The Kirschenbauer, H.G. 1960. Fats and oils, an
stalks from fields are burned for fuel in India and outline of their chemistry and technology.
have been shown to be suitable for short-fiber pulp Reinhold Publishing Corporation, New York.
(Kadambi and Dabral 1955). The species has been 240 p.
planted for dune stabilization (CISR 1972). Castor
bean is widely planted as an ornamental. Its large, Little, E.L., Jr., R.O. Woodbury, and F.H.
star-shaped leaves make it a bold foliage plant. Wadsworth. 1974. Trees of Puerto Rico and the
Some varieties have red- or purplish-colored leaves Virgin Islands. Vol. 2. Agriculture Handbook.
and stems. 449. U.S. Department of Agriculture, Forest
Service, Washington, DC. 1,024 p.
Detrimental Effects.—Castor bean may become a
weed in neglected cropland and pasture. It is not NF-2000 Online Database Information Items.
difficult to control through cultivation and mowing. 2000. Crops: castor oil (Ricinus communis).
Of greater concern than its weedy potential is the http://www.nf-2000.org/secure/Crops/S600.htm
high toxicity of its seeds, which contain ricin, a [not paged].
water-soluble protein. Even a small amount of
masticated seed is likely to cause death. Humans
644
Rondeletia inermis (Spreng.) Krug & Urban cordobancillo
RUBIACEAE
John K. Francis
645
on plantation establishment or management of References
natural stands.
Liogier, H.A. 1997. Descriptive flora of Puerto
Benefits.—Cordobancillo helps protect the soil, Rico and adjacent islands. Vol. 5. Editorial de la
furnishes cover for wildlife, and adds to the Universidad de Puerto Rico, Río Piedras, PR.
aesthetics of Puerto Rican forests. Too small for 436 p.
lumber or poles, the wood is occasionally used for
fuel. Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
646
Rondeletia pilosa Sw. cordobancillo peludo
RUBIACEAE
John K. Francis
647
Benefits.—Cordobancillo peludo contributes to Universidad de Puerto Rico, Río Piedras, PR.
the aesthetics of the forest, it helps protect the soil, 436 p.
and furnishes cover for wildlife. The wood is
useful for small-diameter fuel. Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
References Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Liogier, H.A. 1997. Descriptive flora of Puerto Washington, DC. 1,024 p.
Rico and adjacent islands. Vol. 5. Editorial de la
648
Rosa woodsii Lindl. Wood’s rose
ROSACEAE
Synonyms: Rosa fendleri Crep
Rosa californica Watson.
Rosa macounii Greene
Rosa chrysocarpa Rydb.
Bruce L. Welch
649
slopes, and on sandhills throughout the prairies and food for a number of wildlife species, and
and on riverbanks and clearings in boreal and watershed protection. Fruits, leaves, or twigs are
subalpine forests. It can be found on the eastern eaten by a number of birds and mammals. These
slopes of the Rocky Mountains and throughout the include pronghorn, elk, mule deer, white-tailed
Great Basin. It is found along roadsides and deer, porcupines, beavers, squirrels, coyotes, bear,
south-facing cutbanks (Pacific Southwest small nongame birds and upland game birds
Experiment Station. 2002). In the Great Basin, (Pacific Southwest Experiment Station 2002).
Wood’s rose prefers moist sites along streams, in Livestock also eat Wood’s rose. Crude protein
seepage areas along fences, irrigation canals, content of Wood’s rose leaves vary from 5.7 (fall)
marsh lands, lake shores, and hillsides in to 16.4 (spring) percent of dry matter and stems
palustrine and lacustrine habitats (Welsh and from 5.4 (winter) to 12.0 (spring) percent (Dietz
others 1987). When conditions are favorable, 1972). Phosphorus content of leaves varied from
nearly impenetrable thickets of Wood’s rose are 0.29 (fall) to 0.48 (spring) percent of dry matter
formed along some mountain streams (Pacific with stems varying from 0.12 (winter) to 0.32
Southwest Experiment Station 2002). It flourishes (spring) percent (Dietz 1972). In vitro digestible
in moderate shade to full sunlight and so makes for Wood’s rose was 65.9 percent of dry matter
good growth on all aspects (Blauer and other digested for spring tissues and 54.5 percent for
1975). Wood’s rose is adapted to a wide range of winter tissues (Dietz 1972). These in vitro
soil types and textures (Pacific Southwest digestible values would rank Wood’s rose as a
Experiment Station 2002). Growth is generally highly digestible spring and winter forage (Welch
best on moderately fertile, well-drained clay loam, 1989, Welch and Andrus 1977). The dense
sandy loam, or sandy soils. It is tolerant of thickets formed by Wood’s rose are used for
moderately acid to weakly basic soils. (Pacific nesting and escape cover by many birds and small
Southwest Experiment Station 2002). It is seldom mammals (Pacific Southwest Experiment Station
found where the average annual precipitation is 2002). The extensive rhizomes and good
less than 260 mm (Blauer and others 1975). survivability and revegetation characteristics
makes this species an effective tool in erosion
Reproduction.—Seeds are produced on 2 to 5 control (Pacific Southwest Experiment Station
year old plants. Seeds of Wood’s rose are ingested 2002). Good results have been obtained with
with the hip and dispersed in the dropping of birds rooting of hardwood cuttings, direct seeding, and
and mammals. The seeds have a seed coat transplanting trials (Pacific Southwest Experiment
dormancy and require warm or cold stratification. Station 2002). Native Americans made extensive
Seed viability ranged from 44 to 64 percent for use of Wood’s rose roots, stems, leaves, flowers,
seed collected in northern Alberta. The seeds and hips for foods and therapeutic purposes. The
remain viable for 2 to 5 years. Seeds will hips are a source of vitamin C and are dried for
germinate within 30 to 40 days (Blauer and others use in flavoring teas, jellies, fruitcakes, and
1975, Gill and Pogge 1974, Pacific Southwest puddings. Wood’s rose is used as an ornamental
Experiment Station 2002). Wood’s rose spreads near homes to attract birds and other wildlife
vegetatively through underground rhizomes, (Pacific Southwest Experiment Station 2002).
sprouting from the root crown, and by layering
(Blauer and other 1975, Pacific Southwest References
Experiment Station 2002).
Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
Management.—Wood’s rose has a fairly high Stevens, and B.C. Giunta. 1975. Characteristics
tolerance to browsing (Pacific Southwest and hybridization of important intermountain
Experiment Station 2002). It is susceptible to shrubs. I. Rose family. Research Paper INT-
various leaf spots, as well as to leaf rusts, gray 169. U.S. Department of Agriculture, Forest
mold, powdery mildew, common gall, and stem Service, Intermountain Forest and Range
cankers (Pacific Southwest Experiment Station Experiment Station, Ogden, UT. 36 p.
2002). The principal forest insect pests are tent
caterpillar, rose leaf hopper, and a subspecies of Cronquist, A., N.H. Holmgren, and P.K.
western tussock moth (Pacific Southwest Holmgren. 1997. Intermountain flora: vascular
Experiment Station 2002). plants of the Intermountain west, U.S.A. Vol. 3.
Part A; Subclass Rosidae (except Fabales). The
Benefits.—Wood’s rose adds to the biodiversity New York Botanical Garden., New York. 446 p.
of a multitude of ecosystems. It provides habitat
650
Dietz, D. R. 1972. Nutritive value of shrubs. In: Welch, B.L. 1989. Nutritive value of shrubs. In: C.
C. M. McKell, J. P. Blaisdell, and J. R. Goodin, M. McKell, ed. The biology and utilization of
tech. eds. Wildland shrubs–their biology and shrubs. Academic Press, Inc., San Diego, CA.
utilization, An international symposium; p. 405-424.
Proceedings; 1971 July; Logan, UT. General
Technical Report INT-1. U.S. Department of Welch, B.L. and D. Andrus. 1977. Rose hips-a
Agriculture, Forest Service, Intermountain possible high-energy food for wintering mule
Forest and Range Experiment Station, Ogden, deer? Research Note INT-221. Intermountain
UT. p. 289-302. Forest Experiment Station, U.S. Department of
Agriculture, Forest Service, Ogden, UT. 6 p.
Gill, J.D. and F.L. Pogge. 1974. Rosa L. Rose In:
C.S. Schopmeyer, tech. coord. Seeds of woody Welsh, S.L, N.D. Atwood, S. Goodrich, and L.C.
plants in the United States. Agric. Handbook Higgins. 1987. A Utah flora. Great Basin
654. U.S. Department of Agriculture, Forest Naturalist Memoirs. No. 9. Brigham Young
Service, Washington, DC. p. 732-737. University, Provo, UT. 894 p.
651
Rubus discolor Weihe & Nees Himalayan blackberry
ROSACEAE
John K. Francis
652
Reproduction.—Himalayan blackberry ecosystems in places such as Hawaii (Starr and
commonly flowers from June to August. The others 2003). Plants can be eliminated by grubbing
flowers are intensely visited by honey bees, but with follow-up treatment a year later, repeated
insect pollination is not necessary for seed mowing, and by pasturing with goats (Cox 2003).
production. In fact, sexual reproduction may be Several herbicides are effective using foliar spray,
rare; 17 samples from Himalayan blackberry stem injection, treatment of cut stumps, and basal
plants collected throughout Australia proved to spray methods (Starr and others 2003, Tirmenstein
have no genetic variation (Evans and others 1998). 1989). Dead canes accumulate in older stands and
Good seed crops are produced nearly every year are as much a nuisance to remove as the plants
(Tirmenstein 1989). There are 324,000 cleaned themselves. Because of dormant seeds in the soil,
seeks/kg. The seeds are dispersed by birds and checking and retreatment for many years may be
mammals as well as by gravity. Passing through required to completely eliminate the species.
birds and mammals is reported to increase
germination by 30 percent in the first year Benefits.—Himalayan blackberry provides food
(Bruzzese 1998). Seeds in the soil seed bank and cover for many mammals and birds. It is little
remain viable for several years (Brinkman 1974). used by domestic livestock except for goats.
Stands in Victoria, Australia have been estimated However, deer, elk, rabbits, porcupines, beavers,
to produce 7,000 to 13,000 seeds/m2/year and mountain beavers consume leaves, buds,
(Bruzzese 1998). Himalayan blackberry seedlings twigs, and cambium, especially during the winter
commonly appear after fires or disturbance that months. A large number of species consume the
exposes the soil and allows sunlight to reach the fruits and a number rely on the thickets of stems
surface. Once seedlings become established, most for escape and reproductive cover (Tirmenstein
subsequent reproduction is vegetative. Plants 1989). Himalayan blackberry fruits are among the
reproduce by sprouts from rhizomes and by most delicious of wild fruits but are difficult to
layering (rooting) at the nodes when stems come in pick because of the spines. They are eaten fresh,
contact with the ground. canned, used to flavor ice cream, and made into
pies, jams, jellies, juices, and wines. Some people
Growth and Management.—Canes do not bloom object to the seeds getting caught in one’s teeth
during their first year in which they make most of when eaten whole, but it is a minor inconvenience
their growth (2 to 8 m of elongation). Canes bloom that can be avoided by juicing the fruit. In spring,
and fruit in their second and sometimes third years the succulent canes can be pealed and eaten, fresh
and die at the end of their second or third years. or cooked (Jacobson 2001). The brambles are
Individual root crowns live a maximum of 7.5 allowed to grow over fences and trellises to create
years in Australia (Bruzzese 1998). By suckering barriers almost impenetrable to people and
from rhizomes and layering, plants (clones) can livestock. While the species does protect the soil
endure almost indefinitely. Live biomass of a stand from erosion and helps revegetate disturbed sites,
in Victoria, Australia totaled 3.1 tons/ha (59 the thickets and mounds of brambles restrict
percent above-ground and 41 percent below- movement of hikers and woods workers, and
ground) and 27 tons of dead canes and litter suppress other vegetation and slows succession.
(Bruzzese 1998). Scarification is required for
prompt germination. Although several methods References
will work, soaking in concentrated H2SO4 for 50 to
60 minutes followed by 90 days of cold Brayshaw, T.C. 1996. Trees and shrubs of British
stratification is recommended. About 33 percent Columbia. UBC Press, Vancouver, BC, Canada.
germination in about 70 days can be expected. 373 p.
Sowing in late summer for spring germination can
substitute for artificial cold stratification. Sowing Brinkman, K.A. 1974. Rubus L., blackberry,
depth is 3 to 9 mm. (Brinkman 1974). The species raspberry. In: C.S. Schopmeyer, tech. coord.
can be propagated by digging up and replanting Seeds of woody plants in the United States.
suckers. However, many more plants can be Agriculture Handbook 450. U.S. Department of
started by using root cuttings. Root pieces between Agriculture, Forest Service, Washington, DC.
3 and 13 mm in thickness and 10 to 18 cm long are p. 738-743.
recommended. They are grown for 1 year in the
nursery bed before outplanting (Shoemaker 1978). Bruzzese, E. 1998. The biology of blackberry in
Himalayan blackberry can be troublesome in many south-eastern Australia. Plant Protection
habitats and is a potential threat to native Quarterly 13(4): 160-162.
653
Ceska, A. 1999. Rubus armeniacus—a correct e&keywordquery=Rubus+discolor&earl=plant_
name for Himalayan blackberries. Botanical search.cgi. 5 p.
Electronic News ISSN 1188-603X. http://www.
ou.edu.cas/botany-micro/ben/ben230.html. 3 p. Shoemaker, J.S. 1978. Small fruit culture. The
AVT Publishing Company, Inc., Westport, CN.
Cox, C. 2003. Nonchemical methods for removing 357 p.
unwanted blackberry plants. Journal of Pesticide
Reform 23(1): 10-11. Starr, F., K. Starr, and L. Loope. 2003. Plants of
Hawai’i: Rubus discolor. http://www/jear/
Evans, K.J., D.E. Symon, and R.T. Roush. 1998. prg/starr/hiplants/reports/html/rubus_discolor.ht
Taxonomy and genotypes of the Rubus m. 6 p.
fruticosus L. aggregate in Australia. Plant
Protection Quarterly 13(4): 152-156. Tirmenstein, D. 1989. Rubus discolor. In: U.S.
Department of Agriculture, Forest Service,
Jacobson, A.L. 2001. Himalaya blackberry: Rubus Rocky Mountain Research Station, Fire Sciences
armeniacus Focke. http://www.arthurleej.com/ Laboratory, Fire Effects Information System.
a-himalayabb.html. 4 p. http://www.fs.fed.us/database/feis/plants/shurb/
rubdis/all.html. 15 p.
Natural Resources Conservation Service. 2003.
Plants profile: Rubus discolor Weihe & Nees, Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Himalayan blackberry. http://plants.usda.gov/ Higgins. 1987. A Utah flora. Great Basin
cgi_bin/plant_search.cgi?mode=Scientific+Nam Naturalist Memoirs 9. Brigham Young
University Press, Provo, UT. 894 p.
654
Rubus parviflorus Nutt. thimbleberry
ROSACEAE
John K. Francis
655
Wyoming (Tirmenstein 1989). Thimbleberry is spring sowing. Also, scarifying with H2SO4 prior
moderately shade tolerant. Although it is most to the cold stratification may enhance germination.
abundant and grows and fruits best in 60 to 100 Tests have given maximum germination estimates
percent sunlight, the species persists in the of 62 percent (Tirmenstein 1989). Hardwood
understory of closed stands (Tirmenstein 1989). It cuttings can be easily rooted. New plants can also
occurs both as scattered individual plants and as be started from rhizome cuttings and plant
large continuous stands. Habitat includes burned divisions. Wildlings survive well but are slow to
sites, logged areas, avalanche tracks, roadsides, become established (Washington State University
natural shrublands, hardwood forest understories, Cooperative Extension 2003). Occasionally, it is
understories of low to moderate basal area conifer desirable to release conifer seedlings in and under
forests, and river overflow terraces and shorelines. thimbleberry thickets or to make way for other,
It is a part of a large number of different plant more palatable forage species. The species is
communities. Thimbleberry is resistant to fire, moderately susceptible to herbicides. Picloram,
readily resprouting from rhizomes. In fact, the 2,4-D, and glyphosate have been effective in
species generally benefits from all kinds of appropriate applications (Tirmenstein 1989).
disturbance. It usually appears during the first year
after disturbance and often dominates the Benefits.—Thimbleberry is an important shrub of
understories of logged areas within 5 years. the understory and forest openings. It helps protect
Moderate to light thimbleberry stands can serve as the soil and adds to the beauty of the forest. The
a good protective cover for conifer seedlings. In species is occasionally planted as an ornamental
recreational sites, the species showed relatively for its fragrant flowers and brilliant orange to
low resistance to trampling (Tirmenstein 1989). maroon fall foliage (Borialforest.org 2003, Rook
1998). It is also occasionally planted in
Reproduction.—Thimbleberry generally blooms conservation plantings in disturbed areas.
in June and July. Its fruits mature in July and Thimbleberry is relatively low in energy and
August (Borialforest.org 2003). However, at the protein (4 to 8 percent), and is little used by cattle
extremes of its range, it may bloom anywhere and horses and is only fair forage for sheep. It is
from May to September and fruit between late sometimes important for deer, elk, and other wild
June through September (Tirmenstein 1989). The ungulates during the summer while the leaves are
flowers are insect pollinated. Good fruit and seed still present. Rodents consume bark, buds, and
crops are usually produced every year except in foliage to a limited extent. On the other hand, the
high-elevation areas where production may be fruits are an important food item for numerous
unreliable (Tirmenstein 1989). The seeds are wild mammals and birds. Thickets of thimbleberry
dispersed by birds and mammals that eat the fruits, are also important escape, resting, and
and by gravity. The seeds accumulate in the soil reproductive cover for many species of wildlife
and duff to form a seed bank and germinate in (Tirmenstein 1989). Fruits are certainly edible to
great numbers following fire and other humans but reports vary greatly on their
disturbances that remove the forest canopy (Rook palatability (Clark 1976, Welsh 1974). The author
1998). After establishment by seeds, stands has eaten them for years and finds them inferior to
thicken through sprouts from rhizomes (Rook cultivated raspberries (Rubus idaeus L.) but well
1998). worth the trouble of picking while hiking in the
forest. The fruits are made into an excellent jelly
Growth and Management.—The stems (or and were once dried for later use by Native
canes) live for 2 or 3 years. In the first year, canes Americans. The tender young shoots are juicy and
grow in height and develop only leaves. During sweet and can be boiled or eaten fresh. The leaves
the second and third years height growth continues are sometimes made into herb teas
and flowers and fruits are produced. The canes (Borialforest.org 2003, Washington State
then die and are replaced by others that sprout Department of Transportation 2003). Native
from rhizomes. Maximum height of current canes Americans applied poultices of leaves to burns and
is reached within 10 years after establishment wounds and took decoctions of roots as a tonic, for
(Tirmenstein 1989). Seeds must be collected by vomiting, and certain internal disorders (Moerman
hand and should be cleaned of adhering fruit. 1986).
Seeds may be sown in the fall for spring
germination or may be warm (20 to 30 °C for 90
days) and cold stratified (2 to 5 °C for 90 days) for
656
References Tirmenstein, D. 1989. Rubus parviflorus. In: U.S.
Department of Agriculture, Forest Service,
Abrams, L. 1944. Illustrated flora of the Pacific Rocky Mountain Research Station, Fire Sciences
States. Vol. 2. Stanford University Press, Laboratory, Fire Effects Information System.
Stanford, CA. 635 p. http://www.fs.fed.us/database/feis/plants/shurb/
rubpar/all.html. 37 p.
Borialforest.org. 2003. Rubus parviflorus,
thimbleberry. http://www.borialforest.org/ Washington State Department of Transportation.
shrubs/shrub41.htm. 2 p. 2003. Environmental affairs: Rubus parviflorus.
http://www.wsdot.wa.gov/environment/eao/
Clark, L.J. 1976. Wild flowers of the Pacific culres/ethbot/q-s/RubusPar.htm. 2 p.
Northwest. Gray’s Publishing Limited, Sidney,
BC, Canada. 604 p. Washington State University Cooperative
Extension 2003. Native plants: Rubus
Moerman, D.E. 1986. Medicinal plants of Native parviflorus var. parviflorus. http://cahedb.
Americans. Technical Report 19. Museum of wsu.edu/nativePlant/scripts/webDisplayText.asp
Anthropology, University of Michigan, Ann ?ID=nv044. 1 p.
Arbor, MI. 534 p.
Welsh, S.L. 1974. Anderson’s flora of Alaska.
Natural Resources Conservation Service. 2003. Brigham Young University Press, Provo, UT.
Plants profile: Rubus parviflorus Nutt. 724 p.
http://plants.usda.gov/cgi_bin/topics.cgi. 4 p.
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Rook, E.J.S. 1998. Rubus parviflorus, Higgins. 1987. A flora of Utah. Great Basin
thimbleberry. http://www.rook.org.earl/bwca/ Naturalist Memoirs 9. Brigham Young
hature/shurbs/rubuspar.html. 7 p. University Press, Provo, UT. 894 p.
657
Rubus rosifolius Sm. roseleaf raspberry
ROSACEAE
John K. Francis
658
shrubs, and herbs. The species is occasionally Howard R.A. 1988. Flora of the Lesser Antilles,
planted from wildlings and rooted layers for Leeward and Windward Islands.
personal use in gardens. It is controlled along with Dicotyledoneae, Part 1. Vol. 4. Arnold
other vegetation by slashing or mowing, and by Arboretum, Harvard University, Jamaica Plain,
tillage to prepare ground for crops. MA. 673 p.
Benefits.—Roseleaf raspberry helps return Liogier H.A. 1985. Descriptive flora of Puerto
disturbed areas to forest vegetation, protects the Rico and adjacent islands, Spermatophyta. Vol.
soil, and furnishes food and cover for wildlife. 1. Editorial de la Universidad de Puerto Rico,
Leaves and tender shoot are browsed by livestock. Río Piedras, PR. 352 p.
The berries are eaten out of hand and made into
drinks and preserves throughout its range. Fresh Liogier, H.A. 1990. Plantas medicinales de Puerto
fruits are sold nearly year-round in Ugandan Rico y del Caribe. Iberoamericana de Ediciones,
markets (Burkill 1997). The fruits contain Inc. 566 p.
relatively high (1.58 and 12.93 mg/100g fresh
weight respectively) of vitamins E and C (Wei and Lu, L. and D.E. Boufford. 2002. Rubus Linnaeus
Payne 2002). Roseleaf raspberry has been grown [Draft]. In: A.R. Brach and Hong S., eds. Flora
as a ground cover in Cameroon plantations of China, Harvard University Herbaria,
(Burkill 1997). A double petaled variety (R. Cambridge, MA. http://hua.huh.harvard.edu/
rosifolius var. coronaries (Sims) Focke) is grown china/mss/volume09/Rosaceae-AGH_Rubus_
as an ornamental (Garden Plant Conservation coauthoring.htm. 200 p.
2002). Infusions of flower petals are used to
control diarrhea, vomiting, and other flu Nortan, A. 2002. Bush tucker: Roseleaf raspberry
symptoms, and as a tonic (Liogier 1990). An (Rubus rosifolius). Rumbalara Environmental
infusion of the leaves was brewed in former times Ecucation Centre, Sydney, NSW, Australia.
to relieve menstrual cramps, morning sickness, and http://rumbalara-e.schools.nsw.edu.au/
labor pains (Nortan 2002). bushtucker/ Rubus_rosifolius.htm. [not paged].
659
Salix arctica Pallas arctic willow
SALICACEAE
660
elsewhere in the Rocky Mountains, a common S. chambers (Jones and others 1999). Overall net
arctica/Polygonum bistortoides Pursh community assimilation was higher in the dry habitat than in
was identified in “water-receiving positions” the wet habitat, and higher in females than in
(Cooper and others 1997). However, the latter sites males, although there appeared to also be some
were also reported to have a good probability of habitat-sex interactions (Jones and others 1999).
experiencing wind scouring, and some of the Elevated temperature enhanced development and
associated vegetation suggested the area could be growth of both male and female catkins, but the
relatively dry (Cooper and others 1997). The response of the leaves to elevated temperatures
elevation at which it occurs is also associated with was more variable particularly among male
latitude. Plants are only found above 3,350 m in individuals (Jones and others 1999). These
New Mexico, whereas they occur at sea level and findings indicate the importance of defining both
up to 700 m in northern parts of its range, such as sex and habitat conditions when making
Greenland (Carter 1997, Argus and others 1999). observations on this dioecious species, and they
It grows in a range of soils, from acidic to may also have significance in predicting the
calcareous (Argus and others 1999, Cooper and response of Arctic willow to global climate
others 1997, Pojar and MacKinnon 1994). A change.
mutualistic association exists between arctic
willow and ectomycorrhizal fungi that are Benefits.—The majority of Salix species provide
primarily agarics, or gilled mushrooms (Cripps important browse for wildlife and livestock within
and Horak 2002). their range (Dayton 1931). Arctic willow,
particularly the young leaves stems and buds,
Reproduction.—Arctic willow reproduces by provide valuable browse for ptarmigan and other
seed and also vegetatively by rooting at the nodes arctic animals including muskox and reindeer
of stems (Fertig and Markow 2001). Flowering is (Kershaw and others 1998, Komarov 1936, Tolven
from June into August. On calm, sunny days the and others 2001). Arctic willow is tolerant of
female catkins can be up to 8.5 oC warmer and the varied conditions, and cuttings have been rooted
male catkins up to 7 oC warmer than the and used in restoration projects (Bittman 1997).
surrounding air temperature (Kershaw and others Willow leaves and bark are commonly used for
1998). The warmer temperatures speed pollen and medicinal purposes and the roots, twigs, and bark
seed development and also attract insects are used for a multitude of fiber work, such as
(Kershaw and others 1998). basketry and clothing by Native American peoples
(Moerman 1998). Arctic willow has been used for
Growth and Management.—Seeds have a similar purposes by peoples native to Asian Arctic
physiological dormancy and require a 30-day regions (Moerman 1998). The Yakuts have used
stratification period for germination (Baskin and an infusion as a substitute for tea called “chai-
Baskin 2001). Seeds germinate at 25 oC although talak” (Komarov 1936). It is used as fuel in some
minimum temperatures for germination are not regions where it is the only source of wood
defined (Densmore and Zasada 1983). Thompson (Kershaw and others 1998). Arctic willow is
(1992) reported that seeds of Salix species are cultivated, and it is especially popular for rock
“normally absent” from the seed bank in alpine gardens (Bailey and others 1976, Pojar and
and arctic communities. Individual shrubs live for MacKinnon 1994).
60 to 85 years although one specimen was reported
to be 180 and another 236 years old (Savile 1979, References.
Raup 1959). Habitat appears to influence the sex
that is most abundant. Dawson (1987) reported Argus, G.W., C.L. McJannet, and M.J. Dallwitz.
female plants were significantly more numerous 1999 onwards. ‘Salicaceae of the Canadian
than male plants in mesic-wet, more fertile, low Arctic Archipelago: Descriptions, Illustrations,
soil-temperature sites, whereas male plants were Identification, and Information. http://www.
most prevalent in drier, less fertile sites. In mun.ca/biology /delta/arcticf/.
addition, both habitat and sex may influence a
plant’s response to environmental conditions such Bailey, L.H., E.Z. Bailey, and the staff of the
as temperature. In a 3-year gas exchange field Liberty Hyde Bailey Hortorium. 1976. Hortus
study, male and female willows from dry and wet Third. Macmillian Publishing Company, New
habitats were subjected to passively enhanced York. 1,290 p.
summer temperature using small open-top
661
Baskin, C.C. and J.M. Baskin. 2001. Seeds, Dick-Peddie, W.A. 1993. New Mexico
ecology, biogeography, and evolution of vegetation. University of New Mexico Press,
dormancy and germination. Academic Press, Albuquerque, NM. 244 p.
New York. 666 p.
Fertig, W. and S. Markow. 2001. Guide to the
Bittman, K.K. 1997. High elevation native species willows of Shoshone National Forest. General
island model for mine reclamation. Quintette Technical Report RMRS-GTR-83. U.S.
Operating Corporation, Tumbler Ridge, BC. Department of Agriculture, Forest Service
http://www.teckcominco.com/presentations/kb- Rocky Mountain Research Station, Fort Collins,
highelv-1997.pdf. CO. 80 p.
Britton, N.L. 1901. Manual of the Flora of the Hitchcock, C.L., and A. Cronquist. 1973. Flora of
northern states and Canada. Henry Holt and the Pacific Northwest. University of Washington
Company, New York. 1,080 p. Press, Seattle, WA. 730 p.
Dayton, W. A. 1931. Important western browse Pojar, J. and A. MacKinnon. 1994. Plants of the
plants. Miscellaneous Publication 101. U.S. Pacific Northwest coast–Washington, Oregon,
Department of Agriculture, Washington DC. British Columbia and Alaska. Lone Pine
214 p. Publishing, Edmonton, Canada. 526 p.
Densmore, R. and Zasada, J.C. 1983. Seed Raup, H.M. 1959. The willows of boreal western
dispersal and dormancy patterns in northern America. Contributions from the Gray
willows: Ecological and evolutionary Herbarium of Harvard University 185: 3–95.
significance. Canadian Journal of Botany 61:
3207-3216
662
Savile, D.B.O. 1979. Ring counts in Salix arctica Tolven, A. J., Schroderus, and G.H.R. Henry.
from northern Ellesmere Island. Canadian Field- 2001. Age- and stage-based bud demography of
Naturalist 93: 81-82. Salix arctica under contrasting muskox grazing
pressure in the High Arctic. Evolutionary
Thompson, K. 1992. The functional ecology of Ecology 15: 4-6
seed banks. In: M. Fenner, ed., Seeds, the
ecology of regeneration in plant communities. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
CAB International, Wallingford, Oxon, U.K. Higgins. 1993. A Utah flora. Second Edition,
373 p. revised. Brigham Young University, Provo, UT.
986 p.
663
Salix exigua Nutt. coyote willow
SALICACEAE
James E. Nellessen
times longer than wide, 2 to 16 cm long, 4 to 15
mm wide, the margins are smooth or irregularly
toothed, with one to five teeth per cm, and the
petioles are short, 2 to 7 mm long (Argus 1986,
Britton and Brown 1913, Carter 1997, Gleason and
Cronquist 1963, Great Plains Flora Association
1986, Martin and Hutchins 1980-81, Newsholme
1992). Upon emergence the young twigs and
leaves are densely silvery hairy, but with time and
aging, the upper leaf surface reduces to a
moderately gray hairy or smooth surface,
sometimes slightly shiny. Plants in moist sites at
higher elevations in the mountains may be
noticeably less hairy (Mozingo 1987). The lower
leaf surface generally remains whitish and much
hairier. Coyote willow is one of few willow
species with stomata on both leaf surfaces (Argus
1986). Stipules are small and sometimes absent.
The chromosome number is 2n = 38 (Dorn 1998).
664
complicating the taxonomy for this species, the (Tamarix ramosissima Ledebour), with 52 percent
entire section, likely a syngameon undergoing of the leaf mass remaining after 142 days
geographic speciation (Brunsfield and others (Pomeroy and others 2000). A number of insects
1992). attack coyote willow, including leaf mining
beetles, lepidopterans, sawflies (Euura spp.), and
Range.—This is a widely distributed species of galling insects. A bud galling midge (Rabdophaga
plains and lower elevations to lower montane sp.) affects both terminal and lateral shoots,
habitats in the Western United States, as well as arresting growth of the immediate shoot (Declerck
throughout Eastern deciduous forests if S. interior and Price 1994). Coyote willow responds by
is considered a subspecies. The Western form producing new lateral buds and shoots, but many
(exigua) occurs from Saskatchewan to Eastern do not survive.
British Columbia in Canada, south through the
United States to Eastern and Southern California, Reproduction.—The male flowers are in densely
Arizona, New Mexico, and into Northern Mexico flowered catkins from 1 to 6 cm in length, each
and the Western Great Plains. The Eastern form or flower with two stamens, the filaments hairy. The
subspecies (interior) occurs from New Brunswick female catkins are loosely flowered, from 2 to 10
in Eastern Canada, westward throughout the Great cm long, the ovaries are smooth to slightly silky
Plains, as far west as Idaho, north to Alaska, south with a short or almost nonexistent style. Each
to Eastern New Mexico, and extends into the ovary bears nine to 15 ovules (Argus 1986).
Southeast along a line from Northern Virginia to Catkin scales are yellow and deciduous. The
Tennessee and Louisiana. The two subspecies catkins appear with the leaves or shortly after the
overlap in the Western Great Plains. In the leaves in spring (March to May) on short leafy
Western mountains, coyote willow can be found as stalks. The flowers are bee pollinated. There may
high as 2,900 m in elevation. even be a second blooming period from June to
August (Argus 1986). The fruits are smooth
Ecology.—Coyote willow commonly forms capsules from 4 to 8 mm long, green, turning
thickets along streams and rivers and is an yellow to reddish brown upon ripening. The fruits
important riparian species stabilizing streambanks. open by two valves releasing tiny seeds (22,000/g
It will commonly colonize sand bars along rivers, [Brinkman 1974]) with long hairs that aid in wind
hence the frequently used alternate common name dispersal. The seeds are short-lived, do not require
sandbar willow. It also occurs in ditches, and on stratification, must fall on a moist substrate, and
the edges of swamps, lakeshores, and other can germinate within 12 to 24 hours under proper
wetland habitats. This species will grow anywhere conditions (Young and Young 1992). Successful
groundwater is close to the surface. This includes germination is reported at 22 C for the Western
sites where human activities result in surface water subspecies exigua (Brinkman 1974) and 5 to 25
availability on a reasonably regular and reliable C for the Eastern subspecies interior (Densmore
basis (exclusive of saline conditions). Coyote and Zasada 1983, Baskin and Baskin 2001).
willow is also one of the few willow species that Flower-and-fruit-bearing shoots have been
can be found in lower elevation, hot desert areas in observed to sprout new leafy shoots and become
the Western United States. But because if its wide new side branches (Argus 1986).
elevational range it is both a floodplain-plains and
montane riparian species (Dick-Peddie 1993). Due Growth and Management.—Along the Rio
to the extensive rootstock and clonal nature of the Grande in New Mexico, clearing riparian areas of
species, cut stands of coyote willow will the invasive, nonnative salt cedar prior to peak
regenerate relatively quickly. The presence of an river flows in the spring can facilitate the
aerenchyma-containing wetland plant, Typha reestablishment of natives such as coyote willow
latifolia L., may facilitate a nonaerenchyma (Taylor and others 1999). In Alberta, Canada,
bearing species like coyote willow to survive in successful seedling establishment of several
low oxygen wetland soils (Callaway and King riparian species, including coyote willow, occurred
1996). Leaf water potentials as low as -2.7 MPa best on deposits 80 to 120 cm above the late
have been measured for high elevation populations summer stream flow (Rood and Mahoney 2000).
in Wyoming (Foster and Smith 1991). Coyote In Manitoba, Canada, average annual height
willow leaves within the Colorado River (Arizona) increases were 30 cm, stem diameter increases
decomposed substantially more slowly than those were 2.6 mm, flowering began in 2 to 3 year old
of Populus fremontii S. Watson and salt cedar stems, stem mortality was greatest in 3 to 6 year
665
old clones, entire plant senescence first occurred at Baskin, C.C. and J.M. Baskin. 2001. Seeds:
12 years, and the oldest plant recorded was 31 Ecology, Biogeography, and Evolution of
years on a point bar in the Assiniboine River Dormancy, and Germination. Academic Press,
(Ottenbreit and Staniforth 1992). San Diego, CA. 666 p.
Benefits.—Because coyote willow forms dense Brinkman, K.A. 1974. Salix L. Willow. In: C.S.
thickets and spreads clonally, it is an important Schopmeyer, tech. coord. Seeds of Woody
stabilizer of streambanks. It is frequently and Plants in the United States. U.S. Department of
successfully used in riparian and wetland Agriculture Forest Service, Handbook 450.
revegetation and restoration. It is an important Washington, DC. p. 746-750.
food for wildlife such as deer and beaver. It is
considered good browse for sheep and fair for Britton, N.L. and A. Brown. 1913 (1970 Dover
cattle, but cattle generally find it more palatable edition). An Illustrated Flora of the Northern
later in the growing season (Mozingo 1987, U.S. and Canada, Vol. 1. Dover Publications
Stubbendieck and others 1997, USDA 1937). Inc., New York. 680 p.
Coyote willow is used in intensive culture coppice
plantations (Aravanopoulos and others 1999), was Brunsfield, S.J., D.E. Soltis, and P.S. Soltis. 1992.
one of 32 woody plants tested for biomass Evolutionary patterns and processes in Salix:
gasification, and was one of three species that evidence from chloroplast DNA. Systematic
exceeded 0.30 liter methane per gram of volatile Botany 17(2): 239-256.
solids (Turick and others 1991). Coyote willow
has been successfully grown in tissue culture with Callaway, R.M. and L. King. 1996. Temperature
roots developing from shoots longer than 1 cm driven variation in substrate oxygenation and the
(Stoehr and others 1989). Native Americans have balance of competition and facilitation. Ecology
used the slender stems in making baskets and have 77(4): 1,189-1,195.
made a tea from the bark to treat fever and
headache. Willow bark, as with most willows, is Carter, J.L. 1988. Trees and Shrubs of Colorado.
bitter and acts as an astringent for diarrhea, has Johnson Books, Boulder, CO. 165 p.
been used for fevers, pain, arthritis, rheumatism,
and a poultice for cuts, ulcers, and rashes. Salicylic Carter, J.L. 1997. Trees and Shrubs of New
acid, commonly used in asperin as acetylsalicylic Mexico. Johnson Books, Boulder, CO. 534 p.
acid, was originally derived from salicin contained
in the bark of willows (Foster and Duke 1990). Declerck, F.R. and P.W. Price. 1994. Impact of a
Salicin is a member of the chemical class of bud-galling midge on bud populations of Salix
phenols and affords some protection against exigua. Oikos 70(2): 253-260.
browsing mammals such as rabbits. It tends to
attain a greater concentration in low growing Densmore, R. and J.C. Zasada. 1983. Seed
willows but becomes lost in larger, taller willows dispersal and dormancy patterns in northern
(Harborne 1988). Oestriol (an oestrogen), a human willows: ecological and evolutionary
sex hormone, has also been found in willow significance. Canadian Journal of Botany 61:
flowers. 3,207-3,216.
666
Foster, S.A. and J.A. Duke. 1990. A Field Guide to willow). Canadian Journal of Botany 70(6):
Medicinal Plants (Eastern/Central). Peterson 1,141-1,146.
Field Guide Series, Houghton Mifflin Co.,
Boston, MA. 366 p. Pomeroy, K.E., J.P. Shannon, and D.W. Blinn.
2000. Leaf breakdown in a regulated desert
Foster, J.R. and W.K. Smith. 1991. Stomatal river: Colorado River, Arizona, U.S.A.
conductance patterns and environment in high Hydrobiologia 434: 193-199.
elevation phreatophytes of Wyoming, USA.
Canadian Journal of Botany 69(3): 647-655. Rood, S.B. and J.M. Mahoney. 2000. Revised
instream flow regulation enables cottonwood
Gleason, H.A. and A.R. Cronquist. 1963. Manual recruitment along the St. Mary River, Alberta,
of the Vascular Plants of Northeastern U.S. and Canada. Rivers 7(2): 109-125.
Adjacent Canada. D. Van Nostrand Co., New
York. 810 p. Stoehr, M.U., M. Cai, and L. Zsuffa. 1989. In-vitro
plant regeneration via callus culture of mature
Great Plains Flora Association. 1986. Flora of the Salix exigua. Canadian Journal of Forest
Great Plains. University Press of Kansas, KS. Research 19(12): 1,634-1,638.
1,392 p.
Stubbendieck, J., S.L. Hatch, and C.H. Butterfield.
Harborne, J.B. 1988. Introduction to Ecological 1997. North American Range Plants, 5th Edition.
Biochemistry, 3rd Ed. Academic Press, London. University of Nebraska Press, Lincoln, NE.
356 p. 501 p.
Kartesz, J.T. 1994. A Synonymized Checklist of Taylor, J.P., D.B. Wester, and L.M. Smith. 1999.
the Vascular Flora of the United States, Canada, Soil disturbance, flood management, and
and Greenland, Vol. 1, 2nd Edition. Biota of riparian woody plant establishment in the Rio
North America Program of the North Carolina Grande floodplain. Wetlands 19(2): 372-382.
Botanical Garden. Timber Press, Portland, OR.
622 p. Turick, C.E., M.W. Peck, D.P. Chynoweth, D.E.
Jerger, E.H. White, L. Zsuffa, and W.A.
Kearney, T.H., R. Peebles, and Collaborators. Kenney. 1991. Methane fermentation of woody
1951 with 1960 supplement. Arizona Flora. biomass. Bioresource Technology 37(2): 141-
University of California Press, Berkeley, CA. 147.
1,085 p.
U.S. Department of Agriculture Forest Service.
Martin, W.C. and C.E. Hutchins. 1980-1981 1937 (1988 Dover edition). Range Plant
(reprinted 2001). A Flora of New Mexico, Vol. Handbook. Dover Publications Inc., New York.
1. Bishen Singh Mahendra Pal Singh (India) and 816 p.
Koeltz Scientific Books (Germany). 1,276 p.
Weber, W.A. and R.C. Wittman. 2001a. Colorado
Mozingo, H.N. 1987. Shrubs of the Great Basin: Flora: Eastern Slope. University Press of
A Natural History. University of Nevada Press, Colorado, Boulder CO. 521 p.
Reno, NV. 342 p.
Weber, W.A. and R.C. Wittman. 2001b. Colorado
Newsholme, C. 1992. Willows: The Genus Salix. Flora: Western Slope. University Press of
Timber Press, Portland, OR. 224 p. Colorado, Boulder CO. 488 p.
Ottenbreit, K.A. and R.J. Staniforth. 1992. Life Young, J.A. and C.G. Young. 1992. Seeds of
cycle and age structure of ramets in an Woody Plants in North America. Dioscorides
expanding population of Salix exigua (sandbar Press, Portland, Oregon. 407 p.
667
Salix scouleriana Barratt ex Hook. Scouler’s willow
SALICACEAE
John K. Francis
668
vary from near sea level to about 3,000 m in temperatures. It is an important browse species for
elevation (Peattie 1953). Scouler’s willow is top- domestic livestock and wild animals. Cattle, sheep,
killed by all but gentle fires, but usually greater and goats all like it as browse. It is sometimes the
than 65 percent of the plants sprout quickly most preferred food species for white tailed, black
afterwards (Anderson 2001). The species is tailed and mule deer, elk, moose, and bighorn
intolerant of shade, and when overtopped by sheep. Small mammals, bears, upland game birds,
conifers and other hardwoods, it begins to decline. and waterfowl feed to a lesser extent on leaves,
buds, and seeds. Fresh browse (twigs and leaves)
Reproduction.—Scouler’s willow flowers from contain 41 percent dry matter, 4 percent protein, 2
April through June, flowers appearing before percent fat, 20.8 percent nitrogen-free extract, 11.2
leaves, often while snow is still on the ground, and percent crude fiber, and good quantities of mineral
fruiting occurs from May through July, depending nutrients (Anderson 2001). The cover provided by
on area. The flowers are insect pollinated. There Scouler’s willow is important for mammals and
are about 14,300 cleaned seeds/g. Germination, birds. The flowers provide pollen and nectar to
which is epigeal, begins to occur in 12 to 24 hours honey bees in early spring (Anderson 2001). The
after seeds alight on wet ground. Germination wood, which is soft and close-grained, is not sawn
usually reaches 95 percent in 1 or 2 days into lumber but is used to a limited extent for
(Brinkman 1974). The seeds are dispersed by the firewood and wood carving (Viereck and Little
wind. Plants sprout from the root collar when cut 1972). The Secwepemc people of British
or top-killed. Pieces of stem and root will root and Colombia used Scouler’s willow wood for
grow if partially buried in moist soil (Forest smoking fish, drying meat, and constructing
Practices Branch 1997). fishing weirs, the inner bark for lashing, sowing,
cordage, and headbands, and decoctions of twigs
Growth and Management.—Annual height for treating pimples, body odor, and diaper rash
growth of sprouts from cut stems varies from 1 to (Secwepemc Cultural and Education Society
3 m/year. Up to 60 sprouts are produced per stem 2003).
(Forest Practices Branch 1997). Maximum height
at 20 years is about 9 m. At higher elevations, References
shrubs reach 4 to 5 m in 15 years after which
growth slows until a maximum height of 10 m is Anderson, M.D. 2001. Salix scouleriana. In: U.S.
reached (Natural Resources Conservation Service Department of Agriculture, Forest Service,
2003). Fruits should be collected by hand or with Rocky Mountain Research Station, Fire Sciences
pruning poles as soon as they turn from green to Laboratory, Missoula, MT. http://www.fs.
yellow. The capsules are air-dried until opening. fed.us/database/feis/plants/tree/salsco/all.html.
Generally, the seeds should be sown as soon as 57 p.
possible because they remain viable for only a few
days. Seed can be stored in sealed containers under Anonymous. 2003. Treatment from the Jepson
refrigeration for 4 to 6 weeks, but germination manual: Salicaceae. University of California,
begins to drop rapidly after 10 days. Seeds are Berkeley, CA. http://ucjeps.berkeley.edu/chi-
broadcast on well-prepared beds that are kept bin/get_JM_treatment.pl?7043,7050,7079. 3 p.
continually moist until germination and seedling
emergence. Light is required for successful Brinkman, K.A. 1974. Salix L., willow. In: C.S.
germination (Brinkman 1974). Recommended Schopmeyer, tech. coord. Seeds of woody plants
spacing using rooted cuttings for erosion control is in the United States. Agriculture Handbook 450.
1.8 m by 1.8 m; for unrooted whips or shorter U.S. Department of Agriculture, Forest Service,
cuttings, 0.6 m. Rooted cuttings can be grown to 3 Washington, DC. p. 746-750.
m tall in containers. Cuttings should be 45 to 60
cm long, and whips (not recommended) should be Department of Ecology. 2003. Plant selection
1.2 m long (Department of Ecology 2003). guide. State of Washington, Department of
Ecology, Olympia, WA. http://www.ecy.wa.
Benefits.—Scouler’s willow protects the soil and gov/programs/sea/pubs/93-30/table3.html. 6 p.
helps return sites to forest cover following
disturbance. When growing along streams, it helps Forest Practices Branch. 1997. Operational
protect the stream banks from erosion and shade summary for vegetation management: willow
the watercourse, thus maintaining cooler water complex. ISBN 0-7726-3166-2. Forest Practices
669
Branch, Ministry of Forests, Victoria, British Secwepemc Cultural and Education Society. 2003.
Colombia, Canada. 11 p. Flood plain garden. Secwepemc Nation,
Kamloops, BC, Canada. http://www.secwepemc.
Harrington, H.D. 1964. Manual of the plants of org/SECethnogardens2/flood_plain_garden.htm.
Colorado. Sage Books, Denver, CO. 666 p. 4 p.
Johnson, F.D. 1995. Wild trees of Idaho. Treeguide.com. 2003. Scouler willow, Salix
University of Idaho Press, Moscow, ID. 212 p. scouleriana Barratt ex Hook. http://www.
treeguide.com/Species.asp?Region=NorthAmeri
Natural Resources Conservation Service. 2003. can&SpeciesID=996. 2 p.
Plants profile: Salix scouleriana Barratt ex
Hook. http://plants.usda.gov/chi_bin/ Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
plant_search.cgi?mode=Scientific+Name&keyw trees and shrubs. Agriculture Handbook 410.
ordquery=Salix+scouleriana&earl=plant_search. Forest Service, U.S. Department of Agriculture,
4 p. Washington, DC. 265 p.
Peattie, D.C. 1953. A natural history of western Welsh, S.L. 1974. Anderson’s flora of Alaska and
trees. Houghton Mifflin Company, Boston, MA. adjacent parts of Canada. Brigham Young
751 p. University Press, Provo, UT. 724 p.
670
Salvia apiana Jepson white sage
LAMIACEAE
Arlee M. Montalvo
671
habitats (personal communication, James Beaver, year (Storms 1999). Seedling survival is expected
Indiana University, Bloomington). to be much lower in dry years and weedy areas.
Fire is a normal disturbance factor of most Most growth occurs after winter rains begin and
plant communities within which white sage continues until the soil becomes dry in summer.
occurs. Following fire, this shrub frequently Leaf growth over the season results in a seasonal
sprouts from distinct basal burls (Keeley 1998), dimorphism in leaf size (Westman 1981b). Smaller
but year-old seedlings are killed by fire, and leaves are produced on short shoots that develop
repeated burning kills mature plants (Zedler and from leaf axils along the main elongated stems and
others 1983). Seedlings can emerge from the are often retained when the older, longer leaves
seedbank in the first season after fire, but reported dehisce during drought. With the return of
densities are low (Zedler and others 1983, Keeley seasonal rains, the short shoots elongate into long
1998), and high intensity fires can kill seeds shoots, and the retained short leaves continue to
(Keeley and Fotheringham 1998). expand and subtend the next generation of short
shoots and leaves.
Reproduction.—Inflorescences arise from the
upper nodes in spring and produce flowers from Hybridization.—Epling (1938) reports that white
late May to July. Based on the floral morphology, sage hybridizes with other Salvias with the same
only some of the larger bees are capable of chromosome number, including S. mellifera
transferring pollen to the highly exerted stigmas. Greene, S. munzii Epl., S. leucophylla Greene, S.
Grant and Grant (1964) report three species of clevelandii (Gray) Greene, S. eremostachya Jeps.,
Xylocopa and a species of Bombus as effective S. pachyphylla Epl. Ex Munz., and S. vaseyi
pollinators. Hummingbirds, honeybees, bombyliid (Porter) Parish. Hybrid zones between S. mellifera
flies, and small species of native bees (primarily (black sage) and white sage are especially
Anthophora, Diadasia, and Osmia) sometimes common. Their hybrids are fully viable but, on
visit flowers (Grant and Grant 1964) but are average, suffer reduced fertility relative to parental
thought to be ineffective as pollinators. The species (Epling 1947a, Meyn and Emboden 1987),
protandrous flowers and presence of interspecific and there are fewer seeds/flower in F1 backcrosses
hybrids are consistent with an outcrossing to both parental species (Grant and Grant 1964).
breeding system. Inflorescences are large and Hybrids exhibit a range of intermediate floral and
many-flowered, making some self-pollination leaf traits (Epling 1947a, b, Anderson and
likely. The small nutlets fall from the dried calyx Anderson 1954) and differ in leaf anatomy (Webb
during mid to late summer but are secondarily and Carlquist 1964). Most hybrids are thought to
dispersed by harvester ants and seed caching be F1 crosses or backcrosses to black sage and
rodents. Seed traps revealed that seeds dispersed they tend to grow close to the parental types.
up to 3 m away and that there was a higher Although black and white sage overlap in range,
secondary removal rate of seeds in coastal sage they have different habitat affinities. Their ranges
scrub (80 percent) than in grassland (0 percent) or overlap in most of coastal southern California, but
the ecotone of the two habitats (about 40 percent) white sage is not found north of Santa Barbara
(DeSimone and Zedler 2001). County and it ranges farther south into Baja
California and eastward into the edge of the desert
Establishment and Growth.—Seedlings emerge than does black sage. Both species can occur
in the cool wet season in late winter to early spring intermingled, but black sage tends to be found in
and can take 2 years or more to reach maturity, flatter and wetter areas, while white sage is usually
depending on rain patterns and growing on drier slopes (Epling 1947a, Anderson and
conditions. Seedling establishment in shrublands is Anderson 1954, Grant and Grant 1964, Gill and
associated with gaps in mature vegetation and Hanlon 1998). As such, most sites contain a single
within grasslands in areas where herbs have been species, but hybrids are present at most sites where
removed by gophers (DeSimone and Zedler 1999, the species co-occur (Meyn and Emboden 1987).
2001). Seedlings show a relatively high growth Authors have speculated about factors that may
rate, little herbivore damage and they fold their limit the success of hybrids and maintain black and
leaves in response to drought (DeSimone and white sage as distinct species. Epling (1947a) first
Zedler 2001). In plots that were hand weeded and suggested that hybrids were less adapted to wet
seeded in a twice-normal rainfall year, about a sites than black sage and less adapted to dry sites
quarter of the seedlings survived to the second than white sage, but that they may be better
672
adapted to recently disturbed conditions. Hybrids trimming back the dried inflorescence stalks to
usually occur on recently disturbed areas, encourage compact growth. Plants can be easily
suggesting that hybrids are better adapted to started from seeds in the fall or planted from
intermediate habitats (Anderson and Anderson containers.
1954, Meyn and Emboden 1987). Within a natural
hybrid zone, Gill and Hanlon (1998) found that Growth and Management.—Because of its
xylem pressure potential was significantly higher historical abundance, fast growth, attractive nature,
in white than in black sage, that hybrids were and importance to the biodiversity of sensitive
intermediate to the two parental species, and that sage scrub habitat, white sage is used in many
white sage drops fewer leaves during drought. restoration and roadside revegetation projects,
These data support that white sage is more drought especially in endangered sage scrub vegetation.
adapted than black sage. It is possible that Both container plants and seeds are used
selection will maintain the integrity of the hybrid successfully in dry-land restoration in southern
zones if the intermediate hybrids do better in California. Seeds are collected from the dry,
intermediate habitats relative to their performance fruiting inflorescences, usually in July and August,
in the main habitats of the parental species. depending on site, and processed to remove chaff
The potential for reproductive isolation of the and insects. S&S Seeds (Carpenteria, CA) and
species within areas of overlap has been examined Mirov and Krabel (1939) report 715,000 to
most thoroughly by Grant and Grant (1964), 750,200 bulk seeds/kg, with often 70 percent
augmenting earlier observations (Epling 1947a, purity and 50 percent germination. Seeds should
Anderson and Anderson 1954). Black sage be planted before the onset of the cool rainy
generally flowers earlier than white sage and the season. If seed germination tests show low
short overlap in flowering time of one to several germination but high seed viability, seeds can be
weeks limits hybridization. In addition, white sage treated with dry or liquid smoke before sowing. If
has large, highly modified flowers that are liquid smoke is used, seeds must be air-dried prior
pollinated almost exclusively by large carpenter to dry broadcasting. Dormant seeds can result in a
and bumblebees (Xylocopa and Bombus), while beneficial seed bank and establishment over
black sage has smaller, unmodified flowers that several years. Hydroseeding can be successful on
are pollinated primarily by honeybees and small steep slopes and road cuts. Shallow planting
native solitary bees. methods such as hydroseeding and imprinting are
likely to produce higher germination than drilling
Seed germination.—Keeley (1987) showed that because seed germination is improved with
exposure to 70 oC for 1 hour followed by exposure to light. Based on seedling survival data
incubation in light substantially increased (Storms 1999, DeSimone and Zedler 2001), even
germination over controls, that heat shock for 5 under well-weeded and rainy conditions, fewer
minutes at 100 oC had no effect, and that exposure than 25 percent of seedlings should be expected to
for 5 min at 120 oC killed seeds. Leachate from survive through their first year. This species has
charred wood did not increase germination decreased dramatically in percent cover in western
(Keeley 1987), but cool smoke treatment increased Riverside County since the early 1930’s (Minnich
germination (Keeley and Fotheringham 1998). and Dezzani 1998). Successful restoration will
Liquid smoke can also be used to significantly depend on controlling the factors associated with
increase germination (author’s observation). its decrease, especially invasive plants and fire
Horticulture.—The generally rounded form with frequency, so that shrubs can establish, mature,
attractive whitish leaves and long graceful, pink- and buildup a seed bank.
tinged inflorescence stalks make this an attractive
shrub for background areas of dry landscape Benefits.—White sage leaves are an important
gardens (Perry 1992, Keator 1994, Clebsch 1997). browse of mountain sheep in the winter and spring
Plants require full sun, well-drained soil, do not (Perry and others 1987). Leaves contain essential
tolerate constantly damp conditions, and are oils and a variety of diterpines and triterpenes,
sensitive to conditions below –7 oC (Keator 1994, including carnosic acid, oleaolic acid, and ursolic
Clebsch 1997), although populations in desert acid, some of which have been linked to the
scrub are likely to be more cold tolerant than those medicinal use of white sage by native Tribes.
from coastal locations. Plants should be kept dry Some may also deter many herbivores (Dentali
all summer long. Clebsch (1997) recommends and Hoffman 1992). A tincture of the leaves has
673
been used internally as a diaphoretic and diuretic, Epling, C., H. Lewis, and P.H. Raven. 1962.
and externally to wash skin. The tincture may have Chromosomes of Salvia: section Audibertia.
antimicrobial properties. Antibacterial and Aliso 5: 217-221.
antifungal activity of several compounds have
been verified in vitro (Dentali and Hoffman 1992). Gill, D.S. and B.J. Hanlon. 1998. Water potentials
In addition to medicinal use, the dried leaves have of Salvia apiana, S. mellifera (Lamiaceae), and
been used as a smudge and crushed as a soapless their hybrids in the coastal sage scrub of southern
shampoo (Bean and Saubel 1972, Moore 1989). California. Madroño 45: 141-145.
Seeds have been used as food and leaves for
flavoring. Grant, K.A. and V. Grant. 1964. Mechanical
isolation of Salvia apiana and Salvia mellifera
References (Labiatae). Evolution 18: 196-212.
Anderson, E. and B.R. Anderson. 1954. Hellmers, H., J.S. Horton, G. Juhren, and J.
Introgression of Salvia apiana and Salvia O'Keefe. 1955. Root systems of some chaparral
mellifera. Annals of the Missouri Botanical plants in southern California. Ecology 36: 667-
Garden 41: 329-338. 678.
Bean, J.L. and K.S. Saubel. 1972. Temalpakh: Keator, G. 1994. Complete Garden Guide to the
Cahuilla Indian Knowledge and Usage of Plants. Native Shrubs of California. Chronicle Books,
Malki Museum Press, Morongo Indian San Francisco, CA. 314 p.
Reservation, CA. 225 p.
Keeley, J.E. 1987. Role of fire in seed germination
Clebsch, B. 1997. A Book of Salvias: Sages for of woody taxa in California chaparral. Ecology
Every Garden. Timber Press Inc., Portland, OR. 68: 434-443.
221 p.
Keeley, J.E. 1998. Postfire ecosystem recovery and
Dentali, S.J. and J.J. Hoffmann. 1992. Potential management: The October 1993 large fire
antiinfective agents from Eriodictyon episode in California. In: J. M. Moreno (ed.)
angustifolium and Salvia apiana. International Large Forest Fires. Backbuys Publishers, Leiden,
Journal of Pharmacognosy 30: 223-231. Netherlands. p. 69-90.
DeSimone, S.A. and P.H. Zedler. 1999. Shrub Keeley, J.E., and C.J. Fotheringham. 1998. Smoke-
seedling recruitment in unburned Californian induced seed germination in California
coastal sage scrub and adjacent grassland. chaparral. Ecology 79: 2,320-2,336.
Ecology 80: 2,018-2,032.
Meyn, O. and W.A. Emboden. 1987. Parameters
DeSimone, S.A. and P.H. Zedler. 2001. Do shrub and consequences of introgression in Salvia
colonizers of southern Californian grassland fit apiana X S. mellifera (Lamiaceae). Systematic
generalities for other woody colonizers? Botany 12: 390-399.
Ecological Applications 11: 1,101-1,111.
Minnich, R.A. and R.J. Dezzani. 1998. Historical
Epling, C. 1938. The California salvias. Annals of decline of coastal sage scrub in the Riverside-
the Missouri Botanical Garden 25: 95-188. Perris plane, California. Western Birds 29: 366-
391.
Epling, C. 1947a. The genetic aspects of natural
populations: actual and potential gene flow in Mirov, N.T. and C. J. Kraebel. 1939. Collecting and
natural populations. The American Naturalist 81: handling seeds of wild plants. Civilian
104-113. Conservation Corps, Forestry Publication No. 5,
United States Government Printing Office,
Epling, C. 1947b. Natural hybridization of Salvia Washington, D.C. 42 p.
apiana and S. mellifera. Evolution 1: 69-78.
674
Moore, M. 1989. Medicinal Plants of the Desert wash in Baja California, Mexico. Journal of Arid
and Canyon West. Museum of New Mexico Environments 24: 155-164.
Press, Santa Fe, NM. 184 p.
Storms, N. 1999. Restoration of a Native Shrubland
Munz, P.A. and D.D. Keck. 1968. A California in an Area of Frequent Disturbance and High
Flora with Supplement. University of California Nitrogen Deposition. M.S. Thesis. University of
Press, Berkeley, CA. 1681 + 224 p. California, Riverside. 60 p.
Perry, B. 1992. Landscape Plants for Western Webb, A.-A. and S. Carlquist. 1964. Leaf anatomy
Regions: an Illustrated Guide to Plants for Water as an indicator of Salvia apiana-mellifera
Conservation. Land Design Publishing, introgression. Aliso 5: 437-449.
Claremont, CA. 318 p.
Westman, W.E. 1981a. Factors influencing the
Perry, W.M., J.W. Dole, and S.A. Holl. 1987. distribution of species of California coastal sage
Analysis of the diets of mountain sheep from the scrub. Ecology 62: 439-455.
San Gabriel Mountains, California. California
Fish and Game 73: 156-162. Westman, W.E. 1981b. Seasonal dimorphism of
foliage in Californian coastal sage scrub.
Poole, D.K. and P.C. Miller. 1975. Water relations Oecologia 51: 385-388.
of selected species of chaparral and coastal sage
communities. Ecology 56: 1,118-1,128. Zedler, P.H., C.R. Gautier, and G.S. McMaster.
1983. Vegetation change in response to extreme
Schmitt, A.K., C.E. Martin, V.S. Loeschen, and A. events: the effect of a short interval between fires
Schmitt. 1993. Mid-summer gas exchange and in California chaparral and coastal scrub.
water relations of seven C3 species in a desert Ecology 64: 809-818.
675
Salvia mellifera Greene black sage
LAMIACEAE
676
Reproduction.—The timing of growth and occurred (E. Allen, L. Egerton-Warburton, A.
flowering is closely adapted to the Mediterranean Montalvo, unpublished report).
climate of southern California, with cool wet
winters and hot dry summers (Grant and Grant Seed germination.—Seeds of black sage often
1964, Gill and Mahall 1986). Common garden have low germination rates unless exposed to light
studies with 12 populations revealed that date of or components of fire such as charred wood,
first flowering varies with source population and smoke, or KNO3 (Keeley 1986, Thanos and
ranged from early February to early May Rundel 1995, Keeley and Fotheringham 1998).
(Montalvo, unpublished data). Also, shifts in Exposure to dry cool smoke for 5 min (Keeley and
flowering time occur in years with contrasting Fotheringham 1998) yields higher germination
rainfall patterns (Meyn and Emboden 1987). than charred wood. A 12 to 15 hour soak in a 1:25
The self-compatible flowers are dilution of Regen 2000 Smokemaster, a liquid
hermaphroditic with stamens dispersing pollen smoke product, also breaks dormancy (author’s
before stigmas are receptive. Flowers are observation). Heat of fire may reduce germination
pollinated by small to meduim-sized solitary bees of seeds from sage scrub populations (Keeley and
in families Andrenidae, Anthophoridae, Fotheringham 1998). Oddly, seeds from desert
Halictidae, Megachilidae, and Xylocopinae as well populations, where fires are less frequent, were
as introduced honey bees (Grant and Grant 1964). stimulated to germinate by heat and charred wood,
Rarer visitors include large-bodied Bombus and but not by light (Keeley 1986).
Xylocopa bees, syrphid and bombyliid flies, and
Anna’s hummingbirds. Each flower produces up to Genetics, Geographic Variation, and Fitness.—
four seeds that are gravity dispersed in June and Black sage is genetically variable over its
July from the dry, persistent calyces, and can be geographic range. In a survey of 12 populations
secondarily dispersed by ants. Seeds accumulate in (Montalvo, Clegg, and Ellstrand unpublished, 14
a dormant seed bank (Keeley 1986). allozyme loci), expected heterozygosity was high
(He = 0.23) and alleles averaged 2.81 per locus.
Establishment and Growth.—Black sage occurs Within sites, a low inbreeding coefficient (f =
in plant communities adapted to fire and drought. 0.072) suggests the protandrous flowers and
Seedlings emerge in clearings between adult pollinators combine to promote outcrossing. There
shrubs, but most emerge in the first 2 years is significant but low structure both among and
following fire, primarily from February to April within populations, and results are consistent with
(Keeley 1986, Westman and O’Leary 1986). a relatively high level of historical gene flow
Plants take two or more growing seasons to reach (Theta-p (~FST, sites relative to total population) =
maturity. After fire, plants sometimes resprout 0.041; Theta-s (plots relative to sites) = 0.057; and
from the base (Went and others 1952, Keeley F (FIT) = 0.125).
1986, 1998). However, where fire intensity is high, Populations cluster by genetic similarity, and
shrubs tend to be killed, and recruitment is geographic floristic associations and this is
primarily by seed. Across a range of conditions, somewhat mirrored by distribution patterns of
resprouting success averaged 15.8 percent, and morphological variation and timing of flowering.
shrubs that resprouted had significantly smaller Flowers from populations in the Santa Monica
basal diameters, suggesting resprouting decreases Mountains and Simi Hills can be strikingly bluish.
with age (Keeley 1998). The seasonal pattern of In the drier, more interior hills and valleys of
growth and dormancy varies both geographically Riverside Co., flowers tend to be white to pale
and between years. Generally, seasonal growth blue or lavender, and plants flower earlier. These
begins after the start of fall rains, most new leaves differences are retained in common gardens.
are produced by mid March, and flowering occurs Differences in timing of flowering could affect
from late winter to late spring. seed production of translocated populations if
Roots are colonized by arbuscular pollinator activity, seed predation, or important
mycorrhizal (AM) fungi. After 4 months of growth physical environmental factors differ from home
in a greenhouse, root and shoot mass was higher in sites. Plants from contrasting populations also
seedlings inoculated with AM fungi than in smell different, likely due to differences in
controls, but in younger plants the opposite composition of volatile chemicals. Such
differences among populations may be adaptive
677
and contribute to their success under particular Within a hybrid zone, Gill and Hanlon (1998)
environments. Montalvo and Ellstrand found that xylem pressure potential was
(unpublished) evaluated seedling survival, growth significantly higher in white than in black sage and
and reproduction of 12 populations in a common that putative hybrids were intermediate to the two
garden. A combination of climatic data and soil parental species. These data support that S. apiana
traits was used to calculate environmental is more drought adapted than S. mellifera, and that
similarity among garden and source sites. After the hybrids are intermediate. In addition,
three growing seasons, populations from sources reproductive isolation is favored by flowering
most similar to the garden site were the most differences. S. apiana flowers later than black
successful (measured as survival x flower sage, limiting opportunities to hybridize (Grant
production). This effect was small (r2 = 0.18) but and Grant 1964). In addition, white sage has
is consistent with the presence of weak adaptive larger, highly modified flowers that are pollinated
differences among source populations. almost exclusively by large bees in Xylocopa and
Bombus. Black sage has smaller, two-lipped
Hybridization.—Black sage hybridizes with the flowers that are rarely visited by these large-
shrubs S. apiana Jeps. (n = 15), S. leucophylla bodied bees.
Greene (2n = 30), and S. clevelandii (Gray) Greene
(2n = 30), and rarely with annuals S. columbariae Growth and Management.—Seeds are collected
Benth (n = 13) and S. carduacea Benth (n = 16) from June-August and should be cleaned to
(Epling 1938, 1947a, Epling and others 1962, remove seed predators. Air separation can be used
Munz and Keck 1968). Most hybridization is with to remove lighter unfilled seeds that are sometimes
S. apiana. These hybrids are fully viable but suffer abundant. There are about 1,375,000 seeds/bulk kg
reduced pollen fertility (Epling 1947a, Meyn and (personal communication with Victor Schaff, S&S
Emboden 1987) and numbers of seeds/flower in Seeds, Carpenteria, CA). Treatment of seeds with
F1 back crosses to parental species (Grant and dry or liquid smoke increases germination
Grant 1964). Hybrids differ from both parental substantially. Seeds should be air-dried before
species by a range of intermediate floral, leaf, and handling. Plants can be sown in flats for
anatomical traits (Epling 1947a, b, Anderson and subsequent transplanting or directly sown into the
Anderson 1954, Webb and Carlquist 1964). Most ground in the fall. Plants can be started from
are thought to be F1s or backcrosses to S. mellifera cuttings, but this should be avoided for restoration
and tend to grow near parental types. Hybrid zones because it reduces genetic variation. Prostrate
can be stable in position but may shift in cultivars (e.g., Clebsch 1997) should not be used
morphology (Meyn and Emboden 1985). This in restoration. Black sage grows in well-drained
suggests differences in the relative success of soil and can tolerate some summer water in a
hybrids and backcrossed progeny over time. The garden.
influence of gene exchange on floral form of black
sage does not appear to extend beyond hybrid Benefits.—Black sage is important for restoration,
zones (Epling 1947a, b). erosion control, and native landscape gardening in
Several authors have speculated on the California. Plants provide cover and seeds for
factors promoting hybridization while also wildlife, and in western Riverside Co. the listed
maintaining black and white sage as distinct California gnatcather frequently nests in
species. While ranges overlap in most of southern vegetation containing black sage (Weaver 1998).
California, S. apiana is not found north of Santa The seeds were eaten and the leaves used for
Barbara County and it ranges farther south into flavoring by native California tribes (Bean and
Baja California and eastward. In areas of contact, Saubel 1972). In addition, the leaves have a long
S. mellifera tends to occur in flatter and wetter history of medicinal use by native tribes (Bocek
microsites, while S. apiana occurs on drier slopes 1984). Their diterpene compounds have
(Epling 1947a, Anderson and Anderson 1954, antimicrobial activity against gram (+) bacteria
Grant and Grant 1964, Meyn and Emboden 1987, (Moujir and others 1996).
Gill and Hanlon 1998). Hybrids also occur in
recently disturbed areas, suggesting that hybrids Decreasing Populations.—Black sage
are adapted to intermediate habitats (Anderson and populations are declining. Increased ignition
Anderson 1954, Meyn and Emboden 1987). sources and invasion of shrublands by annual
grasses have increased fire frequency, jeopardizing
678
recovery of seed banks (Westman and O’Leary their hybrids in the coastal sage scrub of
1986, Haidinger and Keeley 1993, Minnich and southern California. Madroño 45: 141-145.
Dezzani 1998). Black sage is also inhibited by
exotic annual grasses, black mustard (Went and Gill, D.S. and B.E. Mahall. 1986. Quantitative
others 1952, Storms 1999), and air pollution phenology and water relations of an evergreen
(Westman 1985, Preston 1988). and deciduous chaparral shrub. Ecological
Monographs 56: 127-143.
References
Anderson, E. and B. R. Anderson. 1954. González, A.G., L.S. Andrés, Z.E. Aguiar, and
Introgression of Salvia apiana and Salvia J.G. Luis. 1992. Diterpenes from Salvia
mellifera. Annals of the Missouri Botanical mellifera and their biogenetic significance.
Garden 41: 329-338. Phytochemistry 31(4): 1,297-1,305.
Arey, J., D.E. Crowley, M. Crowley, M. Resketo, Grant, K.A. and V. Grant. 1964. Mechanical
and J. Lester. 1995. Hydrocarbon emissions isolation of Salvia apiana and Salvia mellifera
from natural vegetation in California's south (Labiatae). Evolution 18: 196-212.
coast air basin. Atmospheric Environment 29:
2,977-2,988. Haidinger, T.L. and J.E. Keeley. 1993. Role of
high fire frequency in destruction of mixed
Bean, J. and K. Saubel. 1972. Temalpakh: Cahuilla chaparral. Madroño 40: 141-147.
Indian knowledge and usage of plants. Malki
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CA. 225 p. Higher Plants of California. University of
California Press, Ltd., Los Angeles, CA. 1,400
Bocek, B. 1984. Ethnobotany of Costanoan p.
Indians, California, based on collections by
John P. Harrington. Economic Botany 38: 240- Keeley, J.E. 1986. Seed germination patterns of
255. Salvia mellifera in fire-prone environments.
Oecologia 71: 1-5.
Clebsch, B. 1997. A Book of Salvias: Sages for
Every Garden. Timber Press, Portland, OR. 221 Keeley, J.E. 1998. Postfire ecosystem recovery
p. and management: The October 1993 large fire
episode in California. In: J. M. Moreno, ed.
Epling, C. 1938. The California salvias. Annals of Large Forest Fires. Backbuys, Leiden, The
the Missouri Botanical Garden 25: 95-188. Netherlands. p. 69-90.
Epling, C. 1947a. The genetic aspects of natural
populations: actual and potential gene flow in Keeley, J.E. and C.J. Fotheringham. 1998. Smoke-
natural populations. The American Naturalist induced seed germination in California
81: 104-113. chaparral. Ecology 79: 2,320-2,336.
Epling, C. 1947b. Natural hybridization of Salvia Kolb, K.J. and S.D. Davis. 1994. Drought
apiana and S. mellifera. Evolution 1: 69-78. tolerance and xylem embolism in co-occurring
Epling, C.,H. Lewis, and P.H. Raven. 1962. species of coastal sage and chaparral. Ecology
Chromosomes of Salvia: section Audibertia. 75: 648-659.
Aliso 5: 217-221.
Meyn, O. and W.A. Emboden. 1987. Parameters
Gigon, A. 1979. CO2-gas exchange, water and consequences of introgression in Salvia
relations and convergence of Mediterranean apiana X S. mellifera (Lamiaceae). Systematic
shrub-types from California and Chile. Botany 12: 390-399.
Oecologia Plantarum 14: 129-150.
Minnich, R.A. and R.J. Dezzani. 1998. Historical
Gill, D.S. and B.J. Hanlon. 1998. Water potentials decline of coastal sage scrub in the Riverside-
of Salvia apiana, S. mellifera (Lamiaceae), and Perris plane, California. Western Birds 29: 366-
391.
679
Moujir, L., A.M. Gutiérrez-Navarro, L. San Weaver, K.L. 1998. Coastal sage scrub variations
Andres, and J.G. Luis. 1996. Bioactive of San Diego County and their influence on the
diterpenoids isolated from Salvia mellifera. distribution of the California gnatcatcher.
Phytotherapy Research 10: 172-174. Western Birds 29: 392-405.
Munz, P.A. and D.D. Keck. 1968. A California Webb, A.-A. and S. Carlquist. 1964. Leaf anatomy
Flora with Supplement. University of California as an indicator of Salvia apiana-mellifera
Press, Berkeley, CA. 1,681 + 224 p. introgression. Aliso 5: 437-449.
Neisess, K.R., R.W. Scora, and J. Kumamoto. Went, F.W., G. Juhren, and M.C. Juhren. 1952.
1987. Volatile leaf oils of California Salvias. Fire and biotic factors affecting germination.
Journal of Natural Products 50: 515-517. Ecology 33: 351-364.
Preston, K.P. 1988. Effects of sulphur dioxide Westman, W.E. 1981. Factors influencing the
pollution on a Californian coastal sage scrub distribution of species of California coastal sage
community. Environmental Pollution 51: 179- scrub. Ecology 62: 439-455.
195.
Westman, W.E. 1983. Xeric Mediterranean-type
Storms, N. 1999. Restoration of a native shrubland shrubland associations of Alta and Baja
in an area of frequent disturbance and high California and the community/continuum
nitrogen deposition. Master of Science in debate. Vegetatio 52: 3-19.
Botany Thesis. University of California,
Riverside. 60 p. Westman, W.E. 1985. Air pollution injury to
coastal sage scrub in the Santa Monica
Thanos, C.A. and P.W. Rundel. 1995. Fire- Mountains, southern California. Water, Air, and
followers in chaparral: nitrogenous compounds Soil Pollution 26: 19-41.
trigger seed germination. Journal of Ecology
83: 207-216. Westman, W.E. and J.F. O'Leary. 1986. Measures
of resilience: the response of coastal sage scrub
to fire. Vegetatio 65: 179-189.
680
Sassafras albidum (Nutt.) Nees sassafras
LAURACEAE
John K. Francis
681
range from 21 to 27 °C. (Griggs 1990). Sassafras Benefits.—Sassafras adds beauty in both summer
grows in a wide variety of soil types and grows and winter to forest lands where it grows. It has
best on well-drained sandy loams with a pH been used to help restore depleted soils of old
between 6.0 and 7.0 (Griggs 1990). Seedlings and fields and is considered superior to black locust
saplings are easily top-killed by fire but readily (Robinia pseudoacacia L.) and pines in this
sprout. Thick-barked trees are injured by fire application (Grigg 1990). Sassafras is browsed
(Sullivan 1993). Sassafras is sometimes attacked year-round by white tailed deer. The plants are
by leaf blight caused by Actinopelte dryina, by leaf also utilized to some extent by woodchucks,
spots caused by Mycosphaerella sassafras, and by rabbits, beavers, and bears. Many species of birds
a Nectria canker. A number of insects also attack and some small mammals consume the fruits.
the species (Griggs 1990). Crude protein ranges from 21.0 percent (leaves in
April) to 6.1 percent (twigs in January). The fruits
Reproduction.—Flowers open in early spring have a high lipid and energy value (Sullivan
with the first leaves. Male and female flowers 1993). Sassafras makes an attractive ornamental
usually occur on different plants (Sargent 1923). small tree and is widely, although not heavily
Fruits mature in August or September. Seed used. The fall foliage colors of yellow, orange, and
production begins when plants are about 10 years red are particularly beautiful. Sassafras heartwood
old (Griggs 1990). There are about 12,800 is pale brown to orange brown; sapwood is
seeds/kg (Bonner and Maisenhelder 1974) or yellowish-white. The wood is ring-porous, coarse-
according to Griggs (1990), 8,800 to 13,200 but straight-grained, brittle, soft, and has a spicy
seeds/kg. About 35 percent of the seeds are sound. aromatic odor. It is resistant to decay making it
Birds are the principal dispersers of seeds, with useful for fence posts and other items exposed to
gravity, water, and mammals playing minor roles. moisture. It is also useful for lumber, millwork,
Germination, which occurs in the spring, is furniture, and small boats. Technical details of the
hypogeal. Seeds may remain viable in the soil seed wood have been published (Center for Wood
bank for up to 6 years (Griggs 1990). Sassafras Anatomy Research 2003). The use of sassafras in
sprouts prolifically from stumps and suckers from herbal medicine has a long history beginning with
lateral roots. Native Americans who used it to treat colds, high
blood pressure, heart troubles, swelling, as a tonic,
Growth and Management.—Growth of sassafras to treat for worms, and to control fever (Moerman
is moderately fast. Sprouts can reach 1.2 m in the 1986). Sassafras tea is still widely drunk both for
first year and 4.5 m in 4 years (Floridata 2003). pleasure and as a tonic. Extracts and infusions of
Rare individual trees reaching the forest canopy the plant are used to treat a wide range of physical
may live as long as 300 years (TreeGuide 2003). complaints. Dry, powdered leaves, called file, are
Fruits are ripe and can be harvested when they turn used to thicken gumbo and other Cajun dishes
from green to dark blue and can be picked by hand (Katzer 1999). Oil of sassafras, extracted from the
or by flailing them from the trees or shrubs onto a root bark, was the original flavoring for root beer
tarp. The seeds can be cleaned of fruit tissue by and was used to flavor candy, chewing gum, and
rubbing against hardware cloth and washing away medicines. Because it contains safrole, a
the pulp. One kg of fruits will yield about 310 g of carcinogen, it is no longer permitted in food, but
seeds. Seeds should be placed in sealed containers may be used to add scent to cosmetics and soap
and stored at 2 to 5 °C. Before planting, seeds (TreeGuide 2003).
should be moistened and held at the above
temperatures for 120 days. Drilling in nursery beds References
in 20- to 30-cm-wide rows and covering with 6 to
12 mm of soil is recommended (Bonner and Bonner, F.T. and L.C. Maisenhelder. 1974.
Maisenhelder 1974). Sassafras can also be Sassafrass albidum (Nutt.) Nees sassafras. In:
reproduced from root cuttings but not from stem C.S. Schopmeyer, tech. coord. Seeds of woody
cuttings (Griggs 1990). Wildlings are difficult to plants in the United States. Agriculture
transplant. Sassafras shrubs and trees can be killed Handbook 450. U.S. Department of Agriculture,
by injecting 2,4-D, picloram, or glyphosate Forest Service, Washington, DC. p. 761-762.
(Sullivan 1993).
Center for Wood Anatomy Research. 2003.
Sassafras albidum. Technical Transsfer Fact
682
Sheet. U.S. Department of Agriculture, Forest Oberle, R. 2003. Sassafras albidum (Nutt.) Nees.
Service, Forest Products Laboratory, Madison, http://www.biologie.uni-ulm.de/systax/
Wisconsin. http://www2fpl.fs.fed.us/ dendrologie/sasalblv.htm. 1 p.
TechSheets/HardwoodNA/htmlDocs/sassafrasal
bi.html. 3 p. Plants For a Future. 2003. Plant portrait-Sassafras
albidum. http://www.scs.leeds.ac.uk/pfaf/
Floridata. 2003. Sassafras albidum. http:// sassafr.html. 3 p.
www.floridata.com/ref/s/sass_alb.cfm. 4 p.
Sanchez de Lorenzo-C., J.M. 2003. Árboles
Griggs, M.M. 1990. Sassafras albidum (Nutt.) ornamentals: Sassafras albidum (Nutt.) Nees.
Nees, sassafras. In: R.M. Burns, and B.H. http://www.arbolesornamentales.com/Sassafrasa
Honkala. Silvics of North America. Vol. 2., lbidum.htm. 1 p.
Hardwoods. Ag. Handb. 654. U.S. Department
of Agriculture, Forest Service, Washington, DC. Sargent, C.S. 1923. Manual of the trees of North
p. 773-777. America (exclusive of Mexico). Houghton
Mifflin, Boston, MA. 910 p.
Katzer, G. 1999. Sassafras [Sassafras albidum
(Nutt.) Nees]. http://www-ang.kfunigraz. Sullivan, J. 1993. Sassafras albidum. In: U.S.
ac.at/~katzer/engl/Sass_alb.html. 4 p. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences
Moerman, D.E. 1986. Medicinal plants of Native Laboratory, Fire Effects Information System.
America. Technical Reports 19. University of http://www.fs.fed.us/database/feis/plants/tree/
Michigan Museum of Anthropology, Ann sasalb/all.html. 17 p.
Arbor, MI. 534 p.
TreeGuide. 2003. Sassafras. http://www.
Natural Resources Conservation Service. 2003. treeguide.com/Species.asp?SpeciesID=953. 3 p.
Plants profile: Sassafras albidum (Nutt.) Nees.
http://plants.usda.gov/cgi_bin/plant_profile.cgi? Vines, R.A. 1982. Trees of North Texas.
symbol=SAAL5. 6 p. University of Texas Press, Austin, TX. 466 p.
683
Schaefferia frutescens Jacq. Florida-boxwood
CELASTRACEAE
John K. Francis
ends. They measure 4 to 7 cm in length and 1.2 to
2.5 cm in width. The flowers are small, pale green
or greenish-yellow or greenish-white, and borne in
small clusters at the leaf axiles. The fruits that
arise from them are spherical to ovoid, 4 to 8 mm
long, and red or orange-red. Each fruit contains
two light-brown seeds (Howard 1989, Little and
others 1974, Liogier 1994, Nelson 1996).
684
and 93 days of sowing in moist peat. Germination Cooperative Extension Service. 3 p.
is epigeal. The fruits are apparently eaten and the
seeds dispersed by birds. Howard, R.A. 1989. Flora of the Lesser Antilles,
Leeward and Windward Islands. Vol. 5. Arnold
Growth and Management.—Florida-boxwood is Arboretum, Harvard University, Jamaica Plain,
reported to have a high growth rate, withstands MA. 604 p.
heavy pruning, and can be formed into hedges and
shaped into small trees (Gilman 1999). Institute of Systematic Botany. 2001. Atlas of
Florida Plants: Schaefferia frutescens.
Benefits.—Florida-boxwood makes superior University of South Florida. http://
quality hedges, screens, and accent plantings. It is plantatlas.usf.edu/main.asp?plantID==2227. 1 p.
also used for reclamation plantings (Gilman
1999). The wood is light brown to yellow, hard Liogier, H.A. 1990. Plantas medicinales de Puerto
and moderately heavy. It has been used in the past Rico y del Caribe. Iberoamericana de Ediciones,
for wood engraving (Little and others 1974). It is Inc., San Juan, PR. 566 p.
also used, when available, for carving, for boxes
(Gilman 1999), and, to a limited extent, for fuel. Liogier, H.A. 1994. Descriptive flora of Puerto
The leaves have been used in the past as Rico and adjacent Islands. Vol. 3. Editorial de la
emergency soap for the body and cloths (Liogier Universidad de Puerto Rico. Río Piedras, PR.
1990). Almost nothing is known about its forage 461 p.
value except that Key deer (Odocolleus virginanus
clavium) will not eat it (Schaus and others 2001). Little, E.L., Jr., R.O. Woodbury, and F.H.
Decoctions are used in herbal medicine for colds, Wadsworth. 1974. Trees of Puerto Rico and the
flu, and chronic cough, and crushed leaves are Virgin Islands. Vol. 2. Agriculture Handbook
used as a soothing bath for skin irritations (Liogier 449. U.S. Department of Agriculture,
1990). Washington, DC. 1,024 p.
685
Schinus terebinthifolius Raddi Brazilian pepper tree
ANACARDIACEAE
John K. Francis
686
2002). Seeds from Brazil were reported to average plant species in suppressing several important
80,600 seeds/kg (Instituto de Pesquisas e Estudos pathogenic bacteria (Martínez and others 1996,
Florestais 2002). Germination of intact fruits Siddiqui and others 1995). Despite these benefits,
(without the exocarp digested or manually Brazilian pepper tree is often undesirable outside
removed) is minimal, and seeds do not remain its native range. It is so well adapted and
viable in the soil bank more than 9 months aggressive in Florida and Hawaii that it suppresses
(Panetta and Mckee 1997). Birds, particularly and replaces native vegetation, including
robins, mockingbirds, and cedar waxwings, are the endangered species. For this reason it is banned
chief dispersers of seeds in Florida (Fire Sciences from cultivation there (Fire Sciences Laboratory
Laboratory 2002). Brazilian pepper tree is 2002, Nelson 1996). Brazilian pepper tree is
propagated from both seeds and cuttings (Little reported to have an allelopathic effect on
and others 1974). It readily sprouts when cut. competing plants (National Parks Service 2002).
The plant causes skin irritation similar to poison
Growth and Management.—Brazilian pepper ivy (Toxicodendron radicans Linn.) and
tree is fast growing and in certain circumstances is respiratory difficulties in some people when it is in
capable of growing to large diameters. The U.S. bloom (Nelson 1996). It is a host of the black twig
Champion tree in Florida measures 134 cm in borer of coffee, Xylosandrus compactus (Eichoff)
diameter at breast height and 10.7 m in height (College of Tropical Agriculture and Human
(American Forests 2002). Brazilian pepper tree Resources 2002). The fruits are eaten by mammals
has shown large growth responses to inoculation and birds, but excessive feeding has been blamed
with mycorrhizal fungi and superphosphate for massive bird kills in Florida (National Parks
(Carneiro and others 1996). Hand pulling, Service 2002), and the unripe fruits can be fatal to
bulldozing, prescribed fire, and spraying with horses (Fire Sciences Laboratory 2002).
herbicides have all been used with varying degrees
of success to control Brazilian pepper tree. So far, References
biological control methods have failed (National
Parks Service 2002). American Forests. 2002. National register of big
trees. American Forests, Washington, DC. http://
Benefits and Detriments.—Widely introduced as www.americanforests.org/resources/bigtrees/
an ornamental, Brazilian pepper tree was popular regester.php?details=2297. 1 p.
for its red berries and bright green foliage. It is
still used for Christmas decorations. Because it Carneiro, M.A.C., J.O. Siqueira, A.C. Davide, L.J.
and the species growing in its understory do not Gomes, N. Curi, and F.R. do Vale. 1996.
burn readily (Fire Sciences Laboratory 2002), Mycorrhizal fungi and superphosphate and the
Brazilian pepper tree has been recommended for growth of tropical woody species. Scientia
planting as fire resistant barriers (Castronovo Forestalis 50: 21-36.
1997). Goats browse on the foliage with no ill
effects (National Parks Service 2002). The species Castronovo, T. 1997. Fire resistive landscaping
is a honey plant (Little and others 1974) and the can save your house and your life.
wood has been used for fuel, lumber, stakes, posts, http://www.themastergardenershow.com/
and railway sleepers (Fire Sciences Laboratory fire_resistive_landscaping_.htm. 5 p.
2002). The dried fruits are marketed in Brazil as a
substitute for black pepper (Piper nigrum) College of Tropical Agriculture and Human
(personal communication with J.A. Parrotta, Resources. 2002. Crop knowledge master:
USDA Forest service, Washington, DC). A Xylosandrus compactus (Eichoff). University of
decoction of bark is used in baths to relieve Hawaii. http://extento.hawaii.edu/kbase/Type/
rhumatisum and back pain (Liogier 1990). The xylosand.htm. 4 p.
essential oils from leaves and flowers are made up
of a mixture of a large number of chemicals, the
Fire Sciences Laboratory. 2002. Fire effects
most important of which are alpha-pinene (24
information database: Schinus terebinthifolius.
percent), limonene (12 percent), and p-cymene (14
Rocky Mountain Research Station, Missoula,
percent) (Singh and others 1998). Extracts of
MT. http://www.fs.fed.us/database/feis/plants/
Brazilian pepper tree were shown to be the most
tree/ schter/all.html. 9 p.
effective of a number aromatic and medicinal
687
Instituto de Pesquisas e Estudos Florestais. 2002. National Parks Service. 2002. Exotic weeds I.
Tabla de preços de sementes de espécies nativas http://www.nature.nps.gov/wv/ipm/exweeds1.ht
e exóticas. http://www.ipef.br/especies/ m. 9 p.
natexoticas.asp. 2 p.
Nelson, G. 1996. The shrubs and woody vines of
Liogier H.A. 1988. Descriptive flora of Puerto Florida. Pineapple Press, Inc., Sarasota, FL.
Rico and adjacent islands, Spermatophyta. Vol. 391 p.
2. Editorial de la Universidad de Puerto Rico,
Río Piedras, PR. 481 p. Panetta, F.D. and J. McKee. 1997. Recruitment of
the invasive ornamental, Schinus
Liogier, H.A. 1990. Plantas medicinales de Puerto terebinthifolius, is dependent upon frugivores.
Rico y del Caribe. Iberoamericana de Ediciones, Australian Journal of Ecology 22(4): 432-438.
Inc., San Juan, PR. 566 p.
Siddiqui, R.R., Uzma-Zafar, S.S. Chaudhry, and
Little, E.L., Jr., R.O. Woodbury, and F.H. Hamid-Ahamad. 1995. Antimicrobial activity of
Wadsworth. 1974. Trees of Puerto Rico and the essential oils from Schinus terebinthifolius,
Virgin Islands. Vol. 2. Agriculture Handbook Cypress sempervirens, Citrus limon, Ferula
449. U.S. Department of Agriculture, assafoetida. Part 1. Pakistan Journal of
Washington, DC. 1,024 p. Scientific and Industrial Research. 38(9-10):
358-361.
Long, R.W. and O. Lakela. 1976. A flora of
Tropical Florida. University of Miami Press, Singh, A.K., J. Singh, K.C. Gupta, and J.J.
Coral Gables, FL. 962 p. Brophy. 1998. Essential oil of leaves and
inflorescence of Schinus terebinthifolius: an
Martínez, M.J., J. Betancourt., N. A. González., exotic plant of India. Journal of Essential Oil
and A. Jauregui. 1996. Screening of some Cuban Research 10(6): 697-699.
medicinal plants for antimicrobial activity.
Journal of Ethnopharmacology 52(3): 171-174.
688
Securidaca virgata Sw. bejuco de sopla
POLYGALACEAE
John K. Francis
689
stems live 5 to 10 years or more; plants may last P. Lohr, and D. Atha. 2000. Medicinal plants
longer by resprouting. Bejuco de sopla is not used by Latino healers for woman’s health
abundant, aggressive, or weedy. Should it be conditions in New York City. Economic Botany
necessary to eliminate individual plants, grubbing 54(3): 344-357.
them out will probably be effective.
Grisebach, A.H.R. 1963. Flora of the British West
Benefits.—Bejuco de sopla contributes to Indian Islands. J. Cramer-Weinheim, New York.
biodiversity, helps protect the soil, and furnishes 789 p.
wildlife cover. It is a honey plant (Marcano-F.
1973). The roots, principally, are used to treat flu, Liogier, H.A. 1988. Descriptive flora of Puerto
laryngitis, edema, urinary infections, and as an Rico and adjacent islands, Spermatophyta. Vol.
expectorant and a diuretic (Liogier 1990). The 2. Editorial de la Universidad de Puerto Rico,
plant is prescribed by traditional Latin healers in Río Piedras, PR 481 p.
New York City to treat uterine fibroids (Balick
and others 2000). Liogier, H.A. 1990. Plantas medicinales de Puerto
Rico y del Caribe. Iberoamericana de Ediciones,
References Inc., San Juan, PR. 566 p.
690
Senecio flaccidus Less. threadleaf groundsel
ASTERACEAE
James E. Nellessen
691
from twig photosynthesis (Comstock and others singing during some ceremonial songs (Elmore
1988). 1976). The Navajo would also use the flower
heads to clean cactus fruits of their spines.
Reproduction.—Threadleaf groundsel generally
blooms in the late summer and fall but will bloom References
earlier in the summer if sufficient moisture is
available. It produces numerous yellow flower Allred, K.W. 2002. A Working Index of New
heads with eight to 17 ray florets, the rays 10 to 15 Mexico Vascular Plant Names. New Mexico
mm long (Carter 1997, Martin and Hutchins 1980- State University. http://web.nmsu.edu/~kallred/
81). Seeds (fruits) are wind-borne with a white herbweb/ [not paged].
fluffy pappus.
Carter, J.L. 1997. Trees and Shrubs of New
Growth and Management.—Threadleaf Mexico. Johnson Books, Boulder, CO. 534 p.
groundsel is a fast growing, short lived perennial.
Plants may live 3 to 6 years or more. It increases in Comstock, J.P., T.A. Cooper, and J.R. Ehleringer.
rangelands as a result of overgrazing pressure. Its 1988. Seasonal patterns of canopy development
abundance may be controlled through proper and carbon gain in nineteen warm desert shrub
grazing management and careful use of herbicides. species. Oecologia, Berlin 75(3): 327-335.
Spraying with 2,4-D has been used to control this
species. Cooper, R.A., R.J. Bowers, C.A. Beckham, and
R.J. Huxtable. 1996. Preparative separation of
Benefits and Disadvantages.—Because this pyrrolizidine alkaloids by high-speed counter
species will colonize open disturbed areas, it helps current chromatography. Journal of
achieve a quick ground cover in a natural Chromatography 732(1): 43-50.
succession process. The perennial nature of the
plant further helps stabilize soil for longer-lived Dick-Peddie, W.A. 1993. New Mexico
perennials to eventually become established. Vegetation: Past, Present and Future. University
Direct human uses are minimal, but it was once of New Mexico Press, Albuquerque, NM. 244 p.
used medicinally by Native Americans. It is a poor
forage plant for cattle and horses and is known to Ehleringer, J.R. and T.A. Cooper. 1992. On the
be toxic (Gay and Dwyer 1998, Stubbendieck and role of orientation in reducing photoinhibitory
others 1997, Warnock 1974) and causes liver damage in photosynthetic-twig desert shrubs.
damage when consumed in large quantities Plant Cell and Environment 15(3): 301-306.
(Kirkpatrick 1992). All parts of the plant are
poisonous, younger leaves more so, and poisoning Elmore, F.H. 1976. Shrubs and Trees of the
is more of a problem on drier range conditions. Southwest Uplands. Southwest Parks and
Symptoms in cattle may include a dry scaly nose, a Monuments Association, Tucson, AZ. 214 p.
rough coat, and continuous walking with no
direction (Phillips Petroleum Company 1963). Epple, A.O. 1995. A Field Guide to the Plants of
Because it has low palatability, it tends to be Arizona. Falcon Publishing Inc., Helena, MT.
avoided except during long droughts or in heavily 347 p.
overgrazed areas where it may increase in
abundance (USDA 1937). Toxicity of threadleaf Gay, C. W., Jr. and D. D. Dwyer. 1998 Reprint.
groundsel may be attributed to the pyrrolizidine New Mexico Range Plants. Cooperative
alkaloids (PA). About 50 percent of all identified Extension Service Circular 374. Revisions by:
PA are toxic, and some are carcinogenic in rodents C. Allison, S. Hatch, and J. Schickedanz. New
(Stegelmeier and others 1999). Threadleaf Mexico State University, Las Cruces, NM. 84 p.
groundsel contains the pyrrolizidine alkaloids:
senecionine, seneciphylline, florosenine, Ivey, R.D. 1995. Flowering Plants of New Mexico,
otonecine-based florosenine, and retrorsine 3rd Edition. Published by the author, NM.
(Cooper and others 1996). As with many toxic 504 p.
plants, concentration and preparation are critical
factors. For example, the Navajo would boil entire Kartesz, J.T. 1994. A Synonymized Checklist of
plants and drink the mixture to assist the voice and the Vascular Flora of the United States, Canada,
692
and Greenland, Vol. 1, 2nd Edition. Biota of Stegelmeier, B.L., J.A. Edgar, S.M. Colegate, D.R.
North America Program of the North Carolina Gardner, T.K. Schoch, R.A. Coulombe, and R.J.
Botanical Garden. Timber Press, Portland, OR. Molyneux. 1999. Pyrrolizidine alkaloid plants,
622 p. metabolism and toxicity. Journal of Natural
Toxins 8(1): 95-116.
Kearney, T.H., R. Peebles, and Collaborators.
1951 with 1960 supplement. Arizona Flora. Stubbendieck, J., S.L. Hatch, and C.H. Butterfield.
University of California Press, Berkeley, CA. 1997. North American Range Plants, 5th Edition.
1,085 p. University of Nebraska Press, Lincoln, NE.
501 p.
Kirkpatrick, Z. M. 1992. Wildflowers of the
Western Plains. University of Texas Press. Turner, B.L. and T.M. Barkley. 1990. Taxonomic
Austin, TX. 240 p. overview of the Senecio flaccidus complex in
North America, including Senecio douglasii.
Martin, W.C. and C.E. Hutchins. 1980-1981 Phytologia 69(1): 51-55.
(reprinted 2001). A Flora of New Mexico. Vol.
2. Bishen Singh Mahendra Pal Singh (India) and U.S. Department of Agriculture, Forest Service.
Koeltz Scientific Books (Germany). p. 1,277- 1937 (1988 Dover edition). Range Plant
2,591. Handbook. Dover Publications Inc., New York.
816 p.
Phillips Petroleum Company. 1963. Pasture and
Range Plants. Phillips Petroleum Co., Warnock, B.H. 1974. Wildflowers of the
Bartlesville, OK. 176 p. Guadalupe Mountains and the Sand Dune
Country, Texas. Sul Ross State University,
Alpine, TX. 176 p.
693
Senecio spartioides Torr. & Gray many-headed groundsel
ASTERACEAE
James E. Nellessen
694
Benefits.—Many-headed groundsel will colonize Ivey, R.D. 1995. Flowering Plants of New Mexico,
open disturbed areas and provide quick ground 3rd Edition. Published by the author, NM.
cover in a natural succession process. Although 504 p.
relatively short-lived the perennial nature of the
plant further helps stabilize soil for longer-lived Kartesz, J.T. 1994. A Synonymized Checklist of
perennials to eventually become established. the Vascular Flora of the United States, Canada,
Many-headed groundsel is toxic to cattle, but it is and Greenland, Vol. 1, 2nd Ed. Biota of North
infrequently eaten (USDA 1937, Kearney and America Program of the North Carolina
others 1951, Warnock 1974). This species should Botanical Garden. Timber Press, Portland, OR.
be utilized more in urban and native plant 622 p.
landscaping. When in bloom it bears, as the name
implies, numerous yellow flower heads, adding Kearney, T.H., R. Peebles, and Collaborators.
considerable color to a desert backyard. It is a 1951 with 1960 supplement. Arizona Flora.
prolific seed producer and will easily regenerate University of California Press, Berkeley, CA.
itself. This may pose a problem for some 1,085 p.
gardeners, managing the abundance of seedlings,
although many of the seedlings will not survive to Martin, W.C. and C.E. Hutchins. 1980-1981
maturity within a natural landscape anyway. (reprinted 2001). A Flora of New Mexico.
Bishen Singh Mahendra Pal Singh, India and
References Koeltz Scientific Books, Germany. Vol. 2, p.
1,277-2,591.
Allred, K.W. 2002. A Working Index of New
Mexico Vascular Plant Names. Available on a U.S. Department of Agriculture Forest Service.
New Mexico State University web page, 1937 (1988 Dover edition). Range Plant
http://web.nmsu.edu/~kallred/herbweb/ [not Handbook. Dover Publications Inc., New York.
paged]. 816 p.
Carter, J.L. 1997. Trees and Shrubs of New Warnock, B. H. 1974. Wildflowers of the
Mexico. Johnson Books, Boulder, CO. 534 p. Guadalupe Mountains and the Sand Dune
Country, Texas. Sul Ross State University,
Elmore, F.H. 1976. Shrubs and Trees of the Alpine, TX. 176 p.
Southwest Uplands. Southwest Parks and
Monuments Association, Tucson, AZ. 214 p. Weber, W.A. and R.C. Wittman. 2001a. Colorado
Flora: Eastern Slope. University Press of
Epple, A.O. 1995. A Field Guide to the Plants of Colorado, Boulder CO. 521 p.
Arizona. Falcon Publishing Inc., Helena, MT.
347 p. Weber, W.A. and R.C. Wittman. 2001b. Colorado
Flora: Western Slope. University Press of
Great Plains Flora Association. 1986. Flora of the Colorado, Boulder CO. 488 p.
Great Plains. University Press of Kansas, KS.
1,392 p.
695
Senna atomaria (L.) Irwin & Barneby flor de San José
FABACEAE
John K. Francis
General Description.—Flor de San José is also Ecology.—Flor de San José is common to rare in
known as yellow candlewood, senna-tree, palo de coastal and upland dry forests at elevations from
chivo, palo de burro, cañafístola cimarrona, near sea level to 1,100 m that receive annual
carbonera, alcaparro, chile perro, palo de zorrillo, rainfalls of about 600 to 1,000 mm. Soils, often
arguchoco, vainillo, chivato, carángano, platanillo, rocky and excessively drained, developed from
bois cabrite, casse marron, and petite casse. It is a both igneous and sedimentary parent materials
flowering, drought-deciduous shrub or small tree 2 with a wide range of textures and with pH’s that
to 12 m in height that occasionally exceeds 20 cm are mildly acid to mildly alkaline. Flor de San José
in stem diameter. The plant may or may not have is moderately intolerant of shade; it grows in the
multiple stems but almost always supports stout open, as a codominant in low forest, and in the
branches relatively low to the ground. The understory of open forests. The species may be
branches are brittle. It is supported by a tap and found in remnant and late secondary forests.
extensive lateral root system that has slender fine
roots. The roots are woody and flexible with a Reproduction.—Flor de San José flowers and
dark gray bark over a pale yellow interior. The fruits throughout the year in Nicaragua (Stevens
crown may be described as spreading and branchy and others 2001). Although flowers and fruits
but not dense. The bark is light gray. Leaves, have been observed at every season of the year in
twigs, and other vegetative parts have an Puerto Rico, individual trees normally flower and
unpleasant odor and are covered with soft hairs. fruit once per year. Because of extended
The alternate, pinnate leaves have petioles 2 to 5 flowering, flowers and fruits may be present on
cm long and two to five pairs of ovate to obovate the same tree. Seeds collected by the author from
entire leaflets, 2 to 11 cm long. The flower Puerto Rico weighed an average of 0.0238 +
clusters are corymbose racemes that bear many 0.0004 g/seed. Sown on moist filter paper, 96
flowers and arise from the bases of the upper percent germinated between 3 and 15 days after
leaves. The flowers have five unequal sepals and sowing. Germination is epigeal. A test of percent
five unequal yellow petals 8 to 15 mm long. The germination of sowings throughout the year in
fruits are flattened, linear, red-brown to black, Mexico resulted in emergence varying from 1.2 to
696
44 percent over 45 day periods depending on Arboretum, Harvard University, Jamaica Plain,
ambient temperature and precipitation (Vera and MA. 673 p.
Sánchez 1995). Reproduction may be common in
areas overgrazed by cattle. Seeds germinate after International Legume Database and Information
pods rot on the ground under mother trees Service. 2002. Senna atomaria (L.) H. Irwin &
(Stevens and others 2001) and may be dispersed to Barneby. http://www.ildis.org/LegumeWeb/
a limited degree by ruminants. 6.00/taxa/15514.shtml. 2 p.
Growth and Management.—Flor de San José Liogier, H.A. 1988. Descriptive flora of Puerto
has a moderate growth rate. Although planted as Rico and adjacent islands, Spermatophyta. Vol.
an ornamental, the author is not aware of any 2. Editorial de la Universidad de Puerto Rico,
reports of wildland plantings. Management and Río Piedras, PR. 481 p.
protection of existing stands are recommended
(Secretaría de Medio Ambiente, Recursos Little, E.L., Jr., R.O. Woodbury, and F.H.
Naturales y Pesca 2002). Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
Benefits.—Flor de San José is planted widely but 449. U.S. Department of Agriculture,
not abundantly as an ornamental for its beautiful Washington, DC. 1,024 p.
floral display and moderate size. The sapwood is
yellow and hard, and the heartwood is brown and Neal, M.C. 1965. In gardens of Hawaii. Special
hard with a density of 0.57 to 0.85 g/cm3 (Timyan Publication 50. Bernice P. Bishop Press,
1996). It is used for fuel and rude construction Honolulu, HI. 924 p.
(Holdridge and Poveda 1975, Secretaría de Medio
Ambiente, Recursos Naturales y Pesca 2002). Northern Prairie Wildlife Research Center. 2002.
Cows eat the pods but do not browse the leaves. Butterflies of North America: Yellow angled-
Some of the common names imply that goats sulphur (Anteos maerula). U.S. Geological
browse the species. Meal made from seed-free Survey. http://www.npwrc.usgs.gov.resource.
pods of flor de San José was eagerly consumed by distr/lepid/bflyusa/usa/723.htm. 3 p.
lambs in an acceptance test (Palma and Román
2002b); the meal contained 6.5 percent crude Olmos, F. 1997. Tapirs as seed dispersers and
protein, 6.8 percent fat, 4.5 percent ash, and 32.1 predators. UNCN/SSC Tapir Specialist Group.
percent crude fiber (Palma and Román 2002a). It http://www.tapirback.com/tapirgal/iucn-ssc/tsg/
has been demonstrated that Baird’s tapir (Tapirus action97/ap97-05.htm. 14 p.
bairdii) kills 100 percent of the flor de San José
seeds when it consumes the pods, apparently Palma, J.M. and L. Román. 2002a. Frutos de
because they germinate within the gut (Olmos especies arbóreas leguminosas y no
1997). Flor de San José is the principal or a key leguminosas para la alimentación de rumiantes.
larval host for the yellow angled-sulphur butterfly, Centro Universitario de Investigación y
Anteos maerula (Fabricius), and the large moth, Desarrollo Agropecuario. http://lead.
Sphingicampa montana (Packard) (Northern virtualcentre.org/es/ele/conferencia2/vbconfe15
Prairie Wildlife Research Center 2002, Savela .htm. 15 p.
2002). A tea made from the leaves is used as a
purgative (Secretaría de Medio Ambiente, Palma, J.M. and L. Román. 2002b. Prueba de
Recursos Naturales y Pesca 2002). selectividad con ovinos de pelo de harinas de
frutos de especies arbóreas. Centro
References Universitario de Investigación y Desarrollo
Agropecuario. http://www.cipav.org.co/
Holdridge, L.R. and L.J. Poveda A. 1975. Arboles redagrofor/memoirias99/PalmaJM.htm. 4 p.
de Costa Rica. Centro Cientifico Tropical, San
José, Costa Rica. 546 p. Savela, M. 2002. Sphingicampa Walsh, 1864.
http://www.funet.fi/pub/sci/bio/life/insecta/lepi
Howard R.A. 1988. Flora of the Lesser Antilles, doptera/ditrysia/bombycoidea/saturniidae/
Leeward and Windward Islands. ceratocampinae/sphingicampa/. 3 p.
Dicotyledoneae, Part 1. Vol. 4. Arnold
697
Secretaría de Medio Ambiente, Recursos Naturales 1. Missouri Botanic Garden Press, St. Louis,
y Pesca. 2002. Especies forestales no MO. 943 p.
maderables y maderables no tradicionales de
zonas áridas y semiáridas: Senna atomeria (L.) Timyan, J. 1996. Bwa yo: important trees of Haiti.
Irwin et Barneby. http://www.semarnat.gob.mx/ South-East Consortium for International
pfnm3/fichas/senna_atomaria.htm. 2 p. Development, Washington, DC. 418 p.
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. Vera-S., F. and M. Sánchez-R. 1995. Epoca de
Montiel, eds. 2001. Flora de Nicaragua. siembra de doce especies forestales de Baja
Monographs in Systematic Botany Vol. 85, No. California Sur. Revista de Ciencia Forestal en
México 20(78): 59-78.
698
Senna occidentalis (L.) Link coffee senna
FABACEAE
John K. Francis
699
dispersed by grazing animals (Sánchez and Senna species from Ethiopia. New Forests
Uranga 1993). 11(2): 155-171.
Growth and Management.—In seasonally cold Guzmán, D.J. 1975. Especies útiles de la flora
or dry climates, the life cycle of coffee senna is salvadoreña. Ministerio de Educación, Dirección
complete in 6 to 9 months, but in warm, de Publicaciones, San Salvador, El Salvador.
continually moist areas plants may last a full year 703 p.
or grow through the second year. Perhaps in Brazil
where extraordinary heights are reached, the Haselwood, E.L. and G.G. Motter, eds. 1966.
species may live a third or fourth year. Growth is Handbook of Hawaiian weeds. Experiment
moderately rapid. Plants add 0.5 to 1.5 m during Station, Hawaiian Sugar Planters’ Association,
the first season. Although coffee senna is planted Honolulu, HI. 479 p.
to yield medicinal materials (Barbadine 2002),
because of its short life and weedy potential, it is Henty, E.E. and G.H. Pritchard. 1975. Weeds of
not advisable to plant it in wildlands. The species New Guinea and their control. Botany Bulletin
can be controlled with broadleaf herbicides (Henty 7. Division of Botany, Department of Forests,
and Prichard 1975). Lae, Papua New Guinea. 189 p.
Benefits and Detriments.—As the name implies, Holm, L., J. Doll, E. Holm, J. Pancho, and J.
the seeds of coffee senna are roasted and used as a Herberger. 1997. World weeds. John Wiley &
coffee substitute (Guzmán 1975). The plant’s Sons, New York. 1,129 p.
tissues contain a host of phytoactive chemicals
that may support its numerous applications in folk Institute of Food and Agricultural Sciences. 2002.
medicine. Extracts or powdered leaves are used as Coffee senna, Cassia occidentalis L. Institute of
an analgesic, antibacterial, anti-hepatotoxic, Food and Agricultural Sciences, Cooperative
antifungal, anti-inflammatory, antiseptic, Extension Service, University of Florida,
antispasmodic, antiparasitic, antiviral, Gainesville, FL. http://edis.ifas.ufl.edu/
carminative, diaphoretic, emmenagogue, BOFY_FW008. 3 p.
febrifuge, insecticidal, immunostimulant, laxative,
purgative, sudorific, and vermifuge. Several of Liogier H.A. 1988. Descriptive flora of Puerto
these effects have been demonstrated in laboratory Rico and adjacent islands, Spermatophyta. Vol.
and clinical tests (Raintree 2002). While coffee 2. Editorial de la Universidad de Puerto Rico,
senna can be a weed of cultivated fields and Río Piedras, PR. 481 p.
plantations, it is principally a problem because it
accumulates in heavily-grazed pastures (Henty Long, R.W. and O. Lakela. 1976. A flora of
and Pritchard 1975). The foliage is poisonous and Tropical Florida. Banyon Books, Miami, FL.
generally avoided by livestock (Stevens and others 962 p.
2001). Ingestion of large amounts of seeds has
been implicated in deaths of cows, horses, and Natural Resources Conservation Service. 2002.
goats (Raintree 2002). Poisoning of pigs fed Plant profile: Senna P. Mill, senna. U.S.
coffee senna seeds resulted in muscle necrosis Department of Agriculture, Natural Resources
(Timm and Riet-Correa 1997). Conservation Service, Washington, DC.
http://plants.usda.gov/cgi_bin/plant_profile.chi?
References symbol=SENNA. 6 p.
700
Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. Timm, C.D. and F. Riet-Correa. 1997. Plants toxic
Montiel, eds. 2001. Flora de Nicaragua. to pigs. Ciencia Rural 27(3): 521-528.
Monographs of Systematic Botany Vol. 85, No.
1. Missouri Botanical Garden Press, St. Louis,
MO. p. 1-943.
701
Senna polyphylla (Jacq.) Irwin & Barneby desert cassia
FABACEAE
John K. Francis
702
Benefits.—Desert cassia is grown by commercial Department of Agriculture, Forest Service,
nurseries and widely planted as a flowering Southern Forest Experiment Station, New
ornamental in the New World Tropics. It is Orleans, LA. 5 p.
employed as an accent plant or trained into a small
background tree. The wood is light brown and Howard R.A. 1988. Flora of the Lesser Antilles,
hard and used occasionally for fence posts and Leeward and Windward Islands.
fuel. Cows apparently do not readily eat the Dicotyledoneae, Part 1. Vol. 4. Arnold
foliage of desert cassia—the species is more Arboretum, Harvard University. Jamaica Plain,
common in heavily grazed rangeland than MA. 673 p.
elsewhere.
Liogier H.A. 1988. Descriptive flora of Puerto
References Rico and adjacent islands, Spermatophyta. Vol.
2. Editorial de la Universidad de Puerto Rico,
Botanics Wholesale. 2001. http://www.botanics. Río Piedras, PR. 481 p.
com/Products/botanics_details.asp?NameAssoc
+153.htm. 1 p. Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
Francis, J.K. and A. Rodríguez. 1993. Seeds of Virgin Islands. Vol. 2. Agriculture Handbook
Puerto Rican trees and shrubs: second 449. U.S. Department of Agriculture,
installment. Research Note SO-374. U.S. Washington, DC. 1,024 p.
703
Serenoa repens (Bartr.) Small saw palmetto
PALMAE
Synonyms: None
704
(Carrington and others 2000). Fruits ripen in increase coverage of saw palmetto (Carrington and
August to November. Saw palmettos flower others 2000).
heavily about every 2 to 4 years. During these
years, individual ramets commonly produce two to Fire.—Saw palmetto naturally occurs in plant
three inflorescences. Due to high flower density communities adapted to periodic burning. Saw
(several thousand flowers/inflorescence), typically palmetto itself is highly flammable due to the
0.4 to 0.5 kg of fruit are produced on each accumulation of dead fronds that remain on the
inflorescence. In some cases, individual plants for several years (Carrington and others
inflorescences can produce up to 12 kg of fruit. 2000) and volatile waxes that cover green leaves.
Average fruit yield for a site is approximately 200 Fires commonly consume all above-ground foliage
kg/ha; however, yields can vary from less than 100 of saw palmettos, and therefore, prescribed fire is
kg/ha to more than 1,500 kg/ha (Carrington and used to reduce leaf coverage for short periods.
others 1997). Fruits are bird and mammal However, leaf growth usually begins a few days
dispersed (Tanner and others 1996). Seed after fires, and saw palmettos can regain 80
germination ranges from 20 percent after 15 percent of crown coverage the first year after
months in field conditions to 55 percent after 6 burning (Hilmon 1968). Fire also benefits fruit
months under lab conditions (Hilmon 1968). Seeds production of saw palmetto, especially within the
can remain viable for up to 1 year, albeit with first year following burns (Carrington and others
reduced germination rates (Carrington and others 2000). However, flowering may be reduced for
2000). Seed germination may be enhanced by several years after burns (Abrahamson 1999) as
passing through animal digestive systems (Tanner palmettos recover from both burning and heavy
and others 1996). fruit production. Frequent burning (every 1 to 4
years) curtails flowering and fruiting by keeping
Growth and Management.—Growth of saw carbohydrate reserves low (Hilmon 1968).
palmetto is slow, ranging between 0.6 and 2.2
cm/year in stem elongation (Abrahamson 1995, Detriments and Benefits.—Saw palmetto was
Hilmon 1968). Based on these estimates, some viewed by early settlers in the Southeastern United
saw palmettos may be 500 to 700 years old States as an obstacle to establishing agricultural
(Abrahamson 1995, Tanner and others 1996). In fields, cattle pastures, and home sites. Ranchers
some cases, land managers may want to reduce the continue to regard it as a competitor of native
density and cover of saw palmetto in order to forage grasses, and foresters cite that it inhibits
improve cattle forage and/or timber production or pine regeneration. Despite these detriments, saw
to reduce fire hazard. Because saw palmetto is palmetto is viewed as a significant pharmaceutical
fire-tolerant, mechanical or chemical treatments resource in Florida and southern Georgia due to
are often needed for control. Roller-chopping the effectiveness of certain fruit compounds (such
reduces abundance of saw palmetto (Hilmon and as free fatty acids, phytosterols) in treating benign
others 1963, Lewis 1970) by crushing vegetative prostatic hyperplasia (Tasca 1985, Braeckman
parts and removing terminal buds. This treatment 1994, Wilt and others 1998), or swelling of the
is particularly effective during periods of low soil prostate gland. In 1995, saw palmetto fruits
moisture (Moore 1974) or when followed by collected for pharmaceuticals sold for over $6/kg.
prescribed burning. Among herbicide treatments, Total estimated value of fruits sold in 1996 was
repeated cycles of 2,4,5-T followed by burning approximately $5 million (Carrington and others
will reduce palmetto cover (Altobellis and Hough 2000). In addition to its medicinal value, saw
1968), but this chemical is no longer available. palmetto can be a significant source of honey
Metsulfuron, often sold as Escort, is the most production (Bennett and Hicklin 1998). Saw
effective herbicide currently available for control palmetto is also a good native plant for
of saw palmetto (personal communication J.L. enviroscaping due to its natural drought- and
Michael, Southern Research Station, Auburn, AL). insect-resistance, and low requirements for
In other cases, land managers may want to fertilization (Tanner and others 1996). Saw
enhance saw palmetto flowering and fruiting. The palmettos serve as nesting and denning habitat for
most cost-efficient practice to increase fruit over 100 animal species, including: the
production is prescribed burning (Carrington and endangered Florida grasshopper sparrow
others 2000). Optimal burning frequency is every (Ammodramus savannarum Gmein), Florida
5 to 8 years. Soil fertilization has been used to panther (Felis concolor Linnaeus), Florida
705
woodrat (Neotoma floridana Ord), wild turkey Fisher, J.B. and P.B. Tomlinson. 1973. Branch
(Meleagris gallopavo Linnaeus), and white-tailed and inflorescence production in saw palmetto
deer (Odocoileus virginianus Zimmermann) (Serenoa repens). Principes 17: 10-19.
(Tanner and others 1996). Fruits are eaten by black
bears (Ursus americanus Pallas), white-tailed deer Hilmon, J.B. 1968. Autecology of saw palmetto
(Odocoileus virginianus), raccoons (Procyon lotor [Serenoa repens (Bartr.) Small]. Ph.D.
Linnaeus), turkeys (Meleagris gallopavo), bob- dissertation. Duke University, Durham, N.C.
white quail (Colinus virginianus L.), gray foxes 191 p.
(Urocyon cinereoargenteus Schreber), opossum
(Dasypus novemcinctus Linnaeus), and gopher Hilmon, J.B., C.E. Lewis, and J.E. Bethune. 1963.
tortoises (Gopherus polyphemus Daudin) (Maehr Highlights of recent results of range research
and Layne 1996). in Southern Florida. Society of American
Foresters Proceedings 1962: 73-76.
References
Lewis C.E. 1970. Responses to chopping and rock
Abrahamson, W.G. 1995. Habitat distribution and phosphate on south Florida ranges. Journal of
competitive neighborhoods of two Florida Range Management 23: 276-282.
palmettos. Bulletin of the Torrey Botanical
Club 122: 1-14. Maehr, D.S. and J.N. Layne. 1996. Florida’s all-
purpose plant the saw palmetto. Palmetto
Abrahamson, W.G. 1999. Episodic reproduction in (Fall): 16 –10, 15, 21.
two fire-prone palms, Serenoa repens and
Sabal etonia (Palmae). Ecology 80:100-115. McNab, W.H. and M.B. Edwards. 1980. Climatic
factors related to the range of saw-palmetto
Altobellis, A.T. and W.A. Hough. 1968. [Serenoa repens (Bartr.) Small]. American
Controlling palmetto with fire and herbicides. Midland Naturalist 103: 205-208.
Georgia Forestry Research Paper 52. Georgia
Forest Research Council, Macon, Ga. Moore, W. H. 1974. Some effects of chopping
saw-palmetto pineland threeawn range in
Bennett, B.C. and J.R. Hicklin. 1998. Uses of saw south Florida. Journal of Range Management
palmetto (Serenoa repens, Arecaceae) in 27(2): 101-104.
Florida Economic Botany 52: 381-393.
Small, J.K. 1926. The saw-palmetto–Serenoa
Braeckman, J. 1994. The extract of Serenoa repens repens. Journal of the New York Botanical
in the treatment of benign prostatic Garden 27: 193-202.
hyperplasia: A multicenter open study.
Current Therapy Research 55: 776-785. Small, J.K. 1933. Manual of the southeastern
flora. John Kunkel Small, New York.
Carrington, M.E., T.D. Gottfried, and J.J. 1,554 p.
Mullahey. (In review) Pollination biology of
saw palmetto (Serenoa repens: Palmae) in Tanner, G.W., J.J. Mullahey, and D. Maehr. 1996.
southwestern Florida. Palms. Saw-palmetto: an ecologically and
economically important native palm. Circular
Carrington, M.E., J.J. Mullahey, G. Krewer, B. WEC-109, Florida Cooperative Extension
Boland, and J. Affolter. 2000. Saw palmetto Service, Institute of Food and Agricultural
(Serenoa repens): an emerging forest resource Sciences, University of Florida, Gainesville,
in the southeastern United States. Southern FL. 7 p.
Journal of Applied Forestry 24(3): 129-134.
Tasca, A. 1985. Treatment of obstructive
Carrington, M.E., J.J. Mullahey, and F. Roka. symptomatology in prostatic adenoma with an
1997. Saw palmetto: A fountain of youth. extract of Serenoa repens. Minerva Urologica
Proceedings American Forage Grassland e Nefrologica 37: 87-91.
Council 6: 233-237.
706
Wilt, T., A. Ishani, and C. Mulrow. 1998. Saw Journal of the American Medical Association
palmetto extracts for treatment of benign 280: 1,604-1,609.
prostatic hyperplasia: a systemtatic review.
707
Serjania polyphylla (L.) Radlk. bejuco de corrales
SAPINDACEAE
John K. Francis
708
spin and fly sideways as they descend and may be a diuretic and purifier of the blood in herbal
travel a considerable distance before reaching the medicine (Liogier 1990).
ground.
References
Growth and Management.—Seedlings grow
about 0.5 m in their first year. Older plants and Acevedo-Rodríguez, P. 1985. Los bejucos de
sprouts grow 2 or 3 m each year. Individual stems Puerto Rico. Vol. 1. General Technical Report
can live for several years; by sprouting, plants SO-58. U.S. Department of Agriculture, Forest
may survive for at least several decades. No Service, Southern Forest Experiment Station,
planting experience for the species has been New Orleans, LA. 331 p.
reported. In the past, farmers controlled bejuco de
corrales by repeated cutting with a machete. These Liogier, H.A. 1990. Plantas medicinales de Puerto
vines could probably be killed faster by spraying Rico y del Caribe. Iberoamericana de Ediciones,
the sprouts that arise after cutting with broadleaf Inc. San Juan, PR. 566 p.
weed killer.
Liogier, H.A. 1994. Descriptive flora of Puerto
Benefits.—The strong and flexible stems of Rico and adjacent Islands. Vol. 3. Editorial de la
bejuco de corrales are suitable for basketry, and Universidad de Puerto Rico, Río Piedras, PR.
some of the common names indicate that it was 461 p.
used for this purpose. Bejuco de corrales is
considered to be a honey plant in the Dominican Marcano Fondeur, E. de J. 1973. La flora apícola
Republic (Marcano Fondeur 1973). Livestock eat de la República Dominicana. http://
young plants and the foliage of older ones that are marcano.freeservers.com/nature/estudios/apicola
within their reach. Bejuco de corrales is reputed to /dicotsp.htm. 11 p.
709
Sesbania sericea (Willd.) Link silky sesban
FABACEAE
John K. Francis
710
However, relatively few of the seedlings progress experimentally intercropped with maize and
past the early seedling stage. resulted in no depression in maize yield. In
Guyana, the species grows in upland rice fields
Growth and Management.—Plants of silky but does not occur in flooded rice paddies (Evans
sesban grow rapidly. Seedlings are about 10 cm and Rotar 1987). The stemwood of silky sesban is
tall at the end of 1 month and reach 2 to 6 m in brash, has a relatively low specific gravity, and
height in about 6 months. Silky sesban was offers few prospects for commercial use.
reported to yield 26.8 Mg/ha (fresh weight) in a
84-day growing period (Evans and Rotar 1987). References
After flowering, height growth of silky sesban
ceases. Plants live from about 8 months to a little Evans, D.O. and P.P. Rotar. 1987. Sesbania in
over a year, depending on conditions. When their agriculture. Tropical Agriculture Series 8.
seeds mature, the plants die. The species may Westview Press, Boulder, CO. 192 p.
grow and die in an annual cycle timed with wet
and dry seasons, or it may simply grow in Howard, R.A. 1988. Flora of the Lesser Antilles,
response to favorable soil conditions as they Leeward and Windward Islands.
occur. If land managers find it necessary to Dicotyledoneae, Part 1. Vol. 4. Arnold
establish silky sesban, sowing into a moist, Arboretum, Harvard University. Jamaica Plain,
prepared seedbed should be sufficient. MA. 673 p.
Benefits.—The species of Sesbania, including Liogier, H.A. 1988. Descriptive flora of Puerto
silky sesban, are used as green manure in India. Rico and adjacent islands, Spermatophyta. Vol.
Silky sesban is readily consumed by cattle in 2. Editorial de la Universidad de Puerto Rico,
pastures. The closely related S. sesban (L.) Merr. Río Piedras, PR 481 p.
has been shown to have moderately good
nutritional values. Some of the species of Sesbania Long, R.W. and O. Lakela. 1976. A flora of
have toxic saponins and canavanine in their seeds. Tropical Florida. Banyon Books, Miami, FL.
So far, the seeds of silky sesban have not been 962 p.
reported to be toxic. Crushed seeds of this species
were fed to chicks at 1 percent of body weight Nelson, G. 1996. The shrubs and woody vines of
each day for 3 days without any signs of toxicity Florida. Pineapple Press, Inc. Sarasota, FL.
(Evans and Rotar 1987). Silky sesban can be 391 p.
weedy but usually causes few problems. It was
711
Sida fallax Walp 'ilima
MALVACEAE
environments.
712
recognized. These included 'ilima-ku-kala, ilima- Anonymous. 2002. ttp://www.anglefire.com/hi4/
lei, 'ilima-ku-kahakai and 'ilima-koli-kukui nhps/news122 k. html [not paged].
(Stephens 2000). 'Ilima-ku-kala is the wild tall Krauss, B. 1998. How to plant a native Hawaiian
form found in the mountains. 'Ilima-ku-kahakai is garden. http://hawaii.gov/health/
the creeping form found along the coastline. oeqc/garden/index.html
'Ilima-lei were cultivated by the Hawaiians for use
in lei making. 'Ilima-koli-kukui was another Krauss, B. 2000. Native plants used as medicine in
cultivar with reddish brown flowers (Stephens Hawaii. http://library.kcc.hawaii.edu/~
2000). 'Ilima is considered a special flower of the soma/krauss/ilima.html. [not paged]
island of Oahu. 'Ilima leis are among the most
treasured leis and are difficult to make, requiring Sherry, K., J.M. Castillo, and R.B Shaw. 1999.
over a thousand blossoms. At one time, only Effects of wildfire on vegetation and rare plants
royalty were allowed to wear an 'ilima lei in arid montane shrubland. Pohakaloa Training
(Anonymous 2001). 'Ilima was also used Area, Hawaii. 1999 Hawaii Conservation
medicinally. The juice squeezed from the flowers Conference, Honolulu, HI.
was used as a mild laxative for children, and the
root bark mixed with flowers was used to treat Stephens, M. 2000. The comparative ecophsiology
asthma (Krauss 2000). Buds were chewed to of mountain and coastal populations of Sida
quench thirst on hot, dry days. Stems were used to fallax Walp. (Malvaceae). M.S. thesis.
make baskets. 'Ilima is said to be one of the forms University of Hawaii, Hilo, HI.
that Laka, the goddess of hula, could take at will
(Quensell 2000). Quensell, N. 2000. Native Hawaiian plants.
http://www.kcc.hawaii.edu/campus/tour/plants/p
Detrimental Effects.―‘Ilima is known to be host ilima.htm [not paged]
to nonnative rust, Puccinia heterospora, which is
now becoming established on native plants. Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
Infection may be heavy and conspicuous. 1990. Manual of the Flowering Plants of
Hawaii. University of Hawaii Press, Honolulu,
References HI. p. 897-898.
713
Sida rhombifolia L. arrowleaf sida
MALVACEAE
John K. Francis
714
failed to induce germination. However, acid an alkaloid content in the root of 0.1 percent and
scarification, heating, and cold storage succeeded the presence of choline, pseudoephedrine, beta-
in breaking dormancy. Some 80 percent phenethylamine, vascin, hipaphorine and related
germination was obtained at the best soil depth for indole alkaloids (Shaman Australis
germination, which was 0.5 to 2 cm (Holm and Ethnobotanicals 2002). Perhaps because of these
others 1997). Arrowleaf sida plants sprout chemicals, arrowleaf sida is unpalatable to cattle
vigorously if cut. The seeds are dispersed by (Kuniata and Rapp 2001). Arrowleaf sida has
water, farm machinery, ruminant animals, and in significant medicinal applications for which it is
impure agricultural seed. Ants disperse the seeds cultivated throughout India. The pounded leaves
in Africa (Holm and other 1997). are used to relieve swelling, the fruits are used to
relieve headache, the mucilage is used as an
Growth and Management.—Arrowleaf sida emollient, and the root is used to treat rheumatism
plants may reach 0.5 m or more in their first year. (Parrotta 2001). Australian Aborigines use the
Growth is most rapid in warm conditions (days of herb to treat diarrhea. Leaves are smoked in
30 °C and nights of 25 °C). Growth nearly ceases Mexico and a tea is prepared in India for the
below 20 °C. Plants survive frosts and winters as stimulation it provides (Shaman Australis
far north as Tennessee (Holm and others 1997). Ethnobotanicals 2002).
Although the species is able to perpetuate itself as
an annual in difficult climates and under cultivated References
agriculture, the aerial portions of individual plants
in favorable climates may live 3 years, and Food and Agriculture Organization. 2002. Cultural
possibly more by resprouting. Control, but not practices. http://www.fao.org/ag/AGP/AGPC/
elimination of arrowleaf sida in plantations and doc/Publicat/FAOGUL2/B204.htm. 2 p.
pastures may be achieved by hand weeding and
herbicide spraying. Hand pulling and mowing are Guzmán, D.J. 1975. Especies útiles de la flora
only partially effective because arrowleaf sida is Salvadoreña. Ministerio de Educación,
difficult to pull and quickly sprouts after cutting. Dirección de Publicaciones. San Salvador, El
The weed problem from arrowleaf sida becomes Salvador. 703 p.
more severe in reduced tillage agriculture (Holm
and others 1997). Mowing or chain slashing is Holm, L., J. Doll, E. Holm, J. Pancho, and J.
recommended in pastures to suppress arrowleaf Herberger. 1997. World weeds. John Wiley and
sida and other unpalatable Sida species and allow Sons, Inc. New York. 1,129 p.
pasture forage plants to grow (Food and
Agriculture Organization 2002). Effective Howard, R.A. 1989. Flora of the Lesser Antilles,
biological control has been obtained in Australia Leeward and Windward Islands. Vol. 5. Arnold
and Papua New Guinea by introducing the leaf- Arboretum, Harvard University, Jamaica Plain,
eating beetle Calligrapha pantherina, which feeds MA. 604 p.
exclusively on three species of weedy Sida’s
(Kuniata and Rapp 2001). Kuniata, L. and G. Rapp. 2001. Biocontrol of Sida
rhombifolia in Papua New Guinea. http://w
Benefits.—Arrowleaf sida stems are used as ww.nt.gov.au/dpif/pubcat/agnotes/ 542.htm. 4 p.
rough cordage, sacking, and for making brooms.
The stems have a high quality fiber and were once Liogier, H.A. 1994. Descriptive flora of Puerto
exported from India and elsewhere as “hemp” Rico and adjacent Islands. Vol. 3. Editorial de la
(Guzmán 1975, Holm and others 1997). Chemical Universidad de Puerto Rico, Río Piedras, PR.
analysis revealed that the leaves contain 461 p.
respectable amounts of nutrients: 74,000 to
347,000 ppm protein, 94,000 to 475,000 ppm Parrotta, J.A. 2001. Healing plants of Peninsular
carbohydrates, 33,000 to 167,000 ppm fiber, India. CABI Publishing, Wallingford, UK and
14,000 to 71,000 ppm fat, and 16,000 to 81,000 New York. 917 p.
ppm ash. However, it was reported that the root
contained 450 ppm alkaloids and the presence of Shaman Australis Ethnobotanicals. 2002. Sida
ephedrine and saponin (Southwest School of rhombifolia, common sida. http://www.shaman-
Botanical Medicine 2002). Another source reports australis.com/Website/Sidarhombifolia.htm. 1 p.
715
Southwest School of Botanical Medicine. 2002.
Constituents. http://www.rt66.com/hrbmoore/
Constituents/Sida_rhombifolia.txt. 1 p.
716
Smilax domingensis Willd. bejuco de membrillo
SMILACACEAE
John K. Francis
717
establishment of the species. The seeds must be Rico y del Caribe. Iberoamericana de Ediciones,
collected by hand and can be cleaned by Inc., San Juan, PR 566 p.
maceration and wet sieving.
Missouri Botanical Garden. 2002. Flora de Costa
Benefits.—Bjuco de membrillo adds to Rica: Smilax domingensis Willd. http://mobot.
biodiversity, helps to protect the soil, and furnishes org/manual.plantas/051162/S051173.html. 10 p.
food and cover for wildlife. Extracts of the plant
are used as a tonic, a sudorific, and a purifier, and Missouri Botanical Garden. 3002. W3 specimen
to treat skin conditions, rheumatism, and venereal data base--28 Aug. 2003. http://mobot.
disease (Liogier 1990) mobot.org/cgi_bin/search_vast. [not paged].
718
Solanum drymophilum O.E. Schulz erubia
SOLANACEAE
Synonyms: none
John K. Francis
719
References Agriculture, Forest Service, Agriculture
Handbook. 449. Washington, DC. 1,024 p.
Liogier, H.A. 1995. Descriptive flora of Puerto
Rico and adjacent islands. Vol. 4. Editorial de la Miner Solá, E. 1999. Arboles y plantas en peligro
Universidad de Puerto Rico, Río Piedras, PR. de extinción en Puerto Rico. Puerto Rico
617 p. Ecológico Vol. 3. First Book Publishing of
Puerto Rico, San Juan, PR. 91 p.
Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the U.S Fish and Wildlife Service. 2002. Species
Virgin Islands. Vol. 2. U.S. Department of accounts: Erubia (Solanum drymorphilum).
http://endangered.fws.gov/i/q/saq50.html. 2 p.
720
Solanum dulcamara L. bittersweet nightshade
SOLANACEAE
Synonyms: none
John K. Francis
721
Reproduction.—Bittersweet nightshade flowers glycoside dulcamarin (responsible for the bitter-
from May to September (Horticulture Purdue sweet taste). Decoctions, usually of the twigs, were
1998). Fruit and seed production can be abundant. once used to treat a variety of ailments. These have
Fresh fruits collected by the author in Utah (n = been largely discontinued because of severe side
50) averaged 0.441 + 0.013 g/fruit. They contained effects. The herb is used today to treat scaly skin
an average of 30.9 + 1.6 seeds/fruit, and the air- afflictions and deficient capillary circulation in the
dried seeds averaged 0.00149 + 0.00002 g/seed or skin (Herbdata New Zealand 2003).
670,000 seeds/kg. Crossley (1974) reports 938,000
seeds/kg. The fruits can be collected by hand, References
cleaned by macerating and wet screening, and
stored at 6 percent moisture for periods of less Abrams, L. 1951. Illustrated flora of the Pacific
than 1 year. When seeds are germinated under States. Vol. 3. Stanford University Press,
light, stratification is not necessary and total Stanford, CA. 866 p.
germination percentages of 61 to 98 percent may
be reached. Stems and branches layer (root) Crossley, J.A. 1974. Solanum dulcamara L. In:
whenever they come in contact with the ground. C.S. Schopmeyer, tech. coord. Seeds of woody
Plants coppice when they are cut or damaged. plants of the United States. Agriculture
Nursery propagation is usually by seeds, but the Handbook 450. U.S. Department of Agriculture,
species can be propagated asexually by root or Forest Service, Washington, DC. p. 777-778.
stem cuttings (Crossley 1974). Seeds of wild
plants are dispersed by birds (Samodien and others Dean, C.C. 1940. Flora of Indiana. Department of
2003). Conservation, Division of Forestry,
Indianapolis, IN. 1,236 p.
Growth and Management.—Bittersweet
nightshade branches grow and die back 1 to 3 m or Herbdata New Zealand. 2003. Solanum
more each year. Larger stems examined by the dulcamara, Linn. http://www.herbdatanz.com/
author had two or three growth rings. Plants Bittersweet.html. 4 p.
probably live much longer by resprouting,
suckering, and layering. Bittersweet nightshade Horticulture Purdue. 1998. Bitter nightshade.
usually is not abundant or aggressive enough to Purdue University, West Lafayette, IN. http://
need control. Occasional plants or patches in www.hort.purdue.edu/newcrop/herbhunters/
gardens, orchards, or landscaped areas can be nightshade.html. 1 p.
eliminated by grubbing or spot spraying with
broadleaf or broad-spectrum herbicides. Kollmann, J. and P.J. Grubb. 1999. Recruitment of
fleshy-fruited species under different shrub
Benefits and Detriments.—Bittersweet species: control by under-canopy environment.
nightshade contributes to the aesthetics of Ecological Research. http://dogwood.ag.
wildlands where it grows and furnishes food and utk.edu/literature/1999/99_kollmann.htm. 1 p.
cover for wildlife. It is a pretty plant, cultivated as
an ornamental for its colorful flowers and fruits Lambo Seeds. 2003. Genus: Solanum. http://www.
and dark green foliage. Several species of moths aros.net/~lambo/dulcamara/dulcamara01.htm.
use it for larval food (Savela 2003). The fruits are 8 p.
eaten by birds, including pheasants (Phasianus
colchicus). Although all parts are poisonous to Natural Resources Conservation Service. 2003.
cattle, horses, and sheep, which normally will not Plants profile: Solanum dulcamara L., climbing
eat it, the stems are eaten by muskrats (Ondatra nightshade. http://plants.usda.gov.cgi_bin/
zibethica). The berries have been used to poison plant_profile.cgi?symbol=SODU. 6 p.
rabbits and dogs (Steyermark 1963). Although the
fruits have an attractive appearance, the flavor is Samodien, J., M. Wood, K. Epps, and M. Thandy.
so disagreeable that it is doubtful that anyone 2003. An introduction to common exotic species
would mistakenly eat enough to be poisoned. Also, to British Columbia’s ecosystems. University
poisonous principals are low in ripe fruits (Lambo College of Fraser Valley, Abbotsford, BC,
Seeds 2003). Bittersweet nightshade contains the Canada. http://www.ucfv.bc.ca/biology/Biol210/
alkaloid solanine (the poisonous principal) and the 1999/Exotic/Exotic_plant.htm. 9 p.
722
Savela, M. 2003. Solanum L. http://www. Stance, C. 1997. New flora of the British Isles. 2nd.
funet.fi/pub/sci/bio/life/plants/magnoliophyta/ Ed. Cambridge University Press, Cambridge,
magnoliophytina/magnoliopsida/solanaceae/ UK. 1,130 p.
solanum/. 5 p.
Steyermark, J.A. 1963. Flora of Missouri. The
Iowa State University Press, Ames, IA. 1,725 p.
723
Solanum persicifolium Dunal berenjena de playa
SOLANACEAE
John K. Francis
are lateral and contain up to 15 small, blue, violet,
or rarely white flowers. The fruits are globose, red
berries about 5 mm in diameter with the calyx still
attached. Each contains several flattened, yellow
seeds (author’s observation, Liogier 1995).
724
been published. Small areas of disturbance in References
suitable habitat would probably encourage natural
establishment. The species is rarely common Liogier, H.A. 1995. Descriptive flora of Puerto
enough to control as a weed in pastures or Rico and adjacent islands. Vol. 4. Editorial de la
plantations. Universidad de Puerto Rico, Río Piedras, PR.
617 p.
Benefits.—Berenjena de playa contributes to the
biodiversity of habitat where it grows, helps to Vázquez, O.J. and D.A. Kolterman. 1998. Floristic
protect the soil, and furnishes food and cover for composition and vegetation types of the Punta
wildlife, particularly birds. Guaniquilla Natural Reserve—Cabo Rojo,
Puerto Rico. Caribbean Journal of Science 34(3-
4): 265-279.
725
Solanum torvum Sw. turkey berry
SOLANACEAE
John K. Francis
726
disturbed ground. Frugivorous birds eat the fruits of the interspecific eggplant hybrid Solanum
and spread the seeds (Pacific Island Ecosystems at melongena x Solanum torvum.
Risk 2001). Turkey berry can be propagated http://www.actahort.org/books/467/467_17.htm.
vegetatively by placing branch cuttings, with or 1 p.
without leaves, in a mist chamber for one month
(Badola and others 1993). CPR Environmental Education Centre. 2001.
Medicinal plants. http://cpreec.org/edu/
Growth and Management.—Turkey berry grows medi-pln.htm. 4 p.
about 0.75 to 1.5 m in height per year. The species
is not long-lived; most plants live about 2 years. Horzog, F. and D. Gautier-Béguin. 2001.
Physical control of the shrub may be done by Uncultivated plants for human nutrition in Côte
grubbing out the plants; lopping will not kill them. d’Ivorie. http://www.fao.org/docrep/W3735e/
They can be killed by translocated herbicides w3735e10.htm. 12 p.
applied to the leaves or the cut stumps (Pacific
Island Ecosystems at Risk 2001). Howard, R.A. 1989. Flora of the Lesser Antilles,
Leeward and Windward Islands.
Benefits.—The wood is soft and light and of little Dicotyledoneae. Part 3. Vol. 6. Arnold
use except for emergency fuel. The fruits are Arboretum, Harvard University, Jamaica Plain,
edible and used in Thai cuisine (Royal MA. 658 p.
Horticultural Society 2001) and incorporated into
soups and sauces in the Ivory Coast (Herzog and Liogier, H.A. 1995. Descriptive flora of Puerto
Gautier-Béguin 2001). Turkey berry contains a Rico and adjacent islands. Vol. 4. Editorial de la
number of potentially pharmacologically active Universidad de Puerto Rico, Río Piedras, PR.
chemicals including the sapogenin steroid, 617 p.
chlorogenin (Badola and others 1993). Aqueous
extracts of turkey berry are lethal to mice or Liogier, H.A. 1990. Plantas mediciales de Puerto
depress the erythrocytes, leukocytes and platelets Rico y del Caribe. Iberoamericana de Ediciones,
in their blood (Tapia and others 1996). A related Inc., San Juan, PR. 566 p.
chemical, cholecalciferol, is the active ingredient
in a number of commercial rodentacides Little, E.L., Jr., R.O. Woodbury, and F.H.
(American Board of Veterinary Toxicology 2001). Wadsworth. 1974. Trees of Puerto Rico and the
Extracts of the plant are reported to be useful in Virgin Islands. Vol. 2. Agriculture Handbook
the treatment of hyperactivity (Null 2001), colds 449. U.S. Department of Agriculture,
and cough (CPR Environmental Education Centre Washington, DC. 1,024 p.
2001), pimples, skin diseases, and leprosy (Liogier
1990). Turkey berry is being crossed with Null, G. 2001. The biochemical activity of plants.
eggplant in an attempt to incorporate genes for http://www.garynull.com/Documents/phytoche
resistance to Verticillium wilt into the vegetable micals/Phytochemicals5.htm. 15 p.
(Bletsos and others 2001).
Pacific Island Ecosystems at Risk. 2001. Invasive
References plant species: Solanum torvum Sw., Solanaceae.
http://www.hear.org/pier/sotor.htm. 2 p.
American Board of Veterinary Toxicology. 2001.
An overview of cholecalciferol toxicosis. Royal Horticultural Society. 2001. Fruits and
http://www.abvt.org/tow.htm. 6 p. vegetables: felty leaves.
http://www.rhs.org.uk/about/mn_pubs_journals_
Badola, K.C., P. Mohinder; H.C.S. Bhanderi, and garden_0898_aubergine_b.asp. 1 p.
M. Pal. 1993. Vegetative propagation of
ranbaigan (Solanum torvum Sw.) by rooting
branch cuttings. Indian Forester 119(12): 1,027-
1,028.
728
Sorbus scopulina Greene Greene’s mountain-ash
ROSACEAE
John K. Francis
729
collection made by the author in Utah, fresh fruits protect the soil, adds to the aesthetics of wildland
weighed an average of 0.333 + 0.009 g/fruit, and sites, especially with its yellow to orange-red fall
air-dried seeds weighed an average of 0.00386 + foliage and red-orange berries, and furnishes cover
0.0001 g/seed (259 seeds/g). There were 24 to 117 for wildlife. The foliage and twigs are browsed by
fruits/cluster and an average of 2.4 seeds/fruit. deer (Odocoileus spp.), elk (Cervus canadensis),
Seed will remain viable in sealed, refrigerated moose (Alces americana), and to a lesser extent,
containers at low relative humidity for 5 years. cattle. The fruits are eaten by black bears
Recommended pretreatment is a 3:1 (Euarctos americanus), martin (Martes
water/hydrogen peroxide soak for 10 minutes americana), fisher (Martes pennanti), rodents,
followed by a 25-hour water soak and a cold American robin (Turdus migratorius), hermit
stratification in peat at 3 °C for 90 to 120 days. (Catharus guttatus), gray-cheeked (Catharus
Germination varies from 90 to 100 percent (Luna minimus) and Swainson’s (Catharus ustulatus)
and Wick 2001). A test of viability can be made by thrushes, pine grosbeaks (Pinicola enucleator),
incubating imbibed, excised embryos at 20 °C for Bohemian waxwings (Bombycilla garrulus),
6 days. Viable embryos either retain their freshly northern flickers (Colaptes auratus), Steller’s jays
excised appearance or become deep green; (Cyanocitta stelleri), and blue grouse
nonviable embryos deteriorate or turn pale yellow (Dendragapus obscurus) (Alaska Department of
green (Umatilla National Forest 2003). In the wild, Fish and Game 2001, Gullion 1964, Stein, 2003,
seeds are dispersed mainly by birds. Greene’s Wier 2003). The fruits are edible to humans, fresh,
mountain-ash may be asexually reproduced using cooked, and dried, but it is necessary to wait until
cuttings. Late-summer semihardwood stem the bitterness disappears after multiple frosts
cuttings 15 to 25 cm long and 0.3 to 1.3 cm in (Moser 2003). A wine is also made from the
diameter are recommended. After hormone berries (Keller 2003). In herbal medicine,
treatment, cuttings placed in a bottom-heated infusions of bark have been used to reduce fevers
mistbed for 6 weeks yielded 47 percent rooted and as a tonic, and infusions of branches have been
cuttings (Luna and Wick 2001). given to children with bed-wetting problems
(Ibiblio 2003). Greene’s mountain-ash is planted
Growth and Management.—This species grows to a limited extent as an ornamental, especially in
slowly as seedlings and only at an intermediate naturalistic landscape settings. The wood is soft
rate later. Plants are able to flower and fruit after 3 and has a specific gravity of 0.59 g/cm3 (Ibiblio
to 5 years (Luna and Wick 2001). Greene’s 2003). It is probably useful for firewood, but other
mountain-ash is usually grown for revegetation uses are not known.
projects from seed in nurseries as container or
bare-root stock. Potted seedlings can be expected References
to reach 15 cm tall with a 2.0 cm caliper by the
end of the first year (Luna and Wick 2001). They Alaska Department of Fish and Game. 2001.
are outplanted as either 1- or 2-year-old stock. Native Alaskan and exotic plants used by
Greene’s mountain-ash planted in seven sites in wildlife. http://www.state.ak.us/adfg/wildlife/
Alberta averaged 1.2 to 2.3 m in height after 4 to 8 geninfo/birds.htm. 9 p.
years of growth (Alberta Agriculture, Food and
Rural Development 2003). Direct seeding is also Alberta Agriculture, Food and Rural Development.
possible. Drilling seed 2 mm deep in fertile, well- 2003. Evaluating woody plants for hardiness and
drained soil is recommended. Many seeds will not landscaping quality in Alberta: Sorbus sp.
germinate until the second or third year (Umatilla (mountain ash). http://www.agric.gov.ab.ca/
National Forest 2003). Although the species is crops/trees/rwptp/ sorbus.html. 3 p.
adapted to many soil conditions, it is sensitive to
high pH in the nursery and the field and is Browse-Shrub and Forb Committee of the
damaged by the pear slug [Caliroa cerasi (L.)] Association of Official Seed Analysis. 1985.
(Alberta Food and Rural Development 2003). Handbook on seeds of browse-shrubs and forbs.
Browsing keeps the plants compact in many Technical Publication R8-TP8. U.S. Department
locations (Clark 1976). The author knows of no of Agriculture, Forest Service, Southern Region,
published data on management of natural stands. Atlanta, GA. 246 p.
Benefits.—Greene’s mountain-ash is an important Clark, L.J. 1976. Wild flowers of the Pacific
component of the Western shrub community and Northwest. Gray’s Publishing, Limited. Sidney,
furnishes a number of benefits. The species helps BC, Canada. 604 p.
730
Davis, R.J. 1952. Flora of Idaho. Brigham Young ash. http://plants.usda.gov/cgi_bin/plant_profile.
University Press, Provo, UT. 836 p. cgi?symbol=SOSC2. 4 p.
Gullion, G.W. 1964. Wildlife uses of Nevada Stein, W.I. 2003. Sorbus L, mountain-ash. In: F.T.
flora. Contr. Toward a Flora of Nevada 49. U.S. Bonner, and R.G. Nisley, eds. Woody plant seed
National Arboretum, Washington, DC. 170 p. manual. U.S. Department of Agriculture, Forest
Service, Washington, DC. http://wpsm.net/
Ibiblio. 2003. Plants for a future: database search Sorbus.pdf. 12 p.
results: Sorbus scopulina. University of North
Carolina, Chapel Hill, NC. http://www.ibiblio. Treeguide. 2003. Greene mountain-ash, Sorbus
org/pfaf.cgi-bin/arr_html?Sorbus+scopulina& scopulina, Greene, Rosaceae.
CAN =LATIND. 6 p. http://www.treeguide.com/Species.asp?Region=
NorthAmerican&SpeciesID=1021. 2 p.
Keller, J. 2003. The winemaking home page:
mountain ash. http://winemaking.jackkeller.net/ Umatilla National Forest. 2003. Umatilla National
mtnash.asp. 2 p. Forest native plant species: Sorbus scopulina.
U.S. Department of Agriculture, Forest Service,
Love, R. 2003. Amelasorbus: An intergeneric Umatilla National Forest, Pendleton, OR. http://
hybred and a new taxon for the Oregon www.fs.fed.us/r6/uma/native/ts100.htm. 8 p.
checklist. http://www.oregonflora.org/ofn/v4n2/
amelasorbus.html. 3 p. Utah State University Horticulture. 2003. Sorbus
scopulina. http://www.usu.edu/natives/shrubs/
Luna, T. and D. Wick. 2001. Propagation protocol scopulina.htm. 1 p.
for production of container Sorbus scopulina
Greene var. scopulina Greene plants (172 ml Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
containers), Glacier National Park, West trees and shrubs. Agriculture Handbook 410.
Glacier, Montana. http://www. U.S. Department of Agriculture, Forest Service,
nativeplantnetwork.org. 8 p. Washington, DC. 265 p.
Moser, R. 2003. Edible Sierra Nevada plants: Welsh, S.L. 1974. Anderson’s flora of Alaska.
Rosaceae-rose family, mountain ash. Brigham Young University Press, Provo, UT.
Backcountry rangers, Fresno, CA. http://www. 724 p.
backcountryrangers.com/edibles/SORBUS.html.
2 p. Wier, S.K. 2003. Greene’s mountain-ash of the
southern Rocky Mountains. http://home.
Natural Resources Conservation Service. 2003. earthlink.net/~swier/MtnAsh.html. 1 p.
Sorbus scopulina Greene, Greene’s mountain
731
Spermacoce verticillata L. botón blanco
RUBIACEAE
John K. Francis
732
but is overcome by dense, tall grass, brush, and treatment of schistosomiasis, and as an abortive.
trees. Because of grazing, mowing, and An essential oil extracted from the leaves has been
cultivation, most plants do not progress beyond shown to inhibit Escherichia coli and
the herbaceous stage. If allowed to grow, they will Staphylococcus aureus (Burkill 2000).
form dense clumps and mats.
References
Reproduction.—In Brazil botón blanco blooms
from February through August (Instituto de Animal Science at Cornell University. 2002. Mind
Biociêcias 2002). In Texas, it flowers from March bending beta-carbolines. http://www.ansci.
through May (Correll and Johnston 1970). cornell.edu/plants/toxicagents/betacarbolines/
Flowering is almost continuous in moist portions bcarb.html. 5 p.
of Puerto Rico. Plants begin blooming in the
nursery at about 9 months. The flowers are Burkill, H.M. 2000. The useful plants of West
pollinated by several species of bees (Instituto de Tropical Africa. Royal Botanic Gardens, Kew,
Biociências 2002). Seeds collected in Puerto Rico UK. 686 p.
averaged 0.00016 g each or 6,250,000 seeds/kg.
Sown on peat without pretreatment, these seeds Correll, D.S. and M.C. Johnston. 1970. Manual of
germinated at 49 percent beginning in 13 days and the vascular plants of Texas. Texas Research
ending at 74 days. The seeds are disbursed by Foundation, Renner, TX. 1,881 p.
grazing animals and farm equipment. Established
plants root readily at the nodes when covered by Environnement et Développement du Tiers-
soil or rotting plant material. monde. 2002. Reconnaisssez, Protégez et
utilisez: Borreria vericillata L. Rubiaceae.
Growth and Management.—Botón blanco http://www.enda.sn/plantesmed/borreria.html.
seedlings grow slowly at first but begin rapid 2 p.
growth after about 6 months. Twenty-three 9-
month-old nursery plants averaged 64 cm in Holm, L., J. Doll, E. Holm, J. Pancho, and J.
height with a maximum of 109 cm. Botón blanco Herberger. 1997. World weeds. John Wiley &
shrubs appears to live at least 4 years and probably Sons, Inc. New York. 1,129 p.
much longer in Puerto Rico. Botón blanco is
controlled in crops and pasture by cultivation, Howard, R. A. 1989. Flora of the Lesser Antilles,
mowing, and spraying with broadleaf herbicides. Leeward and Windward Islands. Vol. 6. Arnold
The importance value of botón blanco in a Arboretum, Harvard University, Jamaica Plain,
Colombian pasture was reduced by 39 percent by MA. 658 p.
simply fertilizing with potassium and sulfur to
increase the vigor of the pasture grasses (Tejos Instituto de Biociências. 2002. Borreria
1981). verticillata. Instituto de Biociêcias,
Universidade de São Paulo, São Paulo, Brazil.
Benefits.—Botón blanco is a forage plant, but not http://ib.usp.br/beeplant/bove.htm. 1 p.
one highly favored by livestock. African material
was found to contain 0.2 percent alkaloid Instituto Botánico Darwin. 2002. Catálogo de las
including borreine and borreverine (Burkill 2000). plantas vasculares de la Argentina. http://
At least part of the alkaloids are beta-carbolines www.darwin.edu.ar/Catalogo/rubiaceae.pdf.
and would represent a poisoning hazard if they 22 p.
were present in higher concentrations (Animal
Science at Cornell University 2002). Botón blanco Jones, F.B. 1975. Flora of the Texas Costal Bend.
has a number of uses in herbal medicine, most Mission Press, Corpus Christi, TX. 262 p.
frequently for skin conditions. In Africa, leaf
extracts are used to treat leprous conditions, Kudremukh Wildlife Foundation. 2002. Checklist
furuncles, ulcers, and gonorrheal sores (Burkill of non-woody plants reported from Kudremukh
2000, Environnement et Développement du Tiers- National Park. http://kudremukh.org/
monde 2002). A lotion is prepared to relieve skin kudremukh/nonwoody.html. 4 p.
itches (Liogier 1990). Other preparations are used
internally to treat diarrhea, as a diuretic in the
733
Liogier, H.A. 1997. Descriptive flora of Puerto http://www.mct.gov.br/prog/ppg7/projetos/
Rico and adjacent islands. Vol. 5. Editorial de la proj991.pdf. 16 p.
Universidad de Puerto Rico, Río Piedras, PR.
436 p. Natural Resources Conservation Service. 2002.
Plants profile: Spermacoce verticillata L.,
Liogier, H.A. 1990. Plantas medicinales de Puerto shrubby false buttonweed. http://plants.usda.
Rico y del Caribe. Iberoamericana de Ediciones, gov/cgi_bin/plant_profile.cgi?symbol=SPVE2.
Inc., San Juan, PR. 566 p. 2 p.
Ministério de Ciencia e Tecnología. 2002. Tejos, R. 1981. Efecto del potasio y azufre sobre el
Recuperação de areas de pastagems pastizal nativo de una sabana. 1. Producción
abandonadas e degradadas através de sistemas contenido de material seca y composicion
agroflorestais na Amazônia Ocidental. botánica. Agronomía Tropical 29(6): 503-516.
734
Strumpfia maritima Jacq. pride-of-Big-Pine
RUBIACEAE
Synonyms: none
John K. Francis
735
References page. http://www.iucn-isg.org/actionplan/ch2/
exumaisland.php. 5 p.
Florida Fish and Wildlife Conservation
Commission. 1997. Florida’s endangered Liogier, H.A. 1990. Plantas medicinales de Puerto
species, threatened species, and species of Rico y del Caribe. Iberoamericana de Ediciones,
special concern. http://floridaconservation.org/ Inc., San Juan, PR. 566 p.
pubs/endanger.html. 18 p.
Liogier, H.A. 1997. Descriptive flora of Puerto
Howard, R.A. 1989. Flora of the Lesser Antilles, Rico and adjacent islands. Vol. 5. Editorial de la
Leeward and Windward Islands. Universidad de Puerto Rico, Río Piedras, PR.
Dicotyledoneae. Part 3. Vol. 6. Arnold 436 p.
Arboretum, Harvard University, Jamaica Plain,
MA. 658 p. Nelson, G. 1996. The shrubs and woody vines of
Florida. Pineapple Press, Inc., Sarasota, FL.
Knapp, C. 2002. Exuma Island iguana, Cyclura 391 p.
cychlura figginsi. Iguana Specialist Group home
736
Styrax americanus Lam. American snowbell
STYRACACEAE
Kristina Connor
737
Benefits.—American snowbell is an attractive Johnson, F.L. and B.W. Hoagland. 1999. Styrax
landscape plant. Its leaves are also a preferred food americanus. Catalog of the woody plants of
of the caterpillar of Callosamia promethea, the Oklahoma, Oklahoma Biological Survey Home
promethea moth (Oehlke [no date]). Page, http://www.biosurvey.ou.edu. 1 p.
738
Suaeda suffrutescens S. Wats. desert seepweed
CHENOPODIACEAE
James E. Nellessen
739
(glycophytes) cannot survive there. Halophytes members of this genus required prior cold
accumulate salts obtained from the soil solution stratification of the seeds with optimal daytime
within their tissues during the transpiration germination temperatures of 25 to 27 °C
process. Many members of the genus Suaeda (summarized in Baskin and Baskin 2001). NaCl
readily translocate salts to upper portions of the concentrations ranging from 0.17 to 0.90 molar
plant. This second group of halophytes can be have depressed maximum germination of 75 to
subdivided into three more categories of tolerance. 100 percent down to 10 percent in four different
Salt enduing species simply tolerate the presence species of Suaeda (summarized in Baskin and
of salt in their cells. Salt excluding species may Baskin 2001).
excrete salt from shoots, including those with
specialized hairs on the plant, or transport them Growth and Management.—Desert seepweed
back to the roots. Salt evading species have can grow relatively quickly. It has been observed
mechanisms to prevent transport to the leaves or to resprout readily in managed and mowed areas
prevent absorption from the soil solution (Waisel adjacent to levees along the lower Rio Grande
1972). Some members of the genus Suaeda fall floodplain in New Mexico. This species is
into the first group requiring salt for survival, restricted to saline and riparian habitats and is not
while others are salt enduring halophytes. Salt highly browsed. Consequently, it does not pose as
(NaCl) and other ions (including sulfate and a major management concern on most rangelands.
potassium) may be sequestered into cell vacuoles It does not typically form dense stands.
where they do not interfere with normal cell
biochemistry. These plants will generally have Benefits.—Desert seepweed leaves and young
higher water potential gradients between the soil shoots have been added to vegetable greens or
and the leaves, which may add to succulence. cacti fruits by Native Americans during cooking to
Waisel (1972) proposed a new order to the add flavor. It has also been added between layers
classification of halophytic plant communities: of cacti fruit and sotol butts (stems of Dasylirion
Suaedetalia--persistent communities in dry desert sp.) during the 24 to 48 hour cooking process in
saline habitats, dominated by annual and perennial mescal pits (Warnock 1974). Pima Indians would
members of the family Chenopodiaceae (as well as eat the plant for greens and the roasted seeds,
other families). Zahran (1982) likewise includes a called pinole. Coahuila Indians extracted a black
Suaeda community classification. Some shrubby dye from the plant used for artwork (Dodge 1985).
members of the genus (S. fruticosa Forsk. and S. This species is not browsed much by livestock but
monoica Forsk.) are reported to accumulate large may be utilized when other vegetation is in short
amounts of salt annually when in dense stands, as supply (Warnock 1974, Dodge 1985).
much as 2 to 4 tons/ha (Waisel 1972). In this
function they may help to desalinate some saline References
habitats.
Allred, K.W. 2002. A Working Index of New
Reproduction.—Flowers are small and greenish, Mexico Vascular Plant Names. http://web.
and occur in spiked clusters of three to nine nmsu.edu/~kallred/herbweb. [not paged].
flowers in the leaf axils. The corolla is absent. The
five-lobed calyx has all parts equal, may be hood Baskin, C.C. and J.M. Baskin. 2001. Seeds:
shaped, and lacks appendages, that is, it is not Ecology, Biogeography, and Evolution of
winged. Seeds are small, black, and 0.7 to 1 mm Dormancy, and Germination. Academic Press,
wide. Desert seepweed blooms from March to Sand Diego, CA. 666 p.
September, depending on local environmental
conditions (Correll and Johnston 1970, Great Chapman, V.J. 1942. The new perspective in the
Plains Flora Association 1986, Kearney and others halophytes. Quarterly Review of Biology 17:
1951, Martin and Hutchins 1980). Many of the salt 291-311.
tolerant members of the Chenopodiaceae have
physiological seed dormancy (Baskin and Baskin Correll, D.S. and M.C. Johnston. 1970. Manual of
2001). Although no seed germination reports the Vascular Plants of Texas. Texas Research
regarding this particular species were located, Foundation, Renner, TX. 1,881 p.
numerous studies have been performed on other
members of the Suaeda genus. Three of four
740
Dick-Peddie, W.A. 1993. New Mexico and Greenland, Vol. 1, 2nd Ed. Biota of North
Vegetation: Past, Present and Future. America Program of the North Carolina
University of New Mexico Press, Albuquerque, Botanical Garden. Timber Press, Portland, OR.
NM. 244 p. 622 p.
Dodge, N.N. 1985. Flowers of the Southwest Kearney, T.H., R. Peebles, and Collaborators.
Deserts. Southwest Parks and Monuments 1951 with 1960 supplement. Arizona Flora.
Association, Tucson, AZ. 136 p. University of California Press, Berkeley, CA.
1,085 p.
Great Plains Flora Association. 1986. Flora of the
Great Plains. University Press of Kansas, KS. Martin, W.C. and C.E. Hutchins. 1980 (reprinted
1,392 p. 2001). A Flora of New Mexico, Vol. 1. Bishen
Singh Mahendra Pal Singh (India) and Koeltz
Hendrickson, J. 1974. Saline habitats and Scientific Books (Germany). 1,276 p.
halophytic vegetation in the Chihuahuan Desert
region. In: R.H. Wauer and D.H. Riskind, eds. Waisel, Y. 1972. Biology of Halophytes.
Transactions of the Symposium of the Academic Press, NY. 395 p.
Biological Resources of the Chihuahuan Desert
Region, United States and Mexico. U.S. Warnock, B.H. 1974. Wildflowers of the
Department of Interior, National Park Service Guadalupe Mountains and the Sand Dune
Transactions and Proceedings Series, No. 3, Country, Texas. Sul Ross State University,
U.S. Goverment Printing Office, Washington Alpine, TX. 176 p.
DC. p. 289-314.
Zahran, M.A. 1982. Ecology of halophytic
Ivey, R.D. 1995. Flowering Plants of New vegetation of Egypt. In: D.N. Sen and K.S.
Mexico, 3rd Edition. Published by the author, Rajpurohit, eds. Contributions to the Ecology of
NM. 504 p. Halophytes.. Dr. W. Junk Publishers, The
Hague, Netherlands. p. 3-20.
Kartesz, J.T. 1994. A Synonymized Checklist of
the Vascular Flora of the United States, Canada,
741
Suriana maritima L. bay-cedar
SURIANACEAE (formerly in SIMAROUBACEAE)
Synonyms: none
John K. Francis
742
extracts of the leaves and bark to treat rheumatism Nelson, G. 1996. The shrubs and woody vines of
and skin ulcers and to stop bleeding (Liogier Florida. Pineapple Press, Inc., Sarasota, FL.
1990). 391 p.
Howard R.A. 1988. Flora of the Lesser Antilles, Schaus, C, S. Wade, and J. Dunan. 2002. Key deer
Leeward and Windward Islands. and plants they won’t eat. Monroe County
Dicotyledoneae, Part 1. Vol. 4. Arnold Extension Service, Key West, FL.
Arboretum, Harvard University, Jamaica Plain, http://monroe.ifas.ufl.edu/key_deer_plants.htm.
MA. 673 p. 4 p.
Liogier H.A. 1988. Descriptive flora of Puerto Smithsonian Institution. 2002. Special status
Rico and adjacent islands, Spermatophyta. Vol. species in the Indian River Lagoon. Smithsonian
2. Editorial de la Universidad de Puerto Rico, Marine Station at Fort Pierce, Smithsonian
Río Piedras, PR. 481 p. Institution, Washington, DC. http://www.
sms.si.edu/IRLSpec/ListedSpec.htm. 3 p.
Liogier, H.A. 1990. Plantas mediciales de Puerto
Rico y del Caribe. Iberoamericana de Ediciones, Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Inc., San Juan, PR. 566 p. Montiel, eds. 2001. Flora of Nicaragua.
Monographs in Systematic Botany Vol. 85, No.
Little, E.L., Jr., R.O. Woodbury, and F.H. 3. Missouri Botanical Garden Press, St. Louis,
Wadsworth. 1974. Trees of Puerto Rico and the MO. p. 1,911-2,666.
Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture, Wildflower Nervana. 2002. Suriana maritima—
Washingtion, DC. 1,024 p. guitarán or temporana (bay-cedar). http://
www.wfnirvana.com/pr/surimari/html. 3 p.
743
Symphoricarpos albus (L.) Blake common snowberry
CAPRIFOLIACEAE
John K. Francis
744
Reproduction.—Common snowberry blooms are on the shrubs. Containing 4 to 13 percent
between early May and late July depending on protein, depending on tissue and season, it
location and matures fruits in August to October provides important early-season browse for cattle
(Grimm 1966). The flowers are pollinated by a and domestic sheep. It is also browsed by most
range of bees, wasps, and syrphids (Gilbert 1995). wild ungulates. It is generally unpalatable to
Fruit and seed production can be abundant, horses and moose. The fruits are consumed by
especially when plants grow in full sun on fertile both black and grizzly bears and are eaten by a
sites. Normally a good seed crop is produced every number of birds and small mammals (McWilliams
year. Fruits remain on stems until midwinter or 2000). Some sources refer to the fruits as being
early spring, if not eaten. Numbers of seeds per toxic to humans (Gilbert 1995, Moerman 1986).
gram vary from 119 to 250 for var. albus and from The toxic agent appears to be the isoquinoline
86 to 144 for var. laevigatus. Germination may be alkaloid chelidonine and causes mild symptoms of
as high as 87 percent if exacting conditions of vomiting, dizziness, and sedation in children
pretreatment are met. Germination is epigeal (Canadian Biodiversity Information Facility 2003).
(Walker 2003). Seeds are dispersed by birds and The fruits were reported to have been eaten but not
mammals. Seedlings are relatively uncommon. favored by Native Americans (McWilliams 2000).
However, vegetative reproduction is common by Various tissues were used by Native Americans as
sprouting of underground stems that run a diuretic, to treat gonorrhea, and for sore eyes
horizontally at a depth of 2 to 5 cm for up to 60 cm (Moerman 1986). Common snowberry is planted
before turning up to form new tops. Stems layer as an ornamental and in conservation plantings for
when they come in contact with the ground, and wildlife amenity, soil stabilization, and strip mine
severed stems lying on the ground will also take reclamation in temperate areas around the world
root (Gilbert 1995). (Gilbert 1995).
Benefits.—Common snowberry helps protect the McWilliams, J. 2000. Symphoricarpos albus. In:
soil, provides food and cover for wildlife, browse U.S. Department of Agriculture, Forest Service,
for livestock, and adds beauty to the landscape, Rocky Mountain Research Station, Fire Sciences
especially in fall and winter when the white fruits Laboratory. Fire Effects Information System.
745
http://www.fs.fed.us/database/feis/plants/shrub/ Stephens, H.A. 1973. Woody plants of the North
symalb/all.html. 32 p. Central Plains. The University Press of Kansas,
Lawrence, KS. 530 p.
Moerman, D.E. 1986. Medicinal plants of Native
America. Technical Reports 19. University of Walker, S.C. 2003. Symphoricarpos Duham.,
Michigan Museum of Anthropology, Ann snowberry. U.S. Department of Agriculture,
Arbor, MI. 534 p. Forest Service, Washington, DC. http://wpsm.
net/Symphoricarpos.pdf. 10 p.
Natural Resources Conservation Service. 2003.
Plants profile: Symphoricarpos albus (L.) Blake Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
http://plants.usda.gov/cgi_bin/plant_profile. Higgins, eds. 1987. A Utah flora. Great Basin
cgi?symbol =SYAL. [not paged]. Naturalist Memoirs 9. Brigham Young
University, Provo, UT. 894 p.
Soper, J.H. and M.L. Heimburger. 1982. Shrubs of
Ontario. Royal Ontario Museum, Toronto,
Ontario, Canada. 495 p.
746
Tabebuia haemantha (Bertol. ex Spreng.) DC. roble cimarrón
BIGNONIACEAE
John K. Francis
747
collected in Salinas, Puerto Rico contained an environmental restoration plantings within its
average of 59.9 ± 2.01 seeds/capsule. The air-dry native range. The light brown, hard wood is
seeds averaged 0.0148 + 0.0003 g/seed or 68,000 suitable for firewood and charcoal but is little used
seeds/kg. Sown on peat moss, these seeds because of the plant’s small size and relative
germinated at 90 percent between 7 and 31 days scarcity.
after sowing. The species grows and survives well
in the nursery, similar to other Tabebuia seedlings. References
In the wild, seedlings are well scattered and
uncommon. Liogier, H.A. 1995. Descriptive flora of Puerto
Rico and adjacent islands. Vol. 4. Editorial de la
Growth and Management.—Roble cimarrón Universidad de Puerto Rico, Río Piedras, PR.
grows about 0.5 m/yr in early years after it is well 617 p.
established. It does not appear to be a long-lived
species. Unfortunately, no plantations are known Little, E.L., Jr. and F.H. Wadsworth. 1964.
and no management experience has been reported. Common trees of Puerto Rico and the Virgin
Islands. Agriculture Handbook 249. U.S.
Benefits.—Roble cimarrón would probably be Department of Agriculture, Forest Service.
suitable as an ornamental for background Washington, DC. 548 p.
plantings in moist and dry frost-free areas. It
would have the advantage of being drought-hardy Vélez, I. and J. van Overbeek. 1950. Plantas
and tolerant of infertile and partially compacted indeseables en los cultivos tropicales. Editorial
soils and should be a good species to plant in Universitaria, Río Piedras, PR. 497 p.
748
Tamarix chinensis Lour. tamarisk
TAMARICACEAE
749
that have 50 percent fewer small mammal species insects have been investigated--from grasshoppers
as well as fewer reptile and amphibian species than to leaf hoppers. Two species specific to tamarisk, a
native riparian stands (Olson 1999). Although bees mealybug (Trabutina mannipara) from Israel and
visit the flowers (Epple 1995) the diversity of a leafbeetle (Diorhaba elongata) from China, have
insects is also low, and nestling birds rely on received advance testing (DeLoach and others
insects brought to the nests by their parents. 1996, DeLoach, 1994). Physical cutting and
Although some birds can nest in the shrubs the removal of tamarisk will only be successful if the
trunks are not large enough to provide habitat for roots are removed or killed. By itself, cutting
animals such as wood-peckers, owls, chickadees, tamarisk by hand (that is by chain saw, weed eater,
and squirrels. The tiny, hairy seeds provide poor or axe) is not often an effective treatment. Cutting
nutrition and are largely indigestible. tamarisk tends to just cause more shoots to grow.
However, there was a report of successful control
Reproduction.—Tamarisk generally flowers in where large tree stumps (20 to 25 cm diameter)
April through August. The pink, fragrant flowers had been cut back to the soil surface and then
are insect pollinated. The seeds, which are small, covered by a black, liner material (Cinnamon
mature in summer and fall and are dispersed by 1990). Purely mechanical removal is also
birds and by water. However, the seed may have sometimes successful if there is enough standing
limited longevity under humid conditions. Seeds vegetation to substantially shade the resprouting
retain high viability if stored in sealed containers tamarisk. For example, in Nevada (Busch and
with desiccant at temperatures between 3 to 27 oC Smith, 1995) when tamarisk had been completely
but lose viability if stored at “moderate humidity” cleared above ground in a stand of willows, there
(Wilgus and Hamilton 1962, Baskin and Baskin was essentially no grow back during the 4 years
2001). Germination temperature appears to be follow up. Tamarisk is sensitive to numerous
flexible. Freshly matured seeds germinated from herbicides (Kunzman and Bennett 1990, Sisneros
87 to 98 percent when kept at constant 1991). At the present time, the most successful
temperatures between 19 to 43 oC. Percent approach to control tamarisk is by cut-stump
germination did not appear related to temperature; herbicide or cut-stump/frill herbicide treatments
96 percent germinated at 19 oC and 91 percent at (Neill 1990, Hughes 1996). This approach is quite
43 oC (Wilgus and Hamilton 1962, Baskin and costly but it is very controlled, does not affect non-
Baskin 2001). Vegetative reproduction by suckers target organisms, and causes the least
and rooting branches is copious. environmental impact. The tamarisk shrubs should
be cut to within 2.5 cm of the ground surface
Growth and Management.—Tamarisk is an (Cinnamon 1990) and a systemic herbicide applied
aggressive, invasive species. Its invasive behavior to the stump within a few minutes after cutting.
is illustrated by its colonization of the Pecos River The time between cutting and herbicide treatment
Valley in New Mexico (Allred 2002). In 1912, a should be as short as possible. Leaving the cut
few seedlings were observed at Lake McMillan. stump for 2 to 5 days before herbicide treatment
By 1915 the shrub covered 600 acres and extended reduces success rate from greater than
up and down the riverbanks. By 1926 it covered approximately 88 percent tamarisk death to less
12,300 acres and extended to 57,000 acres by 1960 than 10 percent (Hays and Mitchell 1990). Fire is
(Allred 2002). It out-competes many native often ineffective. The high water and salt content
species and is difficult to eradicate once of tamarisk makes it difficult to burn. Too light
established (USDA Forest Service 1988). The (low temperature) a fire will permit, or even
extremely deleterious impact of this species on the encourage, tamarisk to re-sprout and become even
riparian and wetland areas of the Southwest has denser. If the fire burn is so hot it kills the
led to intense study on how to control and reverse tamarisk, it is likely that the fire will leave the soil
its spread. There are four general methods-- bare and in such a poor condition that it cannot
biological control, physical removal by hand or by support grasses and desirable vegetation. In the
machine, herbicides, and fire--that can be used to horticultural trade, tamarisk is propagated from
control tamarisk, and each has shortcomings. The hardwood cuttings in the spring or softwood
most appropriate method, or combination of cuttings in the summer (Tykač 1990). Plants
methods, depends upon the particular site. grown for a long time in containers often do not
Biological controls for tamarisk are not yet establish after transplanting (Tykač 1990).
commercially available, although a variety of
750
Benefits.—Tamarisk were once planted for Group for the Introduction of Biological
erosion control and as wind breaks (Baum 1978, Control Agents of Weeds, USDA-APHIS. 21
Allred 2002). It is not significantly grazed or March.
browsed. The tannin substances in the vegetation
are likely to make it unpalatable. Tamarisk has DeLoach, C.J., D. Gerling, L. Fornasari, R.
commercial value in the landscape and Sobhain, S. Myartseva, I.D. Mityaev, Q.G. Lu,
horticultural trade. It is a popular ornamental shrub J.L. Tracy, R. Wang, J.F. Wang, A. Kirk, J.W.
although its use in the United States is now Pemberton, V. Chikatunov, R.V. Jashenko, J.E.
generally discouraged. Many cultivars exist that Johnson, H. Zheng, S. L. Jiang, M.T. Liu, P.A.
have been chosen for flower color and growth Liu, and J. Cisneros. 1996. Biological control
form (Bailey and others 1976, Tykač 1990). program against salt cedar (Tamarix spp.) in the
Tamarix species, although not specifically T. U.S.A. In: V. C. Moran and J.H. Hoffman, eds.
chinensis, have been used for fuel and building Proc. of the IX International symposium on the
materials by Native American tribes in the control of weeds, Cape Town, South Africa. p.
Western United States (Moerman 1998). 253-260.
DeLoach, C.J. 1994. Petition to release into the Moerman, D.E. 1998. Native American ethno-
field the leaf beetle Diorhaba elongata from botany. Timber Press, Portland, OR. 927 p.
China for biological control of saltcedar,
Tamarix ramosissima, a weed of riparian areas Neill, W.M. 1990. Control of Tamarisk by cut-
of the Western United States and Northern stump herbicide treatment. In: Tamarisk
Mexico. Submitted to the Technical Advisory control in Southwestern United States. Special
751
Report 9. Cooperative National Park Resources Tykač, J. 1990. The illustrated guide to ornamental
Studies Unit. p. 91-98. shrubs. Treasure Press, London. 224 p.
Olson, BE. 1999. Impacts of noxious weeds on United States Department of Agriculture, Forest
ecologic and economic systems. In: R.L Sheley Service. 1988. Range Plant Handbook. Dover
and J.K. Petroff, eds. Biology and Management Publications, Inc. New York. 838 p.
of noxious rangeland weed. Oregon State
University Press, Corvallis, OR. 528 p. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Higgins. 1993. A Utah flora. Brigham Young
Sisneros, D. 1991. Herbicide analysis: Lower University, Provo, UT. 986 p.
Colorado River saltcedar vegetation
management study. Publication R-91-06. Wilgus, F. and K.C. Hamilton. 1962. Germination
Bureau of Reclamation, Denver, CO. 165 p. of salt cedar seed. Weeds 10: 332-333.
752
Tamonea boxiana (Moldenke) Howard crow broom
VERBENACEAE
John K. Francis
753
Benefits and Detriments.—Crow broom Liogier, H.A. 1995. Descriptive flora of Puerto
contributes to biodiversity in the habitat where it Rico and adjacent islands. Vol. 4. Editorial de
grows, helps protect the soil, and furnishes cover la Universidad de Puerto Rico, Río Piedras, PR.
for wildlife. It may become a minor weed in 617 p.
rangelands and the clinging seeds can be a
nuisance. Vasquez, O.J. and D.A. Kolterman. 1998. Floristic
composition and vegetation types of the Punta
References Guaniquilla Natural Reserve—Cabo Rojo,
Puerto Rico. Caribbean Journal of Science 34(3-
Howard, R.A. 1989. Flora of the Lesser Antilles. 4): 265-279.
Vol. 6. Arnold Arboretum, Harvard University,
Jamaica Plain, MA. 658 p.
754
Taxus brevifolia Nutt. Pacific yew
TAXACEAE
John K. Francis
755
releases seeds in October. The pollen is dispersed mauls, splitting wedges, gunstocks, boat decking,
by the wind. There are 32,000 to 36,000 seeds/kg fence posts, musical instruments, carved figurines,
(Washington source). About 50 to 99 percent will novelty items, furniture, oriental ceremonial
eventually germinate under proper conditions. The “Toko” poles, turnery, cabinetry, and firewood
seeds are dispersed primarily by birds. Natural (Bolsinger and Jaramillo 1990, Termenstein 1990).
germination usually takes place in the second Infusions, decoctions, and poultices of leaves,
spring after dispersal (Rudolf 1974). Development twigs, and bark were used by Native Americans to
of seedlings is slow. Pacific yew sprouts from treat lung problems, stomachache, wounds, and
stumps and rootstalks after being cut or top-killed. pain (Moerman 1986). Pactitaxel (often called by
It commonly layers when branches or stems are the trade name Taxol) was discovered during a
pressed to the ground by snow or falling debris massive anticancer activity screening program by
(Bolsinger and Jaramillo 1990). the National Cancer Institute during the 1960’s
(21cecPharm 2003). Paclitaxel is a white
Growth and Management.—Growth of Pacific crystalline powder extractable from any of the
yew is slow. Trees in Idaho took 25 years to reach Taxus species. It is a cytotoxic anticancer drug
a stem diameter (15 cm above the ground) of 2.5 used to treat ovarian, breast, and lung cancers, and
cm and 100 years to reach 15 cm. Height growth is Kaposi’s sarcoma (SFT Enterprise 2003).
correspondingly slow (Bolsinger and Jaramillo Harvesting for peclitaxel extraction threatened to
1990). Even in moist, rich soils, few plants live decimate the species’ populations until it was
long enough to reach tree sizes. Fruits should be discovered that other Taxus species were better
picked by hand as soon as they are ripe. The fruit sources and that the drug could be produced by
pulp is removed by maceration and floating off the semi-synthesis and by cell cultures (21cecPharm
pulp, or by soaking in water at room temperature 2003). Pacific yew is used to a limited extent as an
for 4 or 5 days, rubbing the fruits on screens, and ornamental shade tree, foundation plant, hedge,
washing away the pulp. The seeds should be air- and topiary plant (Bolsinger and Jaramillo 1990).
dried for 1 to 2 weeks and should be sown or
stored as soon as drying is complete. Air-dried References
seeds can be stored for 5 to 6 years in sealed
containers at 1 to 2 °C. Seeds sown in nursery beds 21cecPharm. 2003. Paclitaxel story. http://www.
should be covered by about 1 cm of mineral soil 21cecpharm.com/px/story.htm. 4 p.
and mulched. Germination primarily occurs in the
second spring afterwards. To obtain germination a Abrams, L. 1940. Illustrated flora of the Pacific
shorter time after sowing, seeds should be States. Stanford University Press, Stanford, CA.
stratified for 90 to 210 days at 16 °C followed by 538 p.
60 to 120 days at 3 to 6 °C (Rudolf 1974).
Bolsinger, C.L. and A.E. Jaramillo. 1990. Taxus
Benefits.—Pacific yew contributes to the brevifolia Nutt., Pacific yew. In: R.M. Burns,
aesthetics of the forest, helps protect the soil, and and B.H. Honkala, eds. Silvics of North
provides food and cover for wildlife. It is a America. Agriculture Handbook 654. Forest
preferred moose browse. Although eaten in all Service, U.S. Department of Agriculture,
seasons, during the winter, moose will eat all the Washington, DC. p. 573-579.
available leaves and twigs, and even strip the bark.
Deer, elk, and rabbits also browse the species. Earle, C.J. 2002. Taxus brevifolia Nuttall 1849.
Livestock make limited use of it during the winter http://www.botanik.uni-bonn.de/conifers/ta/ta/
and when other food is lacking. The fruit is sweet brevifolia.htm. 3 p.
and eaten by many species of birds (Tirmenstein
1990). Plants growing along streams help protect Moerman, D.E. 1986. Medicinal plants of Native
against streambank erosion and shade the water America. Technical Reports 19. University of
and help maintain cool stream temperatures Michigan Museum of Anthropology, Ann
(Termenstein 1990). Pacific yew wood is fine- Arbor, MI. 534 p.
grained, heavy, hard, elastic, and strong. It has
been used and still is used to a limited extent (due Rudolf, P.O. 1974. Taxus L., yew. In: C.S.
to scarcity) for archery bows, harpoon shafts, Schopmeyer, tech. coord. Seeds of woody plants
canoe paddles, household utensils, tool handles, in the United States. Agriculture Handbook 450.
756
U.S. Department of Agriculture, Forest Service, Tirmenstein, D.A. 1990. Taxus brevifolia. In: U.S.
Washington, DC. p 799-802. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences
Sargent, C.S. 1923. Manual of the trees of North Laboratory, Missoula, MT. Fire Effects
America (exclusive of Mexico). Houghton Information System. http://www.fs.fed.us/
Mifflin, Boston, MA. 910 p. database/feis/plants/tree/ taxbre/all.html. 19 p.
SFT Enterprise. 2003. Natural pactitaxel, full Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
information. http://www.21cep.com/sft/ trees and shrubs. Agriculture Handbook 410.
pxsft.htm. 9 p. U.S. Department of Agriculture, Forest Service,
Washington, DC. 265 p.
757
Taxus canadensis Marsh. Canada yew
TAXACEAE
General Description.—The common name most Range.—Canada yew is native to Eastern Canada
often used for Taxus canadensis is Canada yew. It (Manitoba, New Brunswick, Newfoundland, Nova
is also known as American yew and ground Scotia, Ontario, Prince Edward Island, Quebec)
hemlock; a common French name used in Quebec and the Northeastern United States (Connecticut,
for this species is buis de sapin. This evergreen Illinois, Indiana, Iowa, Kentucky, Maine,
species is usually a low-growing, straggling non- Maryland, Massachusetts, Michigan, New
aromatic shrub, with ascending branches up to 2 m Hampshire, New Jersey, New York, North
tall. Branches often extend two-thirds of their Carolina, Ohio, Pennsylvania, Rhode Island,
length laterally before curving upwards. The bark Tennessee, Vermont, Virginia, West Virginia,
is thin, scaly and red to reddish-brown. Twigs are Wisconsin) (Hils 1993, Natural Resources
green and alternately arranged, and winter buds are Conservation Service 2003). It becomes rare above
composed of imbricate (overlapping), lanceolate, 50 °N (Soper and Heimburger 1982).
keeled scales. The leaves are needle-like, on short
stalks, and persist for several years. The leaf Systematic Botany.—The genus name Taxus is
blades are spirally arranged, flattened, often in two from the Greek word, taxos, for yew.
ranks, linear, abruptly narrowed into a fine point, Approximately eight species of Taxus are
1.2 to 2.2 cm long, with pale green bands on the recognized, and all of them are similar
under surface. The midrib is slightly elevated on morphologically, even though they are well-
the top surface. The leaves lack resin ducts. In separated geographically (Hils 1993). Canada yew
758
is the only native yew throughout its range. are suggested to promote more rapid germination
However, it can be confused with two other yew of seeds (Dirr and others 1987). Seeds may be
species--English yew (T. baccata L.) and Japanese sown in nursery beds in the fall and covered by 10
yew (T. cuspidata Sieb. & Zucc.)--and a hybrid to 12 mm of mineral soil topped by mulch.
between them, (T. media Rehd.) that occasionally However, fall-sown seeds usually germinate the
escapes from cultivation. Canada yew can be second spring (Rudolf 1974). Seedlings should be
distinguished from these by its low-growing, transplanted while still small. However, they are
straggling nature. Immature saplings are slow growing and they must be maintained for 2
unidentifiable to species. Canada yew has 2n = 24 years in nursery beds or pots at a minimum.
chromosomes (Hils 1993). Seedlings should be outplanted in late spring or
early summer after the last frost has occurred.
Ecology.—Canada yew, a slow-growing species, Established plants will tolerate pruning (Brand
is tolerant of shade, which gives it a competitive 2003).
advantage over other species with which it must
compete. It grows best in at least partial shade Benefits.—Canada yew is an important
(Sullivan 1993). Canada yew is found in rich soil component of the forest understory in Northeastern
in a variety of habitats including bogs, swamps, forests, helping to protect the soil and imparting
and rocky banks at elevations up to 1,500 m. It beauty to forest landscapes. The species is used as
prefers soils that are well-drained and that have an ornamental for ground cover and mass
pH’s between 5.0 and 7.5. The climate is humid plantings, and for parental stock in yew breeding
and cool. Exposure to winds from the ocean is not (Sullivan 1993). The toxic compound that is
tolerated. The species grows in many forest present in the plant is the alkaloid taxine
associations and is taken as an indicator of old- (Kingsbury 1964). The seeds and the dried foliage
growth forest. It does not occur in seral have been fatal to livestock, but the fresh foliage is
communities. Canada yew is apparently easily browsed by deer (Fernald 1950, Hils 1993). Moose
killed by fire. Wildfires and other influences that also browse the species and have seriously reduced
open the forest canopy reduce the competitive Canada yew abundance on Isle Royale, Michigan,
ability of the species (Sullivan 1993). since their introduction. The fleshy aril is eaten by
numerous species of birds (Sullivan 1993). The
Reproduction.—Canada yew flowers in April and pulp of the arils is sweet and edible to humans
May and is wind pollinated. Individual plants are (Fernald 1950) but has a slimy texture (Soper and
not self-fertile (Plants For a Future 2003). It Heimburger 1982). A number of Native American
produces seed almost every year (Sullivan 1993). groups made decoctions of leaves and twigs to
The seeds mature in late summer and autumn. treat rheumatism. Infusions and decoctions were
There are 33,000 to 62,400 seeds/kg (Rudolf also used to treat numbness of fingers and legs,
1974). The seeds are dispersed by birds (Sullivan colds, gonorrhea, and as a diuretic (Moerman
1993). Canada yew spreads vegetatively by 1986). New foliage and green stems are
layering (Rook 1998). The connections between commercially harvested for the extraction of
plants of a clonal group usually rot away (Sullivan paclitaxel (Public Forest Council 2003). Paclitaxel
1993). Small plants tend to be males and browsing is a white crystalline powder extractable from any
and other stresses on a population tends to increase of the Taxus species. It is a cytotoxic anticancer
the proportion of male plants in the population drug used to treat ovarian, breast, and lung
(Sullivan 1993). cancers, and Kaposi’s sarcoma (SFT Enterprise
2003).
Growth and Management.—Canada yew is slow
growing (Public Forest Council 2003). It is the References
hardiest of all yew species; however, it is the least
attractive yew species used as an ornamental due Brand, M.H. 2003. Plant UConn database of trees,
to its low-growing, straggling habit (Rehder, shrubs and vines: Taxus canadensis.
1940). The species may be propagated with seeds http://www.hort.uconn.edu/plants/t/taxcan/
and by cuttings of half-hardened or hardened taxcan3/html. 2 p.
terminal shoots that root with good success (Plants
For a Future 2003). Warm stratification for 4 Britton, N.L. and A. Brown. 1913. Illustrated flora
months followed by 4 months of cold stratification of the northern states, Canada, and the British
759
possessions. Vol. 2, 2nd Ed. Scribner, New York. Public Forest Council. 2003. Ground hemlock
735 p. (Taxus canadensis). Public Forest Council Fact
Sheet. Charlottetown, Prince Edward Island,
Dirr, M.A., M.W. Heuser, Jr., and B.L. Dirr. 1987. Canada http://www.gov.pe.ca/af/agweb/
The reference manual of woody plant library/factsheets/ground_hemlock.pdf. 2 p.
propagation. Varsity Press, Athens, GA. 239 p.
Rehder, A. 1940. Manual of cultivated trees and
Fernald, M.L. 1950. Gray’s manual of botany. shrubs. Macmillan Publishing Co., New York.
American Book Co., New York. 1,632 p. 996 p.
Hils, H.J. 1993. Taxaceae. In: Flora of North Rook, J.S. 1998. Taxus canadensis, Canada yew.
America Editorial Committee, eds. Flora of http://www.rook.org/earl/bwca/nature/shrubs/
North America. Vol. 2. Oxford University Press, taxuscan.html. 3 p.
New York. p. 423-427.
Rudolf, P.O. 1974. Taxus L., yew. In: C.S.
Kingsbury, J.M. 1964. Poisonous Plants of the Schopmeyer, tech. coord. Seeds of woody plants
United States and Canada. Prentice-Hall, Inc., in the United States. Agriculture Handbook 450.
Englewood Cliffs, NJ. 626 p. U.S. Department of Agriculture, Forest Service,
Washington, DC. p. 799-802.
Moerman, D.E. 1986. Medicinal plants of Native
America. Technical Reports 19. University of SFT Enterprise. 2003. Natural pactitaxel, full
Michigan Museum of Anthropology, Ann information. http://www.21cep.com/sft/
Arbor, MI. 534 p. pxsft.htm. 9 p.
Natural Resources Conservation Service. 2003. Soper, J.H. and M.L. Heimburger. 1982. Shrubs of
Plants profile: Taxus canadensis Marsh. http:// Ontario. Royal Ontario Museum, Toronto,
plants.usda.gov/cgi_bin/plant_search.cgi?mode= Ontario, Canada. 495 p.
Scientific+Name&keywordquery=Ta...
[not paged]. Sullivan J. 1993. Taxus canadensis. In: U.S.
Department of Agriculture, Forest Service,
Pilger, R.K.F. 1903. Taxaceae. In: H.G.A. Engler, Rocky Mountain Research Station, Fire Sciences
ed. Das Pflanzenreich. V. 18(IV,5). Laboratory, Fire Effects Information System.
p. 1-124. http://www.fs.fed.us/database/feis/plants/shrub/
taxcan/all.html. 10 p.
Plants For a Future. 2003. Taxus canadensis.
http://www.ibiblio.org/pfaf/chi-bin/arr_html?
Taxus+canadensis. 7 p.
760
Tecoma stans (L.) Juss. ex Kunth ginger-thomas
BIGNONIACEAE
John K. Francis
761
germinated (Francis and Rodríguez 1993). No Bailey, L.H. 1941. The standard cyclopedia of
pregermination treatments are necessary. horticulture. Vol. 3. The MacMillan Company,
New York. p. 2,423-3,639.
Growth and Management.—Early growth is
relatively rapid. About 1 m of height growth can be Francis, J.K. and A. Rodríguez. 1993. Seeds of
obtained during the first year. Depending on the Puerto Rican trees and shrubs: second
environment, maximum height of plants may vary installment. Research Note SO-374. U.S.
from 1.5 m to 7 m, and diameters of 8 cm are Department of Agriculture, Forest Service,
sometimes achieved. Establishment activities may Southern Forest Experiment Station, New
include removal of competition and disturbance of Orleans, LA. 5 p.
the soil to encourage natural regeneration, planting
(more reliable for smaller numbers of plants), and Kranz, W.M. and T. Passini. 1997. Amarelinho:
control of competing vegetation. When control of biologia e control. Informe da Pesquisa
ginger-thomas is needed, cutting, grubbing out No.121.Instituto Agronomico do Parana.
stumps, or poisoning is recommended. In Brazil, Londrina, PR, Brazil. 19 p.
tebuthiuron was found to effectively kill plants, but
picloram and 2, 4-D were not (Passini and Kranz Kumar, R. and G. Singh. 1988. Investigations into
1997). the cause of sterility. Tecoma stans L. Letters
Botaniques (France) 135(2): 131-135.
Benefits.—Ginger-thomas is planted as an
ornamental throughout the topics and subtropics. It Little, E.L., Jr. and F.H. Wadsworth. 1964.
is especially prized as a flowering hedge plant. The Common trees of Puerto Rico and the
shrub is planted and managed to enhance the VirginIslands. Agriculture Handbook 249. U.S.
beauty of green belts and natural forests used for Department of Agriculture, Forest Service.
recreation. Used in combination with trees, ginger- Washington, DC. 548 p.
thomas contributes to effective windbreaks and
sound breaks. Growing in thick patches that shade Liogier, H.A. 1990. Plantas medicinales de Puerto
out grass, the species has become a serious weed in Rico y del Caribe. Iberoamericana de Ediciones,
Brazilian pastures (Kranz and Passini 1997). Inc., San Juan, PR. 566 p.
Browsing cattle and goats in Mexico consume up
to 20 percent of the leaves and 100 percent of the Lozoya-Meckes, M. and V. Mellado-Campos.
available flowers (Susano Hernandez 1981). The 1985. Is the Tecoma stans infusion an anti-
foliage in one study in India contained 17 percent diabetic remedy? Journal of Ethnopharmacology
crude protein, 6 percent ash, 18 percent fat, 25 14(1): 1-9.
percent fiber, and 14 percent total polyphenols
(Nag and others 1994). Ginger-thomas leaves, Neg Datta, A., A.K. Lehiri, S. Matai, and L. Si.
bark, and roots contain many biologically active 1994. Proximate composition and polyphenol
chemicals, and extracts from those tissues have content of some tree leaves. Indian Forester
been used in traditional folk medicine to treat many 120(12): 1,122-1,125.
diseases and conditions (Liogier 1990). Perhaps the
most promising compounds are monoterpine Passini, T. and W.M. Kranz. 1997. Eficacia de
alkaloids, which have been shown to effectively herbicidas no controle de amarelinho (Tecoma
reduce the symptoms of diabetes mellitus in rats, stans) em pastagem. Planta Daninha 15(2): 190-
dogs, and mice (Aguilar and others 1993, Lozoya- 197.
Meckes and Mellado-Campos 1985, Perez and
others 1984) Perez G., R.M., A. Ocegueda Z., J.L. Muñoz L.,
J.G. Avila A., and W.W. Morrow. 1984. A study
References of the hypoglycemic effect of some Mexican
plants. Journal of Ethnopharmacology 12(3):
Aguilar, L.C., S. Macias, A. Chagoya, A. 253-262.
Cardenas, P. Díaz, and J M. Cantu. 1993.
Antidiabetic activity of Tecoma stans in rats. Susano Hernandez, R. 1981. Especies arboreas
Fitoterapia 64(4): 304-305. forestales susceptibles de aprovecharse como
forraje. Ciencia Forestal 6(29): 31-39.
762
Tipton, J.L. 1994. Relative drought resistance
among selected southwestern landscape plants.
Journal of Agriculture 20(3): 150-155.
763
Tetrapterys inaequalis Cav. bejuco de paralejo
MALPIGHIACEAE
John K. Francis
764
samaras from Puerto Rico collected by the author References
in April averaged 0.0953 + 0.0014 g/fruit. Planted
on commercial potting mix without any Acevedo-Rodríguez, P. 1985. Los bejucos de
pretreatment, 79 percent germinated between 34 Puerto Rico. Vol. 1. General Technical Report
and 82 days after sowing. Samaras spiral sideways SO-58. U.S. Department of Agriculture, Forest
a few meters from fruiting adult plants. While Service, Southern Forest Experiment Station,
young plants are relatively common in many New Orleans, LA. 331 p.
areas; fruiting plants are relatively rare. Layering
(rooting in contact with the ground) is common Howard R.A. 1988. Flora of the Lesser Antilles,
and appears to be an important means of Leeward and Windward Islands.
perpetuating established plants. Dicotyledoneae, Part 1. Vol. 4. Arnold
Arboretum, Harvard University, Jamaica Plain,
Growth and Management.—Bejuco de paralejo MA. 673 p.
vines extend rapidly from sprouts. A large vine
(7.7 cm in diameter) cut by the author had 27 Liogier, H.A. 1988. Descriptive flora of Puerto
growth rings. Although large plants are capable of Rico and adjacent islands, Spermatophyta. Vol.
smothering small trees, the species causes few 2. Editorial de la Universidad de Puerto Rico,
problems. No planting or management experience Río Piedras, PR. 481 p.
has been published.
Liogier, H.A, and L.F. Martorell. 2000. Flora of
Benefits.—Bejuco de paralejo contributes to the Puerto Rico and adjacent islands, a systematic
biodiversity of forests, helps protect the soil, and synopsis. 2nd ed. Editorial de la Universidad de
furnishes cover for wildlife. Puerto Rico, Río Piedras, PR. 382 p.
765
Thrinax morrisii H. Wendl. brittle thatch palm
ARECACEAE
John K. Francis
766
to favorable conditions in Puerto Rico (Little and squirrel (Sciurus aureogester (Cowley 2002) and
others 1974) and flowers in the spring in Florida the eastern box turtle (Terrapene carolina) (Liu
(Gilman and Watson 1994). The flowers are insect and others 2002), and in the Bahamas by the
pollinated. A collection of seeds in Puerto Rico Bahamas amazon parrot (Amazona leucocephala
averaged 0.0396 + 0.0007 g/seed or 25,000 bahamensis) (Attril 2002).
seeds/kg. Sown without pretreatment on moist
blotter paper, 85 percent germinated in 17 months References
(author’s observation). The seeds are reported to
begin germination 50 to 70 days after sowing Attril, R. 2002. Bahamas wildlife pages: The status
(Hoyos-F. and Braun 2001). The seeds can stand and conservation of the Bahamas amazon.
storage at -40 °C and 66 °C for at least 1 week http://www.bahamaswildlife.fsnet.co.uk/
without loss of viability (Broschat and Meerew parrotconservation.htm. 8 p.
2000). Because seeds have such a long dormant
period, they can be collected by hand from plants Broschat, T.K. and A.W. Meerew. 2000.
or from the ground at almost any time of the year. Ornamental palm horticulture. University Press
The seeds are dispersed by animals. Surviving of Florida, Gainesville, FL. 255 p.
seedlings are not usually common, but the plants
are long-lived and populations can build to high Cowley, M. 2002. Florida native plants: palm
levels. family. http://www.nsis.org/garden/family/palm.
html. 3 p.
Growth and Management.—Brittle thatch palm
is slow growing. A specimen in the Fairchild Desert-tropicals. 2002. Broom palm. http://desert-
Tropical Garden, Florida was 42 years old and tropicals.com/Palm/Thrinax_morrisii.html. 3 p.
1.16 m in height (Zona and Maidman 2000). Plants
mature and begin flowering and fruiting with only Gilman, E.F. and D.G. Watson. 1994. Thrinax
a rosette of basal leaves, long before they develop morrisii, Key thatch palm. Fact Sheet ST-628.
a discernable stem. Plants with long stems are U.S. Department of Agriculture and Southern
uncommon and are probably more than 50 years Group of State Foresters. Winder, GA. 3 p.
old. Wildland plantings are not documented but
presumably could be done with container stock. Howard, R.A. 1979. Flora of the Lesser Antilles,
Either they must be established in poor sites where Leeward and Windward Islands. Vol. 3. Arnold
few other species will grow, or weed protection Arboretum, Harvard University, Jamaica Plain,
must be given almost in perpetuity. Management MA. 586 p.
of existing stands should consist mainly of
protection from fire, development, or heavy Hoyos-F., J. and A. Braun. 2001. Palmas en
grazing. Venezuela. Monograph 47. Sociedad de
Ciencias Naturales La Salle, Caracas,
Venezuela. 424 p.
Benefits.—Brittle thatch palm helps protect the
soil, contributes to the aesthetics of natural stands, Institute of Systematic Botany. 2002. Thrinax
and furnishes food and cover for wildlife. The morrisii. Institute of Systematic Botany,
species is grown and sold as an ornamental. It is University of South Florida, Tampa, FL.
used as a container plant and for buffer strips in http://www.plantatlas.usf.edu/main.asp?plantID
parking lots and medians on highways as well as =555. 2 p.
background and specimen plantings (Gilman and
Watson 1994). Leaves are still used to make Little, E.L., Jr., R.O. Woodbury, and F.H.
brooms, thatch, and mats (called petate) (Osvaldo Wadsworth. 1974. Trees of Puerto Rico and the
2002), although now for ornamentation rather than Virgin Islands. Vol. 2. Agriculture Handbook
necessity. The fruit pulp contains 4,083 µg/g of 449. U.S. Department of Agriculture,
oxalate—capable of causing a burning sensation Washington, DC. 1024 p.
on the skin of sensitive people—that may reduce
its palatability to some species of wildlife Liu, H., S.G. Platt, and C.K. Borg. 2002. Seed
(Broschat and Meerew 2000). However, it is eaten dispersal by the eastern box turtle (Terrapene
in Florida by the introduced Mexican red-bellied carolina) in subtropical pine rockland of the
767
Lower Florida Keys. Botany 2002 Conference, Florida. Pineapple Press, Inc. Sarasota, FL.
August 2-4, Madison, WS. Abstract. 391 p.
http://www.botany2002.org/section3/abstracts/3
6.shtml. 1 p. Osvaldo, J. 2002. El petate. http://joseosvaldo.
freeyellow.com/Petate/html. 12 p.
Long, R.W. and O. Lakela. 1976. A flora of
Tropical Florida. Banyan Books, Miami, FL. Zona, S. and K. Maidman. 2000. Growth rates of
962 p. palms in Fairchild Tropical Garden. Palms
45(3): 151-154.
Nelson, G. 1996. The shrubs and woody vines of
768
Toxicodendron radicans (L.) Kuntze eastern poison ivy
ANACARDIACEAE
John K. Francis
769
of its range. There is an average growing season of Poison Effects.—The pale yellow oil called
240 days in southern Florida, and a mean annual urushiol, present in all plant parts, is the source of
temperature of 19 °C in Arizona (Pavek 1992). all the allergenic pain and suffering associated
The mean annual temperatures are probably with this species. Even in minute amounts, if it
higher, precipitation greater, and growing seasons stays in contact with human skin for more than 5
may approach 365 days in Central America. minutes, it will be absorbed and metabolized. The
Eastern poison ivy is moderately tolerant of shade. metabolites bind with proteins, forming
It is often the most important understory plant, compounds that the immune system recognizes as
climbs nearly every tree in many Southern river “foreign” and attacks. The immune response
floodplain forests, and is a component of most results in rash, blisters, and an intense itch
forest and vegetation types in its ample range. It (Borialforest.org 2003) that lasts for 1 to 3 weeks.
also grows in full sunlight in forest openings, Sensitivity can vary over the life of a person. A
fencerows, bluffs, railroad tracks, and vacant lots. few people have life-threatening reactions to the
Eastern poison ivy grows in old-growth forests in allergen (Schwartz 1999). The chemical is the
some portions of its range and is an early invader same and the effect (called rhus dermatitis) is the
during succession in many environments. It is same for all Toxicodendrons. Ten to 15 percent of
tolerant of fire, browsing, insects, and disease, and the U.S. population is completely immune to rhus
competes aggressively with weeds and brush. dermatitis, 25 to 35 percent react only to high
doses, and 50 percent have a consistent reaction to
Reproduction.—Eastern poison ivy blooms from relatively small doses of the allergen (Family
May through June in Virginia (Von Essen 2000) Practice Notebook.com 2003). The best defense is
and from March through May in Mexico to avoid contact with the plant, especially with the
(Secretaría de Madio Ambiente y Recursos sap or smoke from burning Toxicodendron
Naturales 2003). The start of flowering material. The next line of defense is to quickly
corresponds with leaves being about half formed wash the exposed skin with soap and copious
in the spring (Pavek 1992). The flowers are amounts of water, or if not available, with mud
pollinated by insects, including honey bees (Apis and water or an alkaline material such as baking
mellifera L.) (Pollen Lab 2003). Good crops of soda or wood ashes and water. After a rash
fruits and seeds are produced annually. Seeds are develops, the sufferer is advised to apply
dispersed by birds and mammals and sometimes hydrocortisone cream or any of several creams
by water. Seeds that have been regurgitated from sold over-the-counter for the treatment of rhus
the crop or passed through the digestive tract of dermatitis. A folk remedy mentioned in the
birds demonstrate excellent germination (Pavek literature (Felter 1922) and found by the author to
1992). Eastern poison ivy spreads extensively by almost instantly and completely (but temporarily)
means of scandent stems that root along their stop the itch and irritation of rhus dermatitis is to
whole lengths and below-ground stems (rhizomes) rub the mashed juicy stems of jewel weed
that send up shoots frequently. (Impatiens spp.) on the affected area.
Growth and Management.—Eastern poison ivy Benefits.—Eastern poison ivy, for its dark green
grows rapidly after establishment and assumes foliage during the summer and bright fall colors,
several forms. It scrambles along the ground and adds considerably to the beauty of forests and
sometimes forms thick mats. When given the shrublands where it grows. It has been used to a
opportunity, it ascends trees as a woody vine. limited extent as an ornamental and for
Plants take 3 years to reach the flowering stage conservation plantings. Eastern poison ivy is an
from seed (Pavek 1992). It often forms diffuse important browse species. It is one of the seven
stands of short upright stems. It can be partially most important food species for white-tailed deer
controlled by grazing, but there is a resurgence of in Indiana (Pavek 1992). Goats are said to like
Eastern poison ivy after secession of grazing eastern poison ivy and have been used to control it
because of the reduction in competition. Spot (Schwartz 1999). The fruits are an important fall,
spraying or wiping with the herbicide glyphosate winter, and early spring food for upland game
during the active growing season can effectively birds and many species of song birds (Pavek
control it (Pavek 1992). Grubbing and mowing 1992). Rabbits and rodents also eat the leaves and
offers some degree of control but puts the worker fruits. In herbal medicine, extracts of the leaves
at risk of contracting dermatitis. have been and still are used to treat herpetic
770
eruptions, palsy, and rheumatism. Small internal Natural Resources Conservation Service. 2003.
doses will act as a sedative but must be used with Plants profile: Toxicodendron radicans (L.)
care (Felter 1992, Grieve 2003). Kuntze. http://plants.usda.gov/cgi_profile.
cgi?symbol=TORA2. 6 p.
Griffiths, M. 1994. Index of garden plants. Timber Von Essen, T. 2000. Poison Ivy, Toxicodendron
Press, Portland, OR. 1,234 p. radicans. Wildwood Park, Radford, VA.
http://www.radford.va.us/Community/wildwood
Long, R.W. and O. Lakela. 1976. A flora of /today/Plant_articles/Poison_ivy.htm. 2 p.
Tropical Florida. Banyan Books, Miami, FL.
962 p.
771
Toxicodendron rydbergii (Small ex Rydb.) Greene western poison ivy
ANACARDIACEAE
John K. Francis
772
Seedlings are necessary to establish new colonies almost instantly and completely (but temporarily)
but vegetative (by sprouts from underground stop the itch and irritation of rhus dermatitis is to
stems) reproduction is more important for local rub the mashed juicy stems of jewel weed
spread. (Impatiens spp.) on the affected area.
Growth and Management.—Western poison ivy Benefits.—Western poison ivy helps protect the
grows at a moderate rate. Clones are relatively soil and adds beauty in the summer with its dark
stable over time. Although it is not as invasive as green leaves and in autumn with bright colors to
eastern poison ivy, it is often desirable to eliminate forest and shrublands. Although it causes browsing
it from areas of heavy human use. Spot spraying or animals no ill effects, western poison ivy is low in
wiping with the herbicide glyphosate during the protein and energy and consequently is only
active growing season can effectively control occasionally grazed by domestic animals and wild
eastern poison ivy (Pavek 1992) and probably will ungulates. However, the fruits are eaten by quail,
be effective for western poison ivy. Grubbing and wild turkeys, and some songbirds. It does provide
mowing offers some degree of control but puts the cover for small mammals and birds (McMurray
worker at risk of contracting dermatitis so that 1988). In herbal medicine, extracts of the leaves of
protective clothing should be worn during the this and other Toxicodendron species have been
operation. Branches and roots should be piled to and still are used to treat herpetic eruptions, palsy,
rot in an out-of-the-way place or bagged and sent and rheumatism. Small internal doses will act as a
to the landfill, never burned. sedative but must be used with care (Grieve 2003).
Pavek, D.S. 1992. Toxicodendron radicans. In: St. John, H. 1963. Flora of Southeastern
U.S. Department of Agriculture, Forest Service, Washington and adjacent Idaho. Outdoor
Rocky Mountain Research Station, Fire Sciences Pictures, Escondido, CA. 583 p.
Laboratory, Fire Effects Information System.
http://www.fs.fed.us/database/feis/plants/vine/ Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
toxrad/all.html. 29 p. Higgins, eds. 1987. A Utah flora. Great Basin
Naturalist Memoirs 9. Brigham Young
University, Provo, UT. 894 p.
774
Trema lamarckianum (J.A. Schultes) Blume West Indian trema
ULMACEAE
John K. Francis
775
trema lives about 10 to 20 years. The species has Arboretum, Harvard University, Jamaica Plain,
not been reported as weedy. Reproduction MA. 673 p.
probably can be promoted by scarifying the soil
near seed-bearing plants before the start of Liogier H.A. 1985. Descriptive flora of Puerto
seasonal rains. Rico and adjacent islands, Spermatophyta. Vol.
1. Editorial de la Universidad de Puerto Rico,
Benefits.—West Indian trema is an important Río Piedras, PR. 352 p.
colonizer of disturbed areas and helps protect the
soil from erosion. It has not been used for, but Little, E.L., Jr. and F.H. Wadsworth. 1964.
might be useful for, site stabilization plantings. Common trees of Puerto Rico and the Virgin
West Indian trema is listed as a nitrogen-fixing Islands. Agriculture Handbook 249. U.S.
species (Winrock International 2002). The wood is Department of Agriculture, Forest Service,
soft and seldom used. The fruits are one of the Washington, DC. 548 p.
most important foods of the endangered Puerto
Rican plain pigeon, Columba inornata wetmorei Long, R.W. and O. Lakela. 1976. A flora of
(Division of Endangered Species 2002). Tropical Florida. University of Miami Press,
Coral Gables, FL. 962 p.
References
Massey, J.R. and J.C. Murphy. 1996. Vascular
Division of Endangered Species. 2002. Puerto plant systematics. Section B. General characters
Rican plain pigeon, Columba inornata and character states: 9. Patterns. http://www.
wetmorei. U.S. Fish and Wildlife Service. ibiblio.org/botnet/glossary/b_ix.html. 2 p.
http://endangered. Fws.gov/i/b/sab40.html. 3 p.
Winrock International. 2002. Nitrogen fixing trees
Howard R.A. 1988. Flora of the Lesser Antilles, and shrubs. http://www.winrock.org/forestry/
Leeward and Windward Islands. factpub/nftlist.htm. 29 p.
Dicotyledoneae, Part 1. Vol. 4. Arnold
776
Trichilia hirta L. broomstick
MELIACEAE
John K. Francis
777
blotter paper, 100 percent of them germinated Institution, Washington, DC. http://ctfs.si.edu/
between 17 to 232 days after sowing. Germination webatlas/spanish/tri2hi.html. 2 p.
is hypogeal, and they appear to be recalcitrant
(lose viability if dried beyond a certain point) Howard, R.A. 1988. Flora of the Lesser Antilles,
(author’s observation). In Panama, the seeds are Leeward and Windward Islands.
animal-dispersed (Center for Tropical Forest Dicotyledoneae, Part 1. Vol. 4. Arnold
Science 2002). In Puerto Rico, seedlings are Arboretum, Harvard University, Jamaica Plain,
relatively common but only an occasional MA. 673 p.
individual lasts more than a few weeks or months.
Killeen, T.J., E. García-E., and S.G. Beck, eds.
Growth and Management.—Broomstick in 1993. Guía de árboles de Bolivia. Herbario
Puerto Rico appears to have a moderate growth Nacional de Bolivia and the Missouri Botanical
rate and to live about 10 to 30 years. Management Garden, La Paz, Bolivia and St. Louis, MO.
information is not available. 958 p.
Benefits.—Broomstick adds to secondary forest Liogier, H.A. 1988. Descriptive flora of Puerto
biodiversity, helps protect the soil, and furnishes Rico and adjacent islands, Spermatophyta. Vol.
food and cover for wildlife. The species is used to 2. Editorial de la Universidad de Puerto Rico,
a limited extent for landscaping and is considered Río Piedras, PR. 481 p.
a honey plant. The wood is of medium density
(specific gravity 0.5), moderately hard and Little, E.L., Jr. and F.H. Wadsworth. 1964.
somewhat brash, and is resistant to attack by dry- Common trees of Puerto Rico and the Virgin
wood termites (Criptotermes brevis (Walker)). Islands. Agriculture Handbook 249. U.S.
Another source lists the wood density as 0.48 to Department of Agriculture, Forest Service,
0.96 at 12 to 15 percent moisture content (World Washington, DC. 548 p.
Agroforestry Centre 2002). It has been used for
oars, broom handles, and rustic furniture; today the Núñez-Meléndez, E. 1982. Plantas medicinales de
wood is mainly used for fuel, stakes, and fence Puerto Rico. Editorial de la Universidad de
posts (Little and Wadsworth 1964). An infusion of Puerto Rico, Río Piedras, PR. 498 p.
leaves, flowers, and roots has been used as a
treatment for skin ulcers and as an emetic that has Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
the unfortunate side effect of causing abortions Montiel, eds. 2001. Flora de Nicaragua.
and bleeding. Young branches were strewn on Monographs in Systematic Botany Vol. 85, No.
floors to control fleas and lice (Núñez Meléndes 2. Missouri Botanical Garden Press, St. Louis,
1982). MO. p. 945-1,910.
778
Trichostigma octandrum (L.) H. Walt. hoop vine
PHYTOLACCACEAE
John K. Francis
unpleasant aftertaste. The seeds are 4 to 5 mm in
diameter, black, and shiny (Acevedo-Rodríguez
1985, Howard 1988, Liogier 1985).
779
seeds/kg. Sown in commercial potting mix, 32 Anonymous. 2002. Contribución a la flora polínica
percent of these seeds germinated between 10 and de Noreste Argentina. http://www.unne.edu.ar/
71 days after sowing. Birds apparently disperse cyt/2001/6-Biologicas/B-048.pdf. [not paged].
the seeds. Natural seedlings are not common.
Stems layer (root) whenever they are covered with Educational Concerns for Hunger. 2002. Tropical
soil or organic material. vegetables. http://www.echonet.org/eln&herbs/
eln_catalog/veggies.htm. 4 p.
Growth and Management.—Growth of hoop
vine is rapid from sprouts, as much as 3 m/year. Gann, G.D., K.A. Bradley, and S.W.
Established plants appear to live for several Woodmansee. 2001. Status of South Florida
decades, about as long as the trees they use for flora. Institute of Regional Conservation.
support. No management experience has been http://www.Regionalconservation.org/CH2.htm.
published. 10 p.
Benefits.—The young leaves of hoop vine are Howard R.A. 1988. Flora of the Lesser Antilles,
cooked and eaten as a vegetable after the cooking Leeward and Windward Islands.
water is discarded to remove the bitter flavor Dicotyledoneae, Part 1. Vol. 4. Arnold
(Educational Concerns for Hunger 2002). Both the Arboretum, Harvard University, Jamaica Plain,
split stems and the bast fibers of hoop vine are MA. 673 p.
used for making baskets. The stems are also used
for making bent furniture and craft items (Ruiz Instituto Nacional de Biodiversidad. 2002. Lista de
2002). Juice from the fruits leaves a purple stain especimenes de Trichostigma octandrum. http://
on whatever it contacts and may have potential as www.inbio.ac.cr.bims/k03/p13/c045/o0245/f015
a natural stain for cloth and crafts. The species 56/g008574/s026880.htm. 2 p.
contributes to biodiversity and soil stability, and
provides food and cover for wildlife. In herbal Liogier H.A. 1985. Descriptive flora of Puerto
medicine, the vegetative parts are used as a Rico and adjacent islands, Spermatophyta. Vol.
treatment for heart palpitations, and a powder of 1. Editorial de la Universidad de Puerto Rico,
the root bark is applied topically to treat colds and Río Piedras, PR. 352 p.
water retention (Liogier 1990).
Liogier, H.A. 1990. Plantas medicinales de Puerto
References Rico y del Caribe. Iberoamericana de Ediciones,
Inc., San Juan, PR. 566 p.
Acevedo-Rodríguez, P. 1985. Los bejucos de
Puerto Rico. Vol. 1. General Technical Report Ruiz, D.L. 2002. Cuba. Food and Agriculture
SO-58. U.S. Department of Agriculture, Forest Organization of the United Nations. http://
Service, Southern Forest Experiment Station, www.fao.org/docrep/t2354s/t2354s0r.htm. 4 p.
New Orleans, LA. 331 p.
780
Triumfetta semitriloba Jacq. burweed
TILIACEAE
John K. Francis
781
chromosome count of 2n = 32 (Long and Lakela Collevatti, R.G., L.A.O. Campos, and J.H.
1976). In Brazil, solitary and social bees visit and Schoereder. 1997b. Foraging behaviour of bee
pollinate burweed (Collevatti and others 1997b). It pollinators on the tropical weed Triumfetta
is an abundant seed producer. Fruits collected in semitriloba: departure rules from flower patches.
Puerto Rico averaged 0.0040 + 0.0016 g/fruit. Insectes Sociaux 44(4): 345-352.
Seeds separated from those fruits averaged 0.0039
+ 0.0001 g/seed or 256,000 seeds/kg. Placed on Haselwood, E.L. and G.G. Motter, eds. 1966.
filter paper, 41 percent germinated between 10 and Handbook of Hawaiian weeds. Hawaiian Sugar
119 days from sowing. Germination is epigeal. Planters Association. Honolulu, HI. 479 p.
Seeds are dispersed when they cling to passing
animals. Howard, R.A. 1989. Flora of the Lesser Antilles,
Leeward and Windward Islands. Vol. 5. Arnold
Growth and Management.—Burweed lives Arboretum, Harvard University, Jamaica Plain,
between 1 and 3 years. In seasonally dry habitat, MA. 604 p.
this species behaves as an annual. The plants
grow about 1 m the first year, with growth slowing Liogier, H.A. 1994. Descriptive flora of Puerto
in subsequent years. Because of its weedy nature, Rico and adjacent Islands. Vol. 3. Editorial de la
planting burweed is unlikely, and ill-advised Universidad de Puerto Rico, Río Piedras, PR.
outside its native range. Indeed, control is often 461 p.
needed in croplands and pastures. Treatments
follow normal practice for coarse broadleaf weeds: Long, R.W. and O. Lakela. 1976. A flora of
mechanical cultivation or grubbing and spraying tropical Florida. Banyan Books, Maimi, FL.
with herbicides such as 2,4-D. 962 p.
Benefits.—The stem wood has moderate strength Moanalua Gardens Foundation. 1999. The mystery
and hardness, but because of the plant’s small size of Rapa Nui. Moanalua Gardens Foundation,
the wood is of little value. Fibers from the stem Honolulu, HI. http://gamma.mhpcc.edu/voyage/
bark were once used by the inhabitants of Rapa moanalua/6_rapa.pdf. 20 p.
Nui to make fishing lines (Moanalua Gardens
Foundation 1999). Burweed is reported to have National Geographic Society. 2001. Rapa Nui and
little or no forage value for domestic livestock Sala-y-Gomez subtropical broadleaf forests.
(Haselwood and Motter 1966). National Geographic Society, Washington, DC.
http://nationalgeographic.com/wildworld/
References profiles/terrestrial/oc/oc0111.html. 3 p.
Collevatti, R.G., M.E.C. Amaral, and F.S. Lopes. Pacific Island Ecosystems at Risk. 2001. Invasive
1997a. Role of pollinators in seed set and a test plant species: Triumfetta semitriloba Jacaquin,
of pollen limitation hypothesis in the tropical Tiliaceae. http://www.hear.org/pier/trsem.htm.
weed Triumfetta semitriloba (Tiliaceae). Revista 2 p.
de Biologia Tropical 45(4): 1,401-1,407.
782
Turbina corymbosa (L.) Raf. Christmas vine
CONVOLVULACEAE
John K. Francis
783
ground, and spraying the resulting sprouts with New Orleans, LA. 331 p.
broad-leaf weed killers, should be effective.
Correll, D.S. and M.C. Johnston. 1970. Manuel of
Benefits.—Christmas vine is sometimes grown as the vascular plants of Texas. Texas Research
an ornamental for the clusters of white, scented Foundation. Renner, TX. 1,881 p.
flowers it produces during the early winter. The
nectar gathered from its flowers makes one of the Howard, R.A. 1989. Flora of the Lesser Antilles,
finest honeys (Woman’s Club of Havana 1952). Leeward and Windward Islands.
Extracts from the seeds are used as an analgesic in Dicotyledoneae. Part 3. Vol. 6. Arnold
herbal medicine (Schultes and Hoffmann 1992). Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
Narcotic Properties.—The seeds of Christmas
vine were valued as a sacred hallucinogen by Liogier, H.A. 1995. Descriptive flora of Puerto
Chinantec, Mazatec, Mixtec, Zapotec, and other Rico and adjacent islands. Vol. 4. Editorial de la
groups in Southern Mexico in Pre-Columbian Universidad de Puerto Rico, Río Piedras, PR.
times and are still cultivated and used today as 617 p.
aids in divination and witchcraft. It was
administered by grinding about 13 seeds, adding Long, R.W. and O. Lakela. 1971. A flora of
water, filtering, and drinking the filtrate in a quiet, Tropical Florida. University of Miami Press,
secluded place. Hallucinations follow that last Coral Gables, FL. 962 p.
about 3 hours, sometimes with aftereffects. The
active ingredients are the ergoline alkaloids, Pacific Island Ecosystems at Risk. 2002. Invasive
lysergic acid amide, and lysergic acid plant species: Ardisia elliptica Thunberg,
hydroxyethylamide that are closely related to LSD Myrsinaceae. http://www.hear.org/pier/
(Schultes and Hoffmann 1992). arell.htm. 2 p.
784
Urena lobata L. Caesar weed
MALVACEAE
John K. Francis
785
percent germinated in 9 months. Germination is References
epigeal (author’s observation). Sulfuric acid
scarification was tested as a means of breaking Bautista, L.M.A. 2000. Inquiry into the anti-
dormancy of seed from Sierra Leone. The best inflammatory activity of the syrup from the
treatment (18 molar solution for 90 minutes) gave glycosides of the leaves of kulutkulutan (Urena
96 percent germination as compared with distilled lobata, Linn., family Malvaceae). Centro
water that gave 4 percent germination (Harris Escolar Universitario, Mandiola, Philippines.
1986). Seeds are dispersed by clinging to fur and http:// www.ceu.edu.ph/research1.htm. 3 p.
clothing. Seedlings are common in disturbed sites
near seed sources. Câmara de Comércio e Indústria Portugal-Angola.
2003. Caracterização económico social de
Growth and Management.—Caesar weed grows Angola. http://www.cciportugal-angola.pt/
rapidly and can reach 0.5 to 2 m by the end of the caract.htm. [not paged].
first year. In Puerto Rico, it can live 2 years,
usually dying back to midheight after the first Fagundes, M.H. 2003. Sementes de juta e malva:
growing season. It is not known whether the algumas observações. Companhia Nacional de
shrubs sprout from the roots multiple times. A Abastecimento, Brazil. http://www.conab.gov.
fiber crop yielding 1,800 kg/ha is ready to harvest br/politica_agricola/Conjunturasemanal/Especiai
after 6 to 7 months and seed crops of 300 to 500 s/SEMENTES%20JUTA%MALVA.doc. 10 p.
kg/ha are ready after 7 or 8 months (Fagundes
2003). Fiber crops are established with seed. FAO. 2003. Food composition table for use in
Because of the aggressive nature of the species, Africa. Food and Agriculture Organization of
wildland plantings are not recommended. The the United Nations, Rome. http://www.fao.
author knows of no published specific control org/docrep/003/x6877e/X687705.htm#ch5. [not
measures. paged].
Benefits and Detriments.—Caesar weed Forest Research Institute of Malaysia. 2003. Plants
colonizes disturbed areas and helps to protect the information: Urena lobata Griff., pulut-pulut,
soil while furnishing cover for wildlife. It has Malvaceae. Forest Research Institute of
attractive flowers and contributes to aesthetics of Malaysia, Kuala Lumpur, Malaysia. http://www.
areas it has colonized. Tens of thousands of tons of frim.gov.my/tu/Urena.htm. 1 p.
a jute-like fiber from Caesar weed called aramina
fiber and Congo jute are produced in Brazil Freedman, R. 1998. Famine foods: Malvaceae.
(Fagundes 2003) and Africa (Câmara de Comércio Perdue University, Perdue, IN. http://www.hort.
e Indústria Portugal-Angola 2003). Various perdue.edu/newcrop/faminefoods/ff_families/M
extracts of leaves and roots are used in herbal ALVACEAE.html. 3 p.
medicine to treat such diverse ailments as colic,
malaria, gonorrhea, fever, wounds, toothache, and Harris, P.J.C. 1986. Dormancy of Urena lobata L.
rheumatism (Forest Research Institute of Malaysia seeds. I. Development of sulphuric acid
2003). A semipurified glycoside obtained from scarification techniques. Ghana Journal of
Caesar weed leaves was 86 percent as effective an Agriculatural Science 14-19: 79-84.
anti-inflammatory as aspirin in rats (Bautista
2000). The leaves and flowers are eaten as famine Howard, R.A. 1989. Flora of the Lesser Antilles,
food in Africa (Freedman 1998). Raw leaves are Leeward and Windward Islands. Vol. 5. Arnold
reported to contain 81.8 percent moisture, 54 cal, Arboretum, Harvard University, Jamaica Plain,
3.2 g of protein, 0.1 g fat, 12.8 g carbohydrates, MA. 604 p.
1.8 g fiber, and 2.1 g ash, 558 mg calcium, and 67
mg of phosphorous per 100 g (FAO 2003). Institute of Systematic Botany. 2003. Atlas of
However, the plant is little browsed by cattle and Florida vascular plants. University of South
can become a severe weed in pastures and Florida, Tampa, FL. http://plantatlas.usf.
plantations. Burs that collect on clothing and in edu/synonyms.asp?plantID=1364&genus=Urena
animal fur are a nuisance. &species=lobata. [not paged].
786
Liogier, H.A. 1994. Descriptive flora of Puerto Pacific Island Ecosystems at Risk. 2003. Urena
Rico and adjacent Islands. Vol. 3. Editorial de la lobata L., Malvaceae. http://www.hear.org/
Universidad de Puerto Rico, Río Piedras, PR. Pier/urlob.htm. 3 p.
461 p.
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Natural Resources Conservation Service. 2003. Montiel, eds. 2001. Flora de Nicaragua.
Plant profile: Urena lobata L., Caesarweed. Monographs in Systematic Botany Vol. 85, No.
http://plants.usda.gov/cgi_bin/plant_profile.cgi? 2. Missouri Botanical Garden, St. Louis, MO.
symbol=URLO. 5 p. p. 945-1,910.
787
Urera baccifera (L.) Gaud. ortiga brava
URTICACEAE
Synonyms: none
John K. Francis
788
All parts of the plant are covered with the spine- Bertoni, J.E. de A., F.R. Martins, J.L. de Moraes,
like hairs that cause considerable pain when they and G.J. Shepherd. 1988. Composicão floristica
penetrate the skin. Normal clothing does not e estrutura fitossociologica do parque de
protect against the stinging effect. However, the Vacununga, Santa Rita do Passa Quatro, Sao
pain is usually short-lived, lasting from a few Paulo, gleba Praxides. Boletim Tecnico do
seconds to a few minutes. Although no permanent Instituto Florestal, Sao Paulo 42: 149-170.
injury results, some people may experience rash or
blisters; swelling, fever, and ulcers can occur in Francis, J.K. and A. Rodríguez. 1993. Seeds of
extreme cases (Allen 1943). The plant has long Puerto Rican trees and shrubs: second
been used in folk medicine. Recent laboratory tests installment. Research Note SO-374. U.S.
using rats have shown anti-inflamatory and Department of Agriculture, Forest Service,
analgesic activity of aqueous extracts of ortiga Southern Forest Experiment Station, New
brava (Badilla and others 1999). Amerindians in Orleans, LA. 5 p.
Costa Rica chastised themselves with branches of
ortiga brava in a toughening ritual (Badilla and von Hagen, V.W. 1943. Mexican paper-making
others 1999). plants. Journal of the New York Botanical
Garden 44: 1-10.
References
Little, E.L., Jr., R.O. Woodbury, and F.H.
Allen, P.H. 1943. Poisonous and injurious plants Wadsworth. 1974. Trees of Puerto Rico and the
of Panama. Journal of Tropical Medicine Virgin Islands. Vol. 2. Agriculture Handbook
23(suppl.): 3-76. 449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
Badilla, B., G. Mora, A.J. Lapa, and J.A. Silva
Emim. 1999. Anti-inflammatory activity of Nascimento, H.E.M., A. da S. Dias, A.A.J.
Urera baccifera (Urticaceae) in Sprague-Dawley Tabanez, and V.M. Viana. 1999. Tree
rats. http://www.ots.duke.edu/tropibiojnl/claris/ population and dynamics of a semidecidous
47-2/badilla.html. seasonal forest fragment in the Piracicaba
region, Sao Paulo State, Brazil. Revista
Brasileira de Biologia 59(2): 329-342.
789
Vernonia albicaulis Pers. Santa María
ASTERACEAE
John K. Francis
790
References Liogier, H.A. 1997. Descriptive flora of Puerto
Rico and adjacent islands. Vol. 5. Editorial de la
Howard, R. A. 1989. Flora of the Lesser Antilles, Universidad de Puerto Rico, San Juan, PR.
Leeward and Windward Islands. Vol. 6. Arnold 436 p.
Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
791
Vernonia borinquensis Urban Puerto Rican ironweed
ASTERACEAE
John K. Francis
792
Disturbance is probably necessary for seedling makes it somewhat more difficult to walk through.
establishment. The plants layer (root) whenever
stems come in contact with the soil. References
793
Vernonia proctori Urbatsch Proctor’s vernonia
ASTERACEAE
Synonyms: none
John K. Francis
794
References de extinción en Puerto Rico. Puerto Rico
Ecológico Vol. 3. First Book Publishing of
Liogier, H.A. 1997. Descriptive flora of Puerto Puerto Rico, San Juan, PR. 91 p.
Rico and adjacent islands. Vol. 5. Editorial de la
Universidad de Puerto Rico, Río Piedras, PR. U.S. Fish and Wildlife Service. 2002. Species
436 p. accounts: Vernonia proctorii. http://endangered.
fws.gov/i/q/saqa9.html. 2 p.
Miner-Solá, E. 1999. Arboles y plantas en peligro
795
Vernonia sericea L.C. Rich. long-shoot
ASTERACEAE
John K. Francis
796
g/seed or 3.2 million seeds/kg. Placed on moist to the aesthetic beauty of forests, and furnishes
filter paper, 69 percent germinated over the course cover for wildlife. The species is one of the nectar
of 2 months, beginning 7 days after sowing. sources for the butterfly, Eurema leuce antillarum
Hall (Torres Bauzá 1999).
Growth and Management.—Long-shoot grows
between 0.5 and 1 m in the first year from sprouts. References
The growth rate of seedlings is undoubtedly much
slower. Individual stems live about 3 years, but Liogier, H.A. 1997. Descriptive flora of Puerto
plants sprout and live through a second, perhaps Rico and adjacent islands. Vol. 5. Editorial de la
several, cycles of tops. No propagation or Universidad de Puerto Rico, Río Piedras, PR.
management experience has been published. 436 p.
Presumably, conditions for healthy populations
can be created by maintaining broken or low Torres Bauzá, J.A. 1999. Ciclo de vida de Eurema
basal-area forest stands. leuce antillarum Hall (Lepidoptera: Pieridae) en
Puerto Rico. Caribbean Journal of Science 35(3-
Benefits.—Long-shoot helps protect the soil, adds 4): 195-200.
797
Waltheria indica L. sleepy morning
STERCULIACEAE
John K. Francis
798
These seeds were sown without any pretreatment ethnobotanical dictionary. http://www.ars-
on filter paper and after 16 weeks yielded 13 grin.gov/duke/dictionary/tico/w.html. 4 p.
percent germination. Reproduction is by seeds.
The seeds are disbursed by water, agricultural Burkill, H.M. 2000. The useful plants of West
equipment (Sánchez and Uranga 1993), and Tropical Africa. Royal Botanic Gardens, Kew,
grazing animals. Seedlings are relatively common UK. 686 p.
in disturbed habitat.
Centro de Desarrollo de Agronegocios. 2000. Guia
Growth and Management.—Sleepy morning prictica de manejo de plagas y enfermedades del
plants in Puerto Rico live for 1 or 2 and chile. CDA Boletín Técnico de Producción 13.
occasionally 3 years. Death usually occurs during http://www.hondurasag.org/fintrac-cda/pubs/
the dry season. Perennial growth is more likely in Prod_13_Esp.pdf. 4 p.
continuously moist habitat. Stands of sleepy
morning in agricultural plantations can be Dick, C. 2001. Amazon plant list. Ecotour
controlled by cultivation and probably by Expeditions, Inc. http://www.naturetours.com/
broadleaf herbicides. Am_lispl.html. 8 p.
Benefits and Detriments.—In the Turks and Faftine, O., A. Alage, and J.P. Muir. 2001.
Caicos Islands, sleepy morning is used to make an Characterization of forage selected by cattle on
herb tea (Wood 2001). The plant produces a fiber communal range in Manhição, Mozambique.
that was formerly used for making cords, sacking, http://stephenville.tamu.edu/~jmuir/faftine.htm.
padding, and sandals (Guzmán 1975). Durawhite, 6 p.
an extract of sleepy morning, is used in a
commercial cosmetic for its ability to inhibit Guzmán, D.J. 1975. Especies útiles de la flora
melanin synthesis and whiten the skin (Janssen Salvadoreña. Ministerio de Educación,
Cosmeceutical Care 2001). The plant contains Dirección de Publiciones, San Salvador, El
steroid derivatives and alkaloids of the adouetine Salvador. 703 p.
group that perhaps make it physiologically active.
Various extracts are used as standard febrifugal, Haselwood, E.L. and G.G. Motter. 1966.
purgative, emollient, tonic, analgesic, and Handbook of Hawaiian weeds. Experiment
astringent herbal medicines in Africa (Burkill Station/Hawaiian Sugar Planter’s Association.
2000). In Hawaii, the root is chewed to relieve Honolulu, HI. 479 p.
sore throat (Neal 1965). Stems are used as a chew
stick, and extracts of the plant are used as an eye Howard, R.A. 1989. Flora of the Lesser Antilles,
bath and a remedy for hemoptysis in Panama Leeward and Windward Islands. Vol. 5. Arnold
(Agricultural Research Service 2001). Seeds are Arboretum, Harvard University, Jamaica Plain,
sold commercially, and the species is cultivated in MA. 604 p.
gardens as a medicinal plant. The plant is browsed
by all types of livestock, especially when young Janssen Cosmeceutical Care. 2001. Supreme
(Burkill 2000). Sleepy morning forage in a secrets. http://www.janssen-beauty.com/
Mozambique valley during the rainy season supreme.htm. 11 p.
contained 6.4 percent crude protein, 0.12 percent
phosphorus, and 0.51 percent calcium (Faftine and Lastres, L.. 2000. Guia practica de manejo de
others 2001). Sleepy morning is considered a plagas y enfermedades del chile. CDA Technical
weed in much of its range, but it is seldom Production Bulletin 13. Centro de Desarrollo de
aggressive enough to be a major problem Agronegocios. http://www/jpmdirasag/org/
(Sánchez and Uranga 1993). It is host for a fintrac-cda/pubs/Prod_13_Esp.pdf. 4 p.
number of insects harmful to agricultural crops
(Centro de Desarrollo de Agronegocios 2000, Liogier, H.A. 1994. Descriptive flora of Puerto
Lastres 2000, Sánchez and Uranga 1993). Rico and adjacent Islands. Vol. 3. Editorial de la
Universidad de Puerto Rico, Río Piedras, PR.
References 461 p.
Agricultural Research Service. 2001. Tico Long, R.W. and O. Lakela. 1976. A flora of
799
Tropical Florida. Banyon Books, Miami, FL. plant checklist: Sterculiaceae. Texas A & M
962 p. University, Department of Botany.
http://www.csdl.tamu.edu/FLORA/ftc/dft/
Neal, M.C. 1965. In gardens of Hawaii. Special ftc_str.htm. 1 p.
Publication 50. Bernice P. Bishop Museum
Press, Honolulu, HI. 924 p. Tropilab, Inc. 2001. Waltheria indica L.
http://www.tropilab.com/sleepy_waltheria.html.
Nelson, G. 1996. The shrubs and woody vines of 2 p.
Florida. Pineapple Press, Inc., Sarasota, FL.
391 p. University of Hawaii. 2001. Plants of Hawaii
Volcanoes National Park.
Sánchez, P. and H. Uranga. 1993. Plantas http://www.botany.edu/b308/bigisland/species/
indeseables de importancia economica el los waltheria/waltheria.htm. 3 p.
cultivos tropicales. Editorisal Científico-
Técnica. La Habana, Cuba. 166 p. Wood, K.M. 2001. Roadside wildflowers.
http://www.timespub.tc/Natural%History/
Texas A & M University. 2001. Texas vascular Archive/Summer2001/wildflowers.htm. 2 p.
800
Wedelia reticulata DC. manzanilla de monte
ASTERACEAE
John K. Francis
801
protect the soil, and furnishes cover and possibly Francis, J.K., S. Alemañy, H.A. Liogier, and G.R.
food for wildlife. Although somewhat rough in Proctor. 1998. The flora of Cañón de San
appearance during dry periods, it has pretty Cristóbal, Puerto Rico. General Technical
flowers and should make a fine ornamental for Report IITF-4. U.S. Department of Agriculture,
natural landscaping, especially in coastal areas and Forest Service, International Institute of Tropical
rocky terrain. Forestry, Río Piedras, PR. 37 p.
802
Ziziphus obtusifolia (Hook. ex Torr. & Gray) Gray graythorn
RHAMNACEAE
James E. Nellessen
803
Reproduction.—Flowers of graythorn are small, potassium, and iron were at levels to meet adult
inconspicuous, and occur in axillary cymes of two goat requirements during all seasons. Phosphorus
to seven flowers. The five-petalled flowers are was low in all seasons except the spring. Native
greenish yellow. Graythorn may bloom at various Americans have used the plant for medicinal
times during the growing season, from May purposes. Pima Indians have steeped the roots in
through September, depending on the availability water and applied them to sore eyes. Seri Indians
of moisture. The fruit is a dark blue to black have powdered the roots for application to skin
globose drupe, 6 to 10 mm long, covered with a and scalp sores. Solutions made from roots also
whitish bloom, and matures from July into early served as a soap substitute (Bowers and Wignall
October. Seeds are nondormant, but the seed coats 1993, Epple 1995, Kearney and others 1951).
are impermeable to water, hence are classified as
physically dormant (Baskin and Baskin 2001). References
Physically dormant seeds need some type of
scarification of the seed coat to initiate water Allred, K.W. 2002. A Working Index of New
penetration and germination. Optimal germination Mexico Vascular Plant Names. Available on a
occurs at 25 °C. Both light and prechilling New Mexico State University web page:
maximized the percent germination (Speer and http://web.nmsu.edu/~kallred/herbweb. [not
Wright 1981, Young and Young 1992). Graythorn paged].
seeds from the lower Rio Grande valley in
southern Texas did not need pretreatment for Baskin, C.C. and J. M. Baskin. 2001. Seeds:
germination (Vora 1989). Ecology, Biogeography, and Evolution of
Dormancy, and Germination. Academic Press,
Growth and Management.—Graythorn stem San Diego, CA. 666 p.
regeneration generally occurs just below the cut or
removed portion. When stems are removed to the Bowers, J.E. and B. Wignall. 1993. Shrubs and
ground line, regeneration will occur from the root Trees of the Southwest Deserts. Southwest
crown, and stems can be regenerated from roots, Parks and Monuments Association. Tucson,
but generally only when the root crown itself is AZ. 140 p.
removed (Flinn and others 1992). Graythorn may
demonstrate rapid growth when resprouting from Carter, J.L. 1997. Trees and Shrubs of New
cut stumps or during periods of good moisture but Mexico. Johnson Books, Boulder, CO. 534 p.
is generally a slow growing shrub. Although this
species may on occasion become locally dense and Correll, D.S. and M.C. Johnston. 1970. Manual
form thickets, it is generally a species that is of the Vascular Plants of Texas. Texas Research
diffusely distributed within its habitat, such that Foundation, Renner, TX. 1,881 p.
management should not be a problem.
Dick-Peddie, W.A. 1993. New Mexico
Benefits.—The fruits are readily eaten by birds Vegetation: Past, Present and Future.
such as Gambel's quail and white-winged doves. University of New Mexico Press, Albuquerque,
Graythorn was among several shrubs in a northern NM. 244 p.
Mexico study testing the preferred browse
composition and digestibility of foliage. Graythorn Epple, A.O. 1995. A Field Guide to the Plants of
was grouped with species having a high crude Arizona. Falcon Publishing Inc., Helena, MT.
protein content, lower cell wall content, and 347 p.
smoother or waxy leaf surfaces with the absence of
trichomes. Range goats and white-tailed deer Flinn, R.C., C.J. Scifres, and S.R. Archer. 1992.
included graythorn as an important part of their Variation in basal sprouting in co-occurring
diets (Ramirez and others 1997). In another shrubs: implications for stand dynamics.
nutrient study in northern Mexico (Ramirez and Journal of Vegetation Science 3(1): 125-128.
others 2001) graythorn was among several shrubs
studied for seasonal mineral levels. General Foster, M.A., C.J. Scifres, and P.W. Jacoby, Jr.
mineral concentrations were higher in the spring 1984. Herbaceous vegetation - lotebush
and summer, but only calcium, magnesium, (Ziziphus obtusifolia var. obtusifolia)
804
interactions in north Texas USA. Journal of Ramirez, R.G., G.F.W. Haenlein, and M.A.
Range Management 37(4): 317-320. Nunez-Gonzalez. 2001. Seasonal variation of
macro and trace mineral contents in 14 browse
Great Plains Flora Association. 1986. Flora of the species that grow in northeastern Mexico. Small
Great Plains. University Press of Kansas, KS. Ruminant Research 39(2): 153-159.
1,392 p.
Ramirez, R. G., J.L. Pineiro-Hernandez, and R.K.
Kartesz, J.T. 1994. A Synonymized Checklist of Maiti. 1997. Nutritional profile and leaf surface
the Vascular Flora of the United States, Canada, structure of some native shrubs consumed by
and Greenland. Vol. 1, 2nd Edition. Biota of small ruminants in semiarid regions of
North America Program of the North Carolina northeastern Mexico. Journal of Applied
Botanical Garden. Timber Press, Portland, OR. Animal Research 11(2): 145-156.
622 p.
Speer, E.R. and H.A. Wright. 1981. Germination
Kearney, T.H., R. Peebles, and Collaborators. requirements of lotebush (Ziziphus obtusifolia
1951 with 1960 supplement. Arizona Flora. var. obtusifolia). Journal of Range Management
University of California Press, Berkeley, CA. 34: 365-368.
1,085 p.
Vora, R.S. 1989. Seed germination characteristics
Martin, W.C. and C.E. Hutchins. 1980-1981 of selected native plants of the lower Rio
(reprinted 2001). A Flora of New Mexico. Vol. Grande valley Texas, USA. Journal of Range
1. Bishen Singh Mahendra Pal Singh, India and Management 42(1): 36-40.
Koeltz Scientific Books, Germany. 1,276 p.
Young, J.A. and C.G. Young. 1992. Seeds of
McCoy, D. 1981. Roadside Trees and Shrubs of Woody Plants in North America. Dioscorides
Oklahoma. University of Oklahoma Press, Press, Portland, OR. 407 p.
Norman, OK. 116 p.
Zitzer, S.F., S.R. Archer, and T.W. Boutton. 1996.
Nelson, J.A., P.W. Barnes, and S. Archer. 2002. Spatial variability in the potential for symbiotic
Leaf demography and growth responses to N-2 fixation by woody plants in a subtropical
altered resource availability in woody plants of savanna ecosystem. Journal of Applied Ecology
contrasting leaf habit in a subtropical savanna. 33(5): 1125-1136.
Plant Ecology 160(2): 193-205.
805
Zuckia brandegei (Gray) Welsh & Stutz ex Welsh siltbush
CHENOPODIACEAE
806
predation is low (Pendleton and others 2000). Between population differences in dormancy
Wind and gravity disperse the fruits over time; and response to temperature. In: Proceedings,
some fruits generally remain on the plant until late Symposium on cheatgrass invasion, shrub die-
winter. There are about 625,000 seeds/kg off, and other aspects of shrub biology and
(Pendleton and others 2000). Germination management. U.S. Department of Agriculture,
experiments conducted with seeds of Z. b. var. Forest Service, General Technical Report INT-
brandegei and Z. b. var. plummeri showed that 276. Intermountain Research Station, Ogden,
seeds of warm winter populations germinated over UT. p. 187-292.
a wide range of constant temperatures (15 to 30
o
C) (Meyer and Pendleton 1990). Seeds of cold- Pendleton, R.L., D.C. Freeman, E.D. McArthur
winter populations were dormant at fall and winter and S.C. Sanderson. 2000. Gender specialization
temperatures and required exposure to overwinter in heterodichogamous Grayia brandegei.
chilling to promote germination. Germination American Journal of Botany. 87:508-516.
generally increased with duration of wet
prechilling at 1 oC for up to 8 weeks, dry Pendleton, R.L., E.D. McArthur, D.C. Freeman,
afterripening for up to 14 months, or bract removal and A.C. Blauer. 1988. Heterodichogamy in
(Meyer and Pendleton 1990). Germination is Grayia brandegei (Chenopodiaceae): Report
epigeal. from a new family. American Journal of Botany.
75: 267-274.
Growth and Management.—Few data are
available. Siltbush is a stress-tolerant species Pendleton, R.L., S.D. Nelson, and R.L. Rodriguez.
capable of establishing on steep, eroded sites 1996. Do soil factors determine the distribution
where the combination of salinity, aridity, and fine of spineless hopsage (Grayia brandegei)? In:
soils preclude establishment of other species Proceedings, Symposium on shrubland
(Pendleton and others 1996). Seedling growth rate ecosystem dynamics in a changing climate.
and palatability of mature plants are moderate General Technical Report INT-GTR-338. U.S.
(personal communication with Steve Monsen, Department of Agriculture, Forest Service,
USDA-FS-RMRS, Provo, UT, and Howard Stutz, Intermountain Research Station, Ogden, UT. p.
Brigham Young University, Provo, UT). Plants 205-209.
may attract rodents, other small animals, and deer.
Shaw, N.L., R.L. Pendleton, and E.G. Hurd. 2001.
Benefits.—Siltbush is a potential revegetation and Zuckia brandegei (Gray) Welsh & Stutz ex
soil stabilization species for disturbances resulting Welsh. In: F.T. Bonner and R.G. Nisley, eds.
from mining and other human activities on sites Woody Plant Seed Manual. U.S. Department of
where it is native. Few other species will grow on Agriculture, Forest Service, Washington, D.C.
these outcrops (Pendleton and others 1996). http://wpsm.net/index. html. 8 p.
807
Glossary of Technical Terms Used in this Volume
808
Disjunct population—a population isolated from the main part of the range.
Diurnal—occurring (blooming) during the day.
Dominant—a crown class in forest or shrub canopy in which the individual is taller than the general
canopy level; may also refer to species with a preponderance of numbers or biomass.
Drupe—fleshy, one-seeded fruit whose seed is encased in a stony endocarp.
Ecotone—a zone of transition between adjacent plant communities.
Ecotype—a population within a species that is genetically adapted to a certain habitat distinct from that
which is general for the species.
Edaphic—pertaining to the soil and its ecological relationships.
Endosperm—starchy or oily tissue in many seeds.
Entire—without teeth or indentations on the margin.
Epigeal—germination and emergence in which the cotyledons raise above the soil surface.
Escapes—introduced populations now established and competing in the wild and spreading beyond the
original point of introduction.
Excessively drained soil—water runs off, infiltrates, or evaporates rapidly so that the soil tends to be
droughty even with ample rainfall.
Floret—individual, usually minute, flower of a compound flower.
Fruticose—being woody or shrub-like.
Galleries—small patches of vegetation (usually trees or shrubs) in protected locations distinct from
surrounding vegetation.
Germinative capacity—the percentage of seeds that germinate during the normal period of germination.
Glabrous—not hairy; hairless or nearly so.
Glaucous—with a waxy bloom or whitish substance that rubs off.
Habit—the growth form or appearance of a plant.
Head—a compact, multiple inflorescence, often disk-shaped.
Hammock—a fertile area in Florida and Southeastern United States that is elevated above the surrounding
terrain.
Hypanthium—a floral tube formed by the fusion of the basal portions of the sepals, petals, and stamens
from which the rest of the floral parts arise.
Hypogeal—germination and emergence in which the cotyledons remain below the soil surface.
Inceptisols—order of soils that are young, frequently rocky, and have a poorly developed profile.
Intermediate—a crown class of tree or shrub canopies in which the individual plant crown is within and
does not quite rise to the level of the top of the canopy.
Lanate—soft, woolly pubescence.
Layering—rooting at a point on a stem where the stem comes in contact with the soil or with artificial
treatment called air layering.
Legume—one to multiple-seeded fruit (pod) common in the Fabaceae family.
Lenticels—lenticular or diamond-shaped dots, pits, or corky protrusions on stems.
Litter—organic debris on the surface of the soil or the uppermost layer of the forest floor organic horizon.
Maceration—the process of soaking and gently grinding fruits to remove the pulp from seeds.
Mesic—moist soil or environment; neither excessively wet nor dry.
Mine spoil—dirt and rock left after a mining operation.
Monoecious—having staminate (male) and pistillate (female) flowers on the same plant.
Naturalized—naturally reproduced progeny of horticultural plants or wildland plantings outside their
native range.
Nerve—vein, usually used in relation to leaves.
Nut—a one-seeded fruit enclosed in an involucre (diminuative=nutlet).
Old growth forest—characterized by old, usually large, and often decadent trees.
Open forest—a forest characterized by low basal area, a large amount of space between crowns, and a lot
of sunlight reaching the forest floor.
Open-grown—growing without significant competition from plants of similar size or taller.
Panicle—an inflorescence with indeterminate branching, the primary axis bearing branched secondary axes
and pedicellate flowers.
Pappus—hairs attached to small seeds that together function as a wing.
Perfect flower—a flower with both stamens (male parts) and carpels (female parts).
Pericarp—the wall of a ripened ovary (fruit).
809
pH—a measure of soil acidity or alkalinity; values above 7.0 are alkaline and values below are acid.
Phenotype—a plant as it exists; the product of the interaction of genetics and environment.
Pioneer—shrub or other plant that establishes on or has the ability to colonize newly disturbed habitat.
Pith—spongy central cylinder of parenchyma tissue in stems.
Ploidy—the degree of repetition of the basic chromosome number.
Poorly drained soil—water runs off or evaporates so slowly that the soil remains wet for a large part of the
year.
Provenance—the original geographic source of a plant’s genetic material.
Puberulent—covered with soft, very fine hairs.
Raceme—an inflorescence with indeterminate single axis and pedicellate flowers.
Rachis—the axis of a pinnately compound leaf.
Remnant (forest)—lightly to moderately disturbed pieces or stands of the original forest.
Reticulate—with a net-like structure.
Rhizome—modified underground stem.
Riparian vegetation—plant associations and stand types growing along streams.
Samara—a one-seeded, winged fruit.
Savannah—tropical or subtropical grassland with scattered trees; usually maintained by periodic fires.
Scabrous—covered with scattered, short, stiff hairs.
Scandent—vine-like habit.
Scarify—to perforate or make a seed coat permeable by mechanical or chemical means.
Scrub—vegetation consisting of shrubs and sometimes trees stunted because of poor site conditions.
Secondary forest—forest that has grown up after clearing or lesser disturbance.
Selfing—pollination of a plant with its own pollen.
Seral—transitional as a site or vegetation association moves toward a climax state.
Serrulate—finely serrate.
Sessile—connected directly to the stem or other structure without a stalk.
Shade intolerant—requiring full or nearly full sunlight to survive, grow and reproduce.
Shade tolerant—the ability to survive, grow, and reproduce in the shade of taller vegetation, as under a
tree canopy.
Skeletal soil—substrate in which stones form the structure of the soil with sand, silt, and clay filling the
voids between.
Slash—the cut stems and limbs of trees and brush, or to cut unwanted trees and brush leaving it in place.
Soil texture—classes defining the proportion of sand, silt, and clay making up a soil.
Spike—indeterminate inflorescence of sessile flowers arranged linearly along a central axis.
Stamen—male or pollen-producing part of the flower.
Stipule—a small leaf-like structure found at the base of some leaf petioles, usually present in pairs.
Stolon—a stem or shoot that bends to the ground and takes root after contact.
Stratify—breaking the physiological dormancy of certain seeds with a period of cold or warm incubation.
Succulent—class of plants with fleshy leaves and usually stems that store water in their tissues and usually
have special metabolic pathways (crassulan acid metabolism).
Sucker—root sprouts or to reproduce by root sprouts; sometimes used to mean an epichormic branch.
Suffruticose—perennial plant with stems that are woody near their base.
Suppress—inhibit the growth (and often eventually kill) by shading and competition for water and
nutrients.
Synonym—an equivalent but superseded species name.
Testa—the outer coat of a seed.
Ultisols—a family of highly weathered soils found in subtropical and tropical environments.
Ultramafic rocks—metamorphic rocks (including serpentine) characterized by low percentages of silica
and elevated percentages of iron and magnesium.
Variety—a subdivision of a species that has one or more inherited morphological characteristics.
Well-drained soil—water runs off, infiltrates, or evaporates readily but not rapidly.
Xeric garden—a garden planted with drought-resistant vegetation that is not irrigated.
810
References Consulted
Burns, R.M. and B.H. Honkala, eds. 1990. Silvics of North America. Vol. 2. Agriculture Handbook 654.
U.S. Department of Agriculture, Forest Service, Washington, DC. 877 p.
Editorial staff. 1984. Webster’s ninth new collegiate dictionary. Marriam-Webster Inc., Publishers,
Springfield, MA. 1,563 p.
Lawrence, G.H. 1955. An introduction to plant taxonomy. The Macmillan Company, New York. 179 p.
Soil Survey Staff. 1951. Soil survey manual. Agriculture Handbook 18. Agriculture Research
Administration, U.S. Department of Agriculture, Washington, DC. 503 p.
811
Common Name Index
Common name page Common name page
‘a’ali’i 293 amourette-grand-bois 487
‘a’ali’i ku makani 293 amourette-rivière 489
‘a’ali’i-ku ma kua 293 amouretto 489
abeyelo 235 angelica tree 40
abrojo Colorado 412 angel’s trumpet 36, 286
abuta 212 angel’s whisper 299
abutilon espinoso 540 anguila 327
acacia bush 21 añil 386
acacia courant 289 añil de pasto 386
acacia savane 289 añil falso 252
acacia zarza 19 anisette 567
aceituna Americana 117 anisillo 564, 567
aceituno 154 annatto 112
acequia willow 664 antelope bitterbrush 610
achiote 112 antelope brush 610
afata 714 antelope buckbrush 610
aguacero 127 antibala 714
aguedita 561 Antilles velvetshrub 570
ague tree 681 Apache-plume 337
aguinaldo blanco 783 aperta ruão 564
agüiste 489 applebush 361
‘aheahea 190 aramina 785
aile à ravet 764 arabo carbonero 323
aji 147 arazá-puitá 597
aji de gallina 144 árbol de navidad 352, 617
aji picante 147 arbre de soie 136
akalakala 658 arctic willow 660
akeake 293 arête-boeuf 529
akulikuli-kai 107 argythamnia 45
alcaparro 696 Arizona queen-of-the-night 181
alcotá 212 Armenian blackberry 652
alcotán 567 Aroeira pimenteira 686
alderleaf mountain-mahogany 178 aroma 14
alfilerillo 459 aromo 21
algodón de seda 137 arrowleaf sida 714
algorobo 580 arrowwood 379
alkali sagebrush 66 ash barberry 464
allamanda 36 Asian bittersweet 161
aloes vert 350 ati popa’a 43
alpine dryad 295 atiu 43
alpine juniper 401 auguey 475
amansa 684 ‘aweoweo 190
American angelica tree 40 azahar 508
American beautyberry 135 azota-caballo 299
American bittersweet 164 azúcares 395
American burning bush 333 baby tea 196
American ivy 532 badoh 783
American plum 586 Bahaman wild coffee 608
American snowball 737 balsam 473, 619
American yew 758 balsaminha 142
amor ardiente 393 bálsamo 372
amourette 19 basbasot 714
812
Common Name Index
bitter bush 196, 561, 619
bamboo piper 564 bitter cane 370
banglin 489 bitter cherry 589
baquiña 432 bitter nightshade 721
barbasco 395 bittersweet nightshade 721
basam 275 bizcocho 395
bashful plant 489 black ambret 487
basket willow 664 blackbead 575
basket wiss 779 blackbead tree 235
basket wood 708 blackberry 43, 652
basora 247 black bush 781
basora prieta 798 black hawthorn 254
bastard guava 329 black mimosa 489
bastard sirio 30 black sage 247, 676
bâtard balengène 726 black sagebrush 72
bayberry 502 black sumac 625
bay-cedar 742 black torch 319
bay grape 229 black widow 473
bayonette 125 black willow 668
beach creeper 321 blackwith 708
beach plum 591 blind-eye-tree 374
beautyberry 135 boater bush 798
bébèl 575 bog myrtle 511
bejuco de berac 192 bois-anisette 432
bejuco de canastas 708 bois bolibri 354
bejuco de corrales 708 bois cabrit 32, 524, 696
bejuco de costilla 538, 708 bois cabroit 32
bejuco de fuego 359 bois candelle 38, 319
bejuco de gato 456 bois canon 137
bejuco de guajanilla 538 bois capable 684
bejuco de indio 359 bois caraibe 761
bejuco de membrillo 717 bois-champignon 473
bejuco de mona 213 bois codine 115
bejuco de nasa 779 bois crabbe 575
bejuco de palma 779 bois de fer 32, 235
bejuco de paloma 538, 779 bois de l’encore 524
bejuco de paralejo 764 bois énivrant 219
bejuco de sarsaparilla 717 bois flambeau 38, 186, 319
bejuco de sopla 359, 689, 764 bois fleche 341
belly-ache bush 399, 619 bois flot 376
bemberecua 769 bois-foufou 354, 524
berenjena cimmarona 726 bois-guillaume 365
berenjena de gallina 726 bois jaune 149
berenjena de playa 724 bois mêche 140
berenjena silvestre 726 bois montagne 561
Bermuda mulberry 135 bois-noir 140
Bigelow sagebrush 50 bois petites feuilles 327
big sagebrush 85 bois poison 561
bija 112 bois-puant 140
bilimbi 505 bois pini 38
birchleaf cercocarpus 178 bois puante 699
birchleaf mountain-mahogany 178 bois tan 154
bird pepper 144, 147 bola de coral 393
biscuitwood 149 bomba 137
bonbon bleu 223
813
Common Name Index
814
Common Name Index
815
Common Name Index
816
Common Name Index
817
Common Name Index
818
Common Name Index
819
Common Name Index
820
Common Name Index
821
Common Name Index
822
Common Name Index
823
Common Name Index
824
Common Name Index
825
Common Name Index
826
Common Name Index
827
Authors of Thamnic Descriptions
G.W. Andrews, USDA Forest Service, Forestry Sciences Laboratory, 520 Devall Drive, Auburn, AL
36830
Eric L. Berlow, Research Scientist, University of California San Diego, White Mt. Research Station, 3000
E. Line St., Bishop, CA 93514
Don C. Bragg, Research Forester, USDA Forest Service, Southern Research Station, P.O. Box 3516 UAM,
Monticello, AR 71656
Mary Carrington, Assistant Professor, Science Division, Governors State University, 1 University
Parkway, University Park, IL 60466
Barton D. Clinton, Research Ecologist, USDA Forest Service, Southern Research Station, Asheville, NC
28802
Kristina Connor, Plant Physiologist, USDA Forest Service, Center for Bottomland Hardwoods Research, Box
9681, Mississippi State, MS 39762
Ann M. DeBolt, Botanist, USD Forest Service, Rocky Mountain Research Station, Boise, ID 83702
Margaret S. Devall, Research Ecologist, USDA Forest Service, Center for Bottomland Hardwoods
Research, Stoneville, MS 38776
John K. Francis, formerly Research Forester, U.S. Department of Agriculture, Forest Service, International
Institute of Tropical Forestry, Jardín Botánico Sur, 1201 Calle Ceiba, San Juan PR 00926-1119, in
cooperation with the University of Puerto Rico, Río Piedras, PR 00936-4984, now retired and volunteering
with the USDA Forest Service, Shrub Sciences Laboratory, 735 N 500 E, Provo, UT 84606
Marshall R. Haferkamp, Rangeland Scientist, Fort Keogh Livestock and Range Research Laboratory,
USDA Agricultural Research Service, Miles City, MT 59301
Emerenciana G. Hurd, Botanist, Rocky Mountain Research Station, U.S. Department of Agriculture,
Forest Service, Boise, ID 83702
Deborah K. Kennard, Research Ecologist, USDA Forest Service, Forestry Sciences Laboratory, 320
Breen Street, Athens, GA 30602
Stanley G. Kitchen, Research Botanist, USDA Forest Service, Rocky Mountain Research Station, Shrub
Sciences Laboratory, 735 N. 500 E. Provo, UT 84606
Juanita A. R. Ladyman, Botanist, JnJ Associates, 6760 S. Kit Carson Cir. E., Centennial, CO 80122
Ariel E. Lugo, Director, International Institute of Tropical Forestry, U.S. Department of Agriculture,
Forest Service, Ceiba 1201, Jardín Botanico Sur, Río Piedras, PR 00926
Jinshuang Ma, Research Taxonomist, Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn,
NY 11225
E. Durant McArthur, Research Geneticist, USDA Forest Service, Rocky Mountain Research Station, Shrub
Sciences Laboratory, Provo, UT 84606
828
Paul A. McMillan, Coordinator, Biology Department, Capilano College, 2055 Purcell Way, North Vancouver,
British Columbia, Canada V7J 3H5
W. Henry McNab, Research Forester USDA Forest Service, Southern Research Station, Asheville, NC
28802
Stephen B. Monsen, retired and volunteering with the USDA Forest Service, Shrub Sciences Laboratory,
735 N 500 E, Provo, UT 84606
Arlee M. Montalvo, Plant Population Biologist, Department of Botany and Plant Sciences, University of
California, Riverside, CA 92521-0124
Gerry Moore, Research Taxonomist, Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, NY
11225
James E. Nellessen, Botanist, Biologist and Environmental Scientist, Taschek Environmental Consulting,
8901 Adams St., NE, Albuquerque, NM 87113
Kenneth Outcalt, Research Ecologist, USDA Forest Service, 320 Green Street, Athens, GA 30602
Paula M. Pijut, Plant Physiologist, USDA Forest Service, North Central Research Station, Hardwood Tree
Improvement and Regeneration Center, 195 Marsteller Street, West Lafayette, IN 47907
John A. Parrotta, Biological Scientist, Research & Development, USDA Forest Service, Washington, DC
20090-6090
Rosemary L. Pendleton, Research Ecologist, USDA Forest Service, Rocky Mountain Research Station,
Albuquerque, NM 87102
Sally A. Reynolds, Botanist, U.S. Fish and Wildlife Service, San Francisco Bay National Wildlife Refuge
Complex, P.O. Box 524, Newark, CA 94560
Christopher Ross, Natural Resource Specialist, Bureau of Land Management, Nevada State Office, P.O. Box
12000, Reno, NV 89520
Stewart C. Sanderson, Research Geneticists, USD Forest Service, Rocky Mountain Research Station,
Shrub Sciences Laboratory, 735 N. 500 E., Provo, UT 84606
Nathan M. Schiff, Research Entomologist, USDA Forest Service, Center for Bottomland Hardwoods
Research, Stoneville, MS 38776
Randy S. Senock, Assistant Professor of Tropical Forestry, College of Agriculture, Forestry and Natural
Resource Management, University of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720
Nancy L. Shaw, Research Botanist, USDA Forest Service, Rocky Mountain Research Station, Boise, ID
83702
Amanda J. Stevens, student, College of Agriculture, Forestry and Natural Resource Management,
University of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720
829
Jeffrey R. Taylor, formerly Biological Technician, USDA Forest Service, Rocky Mountain Research
Station, Shrub Sciences Laboratory, 735 N. 500 E., Provo, UT 84606, now student, University of Utah, Salt
Lake City, UT
Sarah A. Taylor Student, College of Agriculture, Forestry and Natural Resource Management, University
of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720
Bruce L. Welch, Plant Physiologist, USDA Forest Service, Rocky Mountain Research Station, Shrub
Sciences Laboratory, 735 N. 500 E., Provo, Utah 84606
830
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