Cabombaceae/Cactaceae 161

Khanna, P. 1965. Morphological and embryological studies in

Nymphaeceae II. Brasenia schreberi Gme!. and Nelumbo
Cactaceae
nucifera Gaertn. Aust. J. Bot. 13: 379-387.
Kosakai, H. 1968. The comparative xylary anatomy of various W.BARTHLOTT and D. R.HUNT
Nymphaeaceae. M. A. Dissertation, Univ. Calif., Santa Bar-
bara.
Langlet, 0., Soderberg, E.1929. Uber die Chromosomenzahlen
einiger Nymphaeaceen. Acta Horti Bergiani 9: 85-104.
Les, D. H. 1988. The origin and affinities of the Ceratophylla-
ceae. Taxon 37: 326-345.
Les, D.H.Garvin, D.K.Wimpee, e.F. 1991. Molecular evol-
Cactaceae A. L. de lussieu (1789) (Cacti), nom. cons.
utionary history of ancient aquatic angiosperms. Proc. N atl.
Opuntiaceae Kunth in Humboldt, Bonpland and Kunth (1823).
Acad. Sci. USA, 88: 10119-10123.
Moseley, M.F., Mehta, I.J., Williamson, P.S., Kosakai, H. 1984.
Morphological studies of the Nymphaeaceae. XIII. Contribu- Succulent perennials of diverse habit, trees, shrubs,
tions to the vegetative and floral structure of Cabomba. climbers, epiphytes or geophytes; roots fibrous or
Amer. J. Bot. 71: 902-924.
Ni, X-M. 1989. Recent development with aquatic plants and
tuberous; stems columnar, terete, globular, tubercled,
water gardens in China. Water Garden Jour. 5: 36-43. ribbed, winged or flattened, often segmented, mostly
Nitzschke, J. 1914. Beitrage zur Phylogenie der Monokotylen, leafless and variously spiny; leaves, where present,
gegriindet auf die Embryosackentwicklung apokarper Nym- spirally arranged, simple, entire, exstipulate, but
phaeaceen Helobien. Cohns Beitr. BioI. Pflanz. 12: 223-267. usually rudimentary or vestigial; axillary buds devel-
Okada, H., Tamura, M. 1981. Karyomorphological study on the oping a persistent cushion-like indumentum of multi-
Nymphaeales. I. lpn. Bot. 56: 367-375.
0rgaard, M. 1991. The genus Cabomba (Cabombaceae) - a tax-
cellular trichomes (areale), and usually leaves trans-
onomic study. Nord. I. Bot. 11: 179-203. formed into spines. Flowers solitary or rarely
Osborn, J. M., Schneider, E. L. 1988. Morphological studies of clustered, usually sessile at the areoles, rarely pedi-
the Nymphaeaceae sensu lato. XVI. The floral biology of cellate in paniculate or cymose inflorescences (Peres-
Brasenia schreberi. Ann. Mo. Bot. Gard. 75: 778--794. kia) or terminal (e. g. Pterocactus), zoophilous, often
Osborn, J.M., Taylor, T.N., Schneider, E.L. 1991. Pollen mor- conspicuous, nearly always bisexual, usually actino-
phology and ultrastructure of the Cabombaceae: Correla-
tions with pollination biology. Amer. J. Bot. 78: 1367-1378.
morphic; receptacle enclosing, and more or less pro-
Raciborski, M. 1894. Die Morphologie der Cabombeen and duced beyond the zone around the ovary ("pericar-
Nymphaeaceen. Flora 78: 244--279. pel") and between ovary and perianth (epigynous
Ramji, M. v., Padmanabhan, D. 1965. Developmental studies hypanthium; "tube"), naked or invested with bract-
on Cabomba caroliniana Gray. 1. Ovule and carpel. Proc. like scales and areoles, areolar trichomes, hairs
Indian Acad. Sci. 62B: 215-223. and/or spines, the upper scales often intergrading
Rao, T. A., Banerjee, B. e. 1979. On foliar sclereids in the Nym-
phaeaceae sensu lato and their use in familial classification.
with the outer tepals; tepals usually numerous in a
Proc.lndian Acad. Sci. 88: 413-422. graded series; stamens often very numerous; anthers
Rataj, K. 1977. A new species of Cabomba of the Negro river, 2-locullar, tetrasporangiate, dehiscing longitudinally;
Amazonas, Brazil. Acta Amazonica 7: 143. ovary inferior (except some Pereskia spp.), unilocu-
Richardson, F.e. 1969. Morphological studies of the Nym- lar, 3-20-carpellate, placentation usually hypanthial,
phaeaceae. IV. Structure and development of the flower of
rarely basal-Iaminal, ovules numerous; style usually
Brasenia schreberi Gmel. Univ. Calif. Pub!. Bot. 47: 1-101.
Schneider. E.L., leter, J.M. 1982. Morphological studies of the long; stigma 3-20-lobed. Fruit juicy or dry, naked,
Nymphaeaceac. XII. The floral biology of Cabomba caroli- scaly, hairy, bristly or spiny, indehiscent or variously
niana. Amer. J. Bot. 69: 1410-1419. dehiscent. Seeds numerous, sometimes strophiolate,
Schrenk, 1. 1888. On the histology of the vegetative organs of in subfam. Opuntioideae arillate, testa variously
Brasenia peltata, Pursh. Bull. Torrey Bot. Club 15: 29-49. sculptured, endosperm present or absent; embryo
Turlier, M-F. 1984. Early leaf organogenesis in heterophyllous
curved, usually strongly so, or nearly straight; coty-
dicotyledons. The case of Cabomba aquatica (Nymphaea-
ceae). Bull. Soc. Bot. Fr. Lett. Bot. 131: 301-314. ledons reduced or vestigial, rarely foliaceous.
Vinogradov, I.S. 1967. Sistema semeistva Nymphaeaceae na The family is characterised by a high degree of paral-
osnove analiza morfologicheskogo stroya. Zap. Centro Kav- lel evolution in vegetative morphology (e. g. the globu-
kazsk Otd. Vsesojuzn. Bot. Obsc. 2: 5-11. lar habit appears to have evolved independently at
Voronkina, N. V. 1974. Anatomical structure of the root apex in least four times) and in flower-structure (e. g. adapta-
Nymphaeales 1. Schaffner. Bot. Zh. (Leningr.) 59: 1417-1424.
tion to hummingbird- and hawkmoth-pollination in
Walker, 1. W. 1974. Aperture evolution in the pollen of primi-
tive angiosperms. Amer. 1. Bot. 61: 1112-1136. non-related taxa). It comprises ca. 100 genera and ca.
Walker, I. W. 1976. Evolutionary significance of the exine in the 1500 species (though due to horticultural interest
pollen of primitive angiosperms. In: Ferguson, I. K., Muller, 1. many more have been proposed: some 12000 bi-
(Eds.) The evolutionary significance of the exine. New York: nomials now exist). Distribution almost exclusively
Academic Press, (Linn. Soc. Symp. Ser. No.1) pp. 251-308. American with centres of diversity in the drier regions
of SW USA and Mexico, and southern S America. One

K. Kubitzki et al. (eds.), Flowering Plants · Dicotyledons

© Springer-Verlag Berlin Heidelberg 1993
162 Cactaceae

zygomorphic (e. g. Schlumbergera) or functionally

dioecious (Mammillaria dioica, Selenicereus innesii).

VEGETATIVE MORPHOLOGY. Perennials with a broad

spectrum of life-forms, ranging from dwarf button-
like forms (ca. 1 cm diam. in Blossfeldia liliputana ) to
giant columns which may exceed 20 m (Carnegiea
gigantea, Neobuxbaumia spp.), large tree-like forms
(Cereus, Pachycereus etc.) or rarely broad-leaved
trees reaching 15 m (Pereskia lychnidiflora, Fig.28,
Quiabentia verticillata). The stem may be simple
or branched, or sometimes densely caespitose or
mound-forming (e. g. Maihuenia, Opuntia spp.,
Fig. 29, Mammillaria spp.). Scandent species occur in
various groups (e. g. Peres kia, Tacinga, Hylocereeae).
Fig. 28. Cactaceae. Pereskia lychnidiflora, a tree with con- About 10 % of the species are forest epiphytes with
spicuous leaves. Near Guatemala City. (Photo W. Rauh) thin, terete stems (e. g. Rhipsalis spp.) or leaf-like flat-
tened phylloclades (e. g. Disocactus, Epiphyllum).
Root-systems in the family are of six types. Most
taxa have long shallow horizontal roots (e.g. Fero-
cactus, Opuntia). In some columnar cacti (e. g.
Pachycereus schottii) this system is combined with a
central root penetrating to deeper soil. Many small
globular cacti exhibit a compact system of short thin
lateral roots enabling the plant to take up water run-
ning off the plant itself. Other "globular" cacti, espe-
cially those where the growing point is almost at
ground level (e.g. Ariocarpus, Neoporteria, Neower-
dermannia), have a single fleshy taproot, often much
larger than the stem itself. Some slender creeping or
climbing plants develop subterranean, single or
multiple storage roots (e. g. Peres kia, Pterocactus,
Peniocereus, Harrisia). Creeping terrestrial species
Fig. 29. Cactaceae. Opuntia floccosa, a cushion-forming opun- (e. g. Stenocereus eruca) and many epiphytes (e. g.
tioid. Cordillera Blanca, Peru, 4000 m. (Photo W. Rauh) Hylocereus) can exhibit an exclusively adventitious
root-system.
The succulent stems characteristic of cacti show a
species of Rhipsalis which is widespread in tropical high degree of morphological diversity (Fig. 31). The
America also extends to Africa, Madagascar and Cey- initial disposition of the leaf-bases (podaria) and are-
lon. Various Opuntia spp. are widely naturalized in oles conforms to helically alternate phyllotaxy, pro-
warmer regions of the Old World and Australia. ducing varying numbers of orthostichies in the Fibo-
nacci series according to the size and shape of the
CHARACTERS OCCURRING IN RELATIVELY FEW GENERA shoot apex. In most cereoid forms (Cereus, Carnegiea
AND SPECIES. Trees with conspicuous persistent leaves etc.), coalescence of the decurrent podaria in individ-
in Pereskia. Epiphytes with leaf-like flattened stems ual or adjacent orthostichies produces vertical ribs
(phylloclades) in several genera of Hylocereeae and which can support a tall fluted stem capable of con-
Rhipsalideae. Geophytes with napiform storage certina-like expansion and contraction according to
roots in Peniocereus and Pterocactus. Areoles sunken water availability. The number of ribs varies from two
into cortical tissue and thus buds developing within in the phylloclades of flat-stemmed epiphytes to
the stem (several Rhipsalideae). Prolongation of are- more than 100 in the multi costate globular genus
oles with continuous growth and annual flower for- Stenocactus, with the lower numbers in the Fibonacci
mation (e. g. Neoraimondia). Multiple flowers per series (3,5,8,13,21) predominating. Many ofthe cac-
areole (e. g. Myrtillocactus). Areoles of fertile zone toid (globular) forms also have ribbed stems, but
modified with trichomes or hair-spines (cephalium) commonly the podaria remain discrete, as spirally
in Discocactus, Espostoa, Melocactus, Coleocepha- arranged tubercles, particularly in the more highly
locereus and various other genera. Flowers strongly neotenous genera such as Rebutia and Mammillaria.
 Cactaceae 163

The simple conspicuous leaves formed in Pereskia

and most genera of the Opuntioideae are somewhat
succulent with pinnate (approaching pseudopalmate)
or obscure venation. In Maihuenia and Opuntia the
leaves are usually small and terete.
The symmetry of the areole may appear super-
ficially radial (when the spines are all similar) but is
generally bilateral, the flowers arising at or towards
the upper (ventral) edge. In several genera, the are-
ole is more or less elongate (spines pectinate in e. g.
Rebutia spp.), or the spiniferous and floral meristems
somewhat disjunct, with the flowers arising at the end
of a ventral groove. In a few genera with bi-partite
areoles (e. g. Ariocarpus, Mammillaria, Pelecy-
phora ), disjunction is complete, with separate dorsal
portion (usually apical on the podaria) bearing tri-
chomes and (usually) spines only, and a ventral por-
tion (usually spineless) which is capable of producing
flowers and or vegetative shoots.
The areoles of all genera produce an indumentum
of multicellular trichomes and usually spines, repre-
senting modified leaves, rarely (in some Pereskia Fig.30A,B. Cactaceae. A,B Pereskia aculeata. A Paniculate
spp.) leaves alternating with spines. inflorescence. B Flower structure. (A after Vaupel 1925,
Spines are present, at least in the seedling stage, in B after Leuenberger 1986)
virtually all genera. They may be diverse in size and
appearance within a single areole, often forming two
series, popularly termed "central" and "radial". Indi-
vidual spines may be thin and hair-like (e. g. Cepha-
locereus senilis, Espostoa spp.), flattened and papery
(e. g. Pediocactus papyracanthus, Opuntia spp.),
hooked (e.g. Mammillaria and Ferocactus spp.), but
are generally bristly (setaceous) or needle-like
(acicular), and straight or curved. The areoles of
Opuntioideae are characterized by a tuft of minute,
microscopically barbed bristle-spines (glochids).
In some genera, the areoles may continue to pro-
duce trichomes and spines over many years (e. g. old
stems of Pereskia, Opuntia, Browningia), and in a
few species can become prolonged in spur-like struc-
tures producing flowers annually (particularly con-
Fig. 31 A-C. Cactaceae. Vegetative morphology of the three
spicuous in Neoraimondia arequipensis). In most cac-
subfamilies. A Pereskia, non-succulent stem with large subper-
toid genera, however, individual areoles cease sistent leaves and axillary areoles with spines. B Opuntia, succu-
activity after the first or second season. lent stem with small, terete deciduous succulent leaves and axil-
lary areoles with spines (glochids). C Cactoideae, leafless
VEGETATIVE ANATOMY. The vascular system of the globular succulent with well-developed areoles. (After Barth-
cactus stem is intricately linked to the formation of lot! 1979)
leaves and areoles; vascularization is closed in Peres-
kioideae and Opuntioideae, and closed or open in
Cactoideae. The sieve-element plastids are of cen- libriform, remammg alive at maturity; rays storing
trospermous type lacking starch inclusions, but with a abundant starch; wood parenchyma para tracheal.
central protein crystalloid and subspherical ring of Overall, the wood structure can be seen to reflect the
protein filaments (type PIlI cf in the classification of growth-habit of the whole plant: trees and erect
Behnke 1981). shrubs have the longest and widest fibres and vessels;
Even in Peres kia, the wood-anatomy is very spe- globular cacti have short and extremely narrow
cialised: short narrow vessels with a single perfor- vessels and fibres may be absent. Anomalous second-
ation plate and multiseriate pitting; fibres narrow and ary growth is absent in the family.
164 Cactaceae

The cortex is transformed into water-storage crypts) or in Blossfeldia (stomata very few in areolar
tissue, composed of large vacuolised parenchyma crypts: the plant is virtually astomatic).
cells, and there is also some similar water-storage
tissue in the pith. Crystal-forming idioblasts are INFLORESCENCE STRUCTURE AND MODIFICATION OF THE
usually present, and also large mucilage cells in the FERTILE PORTION OF STEMS. True inflorescences occur
cortex (and occasionally the pith) of most genera only in Peres kia, where they may be racemose-
(exceptions include Ferocactus). Mucilage is a very paniculate (P. aculeata) or cymose-paniculate (often
slippery, complex and indigestible carbohydrate. subcorymbose with up to 50 flowers in P. grandifolia).
Extreme mucilage formation occurs in Uebelmannia In Pereskia and various Opuntia spp. additional
gummifera, where lineages of cells form conspicuous flowers may arise from proliferous areoles on the
mucilage ducts in the cortex, visible to the naked pericarpel. Elsewhere in the family, including some
eye. Pereskia species (e. g. P.lychnidiflora) flowers are
Probably derived from mucilage cells are the lati- solitary. Usually they occur at lateral or subapical
cifers present in most species of Mammillaria: their areoles. Terminal flowering at the apex of shoots is
development is significantly different from laticifers restricted to a few taxa (e.g. Pterocactus). Normally
in other dicotyledonous families. They arise from one flower is produced per are ole, rarely two or more
lysis of at least two layers of cells and are encircled by simultaneously (up to 10 in, e.g. Myrtillocactus geo-
a parenchymatous epithelium. The white milky exsu- metrizans) or annually in areoles with indefinite
date termed latex appears to be a polysaccharide growth (e. g. Neoraimondia).
chemically more similar to mucilage than to either In many taxa, floriferous areoles are restricted to
rubber or resin. particular portions of the stem. Areoles at the shoot
Calcium oxalate crystals of very diverse form occur apices of Hatiora and Schlumbergera remain aggre-
in almost all parts of the plant (most frequent in cor- gated as a composite areole, producing subterminal
tex and hypodermis), mostly as solitary prisms or ag- flowers. Fertile areoles are often characterised by
gregated into druses. Occasionally multiple crystals thinner spines and/or a rich production of trichomes
("crystal sand") are found and calcium oxalate ra- (e. g. Cleistocactus spp., Pachycereus schottii).
phids occur in several Hylocereeae. Silica bodies are If the flowering zone is conspicuously modified
only known from the epidermis and hypodermis of and demarcated, it is termed a cephalium. Cephalia
Stenocereus spp. may be developed in a lateral, apical or terminal po-
The epidermis is uniseriate, possibly mUltiple in sition. Lateral cephalia are confined to one or several
Pachycereus; the outer periclinal walls are heavily of the ribs of columnar species, beneath the apex.
cutinized, often strongly impregnated with intracuti- They may be limited to modification of the areoles
cular waxes forming massive cutin-wax layers (e. g. (e. g. Pilosocereus) or modification (reduction) of the
Ariocarpus, Copiapoa); a glaucous appearance is ribs as well, resulting in the production of a groove
caused by epicuticular wax platelets or rodlets (many filled with trichomes, and usually bristles, from which
species, e. g. the "blue" Brazilian Cereeae) or pow- flowers emerge (e. g. Coleocephalocereus). Apical
dery white triterpenoid exsudates (Stenocereus growth continues after the cephalium begins to de-
beneckei); and the outer surface of epidermal cells velop, so that the cephalium may become beard- or
may be flat to papillate (e. g. Opuntia microdasys). fleece-like.
The epidermis of spines also exhibits wide diversity: In species which produce apical cephalia, normal
the most common features are excentric solitary pa- vegetative growth of all the ribs is interrupted an-
pillae on each cell (causing e. g. the plumose spines of nually or permanently and the apex produces modi-
Mammillaria plumosa or the barbed glochids and fied flowering areoles instead. Annual ring- or collar-
spines of Opuntioideae). like cephalia of this type occur in a variety of genera
In several species the dead, outer periclinal walls (e. g. Arrojadoa, Cephalocereus (Neodawsonia) spp.,
disintegrate and form sponge-like ectohydric water- Stephanocereus leucostele, Rhipsalis pilocarpa). Al-
conducting structures on the spine surfaces (e. g. Dis- ternatively, the apex may continue to elongate slowly
cocactus horstii, Pelecyphora aselliformis, many Neo- over a long period, developing into a bristly cap or
lloydia (Turbinicarpus) spp.). In the most strongly brush-like structure (e. g. Pachycereus militaris, Ste-
xeromorphic species, a multiple collenchymatous phanocereus luetzelburgii). In species with terminal
hypodermis is present, providing mechanical stability cephalia, vegetative growth ceases permanently
to the skin. The paracytic or parallelocytic stomata when maturity is reached, and the stem-apex
may be distributed evenly over the entire stem sur- becomes purely fertile and non-assimilatory (Disco-
face or restricted to the lateral sides of ribs; an ex- cactus, Melocactus).
treme discontinuous distribution of stomata occurs, The occurrence of cephalia, and their structure,
for example, in Aztekium (stomata restricted to deep were once considered important systematic charac-
 Cactaceae 165

ters, but it is now apparent that much parallel evol- Fig. 32 A-E. Cactaceae. A,B Tacinga lunalis. Flower with
ution has occurred in the family, and that the cepha- massive tube formed by fleshy pericarp, nectar chamber pres-
ent. C Opuntia subulata, succulent stem-like pericarpel with
lium-types described above are connected by inter- conspicuous terete leaves, pericarpel forming only a short tube,
mediate forms. nectar chamber absent. D Opuntioid seed. E Pereskioid/cac-
toid seed. Curved embryo enclosing perisperm; seed coat black;
FLOWER STRUCTURE AND ANATOMY. FLOWER STRUC- hilum and funicle dotted, the latter forming a bony aril around
TURE IN THIS FAMILY IS HIGHLY DERIVED, BUT DESPITE seed in Opuntioideae. (A-C after Krainz 1956-76; E after
CONSPICUOUS, SUPERFICIAL DIVERSITY IT IS ALSO HIGHLY Barthlott and Voit 1979; D orig.)
UNIFORM. IN Peres kia, five or more carpel primordia
originate separately on the convex floral apex and
subsequently jointly form a coenocarpous gynoe- of the five primordia is still discernible (Leins and
cium. In all other genera the unilocular ovary is Schwitalla 1985). The numerous (ca. 20--4000)
derived from four to more than 20 carpels. Two types stamens bear anthers of the regular two-locular tetra-
of placentation (with transitional links) can be distin- sporangiate type, dorsifixed with complete longitudi-
guished (Leins and Schwitalla 1988): basal-laminal nal dehiscence. Stamens may be inserted in one series
(e. g. Pereskia), with placentae formed on the septa at (e. g. Peres kia, Rhipsalis), evenly distributed over the
the connection to the floral axis, or hypanthial lower, inner part of the hypanthial tube (e. g. Rebu-
placentation, the more common condition, with tia) or arranged in two separate series (e. g. Echi-
placentae formed separately from the septa and alter- nopsis). Staminode-like structures are present in
nating with them. Hypanthial placentae may fuse various genera (e. g. Tacinga).
(e. g. Rhipsalis). The funicles are simple or multiple- The inferior ovary results from intercalary growth
branched. In Opuntioideae the enlarged funicle en- beneath the perianth, forming a more or less pro-
closes the whole ovule and forms the bony aril of the nounced hypanthium (commonly called the tube). In
ripe seed. The numerous ovules are campylotropous, some Cactoideae the hypanthium is very con-
bitegmic, and crassinucellate. spicuous, forming a long tube (e. g. Echinopsis, Epi-
The multistaminate androecium develops basipe- phyllum) which may become very thin and petaloid,
tally (centrifugally). In Pereskia, spiral arrangement resembling a true perianth tube (Schlumbergera). In
166 Cactaceae

Fig. 33 A-G. Cactaceae-Cactoideae. A Epiphyllum phyllan- on the pericarpel and hypanthium of scaleleaves
thus, nocturnal salverform flower pollinated by hawkmoth. (sometimes called bracts or bracteoles), areoles and
B Schlumbergera truncata, diurnal red zygomorphic flower pol-
linated by hummingbird. C Hylocereus sp., scaly fruit with
spines. The pericarpel itself (stem-tissue enclosing
seeds embedded in fleshy pulp. D,E Cleistocactus brookei, tu- the carpels) can be succulent and assimilatory (e. g.
bular slightly zygomorphic flower, pericarpel and hypanthium Opuntioideae). In some genera the transition from
with imbricate scales, stamens inserted in throat of hypanthial pericarpel to tube is abrupt (e. g. Seienicereus, Ste-
tube in two series, the lowermost on a collar practically closing nocereus), and in some the pericarpel may be nearly
a nectar chamber. F,G Rhipsalis baccifera. Small rotundate or completely naked (e. g. Mammillaria, Rhipsalis).
flower, pericarpel naked, stamens in one series surrounding an-
nular nectariferous disk. (A, B after Barthlott 1979; D, E after
The scale-leaves are usually larger and more tepaloid
Rauh 1979; others orig.) towards the apex of the hypanthium, forming a more
or less gradual transition to the numerous tepals
(perianth-segments). The terms sepal and petal can-
not be used here, as the perianth is rarely bi-seriate,
Pereskia and in some Cactoideae with small reduced and then only superficially (e. g. Disocactus bi-
flowers (e, g, Rhipsalis) it is not developed. Nectar is formis). Flower-size varies between 6 mm (some
secreted by a disc (Peres kia, Rhipsalis) or along the Rhipsalis and Pseudorhipsalis spp.) and 40 cm (Hy-
basal portion of the hypanthium. In the latter case locereus, Selenicereus spp.), the latter ranking among
distinct nectar chambers may occur, more or less the largest angiosperm flowers. Frequent colours are
closed by the formation of a dense ring of filament yellow and all shades of red in day-flowering species,
bases, or filamental or hypanthial appendices (e. g. and white in those flowering at night. Blue is absent
Schlumbergera), and in some Opuntia species by an (apart from blue-tinged flowers in some Disocactus
annular or even cup-like outgrowth at the base of the spp.). The flowers are commonly funnelform or cam-
style. panulate, less frequently urceolate, salverform or tu-
In Cactaceae the nature of the flower as a modified bular. Most laterally oriented flowers exhibit a slight
shoot is demonstrable in most genera by the presence bilateral zygomorphy, at least in the androecium
 Cactaceae 167

(Cereus, Hylocereus, Echinopsis), and those which kia zinniiflora, Mammillaria dioica and Selenicereus
are ornithophilous are usually strongly zygomorphic innesii, and incomplete in Coryphantha odorata.
in the perianth and tube as well (e. g. Schlumber- Cleistogamy occurs in Frailea and some polyploid
gera ). Rhipsalis forms in Madagascar. Various natural hy-
brids between related species and related genera are
POLLEN MORPHOLOGY (based on Leuenberger 1976). known, and the relative ease of hybridization has
Pollen consists of isopolar or rarely heteropolar been extensively exploited in horticulture, especially
monads (tetrads in Pseudorhipsalis himantoclada); using members of the Hylocereeae.
grains are medium-sized (Gymnocalycium 36 /lm) to
very large (Opuntia 120/lm), spheroidal to subpro- POLLINATION BIOLOGY. The family is entirely zoophi-
late, rarely suboblate; apertures are frequently 3, 6 lous (Porsch 1938/39). Visitors to entomophilous
or 12 (rarely 15 or up to 31); triaperturate pollen is flowers include Hymenoptera and Lepidoptera; orni-
zonotremous, hexa- to polyaperturate pantotremous; thophilous: Trochilidae; chiropterophilous: Glosso-
apertures are usually colpate, in most Opuntioideae phaginae. Adaptations to beetle-pollination and
porate. Exine is tectate to semi-tectate, nexine pos- deceptive flowers seem to be absent. Abundant re-
sibly with endexine, sexine differentiated into colu- ward-pollen and nectar is produced by some species.
mellae and tectum. Columellae are up to twice as Entomophilous (bee-pollinated) flowers seem to be
high as the thickness of the tectum, more or less the plesiomorphic condition within the family (most
evenly dispersed or almost reticuloid; in reticulate Pereskioideae, Opuntioideae and many Cactoideae).
pollen reticolumellate (almost simplicolumellate). Some yellow-flowered Cactoideae (e. g. Astrophy-
Tectum is perforate (punctate, anulopunctate, foveo- tum) exhibit unique behaviour in the ultraviolet
late) to reticulate in most Opuntioideae, spinulose, range, reflexion or absorption depending on the
rarely verrucose or flat. angle of the tepals towards the light-source, produc-
In general, the pollen exhibits common features as- ing nonpigmented ultraviolet patterns. Psychophi-
sociated with the Centrospermae. Palynology con- lous (butterfly-pollinated) flowers occur in several
firms the isolated and derived position of the Opun- genera (e. g. Rebutia ): these are salverform (hypocra-
tioideae but has had rather limited influence on the teriform), with the tepals at more or less right angles
current classification of the Pereskioideae and Cac- to the slender, tubular hypanthium. Hummingbird-
toideae. flowers (unscented with high nectar production,
mostly red, tubular and often zygomorphic) have
KARYOLOGY. The basic chromosome number (surveys evolved in parallel in most tribes of Cactoideae (e. g.
in Ross 1981, Pinkava et al. 1985) is x = 11 (aneu- Cleistocactus, Disocactus, Schlumbergera, Mammilla-
ploidy n = 12 is reported in Selenicereus testudo); ria poselgeri) and even in some Pereskioideae (P. ste-
most species studied are diploids (2n = 22). No poly- nantha) and Opuntioideae (0. cochenillifera).
ploidy is known in Pereskioideae, but about 15 % of Nocturnal white flowers are widespread in the Cac-
species studied in Opuntioidieae and Cactoideae are toideae, but appear to be absent in the tribes Rhip-
polyploid (mainly tetraploid 2n = 44). Hybridization salideae and Cacteae, and in the subfamilies Opun-
and polyploidy are a source of taxonomic complexity, tioideae and Pereskioideae. Such flowers are either
notably in Opuntia, where polyploidy is often corre- sphingophilous (pollinated by hawkmoths), and
lated with self-fertility, adventive embryony, profuse sometimes very highly adapted (e. g. Epiphyllum
branching and vegetative reproduction. Rhipsalis phyllanthus, Selenicereus wittii, with tube up to 30 cm
baccifera ssp. baccifera is diploid in South America long), or chiropterophilous (bat-pollinated). Bat pol-
but tetraploid towards the northern margin of its neo- lination is common in the Cactoideae, though re-
tropical distribution (e. g. in Florida). The widespread stricted to various cereoid genera (e. g. Carnegiea,
palaeotropical ssp. mauritiana is always tetraploid, Pilosocereus) and hylocereoid epiphytes (e. g. Webe-
and some of the almost cleistogamous populations of rocereus ).
the Madagascan ssp. horrida are octoploid (2n = 88).
The more northern mainland popUlations of Mam- PHYTOCHEMISTRY (Hegnauer 1964, 1989). The cen-
millaria prolifera are tetraploid (2n = 44), and its trospermous betalains are responsible for the bright
Caribbean populations hexaploid (2n =66). A single colours in flowers and fruits; anthocyanins are absent.
report of 2n = 264 (24x) in M. capensis is uncon- At least 60 different alkaloids are known to occur
firmed. in the family (Matta and McLaughlin 1982). Most
common are tyramine alkaloids and phenethyl-
REPRODUCTIVE SYSTEMS. Both allogamy and auto- amines. Mescaline and a number of related peyote al-
gamy occur within the family (often within closely re- kaloids occur in high concentrations in Lophophora
lated species). Functional dioecy is present in Peres- (Anderson 1979), and may be present in several other
168 Cactaceae

genera of all three subfamilies. Tetrahydroisoquino- tral portion of each cell (par-concave: "pitted"). The
line alkaloids occur in Lophophora and most Pa- cuticle may be flat or micropapillate (Cactinae), and
chycereeae. cuticular striations are common in many genera. In
Triterpenes (usually sapogenins) are frequent in all some genera, the hilum and micropyle are separated
groups, some are known only from this family. In by a band of sclerified testa tissue (Peres kia, Echino-
some columnar cacti over 50 glycosidic compounds cactus), but usually in Cactoideae they are conjunct
exist. Sterols are common in the giant columnar Pa- or fused into a single complex, forming the hilum-
chycereeae and can make up almost 20 % of the dry micropylar region (HMR). A mucilage sheath may
weight of the soft tissue. Abundant mucilage (a fi- envelope the entire seed (e. g. Callymanthium) or
brous polysaccharide) is present (see vegetative anat- just the HMR (e.g. Schlumbergera). The HMR is ap-
omy) in many genera; a particular kind of white latex pendaged with a strophiolar pad or strophiole in sev-
(probably derived from mucilage) is present in many eral genera (e. g. Blossfeldia ). In all Opuntioideae the
Mammillaria species. entire seed is covered by a bony aril.
Phenols are present in traces only (e. g. quercetin,
kaempferol, ferulic and sinapic acid). Tannins seem DISPERSAL BIOLOGY. Ornithochory is common
to be absent (except possibly in seed coats). Abun- throughout the family: the juicy, sweet fruits are dis-
dant oxalic acid is produced and appears in calcium persed by birds. Myrmecochory (ant-dispersal) is the
oxalate crystals. rule in several dwarf, globular genera with dry fruits
and strophiolate seeds (e. g. Blossfeldia). Many gen-
FRUIT AND SEED (Buxbaum 1957-60). The cactus fruit era show a double, ornitho-myrmecochorous strategy
is usually a conspicuous, globose to oblong, indehis- (e. g. Cereus): if the mature fruit is not removed by
cent berry with numerous seeds embedded in a fleshy birds it becomes dehiscent and the seeds attract ants
pulp. Colours range from green to white, yellow, all by their fleshy funicles. The extremely spiny fruits of
shades of red, and bright blue to almost black. De- many Opuntia spp. are epizoochorous (mammals)
pending on the occurrence and nature of areoles on and function as burrs. Some taxa also have a success-
the pericarp, the fruit may be naked (Mammillaria), ful vegetative dispersal by easily detached stem-seg-
woolly (Selenicereus spp.), spiny (Armatocereus) or ments with barbed or hooked spines that function as
scaly (Hylocereus). In Pereskia and Opuntia, the are- burrs (Opuntia spp., Mammillaria spp.).
oles of the pericarpel can be proliferous and thus Fruits of some species (e. g. Pereskia bahiensis,
"chains" of fruits may be formed. Spiny areoles may Acanthocereus brasiliensis) ripen after falling to the
be shed when the fruit is ripe (Pachycereus pringlei). ground, become scented, and are eaten by larger
The hypanthium and perianth (floral remnant) can mammals. Anemochory occurs occasionally in sev-
be persistent or deciduous. In many taxa fruits eral taxa within the family, for example the seeds of
become dehiscent when mature. Various types of de- Pterocactus and Espostoa blossfeldiorum are wind-
hiscence are found, from simple longitudinal slits in dispersed. The whole, balloon-like fruits of some
Cereus, to operculum formation in Weberocereus etc. Neoporteria species are also blown along by the wind.
The non-pulpy fruits of Ferocactus spp. and Oreo- Anemochory, in combination with hydrochory, plays
cereus open at their base. Mature fruits of Neobux- a role in the minute ombrohydrochorous seeds of
baumia polylopha and other Pachycereeae open star- some Parodia species and other globular genera, with
like, with several longitudinal slits. the strophiole probably functioning as a floating de-
Seed morphology, in particular the configuration vice. Selenicereus wittii, an epiphyte restricted to the
of the hilum and micropyle, and the microsculpture periodically inundated igap6 forests of Amazonia,
of the testa (seedcoat), exhibits a high level of diver- has modified seeds with enlarged air-filled testa cells:
sity which is of systematic importance (Barthlott and possibly the most peculiar case of an hydrochorous
Voit 1979; Barthlott and Hunt in prep.). The shape epiphyte.
and symmetry of seeds in lateral view vary from sub-
orbicular-lenticular to hat-shaped, with mussel- DISTRIBUTION AND HABITATS. With the exception of
shaped predominating. Size varies from 0.4-7.5 mm Rhipsalis baccifera, the family is exclusively Ameri-
(to 12 mm in Pterocactus spp., where the seed-aril can. The distribution ranges from 57 degrees N in Ca-
forms a samara-like wing), averaging 1-2 mm. The nada (Opuntia) to 50 degrees S in Patagonia (Opun-
testa usually appears black or dark brown and can be tia, Pterocactus, Maihuenia); and across the whole of
rugose or ruminate in overall appearance. The single South America from the Brazilian islands of Fer-
cells show features of taxonomic significance, such as nando de Noronha in the east to the Galapagos
anticlinal undulations (only in the subtribe Cactinae), Archipelago in the west. From all kinds of coastal
and depressions of the cell-junctions ("interstices", in habitats, the vertical distribution ranges up to c.
all Cactoideae apart from the Cactinae) or of the cen- 4500 m in the Central Andes in Peru and Chile (e. g.
 Cactaccae 169

Opuntia fioccosa, Q.lagopus). Few of the genera are the USA and Japan propagating and selling a very
wide-ranging and represented in both North and wide selection of the known species as ornamentals.
South America, notably Opuntia, which occurs Regrettably, many of the more slow-growing species
throughout the area of the family, and Peres kia, Pilo- desired by cactus-fanciers are still imported from
socereus, Harrisia, Hylocereus, Selenicereus, Rhip- their native habitats.
salis and Melocactus, which are more or less wide-
spread in the tropical latitudes. The remaining genera CONSERVATION. Like other plants, many cacti are
are mostly associated with three principal centres of threatend by the destruction of their natural habitats.
diversity, one in North America and two in South Agricultural development is a threat to the highly
America, and many of the genera are of limited dis- adapted and slow-growing species of semi-arid re-
tribution. gions, and forest destruction the main reason for the
The northern centre of diversity is the dry SW disappearance of epiphytic species, many Rhipsali-
United States and Mexico, characterized by the tribes deae being seriously threatend by the destruction of
Cacteae (e. g. Echinocactus, Ferocactus, Mammillaria the Brazilian Atlantic rain forest. Erosion and in-
etc.), the giant columnar Pachycereeae (e.g. Carne- creasing desertification are a threat to even the
giea, Pachycereus etc.) and some Echinocereeae toughest "desert" cacti. However, for many of the
(Echinocereus J. more slow-growing and curious ornamental species,
In South America, one main centre of diversity is commercial exploitation is still regarded as a particu-
in the arid and semi-arid regions of SW Andean lar threat. For this reason the whole family Cactaceae
South America (Peru, Chile, Argentina) charac- is listed under Appendix II of the Convention of In-
terized by the tribes Trichocereeae, Notocacteae and ternational Trade in Endangered Species (CITES),
Browningieae. The other is E Brazil, where both the and some very vulnerable species (e. g. from the gen-
xerophilous (caatinga) vegetation and mountainous era Ariocarpus, Aztekium, Discocactus, Obregonia,
rocky vegetation (campo rupestre) have a rich cactus Pediocactus, Pelecyphora, Uebelmannia) are in-
flora, principally members of the tribe Cereeae. cluded in Appendix 1.
Finally, there are two centres of diversity for epi-
phytic cacti. The Hylocereeae are characteristic of AFFINITIES OF THE CACTACEAE. The position of Cacta-
the forests of Central America (with a few species ex- ceae within the betalain-containing Caryophyllales is
tending throughout tropical South America), whilst supported by all data and is no longer seriously ques-
the RhipsaJideae are most numerous in the Atlantic tioned. The family may have common ancestry with
rain forest zone of SE Brazil. As noted already, a few the Phytolaccaceae, together with Aizoaceae, Didi-
species of Rhipsalis occur throughout tropical ereaceae, and Portulacaceae. Ciadistically, it shares
America, and the most widespread, R. baccifera also several synapomorphies with the Portulacaceae, and
occurs naturally in polyploid populations throughout is possibly its sister group. No fossil records are
tropical Africa, Madagascar and Ceylon (Barthlott known, an earlier report now being discredited.
1983).
SUBDIVSION AND RELATIONSHIPS WITHIN THE FAMILY.
ECONOMIC USES. Several species (e. g. Opuntia ficus- All data suggest that the family is monophyletic. It is
indica, Selenicereus megalanthus (= "Mediocactus usually divided into three groups, which are not
coccineus"), Hylocereus spp., and Stenocereus spp.), linked by intermediate taxa. Following Schumann
are cultivated for their edible fruits. Spineless culti- (1897-99), they are here treated as the subfamilies
vars of Opuntia are grown as fodder plants. Agricul- Pereskioideae (two genera), Opuntioideae (five gen-
tural uses such as these are increasing, since these era), and Cactoideae (91 genera); alternatively these
succulent CAM plants may be grown under arid con- taxa may be classified as tribes (Britton and Rose
ditions with little or no irrigation. The local uses of 1919-23; Hunt 1967; Benson 1982). The Pereskioi-
cacti are many and varied: spines are used as fish- deae (as to the genus Pereskia itself) exhibit many an-
hooks or needles; Echinopsis pasacana has been an cestral features; the other two subfamilies are more
important source of timber where other wood is highly derived.
scarce; the peyote, Lophophora williamsii, which The following arrangement of the genera (see also
contains high concentrations of mescaline alkaloids, the diagram Fig.34) is based with minor modifica-
is an hallucinogenic drug; the formerly important red tions on the list recommended by a Working Party of
pigment, cochineal is produced from the insect Dac- the International Organization for Succulent Plant
tylopius coccus feeding on Opuntia spp. Study (lOS) (Hunt and Taylor 1990). As in other
Currently the most important economic aspect of "natural" families, generic delimitation is proble-
cacti, however, is their horticultural value. There are matical (many genera are weakly defined; intra-
a large number of specialized nurseries in Europe, generic hybridization is often possible), and a defini-
170 Cactaceae

CACTOIDEAE
PERESKIOIDEAE

Mammillaria

Fig. 34. Cactaceae. Presumed phylogenetic relationships within KEY TO THE SUBFAMILIES
the family, the size and arrangement of compartments reflecting
the number of species and relationships of the groups 1. Seed encased in a usually whitish or brownish. bony, tricho-
matous, alveolate or wing-like aril; areoles tufted with small,
minutely barbed spines (glochids) Opuntioideae (p.l71)
- Seeds black or brown, usually exarillate, rarely wholly or
tive suprageneric division of the principal subfamily partly enveloped by a mucilage sheath or the hilum with a
corky strophiolc or strophiolar pad; glochids abscnt 2
Cactoideae has not yet been achieved. Many prob- 2. Plants with functional leaves, at least on new growth
lems arise from the extent of neoteny and parallelism Pereskioideae (p.170)
(in vegetative and floral characters) in different - Plants virtually leafless (apart floral scales), though the stems
groups. The current tribal arrangement of the Cactoi- sometimes flattened, leaf-like Cactoideae (p. 172)
deae derives from the subtribes of Britton and Rose
(1919-23), considerably refined by the evolutionary
approach of Berger (1926) and more especially Bux- I. Subfam. Pereskioideae Schumann (1898)
baum (1958). The latter's extensive morphological
survey of the family, with particular attention to floral Tree-like, shrubby, caespitose or scandent; stems
and seed-testa characters, and the phylogenetic hypo- terete, not ribbed or tubercled; fertile zone inflores-
theses he based on his observations, have greatly in- cence-like or undifferentiated; leaves present; spines
fluenced modern cactology. Another major influence present, but glochids absent. Flowers solitary or in in-
on the taxonomy of the family has been the mono- florescences, diurnal; pericarpel with leaves or scales,
graph by the horticulturist Backeberg (Backeberg sometimes persistent; floral areoles with wool, often
1958-62), characterized by a very narrow concept of hairs, and rarely spines; tube absent; tepals free;
genera and species, and marred by failure to observe stamens numerous, inserted at base of perianth;
nomenclatural conventions. Backeberg's own classifi- ovary a cavity at the style-base or inferior, Fruits
cation, based mainly on geographic and phenetic berry-like to pear-like; pericarp juicy or tough, inde-
data, is largely unacceptable (see the survey of classi- hiscent; pulp present or absent. Seeds more or less
fications of the family in Barthlott 1988). circular, 1.7-7.5 mm, black-brown, shiny, not wrin-
 Cactaceae 171

kled, not keeled, not expanded around hilum; testa II. Subfam. Opuutioideae Schumann (1898)
cells uniform, slightly elongate, boundaries straight,
slightly channelled, interstices undifferentiated, relief Tree-like, shrubby or caespitose, rarely scandent;
flat to low-domed, microrelief none (occasionally stems usually segmented;! fertile zone undifferen-
weakly micropapillate); hilum-micropylar region tiated; leaves present, usually small, terete, de-
small, slightly oblique; hilum superficial; micropyle ciduous; glochids present, rarely concealed (Opuntia
disjunct; appendages none. clavarioides Pfeiffer); spines acicular, subulate or
This subfamily constitutes a small primitive group papery. Flowers lateral or rarely terminal, sessile,
combining the most plesiomorphic features in the usually solitary, diurnal; pericarpel stem-like, with
family. There are no transitional forms connecting it leaves, areoles, glochids and often spines; tube short
to the other subfamilies. The large seeds are of com- or absent (but the pericarpel often produced beyond
mon ancestral centrospermous type (resembling, for the ovary); stamens numerous, pollen mostly porate
instance those of Phytolaccaceae), and lack any apo- with reticulate tectum; ovary inferior. Fruits berry-
morphies for the Cactaceae as a whole. like to pear-like; pericarp fleshy, indehiscent, rarely
dry, dehiscent; pulp present, juicy or absent. Seeds
KEY TO THE GENERA OF PERESKIOIDEAE circular to broadly oval, 3-12 mm, covered by a bony
aril formed from the funicle, which envelops the en-
- Tree-like, shrubby or scandent; leaves broad and flat (trop.
America) 1. Pereskia
tire seed; aril brownish, whitish, light grey or pur-
- Caespitose shrubs; leaves terete (Andes of S Argentina, plish, surface bony, trichomatous, alveolate or almost
S Chile) 2. Maihuenia fleshy, winged in Pterocactus.
Seed architecture (including the aril, which is of a
type unique in the Centrospermae) and the mostly
1. Pereskia Miller porate and reticulate pollen emphasize the isolated
Pereskia Miller, Gard. Diet. Abr. ed.4, sine pag. (1754); Leuen- and derived position of this subfamily.
berger, Mem. N. Y. Bot. Gard. 41: 1-141 (1986).
Rhodocactus (A. Berger) F Knuth (1935).
KEY TO THE GENERA OF OPUNTIOIDEAE
Trees, shrubs and woody climbers, some with tube- 1. Leaves broad and flat 2
rous roots; stems not conspicuously succulent, terete, - Leaves terete 3
elongate; leaves broad, thin, deciduous or subpersis- 2. Flowers yellow, usually lateral; seeds 4-5 mm (Mexico,
tent; spines usually numerous. Inflorescence panicu- Guatemala) 3. Pereskiopsis
- Flowers red or pink, usually terminal; seeds 5-8 mm (S
late or corymbose, up to 50-flowered, or flowers clus-
America) 4. Quiabentia
tered or solitary; flowers stalked or sessile; pericarpel 3. Seeds not appreciably winged 4
areoles with wool, often hairs, and rarely spines; tube - Seeds winged all round 7. Pterocactus
none; perianth rotate, spreading or rarely erect, red, 4. Stems slender-cylindric, scrambling, cane-like or slightly flat-
orange, purplish, pink, yellow or white. Fruits some- tened, scarcely fleshy; pith becoming hollow or chambered;
times with persistent scales, sometimes proliferous. tepals and stamens separated by a ring of hairs (E Brazil)
5. Tacinga
Sixteen spp. in tropical America, from S Mexico and - Stems various, not cane-like. fleshy; pith not as above; tepals
the Caribbean region to N Argentina. For discussion and stamens not separated by a ring of hairs 6. Opuntia
of infrageneric classification, see Leuenberger (I.c.,
51-53).
3. Pereskiopsis Britton & Rose
Pereskiopsis Britton & Rose, Smithson. Misc. Collect. 50: 331
2. Maihuenia (Philippi ex F. A. C. Weber) Schumann (1907).
Maihuenia (Philippi ex FA. C. Weber) Schumann, Gesamt.
Kakt.: 754 (1898); Kiesling in Flora Patag6nica 5: 226-227 Sparsely branched or scrambling shrubs; stems terete,
(1988). unsegmented; leaves elliptic, obovate to spathulate
or almost circular, more or less succulent, deciduous
Low, caespitose shrubs, resembling Opuntia (Mai- to persistent; spines usually one to several, acicular.
hueniopsis) spp.; stems succulent, globose or short- Flowers usually lateral, Opuntia -like, verspertine
cylindric; leaves small, terete, persistent; spines
usually 3. Flowers terminal, solitary; perianth spread-
! In genera of all the subfamilies, stem-growth may be peri-
ing, yellow or white. Fruits obovoid or oblong, some-
odically constricted and articulate or segmented. The term
what fleshy; areoles of pericarpel and tube with small "segmented" is only used where the constrictions produce seg-
scales. Seeds almost circular, 3-5 x 2.5-4 mm. Two ments or "joints" of a consistent (i. e. genotypically determined)
spp. in Argentina and Chile. size.
172 Cactaceae

where known; areoles of pericarpel and tube with sometimes sheathed. Flowers lateral or subterminal;
leaves, areoles and glochids; ovary inferior; tube perianth rotate or spreading, rarely erect or slightly
none; perianth yellow, pink or red. Fruits fleshy, seeds zygomorphic (subg. Napalea), yellow, pink, red or
few. Seeds broadly oval, 4-5 mm; aril surface tricho- off-white; stamens sometimes touch-sensitive; style
matous. About nine spp. in Mexico and Guatemala. often more or less expanded near base. Fruits fleshy
or dry, umbilicate. Seeds circular to broadly oval, 3-
9 mm; aril surface bony, trichomatous or alveolate,
4. Quiabentia Britton & Rose
rarely almost fleshy.
Quiabentia Britton and Rose, The Cactaceae 4: 252 (1923). A large genus of> 200 species, occurring through-
out America, from S Canada to Patagonia. There is
Shrubby or tree-like; branches often verticillate, cy- no recent, conservative and comprehensive treat-
lindric-terete; leaves broadly ovate, obovate or spa- ment, though there are useful regional accounts for
thulate, to 7 em, flat, fleshy; areoles with glochids and several countries. The genus is divisible into
(0-)l-numerous acicular spines. Flowers subterminal numerous subgenera or sections corresponding to the
or terminal, diurnal; pericarpel with leaves, areoles, principal genera listed as synonyms. Various species
glochids and spines; tube none; perianth rotate, are grown for forage or edible fruits and some of
showy, red or pink. Fruits, where known, oblong, those introduced to Australia and South Africa have
terete, fleshy. Seeds 5-8 mm; aril surface trichoma- become noxious pests.
tous-fibrous. Two or more spp. in Brazil, Bolivia,
Paraguay, N Argentina.
7. Pterocactus Schumann
Pterocactus Schumann, Monatsschr. Kakt. 7: 6 (1897); Kiesling,
5. Tacinga Britton & Rose Cact. Succ. J. Gr. Brit. 44: 51-56 (1982).
Tacinga Britton and Rose, The Cactaceae 1: 39 (1919).
Dwarf, almost geophytic shrubs; rootstock usually
Weakly scandent shrubs, few branched; stems cane- large, tuberous; stem segments globose, cylindric or
like, terete or rarely compressed, not segmented, at clavate; spines few. Flowers terminal, immersed in the
first green and succulent, eventually woody, usually apex of stem segments, pale yellow or pink; perianth
with chambered pith; spines few or none. Flowers rotate; stamens touch-sensitive. Fruits dry, umbilicate,
usually terminal; tube thick, stem-like, persistent; te- dehiscent, splitting transversely near the top. Seeds
pals spreading to recurved, pale yellow, tinged green, samara-like, more or less circular, flat, 6-12 mm; aril
violet or brown, separated from the stamens by erect forming a broad wing surrounding the seed, pale
hairs; stamens and style erect, exserted. Fruits oblong. beige, papery. Nine closely related spp. in Argentina.
Seeds subglobose, 3-4 mm; aril present, enclosing en-
tire seed; aril white, surface bony. Two spp. in E Brazil.
III. Subfam. Cactoideae
6. Opuntia Miller
Tree-like, columnar, shrubby, caespitose, scandent,
Opuntia Miller, Gard. Diet. Abr. ed.4, sine pag. (1754). globular or epiphytic; roots usually fibrous, some-
Nopalea Salm-Dyck (1850). times tuberous; branching usually mesotonic, some-
Consolea Lemaire (1862).
times basitonic or acrotonic; stems usually unseg-
Tephrocactus Lemaire (1868).
Grusonia Schumann ex Britton & Rose (1919). mented, ribbed or tuberculate, at least in the juvenile
Maihueniopsis Spegazzini (1925). phase, depressed to globular or cylindric; fertile zone
Brasiliopuntia (Schumann) A. Berger (1926). undifferentiated or differentiated; leaves vestigial or
Corynopuntia E. Knuth (1935). lacking (except as scales on the pericarpel); glochids
Cylindropuntia (Engelmann) F.Knuth (1935). absent. Flowers sessile, diurnal or nocturnal; pericar-
Austrocylindropuntia Backeberg (1938).
pel usually with scales and/or areoles, or naked; tube
Platyop un tia (Engelmann) Ritter (1979).
Marenopuntia Backeberg (1950). very short to elongate, with few to numerous scales
Cumulopuntia Ritter (1980). and/or areoles; scales well-developed to rudimentary,
Puna Kiesling (1982). sometimes decurrent; floral areoles with wool and
spines, bristles, hair-spines or hairs, sometimes naked
Trees and shrubs, some low and caespitose; stems or undeveloped; tepals usually decurrent into the
usually segmented; stem-segments cylindric, clavate, tube; stamens usually very numerous, variously in-
subglobose, or more or less flattened, sometimes serted in the tube, sometimes with a separate stami-
tuberculate, very rarely ribbed; spines acicular, sub- nal throat-circle; ovary inferior. Fruit globose to ob-
ulate or papery, usually one to many, rarely none, long or ovoid, small to large; peri carp more or less
 Cactaceae 173

leathery, fleshy or dry, indehiscent or dehiscent; pulp - Peri carpel (at anthesis) nearly naked. or with scales only
present or absent. Seeds very diverse, oval to almost GroupE
6. Pericarpel with spines or bristles Group C
circular, 0.4-5.0 mm, black, black-brown or brown, - Pericarpel with hair-spines or hairs Group D
matt to glossy, not expanded or expanded around and 7. Stem ribbed. or if tubercled the tubercles disposed in more
rarely constricted above hilum; testa-cells uniform to or less vertical rows 8
abruptly smaller near hilum, isodiametric to elon- - Stem tubercled. the tubercles spirally disposed. or very
gate, boundaries straight to undulate, channelled, rarely both ribs and tubercles absent 10
raised or indistinct, interstices undifferentiated to 8. Flowers arising from a terminal cephalium Group G
- Cephalium not developed. although the stem-apex some-
minutely pitted or cratered, relief flat to convex, times densely woolly 9
rarely par-convex or par-concave, very rarely con- 9. Pericarpel and tube bearing areoles with wool, hairs. bristles
cave; microrelief none or micropapillate, or cuticle or spines Group H
weakly striate to strongly folded; hilum and micro- - Pericarpel and tube naked or with hairless scales only
pyle tending to be integrated in a single complex, Group J
sometimes appendaged with a mucilage sheath or a 10. Tubercles somewhat leaf-like or scale-like. arranged as if in
a roselle or cone Group K
strophiolar pad or strophiole; aril absent. - Tubercles not leaf-like, usually gibbous. or absent Group L
The largest and most complex subfamily. The inter-
stitially pitted or cratered seed-testa is probably
unique in the Angiospermae as is the extreme cuticu-
lar ornamentation ("loose cuticle") developed in Group A [Rhipsalideae, Hylocereeae
some genera of tribe Notocacteae. The tribal classifi- and Echinocereeae in part]
cation of the Cactoideae used in this treatment (see
diagram, Fig. 34) is developed from that of Buxbaum 1. Flowers usually less than 4 cm in diameter. tube very short
or none. rarely to 5 em. or the stem divided into short seg-
(1958), incorporating refinements proposed by the ments less than 8 cm 2
lOS Working Party (Hunt and Taylor 1986, 1990) and - Flowers large. usually more than 10 cm long. with a well-
by Barthlott (1988). The number of tribes (9) is the developed tube; stem not divided into short segments 8
same as in Buxbaum's classification, but the tribe 2. Flowers less than 2 cm long. creamy white, arising at the
Rhipsalideae (treated as Hylocereeae subtr. Rhipsa- edges of the stem-segments 3
linae by Buxbaum) is recognized here rather than - Flowers more than 2 cm long, brightly coloured (at least the
style). arising at or near the tips of the stem-segments 7
Buxbaum's Leptocereeae, the genera of which are 3. Stems slender-cylindric or angled 4
referred to other tribes. - Stems flat (2-winged), at least in part 5
4. Stem-segments mainly arising singly from the sides of older
segments, never in apical clusters 57. Lepismium
KEY TO GENERA OF CACTOIDEAE - Stem segments mainly arising at the apices of older seg-
ments. often in clusters. or main stems producing short lat-
1. Stem flat (leaf-like) or 2-winged, at least in part; or if 3-5- eral segments only 7-15 mm (R. mesembryanthoides)
ribbed or cylindric then less than 12 mm in diameter and not 58. Rhipsalis
continuously ribbed; usually spineless or with fine bristles 5. Adult stems all subsimilar. or if of 2 forms then the flat seg-
only Group A ments coarsely toothed (L. houlletianum); seeds less than
- Stems 3-4-winged or ribbed. or else 5 to many-ribbed or 1.5 mm. black or brown 6
tubercled 2 - Adult stems of 2 forms. the main axes terete at least basally.
2. Stems scrambling. trailing or pendent. slender. oftcn scg- the laterals flat. crenate; seeds usually 1.5-2 mm. black
mented. often emitting aerial roots. 3-4-winged or angled. or 18. Pseudorhipsalis
5-12-ribbed; spines usually weak and bristly or very short. or 6. Stem segments mainly arising singly from the sides of older
nonc Group B segments. never in apical clusters 57. Lepismium
- Stems erect or ascending. not scrambling. slender to very - Stem segments mainly arising at the apices of older seg-
stout. not producing aerial roots; 3-4-winged or angled. or ments. sometimes 2 or more together 58. Rhipsalis
few- to many-ribbed. or tubercled; spines often well de- 7. Flowers regular. tube 5 mm or less; lowermost stamens not
veloped 3 forming a ring round the style-base 59. Hatiora
4. Stem elongate. cylindric or columnar. 3-4-winged or angled, - Flowers more or less zygomorphic. tube 8 mm or more;
or ribbed, at least twice as long as thick before flowering; lowermost stamens forming a ring round the style-base
flowers nocturnal or diurnal 4 60. Schlumbergera
- Stem globular or depressed. ribbed or tubercled, less than 8. Flowers diurnal, pink or red
twice as long as tall (above ground) when flowering; flowers 17. Disocactus (Heliocereus, Nopalxochia)
mostly diurnal 7 - Flowers nocturnal. white 9
4. Flowering areoles similar to the non-flowering. or less spiny 9. Areoles of flattened stems spineless 16. Epiphyl/um
5 - Areoles of flattened stems spiny 10
- Flowering areoles different from the non-flowering. more 10. Epiphyte; stems clinging by aerial roots
hairy and/or bristly Group F 15. Selenicereus (Strophocactus)
5. Pericarpel with spines. bristles, hairs or felt; scales con- - Terrestrial shrubs; stems not producing aerial roots
spicuous to obsolete 6 10. Acanthocereus
174 Cactaceae

Group B [Hylocereeae in part, Echinocereeae] 62. Armatocereus

5. Low shrubs, usually < 1 m unless supported; stems < 6 cm
1. Flowers nocturnal, white, pale yellow or pale pink 2 diam. 6
- Flowers diurnal, yellow, orange, red, pink or purple 7 - Larger shrubs, or tree-like or columnar plants 11
2. Plants creeping or climbing, often emitting aerial roots 3 6. Offsets and buds bursting through the epidermis; stigmas
- Plants shrubby, prostrate or scrambling, not rooting green 12. Echinocereus
aerially (except occasionally at tip) but rootstock often - Offsets and buds not bursting through the epidermis;
tuberous 5 stigmas not bright green 7
3. Pericarpel with conspicuous triangular scales, enlarging in 7. Flowers narrowly tubular-funnelform 8
fruit; stems generally 3-winged and with very short spines - Flowers rotate-campanulate 9
14. Hylocereus 8. Flowers orange-red. Mexico (S Baja California)
- Pericarpel without conspicuous triangular scales; stems and 12. Echinocereus (E. pensilis)
spines various 4 - Flowers white (Brazil) 28. Leocereus
4. Flowers shortly funnelform, the tube less than 5 cm 9. Ribs 10-25; flowers yellow or white 10
13. Weberocereus - Ribs 5-12; flowers red or yellow
- Flowers elongate-funnelform, the tube more than 5 em (ex- 46. Corryocactus inc!. 47. Austrocactus
cept S. innesii) 15. Selenicereus 10. Ribs 20-25 (Mexico: Baja California; adjacent USA)
5. Roots with a napiform or Dahlia-like tuberous rootstock; 67. Bergerocactus
stem slender, often minutely pubescent, dark-coloured and - Ribs 10-13 (Peru) 43. Mila
striate or speckled, not rooting aerially, spines generally 11. Flowers campanulate, diurnal (Peru, Bolivia, Chile) 12
very short, appressed 11. Peniocereus - Flowers various, not campanulate, mostly nocturnal 13
- Roots not tuberous, or tubers not as above; stem various, 12. Areoles conspicuously woolly or hairy; flowers white or pale
not striate or speckled, sometimes rooting aerially; spines pink 48. Eulychnia
well-developed or short and weak 6 - Areoles not woolly or hairy; flowers brightly coloured
6. Stems sharply 3-5-winged or angled; tube stout, rigid, mar- 46. Corryocactus
kedly flared, with scattered scales felted and sometimes 13. Flowers narrowly tubular, bright red, 4--12 cm, with oblique
spiny but not hairy in their axils 10. Acanthocereus limb (NW Mexico) 73. Rathbunia
- Stems 3-4-angled or with 5-12 low ribs; tube relatively slen- - Flowers various, not bright red, regular 14
der, not rigid or markedly flared, with numerous scales hairy 14. Tube longer than pericarpel 15
in their axils 9. Harrisia - Tube very short 74. Polaskia
7. Stigmas green 12. Echinocereus (Wilcoxia) 15. Flowers tubular with narrow limb; ribs 4--6; staminal throat-
- Stigmas not green 8 circle present 34. Samaipaticereus
8. Flowers nearly rotate; tube and tepals very short - Flowers various; ribs (5-)6-30 or more; staminal throat-
8. Leptocereus (see also Corryocactus (Erdisia) spp.) circle absent 16
- Flowers funnelform; tube and tepals elongate 9 16. Flowering areoles confluent or discrete; flowers tubular,
9. Stems 3-5-winged, angled or ribbed with narrow limb; transition from periearpel to tube indis-
17. Disocactus (Heliocereus) tinct; areoles of pericarpel and tube with bristles and hairs
- Stems very slender, with 6-12 low ribs 10 68. Pachycereus
10. Rootstock tuberous, turnip- or Dahlia -like; flowers white or - Flowering areoles discrete; flowers tubular-funnelform, fun-
epidermis minutely papillate-hairy 11. Peniocereus nelform or campanulate; transition from pericarpel to tube
- Rootstock not tuberous; flowers coloured; epidermis usually abrupt; areoles of tube, except the lowest, without
smooth or pitted 11 bristles or hairs 17
11. Areoles < 1.4 cm apart on the ribs 12 17. Flowers tubular-funnelform; plants columnar, few-branch-
- Areoles 1.4--2.2 cm apart on the ribs ed; ribs 13-17 or more 70. Neobuxbaumia (N. tetetzo)
12. Echinocereus (E.pensilis) - Flowers funnelform or campa nul ate; plants shrubby or tree-
12. Stem segments elongate, > 30 cm; flowers ± zygomorphic like; ribs 4--13(-19 in S. thurberi) 72. Stenocereus
17. Disocactus (Aporocactus)
- Stem segments short, usually < 10 cm; flowers regular
40. Echinopsis (E. chamaecereus) Group D [Cereoid Trichocereeae, with Eulychnia in
part; no cephalium]
Group C [Cereoids with spiny or bristly pericarpel;
1. Flower-tube very short; ribs 9-17 48. Eulychnia
no cephaJium] - Flower-tube well-developed to elongate 2
1. Perianth concealed during bud-development; stem 3-4- 2. Stamens inserted in the tube in a single series
ribbed, light green 61. Calymmanthium 29. Haageocereus inc!. 30. Brachycereus
- Perianth visible during bud-development; stem various 2 - Stamens inserted in two series 3
2. Flowering areoles becoming elongate, proliferous, densely 3. Limb narrow; flowers nocturnal or diurnal, white or coloured 4
felted; ribs 4--8 64. Neoraimondia - Limb broad; flowers nocturnal, white 6
- Flowering areoles remaining cushion-like, not proliferous; 4. Flowers nocturnal, white 33. Facheiroa (Zehntnerella)
ribs 3-30 or more 3 - Flowers diurnal, mostly red (humming-bird flowers) 5
3. Stem segmented or with abrupt constrictions; perianth-limb 5. Areoles not clothed with long white hairs; fruit 1-4 cm in
ili~ 4 diameter, pulpy, indehiscent or bursting at one side
- Stem not segmented or constricted; perianth-limb various 5 35. Cleistocactus
4. Flowers small, 2-4 cm (Cuba, Hispaniola, Puerto Rico) - Areoles usually clothed with long white hairs (hair-spines);
8. Leptocereus fruit hollow, seeds escaping at base, or if pulpy then 4--5 cm in
- Flowers larger, 6-11 cm (Colombia, Ecuador, Peru) diameter 37.0reocereus
 Cactaceae 175

6. Stems often stout, not segmented; habit various (S America) 7. Cephalium composed of long or very dense bristle-spines 8
40. Echinopsis (see also Harrisia) - Cephalium composed of wool and hairs or bristles
- Stems slender-elongatc, 1-2 m x 2-4.5 cm, or shortly seg- 37. Oreocereus (0. doelzianlls)
mented (SE and W Brazil) 41. Arthrocereus (see also Leptocereus paniclliatus)
8. Ribs 5-15 68. Pachycereus
- Ribs 17-26 70. Neobuxbaumia (N. macrocephala)
Group E [Cereoids with scaly or naked pericarpel; 9. Fertile zone not clearly defined, but the flowers immersed in
no cephalium] long woolly hairs 22. Pilosocereus
- Fertile zone clearly defined by massive development of
1. Flowers very small, less than 2.5 cm, up to 9 per areole; matted wool and/or bristles 10
ribs 5-9 76. Myrtillocactus 10. Pericarpel and fruit scaly, the scales conspicuous or minute,
- Flowers larger, 2.5 cm or more, usually solitary; ribs 3-30 or hairy in their axils 11
more 2 - Pericarpel and fruit naked, or if with minute scales these
2. Pericarpel and tube more or less scaly 3 hairless in their axils 14
- Pericarpel and tube naked or nearly so 9 11. Flowers diurnal, brightly coloured
3. Scales chartaceous 75. Escontria 35. Cleistocactus spp. (see also Denmoza)
- Scales not chartaceous 4 - Flowers nocturnal, pale or dUll-coloured 12
4. Scales of tube distant, but the bases sometimes strongly de- 12. Flowers produced only on stems ca. 5 m; pericarpel and tube
current 5 with minute, almost hairless scales 71. Cephalocereus
- Scales of tube imbricate 8 - Flowers produced on stems 2 m or less; pericarpel and tube
5. Scales not decurrent; flowers funnelform or salverform, 4- with conspicuous scales or hairs 13
15cm 6 13. Stem masked by dense white spines and/or hairs; stem-tissue
- Scales strongly decurrent; flowers tubular to funnelforrn- not discolouring on exposure to air; flowers tubular-camp a-
campanulate, 2.5-8 cm 7 nul ate; scales not imbricate 31. Espostoa
6. Flowers salverform, 4-9 cm (Galapagos Islands) - Stem green, not masked by dense spines and/or hairs; stem-
63. Iasminocereus tissue turning brownish when exposed to air; flowers tubu-
- Flowers funnelform, 12-15 cm (Bolivia, Argentina) lar; scales imbricate 33. Facheiroa
66. Stetsonia 14. Fruit depressed-globose to broadly ovoid, depressed at the
7. Fruit pulp red, relatively juicy 69. Carnegiea point of attachment of the dried perianth, or laterally dehis-
- Fruit pulp white, not juicy 70. Neobuxbaumia cent, variously coloured 15
8. Flowers shortly tubular-campanulate; staminal throat -circle - Fruit obovoid-clavate to elliptic or globose, not depressed at
present (Brazil) 21. Brasilicereus apex, red, sometimes basally dehiscent, or < 15 mm dia-
- Flowers tubular or tubular-funneiform; staminal throat-circle meter 17
absent (Peru, Bolivia, N Chile) 65. Browningia 15. Flowering zone with more or less conspicuous hair-tufts, but
9. Fruits red or yellow, often slightly blue-waxy; stem-tissue not not forming a dense mat 16
darkening when cut 19. Cereus - Flowering zone forming a dense mat or fleece-like cepha-
- Fruits intensely blue-waxy; stem tissue darkening when cut lium on one side of the stem only 32. Espostoopsis
20. Cipocereus 16. Stems higly mucilaginous (C and S America)
22. Pilosocereus
- Stems lacking mucilage (NW Venezuela)
Group F [Cereoids with cephalium, 19. Cereus (c. mortensenii)
and Neoraimondia] 17. Fruit not expressed from the cephalium, indehiscent; peri-
anth-remnant tardily deciduous 25. Micranthocereus
1. Flowering areoles elongating when old, bearing flowers - Fruit expressed from the cephalium, opening by a basal
repeatedly 64. Neoraimondia pore; perianth-remnant persistent 26. Coleocephalocereus
- Flowering areoles not elongating, bearing flowers once only
2 Group G [Globular genera with terminal cephaJium]
2. Fertile zone discontinuous, alternating with vegetative
growth and marked by rings of bristles 3 1. Flowers diurnal, not fragrant, usually pink or red and less
- Fertile zone apical or lateral, continuous or discontinuous than 4 x 2.5 cm; fruit indehiscent 27. Melocactus
but not ringing the stem 5 - Flowers nocturnal, fragrant, white, 4-9 x 4-8 cm; fruit dehis-
3. Stems slender, constricted at each flowering zone 4 cent by lateral splits 45. Discocactus
- Stems columnar, not constricted at the flowering zones
71. Cephalocereus (Neodawsonia) Group H [Notocacteae, Trichocereeae and Cacteae
(see also Haageocereus zonatus)
in part; Echinocereus spp.]
4. Flowers> 3 cm diam., white with green tube; fruits blue-
green; ribs 13-18 23. Stephanocereus (S.leucostele) 1. Peri carpel and tube bristly or spiny; flower buds bursting
- Flowers < 3 cm diam., red, pink or yellow; fruits red, pink, through the stem-surface as they appear and/or stigmas
purple or brownish; ribs 7-14 24. Arrojadoa green and fruits juicy to fleshy 12. Echinocereus
5. Fertile zone covering the whole apex 6 - Peri carpel and tube not spiny, or the tube with some bristles;
- Fertile zone restricted to one side or discontinuous 9 flower buds not bursting through the stem surface; stigmas
6. Stem bottle-shaped, to 1.5 m, usually unbranched; flowers not green, or if green then fruits not juicy or fleshy 2
white within 23. Stephanocereus (Sluetzelburgii) 2. Flowers regular, with distinct stamina I throat -circle 3
- Stems not bottle-shaped; large shrubs or tree-like plants, or - Flowers regular or bilaterally symmetric, lacking a staminal
if less than < 40 cm, flowers red 7 th roa t -circle 5
176 Cactaceae

3. Flowers mostly lateral, more than 4 cm, variously coloured Group J [N otocacteae, Trichocereeae and Cacteae
40. Echinopsis in part]
- Flowers apical to subapical, 2.5-4( -{j) cm, yellow to reddish
orange 4 1. Stem ridged and wrinkled between the ribs; seeds minute,
4. Stems globose or somewhat elongate, simple or sparsely 0.5-D.7 mm, strophiolate 91. Aztekium
branched, not densely spine 42. Rebutia - Stem and seeds not as above 2
- Stems cylindric, freely branched, covered with bristly spines 2. Plant spineless, areoles tufted with woolly hairs only
43. Mila 88. Lophophora
5. Flowers at the shoulder of the stem or below 42. Rebutia - Plant more or less spiny 3
- Flowers near the stem-apex 6 3. Flowers regular 4
6. Scales of flower-tube (at least the uppermost) with wool and - Flowers bilaterally symmetric, the tube curved
bristles in the axils 7 38. Matucana
- Scales of tube only hairy or woolly in the axils 11 4. Fruit dry, opening at apex, the base of the dried perianth
7. Fruit woolly; apical scales spinescent 51. Eriosyce detaching like a lid; flowers yellow with very short tube
- Fruit only sparsely woolly, or glabrous; apical scales not spi- 49. Copiapoa
~=m 8 - Fruit dry or juicy, opening basally or laterally or disintegrat-
8. Fruit clavate to oblong, expanding and becoming partly hol- ing irregularly, or indehiscent 5
low at maturity, opening at the base; seeds with relatively 5. Ribs very thin, more or less wavy, or if few then the areoles
small hilum and no strophiole, the seedcoat often folded with 1-3 large, flattened spines above and 1-2 much smaller
and ridged 52. Neoporteria terete spines below 95. Stenocactus
- Fruit globose to ovoid or cylindric, indehiscent, splitting ver- - Ribs not thin and wavy; spines not as above 6
tically or disintegrating irregularly; or if clavate and ex- 6. Fruit fleshy, usually bursting laterally; ribs usually chinned
panding then the seeds with a broad hilum (equalling the beneath the areoles; areolar glands absent
diameter of the seed) and strophiole 9 44. Gymnocalycium
9. Stem dark reddish brown with surface roughened and some- - Fruit dry, juicy but indehiscent; ribs not "chinned"; areolar
times obscured by waxy whitish scales; large mucilage glands often present 7
bodies present in the cortex or with conspicuous mucilage 7. Mature stem large, more than 20 cm in diameter, or smaller
ducts (Brazil: Minas Gerais) 56. Uebelmannia and the seeds pitted; ribs well defined 93. Ferocactus
- Stem not as above; epidermis green or stem < 4 cm diam., - Mature stem 4-20 cm in diameter; seeds smooth, reticulate or
fruit not usually red 10 tuberculate; ribs poorly defined, strongly tubercled 8
10. Spines never hooked; flowers either cleistogamous or more 8. Spines obscuring the stem, or the central spines strongly re-
than half the diameter of the stem; seeds with depressed curved to hooked; flowers never yellow with red throat;
hilum and no strophiole 55. Frailea stigmas sometimes green; hilum lateral or obligue to main
- Spines hooked or not; flowers never cleistogamous, less axis of seed 80. Sclerocactus
than half the diameter of the stem; seeds not as above - Spines not as above, or flowers yellow with red throat;
53.Parodia stigmas never green; hilum terminating long-axis of seed
11. Scales of pericarpel and tube, and apex of outermost tepals, 82. Thelocactus
narrow, sharply pointed, light to dark brown; flowers
various, regular, not tubular nor scarlet 12
- Scales and outermost tepals not as above; flowers various, Group K [Cacteae in part; stems with
sometimes bilaterally symmetric and/or scarlet 15 leaf- or scale-like tubercles]
12. Stem-surface more or less spotted or covered with small
1. Tubercles vcry long (to 15 em) and thin, 3-angled, with long
tufts of whitish trichomes; seeds cap-shaped, with deeply
papery spines 94. Leuchtenbergia
sunken hilum; flowers yellow, sometimes red in the throat
- Tubercles various, not as above, spineless or the spines cadu-
79. Astrophytum
cous or bristle-like 2
- Stem-surface and seeds not as above; flowers various 13
2. Tubercles rather thin, triangular, scale-like, closely overlap-
13. Plant nearly spineless, except near apex; flowers purplish
ping; spines conspicuous 3
pink; pericarpel and lower part of tube naked; outer tepals
- Tubercles thick, triquetrous and subulate or ± pyramidal,
not aristate but with dense trichomes in the acils
not closely overlapping; spines inconspicuous or absent
78. Geohintonia
87. Ariocarpus
- Plants usually very spiny; flowers yellow or pink to nearly
3. Flowers purplish pink; stamens not sensitive; tubercles ap-
white; pericarpel with scattered scales, these and the outer
pressed throughout, as in a bulb
tepals aristate or spinose-tipped 77. Echinocactus
91. Pelecyphora (P. strobiliformis)
14. Scales of pericarpel and tube, and apex of outermost tepa Is,
- Flowers pale yellow; stamens sensitive; tubercles, except the
aristate or spinose-tipped
youngest, spreading 89. Obregonia
40. Echinospis (E.leucantha, E. spiniflora)
- Scales and outcrmost tepals not as above 15
15. Flowers tubular, the limb not expanded, the stamens and Group L [N otocacteae, Trichocereae and Cacteae
style protruding; fruit splitting laterally when ripe, fleshy in-
in part]
side 36. Denmoza
- Flowers not as above; fruit opening at or splitting from the 1. Stems button-like, ca. 1 cm diam., without ribs or tubercles
base, dry inside 16 54. Blossfeldia
16. Flowers regular; fruit opening at base 39.0roya - Stems tuberculate 2
- Flowers usually bilaterally symmetric; fruit splitting laterally 2. Areoles situated between the tubercles; tubercle apex
or from the base 38. Matucana naked 50. Neowerdermannia
 Cactaceae 177

- Areoles situated at the tubercle apex, sometimes extending - Seed coat cells tabular-concave or par-concave (pitted);
via a groove to the "axil" above, or bi-partite (with a dis- outer tepals fringed; areolar glands absent 96. Escobaria
crete axillary portion) 3 18. Tubercles low, pale grey-green, punctate; flowers 2-3 cm,
3. Flowers arising at the tips of the tubercles, or in short groove yellow, with deep green stigmas 97. Ortegocactus
towards the "axil" 4 - Not as above 19
- Flowers arising in the tubercular "axils" 18 19. Flowering areoles grooved or ridged between the sterile
4. Flowers borne at or near stem-apex 5 apical and fertile axillary portions 20
- Flowers borne on the "shoulder" of the stem or below - Flowering areoles bi-partite, the apical and axillary portions
42. Rebutia discrete 22
5. Pericarpel and fruit bearing scales, wool and sometimes 20. Tubercles hatchet-shaped; spines pectinate
bristles; fruit dry inside, indehiscent, disintegrating irregu- 92. Pelecyphora (P aselliformis)
larly or partly hollow 6 - Tubercles cylindric, conic or gibbous; spines various 21
- Pericarpel and fruit naked or bearing hairless scales only; 21. Seedcoat cells tabular; outer tepals entire, or areoles with
fruit juicy, indehiscent, or dry, opening at base, apex or lat- glands 83. Coryphantha
erally 8 - Seed coat cells tabular-concave or par-concave (pitted);
6. Flowers yellow, but often cleistogamous, the perianth then outer tepals fringed; areolar glands absent 96. Escobaria
scarcely developing; seeds 1-3 mm, shiny, with sunken 22. Seeds black; cells of seedcoat tabular or somewhat convex
hilum 55. Fraileas 85. Mammilloydia
- Flowers yellow, red, pink or whitish, never cleistogamous; - Seeds black or brown; if black, the seedcoat cells par-con-
seeds ca. 0.5-1.2 mm, shiny or dull, the hilum not sunken 7 cave (pitted) 98. Mammillaria
7. Fruit more or less globose, 5-10 mm; seeds 0.5-1 mm, stro-
phiolate 53. Parodia
- Fruit clavate, 10-25 mm; seeds 0.8-1.9 mm, not strophiolate 1. Tribe Echinocereeae (Britton & Rose)
52. Neoporteria
E Buxbaum (1958) (incl. Leptocereeae EBuxbaum)
8. Fruit opening near apex and sometimes splitting longitudi-
nally also, the dried perianth-base raised like a lid 9
- Fruit indehiscent or opening laterally or below, the dried Tree-like or shrubby, sometimes scandent, rarely
perianth-base or its sear remaining firmly attached to the small, globular; roots often tuberous; stems seg-
peri carp on all sides 10 mented or not, elongate to globose, ribbed; fertile
9. Fruit splitting laterally; roots fibrous; flowers pink, white or zone undifferentiated (except in Leptocereus panicu-
yellow; seeds 1.5-5 mm (USA) 81. Pediocactus latus). Flowers usually lateral, rarely subapical or ter-
- Fruit cup-like; roots tuberous; flowers yellow; seeds 0.8-
2.1 mm (Chile) 49. Copiapoa
minal, small to large, nocturnal, white, or small to
10. Pericarpel and fruit always visible, the latter dry, opening medium-sized, diurnal, usually brightly coloured;
laterally or basally, dull green or brownish 11 areoles of pericarpel and tube with spines, bristles or
- Pericarpel hidden between the tubercles or by spines and hair-spines. Fruit usually berry-like, juicy, sometimes
wool at flowering time; or fruit juicy, indehiscent, bright splitting longitudinally, rarely almost dry. Seeds
green, pink or red, or dry and remaining hidden when ripe
medium-sized to large, periphery undifferentiated to
14
11. Spines flattened, resembling dried grass leaves; stem cla-
keeled or crested with larger cells; cell-boundaries
vate-cylindric; stigmas green 80. Sclerocactus straight, interstices usually undifferentiated, rarely
- Spines not as above or stem depressed-globose; stigmas not pitted or cratered; hilum and micropyle conjunct or
green 12 fused; appendages none. Distribution: mostly Carib-
12. Pericarpel and fruit naked, rarely with 1 or 2 scales; flowers bean region, Mexico, SW United States.
to 3.5 x 4 cm; seeds 1-2 mm, black, not strophiolate
84. Neolloydia
- Pericarpel and fruit with 2 or more scales; flowers to 8. Leptocereus (A. Berger) Britton & Rose
6 x 8 cm or seeds only 0.5 mm, brown, strophiolate 13
13. Fruit opening by a basal pore; seeds 1.5-2.5 mm, black, not Leptocereus (A. Berger) Britton and Rose, Contrib. U.S. Natl.
strophiolate; spines strong or dense 82. Thelocactus Herb. 12: 433 (1909).
- Fruit opening by 1-3 vertical splits; seeds 0.5 mm, brown, Neoabbottia Britton & Rose (1921).
strophiolate; spines few, weak, caducous
90. Strombocactus Trees and shrubs, sometimes prostrate or scandent, to
14. Plant spineless, areoles tufted with woolly hairs only
8-10 m, much branched; stems usually segmented,
88. Lophophora
- Plant spiny 15 ribbed; ribs 3-8. Flowers tubular-campanulate, diur-
15. Tubercles conspicuous, more than 1 mm; areolar groove nal or nocturnal, in one species clustered at the felted
short or extending to the base 16 apex of terminal segments, 2-5 x 1.5-3 em, almost
- Tubercles minute, ca. 1 mm, not grooved 86. Epithelantha white or pale green, yellow or pink; areoles of pericar-
16. Fruit juicy, often brightly coloured; seedcoat smooth or pel and tube spiny to nearly naked; tube short; peri-
pitted, black or brown 17
anth short, spreading or rotate. Fruits globose to ob-
- Fruit dry, dull-coloured; seeds tuberculate, black
84. Neolloydia long, 1.5-10 x 1.5-6 cm, fleshy, spiny to nearly naked.
17. Seedcoat cells tabular; outer tepa Is entire, or areoles with Seeds oval or broadly oval, rugose or ruminate. Up to
glands 85. Coryphantha 12 spp., Cuba, Hispaniola and Puerto Rico.
178 Cactaceae

9. Harrisia Britton nocturnal or diurnal, occasionally terminating the

Harrisia Britton, BulL Torrey Bot, Club 35: 561 (1908), stem; tube long and slender; areoles of peri carpel and
Eriocereus (A. Berger) Riccobono (1909), tube with bristles or spines. Fruits narrowly ovoid,
Roseocereus Backeberg (1938), tapered at apex, fleshy red, the spines or bristles more
or less deciduous. Seeds broadly oval, ruminate or
Shrubs, some tree-like, to 7 m, or scandent; roots not. About 20 spp., C America, S to NW Mexico and
sometimes tuberous; stems usually slender, ribbed, the SWUSA.
not segmented, not rooting aerially; ribs 4-12,
Flowers nocturnal, funnelform, 12-22 x ca, 8-12 cm,
white; areoles of pericarpel and tube with hair-spines 12. Echinocereus Engelmann
or merely felted; tube elongate; perianth-limb broad, Echinocereus Engelmann in Wislizenus, Mem. Tour N Mexico,
Fruits areolate and scaly and/or spiny, fleshy, yellow 91 (1848); Taylor, The genus Echinocereus (1985), et in Brad-
or orange, not splitting (subg, Harrisia), or red, ley a 6: 65-84 (1988), ibid. 7: 73-77 (1989).
usually splitting (subg, Eriocereus), Seeds broadly Wilcoxia Britton & Rose (1909), as to type.
Morangaya Rowley (1974).
oval, periphery crested with larger cells; hilum deeply
impressed, forming a chamber, About 20 spp" Carib-
bean region to Argentina, The genus consists of two Low-growing shrubs; roots fibrous or tuberous; stems
well-marked and disjunct subgenera distinguished by simple or clustering, self-supporting, procumbent or
their fruits but united by the unique seed-type, Subg, rarely climbing, globose to cylindric, ribbed; lateral
Harrisia occurs in SE USA (Florida) and the Carib- shoots usually erumpent (bursting through the
bean region, subg. Eriocereus in Brazil, Bolivia, Para- epidermis above an areole). Flower buds developing
guay and Argentina. at the upper edge of the stem areoles or erumpent;
flowers funnelform, small to rather large; areoles of
pericarpel and tube with spines, bristles and some-
10. Acanthocereus (A. Berger) Britton & Rose times abundant wool; stigmas greenish, rarely white.
Acanthocereus (A. Berger) Britton and Rose, Contrib. U.S. Fruits dehiscent, or indehiscent, juicy, fleshy or dry,
Nat!. Herb. 12: 432 (1909); Hunt, Bradleya 9: 82-83 (1991). bearing spiny areoles or these falling off at maturity.
Dendrocereus Britton & Rose (1920). Seeds broadly oval, sometimes ruminate. A genus of
Monvillea Britton & Rose (1920), as to type only. 47 spp., Mexico and SW USA. The genus is divided
? Pseudoacanthocereus Ritter (1979), non Peniocereus subg.
by Taylor (I. c. 1985) into seven sections according to
Pseudoacanthocereus Sanchez-Mejorada (1974).
habit, flower and fruit characters.
Mostly shrubs or weakly scandent, one tree-like with
trunk to 60 cm diam.; stems segmented or not, some-
times rooting at the tips, 3-5-ribbed or rarely flat- 2. Tribe Hylocereeae (Britton & Rose)
tened. Flowers funnelform, 12-25 x ca. 6-12 cm, noc- F. Buxbaum (1958)
turnal, white; areoles of peri carpel and tube with felt
and usually short stiff spines; tube elongate, typically Scandent or epiphytic shrubs; roots mostly adventi-
stiff, erect; perianth broad. Fruit globose or ovoid, 4- tious; stems few-ribbed or flattened; fertile zone un-
8 cm, red or green, spiny, splitting when ripe, in some differentiated. Flowers lateral, medium-sized to very
spp. 8-12 cm, green, naked, indehiscent. Seeds large, nocturnal, white, or diurnal, red or pink, rarely
broadly oval, up 4.8 mm. The genus currently in- small to very small, diurnal, almost white (Pseudo-
cludes six or more species in tropical America and the rhipsalis); areoles of pericarpel and tube with spines,
Caribbean region but is heterogeneous. bristles, hair-spines or naked. Fruits juicy, indehis-
cent. Seeds medium-sized to large, border not ex-
panded; boundaries straight, interstices undifferen-
11. Peniocereus (A. Berger) Britton & Rose
tiated to minutely pitted or rarely cratered
Peniocereus (A. Berger) Britton and Rose, Contrib. U.S. Nat!. (Epiphyllum); hilum and micropyle fused, often
Herb. 12: 428 (1909). band-like; mucilage sheath present, covering entire
Nyctocereus (A. Berger) Britton & Rose (1909).
seed. Distribution: mainly forests of Central
Wilcoxia Britton & Rose (1909), in part.
Neoevansia W.Marshall (1941). America, a few species extending throughout tropical
Cullmannia Distefano (1956). America. The affinities of the Hylocereeae are
probably with the Echinocereeae (Harrisia, Acan-
Prostrate or scandent shrubs; roots thickened, turnip- thocereus) and Cereeae (Cereus); the similarity of
like or Dahlia-like; stems slender, ribbed, sparingly Pseudorhipsalis to Rhipsalis is superficial and due to
branched; epidermis sometimes papillose-downy; parallelism. The genera of Hylocereeae are weakly
spines conspicuous, or appressed and short. Flowers defined, and intergeneric hybrids are numerous.
 Cactaceae 179

13. Weberocereus Britton & Rose 16. Epiphyllum Haworth

Weberocereus Britton & Rose, Contrib. U. S. Nat!. Herb. 12: Epiphyllum Haworth, Syn. P!. Succ. 197 (1812).
431 (1909). Phyllocactus Link (1831).
Werckleocereus Britton & Rose (1909).
Eccremocactus Britton & Rose (1913). Epiphytic or rarely epilithic shrubs; stems initially
terete, mostly becoming 2-ribbed, leaf-like, margins
Epiphytic or epilithic shrubs; stems terete, angled or crenate, serrate, lobed orpinnatisect between the are-
flattened, sometimes rooting aerially; ribs 2-5; spines oles; spines absent, or present on terete stems only.
short, bristly or absent. Flowers nocturnal, funnel- Flowers funnelform or salverform, mostly nocturnal,
form, 3-10 cm; areoles of pericarpel and tube with 10-30 cm; tube elongate; scales small, sparse; areoles
bristles or hair-spines; tepals pink, yellow-white or of pericarpel and tube naked or rarely with hair-spines
green-tinged; staminal throat-circle present. Fruits or bristles; outer tepals white, pale yellow or pink-
globose to oblong, red or yellow, fleshy, bristly or tinged; inner tepals pale yellow or white. Fruits ovoid
naked; pulp white or purple. A genus of nine spp., or oblong, 4-9 x 2-5 cm, often somewhat ridged,
S Mexico, C America (mainly Costa Rica), Panama spineless. About 15 spp., mainly in C America, a few
and Ecuador. species extending to the West Indies and S America.

14. Hylocereus (A. Berger) Britton & Rose

17. Disocactus Lindley
Hylocereus (ABerger) Britton & Rose, Contrib. U. S. Nat!.
Disocactus Lindley, Bot. Reg. 31: t. 9 (1845); Barthlott in Brad-
Herb. 12: 428 (1909).
ley a 9: 86-88 (1991).
Wilmattea Britton & Rose (1920).
Aporocactus Lemaire (1860); Hunt in Bradleya 7: 93-96 (1989).
Wittia Schumann (1903), non Pantocsek.
Scandent or epiphytic shrubs; stems often to 5 m or Heliocereus (A Berger) Britton & Rose (1909).
more, usually 3-winged or angled, segmented, green Chiapasia Britton & Rose (1923).
or glaucous, the margins often horny, often producing Nopalxochia Britton & Rose (1923).
aerial roots; areoles with short spines or rarely spine- Bonifazia Standley & Steyermark (1944).
less. Flowers usually very large, funnelform, noctur- Lobeira Alexander (1944).
Pseudonopalxochia Backeberg (1958).
nal, white or rarely red; pericarpel and tube stout; Wittiocactus Rauschert (1982).
scales usually broad, triangular; areoles of pericarpel
and tube usually naked or nearly so; stamens in a con- Epiphytic or epilithic shrubs; stems ribbed or flat-
tinuous series; style thick, stigmas sometimes bifid. tened and leaf-like, or the main stems terete at base,
Fruits large, globose, ovoid or oblong, fleshy, with flattened above, and the lateral flattened; areoles
broad scales. A genus of ca. 16 spp., C America, the proliferous in some species; spines bristly or absent.
West Indies and N South America. Flowers usually 1 per areole, diverse in size and form,
diurnal, red (occasionally tinged blue), pink, orange,
15. Selenicereus (A. Berger) Britton & Rose rarely pale yellow or white; stamens usually in two
series. Fruit berry-like, with few small scales or al-
Selenicereus (A Berger) Britton & Rose. Contrib. U.S. Nat!.
Herb. 12: 429 (1909); Hunt, in Bradleya 7: 89-92 (1989). most naked. Sixteen spp., mainly in C America, with
Strophocactus Britton & Rose (1913). two species in the West Indies and one extending to N
Deamia Britton & Rose (1920). South America. Barthlott (l. c.) recognises five sub-
Mediocactus Britton & Rose (1920), except the type. genera.
Cryptocereus Alexander (1950).
Marniera Backeberg (1950).
18. Pseudorhipsalis Britton & Rose
Scandent, epilithic or epiphytic shrubs; stems often to Pseudorhipsalis Britton & Rose, The Cactaceae 4: 213-214
5 m or more, slender, often bearing aerial roots; ribs (1923); Barthlott, Bradleya 9: 90 (1991).
2-12; spines short, bristly or hair-like, rarely acicular
or absent. Flowers funnelform, rarely salverform, Epiphytes; stems initially terete, later flattened, leaf-
medium-sized to very large, 12-40 x 10--20 cm, noc- like, spineless. Flowers short-funnelform or rotate,
turnal; areoles of pericarpel and tube with hair-spines, small, 0.7-2 cm (up to 3.5 cm in P. himantoclada),
bristles or spines; tepals white (red-tinged in S. werck- white or yellowish white, diurnal; tube short, 2-
lei), or the outer yellow, pink or brown; staminal 10 mm; stamens in one series. Fruit berry-like, ovoid
throat-circle present. Fruits globose, ovoid or oblong, or globular, ca. 1 cm, whitish, often tinged purplish,
6-8 cm, fleshy, usually red, spination persistent. more or less naked. Up to five spp., C America and
About 20 spp., tropical America and the Caribbean the Caribbean region, one extending throughout
region. Hunt (l. c. 1989) recognises five sections. tropical America.
180 Cactaceae

3. Tribe Cereeae Salm-Dyck (1840) modified or more woolly than the non-flowering;
(as "Cereastreae") flowers tubular, nocturnal or partly diurnal; pericar-
pel and tube ribbed or nearly terete, thickly blue-
Tree-like or shrubby, rarely scandent, rarely globular; waxy and with small, widely spaced scales naked or
roots rarely tuberous; stems usually unsegmented, subtending small areoles, wool and sometimes fine
elongate to globose, ribbed, spiny; fertile zone usually spines; tepals short, usually white, the outermost dark
differentiated, forming lateral, apical or terminal ce- blue-waxy, pale green or yellow. Fruit indehiscent,
phalia. Flowers lateral or subapical, regular or almost globose to ovoid, intensely blue-waxy; floral remnant
so, small to large, nocturnal, white, or diurnal, red, persistent, erect; pulp semi-liquid, translucent. Five
pink or rarely yellow; pericarpel and tube usually spp., endemic to Brazil (Minas Gerais).
naked except for scales. Fruit berry-like, juicy, inde-
hiscent or variously dehiscent, rarely almost dry; floral
21. Brasilicereus Backeberg
remnant often persistent and/or blackening. Seeds
small to large, oval, without conspicuous surface Brasilicereus Backeberg, Blatter f. Kakteenforschung 1938 [8,
sculpturing; cell boundaries straight, interstices undif- 22] (1938).
ferentiated or minutely pitted, rarely cratered towards
periphery; hilum and micropyle conjunct, rarely al- Shrubs; stems cylindric, rather woody; ribs 8-14; are-
most disjunct; appendages none. Distribution: Mainly oles small, close-set; spines acicular. Flowers noctur-
E South America, Melocactus extending to Peru, nal, shortly tubular-campanulate; tube short, stout;
Mexico and the Caribbean region. Apparently a scales conspicuous, fleshy; areoles of pericarpel and
derived group, perhaps related to tribe Trichocereeae. tube naked; tepals widely spreading; staminal throat-
A cladistic treatment of the genera has been given by circle present. Fruit indehiscent, ovoid, green to pur-
Taylor & Zappi, Bradleya 7: 13-40 (1989). plish; floral remnant persistent; pulp whitish. Two
spp., endemic to E Brazil.
19. Cereus Miller
Cereus Miller, Gard. Diet. Abr. ed.4, sine pag. (1754). 22. Pilosocereus Byles & Rowley
Monvillea Britton & Rose (1920), except the type. Pilocereus Schumann (1894), non Lemaire.
Subpilocereus Backeberg (1938). Pilosocereus Byles & Rowley, Cact. Succ. 1. Gr. Brit. 19: 66
Praecereus Buxbaum (1968). (1957).
Mirabella Ritter (1979). Pseudopilocereus F.Buxbaum (1968).

Tree-like or shrubby, usually much branched; stems Tree-like or shrubby, usually more or less branched
erect or ascending, strongly ribbed, sometimes with from the base or trunk, to 10 m; ribs (3-)4-30, often
annual constrictions; often glaucous; ribs 3-14, usually cross-furrowed; areoles, at least the flowering, with
pronounced; spines often numerous, acicular. Fertile more or less abundant woolly hairs (but lacking in
zone undifferentiated or the flowering areoles more or a few well-known spp.), sometimes as long as 5 cm
less spineless; flowers medium-size to large, funnel- and forming skeins covering the ribs. Flowers tubu-
form, nocturnal, usually white; pericarpel and tube lar-campanulate, 4-10 cm, nocturnal, chiropterophi-
elongate, thick, naked or nearly so below and with lous; pericarpel and tube fleshy, naked (or with
scattered small scales above; perianth-limb broad or minute scales), often tinged brown or purple; limb
moderately so. Fruit fleshy, globose, ovoid or oblong, rather narrow, usually white or pale pink; stamens
usually red, sometimes yellow, naked, splitting along very numerous. Fruits dehiscent, ovoid to depressed-
one side when mature to reveal white or rarely pink or globose, smooth, fig-like, usually with red pulp, the
red pulp; floral remnant, or at least the style, usually perianth persistent, blackened, deflexed. Some
persistent. About 35 species in the West Indies and 35 spp., Mexico, the Caribbean region, and trap.
S America. The genera listed as synonyms are recog- S America (especially Brazil).
nizable as subgenera or sections, though, differen-
tiated by vegetative, floral and fruit characters.
23. Stephanocereus A. Berger
20. Cipocereus Ritter Stephanocereus Berger, Entwicklungslinien Kakt.; 97 (1926).
Cipocereus Ritter, Kakteen in Siidamerika 1: 54 (1979); Zappi
& Taylor in Bradleya 9: 86 (1991). One species tree-like or sparsely branched, 1-6 m
Floribunda Ritter, Kakteen in Siidamerika 1: 58 (1979). (5. leucostele), the other unbranched (S. luetzelbur-
gii); stems cylindric or ovoid; ribs 12-20, low. Flowers
Shrubs; stems cylindric, rather woody; ribs 4-21; in ring-like cephalia, alternating with vegetative
spines absent to numerous. Flowering areoles un- growth (S. leucostele) or apical (5. luetzelburgii);
 Cactaceae 181

flowering areoles with dense bristles (to 7 cm in S. confluent and bearded with wool and bristles, form-
leucostele ); flowers tubular-funnelform, 5-10 cm; ing an apical and lateral cephalium on several re-
scales of pericarpel and tube few, scattered, minute, duced and depressed ribs. Flowers tubular to campa-
areoles naked or with very sparse wool; tepals short, nulate-funnelform, to 6 cm, nocturnal; pericarpel and
spreading, white. Fruit ovoid-ellipsoid, indehiscent; tube naked. Fruit berry-like, obovoid-clavate or glo-
floral remnant blackening, persistent; pericarp thick, bose, red, opening by a small basal pore. Six spp.,
blue- to purple-green, naked; pulp white or red. Two E and SE Brazil.
spp., endemic to E Brazil (Bahia).
27. Melocactus Link & Otto
24. Arrojadoa Britton & Rose
Melocactus Link & Otto, Verh. Ver. Befbrd. Gartenb. Preuss.
Arrojadoa Britton & Rose, The Cactaceae 2: 170 (1920). Staaten 3: 417 (1827); Taylor in Bradleya 9: 1-80 (1991).
Cactus sensu Britton & Rose (1922).
Low shrubs, few-branched; stems cylindric or slen-
der-cylindric, sometimes segmented; ribs 7-17; are- Unbranched (unless demaged); stem depressed glo-
oles close-set; spines small or bristly. Flowering are- bose to short-columnar, rarely more than 1 m high,
oles different from the non-flowering, tufted with strongly ribbed, spiny. Flowering zone a terminal ce-
bristles and forming intermittent ring-like apical ce- phalium with wool and usually bristles; flowers small,
phalia through which vegetative growth can continue tubular, red to pink, more or less immersed in the
after flowering, leaving a collar of bristles. Flowers cephalium; pericarpel and tube naked. Fruit a juicy
small, often numerous, tubular, diurnal, red, pink, berry, usually clavate, red, pink or white. A genus of
violet or yellow; pericarpel and tube naked; limb very 31 species in tropical America, especially E Brazil
short, erect or spreading; stamens and style included. and Amazonia, Peru, Venezuela, Central America
Fruit small, berry-like, juicy. Three spp., E Brazil. and the Caribbean.
True stem-tubers occur in A. dinae.

4. Tribe Trichocereeae F. Buxbaum (1958)

25. Micranthocereus Backeberg
Micranthocereus Backeberg, Blatter f. Kakteenforschung 1938 Tree-like, columnar, shrubby or globular, rootstock
(6): [22] (1938).
sometimes thickened; stems usually unsegmented,
Austrocephalocereus Backeberg (1938), in part (inc!. type).
Siccobaccatus Braun & Esteves Pereira (1990).
usually ribbed, sometimes tuberculate-ribbed or sim-
ply tuberculate; fertile zone undifferentiated or dif-
Shrubby to tall columnar, unbranched or branching ferentiated as lateral (Espostoa, Espostoopsis, Fa-
from the base; stems cylindric, ribbed; ribs 10-30 or cheiroa) or terminal (Discocactus) cephalia. Flowers
more, narrow; areoles close-set, with spines and lateral to subapical, small to very large, nocturnal,
sometimes long wool. Fertile zone a continuous or white, or small to large diurnal, brightly coloured,
discontinuous, superficial or sunken, woolly/bristly regular or more or less zygomorphic; pericarpel and
cephalium; flowers from the cephalium, tubular, 2- tube usually with numerous scales and hairy areoles.
5 cm, diurnal or nocturnal; pericarpel and tube naked Fruit usually berry-like, juicy, sometimes splitting
except for minute scales; tepals short, erect or spread- longitudinally, rarely almost dry. Seeds small to me-
ing, variously coloured. Fruits indehiscent, red, con- dium-sized, diverse in shape, oval to almost circular,
spicuous or disintegrating in the cephalium; floral rarely hat-shaped, smooth to rugose or ruminate; cell
remnant tardily deciduous or persistent, rarely black- boundaries straight, interstices undifferentiated, min-
ening; pulp scanty, nearly white. Nine spp., endemic utely pitted or cratered, relief flat to rarely high-
to C and E Brazil. domed or high-conical towards periphery, cuticle
often weakly to regularly striate, sometimes strongly
folded and becoming detached; hilum and micropyle
26. Coleocephalocereus Backeberg conjunct or fused; appendages usually none, strop hi-
Coleocephalocereus Backeberg, Blatter f. Kakteenforschung olar pad rarely present (Rebutia). Distribution:
1938(6): [22] (1938). S America, S of the equator. Weakly differentiated
Buiningia F.Buxbaum (1971). from N otocacteae.

Branching from the base or unbranched; stems elon-

gate-globose to cylindric or columnar; ribs few to
many; areoles close-set; spines weak or strong.
Flowering areoles different from the non-flowering,
182 Cactaceae

28. Leocereus Britton & Rose fleshy, very spiny. The single species is endemic to the
Leocereus Britton & Rose, The Cactaceae 2: 108 (1920); Taylor Galapagos Islands, where it occurs on lava.
& Zappi, Bradleya 8: 107-108 (1990).
31. Espostoa Britton & Rose
Few-branched; rootstock woody; stems slender, erect,
Espostoa Britton & Rose, The Cactaceae 2: 60 (1920).
to 2(-3) m x 1-2.5 cm, ribbed; ribs (10-)12-20, low, Binghamia Britton & Rose (1920).
rounded; spines slender acicular, yellow to dark Pseudoespostoa Backeberg (1934).
brown. Flowers tubular, nocturnal, 4-7 cm; scales Thrixanthocereus Backeberg (1937).
numerous, small, acute, tinged brown; areoles of peri- Vatricania Backeberg (1950).
carpel with spines to 4 mm; areoles of tube with dark
hairs and bristles to 12 mm; inner tepals short, white. Shrubs or tree-like; stems cylindric or columnar,
Fruits globose to ovoid, 2-3 cm, red indehiscent, with many-ribbed, spiny. Fertile zone a lateral cephalium,
deciduous spine-clusters; pulp purple. A single vari- formed over several ribs, these more or less reduced
able species, endemic to E Brazil. and modified. Flowers tubular-campanulate, usually
nocturnal, pale, in some species rather small; tube
short; pericarpel and tube with small acute scales and
29. Haageocereus Backeberg
hairs but not spines; tepals short, spreading to re-
Haageocereus Backeberg, Blatter f. Kakteenforschung 1934(6): curved; lowermost stamens inserted on a diaphragm
[1] (1934). partially closing the nectar chamber. Fruits globose to
Loxanthocereus Backeberg (1937), in part. obovoid, more or less naked or with tufted hairs.
Weberbauerocereus Backeberg (1942).
Seeds diverse, oval to almost hat-shaped. A genus of
up to ten spp., S Ecuador, Peru and Bolivia.
Shrubby or tree-like; stems erect, ascending, decum-
bent or creeping; ribs usually numerous; areoles close-
set; spines numerous; additional bristles sometimes 32. Espostoopsis F. Buxbaum
present at flowering areoles. Flowers nocturnal Espostoopsis EBuxbaum in Krainz, Die Kakteen Lfg. 38/39,
(though often remaining open the following morning), sine pag. (1968).
regular or nearly so, tubular-funnelform, mostly white Austrocephalocereus Backeberg (1938), as to A. dybowskii.
or dull red; tube stout, fleshy; scales numerous; areoles Gerocephalus Ritter (1968).
of pericarpel and tube weakly to densely hairy; peri-
anth spreading, relatively narrow; stamens in a single Shrubby, mainly branching near the base; stems
series; style exserted. Fruits globose to ovoid, fleshy, cylindric, 2-4 m x ca. 8 cm; ribs numerous, low; spines
sparsely scaly and hairy; floral remnant persistent. up to 3 cm, pale yellow or brown, accompanied by
Perhaps ten spp. (out of more than 50 proposed), white hairs covering the whole stem. Fertile zone a
mainly in Peru and extending to N Chile and Bolivia lateral cephalium; flowers shortly tubular-campanu-
(Yungas). The relationships of H. weberbaueri, some- late, ca. 4 cm, nocturnal; peri carpel naked; scales of
times placed in the separate genus Weberbauero- tube less than 1 mm; areoles of tube naked; tepals
cereus, await elucidation, and several other small gen- short, white. Fruits broadly ovoid, 2.5 cm, nearly
era have been proposed which may belong in or near naked, pale pink, indehiscent; pericarp thick; floral
Haageocereus (Lasiocereus Ritter (1966), Pygmae- remnant blackening, persistent; pulp white. The
ocereus Johnson & Backeberg (1957), Yungasocereus single species is endemic to E Brazil (Bahia).
Ritter (1980), Maritimocereus Akers & Buining
(1950), Peruvocereus Akers (1947), Rauhocereus 33. Facheiroa Britton & Rose
Backeberg (1957)).
Facheiroa Britton & Rose, The Cactaceae 2: 173 (1920); Braun
& Esteves Pereira (1987-89), Kakt. and. Sukk. 38: 26-33, 83-
30. Brachycereus Britton & Rose 86, 184-187 (1987); ibid., 39: 64-68, 126-131 (1988); ibid. 40:
198-203 (1989).
Brachycereus Britton & Rose, The Cactaceae 2: 120 (1920). Zehntnerella Britton & Rose (1920).

Low shrub, forming clumps; stems erect or ascending, Shrubs or tree-like; stems cylindric; ribs 12-25 or
cylindric, not segmented, to 60 x 3-5 cm; ribs 16-22, more, low, narrow. Fertile zone, where differentiated,
low; areoles 6-9 mm apart; spines numerous, up 5 cm, a lateral cephalium sunken or superficial, bristly;
usually shorter, yellow at first, darkening with age. flowers tubular, nocturnal; scales numerous, small,
Flowers narrowly funnelform, ca. 9 cm, creamy imbricate; areoles of pericarpel and tube with more
white; pericarpel and tube with small scales and spiny or less copious hairs; tepals short; lowermost stamens
areoles; limb short. Fruit subglobose to oblong, inserted on a diaphragm partially closing the nectar
 Cactaceae 183

chamber. Fruits indehiscent, globose, fleshy, green to "closed" flowers of Cleistocactus sens. str., where the
brown or purple; pulp juicy, semi-translucent. A NE tepals hardly expand, to those of the former genera
Brazilian genus of about three spp. Bolivicereus, Borzicactus, Loxanthocereus, etc.,
where the limb is expanded and more or less oblique.
All are adapted to pollination by humming-birds.
34. Samaipaticereus Cardenas
Samaipaticereus Cardenas, Cact. Succ. 1. (U.S.) 24: 141 (1952).
36. Denmoza Britton & Rose
Tree-like, ca. 3 m; branches erect, unsegmented,
Denmoza Britton & Rose, The Cactaceae 3: 78 (1922).
ribbed; ribs 4-6; spines few, subulate. Flowers nar-
rowly funnelform, ca. 5 cm, nocturnal; tube slightly
Globose to shortly columnar; stem simple, eventually
curved; scales to 1.5 cm; areoles of pericarpel and
many-ribbed. Flowering areoles with or without extra
tube with short hairs and sparse bristles; tepals ca.
bristles; flowers tubular, slightly zygomorphic, diur-
1 cm, white; staminal throat-circle present. Fruit
nal, scarlet; tube straight or curved and dilated above
globose, truncate, tuberculate, pink-red, splitting
the pericarpel; scales numerous; areoles of peri carpel
lengthwise; pulp red-orange; floral remnant persis-
tent. A monotypic genus, endemic to Bolivia. and tube with numerous hair-spines; perianth unex-
panded; tepals short, erect; stamens inserted in the
throat and tube; filaments and style long-exserted;
35. Cleistocactus Lemaire nectar-chamber plugged with a staminodial collar
Cleistocactus Lemaire, Illustr. Hort. 8: Misc. 35 (1861). and hairs. Fruit globose, tufted with short hair-spines,
Borzicactus Riccobono (1909); Kimnach, Cact. Succ. 1. (D. S.) splitting and drying when ripe; pulp white. A single
32: 8-13,57-60,92-96,109-112 (1960). variable species; endemic to Wand NW Argentina.
Loxanthocereus Backeberg (1937), in part.
Seticereus Backeberg (1942).
Bolivicereus Cardenas (1951). 37. Oreocereus (A. Berger) Riccobono
Cephalocleistocactus Ritter (1959).
Akersia Buining (1961). Oreocereus (A. Berger) Riccobono, Boll. R. Ort. Palermo 8: 258
Winteria Ritter (1962), non Murray. (1909).
Seticleistocactus Backeberg (1963). Arequipa Britton & Rose (1922).
Winterocereus Backeberg (1966). Morawetzia Backeberg (1936).
Hildewintera Ritter (1966).
Borzicactella Ritter (1981).
Shrubs; stems erect or ascending, cylindric; areoles,
especially the flowering, often developing long white
Shrubby or rarely tree-like; stems erect, ascending, hairs; spines numerous. Flowers tubular-funnelform,
decumbent, procumbent or pendent, cylindric, usual- more or less zygomorphic, diurnal, usually orange or
ly slender; ribs 5-30, low. Fertile zone undifferen- red or purple; tube straight to somewhat curved; are-
tiated or rarely the flowering areoles with more wool oles of pericarpel and tube numerous, more or less
and/or bristles; flowers narrowly tubular, regular or hairy; perianth-limb narrow, oblique; upper tepals
somewhat zygomorphic, diurnal, red, orange, yellow suberect, lower spreading or recurved; stamens
or green; tube straight or kinked beyond the ovary numerous inserted in the throat and tube, the lower-
and more or less S-shaped; scales numerous, small, most filaments coalescent at the base to form a dia-
sometimes imbricate; areoles of pericarpel and tube phragm over the nectar-chamber, sometimes the dia-
woolly or hairy; perianth-limb unexpanded, the te- phragm invested with staminodial hairs; style and
pals scarcely larger than the scales of the tube and not stamens exserted. Fruits globose to ovoid, hollow,
or only slightly spreading, or expanded and more or dehiscing by abscission at the base; pericarp fleshy;
less strongly oblique, the upper tepals suberect and pulp none. Seeds broadly oval, ruminate. Six to seven
the lower spreading or recurved; stamens variously spp., W South America (Andes of S Peru, S Bolivia,
included or exserted, inserted in the throat and tube N Chile and N Argentina).
in two series, the lowermost on a collar partially clos-
ing the nectar-chamber; style more or less exserted.
Fruit small, globose, tufted with sparse hairs or gla- 38. Matucana Britton & Rose
brescent, indehiscent or bursting open when ripe;
floral remnant usually persistent; pulp fleshy. Matucana Britton & Rose, The Cactaceae 3: 102 (1922); Breg-
man, R. et a\., Succulenta (Netherlands) 65-69 (1986-90).
A genus of 40-50 species in S Ecuador, Peru, Boli-
[Revision of the genus, in 32 parts. In Dutch.]
via, N Argentina, Paraguay, W Brazil and Uruguay. Arequipiopsis Kreuzinger & Buining (1941).
As now defined, the genus displays a variety of inter- Submatucana Backeberg (1959).
grading flower-types, from the strictly tubular, Eomatucana Ritter (1965).
184 Cactaceae

Low-growing; stems globose to short-cylindric; ribs setose; stamens numerous, throat-circle usually pre-
few to numerous, broad, low, more or less tubercu- sent, sometimes the filaments of the throat-circle con-
late; spines numerous, fine, to few or absent. Flowers trasting in colour with the perianth and decurrent
subapical, funnelform to narrowly tubular-funnel- into a distinct membranous ring (hymen); nectar-
form, diurnal, in various colours; areoles of pericar- chamber rarely plugged with a tuft of staminodial
pel and tube naked or hairy; limb usually more or less hairs. Fruits globose to narrowly ovoid or oblong,
zygomorphic, often narrow, rarely regular (M. aurei- fleshy to dry.
flora); filaments of lower stamens coalescent at base A complex genus with 50-100 species in central
to form a diaphragm over the nectar-chamber (except and southern South America. The genera listed as
in M. madisoniorum and M. oreodoxa; staminodial synonyms fall into two groups, one night-flowering
hairs sometimes also present. Fruits hollow, splitting (Echinopsis sens. str., Trichocereus, Setiechinopsis,
longitudinally from the base, the pericarp somewhat Pseudolobivia), the other day-flowering (Acantho-
fleshy at first, later dry. Seeds diverse, oval to hat- calycium, Lobivia, Soehrensia, Helianthocereus, Cha-
shaped, ruminate or not. About six to seven spp. in maecereus).
Peru. The genus ist only weakly differentiated
(mainly by habit) from Oreocereus. Bregman et al.
(1. c.) recognise 19 species. 41. Arthrocereus (A. Berger) A. Berger
Arthrocereus (A. Berger) A. Berger. Kakteen, 146, 327 (1929).
39. Oroya Britton & Rose
Shrubs; stems cylindric; ribs 10--18, low and narrow;
Oroya Britton & Rose, The Cactaceae 3: 102 (1922).
areoles small; spines numerous, thin. Flowers elon-
gate-funnelform, nocturnal; scales sparse; areoles of
Low-growing; stem simple or offsetting, flattened-
pericarpel and tube with wool and hair-spines; limb
globose to very short cylindric, many-ribbed; areoles
broad; staminal throat-circle sometimes present but
elongate; spines pectinate. Flowers subapical, shortly
indistinct. Fruit globose or ovoid, green, with scales
funnelform or campanulate, regular, yellow to red;
and hair-spines; pulp off-white. Up to five spp. ende-
tube very short; scales small; areoles of pericarpel
mic to Wand SE Brazil.
and tube sparsely woolly; outer tepals spreading,
inner suberect; stamens inserted in the throat and
tube, the lowermost on a collar partly enclosing the 42. Rebutia Schumann
nectar-chamber; filaments and style not exserted be-
Rebutia Schumann, Monatsschr. Kakteenk. 5: 102 (1895).
yond the perianth. Fruits obovoid, slightly fleshy with
Aylostera Spegazzini (1923).
small scales. One to two spp., confined to Peru. Mediolobivia Backeberg (1934).
Weingartia Werdermann (1937).
Sulcorebutia Backeberg (1951).
40. Echinopsis Zuccarini
Echinopsis Zuccarini, Abh. Baycr. Akad. Wiss. Mtinchen 2: 675 Low-growing; stems simple or more often freely clus-
(1837).
Lobivia Britton & Rose (1922); Rausch, Lobivia 1985 (1986)
tering, small, globose to shortly cylindric, tuberculate
(inc!. Chamaecereus etc.). or weakly ribbed; areoles circular or oval to elliptic-
Trichocereus Riccobono (1909); Kiesling, in Darwiniana 21: linear; spines relatively weak, often scarcely differen-
263-330 (1978). tiated into radial and central. Flowers diurnal, freely
Chamaecereus Britton & Rose (1922). produced, usually arising towards the stem-base, fun-
Acanthocalycium Backeberg (1935). nelform, comparatively small (less than 5 cm), vari-
Soehrensia (Backeberg) Backeberg (1938).
ously coloured; pericarpel and tube with small scales
Setiechinopsis (Backeberg) Dc Haas (1940).
Pseudolobivia Backeberg (1942). naked or with hairs and sometimes bristles in their
Helianthocereus Backebcrg (1949). axils; tube short or elongate, often slender, often
curved, sometimes occluded ("fused with the style");
Tree-like, columnar, shrubby or globular; stems stamens usually in a single series. Fruits small, sub-
usually distinctly ribbed, very spiny to nearly spine- globose; peri carp juicy at first, drying papery; with-
less; ribs few to numerous, often somewhat tubercu- ered perianth persistent. Seeds oval; testa relief flat to
late beneath or between the areoles. Flowers lateral high-conical, especially at distal end; strophiolar pad
or subapical, funnelform to salverform or subcampa- present in some species. Thirty to 40 species, E cor-
nulate, nocturnal or diurnal, often large; scales of dilleras of the Andes, from Bolivia (Cochabamba to
pericarpel and tube relatively narrow, often Tarija) to NW Argentina (Jujuy to Tucuman). It con-
numerous, rarely bristle-tipped; areoles of pericarpel sists of five intergrading groups, corresponding to the
and tube more or less densely hairy and very rarely genera listed as synonyms.
 Cactaceae 185

43. Mila Britton & Rose 5. Tribe Notocacteae F. Buxbaum (1958)

Mila Britton & Rose, The Cactaceae 3: 211 (1922); Donald,
Ashingtonia 3: 31-35, 38--D2 (1978). Mostly globular, small to medium-sized, rarely
shrubby to tree-like (Corryocactus, Eulychnia); root-
stock sometimes napiform; stems unsegmented,
Low shrub; rootstock stout; stems clustering, short-
ribbed, tuberculate-ribbed or tuberculate; fertile
cylindric; ribs 10--13, low; spines variable in number,
zone undifferentiated, but the flowers mostly subapi-
length, texture and colour, usually more than 20, 1-
cal and the stem-apex often densely felted. Flowers
3 cm. Flowers subapical, small, funnelform-campanu-
small to medium-sized, always diurnal, usually
late, 2-3.3 cm, yellow; scales of pericarpel and tube
brightly coloured, commonly yellow, regular, rarely
small, narrow; areoles of pericarpel and tube woolly
somewhat zygomorphic (Neoporteria spp.); scales
or bristly. Fruits small, globose, to 12-15 mm, fleshy,
usually numerous, small; areoles of pericarpel and
red-tinged, with scales and woolly hairs. A single
tube usually with numerous bristles and/or hairs.
variable species in central Peru.
Fruit usually dry, variously dehiscent or disintegrat-
ing, rarely berry-like, juicy, indehiscent. Seeds small
to medium-sized, rarely large, diverse in shape, oval
44. Gymnocalycium Pfeiffer to almost circular, rarely hat-shaped, smooth or
Gymnocalycium Pfeiffer, Abbild. Beschr. Cact. 2: sub tt. 1, 12 rarely rugose or ruminate; cell boundaries straight,
(1845); Schlitz, Friciana 7(46) (1968). interstices usually undifferentiated, rarely minutely
Brachycalycium Backeberg (1942). pitted, relief flat to high-domed or with minute hair-
like projections, cuticle sometimes striate, sometimes
Low-growing globular cacti, mostly unbranched, strongly folded and becoming detached; hilum and
some clustering; stems often strongly depressed, or micropyle usually conjunct, rarely disjunct or fused;
globose to short-cylindric, umbilicate at apex; ribs 4- strophiole or strophiolar pad often present, or very
15 or more, sometimes spiralling, variously tubercu- rarely mucilage sheath present covering entire seed.
late, spiny; fertile zone undifferentiated. Flowers Distribution: S South America.
usually subapical, funnelform to campanulate, diur-
nal, white or pale pink, or sometimes red or yellow; 46. Corryocactus Britton & Rose
scales of pericarpel and tube characteristically broad,
obtuse, with hyaline margins; areoles of pericarpel Corryocactus Britton & Rose, The Cactaceae 2: 66 (1920).
Erdisia Britton & Rose (1920).
and tube naked; perianth spreading or suberect;
stamens often in two groups, the lowermost sur-
rounding the style immediately above the nectar- Shrubby or rarely tree-like; stems erect, ascending or
chamber, the remainder inserted nearer the mouth of procumbent, cylindric, stout to slender; ribs 4-10;
the tube; throat-circle absent. Fruits oblong-obovoid spines often fierce. Flowers campanulate, diurnal;
to globose, dry or fleshy, splitting, cracking or deli- tube short; scales numerous, small; areoles of pericar-
quescent when ripe. Seeds diverse. Up to 50 spp., pel and tube felted and spiny; limb spreading; tepals
S Brazil, Paraguay, Bolivia, Uruguay, Argentina. short, yellow, orange or purple-red; stamens inserted
Usually classified according to seed characters. in the broad throat. Fruit globose, spiny; floral rem-
nant caducous; pulp juicy. Seeds broadly oval,
smooth to tuberculate, sometimes wrinkled; mucilage
45. Discocactus Pfeiffer sheath sometimes present, covering entire seed. Up
to 20 spp., Peru, Bolivia and Chile.
Discocactus Pfeiffer, AUg. Gartenz. 5: 241 (1837); Buining, The
genus Discocactus Pfeiffer (1980); Taylor, Cact. Succ. J. Gr.
Brit. 43: 37-40 (1981). 47. Austrocactus Britton & Rose
Austrocactus Britton & Rose, The Cactaceae 3: 44 (1922).
Low-growing; stems mostly simple, depressed-glo-
bose to globose, ribbed, spiny; ribs distinct or strongly Low-growing; stem simple or branched at base, slen-
tuberculate. Flowering-zone (cephalium) terminal, der-cylindric to cylindric, up to 50 x 8 cm; ribs 6--12,
depressed, densely woolly and more or less bristly; somewhat tuberculate; central spines hooked or
flowers developing very fast, tubular-salverform, noc- straight. Flowers subapical (lateral in A. spiniflorus),
turnal, white; tube slender, scaly. Fruit globose-cla- urceolate or campanulate, 3-6 cm; pericarpel broadly
vate to oblong, naked, slightly fleshy. Seeds broadly obconic; scales triangular, mucronate; tube short; are-
oval to subglobose, papillate-tuberculate. Eight spp., oles of pericarpel and tube with woolly hairs and
Brazil, E Bolivia and N Paraguay. usually bristles; stamens in two series, the lower
186 Cactaceae

forming a ring round the style; stigmas typically 51. Eriosyce Philippi
purple-red. Fruit typically dry, with wool and bristles, Eriosyce Philippi, An. Univ. Chile 41: 721 (1872).
splitting irregularly or at base; floral remnant persist- Rodentiophila Ritter (1959).
ent. Seeds broadly oval, ca. 2.5 x 2 mm, black-brown,
wrinkled. About five spp., Wand S Argentina and C Stem simple, globose to shortly columnar, to 70 x
and S Chile. 55 cm, ribbed; ribs 19-40; spines numerous. Flowers
arising in the woolly apex, campanulate, orange-red
48. Eulychnia Philippi to purple; areoles of pericarpel and tube with matted
white hair-spines and the upper with bristly spines;
Eulychnia Philippi, Florula Atacamensis 23: 23 (1860); Ritter, perianth limb short. Fruits oblong, densely woolly
Kakteen in Siidamerika 3: 892-903 (1980).
and spiny apically, eventually dry, detaching. Seeds
Philippicereus Backeberg (1942).
broadly oval, relatively large, ca. 2.5 x 2 mm, slightly
keeled. Two spp., endemic to Chile. It is arguable that
Shrubs or tree-like; stems ascending or procumbent;
Eriosyce (which is the older name) should be com-
ribs 9-16; spines often strong, elongate. Flowers
bined with Neoporteria, but for the purpose of this ac-
borne near tips of branches, small, broadly campanu-
count the genus is treated in its prevalent, strict sense.
late, diurnal; tube short, with numerous small scales;
areoles of pericarpel and tube with woolly hairs (and
bristly spines in E. castanea); perianth erect or some- 52. Neoporteria Britton & Rose
what spreading; tepals short, pink or white; style Neoporteria Britton & Rose, The Cactaceae 3: 94 (1922); Donald
short and thick. Fruits globose, fleshy and scaly or and Rowley, Cact. Succ. J. Gr. Brit. 28: 54-58, 74-77, 83 (1966).
hairy, rarely spiny (E. castanea); floral remnant per- Islaya Backeberg (1934).
sistent. Seeds broadly oval, often grey from the de- Pyrrhocactus (A. Berger) Backeberg & F. Knuth (1935).
taching cuticle. About six spp., coastal Chile and Horridocactus Backeberg (1938).
Peru. Neochilenia Backeberg ex Doelz (1942).
Thelocephala Ito (1957).

49. Copiapoa Britton & Rose Stems usually unbranched, globose to short-cylindric,
ribbed; ribs usually divided into prominent tubercles.
Copiapoa Britton & Rose, The Cactaceae 3: 85 (1922); Ritter,
Kakteen in Siidamerika 3: 1044-1107, Figs. 963-1058 (1980). Flowers subapical, tubular-funnelform, broadly fun-
Pilocopiapoa Ritter (1961). nelform or campanulate, diurnal, in various colours;
tube distinct or very short, with small scales; areoles
Low-growing or mound-forming, very small or to of pericarpel and tube variously felted and hairy or
1 m; rootstock fibrous or with greatly enlarged tap- bristly, the uppermost more strongly developed; te-
root, sometimes connected to stem by a slender neck; pals spreading, or the outermost spreading and the
stems globose to stout -cylindric, strongly or weakly inner erect, concealing the stamens; stamens inserted
ribbed or tuberculate; apex often immersed in dense in the throat and tube in one series. Fruits globose to
areolar wool; spines few to numerous. Flowers short- ovoid, sometimes balloon-like (N. isfayensis); peri-
funnelform to campanulate, small, yellow (or very carp initially fleshy but the interior always dry when
rarely red); pericarpel and tube almost naked or mature, and usually releasing the seeds through a
scales few, small, and areoles naked (hairy in C. so- basal opening. Seeds broadly oval, ruminate or not.
faris). Fruit small, globose to turbinate, dehiscent at About 25 spp. in Chile, S Peru und W Argentina.
apex, usually with 1-2 scales near apex. About Following Donald and Rowley (I. c.), the genus has
20 spp., coastal deserts of N Chile. been taken to comprise a number of subgenera and
sections corresponding to the segregate genera pro-
posed by Backeberg and others. The component
50. Neowerdennannia Backeberg groups are distinguished by flower and fruit morpho-
Neowerdermannia Fric, Kaktusar 1(11): 85 (1930). logy, but intergrade.

Low-growing, with stout taproot; stem simple, globose

53. Parodia Spegazzini
to depressed; ribs indistinct, ca. 16, spiralling, deeply
divided into triangular tubercles; areoles at base of Parodia Spegazzini, An. Soc. Cient. Argent. 96: 70 (1923).
upper side of tubercles. Flowers funnelform, diurnal; Malacocarpus Salm-Dyck (1850), non Fischer & Meyer.
Notocactus (Schumann) Fric (1928).
pericarpel and tube with few fleshy scales, areoles Brasilicactus Backeberg (1942).
naked; limb spreading, white or lilac-pink. Fruit lat- Eriocactus Backeberg (1942).
erally dehiscent. Seeds broadly oval, ruminate. Two Wigginsia Porter (1964).
spp., N Argentina, S Bolivia, Peru and N Chile. Brasiliparodia Rittcr (1979).
 Cactaceae 187

Low-growing, simple or clustering; stems mostly 56. Uebelmannia Buining

small, globose to cylindric, ribbed or tuberculate. Uebelmannia Buining, Succulenta (NL) 46: 159 (1967).
Flowers subapical, shortly funnelform, diurnal,
brightly coloured; scales narrow; areoles of pericar-
Unbranched; stems mostly small, globose to cylin-
pel and tube with hairs and bristles, or bristles re-
dric, ~ibbed, spiny;. epidermis smooth or papillate,
stricted to the uppermost areoles. Fruits globose to
sometimes covered m wax platelets; abundant mucil-
clavate-cylindric, woolly and/or bristly, dry or nearly age-cells present often forming conspicuous ducts in
so, mostly thin-walled and disintegrating at or near
the cortex. Flowers subapical, shortly funnelform,
base, or thick-walled and splitting laterally, or pink small, diurnal, yellow; areoles of pericarpel and tube
and fleshy at first, later hollow and dry. Seeds diverse.
with dense wool and a few inconspicuous bristles.
A genus of perhaps 50 spp., in S South America Fruits globose to cylindric, yellow or red, naked
(S Brazil, Uruguay, NE Argentina, S Paraguay and
below, woolly and bristly at apex, thin-walled and dry
the E Andes of Bolivia and NW Argentina). A great
at maturity, disintegrating. Five spp., endemic to the
many ill-defined species have been described. The se-
mountains of E Brazil (Minas Gerais); its systematic
gregate genera listed have been based on for the most position is uncertain.
part on fruit and seed characters.

54. Blossfeldia Werdermann 6. Tribe Rhipsalideae De Candolle (1828)

Blossfeldia Werdermann, Kakteenkunde 1937: 162 (1937).
Epiphytic or epilithic, usually pendulous, rarely
~iny, button-like; stems simple or caespitose, not
creeping or shrubby, never scandent; stems terete,
nbbed or tuberculate, poikilohydric; spines absent. angled or flattened, usually segmented; areoles more
Flowers subapical, small, 0.6-1.5 cm, diurnal, almost or less sunken. Flowers usually lateral, diurnal and
white; scales few, minute; areoles of pericarpel and remaining open at night, rarely subapical or from api-
tube sparsely woolly. Fruit globose, red. Seeds very cal, composite areoles, rotate, regular, usually small
small, 0.4 x 0.4 mm, brown, with minute hair-like cell- or very small, white or pale yellow or pink, rarely me-
projections; strophiole present, large. A single dium-sized, red or purple, regular or more or less tu-
species native to S Bolivia and NW to N Argentina, bular and zygomorphic; pericarpel usually naked,
where it occurs in rock crevices. It is remarkable not rarely with areoles and hairs or spines. Fruit berry-
only for its small size but for its ability to withstand like, juicy, indehiscent. Seeds small to medium-sized
desiccation. Stomata are very few and confined to the without pronounced surface sculpturing; cell boun~
immediate vicinity of the areoles. daries straight to irregularly curved, interstices undif-
ferentiated to minutely pitted; hilum and micropyle
fused; mucilage sheath present, usually restricted to
55. Frailea Britton & Rose hilum area. Mainly E South America, with centres of
Frailea Britton & Rose, The Cactaceae 3: 208 (1922). diversity in E Bolivia and SE Brazil, a few species
extending to C America, one to the Old World.
Low-growing, caespitose or unbranched; stems de- The tribe is well-defined and is not related to the
pressed-globose to cylindric, usually weakly ribbed other epiphytic group in the family, the Hylocereeae.
or tuberculate. Flowers shortly funnelform, yellow, Affinities are discernible with the Notocacteae (Cor-
diurnal, but opening only briefly, or cleistogamous; ryocactus).
areoles of peri carpel and tube with dense wool and
bristles. Fruits thin-walled, dry, densely packed with 57. Lepismium Pfeiffer
seeds, indehiscent or rupturing irregularly; floral
remnant (inc!. wool and bristles) persistent. Seeds Lepismium Pfeiffer, AUg. Gartenz. 3: 315 (1835); Barthlott,
Bradleya 5: 97-100 (1987).
broadly oval to hat-shaped, sometimes with minute, Pfeiffera Salm-Dyck (1845).
hair-like cell projections; strophiole absent. Some Rhipsalis subg. Ophiorhipsalis Schumann (1898).
15 species in E Bolivia, S Brazil, Paraguay, Uruguay Acanthorhipsalis Britton & Rose (1923).
and NE Argentina. Lymanbensonia Kimnach (1984).

Shrubs, epiphytic or epilithic, creeping or pendulous;

stems cylindric, ribbed, angled, winged or flat,
u.sually segmented, the younger segments arising
sm~ly from the sides or apices of older segments;
rudimentary scale-leaves often clearly visible; spines
188 Cactaceae

present or absent. Flowers small, rotate, campanu- 5 cm; new segments arising singly or in clusters from
late or rarely tubular-campanulate; pericarpel often a composite areole at the apices of older segments;
tuberculate and spiny or angled and with or without spines 0 or soft, bristly. Flowers apical, regular, cam-
spines, rarely almost terete; tube usually very short panulate, intense yellow, pink or red; pericarpel
or none. Fruits berry-like, naked or rarely spiny, angled or terete, naked; tube short. Fruits small, obo-
usually red to black. void, naked. Five spp., endemic to SE Brazil. Two
As now understood, a genus of 16 spp., mainly in subgenera: subg. Hatiora (stems cylindric; two spp.)
E Bolivia, extending to Brazil and Argentina. The and subg. Rhipsalidopsis (Britton & Rose) Barthlott
species lack the characteristic acrotonic branching (stems flattened; three spp., including the popular
pattern of other Rhipsalideae, and are often more or ornamental "Easter cacti").
less spiny. Barthlott (I. c.) recognises five subgenera.
60. Schlumbergera Lemaire
58. Rhipsalis Gaertner Schlumbergera Lemaire, Rev. Hart. ser.4, 7: 253 (1858); Hunt,
Rhipsalis Gaertner, Fruct. Sem. 1: 137 (1788); Barthlott in Son- Kew Bull. 23: 255-263 (1969); Barthlott & Rauh, Kakt. and.
derb. Naturw. Ver. Hamburg 7: 241-248 (1983) and in Brad- Sukk. 28: 273-278 (1978); McMillan, Christmas Cacti/Weih-
leya 5: 97-100 (1987). nachtskakteen (1985).
Erythrorhipsalis A. Berger (1920). Epiphyllum Pfeiffer (1837), non Haworth (1812).
Zygocactus Schumann (1890).
Epiphyllanthus A.Berger (1905).
Epiphytic, rarely epilithic cacti, often pendulous;
stems cylindric, ribbed, angled, winged or flat, usually Epiphytic or epilithic shrubs; stems segmented; seg-
segmented and spineless, the younger segments ments flattened, compressed or terete, oblong or
usually arising singly or in clusters (verticils) at the obovate, new segments arising from a composite are-
apices of older segments. Flowers small to very small, ole at the apices of older segments; spines setaceous,
rotate, mostly almost white; pericarpel terete, usually short, or none. Flowers apical, nearly regular to
naked, rarely with soft, bristly spines; tube very short strongly zygomorphic; tube distinct, bearing tepaloid
or none; disk, where present (subg. Rhipsalis) annu- scales; perianth usually purple, pink or white, red,
lar, nectariferous. Fruits small, berry-like, usually orange or yellow in some cultivated forms; stamens
naked. inserted on the tube, the lowermost united at the base
A genus of up to 50 spp. distributed throughout to form a short tube around the style; stigmas erect,
tropical America, with one species (R. baccifera) ex- connivent. Fruits berry-like, globose to obconic,
tending to tropical Africa, Madagascar and Ceylon ribbed or terete. Six spp., SE Brazil, divisible into two
(Sri Lanka). This is the only cactus species which oc- groups. In Schlumbergera sens. str. (inc\. Zygocactus
curs spontaneously outside America. Rhipsalis is the Schumann), the younger stem segments are flat-
principal South American genus of epiphytic cacti, tened, with the areoles confined to the margins and
with its centre of distribution in E Brazil. It is cur- apex; in the other (Epiphyllanthus A. Berger), the
rently restricted to small-flowered species in which younger stem segments are cylindric or compressed
the stem segments, whether terete or flat, branch ac- and Opuntia -like, with areoles distributed generally
rotonically, i. e. predominantly from the apex. There over the surface.
are three subgenera, subg. Erythrorhipsalis A. Berger
(stems terete; flowers campanulate-funnelform;
seven spp.), subg. Rhipsalis (stems terete or angled, 7. Tribe Browningieae EBuxbaum (1966)
rarely flattened; flowers rotate; about 35 spp.) and
subg. Phyllarthrorhipsalis E Buxb. (stems flattened; Large tree-like, shrubby or columnar; stems ribbed,
flowers rotate; about seven weakly defined spp.). usually stout, segmented or unsegmented, usually
becoming very fiercely spiny; fertile zone undifferen-
59. Hatiora Britton & Rose tiated, or sometimes less spiny. Flowers lateral,
medium-sized to large, tubular, funnelform or salver-
Hatiora Britton & Rose, Bailey, Standard Cycl. Hart. 3: 1432
form, usually nocturnal, pale-coloured; perciarpel
(1915); Barthlott in Bradleya 5: 97-100 (1987).
Hariota De Candolle (1834), non Adanson (1763).
and tube often with conspicuous sometimes imbri-
Rhipsalidopsis Britton & Rose (1923). cate scales, or with areoles with spines or bristles.
Epiphyllopsis (A. Berger) Backeberg & F. Knuth (1935). Fruits fleshy, indehiscent, scaly, spiny or almost
Pseudozygocactus Backeberg (1938). naked, usually with white funicular pulp. Seeds me-
dium-sized to large, sometimes rugose to strongly
Epiphytic or epilithic shrubs; stems cylindric, angled, ruminate; cell boundaries straight or rarely irregu-
winged or flat, segmented; segments usually less than larly curved, interstices undifferentiated to minutely
 Cactaceae 189

pitted and cratered towards periphery, relief flat to carp thick, dull red, almost naked, or with an occa-
high-domed at periphery, cuticle usually weakly stri- sional spine. A single variable species endemic to the
ate to strongly folded; hilum and micropyle conjunct; Galapagos Island (Ecuador).
appendages none or mucilage sheath present, cover-
ing entire seed (Calymmanthium, Neoraimondia).
Distribution: Andean S America, from Ecuador to 64. Neoraimondia Britton & Rose
N Chile and Argentina. Neoraimondia Britton & Rose, The Cactaceae 2: 181 (1920).
Here treated in an amplified sense, following Neocardenasia Backeberg (1949).
Barthlott (1988) and Hunt and Taylor (1990). The
group is reasonably coherent, but doubtfully sepa- Shrubby or tree-like; stems erect, ribbed; ribs 4-8;
rable from Pachycereeae.
non-flowering areoles large, brown-felted, usually
spiny and one or more of the spines very long; flower-
61. Calymmanthium Ritter ing areoles felted, nearly spineless, enlarged and pro-
longed into a spur by continuous growth, flowering
Calymmanthium Ritter, Kakt. and. Sukk. 13: 25 (1962).
annually. Flowers 1-2 per areole, small, pink or off-
white, nocturnal (?); pericarpel and tube bearing
Shrubby or arborescent; stems segmented, 3-4 wing- small scales and felted areoles with or without
ed, spiny, especially the old stems. Flowers nocturnal bristles; limb short; stamens inserted in the upper part
(?), narrowly tubular-campanulate; tube partially of the tube. Fruits globose to oblong, felted and more
sheathing the perianth in bud; scales small; external or less spiny. Seed broadly oval, rugose or ruminate;
areoles of pericarpel and tube with felt and bristly mucilage-sheath present, covering entire seed. A
spines, internal naked; perianth-limb narrow, spread- genus of two species in Peru, N Chile and Bolivia.
ing, the lowest part sheathed by the tube; outer tepals
tinged red, inner white; stamens inserted in the throat
and tube. Fruit fleshy, indehiscent, spineless or nearly 65. Browningia Britton & Rose
so. Seeds broadly oval; mucilage sheath covering the
entire seed. A single species, endemic to N Peru. The Browningia Britton & Rose, The Cactaceae 2: 63 (1920).
Gymnanthocereus Backeberg (1937).
concealed development of the perianth is a feature Azureocereus Akers & Johnson (1949).
unique to this genus. Castellanosia Cardenas (1951).
Gymnocereus Backeberg (1959).
62. Armatocereus Backeberg
Shrubby or tree-like, to 10 m; stems cylindric, ribbed;
Armatocereus Backeberg, Blatter f. Kakteenforschung 1938
ribs 7-34; flowering areoles usually less spiny than the
(6): [21] (1938); Hunt, Brad1eya 9: 83-84 (1991).
non-flowering. Flowers tubular-funnelform, noctur-
nal; tube somewhat curved; scales conspicuous, over-
Shrubby or tree-like; stems stout, erect or ascending,
lapping; areoles of pericarpel and tube naked or al-
cylindric, ribbed, constricted annually and thus more
most naked; tepals spreading, rather short, white.
or less segmented; ribs 3-11. Flowers tubular, noctur-
Fruit small, globose to ovoid, somewhat fleshy, or be-
nal; tube not markedly flared; areoles of pericarpel
coming dry, covered with deciduous scales. Seeds
and tube felted, often spiny; limb short; tepals erect
diverse, smooth to ruminate. Seven species, Bolivia,
to spreading, white or rarely red; stamens inserted in
Peru and N Chile.
the throat and upper part of tube. Fruit globose to
ovoid, fleshy, spiny. Up to ten species, Colombia,
Ecuador and Peru. 66. Stetsonia Britton & Rose
Stetsonia Britton & Rose, The Cactaceae 2: 64 (1920).
63. Iasminocereus Britton & Rose
fasminocereus Britton & Rose, The Cactaceae 2: 146 (1920). Tree-like, 5-10 m; stems cylindric, ribbed, spiny; ribs
8-9. Flowers funnelform, 12-15 cm, nocturnal; peri-
Tree-like, to 8 m; branches segmented, ribbed; ribs carpel with numerous broad, imbricate apiculate
11-22; spines numerous, long. Flowers salverform, scales; tube elongate, with scattered scales; perianth
4-9 x 2-D cm, nocturnal; pericarpel and tube slender, limb broad, spreading to rotate; inner tepals white;
creamy white; scale numerous, small, triangular; stamens numerous, inserted in the tube. Fruits ob-
areoles of pericarpel and tube naked; tepals nar- long or globose, green or tinged red. Seeds rugose. A
rowly oblong-spathulate, sometimes apiculate, single species, native to Paraguay, NW Argentina and
creamy white. Fruits oblong-ovoid, to 6 x 4 cm; peri- S Bolivia.
190 Cactaceae

8. Tribe Pachycereeae EBuxbaum (1958) fluent, or connected by a groove, densely felted,

spineless, or furnished with numerous long, setaceous
Large, tree-like, shrubby or columnar; stems ribbed, spines; flowers small to medium-sized, usually noc-
unsegmented, usually stout; fertile zone undifferen- turnal, shortly tubular, funnelform or campanulate;
tiated or commonly differentiated and forming an tube with scales; areoles of pericarpel and tube naked
apical or lateral cephalium. Flowers lateral, subapical or woolly and/or bristly. Fruits subglobose, fleshy
or restricted to the cephalia, small to medium-sized, with red or purple pulp or soon dry, generally densely
tubular, campanulate or funnelform, usually regular, bristly, at least the apex, sometimes the bristles cadu-
nocturnal, pale-coloured, rarely diurnal, yellos (Ber- cous. Twelve species, endemic to Mexico.
gerocactus) or zygomorphic, red (Rathbunia); peri-
carpel and tube variously scaly; areoles of peri carpel 69. Camegiea Britton & Rose
usually with spines or bristles, rarely naked, those of
the tube commonly naked. Fruits fleshy, usually Carnegiea Britton & Rose, J. N. Y. Bot. Gard. 9: 187 (1908).
spiny, or the areoles and spines caducous, rarely scaly
or almost naked, indehiscent or commonly dehiscent Columnar or few-branched, to 16 m; stems erect,
by splits from the apex, usually with red or white stout, ribbed; ribs 12-30, spiny. Flowers subapical,
funicular pulp. Seeds diverse, medium-sized to very funnelform-campanulate, 9-12 x 5-6 cm; pericarpel
large, ruminate or not; cell boundaries straight, rarely and tube with numerous pronounced decurrent
irregularly curved, interstices undifferentiated to scales; areoles of pericarpel and tube felted, without
minutely pitted or cratered, relief flat to domed, spines; inner tepals white; stamens very numerous.
cuticle rarely striate; hilum and micropyle conjunct or Fruit obovoid, red or red-green, scaly, splitting ir-
fused; appendages none. Distribution: Mainly Mexi- regularly from the apex; pulp red. A single species,
co, extending to SW USA, Caribbean region and the picturesque "saguaro", C. gigantea, native to SW
C America to Venezuela. USA and NW Mexico.

67. Bergerocactus Britton & Rose 70. Neobuxbaumia Backeberg

Neobuxbaumia Backebcrg, Blatter f. Kakteenforschung 1938:
Bergerocactus Britton & Rose, Contrib. U. S. Nat!. Herb. 12:
[8,21 J (1938).
435 (1909).
Rooksbya Backeberg (1959).

Shrub; rootstock thickened; stems erect, to 60 x 3-6 Massive columnar or tree-like cacti, simple or branch-
cm, branching from base, rather woody; ribs 20-25, ed; stems stout cylindric, ribbed; ribs numerous,
low, spiny. Flowers small, 2 x 2 cm, diurnal, pale yel- usually low. Fertile zone usually unmodified, or api-
low; areoles of peri carpel and tube felted and spiny; cal, with larger areoles with numerous bristles and
tube short; tepals numerous, spreading. Fruit glo- bristly spines; flowers tubular-campanulate or tubu-
bose, densely spiny. A single species of uncertain af- lar-funnelform, nocturnal, white or pink; scales fleshy;
finity, endemic to SW USA (S California) and ad- areoles of pericarpel and tube naked or with a few
jacent NW Mexico (NW Baja California). Natural weak bristles. Fruits ovoid, spiny, dehiscent by verti-
hybrids with species of Myrtillocactus and Pachy- cal slits; pulp non-juicy, white. Seven species, endemic
cereus have been described. to Mexico, perhaps better combined with Carnegiea.

68. Pachycereus (A. Berger) Britton & Rose 71. Cephalocereus Pfeiffer
Pachycereus Britton & Rose, Contrib. U.S. Nat!. Herb. 12: 420 Cephalocereus Pfeiffer, Allg. Gartenz. 6: 142 (1838).
(1909). Pilocereus Lemairc, (1839), as to type.
Lemaireocereus Britton & Rose (1909), as to type. Haseltonia Backeberg (1949).
Lophocereus (A. Berger) Britton & Rose (1909). Neodawsonia Backeberg (1949).
Marginatocereus Backeberg (1942).
Mitrocereus Backeberg (1942).
Backebergia Bravo (1953).
Columnar; stems unbranched or few-branched at
?Anisocereus Backeberg (1938). base, erect, to 10-12 m, ribbed; ribs 12-30 or more.
? Pterocereus MacDougall & Miranda (1954). Flowering areoles different for the non-flowering,
? Pseudomitrocereus Bravo & F. Buxbaum (1961). usually with numerous hair-like bristles. Flowers me-
dium-sized, nocturnal, tubular-campanulate; pericar-
Tree-like or shrubby, often massive; stems stout, pel and tube with minute scales; areoles of pericarpel
erect. Non-flowering areoles spiny; flowering areoles and tube with sparse wool and short hairs; stamens
similar to the non-flowering or often dissimilar, con- numerous, inserted in the tube and throat, the lower-
 Cactaceae 191

most arising from a rim nearly covering the nectar- 75. Escontria Britton & Rose
chamber. Fruit ovoid, with small scales; floral rem- Escontria Britton & Rose, Contrib. U.S. Natl. Herb. 10: 125
nant persistent. (1906).
Now restricted to about three spp., endemic to
Mexico. Most of the species formerly included in Tree-like, to 4 m; stems ribbed, spiny; ribs 7-8.
Cephalocereus are now referred to Pilosocereus Flowers subapical, tubular-campanulate, 3 cm, diur-
Byles & Rowley (Pilocereus Schumann, non nal; tube short, with prominent imbricate, charta-
Lemaire). The relationships of those that remain are ceous scales; tepals pale yellow. Fruits globose,
debatable. purple-brown, fleshy. A single species, endemic to
S Mexico.
72. Stenocereus Riccobono
Stenocereus Riccobono, Boll. R. Ort. Palermo 8 [pub!, 191O?]: 253
76. Myrtillocactus Console
(1909); Gibson, Cact. Succ. 1. (U. S.) 60: 11-16 (1988), et seq.
Lemaireocereus Britton & Rose (1920), in part, exc!. typo Myrtillocactus Console, Boll. R. Ort. Bot. Palermo 1: 8 (1897).
Machaerocereus Britton & Rose (1920).
Isolatocereus Backeberg (1942).
Ritterocereus Backeberg (1942).
Tree-like or shrubby; branches numerous, ascending,
Hertrichocereus Backeberg (1950). few-ribbed, spiny. Flowers diurnal, small, up to 9 per
Marshallocereus Backeberg (1950). areole; scales small; areoles of pericarpel and tube
slightly woolly; tube very short; perianth rotate;
Tree-like or shrubby, sometimes thicket-forming or stamens relatively few. Fruits small, globose, berry-
creeping; stems stout, ribbed, often heavily spined; like, fleshy. Four closely related spp., Mexico and
flowering areoles usually discrete. Flowers funnel- Guatemala.
form or campanulate, usually nocturnal; pericarpel
with numerous areoles and usually small spines; tube
flared or not, with decurrent scales. Fruits globose or 9. Tribe Cacteae
ovoid, fleshy, often with deciduous spines. About
25 spp., Mexico, C America, the West Indies, Vene- Mostly globular, small to large, rarely short columnar;
zuela and Colombia. rootstock sometimes napiform; stems unsegmented,
ribbed, tuberculate-ribbed or tuberculate; areoles
73. Rathbunia Britton & Rose tending to be oval, bandlike and more or less
Rathbunia Britton & Rose, Contrib. U.S. Nat!. Herb. 12: 414
grooved, or bipartite (the abaxial and adaxial por-
(1909). tions disjunct), the abaxial portion producing only
spines, the adaxial vegetative and fertile. Fertile zone
Sprawling or thicket-forming shrubs; stems ribbed, undifferentiated (but the flowering areoles con-
very spiny. Flowers narrowly tubular, zygomorphic, tiguous or confluent in Echinocactus spp.). Flowers
diurnal, red; areoles of pericarpel slightly felted and subapical, small to medium-sized, always diurnal
with a few bristly spines; tube elongate, its areoles (closing at night), often brightly coloured, regular,
usually spineless, perianth somewhat oblique; tepals rarely zygomorphic; pericarpel and tube scaly, or
short, spreading or reflexed; stamens numerous, ex- pericarpeJ naked; areoles of pericarpel and tube
serted. Fruits globose, with few deciduous, spines. usually naked, rarely with bristles and hairs. Fruit
A genus of two species confined to NW Mexico. berry-like, juicy, indehiscent, or soon dry, variously
These are Stenocereus spp. with flowers adapted to dehiscent or disintegrating. Seeds broadly oval to al-
bird pollination, but cannot be included in that genus most circular, very small to very large, rarely rugose;
until the generic name Stenocereus is conserved. cell boundaries straight or variously undulate, inter-
stices undifferentiated, relief flat to low-domed or
low- to high-conical, or concave or par-concave
74. Polaskia Backeberg ("pitted"), microrelief none to micropapillate, cuticle
Polaskia Backeberg, Blatter f. Sukkulentenkunde 1: 4 (1949). rarely weakly striate; hilum and micropyle disjunct or
Heliabravoa Backeberg (1956). rarely conjunct; unappendaged or rarely strophiolate.
Distribution: USA and Mexico, extending to S Ca-
Tree-like, 4-5 m; stems ribbed, spiny; ribs 7-12. nada, Caribbean region, Venezuela and Colombia.
Flowers small, urceolate or campanulate, diurnal,
creamy white or pale pink. Fruits globose, 2-4 cm,
red, juicy, edible. Two spp., S Mexico, apparently
linking Myrtillocactus and Stenocereus.
192 Cactaceae

77. Echinocactus Link & Otto scales pointed; areoles of pericarpel and tube woolly.
Echinocactus Link & Otto, Verh. Ver. Befbrd. Gartenb. Preuss. Fruit globose, with persistent scales and wool; inte-
Staaten 3: 420 (1827). rior almost dry. Seeds hat-shaped, ca. 2.5 x 2 mm,
Echinofossulocactus Lawrcnce (1841), in part, incl. lectotype. black-brown, shiny, the testa expanded and inrolled
Homalocephala Britton & Rose (1922). the sunken hilum. Four spp., E Mexico and SW USA
(S Texas).
Discoid, globose or shortly columnar cacti; stems
strongly ribbed; areoles large, elongate, discrete, or
confluent in adult plants of typical species; forming a 80. Sclerocactus Britton & Rose
broad woolly crown at stem apex; spines stout; nec- Sclerocactus Britton & Rose, The Cactaceae 3: 212 (1922).
tar-secreting glands absent. Flowers apical, shortly Echinomastus Britton & Rose (1922).
funnelform to campanulate; pericarpel and tube Toumeya Britton & Rose (1922).
short with numerous, narrow, pointed scales; areoles Ancistrocactus Britton & Rose (1923).
Glandulicactus Backeberg (1938).
of pericarpel and tube desely woolly. Fruits turbinate Coloradoa Boissevain & Davidson (1940).
to oblong-clavate, with narrow scales and dense wool;
juicy pulp or nearly dry, indehiscent or dehiscing
Low-growing cacti, usually unbranched; stems small,
basally. Seeds large, broadly oval to almost circular,
depressed-globose to clavate or cylindric, tubercu-
2.8-4.0 x 1.9-2.9 mm, brown or black-brown, shiny or
late-ribbed, spiny; areoles more or less extended
dull, smooth or tuberculate, rarely rugose or keeled;
above the spine-bearing part, often with nectar-se-
appendages none. Now restricted to six spp., SW
creting glands. Flowers apical, shortly funnelform or
USA and Mexico. The genus formerly accommo-
campanulate; areoles of pericarpel and tube naked.
dated nearly all cacti with ribbed stems other than
Fruits ovoid to cylindric, clavate or barrel-shaped,
those assigned to Cereus or Melocactus.
usually scaly, either fleshy to juicy and indehiscent, or
becoming dry and opening irregularly at or towards
78. Geohintonia Glass & Fitz Maurice base or by means of 1-3 vertical splits in the pericarp;
floral remnant persistent. Seeds broadly oval, 1.5-
Geohintonia Glass & Fitz Maurice, Cact. Suc. Mex. 37(1): 16-19
('1992', publ. 1991), Cact. Succ. J. (U.S.) 64: 141-147 (1992).
4.0 x 1.0--3.8 mm, brown or black-brown, shiny or
matt, periphery keeled or not; relief flat to low-
domed or high-conical; hilum medium to large, basal
Globose to shortly columnar, to 10(-20) x 11 cm,
or oblique, superficial or impressed, appendages
strongly ribbed; ribs 18-20; areoles large, discrete,
none. About 15-18 spp. in N Mexico and SW USA
initially very woolly with long trichomes, later nearly
broadly united by flower-, fruit- and seed morpho-
glabrous; spines usually 3, to 15 mm, curved, flattened,
logy.
soon papery and brittle, deciduous. Flowers apical,
funnelform, ca. 2 x 2-4 cm; pericarpel and lower part
of tube naked, white; scales of upper tube small, acute, 81. Pediocactus Britton & Rose
densely hairy in the axils with white trichomes to ca.
Pediocactus Britton & Rose, Britton & Brown, Ill. Flowers
1 cm; tepals deep pink to magenta; stigmas 5-6, 2 mm, ed.2, 2: 569 (1913); Heil, Armstrong, & Schleser, Cact. Succ.
yellowish white. Fruit hidden in apical wool, ca. 9 x 1. (U.S.) 53: 17-39 (1981).
4-5 mm, naked, thin-walled, pale, drying and break- Utahia Britton & Rose (1922).
ing off below the middle; seeds broadly oval, Navajoa Croizat (1943).
1.2 x 0.7 mm, black-brown, shiny, relief flat to low- Pilacanthus B. W. Benson & Backeberg (1957).
domed; hilum large, basal, superficial. Monotypic;
endemic on gypsum outcrops in NE Mexico. Dwarf or low-growing cacti, simple or clustering;
stems tuberculate. Flowers from near the stem-apex,
campanulate; pericarpel more or less naked; tube
79. Astrophytum Lemaire very short, scaly. Fruits mostly top-shaped, nearly
Astrophytum Lemaire, Cact. Gen. Nov. Sp. Hort. Monv: 3 naked, dull-doloured, the dried perianth partly de-
(1839). ciduous leaving a cap which opens like a lid as the
pericarp splits vertically. Seeds broadly oval, 2.3-
Hemispheric, globose or shortly columnar cacti; 2.9(-5.0) x 1.0-2.1(-3.5) mm, black-brown, matt, ru-
stems simple, densely to sparsely covered with gose or not, keeled or not; relief low- to high-domed;
minute tufts of whitish trichomes, or rarely glabrous, hilum medium, basal or oblique, impressed. Six spp.,
green; ribs 4-10; areoles large, sometimes close-set, confined to Wand SW USA (Colorado Plateau,
but distinct; spines present or absent. Flowers apical, Columbia River and Great Basin, and the Rocky
shortly funnelform, yellow or yellow with red throat; Mountains ).
 Cactaceae 193

82. Thelocactus Britton & Rose 84. Neolloydia Britton & Rose
Thelocactus Britton & Rose, Bull. Torrey Bot. Club 49: 251 Neolloydia Britton & Rose, Bull. Torrey Bot. Club 49: 251
(1922); Anderson in Bradleya 5: 49-76 (1987). (1922); Anderson, Bradleya 4: 1-28 (1986).
Hamatocactus Britton & Rose (1922). Turbinicarpus (Backeberg) E Buxbaum & Backeberg (1937).
Gymnocactus Backeberg (1938).
Low-growing cacti; simple or clustering; stems de- Rapicactlls E Buxbaum & Oehme (1942).
Normanbokea Kladiwa & E Buxbaum (1969).
pressed-globose to short-cylindric, ribbed or tubercu-
late; areoles with or without nectar-secreting glands,
sometimes extending at the upper edge to form a Low-growing or dwarf cacti, simple or clustering;
short groove; spines few to numerous. Flowers apical, rootstock fibrous to napiform; stems depressed-glo-
funnelform, small to large; scales of pericarpel and bose to short cylindric, tuberculate; areoles usually
tube conspicuous. Fruits globose, dark green or entire, apical, rarely bipartite, with apical and axillary
tinged brown, dry and dehiscent by a basal pore, or portions connected by a groove (N. conoidea); spines
rarely bright red, somewhat fleshy and indehiscent. various but never hooked. Flowers mostly small,
Seeds broadly oval, 1.3-2.5 x 1.0-1.5 mm, black- short funnelform, at upper edge of areole (axillary in
brown, shiny or not, keeled; relief flat, concave or N. conoidea); pericarpel naked or rarely with 1-2
par-concave; hilum medium to large, basal or ob- small scales; tube narrow, short. Fruits globose to tur-
lique, impressed, appendages none. Eleven variable binate, naked or almost so, dry and papery when ma-
species, C and N Mexico and SW USA (Texas). ture (N. conoidea) or more or less fleshy, splitting ver-
tically or disintegrating to release the seeds; floral
remnant caducous (N. conoidea) or persistent. Seeds
83. Coryphantha (Engelmann) Lemaire nearly circular to broadly oval, 1.2-1.9 x 0.9-1.3 mm,
black-brown or brown, matt, not wrinkled or ridged,
Coryphantha Lemaire, Les Cactees: 32 (1868).
Lepidocoryphantha Backebcrg (1938).
periphery keeled or not; relief low- to high-domed;
Cumarinia EBuxbaum (1951). hilum medium to large, basal or oblique, superficial
or impressed.
Mostly globose cacti; stems simple or clustering, glo- As amplified by Anderson (I.c.), the genus con-
bose to cylindric, to 50 cm, tuberculate; tubercles tains 14 species native to E and NE Mexico and
terete, cylindric, gibbous or pyramidal, grooved SW Texas. As noted in the description, however,
above except in seedlings and juvenile plants; are- there are significant differences between the type
oles of juvenile plants either apical only, later with a species (N. conoidea) and the remainder, and it may
rudimentary groove (subg. Coryphantha) or two- be preferable to uphold Turbinicarpus as a separate
parted (apical and "axillary") but not connected by a genus. Many of the species are semi-geophytic in
groove (subg. Neocoryphantha Backeberg); spines nature.
normally developed, or in some species some of the
spines modified into coloured glands. Flowering are- 85. Mammilloydia F. Buxbaum
oles two-parted, consisting of an apical (abaxial),
spine-bearing but sterile part and an "axillary" (adax- Mammilloydia E Buxbaum, Bot. Stud. 12: 64, 65 (1951).
ial) fertile part, connected by the areolar groove;
flowers funnelform or campanulate, up to 6.5 x Low-growing; stems depressed-globose to shortly
10 cm; pericarpel naked or with small, sparse scales cylindric, simple or clustering, to ca. 14 cm diam.,
only. Fruit globose to ovoid, oblong or clavate, tuberculate; tubercles broadly cylindric, obtuse, not
naked, green or yellowish, juicy, indehiscent; floral grooved; areoles bipartite, as in Mammillaria; spines
remnant persistent. Seeds oval to broadly oval, ca. 1- numerous in several series, white, sometimes pinkish
2 x 1-1.5 mm, brown or rarely black-brown (c. odo- or brown-tipped. Flowers arising singly at "axillary"
rata), shiny or dull. About 45 spp., SW USA, Mexico. areoles, ca. 2-3 cm, pinkish; pericarpel and tube
The plants are superficially like Mammillaria, but naked. Fruits cylindric, pink, juicy, indehiscent, the
have grooved tubercles and different seeds. Also, the withered perianth deciduous. Seeds ca. 1 mm, black,
flowers are usually much larger, but less freely pro- relief flat or domed (not pitted). A single species,
duced, and are accompanied in some species by endemic to NE Mexico. Closely resembling Mam-
coloured, nectar-secreting glands. millaria, but almost certainly convergent, as B ux-
baum argued.
194 Cactaceae

86. Epithelantha Britton & Rose 89. Obregonia Fric

Epithelantha Britton & Rose, The Cactaceae 3: 92 (1922); Glass Obregonia Fric, Zivot v Prirode 29(2): 1-4 (1925).
& Foster, Cac!. Succ. 1. (U.S.) 50: 184-187 (1978).
Low-growing cacti; stem mostly simple, depressed, 5-
Dwarf species; stems globose to obovoid, simple or 20(-30) cm diam., pale green or tinged brown, tuber-
clustering, 1-6 cm diam., tuberculate; tubercles culate, becoming tuberous at base and with carrot-
minute, often obscured by spines; spines 19-38, like taproot; tubercles 5-15 x 7-15 mm, triangular,
1-8 mm, white to pale grey or yellowish, the upper- arranged as in a rosette, acuminate; areole apical;
most clavate, functioning as glands at first, the tip spines 0-5, to 15 mm, pale, weak, soon deciduous.
later deciduous. Flowers arising in the woolly stem- Flowers in woolly stem-apex, 2-3.6 x 2.5 cm, almost
apex, campanulate, 3-12 mm diam., almost white or white; pericarpel and tube well-developed, naked;
pale orange to pink; pericarpel and tube naked; tepals tepals narrow, 1-1.5 mm wide; stamens sometimes
and stamens few; stigma-lobes mostly 3(-4). Fruits sensitive as in Lophophora; filaments pink. Fruits 16-
3-18 mm, clavate-cylindric, red, rather juicy, few- 25 x 3-6 mm, clavate, white, fleshy then dry; floral
seeded, indehiscent, the withered perianth deciduous. remnant deciduous. Seed almost circular, ca.
Seeds oval, 1-1.5 x ca. 1 mm, black-brown, shiny, 1.3 x 1.5 mm, black-brown, shiny; relief low-domed;
relief low-domed; hilum medium-sized, basal, im- hilum medium, oblique, impressed. A monotypic
pressed. One to two spp., SW USA and NE Mexico. genus, endemic to NE Mexico.

87. Ariocarpus Scheidweiler 90. Strombocactus Britton & Rose

Ariocarpus Scheidweiler, Bull. Acad. Sci. Brux. 5: 491 (1838). Strombocactus Britton & Rose, The Cactaceae 3: 106 (1922).
Roseocactus A. Berger (1925).
Neogomesia Castaneda (1941). Low-growing cacti; simple; stems depressed to glo-
bose or turbinate, tuberculate, 2-8 x 3-17 cm; tuber-
Slow-growing, often semi-geophytic; stem-base and cles spiralled, irregularly rhomboid, pale grey-green;
roots stout; stems usually unbranched, tuberculate; areoles soon losing their wool; spines weak, 1-5, to
tubercles spiralled or rosulate, more or less triangu- 15 mm, erect, off-white, darker near apex, often early
lar, short or elongate; areoles bipartite, varied in deciduous. Flowers apical, short-funnelform, to
structure; spines usually absent. Flowers short-fun- 3.2 x 3.2 cm, pale yellow to almost white with red
nelform; tube naked; perianth-limb spreading, pink, throat; scales few, green- or red-tinged; stigmas 7-11.
purple, pale yellow or white. Fruit inconspicuous, Fruits 7-10 x 6-7 mm, red-brown to green, dehiscing
drying and disintegrating in situ. Seeds broadly oval, by 2-4 longitudinal splits. Seeds very small, almost
1.5-2 x ca. 1.5 mm, black, tuberculate. Six spp., N globose, 0.4-0.5 x 0.3-0.4 mm, brown, shiny; relief
and E Mexico and SW USA (S Texas). low-conical; hilum medium, basal, superficial; stro-
phiole present, large. Monotypic; endemic to central
88. Lophophora 1. Coulter E Mexico.

Lophophora 1. Coulter, Contrib. U.S. Natl. Herb. 3: 131 (1894);

Anderson, E. F. (1979). 91. Aztekium Bodeker
Aztekium Bbdeker, Monatsschr. Deutsch. Kak!. Ges. 1: 52
Low-growing, almost geophytic cacti; rootstock napi- (1929); Glass & Fitz Maurice in Cac!. Succ. J. (U.S.) 64: 141-
form or carrot-like; stems depressed-globose, usually 147 (1992).
clustering, weakly tuberculate-ribbed, spineless.
Flowers arising in the densely woolly stem apex, cam- Low-growing; stems up to 20 x 10 cm, simple or clus-
panulate, self-fertile; pericarpel and tube naked; tering, ribbed, weakly spined, pale grey-green; ribs 8-
stamens sensitive, closing around the style when 15, transversely wrinkled and with longitudinal rib-
touched. Fruits cylindric to clavate, pink or red, like ridges in between; areoles contiguous, woolly;
naked, juicy when ripe, but soon drying; floral rem- spines 1-3, to 13 mm, upward-curved and appressed,
nant deciduous. Seeds broadly oval, 1.5 x 1.1 mm, flattened, yellowish to grey, brittle deciduous. Flow-
black-brown, matt; relief low-domed; hilum large, ers apical, funnelform, c. 12 x 30 mm; tepals whitish to
basal, impressed. Two spp., E and N Mexico and SW purplish pink; pericarpel and tube slender, naked;
USA (S Texas). stigmas c. 4. Fruits hidden in apical wool, berry-like, c.
3-8 mm. Seeds very small, c. 0.7 x 0.5 mm, brown,
shiny, tuberculate; strophiole present, large. A genus
of 2 species endemic to NE Mexico.
 Cactaceae 195

92. Pelecyphora Ehrenberg 95. Stenocactus (Schumann) A. W. Hill

Pelecyphora Ehrenberg, Bot. Zeit. 1: 737 (1843). Stenocactus Hill, Index Kewensis, supp!. 8: 228 (1933).
Encephalocarpus Berger (1929). Echinofossulocactus Lawrence (1841), in part, exc!. lectotype,
and scnsu Britton & Rose (1922).
Low-growing; stems simple or clustering, globose or
obconic, tuberculate; tubercles hatchet-shaped and Low-growing cacti, usually unbranched, allied to
laterally compressed, or scale-like and dorsiventrally Ferocactus; stems subglobose to short cylindric,
compressed and then acute, keeled and somewhat in- ribbed; ribs acute, usually very numerous, thin and
curved; areoles bi-partite, the vegetative, spine-bear- often sinuate (except in S. coptonogonus); areoles
ing part abaxial, at the tubercle-apex, the woolly usually widely spaced; spines commonly differen-
flowering part adaxial, in the tubercular "axil", the tiated into a large upper series (central spines) and
two parts interconnected by a narrow groove or shal- small lower series (radials), the former often strongly
low corky ridge. Flowers subapical, shortly funnel- flattened. Flowers small, shortly funnelform or cam-
form or campanulate; tube naked; perianth purple- panulate; pericarpel with scales; areoles of pericarpel
pink. Fruits small, dry. Seeds almost circular, ca. and tube naked; tube usually short. Fruits small,
1.2 x 1.0 mm, brown, matt; relief par-concave; hilum mostly globose, pale green, scaly, dry, dehiscent at one
small, oblique, superficial, appendages none. Two side. Seeds broadly oval, 1.4 x 1.3 mm, black-brown,
species, endemic to NE Mexico. shiny; relief concave; hilum medium, basal, impressed,
appendages none. About ten spp., native to Mexico.
93. Ferocactus Britton & Rose
Ferocactus Britton & Rose, The Cactaceae 3: 123 (1922); Tay-
96. Escobaria Britton & Rose
lor, Bradleya 2: 19-38 (1984). Escobaria Britton & Rose, The Cactaceae 4: 53 (1923); Taylor
Bisnaga Orcutt (1926). in Brit. Cact. Succ. J. 4: 36-44 (1986).
Neobesseya Britton & Rose (1923).
Barrel-cacti, unbranched or caespitose; stems often Cochiseia Earle (1976).
large or very large, depressed-globose to cylindric,
ribbed; ribs few to numerous; areoles with nectar- Low-growing cacti, simple or clustering; stems de-
secreting glands; spines often fierce. Flowers short- pressed-globose to cylindric, tuberculate; areoles
funnelform, funnelform or campanulate; scales con- elongate (except in sect. Acharagma N. P. Taylor), ex-
spicuous; areoles of pericarpel and tube naked; tepals tending via a groove from the tubercle-apex to its
and stamens separated by a ring of hairs. Fruits glo- axil; spines present; nectar-secreting gland-spines ab-
bose to oblong, thick-walled, dry and dehiscent at sent. Flowers arising at upper (adaxial) edge of are-
base (sect. Ferocactus), or juicy and indehiscent or oles and thus between the tubercles (at their tip in
splitting irregularly (sect. Bisnaga). Seeds broadly sect. Acharagma); pericarpel naked; tube short; outer
oval, 1.4-2.4 mm, black-brown, shiny, relief flat to tepals usually ciliate-margined. Fruits berry-like,
slightly concave or almost par-concave ("pitted"). ovoid, shortly clavate or cylindric, naked or with one
Twenty three spp., SW USA, Mexico. or more small scales near apex, green, pink or red.
Seeds broadly oval to almost circular, 1.0-1.7 mm,
brown or black-brown, matt, usually deeply pitted
94. Leuchtenbergia Hooker (par-concave), only slightly so in sect. Pleurantha).
Leuchtenbergia Hooker, Curtis's Bot. Mag. 74: t.4393 (1848). About 15 spp. in W USA, S Canada, N Mexico and
Cuba. Related to Mammillaria, but considered to be
Low-growing, rarely to 70 cm high, simple or occa- less highly evolved, with the areoles not developing
sionally clustering; rootstock fleshy; stem globose to discrete spiniferous and flowering portions, and the
short -cylindric, tuberculate; tubercles 10-12 cm, glau- flowers arising mostly near the stem-apex. Some
cous, triangular; areoles apical; spines to 15 cm, authors combine the genus with Coryphantha, which
papery, flattened and flexuous. Flowers borne at the similarly has grooved tubercles, but it seems more
ventral edge of areoles of young tubercles, otherwise likely that the groups are convergent.
like those of Ferocactus, to 8 x 5-6 cm, yellow. Fruits
as in Ferocactus subg. Ferocactus, ovoid-oblong, ca.
97. Ortegocactus Alexander
3 cm, excluding perianth, grey-green. Seed broadly
oval, 2.4 x 2.0 mm, black-brown, matt, periphery Ortego cactus Alexander, Cact. Suce. J. (U.S.) 33: 39 (1961).
keeled; relief low-domed; hilum small, basal, deeply
impressed. A single aberrant species, endemic to Low-growing; stems clustering, reminiscent of M.
NMexico. schumannii, globose to short-cylindric, 3-4 cm diam.,
196 Cactaceae

pale grey-green, tuberculate; tubercles low, rhom- 3S0 species. Many of these are only varieties or local
boid, 10-12 mm diam., minutely punctate, the flower- forms. As argued by Buxbaum, the seed structure of
ing grooved or not; areoles bipartite as in Mammilla- M. candida indicates that its overall resemblance to
ria, the flowering sometimes grooved between the Mammillaria is due to convergence, and his genus
abaxial vegetative part and the adaxial floriferous Mammilloydia is here upheld (see no.8S). Hunt
part, as in Escobaria; spines, at least the tip, almost (1971 etc) recognises a further four small subgenera
black; central spine 1, 4-S mm; radial spines 7-8, (Oehmea, Dolichothele, Cochemiea, Mamillopsis)
S-10 mm. Flowers axillary, diurnal, funnelform, and divides subg. Mammillaria into three sections
2-3 x 1.8-2.5 cm; pericarpel without scales, but im- and several series.
mersed in areolar wool; tube short, pale green; tepals
yellow, or the outermost tinged purple outside;
stigmas deep green. Fruits globose-ellipsoid, dull red, Selected Bibliography
soon dry; floral remnant persistent. Seed almost glo-
bose, 0.8-0.9 x 0.7 mm, black-brown, matt; relief par- Anderson, E. F. 1979. Peyote, the divine cactus. Tucson: Univ.
concave; hilum large, basal, superficial. A single Arizona Press.
species of restricted distribution SE Mexico (Oax- Backeberg, C. 1958-62. Die Cactaceae. 6 Vols. J ena: G. Fischer.
Backeberg, C. 1966. Das Kakteenlexikon. Jena: G.Fischer.
aca). It should probably be included in either Esco-
Backeberg, C. 1977. Cactus lexicon. English cd., trans I. L. Glass,
baria or Mammillaria. Poole: Blandford Press.
Barthlott, W. 1983. Biogeography and evolution in neo- and
98. Mammillaria Haworth palaeotropical Rhipsalinae. In: Kubitzki, K (Ed.) Dispersal
and distribution. Sonderbd. naturbd. naturwiss. Ver. Ham-
Mammillaria Haworth. Syn. PI. Succ: 177 (1812); Craig, The burg 7. Hamburg: P.Parey, pp.241-248.
Mammillaria handbook (1945); Hunt, Cact. Succ. J. Gr. Brit. Barthlott, W 1988. Uber die systematischen Gliederungen der
3: 53-72 (1971) and Bradleya 1: 105-128 (1983), 2: 65-96 Cactaceae. Beitr. BioI. Pflanz. 63: 17-40.
(1984),3: 53-66 (1985),4: 39-64 (1986). 5: 17-48 (1987). Barthlott, W, Hunt, D. 1992. Micromorphology and descriptive
Cactus Linnaeus (1753), in part, as to type. terminology of the seeds of Cactaceae. in prep.
Cochemiea Walton (1899). Barthlott, W, Voit, G. 1979. Mikromorphologie der Samen-
Bartschella Britton & Rose (1923). schalen und Taxonomie der Cactaceae. PI. Syst. Evol. 132:
Dolichothele Britton & Rose (1923). 205-229.
Mamillopsis Britton & Rose (1923). Behnke, H.-D. 1981. Sieve-element characters. Nord. J. Bot. 1:
Neomammillaria Britton & Rose (1923). 381-400.
Phellosperma Britton & Rose (1923). Benson, L. 1982. The cacti of the United States and Canada.
Solisia Britton & Rose (1923). Stanford: Stanford University Press.
Chilita Orcutt (1926). Berger, A 1926. Die Entwicklungslinien der Kakteen. Jena:
Porfiria Boedeker (1926). G.Fischer.
Krainzia Backeberg (1938). Boke, N. H. 1980. Developmental morphology and anatomy in
Oehmea Buxbaum (1951). Cactaceae. Bioscience 30: 605-610.
Pseudomammillaria F. Buxbaum (1951). Bravo-Hollis, H., Sanchez-Mejorada, H. 1978-91. Las Cacta-
Leptocladodia F. Buxbaum (1954). ceas de Mexico, 2nd edn. 3 Vols. Mexico: Universidad Nacio-
nal
Low-growing cacti, simple or clustering, sometimes Britton, N. L., Rose, 1. N. 1919-23. The Cactaceae. 4 Vols.
forming mounds to 1 m across, sometimes almost Washington, D.C.: Carnegie Institution.
geophytic; stems depressed-globose or globose to Buxbaum, F. 1950-54. Morphology of cacti. Parts I-III. Pasa-
dena: Abbey Garden Press.
cylindric, tuberculate, some species laticiferous;
Buxbaum, F. 1957-60. Morphologie der Kakteen. In: Krainz, H.
tubercles terete, conic, pyramidal or gibbous, not (Ed.) Die Kakteen. Stuttgart: Franckh.
grooved; areoles bipartite, as in Coryphantha and Buxbaum, F. 1958. The phylogenetic division of the subfamily
Escobaria, but with no interconnecting groove; dor- Cereoideae, Cactaceae. Madrofio 14: 177-206.
sal (distal) portion of areoles at tubercle-apex, spiny, Eggli, U., Taylor, N. P. 1991. Index of names of Cactaceae pub-
normally non-flowering; ventral portion "axillary", lished 1950-1990. Royal Botanic Gardens Kew.
Endler, J., Buxbaum, F. 1974. Die Pflanzenfamilie der Kakteen.
naked, felted, hairy, or with bristle-spines; gland-
Minden: A. Philler.
spines absent. Flowers arising singly at "axillary" are- Gibson, A.C., Horak, K.E. 1979. Systematic anatomy and phy-
oles, mostly small, campanulate to funnelform; peri- logeny of Mexican columnar cacti. Ann. Mo. Bot. Gard. 65:
carpel naked; tube usually short. Fruit berry-like, 999-1057.
oblong or clavate, often bright red, rarely immersed Gibson, A. c., Nobel, P. S. 1986. The cactus primer. Cambridge
in stem. Seeds diverse; relief par-concave. (Mass.): Harvard Univ. Press.
Hegnauer, R. 1964, 1989. Chemotaxonomie der Pflanzen, Vol. 3
A large genus of perhaps lS0-200 spp. in SW USA, (1964), Vol. 8 (1989). Basel: Birkhauser.
Mexico, C America, the Caribbean region, Colombia Hunt, D. R. 1967. Cactaceae. In: Hutchinson, J. (Ed.) The gen-
and Venezuela. The genus has long been popular era of flowering plants, Vo1.2. Oxford Univ. Press, pp.427-
with collectors, and horticultural treatments list 2SO- 467.
 Calycanthaceae 197

Hunt, D, R. 1992. CITES Cactaceae Checklist. Kew: Royal Bo-

tanic Gardens and International Organization for Succulent
Calycanthaceae
Plant Study (lOS).
K.KUBITZKI
Hunt, D., Taylor, N. (Eds.) 1986. The genera of the Cactaceae:
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Hunt, D., Taylor, N. (Eds.) 1990. The genera of Cactaceae: pro-
gress towards consensus. Bradleya 8: 85-107.
Hunt, D., Taylor, N. (Eds.) 1991. Notes on miscellaneous genera
of Cactaceae. Bradleya 9: 81-92.
Hunt, D. R. et al. 1989. Cactaceae. In: Walters, S. M. et al. (Eds.)
The European garden flora, Vol. 3. Cambridge: Cambridge
Calycanthaceae Lind!., Bot. Reg. 5: sub t. 404 (1819) ("Calycan-
Univ. Press.
theae"), inc!. Idiospermaceae S. T. Blake (1972).
Krainz, H (Ed.) 1956-76. Die Kakteen. Issued in fascicles.
Stuttgart: Franckh.
Leins, P., Schwitalla, S. 1985. Studien an Cactaceen Bliiten 1. Shrubs or trees with aromatic bark and ethereal oil
Beitr. BioI. Pflanz. 60: 313-323. cells in parenchymatous tissue; evergreen or deci-
Leins, P., Schwitalla, S. 1988. Placentation in Cactaceae. In:
Leins, P., Tucker, S. C, Endress, P. K. (Eds.) Aspects of floral
duous; hairs unicellular; leaves opposite, simple, en-
development. Berlin: 1. Cramer, Borntraeger, pp. 57--68. tire; stipules absent; flowers solitary, or in few-flower-
Leuenberger, B. E. 1976. Die Pollenmorphologie der Cactaceae ed terminal inflorescences, perfect, perigynous, with
und ihre Bedeutung flir die Systematik. Diss. Bot. 31. Vaduz: a cup-shaped or urceolate receptacle; all floral
J.Cramer. phyllomes arranged in a continuous spiral; perianth
Madsen, J.E. 1989. Cactaceae. In: Harling, G., Andersson, L. of 15-40 petaloid tepals spirally attached to the out-
(Eds.) Flora of Ecuador, fasc. 35. Copenhagen: Nord. J. Bot.
79pp.
side of the receptacle; stamens 5-30, inserted on the
Matta, R, McLaughlin, J. L. 1982. Cactus alkaloids. A compre- rim of the receptacle; filaments short or absent; pol-
hensive tabular summary. Rev. Latinoameric. Quim. 12: 95- len sacs 2, abaxial, extrorse, opening by longitudinal
117. slits; connectives prolonged beyond the pollen sacs;
Mauseth, J.D. 1982-84. Introduction to cactus anatomy, staminodes numerous, lining the inner surface of the
parts 1-11. Cact. Succ. J (U.S.) 54: 263-266. 55: 18-21,42,84- receptacle; gynoecium of one to few or of many free
89, 113-118, 171-175, 272-276. 56: 33-37, 131-135,181-184,
212-216,250-255.
carpels; ovule solitary, basal, anatropous, crassinucel-
Nobel, P.S. 1988. Environmental biology of agaves and cacti. lar, bitegmic, sometimes a second, upper, aborting
Cambridge: Cambridge Univ. Press. ovule present in each carpel. Fruit or fruits indehis-
Pink ava, D.J., Baker, M. A., Parfitt, B. D., Mohlenbrock, M. w., cent, dry, enclosed in the receptacle; seeds poisonous,
Worthington, R.D. 1985. Chromosome numbers in some exalbuminous; cotyledons 2-4, massive, or spirally
cacti of Western North America (V). Syst. Bot. 10: 471-483. twisted.
Porsch, O. 1938-39. Das Bestaubungsleben der Kakteenbliite.
lahrb. Deutsch. Kakteenges. 1938/1,1939/1. Neudamm: Neu-
A family comprising four genera and about eight
mann. species, confined to temperate China, temperate N
Rauh, W. 1958. Beitrag zur Kenntnis der peruanischen Kak- America and tropical NE Australia.
teenvegetation. Sitz. Ber. Heid. Akad. Wiss. Berlin Heidel-
berg New York: Springer. VEGETATIVE STRUCTURES. The primary vascular cy-
Rauh, W. 1979. Kakteen an ihren Standorten. Berlin: Parey.
linder of the Calycanthaceae is notable for posses-
Ritter, F. 1979-81. Kakteen in Siidamerika, 4 Vols. Spangen-
berg: Selbstverlag. sing a peculiar system of four inverted cortical
Ross, R. 1981. Chromosome counts, cytology, and reproduction bundles. It was studied by Fahn and Bailey (1957)
in the Cactaceae. Amer. J. Bot. 68: 463-470. and by Wilson (1979), who described the nodal
Schumann, K.M. 1897-99. Gesamtbeschreibung der Kakteen. anatomy as of the double-trace unilacunar type.
Neudamm: Neumann. Secretory cells are abundantly present. The stomata
are rubiaceous; the hairs are unicellular. A compre-
hensive study of the wood anatomy was undertaken
by Carlquist (1983). The vessels, as seen in transsec-
tion, are arranged in diagonal bands (Calycanthus,
Chimonanthus), in radial chains (Idiospermum), or
solitary. Common features include simple perfora-
tion plates, tyloses, paratracheal and apotracheal
parenchyma, uniseriate and multiseriate (mostly bi-
seriate) rays, and fiber tracheids with few, small pits.
Anatomical differences between the genera (and
species) seem to be related to differences in habitat
(Carlquist 1983). Sieve element plastids contain pro-
tein filaments and protein crystals. Behnke (1988)