The Encyclopedia of Fruit & Nuts
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The Encyclopedia of Fruit & Nuts
Edited by
Jules Janick
Department of Horticulture and Landscape Architecture
Purdue University
West Lafayette, Indiana, USA
and
Robert E. Paull
Department of Tropical Plant and Soil Sciences
University of Hawai’i at Manoa
Honolulu, Hawai’i, USA
CABI is a trading name of CAB International
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A catalogue record for this book is available from the British Library,
London, UK.
Library of Congress Cataloging-in-Publication Data
The Encyclopedia of fruit and nuts / edited by Jules Janick and Robert E. Paull.
p. cm.
Includes bibliographical references and index.
ISBN 0-85199-638-8 (alk. paper)
1. Tropical fruit--Varieties--Encylopedias. 2. Tropical nuts--Varieties-Encyclopedias. I. Janick, Jules, 1931- II. Paull, Robert E. III. Title.
SB359.E56 2006
634⬘.603--dc22
2006027763
ISBN 978 0 85199 638 7
Typeset by Columns Design Ltd, Reading, UK
Printed and bound in the UK by Cambridge University Press, Cambridge
Contributors
Akinnifesi, Festus K., World Agroforestry Centre, Southern
African Development Community (SADC) International Centre
for Research in Agroforestry (ICRAF) Programme, Chitedze
Agricultural Research Station, PO Box 30798, Lilongwe, Malawi.
f.akinnifesi@cgiar.org
Allen Stevens, M., 21715 County Road 97, Woodland, CA 95695,
USA
Alves, Rafael Moysés, Empresa Brasileira de Pesquisa
Agropecuária, EMBRAPA Amazônia Oriental, Travessa Eneas
Pinheiro s/n, Laboratório de Fitomelhoramento Marco, 66095100 Belem, Pará, Brazil – Caixa-Postal: 48
Andall, Reginald, Caribbean Agricultural Research and
Development Insititute, St George’s, Grenada
Ang, B.N., Applied Agricultural Research Sendirian Berhad, Buloh,
47000 Selangor, Malaysia
Arce, Rogerene K.M., Department of Tropical Plant and Soil
Sciences, University of Hawai’i at Manoa, 3190 Maile Way,
Honolulu, HI 96822, USA
Atangana, A.R., World Agroforestry Centre (ICRAF), Regional
Office African Humid Tropics, PO Box 2067, Yaoundé, Cameroon
Ballington, James R., Department of Horticultural Science, North
Carolina State University, Raleigh, North Carolina, USA
Bartholomew, Duane, Department of Tropical Plant and Soil
Sciences, University of Hawai’i at Manoa, 3190 Maile Way,
Honolulu, HI 96822, USA
Batlle, Ignasi, Institut de Recerca i Tecnología Agroalimentàries
(IRTA), Centre Mas Bové, Dept. d’Arboricultura Mediterrània,
Apartat 415, 43.280 Reus, Spain
Bayogan, Emma Ruth V., Davao Oriental State College of Science
and Technology, Mati, Davao Oriental, Philippines
Bayuelo-Jiménez, Jeannette S., Department of Plant Nutrition,
Universidad Michoacana de San Nicolás de Hidalgo, Morelia,
México
Bell, Richard L., United States Department of Agriculture,
Agricultural Research Service, Appalachian Fruit Research
Station, Kearneysville, West Virginia, USA
Benzioni, Aliza, The Institutes for Applied Research, Ben-Gurion
University of the Negev, Beer-Sheva, Israel
Bittenbender, H.C., Department of Tropical Plant and Soil
Sciences, University of Hawai’i, 3190 Maile Way, Honolulu, HI
96822, USA
Bosland, Paul W., Department of Agronomy and Horticulture,
New Mexico State University, Las Cruces, NM 88003-0003,USA
Carnelossi, Marcelo A.G., Department of Chemical Engineering,
Federal University of Sergipe, Av. Marechal Rondon s/n, São
Cristóvão, SE 49100-000, Brazil
Casas, Alejandro, Centro de Investigaciones en Ecosistemas,
Universidad Nacional Autónoma de México, Apartado Postal 27-3
(Santa María de Guido) Morelia, Michoacán CP 58089, México
Chan, Ying Kwok, Malaysian Agricultural Research and
Development Institute, PO Box 12301 GPO, 50774 Kuala
Lumpur, Malaysia
Chao, Chih-Cheng T., Department of Botany and Plant Sciences,
University of California-Riverside, Riverside, California, 925210124, USA. ctchao@citrus.ucr.edu
Chaplin, Grantley R., 696 Mt Sylvia Road, Gatton, Queensland,
Australia 4343
Chen, Ching Cheng, Department of Horticulture, National Chung
Hsing University, Kuo Kuang Road, Taichung, Taiwan, Republic
of China
Chen, Kunsong, Department of Horticulture, Huajiachi Campus,
Zhejiang University, Hangzhou, 310029, PR China
Chilanga, T., World Agroforestry Centre, Southern African
Development Community (SADC) International Centre for
Research in Agroforestry (ICRAF) Programme, Chitedze
Agricultural Research Station, PO Box 30798, Lilongwe, Malawi
Clement, Charles R., Instituto Nacional de Pesquisas da Amazônia
(INPA) Cx. Postal 478, Manaus, Amazonas, Brazil
Cohen, Jane E., Department of Life Sciences, University of the
West Indies, Mona, Kingston, Jamaica
Coronel, Roberto E., Institute of Plant Breeding, College of
Agriculture, University of the Philippines Los Baños, College,
Laguna, Philippines
Cousins, Peter, United States Department of Agriculture (USDA)
Agricultural Research Service (ARS), Plant Genetic Resources
Unit, New York State Agricultural Experiment Station, Geneva,
NY 14456, USA
Crane, Jonathan H., Tropical Research and Education Center,
University of Florida, Institute of Food and Agricultural Sciences
(IFAS), Gainesville, FL 33031-3314, USA
da Silva, Marcicleide L., Universidade Federal do Amazonas,
Manaus, Brazil
Daunay, Marie-Christine, Institut National de la Recherche
Agronomique (INRA), Unité de Génétique et Amélioration des
Fruits et Légumes, Domaine St Maurice, BP 94 84143 Montfavet
Cedex, France
Davis, Anthony S., Hardwood Tree Improvement and Regeneration
Center, Department of Forestry and Natural Resources, Purdue
University, 715 W. State Street, West Lafayette, IN 47907, USA
d’Eeckenbrugge, Geo Coppens, Centre de coopération internationale
en recherche agronomique pour le développement (CIRAD) –
FLHOR/ENSIA-SIARC, Av. Agropolis, TA 50/PS4, 34398
Montpellier Cedex 5, France
de Souza, Francisco Xavier, EMBRAPA Agroindústria Tropical,
PO Box 3761, CEP 60511-110 Fortaleza, Ceara State, Brazil.
xavier@cnpat.embrapa.br
Diczbalis, Yan A., Queensland Department of Primary Industry
and Fisheries, Horticulture and Forestry Science, Centre for Wet
Tropics Agriculture, South Johnstone Road, 4859, Queensland,
Australia
Dornier, Manuel, Centre de coopération internationale en
recherche agronomique pour le développement (CIRAD) –
FLHOR/ENSIA-SIARC, Av. Agropolis, TA 50/PS4, 34398
Montpellier Cedex 5, France
v
vi
Contributors
Doyle, James F., Staff Research Associate, Department of
Pomology, University of California, Davis, California and
Kearney Agricultural Center, Parlier, California, USA
Drew, Roderick A., School of Biomolecular and Biomedical
Science, Griffith University, Nathan, Queensland 4111, Australia
Duarte, Odilo, Escuela Agrícola Panamericana – El Zamorano, PO
Box 93, Tegucigalpa, Honduras
Enujiugha, Victor N., Department of Food Science and
Technology, Federal University of Technology, Akure, Nigeria
Erickson, Homer T., 1409 North Salisbury Street, West Lafayette,
IN, 47906, USA
Ferguson, A.R., The Horticulture and Food Research Institute of
New Zealand Ltd, Private Bag 92 169, Auckland, New Zealand
Ferguson, Ian, The Horticulture and Food Research Institute of
New Zealand, Private Bag 92 169, Auckland, New Zealand
Ferguson, Louise, University of California at Davis, Kearney
Agricultural Center, 9240 South Riverbend Ave., Parlier, CA
93648, USA
Figueira, Antonio, Centro de Energia Nuclear na Agricultura,
Universidade de São Paulo, São Paulo, Brazil
Filgueiras, Heloisa Almeida Cunha, EMBRAPA Agroindústria
Tropical, PO Box 3761, CEP 60511-110 Fortaleza, Ceara State,
Brazil. heloisa@cnpat.embrapa.br
Finn, Chad, United States Department of Agriculture – Agricultural
Research Service, Horticultural Crops Research Laboratory, 3420
NW Orchard Ave, Corvallis, OR 97330, USA
Foidl, Nikolaus, Carretera Sur Km 11, PB432, Managua, Nicaragua
George, Alan, Queensland Department of Primary Industry, PO Box
5165 Sunshine Coast Mail Centre 4560, Queensland, Australia
Gmitter, Frederick G. Jr, University of Florida, Institute of Food
and Agricultural Sciences (IFAS), Citrus Research and Education
Center, 700 Experiment Station Road, Lake Alfred, FL 33850,
USA
Goh, K.J., Applied Agricultural Research Sendirian Berhad, Buloh,
47000 Selangor, Malaysia
Gradziel, Thomas M., Department of Pomology, University of
California, Davis, CA 95616, USA
Graham, Charles J., Louisiana State University Agricultural Center,
Pecan Research/Extension Station, Shreveport, Louisiana, USA
Grant, Joseph A., University of California Cooperative Extension,
420 S. Wilson Way, Stockton, CA 95205, USA
Grauke, L.J., United States Department of Agriculture (USDA)
Agricultural Research Service (ARS) Pecan Breeding and
Genetics, College Station, TX 77879, USA
Gutiérrez, Mario Mejía, Carrera 43 Nro. 10-50, Apto 502, Cali,
Colombia
Hall, John B., School of Agricultural and Forest Sciences,
University of Wales, Bangor, UK
Hancock, James F., Department of Horticulture, Michigan State
University, East Lansing, MI, 28824-1325, USA
Haq, Nazmul, International Centre for Underutilised Crops (ICUC),
University of Southampton, Southampton SO17 1BJ, UK
Harries, Hugh C., Centro de Investigación Cientifica de Yucatán,
Mérida, México
Heiser, Charles, Department of Biology, Jordan Hall, Indiana
University, Bloomington, IN 47405, USA
Huang, Shu, University of Florida, Institute of Food and
Agricultural Sciences (IFAS), Citrus Research and Education
Center, 700 Experiment Station Road, Lake Alfred, FL 33850,
USA
Hughes, Angela, International Centre for Underutilised Crops
(ICUC), University of Southampton, Southampton SO17 1BJ,
UK
Iezzoni, Amy, Department of Horticulture, Plant and Soil Science
Building, Michigan State University, East Lansing, MI 48824,
USA
Janick, Jules, Department of Horticulture and Landscape
Architecture, Purdue University, 625 Agriculture Mall Drive,
West Lafayette, IN 47907-2010, USA
Jeppsson, Niklas, Gamla Fjälkingevägen 34F, 29150 Kristianstad,
Sweden. jeppsson.niklas@telia.com BioUmbrella Available at:
www.bioumbrella.net
Kee, K.K., Applied Agricultural Research Sendirian Berhad, Buloh,
47000 Selangor, Malaysia
Ketsa, Saichol, Department of Horticulture, Kasetsant University,
Chatuchuk, Bangkok, Thailand
Kitajima, Akira, Graduate School of Agriculture, Kyoto
University, Sakyo-ku, Kyoto 8502, Japan
Krueger, Robert R., United States Department of Agriculture –
Agricultural Research Service National Clonal Germplasm
Repository for Citrus and Dates, 1060 Martin Luther King Blvd,
Riverside, CA 92507-5437, USA. rkrueger@citrus.ucr.edu
Kwesiga, F., International Centre of Research in Agroforestry
(ICRAF) Southern Africa Regional Programme, PO Box 128,
Mount Pleasant Harare, Zimbabwe
Lara, Teresa Rojas, Universidad Nacional Agraria, La Molina,
Lima, Peru
Layne, Desmond R., Clemson University, Department of
Horticulture, Clemson, South Carolina, USA
Layne, Richard E.C., 127 Baldwin Avenue, Harrow, Ontario NOR
1GO, Canada
Lin, Shunquan, College of Horticulture, South China Agriculture
University, Guangzhou 510642, China
Lindstrom, Anders Jan, Nong Nooch Tropical Botanical Garden,
34/1 Sukhumvit Highway, Najomtien, Sattahip, Chonburi 20250,
Thailand
Love, Ken, PO Box 1242, Captain Cook, HI 96704, USA.
kenlove@hawaii.edu
Lyrene, Paul, Department of Horticultural Sciences, University of
Florida, 2135 Fifield Hall, Gainesville, FL 32606, USA
Macía, Manuel J., Real Jardín Botánico de Madrid, Plaza de
Murillo 2, E-28014 Madrid, Spain. mmacia@ma-rjb.csic.es
Maynard, Donald N., University of Florida, Gulf Coast Research
and Education Center, Bradenton, Florida, USA
McGranahan, Gale, Department of Pomology, University of
California, Davis, CA 95616, USA
Meerow, Alan W., United States Department of Agriculture –
Agricultural Research Service, 13601 Old Cutler Road, Miami,
FL 33158, USA
Mehlenbacher, Shawn A., Department of Horticulture, Oregon
State University, Corvallis, OR 97331, USA
Moore, Patrick P., Department of Horticulture and Landscape
Architecture, Washington State University, Puyallup, WA 98371,
USA
Mshigenio, Keto E., Zero Emissions Research and Initiatives
(ZERI) Regional Office for Africa, University of Namibia, Private
Bag 13301, Windhoek, Namibia
Nagao, Mike A., Department of Tropical Plant and Soil Sciences,
Beaumont Research Center, University of Hawai’i at Manoa,
Hilo, HI 96720, USA
Contributors
Nageswara Rao, M., University of Florida, Institute of Food and
Agricultural Sciences (IFAS), Citrus Research and Education
Center, 700 Experiment Station Road, Lake Alfred, FL 33850,
USA
Narain, Narendra, Department of Chemical Engineering, Federal
University of Sergipe, Av. Marechal Rondon s/n, São Cristóvão,
SE 49100-000, Brazil
Nelson, Scot C., Department of Plant and Environmental
Protection Sciences, Beaumont Research Center, University of
Hawai’i at Manoa, Hilo, HI 96720, USA
Nerd, Avinoam, Ben-Gurion University of the Negev, POB 653,
Beer-Sheva 84105, Israel
Nobel, Park S., Department of Ecology and Evolutionary Biology,
University of California, Los Angeles, CA 90095-1606, USA
Okie, W.R., United States Department of Agriculture – Agricultural
Research Service, Southeastern Fruit and Tree Nut Research
Laboratory, Byron, Georgia, USA
Ooi, L.H., Applied Agricultural Research Sendirian Berhad, Buloh,
47000 Selangor, Malaysia
Ortiz, Rodomiro, International Institute of Tropical Agriculture,
Ibadan, Nigeria (International mailing address: Lambourn Ltd
(UK), Carolyn House, 26 Dingwall Road, Croydon CR9 3EE,
UK)
Oudhia, Pankaj, 28-A, Geeta Nagar, College Road, Raipur -492001,
Chattisgarh, India
Paris, Harry S., Agricultural Research Organization, Newe Ya’ar
Research Center, Ramat Yishay, Israel
Paull, Robert E., Department of Tropical Plant and Soil Sciences,
University of Hawai’i at Manoa, Honolulu, HI 96822, USA
Pomper, Kirk W., Kentucky State University, Land Grant Program,
Frankfort, Kentucky, USA
Ragone, Diane, National Tropical Botanical Garden, 3530 Papalina
Road, Kalaheo, Hawai’i, HI 96741, USA
Rapoport, Hava, Institute of Sustainable Agriculture, Consejo
Superior de Investigaciones Científicas, Córdoba, Spain
Reich, Lee, 387 Springtown Road, New Paltz, NY 12561, USA
Ruales, Jenny, Escuela Politecnica Nacional, Dept. Ciencia de los
Alimentos y Biotecnología, PO Box 17012759, Quito, Ecuador
Rumpunen, Kimmo, Department of Crop Science, Swedish
University of Agricultural Sciences, SE 291 94 Kristianstad,
Sweden. kimmo.rumpunen@vv.slu.se
Ryugo, Kay, Department of Pomology, University of California at
Davis, Davis, California, USA
Salakpetch, Surmsuk, Chanthaburi Horticultural Research Center,
Khlung, Chanthaburi, 22110, Thailand
Santos, Ronaldo Pereira, Instituto Nacional de Pesquisas da
Amazônia (INPA) – Coordenação de Pesquisas em Ciências
Agronômicas (CPCA), Sala Dr Charles Clement, Caixa Postal
478, CEP 69011-970, Manaus, AM, Brazil
Sauco, Victor Galan, Instituto Canario de Investigación Agraria
(ICIA), Consejeria de Agricultura, Tenerife, Spain
Shu, Zen-Hong, Meiho Institute of Technology, 23 Pingkuang
Road, Neipu, Pingtung, Taiwan 912, Republic of China
vii
Sivakumar, A., T.A. Pai Management Institute, Manipal 576 104,
Udupi District, Karnataka State, India
Soh, A.C., Applied Agricultural Research Sendirian Berhad, Buloh,
47000 Selangor, Malaysia
Soneji, Jaya, University of Florida, Institute of Food and
Agricultural Sciences (IFAS), Citrus Research and Education
Center, 700 Experiment Station Road, Lake Alfred, FL 33850,
USA
Sousa, Nelcimar Reis, EMBRAPA Amaz ia Ocidental, C. Postal
319, Cep. 69011-970, Manaus, Amazonas, Brazil
Sozzi, Gabriel O., Cátedra de Fruticultura, Facultad de Agronomía,
Universidad de Buenos Aires – Consejo Nacional de
Investigaciones Científicas y Técnicas (CONICET), Avda, San
Martín 4453, C 1417 DSE Buenos Aires, Argentina
Steele, Orlo C., Department of Botany, University of Hawai’i at
Manoa, Hilo, HI 96720, USA
St-Pierre, Richard, Plant Gene Resources of Canada, Agriculture
and Agri-Food Canada, Saskatoon Research Centre, 107 Science
Place, Saskatoon, Saskatchewan S7N 0X2, Canada
Tchoundjeu, Z., World Agroforestry Centre (ICRAF), Regional
Office African Humid Tropics, PO Box 2067, Yaoundé,
Cameroon. z.tchoundjeu@cgiar.org
Thomas, Michael B., Department of Botany, University of Hawai’i
at Manoa, 3190 Maile Way, Honolulu, HI 96822, USA
Thompson, Maxine M., Department of Horticulture, Oregon
State University, Corvallis, Oregon, USA
Thompson, Tommy E., United States Department of Agriculture
(USDA) Agricultural Research Service (ARS) Pecan Breeding
and Genetics, College Station, TX 77879, USA
Thorp, Grant, The Horticulture and Food Research Institute of
New Zealand Ltd, Mt Albert Research Centre, Private Bag 92
169, Auckland, New Zealand
Vieira de Freitas, Danival, Av. Efigênio Salles, 2224 – Aleixo,
Condomínio Parque dos Rios II, Bloco 2A/Apto 101, 69060-020
Manaus, Amazonas, Brazil
Webster, Tony, 1 Pine Grove, Maidstone, Kent ME14 2AJ, UK
Werlemark, Gun, SLU Balsgård, Fjälkestadsvägen 459, 291 94
Kristianstad, Sweden
Wilson, Barrett C., Hardwood Tree Improvement and Regeneration
Center, Department of Forestry and Natural Resources, Purdue
University, 715 W. State Street, West Lafayette, IN 47907, USA
Xu, Changjie, Department of Horticulture, Huajiachi Campus,
Zhejiang University, Hangzhou, 310029, PR China
Yen, Chung-Ruey, Department of Plant Industry, National
Pingtung University of Science and Technology, Neipu,
Pingtung, Taiwan, PR China
Yonemori, Keizo, Graduate School of Agriculture, Kyoto
University, Sakyo-ku, Kyoto 8502, Japan
Zee, Francis, United States Department of Agriculture, Curator –
Tropical Fruit and Nuts, Pacific Basin Tropical Plant Genetic
Resources Management Unit, Hilo, Hawai’i
Zhang, Bo, Department of Horticulture, Huajiachi Campus,
Zhejiang University, Hangzhou, 310029, PR China
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Preface
This work is a horticultural encyclopedia of fruit and nut
crops in the broad sense. Entries have been contributed by
authorities and we have attempted to adopt a uniform
approach although this has not been possible in all cases.
Entries are grouped alphabetically by family and then by
species. Common names vary widely by language, region and
locality and are therefore given in the index at the end of the
volume. To find individual entries the reader is advised to
check the combined index of species and common names.
The encyclopedia is intended as an authoritative
compendium of information on economic, mostly edible,
temperate, subtropical and tropical fruit and nuts. There are
literally thousands of species with edible fruit and we could not
include all of them but we have tried to include those that are
considered to be economically significant. Coverage includes
palms and cacti as well as vegetable fruit of the Solanaceae and
Cucurbitaceae, often considered as vegetables in some cultures.
The volume is intended for libraries, researchers, students and
serious amateurs; it is not specifically intended for the average
home gardener although many will find it useful.
It may be helpful to define fruit both botanically and
horticulturally. Fruit in the botanical sense are mature ovaries
and associated flower parts. Their morphological classification
and terminology is complex and somewhat bewildering. When
a single ovary is involved, they are defined as simple fruit that
may be fleshy (succulent) or dry, that is made up of non-living
scherenchyma cells with lignified or suberized walls. The
ovary wall or pericarp is composed of three distinct layers
from outer to inner as exocarp, mesocarp and endocarp.
When the entire pericarp of simple fruit is fleshy (Plate 1),
the fruit is referred to as a berry (grape and tomato for
example). A berry with a hard rind is called a pepo (squash).
When the exocarp and mesocarp form a rind the fruit is called
a heperidium, as in citrus, where the edible juicy portion is
the endocarp. Simple fleshy fruit having a stony endocarp
(peach, cherry, plum and olive) are known as drupes. Here the
edible portion is the mesocarp. When the inner portion of the
pericarp forms a dry paper-like core the fruit are known as
pomes. The dry, dehiscent simple fruit (Plate 2) include such
types as pods (pea), follicles (milkweed), capsules (jimson
weed) or siliques (crucifers). Dry, simple fruit that do not
dehisce (Plate 3) when ripe include achenes (sunflower),
caryopses (maize), samaras (maples), schizocarps (carrot)
and nuts (walnut). Aggregate fruit (Plate 4) are derived from a
flower having many pistils on a common receptacle. Individual
fruit may be drupes (stony) as in blackberries, or achenes (that
is, one-seeded dry fruits) as in strawberry (note that in the
strawberry the edible portion is the receptacle). Multiple fruit
(Plate 4) are derived from many separate but closed clustered
flowers in which the fruitlets coalesce and fuse such as in the
pineapple, fig and mulberry.
Fruit crops in the horticultural sense are those plants
bearing more or less succulent fruit or closely related
structures. They are most often perennial and are usually
woody but there are exceptions such as the banana, which is
the fruit of a tropical herbaceous plant but appears tree-like.
Horticultural groupings are varied. Temperate tree fruit of the
Rosaceae are often referred to as pome fruit when their fruit
are botanical pomes (apple, pear, quince, medlar) or stone
fruit when their fruit are drupes (apricot, cherry, peach,
plum). Fruit of small stature (such as strawberry, raspberry,
blackberry, blueberry and currants) are often called small or
berry fruit. Fruit from vines such as grape and kiwifruit are
known as vine fruit. Nuts are a specialized category of fruit
characterized by a hard shell that is separable from a firmer
inner kernel.
Fruit are an important part of human culture, religious
practices, mythology and art. Horticultural fruit are among
the most beloved of plant products, acclaimed and desired for
their delectable flavour, pleasing aroma and beautiful
appearance. As a result, fruit have become important world
industries. Fruit by their diversity are used in a myriad of
forms: mostly fresh but also dried, canned, frozen, pickled and
fermented. Most are consumed out-of-hand as snacks or
desserts, but are often used in salads, cooked as a side dish,
made into jams and preserves or consumed as wine or
liqueurs. The great majority of fruit are considered an
important source of calories and are often classed as functional
foods or neutriceuticals. A few are rich in fats and oils (oil
palm and avocado for example); most are low in protein. A few
are sources of industrial products, olive for example.
Of the 30 most important world crops in terms of tonnes of
fresh product, five are fruit crops (banana/plantain, orange,
grape, apple and mango). Many fruit are considered important
enough to be included in Food and Agriculture Organization
(FAO) world agricultural statistics.
This work includes 292 entries covering information on
over 322 fruit. We thank all the authors who have contributed
entries and appreciate the helpful editing of Dr Priscilla
Sharland that has helped make this work a reality.
Jules Janick and Robert E. Paull
References
Flores, E.M. (1999) La Planta: Estructura y Funcion. Editorial Libro
Universitario Regional, Cartogo, Costa Rica, pp. 1–884.
Foster, A.S. and Gifford, E.M. (1974) Comparative Morphology of
Vascular Plants. W.H. Freeman & Co., San Francisco, California,
pp. 1–555.
Vozzo, J.A. (ed.) (2002) Tropical Tree Seed Manual. Agriculture
Handbook No. 721. United States Department of Agriculture
(USDA) Forest Service, Washington, DC.
ix
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Plate 1
Plate 2
Plate 1. Types of fleshy fruit (source: Vozzo, 2002).
Plate 2. Types of dry dehiscent fruit (source: Vozzo, 2002).
Plate 3
Plate 4
Plate 3. Types of dry indehiscent fruit (source: Vozzo, 2002).
Plate 4. Types of aggregate and accessory type fruit (source: Vozzo, 2002).
Glossary
Abaxial: the side or face away from the axis.
Abiotic: the absence of life or living organisms.
Abscission: separation of flowers, fruit and leaves from plants.
Accessory buds: buds found beside or above the true bud at a node.
Achene: small single-seeded dry fruit.
Acicular: needle-shaped.
Acorn: the fruit of oaks, a thick-walled nut with a woody cup-like
base.
Acropetally: developing toward the apex.
Aculeate: armed with prickles as distinct to thorns.
Acuminate: elongated tapering gradually to a long, thin point.
Acute: applied to tips and bases that end in a sharp point less than 90°.
Adaptation: the evolutionary adjustments (genetic, structural,
functional) that fit an individual or groups of individuals to their
environment.
Adaxial: the side or face next to the axis.
Adnate: one organ united to another organ, e.g. ovary and calyx tube.
Adventitious: a plant organ that arises from an unexpected position,
e.g. shoots that arise directly from true roots as in raspberry.
Adventitious buds: buds that form along a root or stem other than in
the leaf axil, often after injury or pruning.
Aerenchyma: parenchyma cells that are surrounded by open air-filled
canals.
Aerial roots: roots produced above ground, often used for climbing.
Aggregated: joined together.
Aggregate flower: a single flower or crowded into a dense flower
cluster on a receptacle.
Aggregate fruit: a fruit formed by the coherence or the connation of
pistils that were distinct in the flower.
Air layering (marcottage): multiplying a plant by inducing rooting on
a root or shoot, often involving girdling and when the roots appear
the stem is cut below the roots. The stem is enclosed in shooting
media held in place by a sleeve closed at the two ends.
Alate: winged as a stem or seed.
Albus: white.
Aliform: wing-shaped.
Alternate: the arrangement of one leaf, bud or branch per node at
opposite sides of the stem.
Ambient: the prevailing environmental conditions especially
temperature.
Ament: spike of unisexual, apetalous flowers having scaly, usually
deciduous, bracts; a catkin.
Anastomosed: joined.
Androecious [plants]: plants that bear staminate flowers only.
Androecium: the collective stamens of a flower as a unit.
Androgynophore: a stalk bearing both stamens and pistil above the
point of perianth attachment.
Andromonoecious [plants]: plants that bear both staminate and
perfect (hermaphroditic) flowers.
Anterior: on the front side, away from the axis, toward the subtending
bract.
Anther: pollen-bearing part of a stamen at the top of a filament (or
may be sessile).
Anthesis: the period when the flower opens, often used to refer to the
bursting of the pollen sacs and pollen release.
Anthocyanin: water-soluble red, blue or purple pigments.
Apetalous: without petals.
Apex (pl. apices): the tip or terminal end of a leaf or stem.
Apical: at the apex or tip of an organ.
Apical buds: buds that produce stems and are located at the tip of the
stem.
Apical meristem: meristem located at the tip of the stem.
Apiculate: ending with a short, sharp, abrupt point.
Apocarpous: carpels separate from each other.
Apogamy: a type of apomixis involving the suppression of
gametophyte formation so that seeds are formed directly from
somatic (body) cells of the parent tissue.
Apomixis: reproduction without fertilization or formation of gametes.
An apomict usually is genetically identical with its source plant
(ortet).
Appressed: lying flat against another organ, but not fused to it.
Approach graft or inarching: two independently growing, selfsustaining plants are grafted together; inarching is often used
when replacing the root system and approach grafting when
replacing the scion.
Arbor: a tree, a plant with distinct stem and branches.
Arboreal and arborescent: tree-like or pertaining to trees (> 6 m).
Arcuate: arched, bent like a bow.
Aril: a fleshy appendage of the seed, either on a seedcoat or arising
from the base of a seed.
Arillate: having an aril or arils.
Articulate: having nodes or joints where separation may naturally
occur leaving a clean scar.
Asexual (vegetative) reproduction: reproduction without fertilization
such as tubers, bulbs or rooted stems, or from sexual parts such as
unfertilized eggs or other cells in the ovule.
Attenuate: tapering gradually to a narrow end or base.
Auricle: small fingers of tissue at the base of the leaf blade of a grass
that extend partially around the stem.
Awn: a stiff or flexible bristle, frequent in grasses.
Axial: located in the axis.
Axil: the angle formed between any two adjoining organs, such as
stem and leaf.
Axillary: in an axil.
Axis: the main stem or central support of a plant.
Bare root: a plant dug up with bare roots for transplanting.
Bark: all tissues lying outward from the vascular cambium.
Berry: fleshy or pulpy indehiscent fruit with one or more seeds
embedded in the fleshy tissue of the pericarp; may be formed
from either a superior or an inferior ovary.
xi
xii
Glossary
Bilateral symmetry: being divided into two equal, mirror-image parts.
Bisexual: having both male and female present and functional in the
same flower or inflorescence, hermaphroditic.
Blade: the flattened lamina and expanded part of a leaf or parts of a
compound leaf.
Bourse: the terminal portion of the shoot or spur that bears flowers
and fruit. Axillary buds (bourse buds) develop below the
flowers/fruit and grow into bourse shoots.
Brackish: somewhat salty.
Bract: reduced leaf, subtending a flower or flower stalk, often small.
Branchlet: small or secondary branch.
Bridge graft: graft used to bridge over girdled areas of a tree.
Bristle: hair-like structure.
Bud: an immature or embryonic shoot, flower or inflorescence,
frequently enclosed in scales.
Budding: grafting by inserting a bud, into a slit or hole made in the
bark of a stock plant.
Bud graft: see Budding.
Bulb: a leaf bud with fleshy scales.
Caducous: parts of a plant that are shed or drop off early in development.
Calcareous: applied to soils containing calcium carbonate.
Callus: a small hard protrusion of undifferentiated (parenchyma)
tissue formed at a wounded surface.
Calyx: collective term for the outer separate or united sepals of a
flower, the outer series of flower parts.
Cambium: the growing or dividing single layer of cells located
between the wood and bark.
Camptodrome: leaf venation where the secondary veins bend
forwards and anastomose before the end of the leaf.
Capitular: having a globular head; collected in a head.
Capsule: dry dehiscent fruit composed of two or more united carpels.
Carinate: having a keel or a longitudinal medial line on the lower
surface.
Carpel: simple pistil or unit of a combined pistil.
Carpellody: stamens develop abnormally into carpel-like fleshy
structures.
Cataphylls: scale-like leaves.
Catkin: a scaly spike of usually unisexual and reduced flowers, not
applied to the male or staminate strobilus of conifers.
Cauliflorus: having flowers on the stem.
Cauliforous: stalk-like.
Chalaza: point of an ovule or seed where the integuments are united
to the nucellus opposite the micropyle, to which the funiculus is
attached.
Chimera: plant or parts of a plant whose tissues are of or from
genetically different layers.
Ciliate: marginal or fringe of hairs along an edge.
Clavate: club-shaped, gradually thickened towards the end.
Clone: a group of individual plants asexually propagated from a
single plant and, therefore, genetically identical.
Clonal test: evaluation of an individual (ortet) or a clone (ramets) by
comparing clones.
Coleoptile: in monocotyledons, a sheath that covers the plumule.
Coleorhiza: in monocotyledons, a sheath that covers the radicle.
Columella: the persistent axis of certain capsules.
Comose: hairy.
Companion cells: phloem cell connected to a sieve-tube member by
numerous plasmodesmata.
Compatible: plant parts (scion and rootstock) capable of forming a
permanent union when grafted.
Complete flower: having all the components: pistils, stamens, petals
and sepals.
Compound: composed of several similar parts (leaflets), or a
paniculate inflorescence (florets) each attached to a petiole-like
structure (rachis) or directly to the top of the petiole.
Compound leaf: divided into two or more blades (leaflets); palmately
compound leaves have three or more leaflets arising from a
common point, while pinnately compound leaves have leaflets
arranged along a common axis.
Cone: mass of ovule-bearing or pollen-bearing bracts or scales
arranged spirally on a cylindrical or globose axis; common to most
conifers.
Conic: cone-shaped.
Conifer: plants with cones and naked ovules; any of an order of trees
and shrubs bearing true cones or with arillate seeds.
Connate: parts of the same whorl grown together, as in sepals.
Coppicing: trees that are cut down or pruned severely to the stump
and re-growth produces multiple stems called poles.
Cordate: heart-shaped in outline, such as a leaf with two rounded
basal lobes.
Coriaceous: having a stiff leathery texture.
Corm: enlarged fleshy base of a stem, bulb-like but solid.
Corolla: the second floral whorl of a complete flower, collective term
for all free or united petals of a flower.
Corymb or corymbose: flat- or round-topped flower cluster, outer
pedicels are longer than inner pedicles with the outer flowers
opening before the inner flowers.
Costa: the extension of the petiole through the lamina of a palmate
leaf.
Costapalmate leaf: palm leaf in which the petiole extends into the leaf
blade.
Cotyledon: primary or rudimentary embryonic leaf of seed plants.
Crenate: having margins with shallow, rounded teeth.
Crenatures: notches or indentations.
Cross-pollination: pollination by a genetically different plant. An
outcross is a cross to an unrelated individual.
Crownshaft: the tightly packed tubular leaf bases of some featherleaved palms sheath each other around the stem forming a
conspicuous neck-like structure.
Crustaceous: having a hard covering or crust.
Cultivar: cultivated variety, synonymous with the term variety in the
International Code of Nomenclature. Cultivars are always graced
with a name and when written are usually capitalized and
separated by single quotes or preceded by the word cultivar or
abbreviation cv. A cultivar may be a clone or an F1 hybrid or be
seed-propagated, if uniform, but practically all cultivars of fruit
are clones. The term cultivar was proposed to avoid confusion
with a botanical variety (see Variety).
Cuneate: wedge-shaped; triangular and tapering to a point at the
base.
Cupola: dome-like structure.
Cutaneous: of, pertaining to or affecting the skin.
Cuticle: waxy covering on the surface of stems and leaves that acts to
prevent desiccation in terrestrial plants.
Cutin: complex fatty or waxy substance found on the surface of
certain seeds or leaves, often making them impermeable to water.
Glossary
Cutting: detached portion of stem or other plant part which, when
rooted, produces a whole plant.
Cyme: an irregular umbellate inflorescence in which the primary axis
bears a single terminal flower that develops first.
D.B.H.: diameter at breast height; generally accepted standard for
measuring trees at 137 cm above the ground.
Deciduous: detaching or falling off very early; usually in reference to
leaves, leaf tips or the sepals and petals of most flowers after
expansion.
Decurrent: continued downwards to the stem and attached to the
stem, forming a wing-like appendage.
Decussate: arranged along the stem in pairs, each pair at right angles
to the pair immediately above or below, as in leaves.
Dehiscence: splitting or opening in a regular manner to let pollen or
spores escape through a valve, slit, cap or other opening.
Dehiscent dry fruit: mature fruit that has a dry pericarp that opens to
let seeds escape.
Dentate: toothed; having triangular teeth that are perpendicular to
the margin.
Denticulate: finely dentate.
Dichogamy: male and female organs mature on the same plant at
different times, ensuring natural cross-pollination.
Dicotyledonous: having two seed leaves.
Digitate: divided into distinct lobes in a radiate manner.
Dioecious: species with staminate and pistillate flowers on separate
individuals, unisexual (staminate or pistillate).
Discolorous: leaves in which the two surfaces are different in colour.
Distal: opposite the point of attachment; apical; away from the axis.
Divaricate: (of a branch) coming off the stem almost at a right angle.
Dormant: physiological condition with no active growth due to
temperature or drought.
Dorsal: appertaining to the back.
Double-worked plant: a plant that has been grafted twice, usually to
overcome incompatibility with an interstock between the scion
and rootstock.
Drupe: a fleshy indehiscent fruit having a hard endocarp and a single
seed; sometimes having more than one encased seed.
Druse: star-shaped crystal.
Dulcis: sweet, agreeable.
Ecotype: a race that is adapted to a particular environment.
Ellipsoid: three-dimensional body whose plane sections are all either
ellipses or circles.
Elliptic: oblong with the ends equally or almost equally rounded.
Emarginate: notched at the apex, as a petal or leaf.
Emasculated: removal of immature staminate flower structures
(anthers) to prevent self-pollination.
Endocarp: inner layer of a pericarp.
Endosperm: in angiosperms, an embryonic nutritive tissue formed
during double fertilization by the fusion of a sperm with the polar
nuclei; which is triploid (3n).
Entire: smooth, without teeth or indentations; applied to margins,
edges.
Eophyll: first leaf above the cotyledons.
Epicalyx: an involucre of bracts below the flower.
Epicarp: outer layer of the pericarp or matured ovary.
Epicotyl: portion of the embryo or seedling above the cotyledons.
xiii
Epidermis: outer layer of cells.
Epigeal: growing on or close to the ground.
Epigeal germination: emergence of cotyledons above the surface of
the ground.
Epigynous: growing on the summit of the ovary, or apparently so.
Epipetalous: a flower in which the stamens are connected to the
petals.
Ester: chemical compound formed between an acid and alcohol.
Etiolate: elongated due to lack of chlorophyll.
Etiolation: elongation, discoloration and poor plant growth due to the
lack of chlorophyll.
Evergreen: plants with live leaves persisting through one or more
winter seasons.
Exfoliate: peeling bark from a branch or trunk.
Exocarp: outermost layer of the fruit wall (pericarp).
Exoderm: outer layer of one or more layers of thickness in the cortex
of some roots.
Exserted: projecting beyond, stamens exceeding the corolla.
Fat: ester of fatty acid and glycerol (or another alcohol) found in
plants and animals; in liquid form, called oil.
Filament: stalk-like portion of a stamen, supporting the anther.
Filiform: thread-like, slender.
Fissure bark: furrowed and ridged, or splitting lengthwise.
Florets: applied to the separate flowers of inflorescences.
Flower: angiosperm reproductive structure composed of calyx
(sepals), petals, stamens and pistil.
Foliaceous: leaf-like.
Foliole: leaflet.
Follicle: dry, one-celled fruit with a single placenta and splitting
along the opposite edge.
Free: separate, not joined together or with other organs.
Funiculus (funicle): stalk of an embryo or seed.
Gamopetalous: having the petals fused around the base.
Genotype: an individual’s hereditary constitution, with or without
phenotypic expression of the one or more characters it underlies.
It interacts with the environment to produce the phenotype.
Geographic race: a race native to a geographic area.
Geographic variation: The phenotypic differences among native trees
growing in different portions of a species’ range. If the differences
are largely genetic rather than environmental, the variation is
usually specified as racial, ecotypic or clinal.
Glabrescent: smooth.
Glabrous: having no hairs, bristles or stalked glands.
Gland: a secretory structure on the surface in a depression, protuberance or appendage on the surface of an organ that secretes a
usually sticky fluid; any structure resembling such a gland.
Gland of salt: hydatode that excretes water with a high salt and
mineral proportion.
Glaucous: surface with a fine white substance (bloom) that will rub
off.
Globose: globe-shaped, spherical.
Glochids: the fine hair-like spines found in the areoles of many cacti.
Glomerule: small, compact cluster.
Glutinous: sticky, gummy, having the quality of glue.
Graft: a finished plant that comes from joining a scion and a rootstock.
xiv
Glossary
Grafting: uniting parts of separate individuals, such as an aerial
portion (scion) that is joined with a rootstock, and a union forms
allowing/re-establishing vascular continuity and growth.
Graft incompatibility: inability of the stock and scion to form or
maintain a union that will allow growth.
Graft union: the junction where the rootstock and scion come
together (fuse).
Gummosis: any of various viscous substances that are exuded by
certain plants and trees, then dry into water-soluble, noncrystalline, brittle solids.
Gutation: process by which water passes from inside the leaf and is
deposited on the outer surface.
Gynoecious [plants]: plants that bear pistillate flowers only.
Gynoecium: whorl or group of carpels in the centre or at the top of
the flower; all the carpels in a flower.
Gynomonoecious [plants]: plants that bear both pistillate and perfect
(hermaphroditic) flowers.
Gynophore: stalk of the pistil.
Halophyte: plant with the capability to grow in saline habitats.
Hapaxanthic: a palm stem that exhibits determinate growth that dies
after flowering and fruiting.
Heteromorphic: having different forms at different periods of the life
cycle.
Heterosis: increased vigour or other superior qualities arising from
the crossbreeding of genetically different plants.
Heterostyly: flower in which styles and stamens are of different
heights relative to each other to ensure cross-pollination. See
Heterotristyly for explanation of tristyly and distyly.
Heterotristyly (tristyly): flower in which the stigma and stamens are
at different heights relative to each other. In tristyly stamens and
stigma are the same length, or stamens are longer than stigma or
stigma longer than stamens. (In distyly flowers have stamens
longer than stigma or stigma longer than stamens.)
Hirsutus: with long soft hairs.
Hispid: harshly or bristly hairy.
Hydathode: structure that secretes water; found in the margins of
leaves.
Hypanthium: floral tube formed by the fusion of the basal portions of
the sepals, petals and stamens from which the rest of the floral
parts emanate.
Hypocotyl: portion of the axis of a plant embryo below the point of
attachment of the cotyledons; forms the base of the shoot and the
root.
Hypogeal: underground, subterranean.
Hypogeal germination: emergence of cotyledons below the surface of
the ground.
Hypogynous: situated on the receptacle beneath the ovary and free from
it and from the calyx; having the petals and stamens so situated.
Imbricate: applied to leaves or to the parts of the flower when they
overlap each other in a regular arrangement.
Imparipinnate: a compound leaf with a terminal pinna.
Imperfect flower: unisexual flower; flower lacking either male or
female parts.
Impressed: having sunken veins as viewed from the upper leaf surface.
Inarching: see Approach graft.
Incised: intermediate between toothed (dentate or serrate) and lobed,
being a sharply inward cut leaf (the inward cuts are called incisions).
Incompatible graft: plants whose parts will not form a permanent
union when grafted together.
Incompatibility: a failure or partial failure in some process of grafting
or fertilization. For example, pollen tube growth may be deficient
even though the egg and sperm cells are potentially functional.
Incomplete flower: flower lacking at least one of the four basic parts:
pistils, sepals, stamens or petals.
Incumbent: describing cotyledons lying with the back of one against
the radicle.
Indehiscent: not opening naturally when ripe.
Indumentum: general term for the hairy or scaly covering of plants.
Induplicate: palms in which the leaflets or segments are folded
upward forming a ‘V’.
Inferior ovary: one with the flower parts growing from above; one
that is adnate to the calyx.
Inflorescence: any complete flower cluster including branches and
bracts; clusters separated by leaves are separate inflorescences.
Infusion: liquid derived from seeping or soaking (leaves, bark, roots,
etc.); extraction of soluble properties or ingredients.
Integument: natural covering, such as skin, shell or rind; also
tegument.
Intercalary: meristem situated between the apex and the base.
Internode: the portion of a stem between two nodes.
Interstock: an intermediate plant part, most often a stem piece that is
compatible with both the scion and the rootstock. Used in cases
where the scion and rootstock are not directly compatible with
each other or where additional dwarfing and cold or disease
resistance is desired.
Involucre: a number of bracts subtending a flower cluster, umbel or
the like.
Irregular: flowers in which the parts of the calyx or corolla are
dissimilar in size or shape; or the flower cannot be divided into
two equal halves in a vertical plane.
Jugate: yoked.
Juvenile: organ or tissue that is not fully developed; a plant that is
unable to produce flowers in contrast with a mature plant which is
capable of flowering and is able to reproduce. Plants in the
juvenile state may have different morphological features from
mature plants.
Lacunae: small spaces.
Lamina: blade or expanded portion of a leaf.
Lanate: having woolly hair.
Lanceolate: lance-shaped, longer than wide, widest below the middle,
tapering toward the apex, or both apex and base; resembling a
lance head.
Laticifer: cell or series of longitudinal cells that contain a specific
fluid called latex.
Leaf primordium: lateral outgrowth from the apical meristem that
develops into a leaf.
Leaf scar: mark left on a twig when a leaf falls.
Leaflet: single segment (blade) of a compound leaf.
Legume: member of the Fabaceae (Leguminosae), the pulse family,
with a dry, dehiscent fruit formed from one carpel and having two
longitudinal lines of dehiscence.
Glossary
Lens: biconvex-lens-shaped.
Lenticel: corky spot on the surface of a twig; sometimes persists on
the bark of a branch to admit air into the interior.
Lignotuber: woody underground stem.
Ligule: strap-shaped corolla, as in the rayflowers of Asteraceae; a thin,
often scarious (scar-like) projection from the summit of the sheath
in grasses.
Linear: narrow and elongated with parallel or nearly parallel edges.
Liners: rooted shoots used for propagation.
Lobe: segment of a leaf between indentations that do not extend to
the midrib or base of the leaf.
Locule: compartment or cavity of an ovary, anther or fruit.
Loculicidal: dehiscing lengthwise, dividing each loculus into two parts.
Loculus: cell of a carpel in which the seed is located; cell of an anther
in which the pollen is located.
Marcottage: see Air layering.
Mature: organ or tissue that is fully developed, for example, ripe; a
plant that is able to produce flowers (reproduce). See Juvenile.
Medium plane: plane that divides the seed into two equal parts.
Mericarp: portion of fruit that seemingly matured as a separate fruit.
Mesocarp: fleshy part of the wall of a succulent fruit; the middle layer
of the pericarp in a drupe.
Mesomorphic: soft and with little fibrous tissue, but not succulent.
Microphyllous: having small leaves that are usually hard and narrow.
Micropyle: opening in integument at apex of ovule.
Midrib: central or main vein of a leaf or leaf-like part.
Monocotyledonous: having one seed leaf.
Monoecious: having stamens and pistils in separate flowers on the
same plant.
Monotypic: having only one representative.
Morphology: study of the form and structure of whole plants, organs,
tissues or cells.
Mucilage: any of various gummy secretions or gelatinous substances.
Mucronate: terminating in sharp point.
Mycorrhiza: symbiotic association of fungus and root.
Naked flower: having no perianth.
Nectary: gland that secretes nectar.
Needle: narrow, usually stiff leaf, as in pines, firs and hemlocks.
Nervation: arrangement of veins, as in a leaf; venation.
Node: the narrow region on a stem where a leaf or leaves are or were
attached.
Nucellus: maternal tissue surrounding the ovule.
Nut: dry, hard, indehiscent, one-celled and one-seeded fruit; usually
resulting from a compound ovary.
Obconic: conical with the apex downwards.
Oblong: elongate in form with sides parallel or nearly parallel, the
ends blunted and not tapering; wider than long.
Obovate: ovate with a narrow end at the base.
Obtuse: blunt point or rounded at the end, the angle of the point
greater than 90°.
Ontogeny: development of an individual organism.
Open pollination: natural pollination effected by wind or insects, and
not directly influenced by humans.
Opposite: leaves or branches growing in pairs, one on each side of the
axis and 180° from each other.
xv
Orbicular: circular in outline.
Ovate: egg-shaped in outline with the wider half below the middle.
Ovoid: three-dimensional structure having the shape of an egg with
the broader half below the middle.
Palmate: radiately arranged, ribbed or lobed, as in the fingers of a
hand.
Palminerved: having lobes radiating from a common point.
Panicle: a compound raceme; an inflorescence in which the lateral
branches arising from the peduncle produce flower-bearing
branches instead of single flowers.
Papilla(ae): a minute nipple-shaped projection.
Pari: prefix meaning equal.
Parietal: on the sides or wall of the carpels.
Paripinnate: having a pair of leaflets at the apex.
Parthenocarpy: production of fruit without viable seeds, as in
bananas and some grapes; may be induced artificially.
Parthenogenesis: type of reproduction in which females produce
offspring from unfertilized eggs; a type of apomixis.
Patch budding: is done on plants with thick bark while the plants are
actively growing; a rectangular piece of bark is removed from the
rootstock and covered with a bud and a piece of bark from the
scion.
Pedicel: the secondary stalks of a compound inflorescence bearing
individual flowers.
Peduncle: main flower stalk of a compound inflorescence, supporting
either a cluster of flowers or the only flower of a single-flowered
inflorescence.
Peltate: having the stalk of a leaf attached to the lower surface of the
blade somewhere within the margin, rather than on the margin.
Pendulous: drooping or hanging loosely.
Pentamerous: grouped in fives.
Perennial: plant that lives for more than 2 years.
Perfect: flowers having both functional stamens and pistils.
Perianth: calyx and corolla collectively, or the calyx alone if the
corolla is absent.
Pericarp: walls of a ripe ovule or fruit; its layers may be fused into one,
or separated or divisible into epicarp, mesocarp and endocarp.
Periclinal: parallel to the surface.
Pericycle: a cylinder of cells that lies just inside the endodermis
forming a cylinder around vascular tissues in many roots and
stems.
Perigynous: when the sepals, petals and stamens are carried up
around the ovary but not attached to it.
Persistent: remaining attached past the expected time for dropping.
Petal: one of the parts of the corolla, the inner set of the perianth;
may be separate or united to another petal.
Petaloid: sepal having colour and texture resembling petals.
Petiole: leaf stalk; sometimes absent.
Petiolule: compound leaves, the stalk of a leaflet.
Phellogen: sheet-like meristem that produces cork.
Phenology: the relation between periodic plant development (such as
leaf development, flowering, root growth) and seasonal climatic
changes (such as temperature, moisture availability or daylength).
Phenotype: the plant or character as described, or degree of
expression of a character; the product of the interaction of the
genes of an organism (genotype) with the environment.
Photoperiodism: response (e.g. flowering, germination) of organisms
to the relative length of the daily periods of light and darkness.
xvi
Glossary
Phyllode: expanded flattened petiole resembling and having the
function of a leaf.
Phylogeny: the evolutionary relationships among organs and taxa.
Phylotaxy: arrangement of leaf order.
Pilose: covered with hair, especially soft hair.
Pinnate: compound leaf, having lobes or blades of a leaf arranged
along the sides of a common axis; also applies to major lateral
veins of a leaf.
Pinnule: foliole of second or third order.
Piscicidal: poisonous to fish.
Pistil: the female organ of a flower, collectively ovary, style and stigma
(gynoecium) consisting of a single carpel or two or more fused
carpels.
Pistillate: with pistils and not stamens; may apply to individual
flowers or inflorescences or to plants of a dioecious species in
angiosperms; female.
Pit: hard endocarp that encloses the seed of a drupe.
Placenta: ovule-bearing part of the ovary and seed-bearing surface in
the fruit.
Placentation: method of attachment of the seeds within the ovary.
Plagiotropic: attracted to one side.
Pleonanthy: a palm stem that grows indeterminately, producing
flowers on specialized axillary branch systems.
Plumule: shoot of the embryo.
Pneumatophores: erect exposed roots that arise from underground
root system.
Pod: any dry, dehiscent fruit.
Polarity: structural and/or physiological difference established in the
plant, embryo, organ, tissue, cell; often in reference to direction.
Pollarded: cut back nearly to the trunk to produce a dense mass of
branches.
Pollen: male spore-like structures produced by anthers in flowers and
by male cones.
Pollen grain: small structure of higher plants that contains haploid
male nuclei (gametes) and is surrounded by a double wall, the exina
and intina; transported from the anther of the stamen to the stigma
or stigmatic portion of the pistil, a process called pollination.
Pollen sac: locus in the anther that contains the pollen grains.
Pollen tube: microscopic tube that grows down the stigma from the
pollen grain; through it the sperm cells are deposited into the
embryo sac.
Pollination: process by which pollen is transferred from the anther
where it is produced, to the stigma of a flower.
Pollinium: mass of pollen grains.
Polymorphic: having, assuming or passing through many or various
forms or stages; polymorphous.
Polysome: ribosome associated with protein syntheses.
Pome: fruit in which the floral cup forms a thick outer fleshy layer
and has a papery inner pericarp layer (endocarp) forming a
multiseeded core (e.g. apple, pear).
Precocious: developing early, flowers appear before leaves.
Propagule: a plant part, such as a bud, tuber, root or shoot, used to
propagate an individual vegetatively.
Prophyll: a leaf-like bract that covers the inflorescence during
development.
Protandry: the termination of the shedding of pollen of a plant or
flower prior to receptivity on the same plant or flower
(proterandrous).
Protogyny: the termination of stigma receptivity prior to the
maturation of pollen on the same plant or flower (proterogynous).
Provenance: the original geographic source of seed, pollen or
propagules.
Pruinose: having a whitish dust on the surface.
Pruning: removal of unwanted parts of a plant.
Pseudocarp: fruit that develops not only from the ripened ovary, or
ovaries, but from non-ovarian tissue as well.
Puberulent: minutely pubescent.
Pubescent: covered with fine, short, soft hairs.
Pulvinule: pulvinus at the base of a petiole.
Pulvinus: swelling at the base of the petiole related to leaf movement.
Punctate: spotted with coloured or translucent dots or depressions,
usually due to glands.
Pyrene: hard or stony endocarp; nutlet.
Pyriform: pear-shaped.
Pyxidium: a seed capsule with a top that comes off as the seeds are
released.
Race: a population within a species that has similar characteristics but
is distinct from other populations, usually interbreeding.
Raceme: an inflorescence in which the single flowers are borne on
pedicels arranged singly along the sides of a flower–shoot axis.
Rachilla (pl. rachillae): the rachis of the spikelet in palms, grasses and
sedges. In palms the branches of the inflorescence that bear
flowers on the terminal segements.
Rachis: main axis of a spike or of a pinnately compound leaf,
excluding the petiole.
Radial symmetry: when cut through the centre along any plane,
produces similar halves.
Radicle: portion of the embryo below the cotyledons that will form
the roots, more properly called the caudicle.
Ramet: an individual member of a clone.
Raphide: sharp-pointed, sometimes barbed, crystals of calcium
oxalate, often present in aroids and palms, sometimes acrid.
Receptacle: portion of the axis of a flower stalk on which the flower is
borne.
Receptivity: the condition of the pistillate flower that permits
effective pollination.
Reduplicate: palms in which the leaflets or segments are folded
downward forming an inverted ‘V’.
Reflexed: abruptly turned or bent backwards.
Refracted: bent sharply backwards.
Regular flower: all flower parts radially symmetrical of similar size
and shape.
Reniform: kidney-shaped in outline.
Reparius: growing on the banks of streams or lakes.
Reticulate: having the veins or nerves arranged in a net-like pattern.
Retrorse: bent abruptly backwards and downwards.
Revolute: leaves where the margins are rolled backwards towards the
midrib.
Rhizome: underground stem usually growing horizontally.
Rhizosphere: soil surrounding the root.
Rhombic: four-margined leaf that is diamond-shaped, having three
prominent tips, two on the side and one at the top.
Rimose: having many fissures.
Rootstock: the portion of a grafted or budded plant that provides the
root system; may include a length of stem.
Rosette: a crown of prostrate leaves radiating at the base of a plant.
Rugose: rough and wrinkled; applied to leaves on which the reticulate
venation is very prominent underneath.
Russetting [of fruit skin]: reddish-brown discoloration.
Glossary
Saccate: pouched or bag-shaped.
Sagittate: shaped like an arrowhead.
Samara: indehiscent, one-seeded winged fruit (e.g. maple, ash).
Samaroid: having the shape of the samara.
Sarcotesta: fleshy testa.
Scabrous: rough or harsh to the touch due to minute stiff hairs or
other projections.
Scale: reduced leaves that are usually sessile and seldom green;
sometimes epidermal outgrowths, if disc-like or flattened.
Scape: long leafless stem that finishes in a flower or inflorescence.
Scion: part of plant with three or four buds inserted into a stalk for
grafting, often a shoot that is grafted onto the rootstock of another
plant.
Sclerous: hardened or toughened.
Seedcoat: outer protective layer of a seed that develops from the
integument of the ovule; testa.
Senescence: ageing, a progression of irreversible change in a living
organism, eventually leading to death.
Sepal: one of the parts of a calyx or outer set of flower parts; may be
separate or united to another sepal.
Septate: divided by a septum or septa.
Septum: dividing wall or membrane in a plant, ovary or fruit.
Sericeous: covered with silky down.
Serotinous: late in occurring, developing and flowering.
Serrate: having sharp, saw-like teeth pointed upwards or forwards.
Serrulate: finely or minutely serrate.
Sessile: without a stalk.
Sessile leaf: leaf that has no petiole.
Setiform: bristle-like.
Sheath: tubular structure surrounding an organ or part, such as the
basal part of a leaf; the circle of scales around the base of pine
needles.
Shield budding: see T-budding.
Shrub: woody plant less than tree size, frequently with several
branches at or near the base.
Silique: dry, dehiscent, elongated fruit formed from a superior ovary
of two carpels, with two parietal placentas, and divided into two
loci by a false septum between the placentas; occurs in plants of
the family Cruciferae.
Simple fruit: fruit that ripens from a single ovary.
Sincarpica flower: flower with its carpels fused.
Sinus: cleft or recess between two lobes of an expanded organ such as
a leaf.
Spadix: spike with a thickened, fleshy axis.
Spathe: large bract at the base of a spadix, that encloses the spadix (at
least initially) as a sheath.
Spatulate: shaped like a spatula (spoon-shaped); somewhat widened
towards a rounded end.
Spicate: having spikes.
Spike: type of inflorescence having sessile flowers on a long common
axis.
Stamen: sporophyll within the flower; in angiosperms, the floral
organ that bears pollen.
Staminate: having pollen-bearing stamens only, on individual plants
of a dioecious species or flowers and inflorescences; male.
Staminode: an aborted or rudimentary stamen in which the anther
remains reduced and sterile.
Sterility: absence or defectiveness of pollen, eggs, embryo or endosperm,
which prevents sexual reproduction. See Incompatibility.
Stigma: pollen-receptive part of a pistil, often enlarged, usually at the
tip of the style.
xvii
Stipe: stalk of a pistil; not the pedicel; stalk under elevated glands.
Stipitate: having a stipe or borne on a stipe.
Stipule: one of a pair of lateral appendages at the base of a leaf
petiole.
Stock or rootstock: the root-bearing plant or plant part.
Stolon: large, indeterminate, underground stem that produces roots
at intervals capable of giving rise to a new plant.
Stone: drupe.
Stool: cluster of shoots or stems springing up from a base or root.
Striate: having longitudinal lines, such as bark.
Style: narrow upper part of ovary that supports the stigma.
Sub: prefix signifying almost, less than completely, somewhat, under.
Subbacate: partly pulpy (bacate means completely pulpy like a berry).
Sub-basal: related to, situated at, or forming the base.
Suberin: complex of fatty substances present in the wall of cork
tissue that waterproofs it and makes it resistant to decay.
Suberize: convert to corky tissue.
Suckers: many stems arising from the base of a tree or shrub and
gradually spreading the diameter of its basal area.
Sulcate: with narrow, deep grooves on the stem.
Superior ovary: ovary with the flower parts growing from below it.
Sympodial: an axis made up of multiple bases.
Syncarpous: having the carpels of the gynoecium united in a
compound ovary.
Synpetalous: without petals.
Taxon (pl. taxa): a formal category of taxonomy, e.g. family, genus,
species.
Taxonomy: classification of organisms, including identification and
nomenclature until recently based upon morphological characters.
T-budding: when a single mature bud is inserted into a T-shaped
incision in the rootstock (also known as shield budding).
Tepal: perianth member or segment; term used for perianth parts
undifferentiated into distinct sepals and petals.
Terete: circular in cross-section.
Terminal bud: apex of the leaf is at the tip end opposite the petiole.
Testa: outer covering of the seed; the seedcoat.
Tetramerous: relating to groups of four.
Thorn: hard, sharp-pointed stem.
Tomentose: having dense soft, fine, matted, short hairs.
Toothed: the condition of a margin broken into small projecting
segments, either serrations, dentations or crenations.
Topworking: cutting back the branches and top of an established tree
or mature plant and then budded or grafted with new scions.
Torus: the part on which the divisions of a flower or fruit are
seated.
Training: orientation of a plant in space, often combined with
pruning and tying.
Trichome: hair, bristle, scale or other such outgrowth of the
epidermis.
Trigonous: three-angled.
Truncate: having an apex or base that is almost or quite straight
across, if cut off.
Tuber: underground stem in which carbohydrates are stored.
Tuff: cluster of short-stalked flowers, leaves, etc., growing from a
common point.
Turbinate: top-shaped; a solid having a tapering base and a broad,
rounded apex.
xviii
Glossary
Umbel: inflorescence having the flower stalks or pedicels, nearly
equal in length, spread from a common centre.
Understock: rootstock.
Unguiculate: having a small hook.
Union: the point where the scion and rootstock are joined.
Valvate: meeting without overlapping, as the parts of certain buds;
opening by valves, as certain capsules and anthers.
Valve: one of the segments into which a capsule dehisces; flap or lidlike part of certain anthers.
Variety: a taxonomic subdivision of a species based on minor
characteristics and often an exclusive geographic range. See
Cultivar for a distinction between botanical variety and cultivated
variety (cultivar).
Vegetative: referring to non-reproductive structures or growth.
Vegetative (asexual) propagation: propagation of a plant by asexual
means (budding, dividing, grafting, rooting and air layering) when
the resulting members of the clone (ramets) are identical with
those of the original plant (ortet). Propagating by apomictic seed
or somatic embryos is a means of vegetative propagation. See
Asexual reproduction.
Vestigial: of or pertaining to a degenerate or imperfectly developed
organ or structure having little or no utility, but which in an
earlier stage of the individual or in preceding organisms
performed a useful function.
Villous: densely covered with soft, fine, unmatted, relatively long hairs.
Viscid: sticky.
Viviparous: seeds germinate on the parent plant.
Whip graft: uses scions and rootstocks in grafting of the same
diameter, often pencil thin.
Whorl: a ring of three or more structures (leaves, stems, etc.) in a
circle, not spiralled.
Wildling: a wild plant transplanted to a cultivated area.
Wing: membrane; or thin, dry expansion or appendage of a seed or
fruit.
The Glossary is based, in part, on:
Hemsley, W.B. (1877) Handbook of Hardy Trees, Shrubs, and
Herbaceous Plants. Longman, Green and Co., London.
Snyder, E.B. (1972) Glossary for Forest Tree Improvement Workers.
Southern Forest Experiment Station, United States Department
of Agriculture (USDA). Available at: http://www.sfws.auburn.
edu/sfnmc/class/glossary.html (accessed 8 November 2006).
Vozzo, J.A. (ed.) (2002) Tropical Tree Seed Manual. Agriculture
Handbook No. 721. United States Department of Agriculture
(USDA) Forest Service, Washington, DC.
A
ACTINIDIACEAE
Actinidia deliciosa
kiwifruit
Introduction
The kiwifruit of international commerce are large-fruited
selections of Actinidia chinensis Planch. and Actinidia deliciosa
(A. Chev.) C.F. Liang et A.R. Ferguson, Actinidiaceae.
Kiwifruit are among the most recently domesticated of all
fruit plants: A. chinensis has been cultivated commercially for
little more than 20 years, A. deliciosa for about 70 years. The
common name ‘kiwifruit’ is itself very recent, being devised in
1959 originally for the fruit of A. deliciosa but now being
increasingly applied to fruit of other Actinidia species, such as
Actinidia arguta (Sieb. et Zucc.) Planch. ex Miq., the ‘hardy’
kiwifruit or ‘baby kiwi’. Kiwifruit has completely replaced the
older name of ‘Chinese gooseberry’. In China, ‘mihoutao’
(monkey peach) is used for all Actinidia species, but as A.
chinensis and A. deliciosa are by far the most important
economically of the various Actinidia species in China, they
are often simply referred to together as mihoutao. More
explicitly, the two species are sometimes referred to together
as zhonghua (Chinese) mihoutao.
Although relative newcomers to the fruit bowl, kiwifruit
have found ready acceptance among consumers. Externally,
kiwifruit are distinctly different from other fruit. Once cut
open, they are strikingly beautiful with a strong radiating
pattern of lighter-coloured rays interspersed by several rows of
small dark brown-black seeds. The core is creamy white and
the outer flesh is a bright green or a sharp yellow. It is thus not
surprising that kiwifruit are often used for food decoration
rather than just being eaten. The fruit flavours are distinctive
but very different in yellow and in green kiwifruit and appeal
to largely different sectors of the consuming public. Ripe fruit
of A. arguta can have a superb, aromatic flavour.
History and origins
Fruit of different Actinidia species have been collected from
the wild for many centuries but widespread cultivation was
first attempted only during the last 100 years. Actinidia arguta
was being grown in gardens in Europe and North America
during the last decades of the 19th century but its commercial
potential has yet to be realized. Actinidia deliciosa was first
grown in orchards in New Zealand in the 1920s, and
cultivation of A. chinensis started half a century later.
Seed and plants of A. deliciosa were sent from China to
Europe and to the USA from 1898 onwards, but establishment
of the plants was restricted by their climatic requirements, the
need for both male and female plants for fruiting, and by the
onset of World War I. Seed of A. deliciosa also went to New
© CAB International 2008. The Encyclopedia of Fruit & Nuts
(eds Jules Janick and Robert E. Paull)
Zealand in 1904 and there the circumstances were better and
the plants prospered. The first small orchard was fruiting by
the early 1930s but cultivation remained for many years on a
very small scale with just enough fruit being produced for the
New Zealand market. Exports, starting in 1953, were at first
considered as a means of absorbing surplus fruit that could
not be sold locally. Promotion and acceptance of kiwifruit on
the export markets resulted in much better returns to growers
from the mid-1960s so that production expanded rapidly and
by 1976 exports of kiwifruit from New Zealand exceeded local
consumption. The subsequent development of the New
Zealand kiwifruit industry was the development of an industry
geared towards export. Growers in other countries were
encouraged to start growing kiwifruit and they used the
cultivars first developed in New Zealand. These cultivars were
all of A. deliciosa and so it was the hairy skinned kiwifruit with
green flesh that became known to consumers throughout the
world.
Domestication of A. chinensis is even more recent. The first
experimental plantings of this species were established in China
in 1957. Subsequently, a survey was made of wild kiwifruit
resources in China and many superior genotypes of both A.
chinensis and A. deliciosa were selected for further evaluation.
These became the basis for most commercial plantings in
China. Seed of A. chinensis were imported into New Zealand in
1977. The populations raised from these seed are probably the
first plants of A. chinensis to have been grown outside of China,
and are certainly the first such plants known to have survived.
Since then seed of A. chinensis or budwood of superior
selections have been introduced into many different countries.
World production
Kiwifruit are still a minor crop compared to other fruit such
as apples, bananas, grapes or citrus. Production of kiwifruit
amounts to perhaps 0.2% of total world production of fruit.
In only a few countries are kiwifruit an important crop and in
New Zealand they are uniquely important, now being the
single most valuable horticultural export.
Actinidia chinensis and A. deliciosa are widespread in China
and appreciable quantities of fruit (100,000–150,000 t) are
collected from the wild each year. About two-thirds of this is
of A. chinensis, one-third of A. deliciosa and, in addition, there
are much smaller quantities of fruit of other Actinidia species.
Harvesting of wild fruit often involves pulling vines down out
of trees and this, together with general deforestation, means
that the natural resource is under serious threat. However,
kiwifruit collected from the wild are often very small or are
otherwise of poor quality and they are becoming less
important as commercial cultivation increases in China.
In 2002, the total area throughout the world planted in
1
2
Actinidiaceae
kiwifruit was about 120,000 ha. China had 60,000 ha of
kiwifruit orchards, Italy 20,000 ha, New Zealand 12,000 ha,
Chile 8000 ha and all other countries (mainly Greece, France,
Japan and the USA in decreasing importance) a total of about
20,000 ha. Considering only cultivated kiwifruit, in 2002, total
world production was 1,350,000 t: China produced 350,000 t,
about the same as Italy, New Zealand 250,000 t and Chile
150,000 t, so together, these four countries accounted for over
80% of world commercial kiwifruit production.
Approximately 7.5% of current commercial production
would be of cultivars of A. chinensis (almost entirely from
China and New Zealand) and the remainder of A. deliciosa, of
which about 80% would be of ‘Hayward’. This cultivar still
dominates international trade but cultivars of A. chinensis,
especially ‘Hort16A’, are rapidly becoming more important.
Actinidia arguta remains potentially important but at
present world plantings amount to only 100–200 ha.
Uses
Kiwifruit are produced primarily for fresh consumption,
either scooped out with a spoon or peeled. Fruit of A. arguta
are much smaller, about the size of a grape, and can be readily
eaten whole. For all kiwifruit, the premium product is the
fresh fruit, particularly if these can be stored for extended
periods while still retaining quality attributes such as flavour.
Processing usually results in major changes to the colour,
aroma, taste and texture of kiwifruit and most processed
products lack the appeal of fresh kiwifruit. In China, fruit of
A. chinensis are often preferred for processing because they are
sweeter and because their fruit flesh has a clear yellow colour
whereas the chlorophyll-based green of A. deliciosa fruit
changes to a ‘dismal’ brown on processing. Increased
production of yellow-fleshed kiwifruit in countries such as
New Zealand may provide the raw material for more
successful processed kiwifruit products.
Processing of kiwifruit is more important in China than in
other producing countries, partly because its storage facilities
and transport systems are still being developed. Even so, most
fruit are eaten fresh, and only 20–30% of the Chinese
kiwifruit production is processed into a variety of products
such as fruit juices, either natural or clarified, juice
concentrates, jams, fruit preserved either whole or sliced in
syrup, dried fruit, soft drinks and wine and spirits.
Processing is even less important outside of China. The
other three major producers of kiwifruit, Italy, New Zealand
and Chile, are dependent on export of fresh fruit. New
Zealand has a particularly small home market and 85–90% of
kiwifruit produced each year are exported as fresh fruit. Chile
and Italy each export about 75% of the kiwifruit they produce.
Processing in these countries is still largely an attempt to make
use of the significant quantities of fruit that do not meet
export standards. Successful processing will probably require
identification of specific compounds or combinations of
compounds that bestow valuable nutritional or textural
advantages on processed products.
Health benefits
Kiwifruit are among the most valuable nutritionally of all
readily available fruit. Most published information is for
A. deliciosa ‘Hayward’ (Table A.1) but the composition of A.
chinensis ‘Hort16A’ is similar.
Kiwifruit are a good source of potassium and have a high
potassium:sodium ratio. They are also a useful source of
magnesium but other minerals are not sufficient to make a
significant contribution to the diet.
They contain about 2–3% dietary fibre owing to pectins,
oligosaccharides and polysaccharides that are not broken down
and absorbed in the small intestine. A 100 g serving of
kiwifruit will supply about 10% of recommended daily
requirements.
Depending on the individual, kiwifruit can be a strong
laxative, both as bulking agents and for stimulation of motility.
Fresh kiwifruit, kiwifruit juice or dried products are often used
to maintain regularity in bowel movement, especially for older
or sedentary people. There is a huge demand throughout the
Western world for natural laxative products and the laxative
content of kiwifruit may well prove to be, next to their high
vitamin C, their single most valuable attribute contributing to
health.
Kiwifruit are outstanding for their vitamin C content.
‘Hayward’ kiwifruit when harvested contain about 85 mg
ascorbate/100 g fresh weight and very little of this is lost on
storage or ripening. Fruit stored for 6 months at 0°C and then
ripened will still contain at least 90% of the vitamin C present
in the fruit at harvest. Two medium-sized ‘Hayward’ kiwifruit,
even after prolonged storage, can therefore easily satisfy
recommended daily requirements (USA) for vitamin C, which
range from 30 mg for a child to 120 mg for a lactating mother.
‘Hayward’, however, contains only relatively modest amounts
of vitamin C compared to many other kiwifruit cultivars: A.
chinensis ‘Hort16A’ usually contains 20–30% more vitamin C
than ‘Hayward’ and many commonly grown kiwifruit cultivars
in China contain at least twice as much. Fruit of other
Table A.1. Composition of A. deliciosa ‘Hayward’ kiwifruit (100 g fresh
weight edible portion).
Proximate
Water
Protein
Lipid
Carbohydrate
Energy (kJ)
Minerals
Calcium
Copper
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
B1 (thiamine)
B2 (riboflavine)
B3 (niacin)
C (ascorbate)
E (tocopherol)
Folate (mg)
g
80–85
1
0.5
15
250
mg
40
0.16
0.4
25
30
300
5
mg
0.02
0.02
0.5
85
1.6
38
Actinidia
Actinidia species can contain much more vitamin C, up to 1%
fresh weight.
Kiwifruit, however, also have some potential disadvantages
such as high oxalate content and allergenic activity. ‘Hayward’
kiwifruit contain appreciable amounts of oxalate but these are
insufficient to cause a nutritional problem, assuming normal
consumption. Nevertheless, eating some processed kiwifruit
products such as nectars, dried slices or fruit ‘leathers’ can
cause irritation of the mucous membranes of the mouth which
is due, at least in part, to mechanical irritation of the
membranes by oxalate raphides.
Kiwifruit also contain allergens that can cause allergic
responses in susceptible consumers, possibly 2–3% of the total
population. The risk should not be exaggerated: a higher
proportion of the population in some countries show some
adverse reactions to apples, among the most common of all
fruit. Fortunately, extreme allergic responses to kiwifruit are
not frequent.
‘Hayward’ kiwifruit contain large amounts of the highly
active proteolytic enzyme actinidin (E.C. 3.4.22.14). Actinidin
has been implicated in both the laxative and the allergenic
properties of kiwifruit but the enzyme activity does not appear
to be a major health hazard for most people. Actinidin can
cause problems if fresh fruit are incorporated into gelatinebased jellies or are mixed with dairy products.
The anticancer and antimutagenic potential of the fruit are
now being studied. The antioxidant capacity of kiwifruit
likewise requires more study.
Botany
TAXONOMY AND NOMENCLATURE Kiwifruit belong to the
genus Actinidia Lindl., an Asian genus of some 70 species. The
defining characteristics of the genus are:
●
●
●
●
all members of the genus are climbers or scramblers;
all species are dioecious;
the ovary of female flowers is formed by fusion of the lower
parts of numerous carpels but the upper parts of the
carpels remain free forming a distinctive circle of radiating
styles;
the fruit are botanically berries with many seeds embedded
in a fleshy pericarp.
The familiar green and hairy kiwifruit belongs to A.
deliciosa. Until recently, A. deliciosa was treated as a variety of
A. chinensis and was raised to species status only in 1984: prior
to that date, most references in the literature to A. chinensis
actually refer to what is now known as A. deliciosa.
MORPHOLOGY Kiwifruit are vigorous vines which, in the
wild, can grow to the tops of trees, 5–6 m high. Cultivated
plants are tightly controlled into a single trunk, usually about
1.8 m high. The main branches form a permanent framework
and the younger shoots are replaced every 2 or 3 years. Shoots
of the current season come from axillary buds on canes
produced in the previous season. Canes with lateral and
second-order lateral shoots are the typical fruiting units of
kiwifruit vines.
The canopy of a typical orchard vine has a leaf surface area
of 30–40 m2 made up of 4000–5000 leaves. Mature leaves are
large, up to 20 cm in diameter, and the lower leaf surface has a
3
thick felting of stellate hairs. Vines are deciduous and the overwintering buds of A. chinensis and of A. deliciosa are
characteristically different. In A. chinensis, the bud base is
small, and the bud is relatively exposed, being protected only
by bud scales; in A. deliciosa, the bud base is large and
protruding and the bud is almost completely submerged in the
bark.
Flowers are borne either singly (as in most female cultivars)
or in small inflorescences of five to seven flowers (as in most
male cultivars). In general, vines do not flower until 3 or 4
years old, but A. chinensis is noticeably more precocious than
A. deliciosa. Flowers are borne in leaf axils towards the base of
flowering shoots, never terminally. Pistillate flowers are
generally larger than staminate flowers. Flowers are cupshaped, facing downwards, with five or more petals which are
white on opening but within a few days become a rather dirty
golden. The gynoecium is surrounded by whorls of stamens
with bright yellow anthers (almost black in A. arguta). Both
pistillate and staminate flowers have a distinct odour.
Flowering occurs about 2 months after budbreak: flowers are
differentiated in spring and weather conditions during this
period can have a marked effect on flower number and
development. Female flowers remain receptive for about 4
days after opening.
The fruit is a berry with many small seeds embedded in the
juicy flesh (about 250 in a 12 g fruit of A. arguta, at least 500 in
an average-sized ‘Hort16A’, more than 1000 in a ‘Hayward’
fruit) (see Plate 5). Fruit shape and hairiness vary greatly but
fruit of A. chinensis are usually covered with soft, downy hairs
which are often shed early in fruit development; those of A.
deliciosa have persistent, long, hard, bristle-like hairs which are
only partially removed during grading and packing. Fruit of
most commercial cultivars have an elongated ovoidal shape, are
the size of a large hen’s egg and weigh, on average, 80–110 g.
The tough hairy skin of fruit of A. deliciosa is certainly
unpalatable, while that of A. chinensis fruit may not be as hairy
but is still unpalatable and would not normally be eaten.
Consumption of these kiwifruit requires a knife and a spoon.
Some Actinidia species, such as A. arguta and Actinidia
kolomikta (Maxim. et Rupr.) Maxim., have fruit with smooth,
hairless, edible skins. They are ‘snack’ fruit, readily eaten
without creating a mess.
The bright green colour of the fruit flesh of A. deliciosa and
some cultivars of A. chinensis is due to chlorophyll which is
largely retained during fruit maturation, storage and ripening.
In most A. chinensis cultivars, such as ‘Hort16A’ and ‘Jinfeng’,
the fruit flesh is yellowish green to yellow owing to partial or
complete loss of the chlorophyll. ‘Hongyang’ (also A. chinensis)
is even more striking as the flesh around the seeds is red,
making a cross-section of the fruit most attractive.
REPRODUCTIVE BIOLOGY Every female kiwifruit flower
normally sets to form a fruit. There is little subsequent fruit
drop so crop load is largely determined by the number of
flowers. Flower numbers are often limiting and much of vine
management is aimed at ensuring that sufficient flowering
wood of the desired type is retained at pruning.
All Actinidia species are functionally dioecious: flowers of
pistillate kiwifruit may look perfect but the pollen produced is
non-viable; staminate plants produce viable pollen but have
4
Actinidiaceae
only a vestigial ovary and poorly developed styles. Dioecism is
not absolute and different states have been identified: male,
inconstant male, hermaphrodite, inconstant female, female
and neuter. Gradients of male or female sterility are also
found. Among the most easily noticed variants are fruiting
males which carry both staminate flowers and bisexual flowers
which have small ovaries, a few ovules and limited stylar
development: these can produce small fruit.
Most taxa within Actinidia are diploid, x = 29, an unusually
high number possibly indicating ancient polyploidy, but some
are tetraploid (4x), hexaploid (6x) or octaploid (8x). There may
also be intrataxon variation in ploidy: for example, most plants
so far studied of A. chinensis are diploid but plants from a
restricted part of the total geographic range of the species are
tetraploid. These ploidy races cannot be distinguished
morphologically and the only consistent difference so far
noticed is that tetraploid genotypes of A. chinensis flower about
2 weeks later than diploid genotypes. Most important A.
chinensis cultivars in China are tetraploid but ‘Hort16A’ is
diploid. All cultivars of A. deliciosa are hexaploid, 2n = 6x =
174. Most cultivars of A. arguta are tetraploid, 2n = 4x = 116,
but several, known as ‘Issai’ are hexaploid.
Dioecism and ploidy variation have important practical
consequences. A kiwifruit orchard must contain both male and
female plants for transfer of pollen and seed set. Each
kiwifruit that is of commercial size contains many seeds and
there is a strong correlation between seed number and fruit
size. Efficiency of pollen transfer is one of the most important
factors in determining crop yield. In commercial orchards,
about 10% of the canopy area is allocated to male vines set out
in a regular array to ensure that male and female flowers are in
close proximity. Male vines must coincide in flowering with
female vines, they should have an extended flowering period
and they should carry heavy flower loads producing large
amounts of viable pollen capable of setting seed. Most pollen
transfer is effected by honeybees which are brought into the
orchard as female vines start flowering. Kiwifruit flowers are
not particularly attractive to bees and many aspects of orchard
layout and management are designed to keep bees working,
thereby ensuring good pollination. Mechanical pollination,
using pollen collected from male vines, is sometimes used to
supplement natural pollination.
Variation in ploidy level had no practical significance when
only cultivars of A. deliciosa were grown. However, the
cultivars of A. chinensis now grown are diploid or tetraploid.
To ensure fertilization and continued seed growth, female
vines and their pollinators must flower at the same time and
should preferably be at the same ploidy level. Nevertheless,
ploidy levels may not always be critical as A. deliciosa pollen,
which had been collected and stored, has been successfully
used for mechanical pollination of diploid ‘Hort16A’.
FRUIT GROWTH AND DEVELOPMENT Fruit of small-fruited
Actinidia species, such as A. arguta, can reach 80% of their
final size after less than 6 weeks growth and can be harvested
only 100–110 days after pollination, whereas those of A.
chinensis and A. deliciosa are typically harvested 180–210 days
after pollination, depending on cultivar and climate. During
development of the ovary into the mature fruit, linear
measurements increase six- to tenfold and fresh weight and
volume increase several hundredfold. About two-thirds of this
increase in volume or in weight occurs in the first 10 weeks
after pollination and kiwifruit growth shows the double
sigmoidal curve typical of many fruit: a period of very rapid
growth for the first 8 weeks (a period during which fruit of
‘Hort16A’ can increase by 1.6 g/day), a subsequent 3 weeks of
slower growth, followed by a second period of more rapid
growth. Cell division in the inner and outer pericarp ceases
after the first 3 or 4 weeks and subsequent growth is almost
entirely due to cell enlargement.
The chemical composition of the fruit changes during
growth and maturation. Total solids increase over much of the
growing season but the proportion of carbohydrates present as
starch or as soluble sugars changes. Starch at its highest can
account for half the total dry weight of the fruit but about 140
days after pollination, starch begins to decrease and there is an
increase in soluble solids owing to conversion of the starch and
to translocation of sugars from the rest of the vine.
During later stages of fruit growth in A. deliciosa, the
internal appearance of the fruit changes little apart from seeds
changing colour and the loss of starch and softening of the
tissues making the fruit seem juicier and greener. Fruit of
many A. chinensis cultivars are green during initial growth and
development but during fruit maturation chlorophyll is lost so
that the fruit flesh becomes yellow.
CLIMATIC REQUIREMENTS Wild A. chinensis and A. deliciosa
occur mostly on steep hills and mountain slopes. They grow in
relatively damp and sheltered areas and are seldom found
where there is little shade or moisture or where they are
exposed to strong winds. Young plants in particular do best in
shade but sun is required for fruiting. Kiwifruit are abundant
in gullies, under the tree canopy or on forest edges where they
can scramble up through the trees. Winter temperatures can
fall well below 0°C but there is need for a long frost-free
period and abundant rain during the growing season. These
conditions indicate the ideal conditions for successful growth
and cropping of kiwifruit, climatic conditions that are
generally restricted to between latitudes 25° and 45°.
Kiwifruit can be grown extraordinarily well under the
temperate conditions in the Bay of Plenty, New Zealand, but
they are by no means restricted to such conditions and can be
successfully cropped under more rigorous conditions, in
California, Chile or southern Italy, if the vine management
techniques first devised for New Zealand are modified.
Actinidia species differ in their climatic requirements, as
indicated by their natural distributions in China. Actinidia
chinensis is found mainly to the east, A. deliciosa more inland in
colder regions and where both species occur in the same area,
they are separated vertically, with A. deliciosa at the higher,
colder altitudes. Therefore A. chinensis is likely to be more
susceptible to winter cold and to spring frosts, especially as it
breaks bud and flowers about a month ahead of A. deliciosa.
Kiwifruit are temperate plants requiring a period of winter
chilling for adequate budbreak and flowering. Sufficient winter
chilling condenses the period of budbreak, budbreak is more
uniform, there are more flowers and a condensed flowering
period which should reduce fruit to fruit variation in growth
and maturity. Inadequate winter chilling can be a serious
problem in areas with relatively mild winters but cool springs.
Actinidia
Sprays of dormancy-breaking chemicals such as hydrogen
cyanamide can then be applied. Winters can, however, be too
cold. In such places, species such as A. arguta will grow and
crop even if mid-winter temperatures drop to ⫺30°C,
conditions that other kiwifruit will seldom survive. This is not
surprising as A. arguta occurs naturally at latitudes much
further to the north or at much higher altitudes in the south
where A. chinensis and A. deliciosa are common.
Kiwifruit are susceptible to spring and autumn frosts. They
flower about 2 months after budbreak and fruit are ready for
harvest 5–6 months after flowering. A frost-free growing
period of 7–8 months is required.
Kiwifruit vines have very large leaves and very high rates of
water conductivity and transpiration. Transpiration rates can
reach 80–100 l/day. During the growing season, vines need
800–1200 mm of water evenly distributed. Vines are prone to
water stress on windy days or hot sunny days and this can
result in reduced fruit growth. If water is limiting during early
fruit growth, any reduction in fruit size is irreversible.
A characteristic feature of kiwifruit orchards in New Zealand
is the shelterbelts. New Zealand is particularly windy but
inadequate shelter is a major limitation to successful kiwifruit
cultivation in many parts of the world. Young vigorous shoots
that eventually form fruiting canes in the following season are
easily blown out in spring and windrub of developing fruit is a
major cause for rejection at grading. Establishment of young
vines can also be affected by wind. However, there must be a
compromise: shelter may be required but living shelterbelts
compete with vines for water and nutrients and excessive
shading can affect vine growth and flower evocation.
Hail can cause severe damage to young shoots and leaves or
fruit. Hail nets are used in some regions.
Kiwifruit may appear to be demanding in their climatic
requirements but these are largely the conditions for which
cultivars were originally selected in New Zealand. These
cultivars may not be well adjusted to other climatic conditions
but likewise cultivars selected under more extreme conditions
in continental China may not be well suited to New Zealand or
other temperate countries. Management practices may have to
be modified to particular environments.
Horticulture
Kiwifruit propagation is easily achieved by
micropropagation, by rooted cuttings or, occasionally, by root
cuttings. Such plants are clonal. Many are also produced by
grafting onto seedling rootstocks. Under good growing
conditions the method of propagation has no important effects
on vine vigour or productivity although it might be expected
that plants grafted onto seedling rootstocks would be more
variable. Very little use has so far been made of the few clonal
rootstocks that have been selected.
Mature plants can also be readily reworked. This allows
rapid conversion of an established orchard to a new, more
profitable cultivar or replacement of males by better
pollinators. In New Zealand, many mature ‘Hayward’ orchards
have been converted to ‘Hort16A’ by decapitating the plants
and grafting onto the stumps. The existing root systems allow
rapid development of the new canopy with good commercial
yields of the new fruit being achieved in the second or third
year after grafting. Such plants usually consist of a seedling
PROPAGATION
5
rootstock (A. deliciosa), an interstock of ‘Hayward’ (also A.
deliciosa) and a canopy of ‘Hort16A’ (A. chinensis).
STRUCTURES Kiwifruit vines are large and
individual plants can carry 100 kg of fruit. Vines are not selfsupporting and they require support structures that are strong
and can last for 50 years or more. It is false economy to skimp
on support structures. Fruiting canes must be firmly held in
position so that they are not blown out or the fruit damaged by
windrub. Two main types of support structure are used: the
pergola maintains fruiting canes in a plane about 1.8 m above
the ground; with T-bars, the structure of the vine is essentially
similar with fruiting arms held in a fixed position but hanging
towards the ground. T-bar systems are somewhat cheaper and
easier to manage but pergolas give higher yields of good
quality fruit as these are less susceptible to wind damage or to
sunburn. Pergolas are more common for ‘Hort16A’ whose
fruit have thinner skins and are consequently more easily
damaged. Generally there are 400–500 plants/ha.
SUPPORT
TRAINING AND PRUNING
The aim is to establish:
● a well-organized framework of permanent branches;
● a balance between vegetative growth and fruit production;
● a canopy that intercepts light efficiently but is open enough
to allow sufficient light through for flower evocation and
fruit quality;
● a canopy open enough to allow ready access by bees and to
reduce the incidence of diseases such as Botrytis but not so
open or uneven that wind can cause fruit damage;
● a canopy that is easily managed and harvested and keeps
vines to their allocated spaces;
● a canopy that allows the ready production of fruit of the
size and quality required by the market.
Flowers are produced only on shoots of the current season
and usually only on shoots growing from 1-year-old wood.
Fruiting canes should therefore be replaced on a regular 2- or
3-year cycle. New canes should be evenly spaced and at winter
pruning sufficient new wood should be left to provide the
appropriate fruit load – in Italy, about 15–20 winter buds/m2
canopy are recommended for ‘Hayward’. The number of
winter buds retained at winter pruning can be modified with
experience: observations over several seasons will indicate the
likely percentage budbreak, the number of shoots that will
carry flowers, and the number of flowers per flowering shoot.
Higher crop loads can reduce average fruit size, but returns
are usually better for larger fruit. Much new vegetative growth
is removed during summer to ensure that, while replacement
canes are retained, the canopy does not become too dense or
tangled. The amount of summer pruning will depend on the
climate and cultivar as well as the need to protect fruit from
sunburn. With ‘Hayward’, typically 60% of the above-ground
mass of the vine is removed in prunings, leaves and fruit each
season.
Individual cultivars respond differently to management.
‘Hayward’ is one of the least precocious and tends to carry
lighter crops and selection of new fruiting wood is therefore
important. Differences in phenology are also important.
‘Hort16A’, when grown under the same climatic conditions as
‘Hayward’, breaks bud and flowers a month earlier. It is
6
Actinidiaceae
particularly vigorous, especially when grafted onto mature
rootstocks, and vegetative growth continues about a month later
in the season with considerable production of secondary shoots.
Canes developing late in the season are less productive than
those that grew earlier and also tend to flower later with the
fruit maturing later. However, very vigorous canes from early in
the season are also not ideal. Management techniques devised
for ‘Hayward’ cannot be simply transferred to other cultivars.
Yields vary greatly according to country and cultivar. New
Zealand orchards produce about 25 t of ‘Hayward’ fruit per
canopy ha which equates to about 6000 trays of export quality
fruit. ‘Hort16A’ under the same conditions typically produces
higher yields, 10,000 to 12,000 trays/ha, because of its growth
habit and because it is more floriferous. The size of ‘Hort16A’
fruit is routinely increased by use of biostimulants such as
Benefit®PZ. This combination of higher yields and large fruit
sizes has made ‘Hort16A’ more profitable for growers.
Male plants are usually pruned rigorously immediately after
flowering so that their vigorous growth does not shade out
neighbouring female plants.
THINNING If required, thinning is carried out immediately
after fruit set with removal of lateral or misshapen fruit and
then any surplus fruit. If too many winter canes have been laid
down, the weaker canes can be removed.
FERTILIZATION Considerable amounts of nutrients are removed
from orchards but the need for fertilizers should be determined
by leaf or soil analysis. Nitrogen deficiency can cause marked
reductions in vegetative growth and yields whereas excesses are
believed to affect fruit quality and storage. Potassium and
calcium also affect fruit quality. In some areas, iron deficiency is
common. There has been little work to determine relative
requirements of different cultivars or at different crop loads.
Similar types of pests occur on
kiwifruit in the countries in which they are grown and all tend
to be generalists affecting a broad range of plants. Armoured
scales are generally the most serious but although the species
involved are cosmopolitan, the abundance of a particular
species varies according to country. The other main group of
pests, the leafrollers, tend to be specific to each country and
are therefore a quarantine problem as well as damaging the
fruit. Nematodes are a problem in some countries.
Kiwifruit are also susceptible to bacterial and fungal
diseases. Pseudomonas species cause bacterial canker, bacterial
necrosis and, potentially most serious, bacterial blossom
blight. Sclerotinia can also affect fruit on the vine but the other
serious fungal diseases are those that develop while the fruit is
in storage (mainly Botrytis cinerea) or after the fruit is taken
from storage (e.g. Botryosphaeria dothidea).
When kiwifruit were domesticated, they were freed of many
of the pests and diseases to which they are prone in China.
However, as plantings have increased so too have the problems.
Fortunately, the number of pests on kiwifruit is still fairly
limited and they can be well controlled by integrated pest
management systems. Organic production is realistic.
MAIN DISEASES AND PESTS
MATURITY INDICES, HANDLING AND POSTHARVEST STORAGE
Kiwifruit harvested prematurely have a poor colour, an
inadequate flavour when ripened, and a shortened storage life.
The criteria used to decide when to harvest depends on the
cultivar or species. In A. deliciosa, there are no useful visible
indications of maturity, but the rapid increase in soluble sugars
at the final stages of fruit growth is a useful indicator of
physiological maturity. Fruit harvested at a soluble solids
content (SSC) of 6.2–6.5% will store well and be acceptable
when ripened. Dry matter is also used as a maturity index. On
the basis of measurements such as SSC and firmness, ‘Hort16A’
fruit reach physiological maturity about 1 month ahead of those
of ‘Hayward’, but they are not harvested until about the same
time as ‘Hayward’ because they are promoted for their yellow
flesh colour. Loss of chlorophyll is very slow once ‘Hort16A’
fruit are picked and in storage, so they must remain on the vine
until the flesh has a hue angle of 103° or less. At this stage,
firmness may be only 4–5 kgf (as compared to about 7 kgf for
‘Hayward’) and SSC more than 10%. Fruit of A. arguta ripen
unevenly on the vine and are therefore harvested when they
reach a dry matter (DM) threshold (approximately 20% DM)
and about 1% of fruit on the vine are soft. Fruit harvested
earlier are firmer, and are therefore easier to handle, but do not
store as well. Fruit harvested too late are unmanageably soft and
susceptible to mechanical injury.
Once maturity has been reached, the whole crop is normally
harvested. Although fruit are still firm, they must be handled
gently. At harvest, ‘Hort16A’ fruit are softer than those of
‘Hayward’ and they are more vulnerable to damage, especially
because they have the added problem of the sharp ‘beak’ at the
distal end of the fruit. The fruit are not cooled immediately
but left at ambient for several days as this curing helps reduce
the incidence of Botrytis stem end rot.
When harvested sufficiently mature, kiwifruit can be stored
for very long periods. ‘Hayward’ is exceptional and can be
stored for up to 6 months in air under refrigeration and the
fruit will still be acceptable. Controlled atmosphere storage
can extend the storage life even further. However, there can be
a loss of flavour on long-term storage. Other cultivars of A.
deliciosa tend to store less well. Cultivars of A. chinensis
generally have relatively short storage lives and ‘Hort16A’ is
one of the better ones with an expected storage life of 12–16
weeks. Kiwifruit are stored at or close to 0°C. They are very
firm at harvest and soften in store. There is a period of very
rapid softening down to about 1 kgf and then a slow and
gradual softening. Excessive softening, low temperature
disorders such as physiological pitting and low temperature
breakdown, shrivelling from water loss and fungal pitting or
stem end rots all mark the end of storage life in different
cultivars. Low temperature disorders can be avoided by
storage at higher temperatures but then the fruit soften more
quickly. They are very sensitive to ethylene and fruit softening
in coolstore is accelerated if even low levels are present. Shelf
life is 3–10 days depending on the preceding storage period.
Fruit of A. arguta are more delicate than fruit of other
commercial kiwifruit species and their storage life is
correspondingly shorter, at most 10–12 weeks.
MAIN CULTIVARS AND BREEDING In most countries with
commercial kiwifruit orchards, ‘Hayward’ (A. deliciosa) is the
only fruiting cultivar grown and it has become the kiwifruit in
the marketplace (Table A.2). Thus ‘Hayward’ currently
Viburnum
Table A.2. Most important kiwifruit cultivars, 2002.
Cultivar
Area (ha)a Main growing districts
Actinidia deliciosa
‘Hayward’
64,400
‘Miliang No. 1’
‘Jinkui’
‘Bruno’
5,500
2,340
2,000
In all countries growing kiwifruit, including
China
Mainly in Shaanxi, Guizhou and Henan,
China
Guizhou, Henan and Fujian, China
Hubei and Fujian, China
Zhejiang, China
Actinidia chinensis
‘Kuimi’
‘Jinfeng’
‘Zaoxian’
‘Hongyan’
‘Hort16A’
3,080
2,720
2,330
2,180
2,000
Jiangxi, China
Zhejiang and Jiangxi, China
Jiangxi and Zhejiang, China
Sichuan, China
New Zealand
‘Qinmei’
All others
17,480
12,120
China
a Areas
include the 10% of orchard canopy allocated to accompanying
pollinators.
accounts for about 75% of world kiwifruit production, 97.5%
of kiwifruit production outside of China. It was originally
selected because its fruit are large, have a good flavour and can
be stored for extended periods while still remaining acceptable
to consumers. The relative flowering times of ‘Hayward’ and
its pollinators can vary according to climate so, although
‘Hayward’ is grown in many different countries, the
accompanying males grown may vary from country to country.
The situation in China is very different. ‘Hayward’ is grown
but, although it is the second most widely planted kiwifruit
cultivar in China, it accounts for only 13% of the total area
planted in kiwifruit. ‘Hayward’ and the eight other most
common cultivars account for only 80% of total plantings.
There is a strong preference for cultivars selected locally and
most are largely restricted to one or two, usually contiguous,
provinces. Thus ‘Qinmei’ (A. deliciosa) comprises about 30%
of all kiwifruit plantings in China but is predominantly
confined to Shaanxi, the province in which it was selected.
The Chinese kiwifruit industry will probably consolidate on
fewer cultivars of more consistent fruit quality.
The many kiwifruit cultivars in China are selections from
the wild. ‘Hayward’ and ‘Bruno’ (also from New Zealand) were
selections from small seedling populations only one or two
generations removed from the wild. Only one successful
cultivar, ‘Hort16A’, has so far resulted from deliberate breeding
programmes. With its distinctive appearance, its golden-yellow
flesh, and its very different, sweeter, ‘subtropical’ flavour,
‘Hort16A’, commercialized under the name ZESPRITM
GOLD Kiwifruit is perceived as giving the New Zealand
kiwifruit industry a competitive advantage. It is the first
cultivar of A. chinensis to be traded internationally and its
success is likely to encourage the development of competitive
cultivars, either some of the existing Chinese cultivars or
cultivars specifically bred for the purpose.
Ross Ferguson
Further reading
Costa, G. (1999) Kiwifruit orchard management: new developments.
Acta Horticulturae 498, 111–119.
7
Ferguson, A.R. (1984) Kiwifruit: a botanical review. Horticultural
Reviews 6, 1–64.
Ferguson, A.R. (1990) Kiwifruit management. In: Galletta, G.J. and
Himelrick, D.G. (eds) Small Fruit Crop Management. Prentice
Hall, Englewood Cliffs, New Jersey, pp. 472–505.
Ferguson, A.R. (1990) Kiwifruit (Actinidia). In: Moore, J.N. and
Ballington, J.R., Jr (eds) Genetic Resources of Temperate Fruit and
Nut Crops. International Society for Horticultural Science,
Wageningen, The Netherlands, pp. 603–653.
Ferguson, A.R. and Stanley, R. (2003) Kiwifruit. In: Caballero, B.,
Trugo, L. and Finglas, P. (eds) Encyclopedia of Food Sciences and
Nutrition, 2nd edn. Academic Press, New York, pp. 3425–3431.
Huang, H. and Ferguson, A.R. (2001) Kiwifruit in China. New
Zealand Journal of Crop and Horticultural Science 29, 1–14.
Patterson, K., Burdon, J. and Lallu, N. (2003) ‘Hort16A’ kiwifruit:
progress and issues with commercialisation. Acta Horticulturae
610, 267–273.
Perera, C.O., Young, H. and Beever, D.J. (1998) Kiwifruit. In: Shaw,
P.E., Chan, H.T. Jr and Nagy, S. (eds) Tropical and Subtropical
Fruits. AgScience Inc., Auburndale, Florida, pp. 336–385.
Steven, D. (1999) Integrated and organic production of kiwifruit.
Acta Horticulturae 498, 345–354.
Warrington, I.J. and Weston, G.C. (1990) Kiwifruit: Science and
Management. Ray Richards Publisher in association with the New
Zealand Society for Horticultural Science, Auckland, NZ,
pp. 1–576.
ADOXACEAE
Viburnum spp.
viburnums
The genus Viburnum, Adoxaceae (formerly Caprifoliaceae),
comprises more than 200 species throughout the northern
hemisphere, primarily Asia and North America, and many
produce copious amounts of fleshy fruit. Many are used as
ornamental plants in the landscape as they can have highly
scented, attractive, showy flowers and masses of variously
coloured fruit borne on attractive plants that may have
deciduous or evergreen foliage. Some are extremely good food
sources and none are known to be toxic, although some will
cause nausea if large quantities of raw fruit are consumed. At
various times, several similar species have been given the name
highbush cranberry and they are commonly harvested from
the wild and consumed.
Several similar edible Viburnum species with tart, red fruit
have been given the name highbush cranberry. They are
commonly harvested from the wild. Thus, Viburnum opulus,
Viburnum edule and Viburnum trilobum are now considered to
be subspecies of V. opulus (V. opulus opulus, V. opulus edule, V.
opulus americanum formerly V. trilobum). Viburnum opulus will
be used here as the general term. In addition to V. opulus,
many other Viburnum species are considered edible. The
highbush cranberry species are native to the northern
hemisphere but have been scattered throughout the temperate
regions of the world due to their highly ornamental
characteristics. While generally a woody shrub 2–3 m tall, at
the northern, very cold edges of its range it may not get any
taller then 0.3 m and in ideal circumstances it may reach 5 m.
Highbush cranberry is adapted to a wide range of soils but is
most productive on moist, reasonably fertile soils.
In the spring, the flowers bloom on large, showy, cymes.
8
Adoxaceae
The individual flowers are small and white but a cyme may
contain hundreds of flowers. The flowers are insect pollinated
and can be fragrant. Flowering takes place in the spring and
the fruit develop over the summer, finally ripening in late
summer and autumn. The fruit are most typically a bright red
colour, or yellowish, and individually are 0.8–1 cm in diameter.
The fruit of V. opulus opulus are usually described as
astringent, whereas the fruit of V. opulus americanum, while
tart, are considered to have a very good flavour. Since the fruit
is very tart, much like a cranberry, it is seldom eaten raw;
rather it is often blended with sugar in processed jelly or sauce
type products.
For commercial production in managed stands, plants
should be established 2–3 m apart within rows. Irrigation is
important for plant establishment and for maximum fruit yield
and quality. Nitrogen fertilization will also be important for
best production and plant health. No chemicals are approved
for weed, insect or disease control. When grown in their native
range, diseases should cause minimal problems if the plants are
healthy, however, bacterial leaf spot (Pseudomonas viburni),
powdery mildew (Sphaerotheca macularis) and shoot blight
(Botrytis cinerea) have been reported as potential problems.
Occasionally aphids are reported as a problem. In the early
1900s, improved cultivars with larger fruit, greater production
and better fruit quality were released, these include ‘Hahs’,
‘Andrews’, ‘Wentworth’, ‘Manitou Pembina’ (syn. ‘Manitou’)
and ‘Phillips’. Nearly all fruit that is now consumed is
harvested from wild stands, with locally important industries
selling to speciality markets.
In Native American culture, the fruit were prized for food
and the roots, bark, twigs and fruit were used to treat various
maladies including use as a cold remedy, pulmonary aid,
cough medicine and throat aid, antidiarrhoeal, cathartic and
others.
Chad E. Finn
Further reading
Darrow, G.M. (1923) Viburnum americanum as a garden fruit.
Proceedings of the American Society for Horticultural Science 21,
44–55.
Moerman, D.E. (1998) Native American Ethnobotany. Timber Press,
Portland, Oregon.
Plants for a Future. Available at: http://www.pfaf.org/index.html
(accessed 3 November 2006).
Stang, E.J. (1990) Elderberry, highbush cranberry, and juneberry
management. In: Galletta, G.J. and Himelrick, D.G. (eds) Small
Fruit Crop Management. Prentice-Hall, Englewood Cliffs, New
Jersey, pp. 363–382.
ANACARDIACEAE
Anacardium occidentale
cashew
Cashew, Anacardium occidentale L. (Anacardiaceae), is one of
the important edible nuts that is consumed worldwide.
Common names include: in Arabic habb al-biladhir; in
Bengali hijlibadam, hijuli; in Chinese yao guo and yao guo
shu; in French acajou a pommes, noix-cajou, noix d’acajou,
pomme d’acajou; in Hindi kaaju; in Malay gajus, jambu golok
and jambu mede; in Nepali kaaju; in Portuguese caju and
cajueiro; in Spanish anacardo, maranon, casho; in Swahili
mbibo and mkanju; in Tamil mindiri; and in Thai mamuang,
yaruang. The Vavilovian South American centre VIII
(Brazil–Paraguay) is considered the centre of origin.
Prior to the Portuguese colonization of Brazil, indigenous
Indians consumed both the nut and the enlarged pedicel
(receptacle), which is referred to as the ‘cashew apple’. Juice
squeezed from the cashew apples was fermented to produce
wine. The Brazilian Indians roasted nuts over a fire thus
burning off the toxic outer covering and the Portuguese
colonizers copied this method. The trees are now often found
growing wild on the drier sandy soils in the central plains of
Brazil and are cultivated in many parts of the Amazon
rainforest. By 1750, the cashew was widely distributed
throughout tropical America. The cashew trees were planted
as a backyard tree, partly for shade and established beyond
their indigenous coastal distribution.
The Portuguese introduced cashew trees to India in the
18th century, where they were initially grown for producing
wine and brandy and later introduced to other Asian countries.
The Portuguese also exported the seeds to their colonies in
East Africa in the late 18th century where they quickly became
naturalized and grew wild along the Mozambique coast. From
there they were introduced and naturalized in other East
African countries such as Kenya and Tanzania. Soon, the
African people started selling the wild harvested nuts to
Portuguese traders, who in turn sold them to merchants in
India for processing.
Cashews have spread widely in the Indian Ocean region and
have become naturalized in seashore habitats. The trees were
planted in all suitable areas of tropical India, and in the 1950s
quite large orchards were planted, chiefly in the Indian state of
Kerala. Trade in cashew nuts started at the beginning of the
20th century and grew particularly fast in the 1930s, being
dominated mainly by Indian production. Since the 1960s,
there has been rapid growth in the industry, particularly in
India, Madagascar and Mozambique.
World production and yield
Cashew now grows all along the sea coasts in tropical regions
starting from southern America to the West Indies, west and
east Africa and India. India is the largest producer of raw
cashew nut in the world (Fig. A.1). In India, the state of
Kerala is the largest producer, processor and exporter of raw
nuts. Other cashew-growing Indian states are Andhra Pradesh,
Orissa, Goa, Karnataka, Maharashtra, Tamil Nadu and west
Bengal. However, in the later part of the 20th century, other
countries such as Brazil, Vietnam, Tanzania, Mozambique,
Guinea Bissau, Nigeria and Indonesia also started developing
cashew plantations.
Global trade in raw cashews now takes place from over 24
countries. In north-eastern Brazil, cashew is grown in the
states of Ceara, Piaui and Rio Grande do Norte which
together account for 90% of Brazil’s cashew production. The
northern province of Nampula is the major contributor to
cashew production in Mozambique. In Tanzania, cashews are
grown in the Mtwara, Lindi, Ruvuma, Tanga and coastal
regions. In Kenya, cashew is grown in the narrow coastal belt
covering the districts of Kilifi, Kwale and Lamu.
India, Brazil, Mozambique, Tanzania and Kenya together
Anacardium
Table A.3. Proximate fruit composition of cashew apple and nut which has
an edible flesh to fruit ratio of 87% (Source: USDA, 2003; Leung et al., 1972).
Area (ha) and production (t)
3,000,000
2,500,000
Area harvested, world
Production, world
Area harvested, India
Production, India
Proximate
2,000,000
1,500,000
1,000,000
500,000
0
1961
1966
1971
1976
1981
9
1986
1991
1996
Year
Fig. A.1. Cashew production and area harvested in the world and India.
contribute more than 80% of the total raw nut production in
the world. While the area planted has increased in many of the
countries over the past three decades, production has
stagnated which resulted in a huge drop in productivity
between 1975 and 1987. Vietnam in the past few years has
gained ground as a significant producer of raw nuts and
exporter of cashew kernels to the international market.
Water
Energy (kcal)
Energy (kJ)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
Apple
(g/100g)
85.4
53
0.8
0.4
13.1
0.4
0.3
Nut
(g/100g)
5.2
553
2314
18.2
43.9
30.2
3.3
2.54
Minerals
mg
mg
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
7
0.6
37
6.7
292
593
660
12
Vitamins
18
124
7
mg
198
0.02
0.01
0.5
50 IU
mg
Uses and nutrient composition
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
0.5
0.42
0.06
1.06
0
The true fruit of the tree is the cashew nut that consists of a
double shell containing a caustic phenolic resin in
honeycomb-like cells, enclosing the edible kidney-shaped
kernel. An interesting feature of the cashew is that the nut
develops first and when it is full-grown but not yet ripe, the
peduncle (receptacle) fills out and becomes plump, fleshy,
pear-shaped or rhomboid-to-ovate, 5 to 12 cm in length. The
fruit has a waxy, yellow, red or red-and-yellow skin and
spongy, fibrous, very juicy, astringent, acid to subacid, yellow
pulp. This swollen receptacle is the ‘cashew apple’. In Brazil,
the cashew apple is sold fresh and the juice is widely available.
However in India the apple is not widely used.
Cashew kernels have the highest protein content among tree
nuts (19.5%). This protein content matches soybean and is
higher than peanut. All the essential amino acids are present.
Its crude fibre is low and it has a high lipid content (46%).
The lipids composition is halfway between peanut and olive
oil and the oil contains all the fat-soluble vitamins. The ratio
of saturated to unsaturated fatty acids is 4:1, close to the ideal
5:1. Cashews, with about 45 g of fat/100 g serving (at least a
quarter of which are monosaturates) and a particularly high
carbohydrate content for a nut at about 30 g/100 g portion,
provides 553 calories/100 g intake (Table A.3). The kernels are
also very high in magnesium, having only slightly less than
almonds.
The cashew apple is used either for juice or preserved in
syrup (candied). The fresh apples are very astringent due to
their high tannin content, and are much more palatable if first
processed to remove the bitter taste. This can be accomplished
either by steaming under pressure (pressure cooked) for 10–15
min, or boiling in salty water for 15 min. The apples are then
pressed to remove excess moisture and boiled in cane sugar
syrup for 2 h. Finally they are sun dried, or placed in an
electric food drier. Cashew apples are also canned in syrup,
used to prepare chutneys and fruit pastes, and because of the
high pectin content, readily set when making jams.
The juice can also be extracted and strained, after which
gelatin is added. The tannins in the juice bind to the gelatin
and form a precipitate, which can then be removed by
filtering. Sugar can then be added to taste. The juice readily
ferments and is used in various countries to prepare wines and
distilled liquors (e.g. Brazil, Guatemala, West Africa, India,
Sri Lanka and the Philippines).
The cashew tree is used for reforestation, to prevent
desertification, and as a roadside buffer tree. Cashew was first
planted in India in order to prevent erosion on the coast. The
wood from the tree is used for carpentry, firewood and
charcoal. The tree exudes a gum called cashawa that can be
used in varnishes or in place of gum arabic. Cashew bark has
about 9% tannin, which is used in tanning leather.
The bark and leaves of the tree are used medicinally, the
cashew nut has international appeal and market value, and
even the shell around the nut is used medicinally. Cashew
apple juice, without removal of tannin, is prescribed as a
remedy for sore throat and chronic dysentery in Cuba and
Brazil. Fresh or distilled, it is a potent diuretic and is said to
possess sudorific properties. The brandy is applied as a
liniment to relieve the pain of rheumatism and neuralgia.
In 16th-century Brazil, cashew apples and their juice were
taken by Europeans to treat fever, to sweeten breath and to
‘conserve the stomach’. The indigenous tribes had used parts
of the cashew tree, the nut and apples for centuries. The
Tikuna tribe in northwest Amazonia use the ‘apple’ juice for
influenza and for warts. In traditional medicine, the fruit juice
is used against diarrhoea. The shell oil (cardol) is used to heal
10
Anacardiaceae
foot wounds. The decoction of bark and leaves is used against
diarrhoea and abdominal pains, inflammation and diabetes.
Scientific tests have shown that sodium anacardate destroys
certain snake venoms as well as tetanus and diphtheria toxins,
and the vegetative form of anaerobic bacteria. Modern
medicine uses cardol for its vesicant properties, as a dye for
skin pigmentation, as respiratory and circulatory analeptic and
as an antagonist of barbiturics.
Cashew apple contains calcium, phosphorus, iron and
vitamin C (ascorbic acid) and is regarded as having skin
conditioning activity due to its proteins and mucilage. Extracts
are used in body care products, as coadjutant in the treatment
of premature ageing and to remineralize the skin. It is also a
good scalp conditioner and tonic, often used in shampoos,
lotions and scalp creams.
The cashew nutshell liquid (CNSL) is used as a protective
agent, in such products as varnishes and cements, and ink is
extracted from the shell. This CNSL oil is composed of
anacardic acid (71.7%), cardol (18.7%), cardanol (4.7%), a
novel phenol (2.7%) and two unknown minor ingredients
(2.2%). Each of the phenolic constituents contains monoene,
diene and triene cardanols. Anacardic acids, 2-methylcardols
and cardols in fruit have been found to exhibit tyrosinase
inhibitory activity.
Industrially, CNSL has been used for many years as a
component in the manufacture of friction dusts. The dual
phenolic/alkenyl nature of CNSL makes it an ideal natural raw
material for the synthesis of water-resistant resins and polymers.
The sap from the bark provides indelible ink. When the
trunks of the trees are tapped, a milky juice exudes, which is
white when fresh, but turns black on exposure, and, in India,
is used as varnish. It stains cotton or linen deep black upon
exposure to air. A tar prepared from the pericarp is used in
tarring woodwork and boats to protect them from the ravages
of insects, and the gum from the stems is employed by
bookbinders to insure the bindings from destruction by
worms and book-pests.
(e.g. deep sandy loams), the tree develops a pronounced
taproot. Cashews rapidly develop an extensive system of
lateral roots that reach far beyond the edge of the canopy.
The simple smooth leaves occur mainly in terminal clusters
and are arranged alternately on the stem on a short petiole
with prominent veins. Each leaf is 10–20 cm long and
5–10 cm wide, oblong–oval or obovate (Fig. A.2). The mature
leaves are green and leathery with young leaves ranging in
colour from golden to red and pliable.
Cashew flowers are borne in 10–20 cm terminal panicles
and consist predominantly of staminate flowers and some
perfect (hermaphroditic) flowers. There are no pistillate
flowers. Individual flowers are sweet-smelling, small with
usually five yellowish-green petals, each about 1.5 cm long.
The true fruit is a kidney shaped nut consisting of a doublewalled shell. The shell has an outer thick exocarp, and an inner
hard endocarp separated by a resinous, cellular mesocarp and
encloses the edible kernel. The nut is green at first, but becomes
a greyish brown as it develops. As the nut approaches maturity,
the pedince (receptacle) becomes swollen and fleshy, forming a
5–12 cm yellow or red, juicy, pear-shaped pseudo-fruit.
ECOLOGY AND CLIMATIC REQUIREMENTS Cashew grows best
in a warm, moist, tropical climate with a well-defined dry
season of 4–5 months during the reproductive phase, followed
by a wet season of 4–5 months (1000–2000 mm rainfall).
Cashew requires an equable environment with a maximum
temperature of 34°C and a minimum of 20°C. Damage occurs
to young trees or flowers below the minimum temperature of
7°C and above the maximum of 45°C. Prolonged cool
Botany
TAXONOMY AND NOMENCLATURE The Anacardiaceae includes
76 genera with over 600 species including Spondias mombin
(hog plum), Mangifera indica (mango), Pistacia vera (pistachio
nut), Semecarpus anacardium (Indian marking nut tree),
Metopium toxiferum, Comocladia dodonaea, Schinus molle
(Peruvian pepper tree) and Toxicodendron vernicifluum (lacquer
tree). About 25 genera contain poisonous species. The
synonyms for cashew, A. occidentale L. are Acajuba occidentalis
Gaertn. and Cassuvium pomiferum Lam.
Under ideal tropical conditions the cashew is
an attractive, erect, 7–15 m evergreen tree, with smooth brown
bark, and a dense symmetrical, spreading canopy. Branching
occurs very low on the trunk, with the lowest limbs often
touching the ground where they can take root. More usual,
where conditions are less than optimal, the tree grows to no
more than 5–7 m and can develop an ill-defined trunk and a
sprawling, straggly growth habit. Such trees are of less
ornamental value to the landscape, though the colourful fruit
remains an attractive feature. Where soil conditions permit
DESCRIPTION
Fig. A.2. Leaf, flower and fruit of Anacardium occidentale (Source:
Verheij and Coronel, 1992).
Anacardium
temperatures will damage mature trees, though such trees can
survive temperatures of about 0°C for a short time. The
optimum sunshine is 1285 h (9 h/day) in the flowering/fruit
set period. It is grown at altitudes from sea level to 700 m and
thrives well between 27°N and 28°S latitudes. Most of the
regions where it is an economically important crop are
between 15°S and 15°N. It grows usually unirrigated but
responds to summer irrigation. Cashew flourishes in the hot,
dry tropics along sea coasts.
Cashew is often grown on marginal soils and also on
wasteland mostly unsuitable for other economic crops. The
tree requires good drainage, friable soils and can tolerate a
wide pH range and even salt injury. It is found along sandy sea
coasts, fairly steep laterite slopes or rolling land with shallow
top soils in India; alluvial soils in Sri Lanka; ferruginous soils
in East and West Africa, Brazil and Madagascar; and volcanic
soils in the Philippines, Indonesia and the Fiji Islands. The
most fertile soils for cashew are virgin forest soils.
Cashew is known as a ‘poor man’s crop’ and is good for
smallholders because it will grow with minimal fertility.
Under these unfavourable conditions it produces a harvest,
although a low one. Many cashew groves are intercropped
with coconut or annual crops.
REPRODUCTIVE BIOLOGY Fruit are produced after 3 years,
during which lower branches and suckers are removed. Full
production is attained by the tenth year and a tree continues to
bear until it is about 30 years old. Cashew flowering is
unaffected by daylength. In a seasonally dry climate, flowers
are produced immediately after the rainy season. In tropical
climates, that have wet and dry periods throughout the year,
flowering can occur at any time.
Cashew is an andromonecious (polygamomonecious) plant
with staminate and hermaphroditic flowers intermixed on the
same panicle. On the same tree, the hermaphroditic or perfect
flowers are bigger in size than the staminate or male flowers.
The flowers produced early in the panicle are mostly male.
Perfect flowers are generally produced about 1 month after the
first flowers are produced in the panicle.
The petals turn from white or creamy white to pink and
become recurved as the flower fully opens. The stigma is
immediately receptive, however, pollen release occurs later,
thereby permitting an opportunity for cross-fertilization. On
opening, flowers are receptive to pollen for only a day. The
presence of scented flowers and sticky pollen is circumstantial
evidence of an important role for insect pollinators. Studies to
date have implicated both wind and a variety of insects as
pollinating agents, but there is no information on their relative
importance. Beehives are placed in or near the orchard to
improve the yield. The flowering period is 2–3 months in
length (during the dry season), with the fruit appearing 2
months later.
FRUIT GROWTH AND DEVELOPMENT The fruit is unusual in
that the kidney-shaped nut, the true fruit, is borne on a greatly
enlarged, fleshy receptacle. This structure, the cashew apple, is
about 5 cm in diameter and 7–10 cm in length. At maturity,
approximately 2–3 months after flowering, the fruit turns
bright yellow or red. Flowers and fruit at various stages of
development are often present on the same panicle.
11
Horticulture
Cashew can be propagated by seed, air layers
and softwood grafts. Seeds germinate in 7–10 days. Since
it is a cross-pollinated crop, vegetative propagation is
recommended to obtain true progeny. Mother trees having the
following characteristics are selected: (i) good health, vigorous
growth and intensive branching habit with panicles having a
high percentage of hermaphroditic flowers; (ii) trees 15–25
years of age; and (iii) nuts of medium size and weight (5–8
g/nut) with an average annual yield of 15 kg nuts, and 7–8
nuts per panicle. Different methods of grafting such as
epicotyl grafting, softwood grafting, veneer grafting, side
grafting and patch budding have been tried in cashew with
varying degrees of success. Softwood grafting is the best for
commercial multiplication of cashew. Softwood grafts can be
prepared almost throughout the year with a mean graft
success of about 60–70%. Higher success is achieved during
the monsoon season. The softwood grafts are ready for
transplanting in 5–6 months after grafting. A successful
micro-grafting technique has been developed using in vitro
germinated seedlings as rootstocks and axenic shoot cultures
from shoot-tip and nodal cultures as micro-scions. Approach
grafting, as well as layering, are most successful if carried out
just prior to the pre-flowering flush of growth. Whip grafting
may be more successful if it is done immediately after the fruit
has ripened. Budding should be done about 1 month after
flowering begins.
Cuttings are sometimes difficult to root, although ringing
the cutting 40 days prior to removal from the parent plant has
sometimes improved rooting. Cuttings are taken from 1–2year-old shoots whose stems are still light-coloured and
somewhat flexible. Field establishment of air layers is poor.
PROPAGATION
Planting is done in pits of 60 ⫻ 60 ⫻ 60 cm
during June–July. Seeds are either sown in situ or seedlings
raised in polybags are transplanted at the onset of the
monsoon. Initial close planting at 3, 4, 5 m apart, results in
early shading of the soil surface, suppresses weeds, conserves
soil and moisture and provides high initial yield per unit area,
but requires thinning to the final spacings of 8, 9, 10 m at 5 or
6 years when canopies and root systems are intermingled with
those of neighbouring trees. Recommended spacing is 10 ⫻
10 m, thinned to 20 ⫻ 20 m after about 10 years, with
maximum planting of 250 trees/ha. Once established, the field
needs little care. Intercropping may be done for the first few
years, with cotton, groundnut or yams. Proper staking of the
plants is required to avoid lodging resulting from wind during
the initial years of planting.
CULTIVATION
Sprouts coming from the rootstock
portion of the graft should be removed frequently during the
first year of planting. Initial training and pruning of young
cashew plants during the first 3–4 years is essential for
providing proper shape. Thereafter, little or no pruning is
necessary. The plants should be allowed to grow by maintaining
a single stem up to 0.75–1.00 m from ground level. This can be
achieved by removing the side shoots or side branches
gradually as the plants start growing from the second year of
planting. Weak and crossed branches can also be removed.
Initial training and pruning of cashew plants facilitate easy
TRAINING AND PRUNING
12
Anacardiaceae
cultural operations such as terrace making, weeding, fertilizer
application, nut collection and plant protection. The flower
panicles that emerge from grafted plants during the first and
second year of planting should also be removed (deblossoming)
in order to allow the plant to continue to grow. The plants are
allowed to flower and fruit only from the third year onwards.
In older cashew plantations, removal of dried or dead wood,
crossed branches, and water shoots should be attended to at
least once in 2–3 years. This allows proper growth of the
canopy and receipt of adequate sunlight on all the branches.
Pruning of cashew plants should be done during May/June in
the northern hemisphere.
Topworking is used to rejuvenate unproductive and mature
trees from 5 to 20 years of age. The unproductive trees are cut
at a height of 0.75–1.00 m from ground level in May–
September. Sprouts emerge in 30–45 days and new, 20–25day-old shoots should be grafted with scions of high-yielding
cultivars using the softwood grafting technique. To ensure at
least six or seven successful grafts, 10–15 grafts are performed
on the new shoots of every tree. The best season for grafting is
July–November. Thinning of the extra shoots arising from the
stumps should be done to obtain better growth of the grafts.
Sprouts below the graft joint should be removed. The topworked trees start fruiting from the second year. A major
disadvantage is the loss of trees due to stem borer attack.
Cashew plantations are usually kept well weeded for ease of
harvesting and to prevent competition for water and nutrients.
In some areas, the cleared ground also acts as a firebreak.
NUTRITION On sandy soils, laterite soils and on sloping land
with heavy rainfall, fertilizer is applied in a circular trench
25 cm wide and 15 cm deep, at a distance of 1.5 m from the
trunk. On red loamy soils with low rainfall, the fertilizer should
be incorporated into the soil in a band 1.5 m wide, at a distance
of 1.5 m (inner edge) to 3 m (outer edge) round each tree.
Cashews respond well to fertilizer, but there are no specific
recommendations and the following are suggested. For mature
trees an annual total of 15–20 kg of fertilizer is applied,
splitting the applications into thirds: one is applied at panicle
emergence, another as the fruit approach maturity and a final
application in August. A fertilizer dose of 750 g N, 325 g P2O5
and 750 g K2O per plant is recommended for cashew. Apply
one-fifth of the dose after the completion of the first year, twofifths of the dose during the second year and thus reaching full
dose from the fifth year onwards.
IRRIGATION Cashew trees are generally grown under rain-fed
conditions. During summer, it is advisable to provide
irrigation of 200 l/plant at fortnightly intervals. Cashew will
not withstand water logging and proper drainage is essential.
The use of black polyethylene sheet for mulching and
fortnightly irrigation increased fruit retention.
HARVESTING AND POSTHARVEST HANDLING The colour of
the nut changes from brownish green at fruit set to light green
when the apple is one-quarter grown and turns grey
thereafter, irrespective of cultivar. The nuts are harvested at
this grey stage to avoid collecting immature nuts. The mature
fruit fall to the ground as the ‘apple’ dries. In wet weather,
fruit are gathered each day and then dried for 1–3 days.
Harvesting of raw nuts is done either by collection after
natural drop, after thrashing the tree with a stick or by tree
shaking. The nuts are dried in the sun on bamboo mats, being
turned for several days until they rattle in the shell. Care is
taken to avoid overheating of the nuts. Overheating can lead to
breakage of the shell liquid structure and the leakage of caustic
oil to the kernel.
Mechanical shelling has been unsuccessful, so hand labour
is required. Cashews are usually roasted in the shell (to make it
brittle) and then cracked, the nuts removed and vacuum
packed. In India, part of the nuts are harvested from wild trees
by people who augment their meagre income from other crops
grown on poor land. Kernels are extracted by people skilled in
breaking open the shells with wooden hammers without
breaking the kernels. Nuts are separated from the fleshy
pedicel and receptacle, the seedcoat removed by hand and the
nuts dried. Fresh green nuts from Africa and the islands off
southern India are shipped to processing plants in western
India.
Processing of cashew is complicated and costly when done
on a large scale because of the caustic oil, CNSL. The
traditional method of processing the nuts is to roast them over
a fire in a perforated pan to burn off the CNSL. The nuts
swell and release the CNSL, which drips through holes of the
pan into the fire. This causes an abundance of thick irritating
smoke. Next the nuts are tumbled in ashes or sawdust to
absorb the rest of the CNSL. Shells are then removed by
hand. The kernels processed this way are of low quality and
used mostly for local consumption. For export-quality kernels
a large infrastructure including machinery, factories and
personnel is needed. A hot-oil bath is used to remove the
CNSL.
The nuts are graded according to size and colour. Miniprocessing factories process 500–1500 kg of raw nuts/day and
employ 25–200 people/t of processed cashew. Humans are
better at separating the nut from the shell than machines and
kernel breakage is less at the mini-factories.
One of the biggest costs is stockpiling cashew to keep the
mini-factories operating. The cashew-harvesting season is only
about 2 months long. To keep the plant operating for 200
days/year, the smallest plants need to store about 100 t of raw
cashew.
Helopeltis anacardii is a sap-sucking
insect that can cause flower damage and is the major insect
pest in South-east Asia, India and East Africa. A severe attack
will result in up to 80% of branches being damaged. Helopeltis
anacardii usually appears with the emergence of new flushes
and panicles and the symptoms are drying of the inflorescence
and dieback of shoots. Pesticide is sprayed three times, first
with the emergence of new vegetative flushes in
October–November, second at the commencement of panicle
emergence in December–January and the third at completion
of flowering/initiation of fruit set in January–February.
The green ant (Oecophylla smaragdina), the meat ant
(Iridomyrmex sanguineus), mantis (Orthoderinae sp. and
Mantidae sp.), predatory bugs (Geocoris australis) and spiders
(Oxyopes sp.) significantly reduce the numbers of Heliopeltis
spp. Green ants were the most abundant predatory species in
cashew plantations. Green ants also significantly reduce the
PESTS AND DISEASES
Anacardium
numbers of several other cashew insect pests such as the fruit
spotting bug (Amblypelta lutescens), the mango tip borer
(Penicillaria jocosatrix) and the leaf roller (Anigraea ochrobasis).
Other insect pests include borers, thrips, mealy bugs, weevils,
caterpillars and leaf miners.
Cashew stem and root borer (Plocaederus ferrugineus) is a
serious pest, which is capable of destroying the cashew tree.
Main symptoms of attack are yellowing of leaves, drying of
twigs, presence of holes at the base of the stem with exuding
sap and frass. To reduce the spread of infestation, it is essential
to remove the dead trees and trees in an advanced stage of
infestation at least every 6 months. Anthracnose
(Colletotrichum gloeosporioides) is prevalent in cashew plantations during the rainy season. The main symptom is the
appearance of white patches on branches followed by drying of
twigs from the tip. The affected parts are chiselled out and
Bordeaux paste is applied.
Powdery mildew (Oidium sp.) kills the flowers and can have
a devastating effect on cashew tree yields. Powdery mildew is a
significant problem in East Africa. Powdery mildew likes cool,
humid conditions and succulent plant growth. It does not
tolerate high temperatures or high ultraviolet light
concentrations. It can reproduce in 48 hours, releasing
millions of spores into the atmosphere. To improve cashew
yields in powdery mildew areas without using chemicals,
pruning suckers on lower branches to let in more sunlight can
help. These tend to be highly infected by powdery mildew and
a source of spores for future infection.
Other diseases of cashew include dieback, damping off and
anthracnose. Anthracnose is caused by the fungus C.
gloeosporioides and, under wet conditions, can cause almost
total crop failure. It also affects other tropical fruit trees such
as mango and citrus.
Pesticide-free plantations would reduce costs and prevent
health hazards and environmental damage. Encouraging an
ecosystem conducive to beneficial organisms, especially ants,
which prey on cashew pests would be one strategy. The
nectaries (pits which secrete nectar) on cashew trees
apparently attract ants to places where the trees are susceptible
to pest damage (especially the young leaves, developing
inflorescences and young fruit). The nectaries also attract
spiders and predatory and parasitic wasps. All of these insects
then prey upon the cashew insect pests. The number of
nectaries increases as the tree becomes larger, and therefore
more ant species are found in older trees. In younger
plantings, it may be advisable to encourage a more diversified
habitat to attract more ants. Creating desirable habitats for
ants and other beneficial insects is done by mixed plantings,
having brush and grass understories, and leaving dead wood
and flat stones in the area.
CULTIVARS AND BREEDING Until recently, cashew breeding in
many countries has been limited to a selection programme,
attempting to find what is best from local material and foreign
introductions. For commercial production, clonal propagation
of superior genotypes must be the basis for the industry. A
number of hybridization programmes are being carried out to
select superior genotypes. Hybridization between local and
introduced material aims to combine complementary qualities
from parents with contrasting characteristics, taking care to
13
prevent inbreeding depression by avoiding parents with a
common ancestry. The seed produced is germinated and
grafted onto a mature seedling rootstock to be appraised.
Objectives include hypersensitivity to powdery mildew, bud
vigour and kernel quality. The selected plants are multiplied
by budding or grafting for trials. The plants are appraised over
3 consecutive crop years for growth and vegetative habit,
Helopeltis tolerance, powdery mildew resistance (with and
without chemical control) and yield in terms of nut weight,
number and quality.
In India more than 35 cultivars of cashew are available that
have been recommended for use in different states where
cashew is grown. In Kerala, new cultiars released include
‘Raghav’, ‘Damodar’, ‘Priyanka’ and ‘Amrutha’. The older
cultivars are ‘Anakkayam 1’ and ‘Madakatthara 1’. In Tamil
Nadu ‘VRI12’ and ‘VRI13’ are popular cultivars. The dwarfprecocious cashew clones ‘CCP06’, ‘CCP09’, ‘CCP1001’,
‘EMBRAPA50’ and ‘EMBRAPA51’ represent popular
cultivars in Brazil.
A. Sivakumar and T. A. Pai
Literature cited and further reading
Agtrans Research (1996) The Cashew Research and Development
Program: Performance and Future Prospects for Industry Development
– Background Report. Agtrans Research, Brisbane, Australia,
pp. 1–26.
Akinpelu, D.A. (2001) Antimicrobial activity of Anacardium
occidentale bark. Filoterapia 72 (3), 286–287.
Anon. (1996) The World of Cashew. Oltremare Spa, Bologna, Italy.
Behrens, R. (1996) Cashew as an Agroforestry Crop: Prospects and
Potentials. Margraf Verlag, Germany.
Bicalho, B. and Rezende, C.M. (2001) Volatile compounds of cashew
apple (Anacardium occidentale L.). Z. Naturforsch 56 (1–2), 35–39.
Boggetti, B. (1997) Development of micropropagation and potential
genetic transformation systems of cashew (Anacardium occidentale
L.). PhD thesis, Wye College, University of London.
Coronel, R. (1983) Promising Fruits of the Philippines. University of
the Philippines, Los Banos.
Das, S., Jha, T.B. and Jha, S. (1996) In vitro propagation of cashew
nut. Plant Cell Reports 15, 615–619.
D’Souza, L. and D’Silva, I. (1992) In vitro propagation of
Anacardium occidentale L. Plant, Cell Tissue and Organ Culture 29,
1–6.
Duke, J. (1989) Handbook of Nuts. CRC Press, Boca Raton, Florida.
Duncan, I. (1992) World Cashew Market: 1992. RIRDC, Canberra,
Australian Capital Territory, pp. 1–100.
Falcone, A.M. and Leva, A.R. (1989) Propagation and organogenesis
in vitro of Anacardium occidentale L. 1st International Symposium
on In Vitro Culture and Horticultural Breeding, 30 May–3 June
1988, Cesana, Italy.
Hegde, M., Kulasekaran, M., Jayasankar, S. and Shamnungavelu,
K.G. (1991) In vitro embryogenesis in cashew (Anacardium
occidentale L.). Indian Cashew Journal 21 (4), 17–25.
Hilton, B. (1998) Our experience with cashew. ECHO Development
Notes Issue 62.
Hilton, B. (1999) Additional comments about cashew. ECHO
Development Notes Issue 63.
Leung, W.-T., Butrum, W.R.R. and Chan, F.H. (1972) Food
Composition Table for Use in East Asia. USA Department of
Health, Education and Welfare, National Institute of Health,
Washington, DC.
14
Anacardiaceae
Leva, A.R. and Falcone, A.M. (1990) Propagation and organogenesis in
vitro of Anacardium occidentale L. Acta Horticulturae 280, 143–145.
Masawe, P.A.L., Cundall, E.P. and Caligari, P.D.S. (1999)
Observations on progenies in a crossing scheme between cashew
clones: establishment characters. Tanzanian Journal of
Agricultural Science 2, 1–6.
Masawe, P.A.L., Cundall, E.P. and Caligari, P.D.S. (1999) Studies on
genotype–environment interaction (G ⫻ E) in half-sib progenies
of cashew (Anacardium occidentale Linn) in Tanzania. Tanzanian
Journal of Agricultural Science 2, 53–62.
Menninger, E.A. (1977) Edible Nuts of the World. Horticultural Books
Inc., Stuart, Florida.
Mitchell, J. and Mori, S. (1987) The cashew and its relatives
(Anacardium: Anacardiaceae). Memoirs of the New York Botanical
Garden 42, 38.
Mneney, E.E. (1998) Development of in vitro techniques for clonal
propagation and genetic fingerprinting of elite disease-free cashew
(Anacardium occidentale L.). PhD thesis, Wye College, University
of London.
NOMISMA [Italian Economic Development Organization] (1994)
The World Cashew Economy, 2nd edn. L’Inchiostroblu, Bologna,
Italy, pp. 1–218.
Philip, V.J. (1984) In vitro organogenesis and plantlet formation in
cashew (Anacardium occidentale L.). Annals of Botany 54, 149–152.
Purseglove, J.W. (1987) Tropical Crops: Dicotyledons. Longman
Scientific and Technical, Harlow, UK.
Rosengarten, F. Jr (1984) The Book of Edible Nuts. Walker and
Company, New York.
Sy, M.O., Martinelli, L. and Scienza, A. (1991) In vitro organogenesis
and regeneration in cashew (Anacardium occidentale L.). Acta
Horticulturae 289, 267–268.
US Department of Agriculture (USDA) (2003) USDA National
Nutritional Database. Available at: http://www.nal.usda.gov.fric/
foodcomp/search (accessed 6 November 2006).
Veeraraghavan, P.G., Celine, V.A. and Balakrishnan, S. (1985) Study
on the fertilizer requirements of cashew (Anacardium occidentale
L.). Cashew Causerie 7 (2), 6–8.
Verheij, E.W.M. and Coronel, R.E. (eds) (1992) Edible Fruits and
Nuts. Plant Resources of South East Asia No. 2. PROSEA
Foundation, Bogor, Indonesia.
Woodroof, J.G. (1979) Tree Nuts: Production, Processing, Products, 2nd
edn. AVI Publishing Company, Westport, Connecticut.
Buchanania lanzan
chironji
Chironji, Buchanania lanzan Spreng. (Anacardiaceae), is
endemic in the dry deciduous tropical forests of India. The
common names are: in Bengali chironji; in English narrowleaved buchanania, cheraunji nut tree, chirauli nut, chirauli
nut tree, chironji nut, cuddapa almond and cuddapah almond;
in Hindi it is known as achaar, baruda, char (fruit), chironji
(cheronjee) and priyala; in Kannada as charpoppu; in
Malayalam as mungapper; in Nepalese as acar and ciraaunji; in
Oriya as charu; in Sanskrit as char, priyalam and rajadana; in
Tamil as morala; in Telugu as morichettu and saara chettu; in
Burma as lambo; and in Thai as mamuang hua maeng wan and
mamuang maeng wan (Porcher, 2005). The fruit of
Buchanania arborescens (Blume) Blume, found from the
Philippines through South and South-east Asia to northern
Australia, is also eaten. In English, it is known as little
gooseberry tree and in Tagalog as balinghasai and the fruit are
0.7–1 cm long with a thin pulp.
World production and yield
Fallen ripe fruit are collected, however sometimes the tree is
cut down. This destruction has made the species vulnerable to
over-exploitation. Yields of chironji are from 1 to 5 kg/tree
with an average weight of 0.27 g (Rai, 1982; Chadhar and
Sharma, 1997). In central India, 1500 t of fruit are collected
from the wild annually.
Uses and nutritional composition
The fruit are considered as one of India’s most delicious wild
fruit. The seeds are also eaten and are regarded as a substitute
for almonds. The fruit is used in sweets and bakery/
confectionery products. The seed contains about 59% fat,
12% starch and up to 22% protein (Table A.4). In many parts
of India, Buchanania oil is used as a substitute to olive oil and
for almond oil in medicinal preparations and confectioneries.
From the seeds a traditional dish ‘Chironji Ki Burfi’ is
prepared. It is considered very beneficial for newly wed
couples. The bark yields a tannin up to 13% of dry weight.
The wood is only used for firewood. The leaves are used as
fodder for sheep, goat and cattle. It is a host of the Kusumi lac
insect. Chironji is also regarded as a valuable herb.
The tree also has numerous medicinal uses. In the different
systems of medicine in India, the chironji roots, leaves, fruit,
seeds and gum are used. An extract of the root is used as an
expectorant, for biliousness and for blood diseases. The leaf
juice is used for digestive complaints, as an expectorant,
aphrodisiac and purgative. The seed oil is used to treat skin
diseases and to remove face blemishes. The oil is also applied
externally on glandular swellings of the neck alone and in
combination with other herbal oils. Buchanania gum is used
internally in treatment of intercostal pain and diarrhoea. The
oil is useful for coating tablets for its delayed release.
Botany
TAXONOMY AND NOMENCLATURE The synonym is Buchanania
angustifolia Roxb. The genus has about 20 species that are
distributed in tropical Asia, Australia and the Pacific Islands.
Table A.4. Proximate kernel composition per 100 g of Buchanania lanzan.
Proximate
Water
Energy (kcal)
Protein
Lipid (fat)
Carbohydrate
Fibre
Minerals
Calcium
Iron
Phosphorus
%
3.0
656
19.0–21.6
59.1
12.1
3.8
mg
279
8.5
528
Vitamins
mg
Ascorbic acid
Thiamine
Riboflavin
Niacin
5.0
0.69
0.53
1.5
Mangifera
This evergreen, moderate sized tree (15 m) has
a straight, cylindrical trunk and tomentose branches. The bark
is rough and dark grey or black, fissured with prominent
squares, 1.25–1.75 cm thick and reddish inside. The leaves are
broadly oblong with a rounded base, 8–20 cm by 4–12.5 cm.
The small, greenish-white flowers (0.6 cm in diameter) are
axillary and on 5–15 cm long terminal panicles. The calyx is
three to five lobed, 1 mm long and the four or five petals are
3 mm long. The ten stamens are inserted at the base of the
fleshy disc. The ovary has five to six free carpels, inside the
disc, although only one carpel is fertile. The black fruit is a
drupe 1–1.5 cm in diameter.
DESCRIPTION
ECOLOGY AND CLIMATIC REQUIREMENTS It is a common tree
in dry open deciduous forests with a monsoonal climate to
500 m. The tree grows most commonly on yellow sandy-loam
soils.
Flowering occurs from January to
March and the harvest season is from April to June.
REPRODUCTIVE BIOLOGY
Horticulture
The seeds are tolerant to desiccation and chilling. The seeds
show 95–100% survival up to 90 days at all storage
temperatures with gradual loss in germination on or after 280
days of storage. The seeds have a hard seedcoat that lowers
germination. Root cutting can be successfully used for
propagation. A tissue culture technique for the rapid clonal
multiplication has been developed.
No known cultivars have been selected. Wide variation
occurs in fruit size, fruit per panicle, panicles per tree, total
soluble solids and fruit yield per tree (Munde et al., 2004).
Pankaj Oudhia and Robert E. Paull
15
priyal: fruit and seeds of Buchanania lanzan Spreng. (Family:
Anacardiaceae). Bulletin of the Botanical Survey of India 22,
68–76.
Munde, V.M., Shinde, G.S., Sajindranath, A.K., Prabhu, T. and
Machewad, P.M. (2004) Variability studies in charoli (Buchanania
lanzan Spreng.). Journal of Soils and Crops 14, 205–206.
Oudhia, P. (2003) Research Articles – Indian Herbal Research and
Methods. Available at: http://botanical.com/site/column_
poudhia/poudhia_index.htm (accessed 6 November 2006).
Porcher, M.H. (2005) Multilingual Multiscript Plant Name
Database. The University of Melbourne, Australia. Available at:
http://www.plantnames.unimelb.edu.au/Sorting/Frontpage.html
(accessed 31 August 2005).
Rai, Y.C. (1982) Buchanania lanzan, Spreng. – studies on methods of
propagation and estimation of fruit yield. Indian Forester 108,
501–511.
Saha, D.P. and Maurya, K.R. (1998) Pomological description and
medicinal value of chiraunjee (Buchanania latifolia Spreng.).
Journal of Applied Biology 8, 67–69.
Samant, S.K. and Rege, D.V. (1990) Some enzymes and enzyme
inhibitors from charoli and cashew nut. Journal of Food Science
and Technology 27, 231–232.
Sebastian, M.K. and Bhandari, M.M. (1990) Edible wild plants of
the forest areas of Rajasthan (India). Journal of Economic and
Taxonomic Botany 14, 689–694.
Shiva, A. (2001) Harvesting of edible plant species: Buchanania
lanzan, Emblica officinalis, Mangifera indica, Moringa oleifera and
Syzygium cuminii. MFP News 11, 10–12.
Singh, J., Banerjee, S.K. and Francis, A. (2002) Vegetative
propagation of Buchanania lanzan Spreng. root cuttings. Indian
Forester 128, 700–704.
Singh, R.P., Rana, B.S. and Garkoti, S.C. (1993) Biomass and
production patterns of three dominant tree species along a girth
series in a natural tropical forest at Chakia, Varanasi (India).
Indian Forester 119, 472–480.
Literature cited and further reading
Anon. (1995) Chironji (Buchanania lanzan). Indian Council of
Forestry Research and Education (ICFRE), Dehra Dun, pp. 1–16.
Arya, R., Babu, V., Ilyas, M. and Nasim, K.T. (1992) Myricetin 3⬘rhamnoside-3-galactoside from Buchanania lanzan (Anacardiaceae).
Phytochemistry 31, 2569–2570.
Banerjee, A. and Jain, M. (1988) Investigations of Buchanania lanzan
seed oil. Fitoterapia 5, 406.
Bhatnagar, P. and Jain, S. (2002) Collection, processing and
marketing of Buchanania lanzan (Chironji) in Chhindwara
District. Vaniki Sandesh 26, 11–15.
Chadhar, S.K. and Sharma, M.C. (1997) Trends in fruit production
in Buchanania lanzan trees. Vaniki Sandesh 21, 1–3.
Chandra, D. and Ghosh, R.B. (1993) Family Anacardiaceae in West
Bengal. Journal of Economic and Taxonomic Botany 17, 587–591.
Duke, J.A. (1989) Handbook of Nuts. CRC Press, Boca Raton,
Florida.
Hemavathy, J. and Prabhakar, J.V. (1988) Lipid composition of
chironji (Buchanania lanzan) kernel. Journal of Food Composition
and Analysis 1, 366–370.
Kumar, S., Pachori, P.K. and Shrivastava, D.C. (2002) Storability of
chironji (Bachanania lanzan L.) kernels. Jawaharlal Nehru Krishi
Vishwa Vidyalaya Research Journal 36, 61–65.
Mitra, R. and Mehrotra, S. (1980) Pharmacognostical studies on
Mangifera indica
mango
Mango, Mangifera indica L. (Anacardiaceae), has been
cultivated for millennia and is one of the premier fruit grown
and consumed throughout the world. There are numerous
common names including: mango in English and Spanish;
mampelam, ampelam and mangga in Malay; mangga, paho
and mango in Philippino; thayet and tharyetthi in
Myanmarese; svaay in Cambodian; mwàngx in Laotian;
mamuang in Thai; xoàl in Vietnamese; pauh, ampelam and
mangga in Indonesian; mangot, mangue and manguier in
French; and mango and mangueira in Portuguese.
Native to Asia, the centre of origin is thought to be in the
region of north-eastern India and Myanmar, however, it was
distributed throughout South-east Asia and the Malay
Archipelago at least 1500 years ago and to parts of Africa about
1000 years ago (Smith et al., 1992). Wild mango is reported
throughout these areas and China and Sri Lanka. Thus mango
was distributed in ancient times throughout all of tropical and
subtropical Asia, and northern and eastern Africa. Mango was
later distributed to the New World during the exploration and
colonization of the New World from the 15th to the 18th
centuries by the Portuguese, Spanish, British and French.
Feral mango populations are now found in parts of West
16
Anacardiaceae
Africa, Mexico and Central and South America (Smith et al.,
1992). Today mango is grown on various commercial scales
throughout the warm to cool subtropical and tropical areas of
the world (Martin et al., 1987).
World production and yield
There is an estimated 3.7 million ha of mangos worldwide
(FAOSTAT, 2005a, b). Mango production in 2004 was
estimated at 26.6 million t, ranked seventh in worldwide fruit
production behind banana, grape, oranges, apple, coconut and
plantain. The top ten mango-producing countries based on
area of production include India, China, Thailand, Mexico,
Indonesia, the Philippines, Nigeria, Pakistan, Guinea and
Brazil. The top five largest mango-exporting countries are
Mexico, India, Brazil, Peru and the Philippines with exports
worldwide valued at US$560.4 million. The top five mangoimporting countries are the USA, the Netherlands, the United
Arab Emirates, Saudi Arabia and Bangladesh with imports
valued at US$703.9 million.
Fruit are available year-round depending upon production
location and cultivar. Production per ha varies greatly with
average yields of 2–6 t/ha being common in some regions and
with highest yields reported to be 10–30 t/ha (Verheij and
Coronel, 1992). Average yields for productive orchards range
from 22 to 25 t/ha (Nakasone and Paull, 1998).
Uses and nutritional composition
Mango is commonly eaten fresh and depending upon the
cultivar may be consumed at an immature (unripe, green peel)
stage or when fully ripe. In addition, the pulp may be cooked,
dried, preserved, frozen or powdered. Mango pulp may be
incorporated into beverages, desserts, ice cream, sorbets,
preserves, jellies, fruit salads, chutneys, pickles, canned in
syrup, pureed and dried.
Mangos are a rich source of vitamins A and C (Table A.5)
and have recently been found to be high in anti-cancer
antioxidants and phenols. Historically there have been many
reported medicinal uses of the sap (latex), flowers, seeds and
leaves for use as astringents, treating diarrhoea, haemorrhages,
fever, hypertension and haemorrhoids.
Botany
The genus Mangifera is one
of 73 genera belonging to the Anacardiaceae (Bompard and
Schnell, 1997). However, the taxonomy of Mangifera remains in
flux with over 60 species found in South and South-east Asia.
The genus Mangifera has been divided into two subgenera,
Limus sp., which are quiet distinct from the common mango
and may be ancestral to the mango, and the subgenera
Mangifera. The subgenus, Mangifera is further divided into five
sections based on differing floral, seed and anatomical attributes
with M. indica belonging to Section Mangifera Ding Hou. This
section is further divided into three groups based on floral
structures and organ number variation with M. indica belonging
to a group with five species characterized by tetra- and
pentamerous flowers. Two main centres of domestication of
mango are recognized, India with monoembryonic cultivars and
TAXONOMY AND NOMENCLATURE
Table A.5. Composition of mango per 100 g edible portion (Source: Morton,
1987; Verheij and Coronel, 1992; USDA, 2005).
Proximate
Energy (kJ)
Protein
Fat
Carbohydrate
Fibre
Ash
Moisture
Minerals
Calcium
Potassium
Phosphorus
Iron
Magnesium
Vitamins
Thiamine
Riboflavin
Niacin
Vitamin C
Vitamin E
Vitamin A (IU)
g
225–350
0.30–0.8
0.10–0.27
13.2–20.0
0.60–1.80
0.50
78–85
mg
9.0–25.0
156.0
10.0–15.0
0.10–0.20
9.00
mg
0.058
0.057
0.584
14–62
1.12
765
Asia, which includes Indonesia, the Philippines and parts of
Vietnam, Thailand and Myanmar, with polyembryonic seeds.
The major mango species in cultivation is M. indica,
however Mangifera foetida (horse mango), Mangifera odorata
(kuini), Mangifera caesia (binjai), Mangifera langenifera (lanjur
or langoot) and Mangifera zylanica are cultivated or gathered
commercially on a small scale in various parts of South-east
Asia (Yaacob and Subhadrabandhu, 1995; Bompard and
Schnell, 1997). Two broad types of mango are recognized,
Indian and Indo-Chinese (Crane et al., 1997). Indian types
have monoembryonic seeds, are usually highly coloured, and
tend to be susceptible to anthracnose (Colletotrichum
gloeosporioides). Indo-Chinese types have polyembryonic seeds,
usually are green to light green to yellow at maturity, and tend
to be more resistant to anthracnose.
There are a number of mango relatives with edible fruit
including cashew (Anacardium occidentale), anacardier géant
(Anacardium giganteum), maprang (Bouea gandaria), Kaffir
plum (Harpephyllum caffrum), bembé (Lannea acida),
ambarella (Spondias cytherea), yellow mombin (Spondias
mombin), purple mombin (Spondias purpurea) and imbu
(Spondias tuberose) (Martin et al., 1987).
Mango trees may grow to 45 m in height with a
broad canopy up to 38 m in width. In deep soils the taproot
may extend downwards to 6 m and the lateral fibrous root
system may extend well beyond the canopy drip-line.
Immature leaves are reddish brown and soft. Mature leaves are
green, simple, spirally arranged, lanceolate to oblong, 8–40 cm
long by 2–10 cm wide, and coriaceous (Fig. A.3). Flowers are
borne on green, yellow or pinkish-coloured large terminal or
axillary panicles up to 60 cm in length. Each panicle may
possess 300–6000 individual flowers (Iyer and Degani, 1997).
There are two flower types on each panicle, hermaphroditic
DESCRIPTION
Mangifera
Fig. A.3. Leaf, flower and fruit of Mangifera indica (Source:
Nakasone and Paull, 1998).
(perfect) and staminate (male). Flowers are small, greenish
yellow to reddish pink, calyx five-lobed and five petals.
Hermaphroditic flowers have one simple pistil with one
functional stamen and four staminodes; staminate flowers have
no ovary, usually one functional stamen and four staminodes.
Fruit an oblong, reniform to oblate, fleshy drupe, 6–30 cm
long, 100–2300 g, with or without a pronounced beak; fruit
are solitary or in clusters (Martin et al., 1987; Yaacob and
Subhadrabandhu, 1995). Fruit possess one seed enclosed in a
stony endocarp; seeds may have one (monoembryonic types)
or multiple (polyembryonic) embryos. The waxy, smooth peel
is somewhat leathery and ranges in colour at maturity from
green to light green with ground colour or blushes of one or
more shades of yellow, orange, red or purple alone or more
commonly in combination. The pulp of immature fruit is hard
and white to light yellow, at maturity the pulp may be more or
less fibrous, soft, juicy and light yellow to orange. Fruit may
emit a mild to strong aroma and the flesh from ripe fruit is
sweet with a mild to strong flavour; fibre content varies. Fruit
possess one monoembryonic or polyembryonic seed.
ECOLOGY AND CLIMATIC REQUIREMENTS There are two main
ecological types of mango – those that evolved in the warm
subtropical climate of India and those that evolved in the hot,
humid, lowland tropical areas of South-east Asia. Mango trees
may be grown in a wide range of climates, however, highest
17
production of quality fruit occurs in those areas with a
distinctive non-freezing cool period and/or an extended dry
period (at least 3 months) prior to flowering, hot temperatures
(30–33°C) during fruit development, and access to water
(ideally from irrigation and not rainfall) from flowering to
harvest (Davenport and Núñez-Elisea, 1997; Whiley and
Schaffer, 1997).
Mango trees are adapted to a wide range of soil types
including various sandy, loamy, clayey, and rocky soils (Yaacob
and Subhadrabandhu, 1995). Mango trees produce best under
full sunlight exposure; constant shading reduces or delays
flower-bud formation. There does not appear to be any effect
of photoperiod on mango flowering. Peel colour development
is dependent upon light exposure. Mango trees are only
moderately freeze tolerant with young trees damaged or killed
at temperatures below ⫺1.1°C to ⫺1.7°C and mature trees
damaged or killed at or below ⫺3.9°C to ⫺6.0°C (Snow, 1963;
Campbell et al., 1977). The minimum temperature for
vegetative growth is about 15°C with optimum growing
temperatures ranging from 24°C to about 30°C (Schaffer et
al., 1994). In contrast, the base temperature for panicle growth
appears to be around 12.5°C. Soil temperatures at 12°C have
been found to limit plant growth. Low temperatures (4.4°C–
15°C) during flowering may result in abnormal flowers and
reduced pollen viability and fruit set (Schaffer et al., 1994;
Whiley and Schaffer, 1997). In contrast, high temperatures
(33–36°C) during pollen meiosis result in reduced pollen
viability.
Mangos are tolerant to periodic and repeated flooding and
this appears to be dependent upon development of
hypertrophied lenticels along the trunk. Mango trees are very
drought tolerant due to their laticifer cells (resin canals),
osmotic adjustment of leaves, their deep and extensive root
system, and coriaceous leaves. However, drought during fruit
set and development may result in reduced yields and fruit
size. Mango trees are intolerant of saline soil and water which
causes necrosis of leaf margins and tips, defoliation, stem
dieback and tree death. However, there is some variability in
the saline tolerance of various cultivars.
Windy conditions during flowering and fruit development
may cause fruit drop and reduce fruit quality through
scarring. Mango trees are moderately tolerant of constant
winds, however, substantial yield increases have been reported
after orchard trees were protected from prevailing winds by
man-made or natural windbreaks. This was attributed to
improved fruit set, less physical damage to leaves and fruit and
reduced disease incidence. Mature mango trees are moderately
wind tolerant although large trees are susceptible to toppling,
major limb and trunk damage, and uprooting due to typhoons
or hurricanes (Crane et al., 1993).
REPRODUCTIVE BIOLOGY The mango inflorescence bears
functionally male and bisexual flowers (termed polgamy) with
the ratio of functionally male to bisexual flowers varying by
genetic predisposition of the cultivar, temperatures during
floral morphogenesis (low temperatures reduce the proportion
of bisexual flowers) and endogenous growth regulators
(Davenport and Núñez-Elisea, 1997). Most mango cultivars
are self-fertile but benefit from cross-pollination (Martin et
al., 1987). Flowers open in the morning and anthesis is
18
Anacardiaceae
generally completed by noon; receptivity of the flowers usually
lasts up to about 72 h (Iyer and Degani, 1997). Fruit set varies
with genetic predisposition of the cultivar for selfincompatibility, weather conditions during and immediately
after flowering, and opportunity for cross-pollination. In
general, fruit set from self-pollination ranges from 0 to 1.7%
whereas from cross-pollination from 6.4 to 23.4% (Nakasone
and Paull, 1998).
Mango flowers may be pollinated by flies, bees, thrips and
other insects, with flies probably the most important. Some
fruit set may occur due to wind pollination. There are two
types of mangos: polyembryonic, forming seed with several
adventitious embryos arising from nucellar tissue; and
monoembryonic, forming seed with a single zygotic embryo.
In order to flower profusely, mango trees require mature,
resting, terminals (stems) and a quiescent period induced by
either cool non-freezing temperatures (8–15°C night/< 20°C
day) and/or dry conditions (Davenport and Núñez-Elisea,
1997). Furthermore, the conditions during flowering
(induction) after shoot initiation also influence the balance
between putative floral (possibly cytokinins) and anti-floral
(possibly gibberellins) endogenously produced compounds.
Exposure of shoots initiated to grow during cool temperatures
result in reproductive growth, in contrast exposure of shoots
initiated to grow during cool temperatures but then
immediately exposed to warm temperatures (25°C night/30°C
day) may produce vegetative growth or mixed (reproductive/
vegetative) growth.
Numerous techniques have been developed for inducing or
enhancing mango flowering, however, not all of them work on
all cultivars, climates and soils. Traditional methods to induce
mango flowering include cincturing the trunk or branches,
root pruning, smudging and applications of saline materials.
More recently, foliar applications of ethylene-releasing
compounds, potassium nitrate and paclobutrazol have proved
to be efficacious on more cultivars and under a wider range of
environmental conditions. Paclobutrazol may also be applied
as a soil drench.
FRUIT DEVELOPMENT Time from flowering to fruit maturity
takes 3–6 months depending upon cultivar and temperatures
(Martin et al., 1987). Mango fruit growth follows a typical
sigmoidal pattern with the seed developing first followed by a
final rapid increase in mesocarp (pulp) as fruit near maturity.
Horticulture
PROPAGATION Mango may be propagated by seed, grafting,
budding, marcottage, rooted cuttings and tissue culture.
Polyembryonic cultivars may be propagated by seed but
grafting or budding is now more common. Monoembryonic
cultivars must be propagated vegetatively; most commonly by
budding or grafting. Flowering and fruiting take 6–10 years
from seed and 3–5 years from grafting depending upon
genetic predisposition of the cultivar, environmental
conditions and culture.
ROOTSTOCKS In general, polyembryonic mangos are used as
rootstocks because of their uniformity. Various rootstocks have
been selected for their tolerance to soil conditions such as high
pH and calcareous soils (‘No. 13–1’, ‘Turpentine’) or reduced
tree vigour (‘Olour’, ‘Vellai Colamban’, ‘Saber’) (Iyer and
Degani, 1997; Morton, 1987). Other polyembryonic
rootstocks include ‘Madu’ in Indonesia, ‘Kaew’ in Thailand
and ‘Kensington Pride’ in Australia (Verheij and Coronel,
1992).
The tree spacing selected depends
upon climate, cultivar, soil type and depth, horticultural
expertise and economics. Plant spacings vary widely and may
range from 4 to 12 m in-row and 6 to 15 m between rows.
Pruning recommendations vary with production region,
cultivar, climate, soil type, inherent vigour of the cultivar,
available technology and labour, and tradition. In general,
trees planted in more closely spaced orchards require initial
tree training to establish two to four main scaffold limbs and
the basic tree shape. Maintenance pruning to remove dead and
damaged limbs, water sprouts, limit tree size and maintain
production is generally recommended for immediately after
harvest and may consist of hand pruning (thinning-out entire
shoots and/or branches), tipping shoots back, and/or
mechanical hedging and topping. Intensive hand pruning to
remove non-flowering shoots after fruit set, followed by
removal of fruiting stems after harvest, may limit tree size and
reduce biennial bearing in some cultivars. Paclobutrazol has
been used to reduce flushing and tree growth successfully in
various mango regions of the world; however, the success of
this practice varies by cultivar and environmental conditions
(Crane et al., 1997).
PRUNING AND TRAINING
NUTRITION AND FERTILIZATION Fertilizer practices are based
on research, observation and experience and should include
the application of all essential elements. A wide range of
nutrient rates and formulations is recommended depending
upon the region, cultivar, soil type and availability of fertilizer
materials. Furthermore, standard leaf nutrient levels vary with
production region and deficiency symptoms and leaf sampling
procedures have been described previously (Samra and Arora,
1997). General recommendations for young trees are about
100–200 g of an NPK (nitrogen, phosphorus, potassium)
material applied two to four times per year with rates
increasing and frequency decreasing as trees mature (1.0–2.0
kg/tree/application) (Crane et al., 1997; Samra and Arora,
1997). Similarly, secondary (magnesium, sulphur) and minor
element (zinc, manganese, iron, boron) fertilization varies
widely based on local environmental conditions and inherent
genetics of the cultivar grown.
Irrigation practices vary widely depending upon soil type,
depth to water table, availability of water, water quality,
available technology and cost, plant spacing and climate
(Crane et al., 1997). In general, newly planted trees should be
planted during the rainy season or irrigated until established
(Crane et al., 1997). The necessity of irrigation for mature
trees varies with climate, soil type, cultivar and tree phenology.
In some areas, irrigating from flowering to near harvest is
beneficial.
DISEASES, PESTS AND WEEDS
There are numerous disease
and insect pests of mango. Important diseases of mango
include anthracnose (C. gloeosporioides), powdery mildew
Mangifera
(Oidium mangiferae) and bacterial black spot (Xanthomonas
campestris pv. mangiferaeindicae) (Dodd et al., 1997; Ploetz and
Prakash, 1997; Ploetz, 2003). Other major diseases in some
production regions include mango malformation (caused by a
number of closely related Fusarium spp.) which affects panicle
formation and fruit set, pink disease (Erythricium salmonicolor)
which kills trees by colonizing the vascular and cambial
tissues, and black spot (Alternaria alternata) which is primarily
a postharvest fruit problem.
Every mango production area has a unique pest complex
and numerous insect pests attack leaves, flowers, fruit, shoots
and the trunk of mango and their importance varies with
production region. Various mango hopper species (e.g.
Idioscopus clypealis, Idioscopus niveosparsus) are of major
importance in South-east Asia and attack the mango
inflorescence and flowers. Numerous Lepidoptera species (e.g.
Chlumetia transversa, Eupithecia sp., various Noctuidae) and
several flower thrips (Frankliniella bispinosa, Frankliniella
kelliae) attack mango flowers reducing fruit set and crop
yields. Some fruit flies of the Tephritidae are major mango
fruit pests including species from the genera Anastrepha,
Bactocera, Dacus and Ceratitis. Other important insect pests
include fruit piercing moths (Eudocima fullonia, Eudocima
maternal, Eudocima salaminia), various midges (e.g. Erosomya
mangiferae, Erosomya indica), the mango seed weevil
(Sternochetus mangiferae), mango seed borer (Deanolis
sublimbalis), various mites (e.g. Oligonychus sp., Aceria
mangiferae), scales (e.g. Aulacaspis tubercularis, Philephedra
tuberculosa, Ceroplastes pseudoceriferus) and mealy bugs
(Rastrococcus invadens, Rastrococcus spinosus) (Peña and
Mohyuddin, 1997; Waite, 2002).
POSTHARVEST HANDLING AND STORAGE The recommended
stage of fruit maturity for harvest depends upon the end use of
the fruit (e.g. green-crunchy or soft-ripe), the distance and
time to markets, available storage conditions and any required
quarantine treatments. For example, ‘Khieo Sawoey’ is
harvested at the immature, green stage and consumed as a
green-crunchy fruit. In contrast the recommended stage of
maturity for harvesting ‘Nom Doc Mai’ for eating ripe is
about 100–102 days from bloom in Thailand and the
Philippines (Yaacob and Subhadrabandhu, 1995). Typically
‘Tommy Atkins’ and ‘Keitt’ mangos may be harvested at early
or late maturity depending upon the distance to markets
within the USA.
Mangos are picked by hand or with the help of various
picking poles with bags or hooks. In general, fruit destined for
export markets are pre-cooled, washed and sometimes treated
with fungicide to reduce postharvest decay, waxed to reduce
water loss, quarantine treated to kill fruit fly larvae and eggs,
and then graded by appearance and size, packed, stored and
shipped (Johnson et al., 1997). Fruit sold in local, regional and
national markets may or may not be extensively graded and
treated prior to marketing. Mature mangos may also be gassed
with ethylene prior to marketing to synchronize fruit ripening
and colour development.
The phytosanitary requirements for exporting mangos
varies by country and ranges from no restrictions to entry into
a country to certification that fruit were grown in a fruit-fly
free area and/or postharvest disinfested in some specific
19
manner (Johnson et al., 1997). Quarantine treatments for fruit
fly disinfestations include various hot water treatments, vapour
heat, hot air and irradiation. Mango storage temperatures vary
by cultivar and stage of fruit maturity. Safe storage
temperatures for a 2–3 week period may range from 8 to 21°C
and 85 to 90% relative humidity (McGregor, 1987; Morton,
1987; Johnson et al., 1997).
MAIN CULTIVARS AND BREEDING Mango is genetically
heterogeneous resulting in seedling progeny with a very wide
range of growth habits and fruit characteristics (Smith et al.,
1992). There are hundreds (perhaps more than a thousand) of
mango cultivars and formal and informal selection from
seedling populations has occurred for millennia throughout
India and South-east Asia and more recently throughout the
tropics and subtropics of the western hemisphere. There are
over 25 field germplasm banks throughout the tropics and
subtropics with responsibility for the conservation of M.
indica and its relatives. Many of them house named mango
cultivars, land races, selections and relatives of mango.
Currently there are formal breeding or selection programmes
in Asia (e.g. Malaysia, Indonesia, India, Pakistan, Taiwan),
Central and South America (e.g. Brazil) and North America
(e.g. Mexico, USA/Florida). Mango breeding or selection
criteria include regular bearing, low to moderate vigour,
disease and insect resistance, freedom from physiological
disorders, increased fruitfulness and storability to name a few.
Selection criteria for rootstocks included polyembrony,
dwarfing, tolerance to adverse edaphic conditions (e.g. high
pH, flooding) and scion compatibility (Iyer and Degani, 1997).
In addition, Mangifera relatives possess various characteristics
such as disease resistance, adaptation to stressful edaphic
conditions (e.g. flooding), dwarfing, and free-stone fruit that
in the future may be utilized successfully to improve cultivated
mango (Litz and Lavi, 1997).
Major cultivars vary in every production area and most
were selected from chance seedlings. Literally hundreds of
mango cultivars are grown in India with ‘Alphonso’, ‘Alampur
Baneshan’, ‘Paheri’ and ‘Neelum’ just to name a few. In
Thailand cultivars are divided into those eaten unripe (called
starch or crispy mango) and those eaten ripe: crispy cultivars
include ‘Khieo Sawoey’ and ‘Rad’ and ripe-eating cultivars
include ‘Okrong’, ‘Nam Dok Mai’, ‘Nang Klangwan’ and
‘Thong Dam’ (Yaacob and Subhadrabandhu, 1995). In the
Philippines, ‘Carabao’ and ‘Pico’ are predominant cultivars.
Major cultivars in the western hemisphere include ‘Manila’
and ‘Altaulfo’ (both from Mexico), ‘Bourbon’, ‘Extrema’ and
‘Coraçao de Boi’ (all from Brazil), ‘Kensington Pride’
(Australia), ‘Tommy Atkins’, ‘Keitt’, ‘Kent’ and ‘Haden’ (all
originally from Florida but now the major export mango
cultivars used in the export trade from Mexico and Latin
America) (Knight, 1997).
Jonathan H. Crane
Literature cited and further reading
Bompard, J.M. and Schnell, R.J. (1997) Taxonomy and systematics.
In: Litz, R.E. (ed.) The Mango: Botany, Production, and Uses.
CAB International, Wallingford, Oxon, UK, pp. 21–47.
Campbell, C.W., Knight, R.J. Jr and Zareski, N.L. (1977) Freeze
damage to tropical fruits in southern Florida. Proceedings of the
Florida State Horticultural Society 90, 254–257.
20
Anacardiaceae
Crane, J.H., Campbell, R.J. and Balerdi, C.F. (1993) Effect of
hurricane Andrew on tropical fruit trees. Proceedings of the Florida
State Horticultural Society 106, 139–144.
Crane, J.H., Bally, I.S.E., Mosqueda-Vazquez, R.V. and Tomer, E.
(1997) Crop production. In: Litz, R.E. (ed.) The Mango: Botany,
Production, and Uses. CAB International, Wallingford, Oxon, UK,
pp. 203–256.
Davenport, T.L. and Núñez-Elisea, R. (1997) Reproductive
physiology. In: Litz, R.E. (ed.) The Mango: Botany, Production,
and Uses. CAB International, Wallingford, Oxon, UK,
pp. 69–146.
Dodd, J.C., Prusky, D. and Jeffries, P. (1997) Fruit diseases. In: Litz,
R.E. (ed.) The Mango: Botany, Production, and Uses. CAB
International, Wallingford, Oxon, UK, pp. 257–280.
FAOSTAT (2005a) FAO database, mangos, export, import, and value
2003. Available at: http://faostat.fao.org/faostat/form?collection
=Trade.CropsLivestockProducts&Domain=Trade&servlet=1&ha
sbulk=&version=ext&language=EN (accessed 17 July 2005).
FAOSTAT (2005b) FAO database, mangoes, area harvested and
production 2004. Available at: http://faostat.fao.org/faostat/
form?collection=Production.Crops.Primary&Domain=Production
&servlet=1&hasbulk=&version=ext&language=EN (accessed 17
July 2005).
Iyer, C.P.A. and Degani, C. (1997) Classical breeding and genetics.
In: Litz, R.E. (ed.) The Mango: Botany, Production, and Uses.
CAB International, Wallingford, Oxon, UK, pp. 49–68.
Johnson, G.I., Sharp, J.L., Milne, D.L. and Oosthyse, S.A. (1997)
Postharvest technology and quarantine treatments. In: Litz, R.E.
(ed.) The Mango: Botany, Production, and Uses. CAB
International, Wallingford, Oxon, UK, pp. 447–507.
Knight, R.J. Jr (1997) Important mango cultivars and their
descriptors. In: Litz, R.E. (ed.) The Mango: Botany, Production,
and Uses. CAB International, Wallingford, Oxon, UK,
pp. 545–565.
Litz, R.E. and Lavi, U. (1997) Biotechnology. In: Litz, R.E. (ed.) The
Mango: Botany, Production, and Uses. CAB International,
Wallingford, Oxon, UK, pp. 401–423.
McGregor, B.M. (1987) Tropical Products Transport Handbook.
Agriculture Handbook No. 668. US Department of Agriculture
(USDA) Agriculture Marketing Service (AMS), Washington,
DC, pp. 1–101.
Martin, F.W., Campbell, C.W. and Ruberté, R.M. (1987) Perennial
Edible Fruits of the Tropics – an Inventory. Agriculture Handbook
No. 642. US Department of Agriculture (USDA) Agriculture
Research Service (ARS), US Government Printing Office,
Washington, DC, pp. 1–14.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resource
Systems, Winterville, North Carolina, pp. 221–237.
Nakasone, H.Y. and Paull, R.E. (1998) Tropical Fruits. CAB
International, Wallingford, Oxon, UK, pp. 208–238.
Peña, J.E. and Mohyuddin, A.I. (1997) Insect pests. In: Litz, R.E.
(ed.) The Mango: Botany, Production, and Uses. CAB
International, Wallingford, Oxon, UK, pp. 327–362.
Ploetz, R.C. (2003) Diseases of mango. In: Ploetz, R.C. (ed.) Diseases
of Tropical Fruit Crops. CAB International, Wallingford, Oxon,
UK, pp. 327–363.
Ploetz, R.C. and Prakash, O. (1997) Foliar, floral, and soil borne
diseases. In: Litz, R.E. (ed.) The Mango: Botany, Production, and
Uses. CAB International, Wallingford, Oxon, UK, pp. 281–325.
Samra, J.S. and Arora, Y.K. (1997) Mineral nutrition. In: Litz, R.E.
(ed.) The Mango: Botany, Production, and Uses. CAB International, Wallingford, Oxon, UK, pp. 175–201.
Schaffer, B., Whiley, A.W. and Crane, J.H. (1994) Mango. In:
Schaffer, B. and Andersen, P.C. (eds) Handbook of Environmental
Physiology of Fruit Crops, Vol. II: Subtropical and Tropical Crops.
CRC Press, Boca Raton, Florida, pp. 165–197.
Sergio, A.R.R. and Russell, R.M. (1999) β-carotene and other
carotenoids as antoxidants. Journal of the American College of
Nutrition 18, 426–433.
Smith, N.J.H., Williams, J.T., Plucknett, D.L. and Talbot, J.P. (1992)
Tropical Forests and Their Crops. Cornell University Press, New
York, pp. 76–95.
Snow, R.E. (1963) Cold tolerance observations during the 1962
freeze. Proceedings of the Florida State Horticultural Society 76,
374–377.
US Department of Agriculture (USDA) (2005) Mango, NDB No.
09176. USDA Agriculture Research Service (ARS), Nutrient
Data Laboratory, USDA National Nutrient Database for
Standard Reference, Release #17. Available at: http://www.nal.
usda.gov/fnic/foodcomp/cgi-bin/list_nut_ edit.pl (accessed 17
July 2005).
Verheij, E.W.M. and Coronel, R.E. (eds) (1992) Edible Fruits and
Nuts. Plant Resources of South East Asia No. 2. PROSEA
Foundation, Bogor, Indonesia, pp. 211–216.
Waite, G.K. (2002) Pests and pollinators of mango. In: Peña, J.E.,
Sharp, J.L. and Wysoki, M. (eds) Tropical Fruit Pests and Pollinators.
CAB International, Wallingford, Oxon, UK, pp. 103–130.
Whiley, A.W. and Schaffer, B. (1997) Stress physiology. In: Litz, R.E.
(ed.) The Mango: Botany, Production, and Uses. CAB
International, Wallingford, Oxon, UK, pp. 147–173.
Yaacob, O. and Subhadrabandhu, S. (1995) The Production of
Economic Fruits in South-East Asia. Oxford University Press, New
York, pp. 145–155.
Pistacia vera
pistachio
The pistachio, Pistacia vera L. (Anacardiaceae), is native to
western Asia and Asia Minor, where it is still found growing
wild in numerous hot, dry locations in Lebanon, Palestine,
Syria, Iran, Iraq, India, Southern Europe and the desert
countries of Asia and Africa. Several species are referred to as
pistachios, but the name is generally reserved for the edible
nut of commerce. The common names are variants on the
Persian name pistaa: in Arabic it is fustuq, in French pistache,
in German pistazie, Hindi pistaa, Italian pistacchio, Spanish
pistacio and Portuguese pistácio or pistácia.
Pistachio was introduced to Europe at the beginning of the
Christian era. The first pistachio introductions to the USA
were by the US Department of Agriculture (USDA) plant
exploration service in 1890. California introductions were first
planted at the Plant Introduction Station in Chico in the
Northern Sacramento Valley in 1904. Commercial pistachio
production differs markedly between the more intensive new
world production practices, primarily in California and the
traditional, less intensive, non-irrigated production practices
followed in the old world, Iran, Turkey and Syria.
A related species Pistacia lentiscus L. yields a mastic that is
used in incense, to fill teeth, as chewing gum and as a varnish.
The fruit are occasionally used to make sweets or liqueurs.
The Pistacia terebinthus yields Cyprus turpentine collected by
Pistacia
21
making incisions in the trunk. The small Pistacia palaestina
fruit are sold in Arab markets as a condiment and have a
resinous taste.
Table A.7. Proximate composition for raw pistachio nut per 100 g with the
nut being 53% of the nut in shell (Source: USDA, 2006).
World production and yield
Water
Energy (kcal)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
Modern world pistachio production is divided between the
new and the old world. Iran is the world’s leading producing
country, followed by California, Turkey, Syria, Greece and
Italy (Table A.6). Greece and Italy both produce less than 1%
of the world’s total production. Plantings have increased
sharply in the three major producing areas, Iran, California
and Turkey. Since 1985 pistachio production has been initiated
in China, Argentina, Australia and South Africa. However,
production in these countries has not reached commercially
competitive levels. Average yield per tree ranges from 2 to 22.5
kg/year. World yields in 2000 varied from 500 kg/ha to 3037
kg/ha with a world average of 1117 kg/ha (Kaska, 2002).
Yields per hectare have increased in all growing areas over the
last 20 years.
Pistachios display three physiological conditions that
decrease commercial yields. The first is alternate bearing, an
annual cycling of high and low crops as a result of premature
fruit bud abscission. The second is blanking, the failure of the
embryo to develop, resulting in an empty nutshell. The third is
non-splitting, the failure of the fully developed nutshell to
split, resulting in a filled but unopened nut. These all appear
to be related to crop load and are therefore probably ultimately
related to carbohydrate competition. Thus far, little is known
about the specific mechanism of each, though the correlation
with crop load is apparent in each case.
Proximate
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
%
3.97
557
20.61
44.44
27.97
10.3
3.02
mg
107
4.15
121
490
1025
1
mg
5.0
0.87
0.16
1.3
553 IU
nut production increases. The nut has 55% of a non-drying oil
that is predominantly 23.3 g/100 g monounsaturated and
13.46 g/100 g polyunsaturated, and only 5.4 g/100 g saturated
(USDA, 2006).
Uses and nutritional composition
Botany
The primary use of pistachio nuts is eating out of hand, fresh in
season, dried, and dried and roasted, with or without salt, or
flavoured with spices and condiments. ‘Red’ pistachios are
roasted and salted and then coloured with a red vegetable dye to
cover blemishes. Nutritionally, pistachios are superior relative to
other tree nuts and peanuts as they are lower in calories and fat,
and higher in potassium, carbohydrates and protein (Table A.7).
Secondary nut uses are in baked goods, nougat candies,
sausages, pâtés and as a flavouring for ice cream, in ground or
powdered form. In the countries where pistachios are
traditionally grown, the uses are more varied than in the
Western countries. The outer husk is used in India for dying
and tanning. The nuts are used for a number of folk remedies
such as for abdominal pains, abscesses, bruises and dysentery.
A limited amount of pistachio oil is produced for eating and
cosmetic purposes. The amount of oil may increase as world
TAXONOMY AND NOMENCLATURE The synonyms of Pistacia
vera L. are P. nigricans Crantz, P. officinarum Ait., P. reticulata
Willd. and P. terebinthus Mill. The pistachio of commerce is
the only edible species among the 11 species of small trees in
the genus Pistacia; all are characterized by their ability to
exude turpentine or mastic.
Table A.6. World’s major pistachio producers (in millions of kilos)
2002–2004 (Source: FAO, 2004).
Production (millions of kilos)
Year of production
Iran
California, USA
Turkey
Syria
2002
2003
2004
300
310
305
137
54
158
40
50
85
40
50
50
The tree can grow to 7.5–12 m and sometimes
with several twisted trunks. The rough bark is grey. The
pinnate leaves (three, sometimes five) are dark green above and
paler below and vary from ovoid to oblong-ovoid. When young
the leaves are sometimes pubescent. Each leaf at a node
subtends a single axillary bud. Most of these lateral axillary
buds differentiate into inflorescence primordia and produce a
nut-bearing rachis in the following year, on 1-year-old wood.
One or two axillary buds do occur distally of the node on new
growth. This dioecious plant’s panicle has 13 primary
branches, each bearing one terminal and 5–19 lateral flowers
(Plate 6). The small apetalous brownish-green flowers have up
to five sepals. The male flowers have five short stamens and the
female flower has a single tricarpellate, superior ovary (Plate
7). The pedicelled fruit is a single-seeded oval-shaped drupe
(2–2.5 cm long). A semi-dry greenish exocarp and mesocarp
surrounds a hard smooth endocarp and this begins to separate
as the fruit begins to mature (Plate 8). The endocarp dehisces
(splits) longitudinally at the apex after the seed has reached
maturity and maximum size. The exocarp changes from green
DESCRIPTION
22
Anacardiaceae
to red at maturity. The seed has a papery seedcoat and two
greenish-yellow cotyledons.
AND CLIMATIC REQUIREMENTS The tree is
classified as a phreatophyte, having an extensive root system
that allows the tree to mine the soil deeply for water and
nutrients. This author has observed pistachio rootstocks at 9 m
in 10-year-old trees. Pistachios are adapted to survive long
periods of drought.
Areas suitable for pistachio production have long, hot, dry
summers and moderate winters. Pistachios grow best in areas
with 2200–2800 heat units calculated as follows.
ECOLOGY
Heat units =
{mean monthly Tmax + mean monthly Tmin} ⫻ number of days
2
(April–October)
Generally, pistachios should not be planted above 750 m
where summer heat is usually insufficient for complete kernel
development. Elevations of 60–250 m above sea level have
proven ideal in the central Californian valleys. In Iran, major
production occurs at 1200 m on a desert plateau. High
humidity through the growing season promotes foliar fungal
diseases that subsequently overwinter on both male and female
trees and reinoculate the tree the following season.
Pistachios can be successfully grown on a number of soil
types. In California, the sandy loams of the southwest San
Joaquin Valley high in lime and boron have produced the best
yields. In areas with shallow hardpan soils, tree size and
productivity are limited. The tree grows best on well-drained
soils and is intolerant of saturated conditions. It appears to
tolerate alkalinity and salinity well. Trees grown in soils with
soil water extracts up 8 dS/m produce well.
REPRODUCTIVE BIOLOGY The tree can have a long juvenile
period, typically bearing few nuts before 5 years. Full bearing
is achieved between 8 and 10 years of age when irrigated, later
if not. Pistachios require at least 750 h of winter chilling at
temperatures below 7.5°C to produce good, even, timely
budbreak, normal inflorescence development, good fruit set
and normal vegetative growth. When cumulative hours below
7.5°C have fallen to 670, as they did in California in 1977/78,
the bloom and foliation was irregular and delayed, the leaves
were deformed and the yields reduced. Varieties vary in their
chilling requirement from 600 to 1050 h below 7.5°C.
This deciduous tree loses its leaves in the autumn and
remains dormant through the winter. The lateral axillary
inflorescence buds on 1-year-old wood begin to swell in late
March. Within the first 2 weeks of April, the 100–300 flowers
per paniculate rachis are pollinated and set. The tree is
dioecious and a ratio of one staminate tree to eight pistillate
trees is often used. The female flower is apetalous (no petals)
and has no nectaries and does not attract bees. The flowers are
wind pollinated and late spring rains, frosts and strong
desiccating winds interfere with pollination.
ALTERNATE BEARING As the trees age, they develop an
alternate bearing pattern with increasingly large and small
crops. Though the specific mechanism of this phenomenon
has not been defined, evidence suggests that it is a problem of
carbohydrate competition, perhaps mediated by growth
regulator signals. During the period of nut fill in July, the fruit
buds distal to fruit clusters die and abscise. The heavier the
currently borne crop, the greater the subtending bud
abscission. Thus, following a heavy crop year, an individual
branch may bear no fruit. Attempts to alleviate the cycle by
nutritional and growth regulator sprays have not been
successful. However, some success in damping the swing has
been achieved with rejuvenation pruning of older trees.
Currently, pruning appears to be the only method available to
mitigate alternate bearing. Alternate bearing has not been
demonstrated as harmful to the tree and may therefore simply
be a marketing problem. Alternate bearing is not unique to
pistachio trees. Several types of fruit trees show alternate
bearing, however, only pistachios appear to possess the
phenomenon of premature bud abscission.
Fruit set typically follows successful
pollination. After fruit set and through April and May the
endocarp, but not the seed, enlarges. The endocarp is soft and
vulnerable to insect attack and premature splitting that
appears to be a result of rain. In June the endocarp hardens,
and from late June through to early August the seed enlarges
until it fills the shell. Through late August and September the
nut ripens, the radial suture around the endocarp long
circumference splits, the exocarp and mesocarp hull degrades,
and abscission of the individual nut from the rachis starts.
Shoot growth is simultaneous with shell growth. Growth
begins in late April and concludes in late May. The new
extension growth produces pinnately compound leaves with
lateral inflorescence buds in the axils and, generally, a single
apical vegetative bud. The buds differentiate throughout April,
May and June, become quiescent in July, August and
September, and resume differentiation in October. Sometimes
there is an additional flush of shoot growth in late June. This
growth produces primarily vegetative lateral buds as opposed
to the inflorescence buds produced by the spring flush. In
August, leaves distal to heavy fruit clusters often display a
marked depletion and senescence. Most leaves drop by the end
of November and the tree remains dormant through to the
following March.
FRUIT DEVELOPMENT
The growth of the pericarp before the seed has
implications for both blank nut production and shell splitting.
Blank nuts result when there is fruit set and ovary growth, but
the embryo fails to grow, leaving the nutshell empty or blank.
Blanking can occur during two different phases of pistachio
nut development, nut setting and nut filling. It can be affected
by crop load and production practices.
The first empty shells (blanks) are produced as a result of
events that occur at the time of fruit set. This can occur if
pollination occurs but fertilization fails either because pollen
tubes do not complete growth to the ovule, or the ovule is not
viable when the pollen tubes do arrive. The stimulus from
pollination and/or pollen tube growth is sufficient to induce
fruit set and parthenocarpy. Parthenocarpy is common in fruit
that have many seeds rather than in single-seeded fruit such as
the pistachio. Boron leaf levels below 120 ppm dry weight
(August leaf sample) are associated with an increased
percentage of blank nuts at harvest.
Blanks may also develop in July during kernel enlargement,
NUT BLANKS
Pistacia
when a certain percentage of the fertilized embryos fail to
enlarge to fill the shell. Thinning a cluster prior to nut growth
results in a higher percentage of filled nuts on the thinned
cluster. However, the thinned and unthinned clusters have
virtually the same absolute number of filled nuts, though the
thinned clusters may have slightly larger nuts. This has been
demonstrated on a whole-tree scale with pruning experiments.
The edible pistachio, unlike
the species used for rootstocks, is characterized by splitting of
the endocarp at maturity. Splitting begins about the end of
July, at least 1 month before fruit maturity, and continues
through mid-September, progressing simultaneously with
seed maturation. Final seed maturity is indicated by
separation of the hull (exocarp and mesocarp) from the
endocarp shell. The exocarp changes at this time from green
to red and is the most obvious indicator of endocarp splitting.
Pistachio nuts may split along the longitudinal ridges of the
shell and at the tip of the shell. Splitting can occur in any
combination of one or both of the longitudinal ridges, with or
without the tip splitting, or at the tip alone. Investigation of
the anatomical structure of the longitudinal and tip split
regions indicates that these parts of the shell differ from one
another structurally suggesting that different mechanisms may
be involved in shell separation at each site.
There is an inverse relationship between tree crop load, the
percentage of nuts with split shells and the percentage of blank
nuts. As crop load increases, the percentage of nuts with split
shells decreases and the percentage of blank nuts decreases.
Thus, in ‘heavy’ crop years the marketable crop is decreased
by non and splits, and in ‘light’ crop years it is decreased by
blanks.
23
The rootstock seedlings are grown outside in the nursery for
4–12 months and planted, unbudded, 8–16 months from
germination. Planting date is primarily a function of the date
of latest historical frost in spring as the young rootstocks are
not tolerant to temperatures below freezing. It is possible to
transplant bare root nursery stock, but because bare rooted
trees are very sensitive to drying, this is not a common
practice.
ENDOCARP (SHELL) SPLITTING
Horticulture
Both male and female trees are propagated via
standard T-budding, and much less frequently by grafting.
When planting a new pistachio orchard, rootstocks are planted
in February or March and budded in late June, July or early
August. Budding is usually done in early summer when the
scion buds are mature, and the bark on the rootstock is
slipping and active. Red leaves on the newest rootstock growth
is a good indicator that the rootstock bark is slipping.
Generally, trees between 0.7 and 1.5 cm are best for T-budding,
chip budding and saddle grafts.
Pistachio rootstocks are produced from seeds germinated in
January, transplanted to 0.5 m pots or plastic sleeves when
they are approximately 0.25 m tall and moved outside in April.
PROPAGATION
In Iran, California and other pistachio-growing
areas in world, the nut-producing species P. vera is grown on
seedling rootstocks of different Pistacia species or interspecific
hybrids. In California five rootstocks have been used by the
California pistachio industry, three different Pistacia species
and two interspecific hybrids. The rootstocks are P. terebinthus,
Pistacia atlantica and Pistacia integerrima and two hybrids with
P. atlantica and P. integerrima parentage (Table A.8).
P. integerrima is now the standard rootstock due to its
Verticillium dahliae resistance. In other pistachio-producing
countries, the rootstocks are considerably more varied as the
rootstock seed are often selected from unsplit edible nuts
culled from the pistachio processing lines.
ROOTSTOCKS
The tree has an upright growth
habit characterized by a strong apical dominance and a lack of
lateral vegetative buds in older trees. As the trees mature, their
strong apical dominance becomes more marked. These
characteristics have strong implications for young tree
training, mature tree pruning and rejuvenation of fruiting
wood in older trees.
Depending upon rootstock, the pistachio trees have low to
moderate vigour and require 5–7 years of growth before
significant fruiting occurs. For rootstocks, three Pistacia
species, P. terebinthus, P. atlantica, P. integerrima, and one
interspecific hybrid, UCB I, have been planted in California.
Of these, P. terebinthus has the least, and P. atlantica ⫻ P.
integerrima the greatest, vigour.
When irrigated, pistachios are generally trained for the first
5 years after budding. This training is primarily done in the
dormant season though if the scion is on a vigorous rootstock,
and the climate is hot enough, summer tipping for training
can also be done. The cuts are primarily heading or tipping
cuts. The goal is to produce a tree with two to five primary
scaffolds that is amenable to mechanical harvest. After 5 years
of age annual dormant pruning is done to maintain a tree
small enough for mechanical harvest and to produce good
annual crops. Mature tree pruning is almost entirely thinning
cuts as heading cuts decrease the potential buds and do not
PRUNING AND TRAINING
Table A.8. Characteristics of pistachio rootstocks from most * to least ****.
Disease resistance
Rootstock
P. terebinthus
P. atlantica
P. integerrima
P. atlantica ⳯ P. integerrima PGII
P. atlantica ⳯ P. integerrima UCB I
Cold tolerance
*
**
****
***
***
Armillaria
*
****
***
***
*
Nutritional efficiency
Verticillium
Vigour
Early
yield
Zn
****
****
*
****
*
**
**
*
*
*
***
***
**
**
*
**
***
*
***
B
**
***
**
***
Cu
**
***
*
***
24
Anacardiaceae
produce branching due to the strong apical dominance. This is
generally done by hand when the trees are dormant.
Mechanical pruning is being adopted in the California
pistachio industry. Research within the past 15 years has
demonstrated moderate pruning into 1–2-year-old wood by
mechanical side hedging, though non-selective, does not
decrease yield. Mechanical topping does decrease yield, but
the lower cost of mechanical topping compensates for the loss
in yield. Severe mechanical pruning, primarily topping, can
mitigate alternate bearing by producing a strongly vegetative
growth flush that gives a better balance of vegetative and
fruiting buds in the subsequent years.
For pistachio, leaf analysis is
more useful in diagnosing mineral deficiencies and toxicities
than is soil analysis. Leaf samples are taken from current
season’s growth on non-bearing shoots in mid-August (see
Table A.9), after the seed of the current crop has fully filled
the endocarp but before the endocarp has split. Pistachios are
relatively free of macronutrient deficiencies, nitrogen being
the only macronutrient that is occasionally deficient. The trees
are more susceptible to zinc, copper and boron micronutrient
deficiencies. The last, boron, is particularly prevalent if the
tree is on a rootstock with P. integerrima parentage. Boron, in
excess can also produce a toxicity that is visible but rarely
growth limiting.
Zinc deficiency symptoms appear early in the season,
especially if the deficiency is severe. Terminal leaves are small,
chlorotic (i.e. yellow) and appear in tufts, giving rise to the
term ‘little-leaf disease’. In highly deficient cases, terminal
dieback sometimes occurs. Leaves and nuts are markedly
reddish.
The boron requirement in pistachio is the highest known
for any tree crop. Boron deficiency in pistachios manifests at
60 ppm in the leaf tissue as a failure to set fruit, later, at 120
ppm or less, it manifests as thickened, twisted and strapped
leaves.
The symptoms of copper deficiency typically appear in
midsummer, commencing with leaf scorch near the shoot tip
and progressing to defoliation. Immature terminal leaves have
tip burn and are somewhat heart-shaped. Slight shrivelling of
the shoots occurs and small dark lesions appear on the shoot
near the tip. Terminal dieback subsequently occurs in late
summer. Some shoot terminals curl downward, resembling a
shepherd’s crook. Kernels are often badly shrivelled.
NUTRITION AND FERTILIZATION
Table A.9. Critical and suggested levels for August leaf samples.
Element
Nitrogen (N) (%)
Phosphorus (P) (%)
Potassium (K) (%)
Calcium (Ca) (%)
Magnesium (Mg) (%)
Chlorine (Cl) (%)
Manganese (Mn) (ppm)
Boron (Bo) (ppm)
Zinc (Zn) (ppm)
Copper (Cu) (ppm)
Critical value
Suggested range
2.3
0.14
1.8
1.3
0.6
–
30
90
7
4
2.5–2.9
0.14–0.17
2.0–2.2
1.3–4.0
0.6–1.2
0.1–0.3
30–80
120–250
10–15
6–10
Chloride and sodium toxicities are not common except in
the most saline situations. Interestingly, boron toxicity
manifests as a marginal leaf burn much more commonly than
visible sodium or chloride damage.
Since pistachios are phreatophytes and drought
tolerant, they can survive harsh climates without irrigation.
Also, the stomata on their leaves, located only on the abaxial
surface are somewhat less sensitive to desiccating conditions
than stomata on many other trees. For economic production,
adequate irrigation is necessary as it has a significant impact
on young tree development, soil-borne and aerial diseases,
crop yield and quality (both current and subsequent years)
and tree growth. Profitable mature pistachio orchards have
crop coefficients, Kc, ranging from 0.07 to 1.19 through the
growing season. The crop coefficient is the fraction of water
lost from a crop relative to reference evapotranspiration.
Pistachio can use as much as 220 l of water per tree/day
during at the hottest part of the season. The more efficient
drip and microsprinkler irrigation systems are most often used
in modern pistachio orchards. Irrigation research has focused
on controlled deficit irrigation; manipulating water delivery to
maximize the current crop year’s yield and nut quality as well
as the shoot growth that produces the crop in the subsequent
season, while decreasing the total amount of water applied.
IRRIGATION
HANDLING AND STORAGE Pistachios are
harvested by hand cluster removal, knocking or mechanical
trunk shaking onto a catching frame. Because of the high
moisture content, 40–50% on a fresh weight basis, fragility of
their ripe hulls, and open shells within, pistachios are
susceptible to mechanical injury and contamination if they
drop to the orchard floor. Aspergillus flavus, the fungus that
produces the carcinogen aflatoxin, is present in wet orchard
soils and has the potential to infest pistachios that contact the
soil.
Mechanical pistachio harvesters consist of two separate,
self-propelled units about 7.5 m in length. One unit contains a
shaker head that is clamped onto the tree trunk about 0.6 m
above the ground. Most mechanical harvesters can do 0.4 ha
(containing 112 female trees)/h. All Western pistachio
production is mechanically harvested while hand harvesting is
done in Iran, Turkey and Syria.
Shell staining can be greatly increased during postharvest
transport and storage, particularly if high levels of hull
damage were sustained during harvest, as damaged hulls
greatly impede air flow through a bed of nuts. Shell staining
generally increases with increased temperatures and increased
holding times. Nuts will show damage after 8 h at 40°C, 24
hours at 30°C and 40 h at 25°C.
Previously, most processors used a single-stage drying
process using air at 60–71°C for 10–14 h, to achieve 4–6%
wet basis moisture. Conditions of drying were dictated by the
initial moisture content of the nuts and the ambient relative
humidity. Drying of pistachios is now generally a two-stage
process. The hulled nuts are initially dried to 12–13%
moisture in a column dryer designed for grain drying or a
continuous belt dryer. This requires about 3 h at temperatures
below 82°C to prevent shells from splitting so widely the nut
drops out. A rotating drum dryer, at the same temperatures,
POSTHARVEST
Pistacia
can also be used for this first stage of drying. In the second
stage the nuts are transferred to flat-bottomed grain bins
where they are further dried to 4–6% moisture with unheated,
forced air, or air heated to less than 49°C. This second stage of
drying requires 24–48 h. The nuts can then be stored in these
bins until needed. Smaller operations may use bin dryers for
single-stage drying. The desired 4–6% moisture can be
produced by 8 h at 60–66°C.
Sun drying and ambient air drying may be used in small
operations. Drying in the sun requires 3–4 days, with
protective covering to prevent predation by birds and rodents.
Drying nuts in bins with ambient air requires 3 days if
ambient temperatures and relative humidity are sufficient.
Nut depth in the bin should not exceed 1.4 m and air should
be circulating at 0.35 m/s. The major disadvantage to this
method is the potential for fungal growth during the early
stage of drying.
Once dried to 4–6% moisture, but before further
processing, nuts can be held at 20°C and 65–70% relative
humidity for up to 1 year. Temperatures between 0 and 10°C
are recommended for long-term storage (> 1 year). Pistachios
are less subject to rancidity and have a longer storage potential
than almonds, pecans or walnuts. Low oxygen atmospheres (<
0.5%) aid in flavour quality maintenance. In storage, this is
done by reducing the oxygen level and in packages by vacuum
packaging or by flushing with nitrogen to exclude oxygen.
Insects in stored pistachios can be controlled by fumigation
with phosphine. Alternatively, insecticidal controlled
atmospheres and irradiation can be used. Storage of nuts in
0.5% oxygen and 10% carbon dioxide kills all instars of stored
product pests after 2–5 days at 27°C. Temperatures near 0°C or
between 40 and 50°C are also effective. After fumigation the use
of insect-proof packaging is essential to prevent reinfestation.
Size and appearance are the most important components of
pistachio quality. Defects that result in downgrading can be
external, the shell, and internal, the kernel. External shell
defects include total or partial non-splitting, adhering hull
material, stained shells and damage by other means, including
deformity and bird damage. Internal kernel defects include
damage from insects and fungal pathogens, small immature
kernels, rancidity and decay. Size, designated by the number of
nuts per ounce (28 g) is also an important quality attribute.
Grower payout is calculated from the fresh weight of
pistachios delivered to the processing plant, corrected for the
weight of foreign material removed prior to hulling. Prior to
hulling, a grading sample is drawn from each delivery. This
sample is processed individually, the fresh:dry weight ratio
calculated, and third party inspected for the percentage by
weight of split nuts, non-split nuts, blank nuts, nuts with
adhering hulls, light and dark stain, and other defects.
Correction factors for these defects, and for hulling and
drying, are applied to the corrected delivery weight to
calculate grower price. The major components in determining
return to the grower are weight delivered, and the percentages
by weight of the filled split, filled non-split and blank nuts.
DISEASES, PESTS AND WEEDS Three foliar fungal diseases have
been identified as the most destructive to pistachios. They are
Botrytis blossom and shoot blight, caused by Botrytis cinerea
Pers.Fr., Botryosphaeria dothidea and Alternaria late blight,
25
caused by Alternaria alternata. These diseases as well as other
minor diseases have been reported in other countries and in
California, Arizona and Texas where pistachios are grown.
Summaries of the most important diseases with diagnostic
symptoms and signs of each disease and of sporadic diseases
recorded on pistachios are given in Tables A.10 and A.11.
Pistachios are susceptible to a number of insect pests. The
most damaging feed on the young and mature fruit and destroy
nut quality. Virtually none harms the tree or destroys their
host. Among the insect pest of pistachios are Phytocoris, a
genus of hemipteran insects, two species of unarmoured scale
in the family Coccida, the European fruit lecanium,
Parthenolecanium corni (Bouch,), and the closely related frosted
scale, P. pruinosum (Coq), the mireds, Neurocolpus longirostrus
Knight (Miridae), and Calocoris norvegicus (Gmelin) and lygus
(Lygus hesperus). Species of Pentatomidae, most notably the red
shouldered stink bug (Thyanta pallidovirens), Uhler’s and Say’s
stink bugs (Chlorochroa uhleri and C. sayi), the flat green stink
bug (Acrosternum hilare), and some species of Coreidae, the
leaf-footed bugs (Leptoglossus clypealis and Leptoglossus
occidentalis) have produced fruit damage. Navel orange worm
(Amyelois transitella) and the oblique banded leaf roller are
more recent pests. Virtually all insect pest control is chemical.
Thus far biological control has not been well developed for
modern commercial pistachio production.
MAIN CULTIVARS AND BREEDING The diversity of scion
cultivars grown in Iran and the paucity of the same in the West
is another major difference between the modern, Western,
Californian pistachio industry and the traditional Eastern
pistachio industry. The male and female scion cultivars in both
industries are P. vera. The Iranian, Turkish and Syrian
industries have multiple local cultivars, often with different
names, generated by feral trees and isolated producers.
As molecular techniques become more sophisticated the
relationships among the diverse cultivars of the eastern
industries is being determined. Currently, ‘Montaz’, ‘Ohadi’,
‘Agah’ and ‘Kalehgouchi’ are the major Iranian cultivars.
‘Uzun’, ‘Krimizi’ and ‘Siirt’ are among the major Turkish
cultivars. California is a virtual monoculture with single female
and male scions: the female ‘Kerman’ and the male ‘Peters’.
This reliance on a single, vegetatively propagated, female scion
is a potentially devastating situation for the Californian
industry if bud borne viruses appear.
Louise Ferguson
Literature cited and further reading
Crane, J.C. and Iwakari, B.T. (1981) Morphology and reproduction
in pistachio. Horticulture Reviews 3, 376–393.
Crane, J.C. and Maranto, J. (1988) Pistachio Production. University of
California, Agriculture and Natural Resources Publication #2279,
pp. 1–15.
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton, Florida.
Ferguson, L. and Arpaia, M. (1990) New subtropical tree crops in
California. In: Janick, J. and Simon, J.E. (eds) Advances in New
Crops. Timber Press, Portland, Oregon, pp. 331–337.
Food and Agriculture Organization (FAO) (2004) World Pistachio
Situation and Outlook. Available at: http://www.fas.usda.gov.htp/
hort circular/2004/12–10–04/12–04%20pistachio.pdf (accessed
6 November 2006).
26
Anacardiaceae
Table A.10. The most important diseases of pistachio (P. vera).
Common name
Fungal pathogen
Main symptoms and signs of disease
Botryosphaeria panicle and shoot blight
Botryosphaeria dothidea
Verticillium wilt
Verticillium dahliae
Alternaria late blight
Alternaria alternata
Botrytis blossom and shoot blight
Botrytis cinerea
Panicle, leaf, shoot and bud blight; black angular or circular lesions on
green fruit and immature leaves; large black lesions on fruit; large brown
lesions on mature leaves with light beige margins
After sectioning, shoots show discoloured xylem tissues very characteristic
for trees infected by V. dahliae; shoot wilting of certain portions of trees;
and defoliation and sudden death of trees
Small black spots on leaves or necrotic large areas with sporulation in the
centre; leaf blight; black lesions surrounded by red-coloured halo on
epicarp of fruit; fruit blight
Blossom and shoot blight; cankers initiated from male inflorescences
common on ‘02–16’ and ‘02–18’ cultivars; blight and flagging of tender
shoots of both male and female trees
Table A.11. Sporadic pistachio diseases recorded in California between 1985 and 1995.
Common name
Fungal pathogen
Main symptoms and signs
Phomopsis blight
Phomopsis sp.
Shoot and leaf wilting (flagging)
1985
Sclerotinia blight
Root and crown rots
Sclerotinia sclerotiorum
Phytophthora sp.
Shoot and leaf wilting and blight (flagging)
Black lesions on the crown of scion but not extending into the rootstocks
1985
1989
Nut moulds
Armillaria mellea
Aspergillus niger
Decay of roots; white mycelial plaques and distinct black rhizomorphs
1990
Soft water-soaked hull covered with black sporulation on the surface touching the shell 1995
Penicillium expansum and
other Penicillium spp.
Disintegration of hull covered with bluish, greenish powdery sporulation.
Greenish, bluish sporulation leading to soft disintegration of the hulls
Kaska, N. (2002) Pistachio nut growing in the Mediterranean Basin.
Acta Horticulturae 591, 443–455.
Spiegel-Roy, P. (1987) Pistacia. In: Halvey, A.H. (ed.) CRC Handbook
of Flowering. Vol 4. CRC Press, Boca Raton, Florida, p. 8398.
US Department of Agriculture (USDA) (2006) Nutrient Database
for Standard Reference – Release 18. Available at:
http://www.ars.usda.gov/main/site_main.htm?modecode=1235
4500 (accessed 10 January 2006).
Woodroof, J.G. (1979) Tree Nuts, 2nd edn. AVI, Westport,
Connecticut, pp. 1–603.
Sclerocarya birrea
marula
Marula, Sclerocarya birrea (A.Rich.) Hochs. (Anacardiaceae),
is one of a select group of outstanding dryland African fruit
trees. Much valued throughout a range encompassing over 30
countries extending at its widest some 8000 km, the species
has been assigned names in numerous languages (Hall et al.,
2002). The best known of these are marula in southern Africa,
sakoa in Madagascar, mngongo in East Africa, nobiga in
Burkina Faso and neighbouring countries, birr (hence birrea)
in Senegal and homeid in Sudan.
World production and yield
Considerable year-to-year fruit crop variation occurs between
individual trees of S. birrea. Crops of over 0.5 t fresh fruit in a
year have often been reported but are not generally
representative. For a typical population, an estimated average
annual crop of 60–90 kg fresh weight/tree (2000–3000 fruit) is
more realistic (Lewis, 1987; Shackleton, 2002).
Products using the fruit flesh as initial raw material are
Year recorded
1995
internationally and regionally (mainly South Africa) marketed
on a small scale. Currently, only a minute fraction (under 2500
t) of the potential annual fruit crop is used for this and the
workforce involved is small (less than 1000 jobs, many
seasonal). Liqueur, distinctive for the flavour imparted by S.
birrea pulp, has been exported from South Africa for around
20 years while fruit juice, jams and chutneys are cottageindustry products in South Africa, Namibia and Botswana.
Commercial interest in the oil is constrained by the difficulty
of separating kernels from the endocarp and the small kernel
weight per fruit (1.5% of fruit fresh weight; c.10% of whole
endocarp). Nevertheless, the high oil quality is prompting
efforts to overcome these complications and pilot initiatives
have led to internationally marketed cosmetic products
containing the oil.
Uses and nutritional composition
The fruit is a drupe, with a fleshy, edible mesocarp enclosing a
hard endocarp containing 2–3 seeds rich in a useful oil.
Archaeological evidence indicates that the fruit have been
exploited in South Africa for over 10,000 years (Wynberg et
al., 2002), and in West Africa for over 1000 years (Neumann et
al., 1998). In the first half of the 20th century analyses
revealed the suitability of the oil for soap making and the high
vitamin C content of the flesh (Shone, 1979). Serious
commercial initiatives are recent, however – since about 1980.
In the diversity of useful roles S. birrea has few rivals, but it
is the nutritional value and famine food significance of the
fruit flesh and kernels that has brought the tree special cultural
status. Other significant uses are as utility timber, as livestock
feed (fruit and foliage) and medicinal (Teichman, 1983; Hall,
Sclerocarya
2002). Fruit are important and the vitamin C-rich beer from
them (Weinert et al., 1990) is a regular rural market
commodity. Both the pulp and the kernels have been
nutritionally evaluated.
Reported values for proximate fractions (of dry weight) of
the fruit pulp are: carbohydrate (68–85%); protein (3–7%); fat
(1–10%); fibre (6–10%); ash (2–9%). Vitamin C concentration
(mg/100 g fresh weight) is high in the skin (150–200), flesh
(150–400) and juice (100–200). The kernel is rich in energy
(2700 kJ/100 g), phosphorus (700–1900 mg/100 g) and
potassium (500–700 mg/100 g). Fresh weight compositional
values are given in Table A.12. Crude fat accounts for 55–65%
of the dry weight and protein accounts for 20–35%. The
dominant fatty acid is the unsaturated oleic acid (64–75% of
total fatty acid content). A saturated acid, palmitic, contributes
a further 11–18%. The main kernel amino acids are glutamic
acid (180–270 mg/g of protein) and arginine (140–160 mg/g
of protein). Lysine (18–24 mg/g of protein) and phenylalanine
(48 mg/g of protein) contents are low. The oil is poor as a
source of vitamin E but noteworthy for its favourable
saturated:unsaturated fatty acid ratio and oxidative stability.
Botany
Sclerocarya is a genus of
two species in the Anacardiaceae (Sapindales, Eurosid II clade
– Soltis et al., 2000; Judd et al., 2002), referred to the tribe
Spondiadeae. The other member of the genus, the Kenya
endemic Sclerocarya gillettii, is the nearest relative.
Collectively, however, the African/Madagascan Spondiadeae
(c.50 species, currently in ten genera) invite taxonomic
reassessment (Wannan and Quinn, 1991; Schatz, 2001).
Three subspecies of S. birrea (syn. Poupartia birrea) are
recognized: subsp. birrea, from Senegal to Ethiopia and south
to Tanzania; subsp. caffra, from Tanzania to South Africa and
TAXONOMY AND NOMENCLATURE
Table A.12. Proximate fruit composition per 100g of Sclerocarpa birrea flesh
and nut on a fresh weight basis.
Proximate
Water
Energy (kJ)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Flesh (%)
85
225
0.5
57.3
12
1.2
0.9
mg
20.1
0.5
25.3
11.5
317
2.24
mg
194
0.03
0.02
0.27
Nut (%)
4
2703
28.3
0.4
3.7
2.9
3.8
mg
118
4.87
462
808
601
3.81
mg
–
0.42
0.12
0.72
27
Madagascar; and subsp. multifoliolata endemic to Tanzania.
The subspecies are separated on inflorescence and leaflet
characters. Consideration here is at specific rather than
subspecific level. The concentration of Sclerocarya and related
genera in Africa suggests an African origin, apparently south
of the equator. Northward migration of subsp. birrea could
have been via an arid corridor near the Indian Ocean coast in
the late Miocene/early Pliocene, with later spread westward.
This deciduous tree usually reaches 9–12 m in
height, occasionally more, with a taproot and widespreading
lateral roots. At maturity the crown is rounded, but spreads
impressively in very old individuals. In full leaf, the foliage is
fairly dense and concentrated at the ends of stout branchlets.
On old trees the bark of the trunk is rough, with roundish
reddish-brown scales initially exposing light patches after
shedding. The bark of young trees and shoots is smoother and
paler. The leaves are alternate and imparipinnate, mostly with
(depending on subspecies) 6–18 opposite, glabrous oblong to
ovate, usually entire, leaflets 2–9 cm long.
With infrequent exceptions, the species is dioecious
(Teichman, 1982). The male inflorescences, which arise in the
axils of new or recently shed leaves, are racemes carrying
several small groups of reddish flowers. The female
inflorescence is reduced to a spike of one to four reddish
flowers. The flowers are pentamerous, with short sepals and
petals c.5 mm long. In each male flower there are 15–25
stamens around 3 mm long, surrounding a fleshy disc. Each
female flower contains a subglobular ovary encircled by a disc
with 15–25 staminodes around it. The ovary is of two to three
cells, each with a single ovule. Three short lateral styles
terminate in capitate stigmas. The drupe, 3–4 cm in diameter
when ripe, is covered by a thick, tough exocarp, beneath which
is a fibrous, fleshy mesocarp adhering to a very hard, obovoid
endocarp. Each of two to three compartments within the
endocarp contains a flattened seed 15–20 mm long.
DESCRIPTION
AND CLIMATIC REQUIREMENTS North of the
equator, S. birrea occurs across Africa from Senegal on the
Atlantic seaboard (17°W) to inland Eritrea and Ethiopia.
Further south, in eastern Africa, the species reaches the
Indian Ocean coast at 40°E, in Kenya. In northern Madagascar
the eastern limit is 50°E. The latitudinal range is from
17°15’N (Aqr Mountains, Niger) to 31°S (Port Shepstone,
South Africa). It is absent from the equatorial humid forest
region. Outside the natural range the most important
introduction has been to Israel for development as a potential
horticultural crop (Mizrahi and Nerd, 1996), and it is widely
thought that presence in Madagascar has also arisen from
introduction in the distant past.
Sclerocarya birrea is mostly recorded from elevations below
1700 m, and associated with mean annual temperature > 19°C.
South of the Tropic of Capricorn some populations experience
occasional frost. In the natural range of the species the rainfall
regime is strongly seasonal, with mean annual totals typically
500–1250 mm, 4–7 months having > 50 mm mean
precipitation. Most occurrences are on deep, well drained,
sandy or loamy soil, but heavier soils free of waterlogging
support thriving populations in parts of southern Africa.
Sclerocarya birrea is typically a woody species of wooded
ECOLOGY
28
Anacardiaceae
grassland, parkland or grassland and individuals tend to be
well separated and with full crown exposure. Natural
populations are sparse, published assessments for areas of 1 ha
or larger indicating numbers of individuals ⭓ 5 cm diameter
to be well below five per ha (Nghitoolwa et al., 2003). Selective
elimination of male individuals for fuel or utility wood has
modified gender ratios in favour of females in some areas.
There is no regional consistency in the commonly associated
woody species in the southern part of the range, but north of
the equator Acacia seyal, Balanites aegyptiaca, Commiphora
africana and Tamarindus indica often occur with S. birrea.
REPRODUCTIVE BIOLOGY Fruiting apparently starts in wild
trees at an age of 7 to 10 years. Fruit are produced in quantity
by an age of 15 to 20 years but the yield increases as the trees
grow, to an age of well over 100 years (Shackleton et al., 2002).
Flowering is a dry season event, with fruit being dispersed as
the succeeding rainy season starts. The flowers are small and
entomophilous, producing little fragrance but a fair quantity
of nectar attractive to bees, the main pollinator. Fully
developed flowers open between dawn and midday. In the male
flowers the anthers dehisce sequentially over 24–36 h, the
pollen grains remaining viable for at least 12 h and often for
more than 48 h. In the female flowers the stigmatic surfaces
are receptive when the flowers open and remain so for up to 72
h. Secretion of nectar ceases on fertilization. The main natural
disperser is the elephant, which swallows the fruit and
transports them considerable distances. Today, however, cattle
and humans are more important.
The anthesis to fruit maturity is 2–5 months, depending on
temperature and plant moisture status. In years with a good
crop, mature fruit fall over a 1–3 month period.
Horticulture
Propagating S. birrea from seed is possible but
effectively restricted to research and demonstration situations.
There is little justification for inclusion in nursery-raised
plants for distribution to farmers because the gender of
individuals usually remains in doubt for 8–10 years and 50%
will be male. There is some tradition of vegetative propagation
using thick cuttings, which retain the gender of the source
individual (Wynberg et al., 2002) but success with
conventional cuttings has been low. Given the dioecious
character, grafting (Soloviev et al., 2004) has received
attention for combining known female scions with robust
juvenile rootstocks but this, too, has proved difficult and
remains at an exploratory stage. Recent experimental work
using Agrobacterium tumefaciens suggests possible future use
and vegetative multiplication of genetically transformed S.
birrea (Mollel and Goyvaerts, 2004).
PROPAGATION
CULTIVATION When nursery seedlings of S. birrea are raised,
the unit sown is the cleaned whole endocarp (200–500/kg,
varying with locality), even though the two to three true seeds
within contribute only 10% of the weight of this. A soaking
treatment (24–48 h) is applied to loosen the caps closing the
germination apertures, so they are pushed aside by pressure
from the germinating seed. The first seeds are likely to
germinate in 4 days but others will take longer. Over 4 weeks,
seedlings should have developed from at least 80% of the
endocarps. A medium of coarse sand enriched with manure is
used, with a regime of watering twice daily. If root pruning or
re-potting is undertaken, shade should be provided for the
next 2 days. In the continuously warm West African
environment, most seedlings should be 30–40 cm tall and
ready for outplanting after 11 weeks. In the southern part of
the range, winter sowing should be avoided and the nursery
period is longer (up to 6 months).
POSTHARVEST HANDLING AND STORAGE Sclerocarya birrea
has the unusual characteristic of dropping the mature fruit in
a green state to complete the ripening process and turn yellow
on the ground over the next few days. No systematic storage
arrangements for fresh fruit are in general use but
experimental investigations have indicated that satisfactory
storage for 16 days is possible at 4°C (Weinert et al., 1990).
Local processing for beer is the dominant use and generally
undertaken at or close to the parent trees. The fruit selected
for processing are those at the appropriate ripening stage
(slightly green). These are collected together in a shaded place
to further ripen to a creamy yellow colour and are then
processed. If commercial enterprises are supplied, fresh fruit
are collected from source or delivered by middlemen. Where
interest is in the kernel as a reserve dietary item, there is a
tradition of storing whole endocarps in bulk off the ground.
This approach preserves the quality of the kernels, which have
a short shelf life once extracted.
PESTS Fruit pests include a beetle (Carpophilus
hemipterus), a moth (Cryptophlebia leucotreta) and a fly
(Ceratitis cosyra). Sclerocarya birrea is infected by several
loranthaceous mistletoes among which Erianthemum dregei and
Pedistylis galpinii are strongly associated with the tree in South
Africa (Dzerefos et al., 2003) but this impact has not been
quantified.
MAIN
A chromosome number of
2n = 26 has been reported for Mozambique plants (Paiva and
Leitao, 1989). In southern Africa, through efforts to find a
population including individuals yielding fruit as large as 100
g fresh weight with high juice, sugar and vitamin C content,
superior wild trees have been identified in various locations.
Some of these are under consideration for registration as
cultivars (Wynberg et al., 2002). On a wider front, variation is
being assessed through regional and national multilocational
provenance trials in eastern and southern Africa. Exploratory
genetic studies have centred largely on germplasm of southern
and East African origin. A possibility that historical events
have modified a basic pattern of geographic variation has
emerged but needs confirmation through more comprehensive
sampling.
John B. Hall
MAIN CULTIVARS AND BREEDING
Literature cited and further reading
Arnold, T.H., Wells, M.J. and Wehmeyer, A.S. (1986) Khoisan food
plants: taxa with potential for future economic exploitation. In:
Wickens, G.E., Goodin, J.R. and Field, D.V. (eds) Plants for Arid
Lands. George Allen and Unwin, London, pp. 69–86.
Dzerefos, C.M., Witkowski, E.T.F. and Shackleton, C.M. (2003)
Spondias
Host-preference and density of woodrose-forming mistletoes
(Loranthaceae) on savanna vegetation, South Africa. Plant Ecology
167, 163–177.
Hall, J.B. (2002) Sclerocarya birrea (A.Rich.) Hochst. In: Oyen,
L.P.A. and Lemmens, R.H.M.J. (eds) Plant Resources of Tropical
Africa. Precursor. PROTA Programme, Wageningen, pp. 127–131.
Hall, J.B., O’Brien, E.M. and Sinclair, F.L. (2002) Sclerocarya birrea:
a Monograph. School of Agricultural and Forest Sciences,
University of Wales, Bangor, 157 pp.
Judd, W.S., Campbell, C.S., Kellogg, C.A., Stevens, P.F. and
Donoghue, M.J. (2002) Plant Systematics: a Phylogenetic
Approach, 2nd edn. Sinauer, Sunderland, Massachusetts.
Lewis, D.M. (1987) Fruiting patterns, seed germination, and
distribution of Sclerocarya caffra in an elephant-inhabited
woodland. Biotropica 19, 50–56.
Mizrahi, Y. and Nerd, A. (1996) New crops as a possible solution for
the troubled Israeli export market. In: Janick, J. (ed.) Progress in
New Crops. American Society of Horticultural Science Press,
Alexandria, Virginia, pp. 37–45.
Mollel, H.N. and Goyvaerts, E.M.A. (2004) Preliminary examination
of factors affecting Agrobacterium tumefaciens-mediated
transformation of marula, Sclerocarya birrea subsp. caffra
(Anacardiaceae). Plant Cell, Tissue and Organ Culture 79, 321–328.
Neumann, K., Kahlheber, S. and Uebel, D. (1998) Remains of woody
plants from Saouga, a medieval West African village. Vegetation
History and Archaeobotany 7, 57–77.
Nghitoolwa, E., Hall, J.B. and Sinclair, F.L. (2003) Population status
and gender imbalance of the marula tree, Sclerocarya birrea subsp.
caffra in northern Namibia. Agroforestry Systems 59, 289–294.
Paiva, J. and Leitao, M.T. (1989) Chromosome counts for some taxa
of tropical Africa. II. Boletim da Sociedade Broteriana 62, 117–130.
Shackleton, C.M. (2002) Growth and fruit production of Sclerocarya
birrea in the South African lowveld. Agroforestry Systems 55,
175–180.
Shackleton, S.E., Shackleton, C.M., Cunningham, T., Lombard, C.,
Sullivan, C.A. and Netshiluvhi, T.R. (2002) Knowledge on
Sclerocarya birrea subsp. caffra with emphasis on its importance as
a non-timber forest product in South and southern Africa: a
summary. Part 1. Taxonomy, ecology, traditional uses and role in
rural livelihoods. Southern African Forestry Journal 194, 27–41.
Schatz, G.E. (2001) Generic Tree Flora of Madagascar. Royal Botanic
Gardens, Kew, 477 pp.
Shone, A.K. (1979) Notes on the marula. South Africa Department of
Forestry Bulletin 58, 1–89.
Soloviev, P., Niang, T.D. and Gaye, A. (2004) Propagation par
greffage du prunier d’Afrique [Sclerocarya birrea (A.Rich.)
Hochst.] au Sénégal. Fruits 59, 275–280.
Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach, D.C.,
Zanis, M., Savolainen, V., Hahn, W.H., Hoot, S.B., Fay, M.F.,
Axtell, M., Swenson, S.M., Prince, L.M., Kress, W.J., Nixon,
K.C. and Farris, J.S. (2000) Angiosperm phylogeny inferred from
18S rDNA, rbcL, and atpB sequences. Botanical Journal of the
Linnean Society 133, 381–461.
Teichman, I. von (1982) Notes on the distribution, morphology,
importance and uses of the indigenous Anacardiaceae: 1. The
distribution and morphology of Sclerocarya birrea (the marula).
Trees in South Africa 34, 35–41.
Teichman, I. von (1983) Notes on the distribution, morphology,
importance and uses of the indigenous Anacardiaceae: 2. The
importance and uses of Sclerocarya birrea (the marula). Trees in
South Africa 35, 2–7.
29
Wannan, B.S. and Quinn, C.J. (1991) Floral structure and evolution
in the Anacardiaceae. Botanical Journal of the Linnean Society 107,
349–385.
Weinert, I.A.G., Wyk, P.J. van and Holtzhausen, L.C. (1990) Marula.
In: Nagy, S., Shaw, P.E. and Wardowski, W.F. (eds) Fruits of
Tropical and Subtropical Origin: Composition, Properties and Uses.
Florida Science Source, Lake Alfred, Florida, pp. 88–115.
Wynberg, R., Cribbins, J., Leakey, R., Lombard, C., Mander, M.,
Shackleton, S. and Sullivan, C. (2002) Knowledge on Sclerocarya
birrea subsp. caffra with emphasis on its importance as a nontimber forest product in South and southern Africa: a summary.
Part 2. Commercial use, tenure and policy, domestication,
intellectual property rights and benefit-sharing. Southern African
Forestry Journal 196, 67–77.
Semecarpus spp.
marking trees and tar tree
The genus Semecarpus L. (Anacardiaceae) consists of about 50
species from Indo-China–Malaysia, Micronesia and the
Solomon Islands. The young fruit have a tarry resin which
when mixed with lime water or alum is used as a marking ink
and dye. The resin from all parts of these trees frequently
causes dermatitis due to phenolics and catechol. The nut of a
number of the species are edible as is the fruit (swollen
pedicel) when ripe.
Five species have fruit and nuts that are consumed.
Semecarpus anacardium L. (synonyms Semecarpus latifolia
Pers., Anacardium latifolium Lam., Anacardium officinarum
Gaertner and Anacardium orientale Auct.) is known as the
Indian marking nut tree, oriental cashew nut, bhilawa, kidney
bean of Malacca and ostindischer tintenbaum (German). This
medium-sized tree (15 m tall) has simple oblong to ovateshaped alternate leathery leaves (20–60 cm long) that have
hairs on the underside. The small greenish-yellow flower
occurs on a panicle. The ripe, sweetish black fruit (2.5 cm
long), which is the swollen pedicel, and the roasted nut are
eaten. Young fruit are pickled in salt and vinegar. The pericarp
of the fruit yields the black tarry juice known as bhilawan oil
which when mixed with lime water yields an ink. The juice or
oil is insoluble in water but soluble in alcohol, and has been
used as an indigenous medicine for treatment of warts and
piles.
Semecarpus australiensis Engl. is called ganyawu, tar tree and
marking nut. This Australian native cashew is a well-known
food source for aboriginal people of rainforest areas of northeastern Queensland and the Northern Territory. The seed
after processing can be eaten. The processed seeds (nuts) are
tasty. The fleshy orange or red fruit can be eaten after baking.
The fruit of Semecarpus cassuvium Roxb. and Semecarpus
longifolius Blume and the nut of Semecarpus atra (Forster) Vieill.
are also reported to be eaten, with precautions. Robert E. Paull
Spondias cytherea
ambarella
Ambarella, Spondias cythera Sonn. (Anacardiaceae), is from
the Indo-Malaysian region to Tahiti and has been spread
throughout the tropics. It is known as ambarella, Otaheite
apple, wi or vi or evi-apple and greater hog plum in English,
also as Tahitian quince or apple, Polynesian plum and golden
apple. In Malaysia and Indonesia, it is known as kedondong; in
30
Anacardiaceae
Thailand as ma kok farang; in Cambodia as mokak; in Vietnam
as coc, pomme de cythere; in Costa Rica as juplon; in
Colombia as hobo de racimos; in Venezuela as jobo de la India
and mango jobo; in Brazil as caya – manga; in the Philippines
heri, in Burma gway; and in Laos kook hvaan.
Three related species from South-east Asia have edible
fruit. Spondias acida Bl. from the western Indo-Malayan area
has acid fruit. Spondias novoguineensis Kostermans from New
Guinea to the Solomon Islands is semi-cultivated and
frequently confused with S. cythera. Spondias pinnata (Keonig
ex. Linn. F) Kurz. is from Burma, India and Thailand and is
cultivated for its edible fruit.
Table A.13. Proximate fruit composition per 100 g of Spondias cytherea
(Source: Leung et al., 1972).
Proximate
%
Edible portion
Water
Energy (kcal)
Protein
Lipid
Carbohydrate
Fibre
Ash
70
86.9
46
0.2
0.1
12.4
1.1
0.4
Minerals
World production
Ambarella has been introduced from Melanesia through
Polynesia to all tropical areas. Fruit are sold on the markets in
Vietnam, Laos, Cambodia, Gabon and Zanzibar. It is
cultivated in Queensland, Australia, Pacific Islands, Cuba,
Haiti, Dominican Republic, Central America, Venezuela and
from Puerto Rico to Trinidad in the West Indies (Morton,
1987).
Uses and nutritional composition
The pulpy flesh is eaten fresh when still firm and crisp, and
also when ripe when the flesh is soft. When firm, the flesh is
still crisp and juicy, and subacid with a pineapple-like
fragrance and flavour. The green fruit is used for salads,
curries, pickles and juices. Ripe fruit are also stewed and used
for jams, jellies, juice and canned. As the fruit ripens, the skin
and flesh turn golden yellow or orange. If soft, the aroma is
musky and flesh is difficult to slice due to the tough fibres
extending from the seed. The fruit’s vitamin C content is
reported at 36 mg/100 g and it is a good source of iron (Table
A.13). Impact aroma volatiles include ethyl(S)(+) 2-methyl
butyrate, ethyl isovalerate, ethyl propionate, ethyl butyrate,
linalol and trans-pinocarveol (Fraga and Rezende, 2001).
Young leaves are eaten raw or steamed as a vegetable with
salted fish.
The fruit, leaves and bark have been reported to have
medicinal value for the treatment of sores, wounds and burns.
The bark is used mixed with other species to treat diarrhoea.
The light-brown wood has a low density and has little timber
value, though is has been used for canoes in the Society
Islands. The gum has a high viscosity with galactose as the
main component.
Botany
AND NOMENCLATURE The synonym for S.
cytherea Sonnerat is Spondias dulcis Soland Ex. Forst.
TAXONOMY
The rapidly growing tree can reach 10–18 m
and is upright and symmetrical with a rounded crown. The
bark is a light greyish brown and nearly smooth, often with
four to five small buttresses. The leaves are pinnate (20–60 cm
long) composed of 9–25 pairs of glossy, elliptic or obovateoblong leaflets (6.25–10 cm long) on a short petiole (Fig. A.4).
The whitish flowers are small and inconspicuous and are
borne in large terminal panicles (50 cm long) appearing before
DESCRIPTION
mg
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
56
0.3
–
67
95
1
Vitamins
mg
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
36
0.05
0.2
1.4
205 IU
the leaves. In each panicle, male, female and perfect flowers
occur on short pedicels (1–4 mm). The calyx lobes are
triangular (0.5 mm) and the petals (2.5 ⫻ 1 cm) are ovateoblong. The fruit (up to 0.45 kg) are borne on long peduncles
in bunches of a dozen or more and have tough thin skin that is
often russetted. The fruit is an ellipsoid or globose drupe
(4–10 ⫻ 3–8 cm) and changes from bright green to bright
orange on ripening. The tough endocarp has irregular spiny
and fibrous protuberances and contains one to five flat seeds.
The tree grows well
in the warm subtropics and tropics and grows up to 700 m in
the tropics. Shaded trees produce little fruit and full exposure
to sun is necessary. The branches are easily broken by strong
winds and require a sheltered location. The tree is drought
tolerant and may briefly lose its leaves under stress. Ambarella
grows on all well-drained soils.
ECOLOGY AND CLIMATIC REQUIREMENTS
The tree grows rapidly and flowers
and bears fruit in 4 years. In the humid tropics, the tree
produces more or less continuously. Vegetative flushing and
flowering occur together. In areas with a dry season, flowering
occurs on trees that are nearly leafless from lack of soil
moisture. Subtropical conditions lead to flowering in spring.
The dwarf selection flowers and produces fruit all year
around. Fruit bats may assist in seed dispersal.
REPRODUCTIVE BIOLOGY
FRUIT DEVELOPMENT
The fruit takes 6–8 months to mature.
Horticulture
PROPAGATION Often the tree is propagated from seeds that
germinate in 1 month. Some seeds are polyembryonic.
Spondias
31
Grenada, attempts have been made to cross the usual
ambarella with the dwarf type that grows to a height of 3 m
and produces fruit of about 65 g weight in about 12 months.
Reginald Andall and Robert E. Paull
Literature cited and further reading
Fig. A.4. Leaf, flower and fruit of Spondias cytherea (Source: Verheij
and Coronel, 1992).
Vegetative propagation is possible by hardwood cuttings and
air layering, while grafting or shield budding on Spondias
rootstocks is also used. Seedlings and grafted trees are more
vigorous and may fruit in 4 years while budded trees and
cuttings have less vigour. Tree spacing varies from 7.5 to 12 m.
It can be grafted on to its own rootstock. In
India, S. pinnata Kurz. is used.
ROOTSTOCKS
PRUNING AND TRAINING Fruit pruning within the panicle
increases the size of the remaining fruit (Andall and Baldeo,
2000).
DISEASES, PESTS AND WEEDS
The larvae of a beetle attacks
the leaves in Indonesia, while in Costa Rica the bark is eaten
by a wasp that causes necrosis. In the Caribbean, a disease
called gummosis results in debarking of the stem and
subsequent death of the tree and also black spots on the fruit.
No major diseases are reported. Weeds are only problematic in
the very young stages of growth and can be managed manually
or with herbicides.
MAIN CULTIVARS AND BREEDING There are selections, but no
recognized cultivars. Large variations in fruit quality (fruit
acidity and spiny seeds) have been reported. Recently in
Andall, R.P. and Baldeo, S. (2000) The effect of fruit and
inflorescence pruning on fruit size and yield of dwarf golden
apples (Spondias cytherea Sonn.). Tropical Fruits Newsletter 36/37,
12–14.
Fraga, S.R.G. and Rezende, C.M. (2001) The aroma of Brazilian
ambarella fruit (Spondias cytherea Sonnerat). Journal of Essential
Oil Research 13, 252–255.
Kostermans, A.J. (1991) Kedongdong, Ambarella, Amra. The
Spondiadeae (Anacardiaceae) in Asia and the Pacific Area.
PROSEA Foundation, Bogor, Indonesia, pp. 1–100.
Leung, W.T.W., Buthum, R.R. and Chang, F.H. (1972) Food
Composition Tables for Use in East Asia, Part 1. National
Instititutes of Health, Bethesda, Maryland.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resources
Systems Inc., Winterville, North Carolina, pp. 240–242.
Persad, C. (1997) Golden apple – Spondias cytherea, Sonn. Tree.
Tropical Fruits Newsletter 23, 11–13.
Pinto, G.L. de, Martinez, M., Sanabria, L., Rincon, F., Vera, A.,
Beltran, O. and Clamens, C. (2000) The composition of two
Spondias gum exudates. Food Hydrocolloids 14, 259–263.
Popenoe, J. (1979) The genus Spondias in Florida. Proceedings of
Florida State Horticultural Society 92, 277–279.
Verheij, E.W.M. (1992) Spondias cytherea Sonnerat. In: Verheij,
E.W.M. and Coronel, R.E. (eds) Edible Fruits and Nuts. Plant
Resources of South East Asia No. 2. PROSEA Foundation, Bogor,
Indonesia, pp. 287–288.
Verheij, E.W.M. and Coronel, R.E. (eds) (1992) Edible Fruits and
Nuts. Plant Resources of South East Asia No. 2. PROSEA
Foundation, Bogor, Indonesia.
Spondias mombin
yellow mombin
Yellow mombin, Spondias mombin L. (Anacardiaceae), is in the
same family as cashew, mango and pistachio. The fruit, also
known as the tropical plum, has several regional names: in
English it is hog plum or yellow mombin in North America
and the Caribbean islands; thorny hog plum in Malaysia;
Ashanti plum in Ghana; in Latin America the Spanish names
are caimito, ciruela agria, jobo, jobo jacote, ciruela de monte,
ciruela amarilla, obo and uvo; in French Guiana the name is
prunier mombin; and in Brazil it is called taperebá, cajá or
cajá-mirim.
Spondias mombin is the most economically significant
species within the genus that occurs in Brazil. No commercial
orchards occur in Brazil and all fruit is collected from wild
plants. The fruit is attracting attention not only to meet local
market needs in the region of occurrence, but also from other
parts of the country where it is highly appreciated and
commercialized as frozen pulp. The frozen pulp is one of the
most prized pulps in Brazilian markets and the price remains
high even during the harvest season. According to pulp
manufacturers, purchasers may even request a supply of
yellow mombin pulp as a condition for buying other pulps.
32
Anacardiaceae
The major limitation for the cultivation is the height of the
tree – up to 30 m – and the lack of cultivars that can be
recommended for commercial orchards.
Table A.15. Proximate analysis of ripe Spondias mombin L. edible pulp per
100 g (Source: Leung and Flores, 1961; Morton, 1987).
World production and yield
Moisture
Energy (kcal)
Protein
Fat
Total carbohydrate
Fibre
Ash
The species is found throughout tropical America and in the
tropical regions of Africa and Asia. In Brazil, the trees are
found either isolated or in groups, in forests, grasslands and
pastures, as well as in backyards of the north and north-east
states. The plant is native to moist lowland forests from
southern Mexico to Peru and Brazil. It is grown commercially
to a limited extent. In Brazil, all fruit is obtained from native
plants. The plant produces once a year, and harvest season
varies according to climate conditions, usually between early
December and late February. Adult trees in Mexico produce
over 100 kg/year.
Uses and nutritional composition
The fruit is a drupe, with a thin layer of yellow, translucent,
fleshy, juicy, sweet-sour and very aromatic mesocarp
surrounding a bulky endocarp. In Brazil, the ripe fruit is eaten
fresh, the edible portion being the exocarp and mesocarp,
which together may represent as much as 81% of the fresh
weight (Table A.14). Extracted pulp is frozen and sold as such
to restaurants, hotels and snack bars to make juice. It is also
used in ice cream and jam. In other Latin American countries,
it is stewed with sugar and made into jam and the green fruit
can be pickled. The fruit has a good supply of ascorbic acid
(Table A.15). The major volatile components are ethyl acetate,
ethyl butynate, ethyl hexanoate, hexyl butynate and linalool.
Table A.14. Characterization of ripe Spondias mombin L. fruit, Fortaleza,
Ceara, Brazil, 2000 (Source: adapted from Filgueiras et al., 2000).
Characteristics
Fruit weight (g)
Pulp + exocarp (%)
Endocarp (%)
Length (mm)
Diameter (mm)
Soluble solids (%)
Titratable acidity (% citric acid)
Soluble solids/titratable acidity ratio
pH
Soluble sugars (%)
Reducing sugars (%)
Starch (%)
Total pectin (%)
Soluble pectin (%)
Pectin fractions in alcohol insoluble solids
High methoxylation (%)
Low methoxylation (%)
Protopectin (%)
Pectin methylesterase (UAE)a
Polygalacturonase (UAE)
Total vitamin C (mg/100g)
Water-soluble phenolics (%)
Methanol-soluble phenolics (%)
50% Methanol-soluble phenolics (%)
aUAE,
units of enzyme activity.
Average figures
19.92
81.65
18.34
43.10
32.20
11.56
1.03
11.23
3.17
8.41
7.65
52
28
7
10.30
2.11
2.21
36.231
18.32
36.86
12
11
14
Proximate
Minerals
Calcium
Iron
Phosphorus
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
g
73–89
22–48
1.3–1.4
0.1–0.6
8.7–10.0
1.2
0.65
mg
31
2.8
31
mg
28–46
0.095
0.05
0.5
71 IU
The yellow to yellowish-brown light and flexible wood of S.
mombin is used to some extent for woodcraft, the bark and
leaves contain ellagitannins with therapeutical properties. Like
other Spondias, the tannin in the bark is used for tanning and
dyeing. An exuded gum is used as glue, and the young leaves
can be eaten as a cooked vegetable.
A decoction of the bark serves as an emetic substance, a
remedy for diarrhoea, dysentery, haemorrhoids, gonorrhoea
and leucorrhoea. The powdered bark and dried leaves are
applied to wounds. A tea made from flowers and leaves is
recommended to relieve stomach aches, biliousness, urethritis,
cystitis and eye and throat inflammation.
Tannins with antiviral activity against the viruses Herpes
Simplex type 1 and Coxsackie B2 were isolated from the
alcohol extracts of leaves and branches of S. mombin. In
Nigeria there are reports of the antimicrobial potential of leaf
extracts being comparable to those of ampicillin and
gentamycin, as well as its use by traditional medical
practitioners in the treatment of various nervous diseases (Abo
et al., 1999). A decoction of leaves is traditionally used in the
Venezuelan Amazon to treat malaria.
Botany
The genus Spondias was
created by Linnaeus in 1753, with S. mombin L. as the only
species. Later, this genus was expanded to include Spondias
purpurea L., Spondias cytherea Sonn. and Spondias pinnata
(L.f.) Kurz. The genus Spondias now has 18 species, of which
nine species occur in Asia and Oceania (Pacific Ocean), and
nine in the neotropics, including a species introduced from
Asia and a new species, called Spondias testudinis Mitch. and
Daly, found in the south-west Amazon.
TAXONOMY AND NOMENCLATURE
Yellow mombin is a woody fruit tree of tropical
climates, still under domestication. It is an erect, tall tree (up
to 30 m) ramified at the top, with deciduous leaves; a trunk
DESCRIPTION
Spondias
covered with thick, rough bark bearing, in young trees, many
blunt-pointed spines or knobs up to 2 cm long; and a wide,
attractive and impressive canopy in the flowering or fruiting
seasons. The most variable characteristics are: division of the
leaves – simple, pinnate or bipinnate; margins of the leaflets –
entire or crenate; intra-marginal leaf vein – present or absent;
inflorescence – precocious or not, terminal and composite or
lateral and almost simple; number of carpels – from one to
four or five; and endocarp shape and structure.
The whitish flowers usually occur in terminal pyramidal
panicles from 20 to 60 cm long. Male, female and hermaphroditic flowers are found in inflorescences of the same plant.
The flower has a calyx about 5 mm wide; a round receptacle
1–4 mm long; five sepals; five petals; ten stamens, five of
which are inserted in a disc and alternate with the petals and
five epipetalous. The number of flowers per panicle may reach
as high as 2000 (Plate 9), and the number of fruit per cluster is
highly variable both within and among plants.
The ovoid to oblong fruit hang in branched terminal
clusters of a dozen or more (Plate 10). Each fruit is 3–4 cm
long and up to 2.5 cm wide, golden yellow with a thin, tough
skin. The medium-yellow translucent pulp is very juicy and
fibrous. The somewhat musky acid flesh clings to the white
fibrous or ‘corky’ stone.
The species occur
in Asia, Oceania and tropical America, and the centres of
diversity are the Atlantic Forest and West Amazon in the State
of Acre, Brazil, and the adjoining areas of Peru and Bolivia. In
Brazil, the trees are found either isolated or in groups, in the
moist topical forests of the Amazon, and the Atlantic Forest.
In the humid zones of the north-east Atlantic states, it is
found mainly along the coast and lowland. The trees are found
in the wild, maintained or in domestic yards.
It is adapted to grow in humid and arid areas and even in
warm subtropical areas with no frost. It does not grow
satisfactorily above 1000 m. In tropical America, the tree
grows in areas with average annual rainfall between 1000 and
2000 mm, even though there is usually a dry season of up to 5
months. The soil types in these areas are oxisols, ultisols and
inceptisols with a pH from as low as 5.0 to above 7.0. The
species tolerates soils with a moderately low nutrient status.
ECOLOGY AND CLIMATIC REQUIREMENTS
33
around 80 days after anthesis and the fruit abscise when fully
ripe. Ripening is associated with a decline in skin chlorophyll
and acidity and increase in carotenoids and soluble solids
(Costa et al., 1998). Changes in acidity and soluble solids are
more pronounced during the last 15 days of maturation. Fruit
ripen over 4 days from the light green maturity stage.
Horticulture
PROPAGATION AND
ROOTSTOCKS Yellow
mombin are
propagated by seeds, or asexually by cuttings, grafting or in
vitro. The fruit endocarp or stone, contains from none to five
seeds in each locule and germination is low and slow. Physical
and chemical scarification of seeds increases germination
percentage. High variation is found in seed-propagated trees
in height, architecture and shape of canopy, physical-chemical
characteristics of fruit and leaves, length of the juvenile phase,
and variation in phenological phases.
Vegetative propagation can be done using cuttings from
stem or roots, air layering and grafting. Cleft and side-cleft
grafts onto rootstocks of Spondias tuberosa, S. cytherea or S.
mombin are used. A high percentage of these grafts are formed
within 60 days of grafting.
PRUNING AND TRAINING Grafted plants have high growth
rates with single-stem trunks that tend to form a high canopy,
branching at the top like seed-propagated trees. Pruning and
training are required during the first year to reduce height and
develop a more ramified canopy.
Fertilizer requirements have
not yet been determined experimentally and adaptations of
recommendations for other perennial fruit trees that grow in
similar conditions are used. Campbell and Sauls (1991)
suggest that under Florida conditions fertilizer should be
applied every 2–3 months during the first year, beginning with
100 g and increasing to 450 g of 6–6-6–3 or similar analysis.
Thereafter, three to four applications per year are sufficient, in
amounts proportional to the increasing size of the tree,
roughly a 450 g application/year of tree age. Bearing trees can
receive 8–3-9–5 or similar analysis fertilizer at the same rates
three to four times a year.
NUTRITION AND FERTILIZATION
REPRODUCTIVE
BIOLOGY The
reproductive system is
polygamous-dioic or monoic and self-incompatible. The yellow
mombin tree bears hermaphroditic and strongly protandrous
flowers, that is the ovary has not fully developed before pollen
release. This sequence leads to cross-pollination and the
undesirable variability found in orchards propagated from seed.
Flower panicles emerge after new vegetative growth which
occurs following rain. In Costa Rica, the tree blooms in
November/December and again in March while in Jamaica
flowering occurs in April, May and June. In the north-east of
Brazil it blooms only once, in the dry season, usually in
August/September and fruit ripen in the rainy season
(January/February).
In Mexico, trees start to produce fruit by the age of 5 years.
POSTHARVEST HANDLING AND STORAGE Fruit are harvested
after the start of chlorophyll breakdown, when the fruit turns
from light green to yellow. If harvested before this turning
stage the fruit will soften and change colour, but there will be
little changes in acidity, soluble solids and starch contents and
the final quality will be poor.
Harvested fruit are transported in 20 kg containers to retail
stores. It is also displayed in polystyrene trays containing
around 200–300 g fruit wrapped with usually 12 m
polyvinylchloride (PVC) plastic film.
Storage life at 23–25°C is about 4 days for fruit at the lightgreen stage and less than 2 days for ripe fruit. Under
refrigeration at 9–10°C the storage life can be increased to 10
days. At low temperature the fruit develops chilling injury.
FRUIT DEVELOPMENT Fruit growth is sigmoid and it takes
3–4 months to develop from anthesis. Maturation starts
PESTS AND DISEASES Commercial cultivation is still not
significant and only the major pests and diseases have been
34
Anacardiaceae
recorded. The economic importance of these pests and
diseases is unknown. In some cases, pests or pathogens only
attack parts of the plant, causing superficial damage.
Fruit flies (Anastrepha spp.) attack the fruit and do reach a
level of economic loss in low density planting. The leaves are
attacked by the sauva ant (Atta) and by Stiphra robusta Leitão.
The terminal branches are attacked by larvae and the seeds
and endocarps are damaged by weevils.
The major pathogens include Glomerella cingulata, which
causes anthracnose in leaves, inflorescences and fruit.
Sphaceloma spondiadis causes round rough-textured lesions on
leaflets and fruit characterized by cream-coloured centres and
light brown borders. Botryosphaeria rhodina causes resinosis,
with the development of dark cankers that are sometimes
cracked, and abundant gum exudation, and when the lesion
surrounds the trunk or branch it causes yellowing, wilting or
death of the branch or the whole plant. Cercosporiosis disease
caused by Mycosphaerella mombin affects the leaves and begins
with small, round pit spots that become darker and coalesce,
causing yellowing and leaflet fall. Nematodes (Meloidogyne
spp.) attack both adult plants and plantlets. The wood is easily
attacked by termites.
There are no reports of
clones or varieties of superior quality that might be recommended for cultivation. EMBRAPA Tropical Agroindustry in
Brazil is evaluating the behaviour of a number of clones with
promising results.
Heloisa Almeida Cunha Filgueiras and
Francisco Xavier de Souza
MAIN CULTIVARS AND BREEDING
Literature cited and further reading
Abo, K.A., Ogunleye, V.O. and Ashidi, J.S. (1999) Antimicrobial
potential of Spondias mombin L., Croton zambesicus and
Zygotritonia crocea. Phytotherapy Research 13, 494–497.
Airy Shaw, H.K. and Forman, L.L. (1967) The genus Spondias L.
(Anacardiaceae) in tropical Asia. Kew Bulletin 21, 1–20.
Ayoka, A.O., Akomolafe, R.O., Iwalewa, E.O. and Ukponmwan, O.E.
(2005) Studies on the anxyolitic effect of Spondias mombin L.
extracts. African Journal of Traditional, Complementary and
Alternative Medicines 2, 153–165.
Barroso, G.M., Morim, M.P., Peixoto, A.L. and Ichaso, C.L.F. (1999)
Frutos e Sementes: Morfologia Aplicada à Sistemática de
Dicotiledôneas. Universidade Federal de Viçosa, Viçosa, pp. 1–433.
Bora, P.S., Narain, N., Holochuh, H.J. and Vasconcelos, M.A. (1991)
Changes in the physical and chemical composition during
maturation of yellow mombin (Spondias mombin) fruit. Food
Chemistry 41, 341–348.
Campbell, C.W. and Sauls, J.W. (1991) Spondias in Florida. Fruit
Crops Fact Sheet FC-63. University of Florida, Florida, pp. 1–3.
Carballo, A., Carballo, B. and Rodriguez-Acosta, A. (2004)
Preliminary assessment of medicinal plants used as antimalarials
in the Southern Venezuelan Amazon. Revista Brasileira de
Medicina Tropical 37, 186–188.
Corthout, J., Pieters, L., Claeys, M., Vanden Berghe, D.A. and
Vlietinck, A.J. (1991) Antiviral ellagitannins from Spondias
mombin. Phytochemistry 30, 1129–1130.
Costa, N.P. da, Filgueiras, H.A.C., Alves, R.E., da Silva, A.Q. and de
Oliveira, A.C. (1998) Development and maturation of yellow
mombin (Spondias mombin L.) in Northeast Brazil. Proceedings of
the InterAmerican Society for Tropical Horticulture 42, 301–306.
Filgueiras, H.A.C., Moura, C.F.H. and Alves, R.E. (2000) Cajá
(Spondias mombin). In: Alves, R.E., Filgueiras, H.A.C. and
Moura, C.F.H. (eds) Caracterização de Frutas Nativas da América
Latina. São Paulo State, Jaboticabal, Brazil, pp. 19–22.
Filgueiras, H.A.C., Alves, R.E., de Oliveira, A.C., Moura, C.F.H. and
Arayo, N.C.C. (2001) Quality of fruits native to Latin America for
industry: jobo (Spondias mombin L.). Proceedings of the
InterAmerican Society for Tropical Horticulture 43, 72–76.
Francis, J.K. (1992) Spondias mombin L. Hogplum. SO-ITF-SM-51.
US Department of Agriculture, Forest Service, Southern Forest
Experiment Station, New Orleans, Louisiana, pp. 1–4. Available in
Spanish at: http://www.fs.fed.us./global/iitf/spondiasmombin.pdf
(accessed 25 June 2005).
Freire, F.C.O. and Cardoso, J.E. (1997) Doenças de Spondias –
cajarana (S. cytherea Sonn.), cajazeira (S. mombin L.) ciriguela (S.
purpurea L.), umbu (S. tuberosas A. Cam.) e umbuguela (Spondia
spp.) no Brasil. Agrotrópica, Itabuna 9, 75–82.
Joas, J. (1982) Mombins: interesting technological possiblities. Fruits
37, 727–729.
Leon, J. and Shaw, P.E. (1990) Spondias: the red mombin and related
fruits. In: Nagy, S., Shaw, P.E. and Wardonski, F.W. (eds) Fruits of
Tropical and Subtropical Origin: Composition, Properties and Uses.
Science Sourse Inc., Lake Alfred, Florida, pp. 117–126.
Leung, W.T.W. and Flores, M. (1961) Food Composition Table for Use
in Latin America. Nutritional Institutes of Health, Bethesda,
Maryland.
Lorenzi, H. (1992) Árvores Brasileiras: Manual de Identificação e
Cultivo de Plantas Arbóreas Nativas do Brasil. Plantarum de
Estudios da Flora, Nova Odessa, São Paulo, Brazil, 370 pp.
Mitchell, J.D. and Daly, D.C. (1995) Revisão das espécies neotropicais
de Spondias (Anacardiaceae). In: Proceedings of the 46th National
Botanics Congress. Universidade de São Paulo/SBB, São Paulo,
p. 207.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resources
Systems Inc., Winterville, North Carolina, pp. 245–257.
Oliveira, A.C. de, Filgueiras, H.A.C., Alves, R.E. and De Sousa, R.P.
(1998) Room temperature storage of yellow mombin (Spondias
mombin L.) in four stages of maturity. Proceedings of the
InterAmerican Society for Tropical Horticulture 42, 307–312.
Sacramento, C.K. do and Souza, F.X. de (2000) Cajá (Spondias
mombin L.). Série Frutas Nativas 4. Fundacao de Estudos e
Pesquisas em Agronomia, Medicina Veterinaria e Zootecnia
(FUNEP), Jaboticabal, São Paulo, Brazil, 42 pp.
Souza, F.X. de (2000) Effect of rootstock and grafting methods on
the development of Spondias mombin L. plants. Revista Brasileira
de Fruticultura 22, 286–290.
Souza, F.X. de, Innecco, R. and Araújo, C.A.T. (1999) Grafting
Methods Recommended for the Production of Yellow Mombin and
Other Fruit Trees of the Genus Spondias. Comunicado Técnico 37.
EMBRAPA-CNPAT, Fortaleza, Ceara, Brazil, 8 pp.
Spondias purpurea
red mombin
The red mombin, Spondias purpurea L. (Anacardiaceae), is an
aromatic fruit, highly valued for local people in the tropics since
old times. The species is probably native to southern Mexico and
Central America, where wild populations are still found. Early
Spanish navigators took the red mombin to the Philippines. In
Jalisco (Mexico) during the 19th century, red mombin was one
of the most important fruit crops. Nowadays, fruit can be readily
Spondias
found in local markets. In Ecuador, it is commercialized and
found in some supermarkets of the big cities.
The most widespread vernacular name in South America
and Philippines is ciruelo, although in Central America and
Mexico it is also widely known as jocote, and in some parts of
South America as ovo. Many orthographic and phonetic
variants of these common names have been recorded.
World production
The red mombin is the most cultivated species in the genus
Spondias. It has been cultivated widely throughout the
neotropics, from central Mexico and the West Indies, to Peru
and Brazil. It is naturalized in the Antilles from cultivation,
including the Bahamas, and has also been cultivated in
Florida, USA (Popenoe, 1979). There are no data for world
production. In Ecuador, yields of more than 4500 t/year were
reported within the period 1987 and 1990–1992. In an
Ecuadorean Andean dry area, the average yield ranged
between 2250 to 5000 kg/ha from dry season varieties (Macía
and Barfod, 2000).
Uses and nutritional composition
The tree is commonly planted as a living fence. The ripe red
mombin fruit are mainly eaten fresh, but sometimes are
harvested green and eaten with salt as a snack. In Mexico, ripe
fruit are sometimes boiled in water with or without salt and
only eaten dried afterwards. In Florida, dried slices of ripe
fruit have been occasionally commercialized. The soft exocarp
is easily injured and so the mesocarp is processed into
marmalade, juice, wine and liquor. The pulp is used as a
flavouring for ice cream.
The fruit of red mombin have a good calorific density
(Table A.16) due to the high concentration of total
carbohydrates (19.1%). Fructose, glucose and sucrose together
account for 65% of the total soluble solids. It is a moderate
source of potassium and starch, and a good source of vitamin
C (Koziol and Macía, 1998). The main flavour compound is 2hexenal.
35
Table A.16. Chemical composition of red mombin (Spondis purpurea) per
100 g (Source: Koziol and Macía, 1998).
Proximate
Moisture
Food energy (kcal)
Protein
Fat
Total carbohydrates
Fibre
Minerals (ash)
Minerals
Calcium
Iron
Phosphorus
Sodium
Potassium
Zinc
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Carotene c
Pulp composition
pH
Total soluble solids
Starch
Pectins
Fructose
Glucose
Reducing sugars
Sucrose
Citric acid
Malic acid
Oxalic acid
Tartaric acid
Range
Average
g
g
65.0–87.0
61–86
0.1–1.0
0.03–0.8
16.0–22.3
0.2–0.7
0.3–1.1
77.6
74
0.7
0.2
19.1
0.5
0.7
mg
mg
6–25
0.09–1.22
32–56
2–9
230–270
–
17
0.72
42
6
250
20
mg
mg
26–73
0.033–0.110
0.014–0.080
0.4–1.8
0.004–0.225
49
0.084
0.040
1.0
0.119
units
%
g
g
g
g
g
g
g
mg
mg
mg
mg
5.97–7.21
3.29
18
2.47
0.22
2.53
2.00
8.08
6.59
30
110
30
20
Botany
TAXONOMY AND NOMENCLATURE Spondias is a pantropical
genus composed of approximately 18 species, with the centre
of diversity in South-east Asia. In tropical America, there are
nine species. Some of the species (Spondias dulcis and Spondias
mombin) have been introduced throughout the tropics to the
drier areas in Africa, Asia and the South Pacific for their edible
fruit. Spondias purpurea is a well-defined and taxonomically
separate species (Barfod, 1987). No taxonomic treatment is
available of the varieties nor of the species’ genetic diversity.
Spondias purpurea is a small deciduous tree,
3–15 m high, with grey and usually smooth bark. A rather
thick and transparent exudate exudes from cuts and bruises.
The imparipinnate leaves are 6–28 cm long and 5–27-foliolate
with a 15–20 cm rachis. The leaflets are usually 3–6 cm long
and 1–2.5 cm wide, elliptic to oblanceolate. The axillary
inflorescences are 1–10 cm long, few-flowered, usually
produced at older and defoliate nodes. The petals are usually
DESCRIPTION
red to purple, 2.5–3.5 mm long at anthesis. The fruit is a
drupe, which is oblong to obovoid or subglobose 1.5–4.5 cm
long and 1–3.5 cm wide. When ripe, the fruit is usually red
but sometimes yellowish or orange. The mesocarp is fleshy and
juicy, and the endocarp is 1.5–3.5 cm long.
ECOLOGY AND CLIMATIC REQUIREMENTS Natural populations
of red mombin in Mexico and Central America are found in
both dry and wet areas, including a wide range of semideciduous forests. It has been cultivated from 0 to 2000 m
elevation with an average annual precipitation varying from
300 to 1800 mm. The tree is able to grow normally on rocky
substrates, slopes or different soil types including those of
little agricultural value due to a wide physiological and
anatomical plasticity (Pimenta-Barrios and RamírezHernández, 2003). A mycorrizal symbiosis can be associated
with the root and this favours plant growth by promoting
phosphorus absorption.
36
Anacardiaceae
REPRODUCTIVE BIOLOGY Flowering time varies with climate,
but usually occurs during the dry season when trees are
defoliated or just as the young new leaves emerge. New
vegetative shoots are produced and may constitute the major
part of the potentially flower-bearing ramets. In areas with
more year-round precipitation, flowering may occur nearly all
year round. In dry areas, depending upon the tree’s
phenology, flowering can be controlled by carefully planned
irrigation (Macía and Barfod, 2000). If trees are treated with
12% urea to induce defoliation, flowering is advanced by
30–40 days (Almaguer-Vargas et al., 1991). There is no
information on pollination.
FRUIT DEVELOPMENT The fruit develop parthenocarpically and
take about 115 days from anthesis to the start of ripening (De
Melo and Pereira, 2001). The new vegetative shoots and the fruit
mature at the same time. Fruit usually ripen during the dry
season, so that a high number of hours of exposure to full
sunlight produces more sugars and hence better fruit quality.
Fruit harvested at the skin-breaker stage (Plate 11) do not
develop the characteristic full red colour and the fruit after
harvest need to be sorted by colour (Plate 12). The best storage
temperature is about 15°C for a maximum of 10 days. Harvested
fruit held at 25°C ripen in 3–5 days (Manzano, 1998).
Horticulture
PROPAGATION Since the tree only infrequently produces
viable seeds (Juliano, 1932), propagation is by vegetative
cuttings. After the harvest, when the leaves have been shed and
flowering has just started, cuttings 1–2.5 m long are obtained
from the best clones. The cuttings are left in the shade for
about a week and then planted 3–7 m apart, at a depth of
30–40 cm. The soil is irrigated after transplanting to stimulate
rooting. Grafting on other rootstocks such as Spondias pinnata
is possible. In Mexico and Ecuador, pruning is mainly done to
keep the trees short and to facilitate harvesting from the
ground. No fertilization requirements have been reported.
DISEASES, PESTS AND WEEDS
No important diseases and
pests have been recorded. Fruit flies may cause serious damage
to ripe fruit. In dry areas just before flowering, the branches
are cleaned to remove epiphytes. In Ecuador orchards have
been fumigated by smoke from fires along the margins of the
orchard, but occasionally chemical insecticides have been
used. Weeding is necessary one or two times a year in dry areas
while three or four times a year in wet regions.
MAIN CULTIVARS AND BREEDING There are two main groups
of non-commercial varieties: dry-season and wet-season red
mombins. Fruit of the first variety are smaller, albeit sweeter
and slightly less acidic than those of the second variety
(Macía, 1997). The fruit of the first variety are 2.7–3.9 cm
long by 1.9–3 cm wide, compared to the larger fruit of the
second variety, which are 3.1–4.5 cm long by 2.4–3.5 cm wide.
Often cultivars bear the name of the area of origin or fruit
characteristics. Dry-season mombin varieties include
‘Tronador’, ‘Crillo’, ‘Nica’, ‘Morado’, and ‘Rojo Ácido’, while
the wet-season mombins are ‘Corona’, ‘Petapa’ and ‘Cabeza de
loro’ (Leon and Shaw, 1990).
Manuel J. Macía
Literature cited and further reading
Almaguer-Vargas, G., Espinoza-Espinoza, J.R., Martines-Bravo, A.
and Amador-Gomes, J. (1991) Efecto de la defoliación química en
el adelanto de la cosecha de guayabo (Psidium guajava L.) y
ciruelo (Spondias purpurea L.). Proceedings of the InterAmerican
Society for Tropical Horticulture 35, 71–75.
Barfod, A. (1987) Anacardiaceae. In: Harling, G. and Andersson, L.
(eds) Flora of Ecuador 30. Arlöv, Sweden, pp. 9–49.
Cuevas, J.A. (1992) Jocote, ciruelo (Spondias purpurea). In:
Hernández-Bermejo, J.E. and Leon, J. (eds) Cultivos Marginados:
Otra Perspectiva de 1492. FAO, Rome, Italy, pp. 109–113.
De Melo, S. and Pereira, L. (2001) Carbohydrate-related changes in
red mombin (Spondias purpurea L.) fruit. Proceedings of the
InterAmerican Society for Tropical Horticulture 45, 38–41.
De Sousa, R., Filgueiras, H.A., Alves, R., Alves, J. and Cordeiro, A.
(1998) Identification of the optimum harvest stage for red
mombin (Spondias purpurea L.). Proceedings of the InterAmerican
Society for Tropical Horticulture 42, 319–324.
Juliano, J.B. (1932) The cause of sterility of Spondias purpurea Linn.
Philippine Agriculturist 21, 15–24.
Kostermans, A.J.G.H. (1991) Kedondong, ambarella, amra: the
Spondiadeae (Anacardiaceae) in Asia and the Pacific area (with
some notes on introduced American species). Foundation Useful
Plants of Asia 1, 1–100.
Koziol, M.J. and Macía, M.J. (1998) Chemical composition,
nutritional evaluation, and economic prospects of Spondias
purpurea (Anacardiaceae). Economic Botany 52, 373–380.
Leon, J. and Shaw, P.E. (1990) Spondias: the red mombin and related
fruits. In: Nagy, S., Shaw, P.E. and Wardowsky, F.W. (eds) Fruits of
Tropical and Subtropical Regions. Florida Science Source (FSS),
Lake Alfred, Florida, pp. 116–126.
Macía, M.J. (1997) El ‘ovo’ (Spondias purpurea L., Anacardiaceae) un
árbol frutal con posibilidades socioeconómicas en Ecuador. In:
Rios, M. and Pedersen, H.B. (eds) Uso y Manejo de Recursos
Vegetales. Memorias del Segundo Simposio Ecuatoriano de
Etnobotánica y Botánica Económica. Ediciones Abya-Yala, Quito,
Ecuador, pp. 271–281.
Macía, M.J. and Barfod, A.S. (2000) Economic botany of Spondias
purpurea (Anacardiaceae) in Ecuador. Economic Botany 54,
449–458.
Manzano, J.E. (1998) Comportamiento de frutos de ‘ciruela huesito’
(Spondias purpurea L.) almacenadas a diferentes temperaturas.
Proceedings of the InterAmerican Society for Tropical Horticulture
42, 313–318.
Mitchell, J.D. (2001) Anacardiaceae. In: Stevens, W.D., Ulloa, C.,
Pool, A. and Montiel, O.M. (eds) Flora de Nicaragua. Missouri
Botanical Garden Press, Saint Louis, Missouri, pp. 83–93.
Pimenta-Barrios, E. and Ramírez-Hernández, B.C. (2003)
Phenology, growth, and response to light of ciruela mexicana
(Spondias purpurea L., Anacardiaceae). Economic Botany 57,
481–490.
Popenoe, W. (1979) The genus Spondias in Florida. Proceedings of the
Florida State Horticultural Society 92, 277–279.
Tapirira guianensis
wild mombin
Wild mombin, Tapirira guianensis Aublet. (Anacardiaceae), is a
tropical species found from Mexico south to Amazonia, to
Peru and Bolivia and Paraguay. In Brazil, it is found on the
Annona
Atlantic coast from Rio in the south to Ceara and Amapa in
the north, and into parts of Guyana, Venezuela, Colombia and
Panama. Common names include tapiriri, tapirirá, cedro-y,
cupiúva, fruta de pombo, guapiruva, pau pombo, peito de
pomba and jobo in Brazil; duka and waramia in Guyana; wetioedoe and ook doka in southern Mexico; piojo and caobina in
Honduras; isaparitsi, huira caspi and huira caspi colorado in
Peru; fresno, cedrillo and cedro macho in Colombia; cedrillo
and capuli in Ecuador; and jobillo and cedro nogal in
Venezuela. In English it is wild mombin and white mombin,
and in French gommier viande biche, mombin sauvage and
tapirier de la Guyane.
Uses and nutritional composition
The fruit is edible and can be dried.
37
ANNONACEAE
Annona cherimola
cherimoya
Cherimoya, Annona cherimola P. Mill. (Annonaceae), is a
subtropical fruit with white, delicate, sweet flesh containing
many large seeds. It is thought to have originated in the
highland Andes’ valleys between Peru and Ecuador at
1500–2000 m. Its antiquity is attested to by ancient artefacts
shaped in the form of the fruit in Peru. Distribution through
Central America and Mexico probably occurred at an early
date as it has become naturalized in the cool highland areas.
Distribution continued from Mexico to the Caribbean islands,
then to the African coast and the Mediterranean. Introduction
to Africa and the Far East is attributed to early Spanish
navigators. Annona cherimola is known as cherimoya,
chirimoya (Spanish), cherimolier (French), anona (Mexico)
and noina ostrelia (Thailand).
Botany
The genus Tapirira is
pantropical with about 15 species that are found in Asia, Africa
and the Americas. Only one species has edible fruit T.
guianensis Aublet and its synonym is T. miryantha Tr. and Pl.
World production and yield
TAXONOMY AND NOMENCLATURE
The tree can reach 30 m in primary forests but
under poor conditions is much less. The trunk has some
buttresses and the bark is clear grey with some reddish brown,
smooth when young becoming irregular with sinuous cracks.
The sapwood is white and exudes a resin that has a sweet
balsamic odour. The leaves are alternate without stipules with
a variable number of leaflets (5–15). The leaflets are oblong (10
⫻ 3.5 cm). The inflorescence is a long terminal panicle,
covered with very small yellowish-white hermaphroditic or
more often unisexual flowers. The five oblong petals are 2 mm
long. The ovoid ovary (1.5 mm) has one ovary with five
disjoined styles (0.3 mm). The fruit is a small drupe (10 ⫻
8 mm) with a smooth yellow skin that turns black upon
ripening. The edible mesocarp encloses one seed.
DESCRIPTION
The tree is found in
the dry plains of Brazil in the low ground forests as a small
tree. In moist forest areas it can reach 35 m height and 70 m
diameter. It can be found from 50 to 900 m in elevation. The
tree prefers deep, well-drained soils.
ECOLOGY AND CLIMATIC REQUIREMENTS
In Honduras, flowering is observed
from April to June and fruit mature 5 months later from
August to September.
REPRODUCTIVE BIOLOGY
Horticulture
The seeds are recalcitrant and need to be planted soon after
removal from the fruit. Insects and disease have not been
reported.
Robert E. Paull
Literature cited and further reading
Gazel, M. (2000) Tapirira guianensis Aublet. Available at:
http://kourou.cirad.fr/silvolab/implantations/campus/denti
er/tapiriraguinensis.html (accessed 5 May 2005).
The cherimoya is considered the best of the annonas and is
cultivated in subtropical regions and in the tropical highlands.
In most areas, it is grown as a backyard tree or as part of a
subsistence farming system at appropriate elevations.
Commercial production occurs in Spain, Bolivia, Chile, Peru
and New Zealand. World production in 1998 was estimated at
about 100,000 t (Van Damme and Scheldeman, 1999). Spain
had 3600 ha that yielded 35,000 t, Peru 1800 ha and 15,000 t
and Chile 1200 ha and 12,000 t. Experience has shown that the
Californian coastal regions are more conducive to cherimoya
production, having higher relative humidity (70–80%) in
spring and summer than the interior valleys where the relative
humidity can drop to 40% and below during the hotter part of
the day during summer.
Uses and nutritional composition
Cherimoya is most often consumed as a dessert fruit eaten out
of the hand, or scooped with a spoon. It can be served in fruit
salads, sorbets, custards and pies. The fruit is high in calories
and fibre. Cherimoya is a fair to good source of niacin and
vitamin C (Table A.17). The seeds of the annonas are toxic.
Botany
TAXONOMY AND NOMENCLATURE The genus Annona is the
most important in the Annonaceae, since among its 100 or more
species, seven species and one hybrid are grown commercially.
The Annonaceae (130 genera, 250 spp.) are considered a
‘primitive’ member of the Magnoliidae. Annona cherimola is
closely related to Annona glabra and Annona reticulata, both
morphologically and allozymes (Samuel et al., 1991).
The tree is erect but low branched and
somewhat shrubby or spreading, ranging from 5 to 9 m in
height with young branchlets that are rusty-hairy. The leaves
are briefly deciduous (just before spring flowering), alternate,
two-ranked, with minutely hairy petioles 6–12.5 mm long;
ovate to elliptic or ovate lanceolate, short blunt-pointed at the
apex; slightly hairy on the upper surface, velvety on the
underside; 7.5–15 cm long, 3.8–8.9 cm wide. New buds
DESCRIPTION
38
Annonaceae
Table A.17. Proximate fruit composition of cherimoya which has an edible
flesh to fruit ratio of 65% (Source: Wenkam, 1990).
Proximate
Water
Energy (kcal)
Energy (kJ)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
%
68.7
110
460
1.54
0.13
28.95
–
0.67
mg
9
0.25
–
24
–
–
mg
12.2
0.11
0.11
1.0
–
cannot sprout until the leaves have shed as the leaf bases grow
over the axillary buds as in sweetsop.
Flowers are fragrant, solitary or in groups of two or three,
on short hairy stalks along the branches, having three outer,
greenish, fleshy, oblong, down petals to 3 cm long and three
smaller, pinkish inner petals. A compound fruit, the cherimoya
is conical or somewhat heart-shaped, 10–20 cm long and up to
10 cm in width, weighing on average 150–500 g but extra large
specimens may weight 2.7 kg or more (see Fig. A.5 in entry for
Annona muricata). The skin, thin or thick, may be smooth with
fingerprint-like markings or covered with conical or rounded
protuberances. The fruit is easily broken or cut open, exposing
the snow-white, juicy flesh, of pleasing aroma and subacid
flavour, and containing numerous hard, brown or black, beanlike, glossy seeds, 1.25–2 cm long.
AND CLIMATIC REQUIREMENTS Cherimoya is
capable of growing in a wide range of soil types from sandy
soil to clay loams. Higher yields occur on well-drained sandy
to sandy loam soils. Drainage is essential to avoid root rot
diseases, hence the interest in A. glabra as a rootstock related
to its tolerance of wet soils.
Cherimoya benefits from uniform soil moisture for good
production with extremes of moisture lowering production.
Rainfall and high humidity during peak flowering season
greatly enhance fruit production by preventing desiccation of
stigmas, prolonging their receptive period and increasing fruit
set and early fruit growth.
Temperature is the limiting factor, with frost killing young
trees, but older trees show some tolerance. Cherimoya
(7–18°C, mean minimum) is more tolerant to low
temperatures than the least tolerant soursop (15–25°C mean
ECOLOGY
minimum). Cherimoya requires chilling periods and does not
do as well under lowland conditions (George and Nissen,
1987b) and benefits from dry periods. Cherimoya is
susceptible to high temperatures with a growing temperature
of 21–30°C (George and Nissen, 1986b). Poor pollination is a
frequent problem and occurs under high temperature (30°C)
and low humidity (30% relative humidity (RH)). Lower
temperature (25°C) and high humidity (80% RH) greatly
improves pollination. Hand pollination is recommended for
cherimoya to achieve more uniform fruit shape. No
photoperiod responses have been reported.
The softwood of the trees makes them susceptible to wind
damage and limb breakage. Tree shaking may also be partially
responsible for collar rot organism penetration. The fruit skin
is easily damaged by rubbing and exposure to drying winds
(Marler et al., 1994). Productivity can be improved by
windbreaks and under-tree sprinkling to raise the RH above
60%.
REPRODUCTIVE BIOLOGY The flowers are hermaphrodite and
are produced singly or in small clusters on the current
season’s growth, although flowers arising from old wood are
common. New flowers continue to appear towards the apex of
the shoot as flowers produced earlier at the basal portions
mature.
Reducing irrigation in late winter to force cherimoya trees
into dormancy for 1–2 months in spring is recommended in
California to induce flowering. Cherimoya generally requires
27–35 days for flower bud development from initiation to
anthesis. Differences in floral behaviour in the various areas
may be attributed to both genetic variability and climatic
differences. Flowering can extend from 3 to 6 months or even
longer, with heavy peaks. Two major flowering periods occur
after periods of vegetative flushes with the second peak
coinciding with the onset of monsoon in India.
The flowers exhibit both dichogomy and a protogynous
nature. This poses a serious problem in obtaining high yields.
Cherimoya flowers are receptive, opening around 7–9 a.m. and
when pollen is shed at 3–4 p.m. if the relative humidity is
above 80% and temperature >22°C. The flower shows
synchrony with that of sweetsop and this together with
complementary functional sexes favours cross-pollination
leading to natural hybridization. This is attested to by the
frequent appearance of hybrid seedlings under the trees of
sweetsop and cherimoya when grown in close proximity.
Nitidulid beetles (Carpophilus and Uroporus spp.) are the
important pollinators of annona flowers with wind and selfpollination being low (1.5%).
Pollen grains appearing early in a flowering season have
thick walls, are high in starch, germinate poorly, and give poor
fruit set (Saavedra, 1997). Pollen of later flowers show a high
proportion of individual pollen grains without starch grains
that germinate well. Hand pollination is frequently practised
to ensure pollination and good fruit shape. Pollen must be
collected in the evening from fully open flowers, when the sacs
have turned from white to cream. The flowers are held in a
paper bag, not a closed container, and should discharge that
afternoon. The flowers are shaken over a shallow tray or paper
to collect the pollen, which is transferred to a small container
and held in the refrigerator for use the next morning. Pollen
Annona
from 20–30 flowers can give enough pollen to pollinate 50–60
flowers. Hand pollination has shown some variable results and
is less successful on very humid, overcast days and on young
vigorous trees. About 150 flowers can be pollinated in an hour
and a success rate of 80–100% can be achieved.
DEVELOPMENT Fruit growth shows the typical
sigmoidal curve with maturation occurring in 16–24 weeks
depending upon growing conditions. Low humidity (<60%
RH) and temperature (<13°C) near fruit maturity can increase
the severity of fruit skin russetting as well as delaying fruit
maturation.
Fruit is harvested when fully mature and firm. The skin
colour changes as the fruit approaches maturity. Skin of
immature cherimoya is greyish green but turns to yellowgreen at maturity. Determining harvest time by dating floral
anthesis is impractical as flowering occurs over many months.
If a rigid hand pollination protocol is used with removal of
naturally pollinated fruit, days from anthesis can be used.
FRUIT
Horticulture
Annona spp. are usually propagated by seed. A
rapid loss of seed viability occurs (6 months) and seeds should
be planted as soon as possible after removal from the fruit
(George and Nissen, 1987c). Seeds can take up to 30 days to
germinate and gibberellic acid (10,000 ppm) can significantly
increase germination and enhance seedling growth. Seedlings
require at least 3–4 years to bear fruit (Sanewski, 1991).
Clonal propagation by cuttings, layering, inarching, grafting
and budding have been tried. Inconsistent results are obtained
with cherimoya when 1-year cuttings are treated with rooting
hormones. Cherimoya is not easily propagated by air layering
(less than 5% success). A modification where the new shoot is
clamped and only the shoot tip is exposed is successful.
Inarching of A. cherimola to A. reticulata rootstock has been
successful with only A. glabra giving less than 70% success.
Although inarching has given good results, it is time
consuming and costly for large-scale propagation (George and
Nissen, 1987c). Grafting is superior to budding in percentage
takes and subsequent growth, with side whip graft and cleft
graft techniques giving the best results (Duarte et al., 1974).
The branches should be defoliated 1–2 weeks before scionwood is cut to induce bud swelling. T-budding and chip
budding methods are successful.
PROPAGATION
There are considerable graft incompatibilities
among Annona and Rollinia species and types. Cherimoya has
been found to be a vigorous rootstock for ‘Pink’s Mammoth’
(atemoya) (Sanewski, 1991). This is complicated by cultivar
differences in compatibility with common rootstocks.
Cherimoya has been successfully grafted onto A. retriculata
and Annona squamosa rootstocks.
ROOTSTOCKS
Training of trees should begin in
the nursery and pruning should continue after transplanting.
It is desirable to train the tree to a single trunk up to a height
of about 90 cm and then headed back to produce lateral
branches. The lateral branches should be spaced 15–25 cm
above each other and be allowed to grow in different directions
PRUNING AND TRAINING
39
to develop a good scaffold. After about 2 m, they could be left
to natural growth. Pruning is carried out when the trees are
dormant and in heavy trees involves removal of lower limbs
touching the ground and branches in the centre that may be
rubbing against each other. The objective is to allow sunlight
access to the centre of the tree (George and Nissen, 1986a).
All lateral buds can have up to two vegetative buds and
three flower buds (Schroeder, 1992). The lateral buds of
cherimoya are normally ‘buried’ (subpetiolar) in the base of
the swollen leaf petiole. Leaf shed must occur prior to the
elongation of ‘buried’ buds (George and Nissen, 1987a).
Removal of leaves mechanically by stripping or chemically
with urea or ethephon releases these buds. Adventitious buds
can arise at any point on a trunk.
NUTRITION AND FERTILIZATION The Annona spp. have an
indeterminate growth habit (axillary flowering) and applying
nitrogen in somewhat excessive amounts does not interfere
greatly with floral initiation, as is the case with plants having
determinate growth habit.
POSTHARVEST HANDLING AND STORAGE Fruit size, shape and
skin colour along with the absence of defects and decay are
used in grading fruit. Fruit are very susceptible to mechanical
injury. Sugar levels can vary from 14 to 18%, with moderate
acid levels. Mature fruit are firm that become very soft during
ripening. Skin changes in colour from dark to light green or a
greenish yellow and is associated with an increased surface
smoothness. Fruit are harvested when mature and allowed to
ripen during marketing and retailing.
There are no US or international standards. Fruit is usually
packed in a single-layer pack in a fibreboard carton with foam
sleeves or paper wrapping to avoid bruising. There are two
carton sizes, 4 and 8 kg, with 12 or 24 fruit per carton,
respectively. Fruit weight varies from 250 to 600 g. The fruit
needs to be pre-cooled as soon as possible after harvest to
about 12–15°C; room cooling or forced air is most often used.
Fruit are stored at 10–13°C and 90–95% RH for 2–3 weeks; if
held at 20°C fruit last only 3–4 days. Storage is limited by skin
darkening, dessication and disease associated with chilling
injury. Ripe soft fruit can be held at 0–5°C. However, fruit
that is not fully ripe is chilling sensitive especially below 10°C,
the extent of injury depending upon duration. Symptoms
include skin darkening and a failure to fully soften and to
develop full flavour.
Controlled-atmosphere storage in 5% oxygen for 30 days at
10°C leads to fruit that ripen in 11 days after removal to air
storage at 20°C, versus 3 days for fruit held in 20% oxygen.
The addition of carbon dioxide at 3% or 6% can also extend
storage life beyond just air storage. Oxygen levels less than 1%
lead to fruit having an off-flavour.
DISEASES, PESTS AND WEEDS Black canker (Phomopsis
anonacearum) and diplodia rot (Botryodiplodia theobromae)
occur mostly on neglected trees and cause similar symptoms
of purplish to black lesions resulting in mummified fruit
(Table A.18). Marginal leaf scorch is also caused by P.
anonacearum and B. theobromae and causes twig dieback.
Diplodia rot has darker internal discoloration and deeper,
more extensive corky rot in fruit. Cylindrocladium fruit and
40
Annonaceae
Table A.18. Major diseases of cherimoya.
Common name
Organism
Parts affected and symptoms
Country/region
Anthracnose
Armillaria root rot
Bacterial wilt
Black canker (diplodia rot)
Black canker
Fruit rot
Purple blotch
Rust fungus
Colletotrichum gloeosporioides (Glomerella)
Armillaria leuteobubalina
Pseudomonas solanacearum
Botryodiplodia theobromae
Phomopsis anonacearum
Glioclacium roseum
Phytophthora palmivora
Phakopsora cherimoliae
Flowers, fruit, leaves, dieback, seedling damping off
Roots, base of trees, decline
Tree wilt
Leaf scorch, twig dieback
Leaf scorch, twig dieback
Fruit
Spots on immature fruit, fruit drop, twig dieback
Leaves
Universal
Australia
Australia
Australia
Australia
India
Australia
Florida
leaf spot is caused by a soil-borne fungus, Cylindrocladium
colhounii. It can cause almost total loss of fruit during years of
persistent heavy rains. Symptoms begin with small dark spots,
primarily on the shoulders of the fruit, that spread along the
sides, enlarge, become dry and crack. Infection is skin-deep
but fruit becomes unmarketable.
Some insect pests occur in numerous growing areas (Table
A.19). One of the most serious insect pests in Trinidad is the
cerconota moth (Cerconota anonella) that lays its eggs on
young fruit. The emerging larvae tunnel into the pulp, causing
blackened, necrotic areas. It is common to find every fruit
larger than 7.5 cm infested. Bagging the fruit is sometimes
done. This moth has been reported in the American tropics as
far south as Brazil and is a major limiting factor in Surinam.
The bephrata wasp (Bephrata maculicollis) is widely
distributed throughout the Caribbean and Mexico, Central
and northern South America. This wasp is considered to be
the most important pest in Florida (Campbell, 1985). The
larvae infest the seeds and cause damage to the pulp as they
bore through the flesh to emerge when the fruit matures. The
thecla moth (Thecla ortygnus) is widespread through parts of
the Caribbean and in the American tropics but it is not
considered to be as serious as the cerconota moth and
bephrata wasp. Primary damage is done to the flowers. The
larvae feed on flower parts such as the perianth, stamen and
stigmas with the flowers failing to set fruit.
Mature green annonaceous fruit have been shown to be
rarely infested by the Mediterranean fruit fly (Ceratitis
capitata) and oriental fruit fly (Dacus dorsalis); but are found
on occasions in tree-ripened fruit. In Australia, the
Queensland fruit fly (Dacus tryoni) infests ripening atemoya
fruit. ‘African Pride’ appears more susceptible than ‘Pink’s
Mammoth’. Use of bait sprays and field sanitation are
recommended measures to minimize fruit fly infestation
(Smith, 1991). Fruit bagging also provides protection.
Mealy bugs and various species of scale insects are found
universally and usually become a serious pest on neglected
trees. The former is reported to be a major pest on marketable
fruit in some areas of Australia (Sanewski, 1991). Red spider
mites can become a serious problem in dry areas or during dry
seasons. Heavy infestations have been observed on soursop
flowers and leaves in the Tecoman area of Mexico during the
prevailing dry period with trees showing heavy flower drop.
Problem weeds especially grass and twining weeds should
be controlled before planting by cultivation and herbicides.
Young trees should to be protected from weed competition by
hand weeding, mulching or contact herbicides. The shallow
root system limits the use of cultivation under the tree. A
translocated herbicide may be needed for perennial weeds and
is applied as a spot spray.
The chromosome number of
A. cherimola is 2n = 14. Existing commercial cultivars show
considerable variation in growth, fruit set, fruit size and quality.
No single cultivar has all the desirable characteristics. The
length of the juvenile period varies with earliest production
occurring in 2 years and full production in 5–6 years. This
juvenile period is extremely variable with scions on seedling
rootstocks. The seedling rootstocks are derived from extremely
heterogenous openly pollinated seeds; hence it is difficult to fix
MAIN CULTIVARS AND BREEDING
Table A.19. Major insect pests of cherimoya.
Common name
Organism
Parts affected
Country/region
Banana spotting
Bephrata wasp (soursop wasp)
Caribbean fruit fly
Cerconota moth (soursop moth)
Citrus mealy bug
Coconut scale
Mealy bug
Potato leaf hopper
Queensland fruit fly
Red spider mite
Scale insects
Southern stink bug
Thecla moth
Wasp
Amblypelta lutescens
Bephrata maculicollis
Anastrepha suspensa
Cerconota anonella
Planocuccus citri
Aspidiotus destructor, other genera and species
Dysmicoccus
Empoasca fabae
Dacus tryoni
Several genera, species
Saissetia coffeae
Nezara viridula
Thecla ortygnus
Bephratelloides paraguayensis
Young fruit
Fruit
Fruit
Fruit
Fruit
Leaves, stem
Stem, leaves
Leaves
Fruit
Leaves, flowers
Leaves, stem
Fruit
Flower, young fruit
Fruit
Queensland
Mexico, Americas, Trinidad, Surinam
Caribbean, Mexico
Americans, Trinidad, Surinam
Queensland
Caribbean
Universal
Caribbean
Australia
American tropics
Universal
Caribbean
Americas, Caribbean
Americas, Barbados
Annona
specific characters in a short period. Breeding programmes have
focused on selections from seedling populations. Early maturity,
better fruit appearance, and in the subtropics greater cold
tolerance are the most frequent objectives.
Cherimoya has very few named clonal cultivars (Table A.20).
Most of the plantings have been of seedlings. In California, some
old cultivars of cherimoya include ‘McPherson’, ‘Deliciosa’ and
‘Bays’. Considerable work in Peru has been done on the
development of cultivars that are not known elsewhere. Chile,
Spain and New Zealand grow the cherimoya as it is more
tolerant to cold temperatures with more successful selfpollination than the atemoya. New Zealand’s principal cultivars
are ‘Reretai’, ‘Burton’s Wonder’, and ‘Burton’s Favorite’.
Chilean cultivars ‘Bronceada’ and ‘Concha Lisa’ have performed
well in Australia; ‘Bronceada’ possesses a postharvest coldstorage life of 3 weeks. In Spain, ‘Fino de Jete’ and ‘Campa’ are
the most extensively cultivated due to their superior yield and
quality (Pascual et al., 1993). Isozyme studies indicated that
these two cultivars showed identical banding patterns for 15
enzymes, indicating that they may be the same cultivar. A cluster
analysis of isozyme patterns showed that Spanish cherimoya
cultivars were distinctly different from cultivars in California
(Pascual et al., 1993) and atemoya (Ellstrand and Lee, 1987).
In order to develop cultivars adapted to cooler environments,
Australia has concentrated on self-progenies and interspecific
crosses of A. cherimola with A. reticulata and Annona diversifolia.
Progenies of A. cherimola ⫻ A. reticulata are late maturing,
showing flowering and fruiting characteristics of A. reticulata
that flower in the autumn and mature fruit in late spring. Four
promising selections possessing most of the fruit qualities of
commercial cultivars have been established in various areas for
further evaluation (George et al., 1992).
Robert E. Paull
Literature cited and further reading
Bonaventure, L. (1999) A Cultura da Cherimoia e de Seu Hibrido a
Atemoia. Edit Nobel, São Paulo, Brazil, 182 pp.
Campbell, C.W. (1985) Cultivation of fruit of the Annonaceae in
Florida. Proceedings of the American Society for Horticultural
Science Tropical Region 29, 68–70.
Table A.20. Selected cultivars of cherimoya.
Name
Origin
‘Andrews’
‘Bays’
‘Booth’
‘Bronceada’
‘Burton’s Wonder’
‘Campa’
‘Concha Lisa’
‘Cristalino’
‘E-8’
‘Fino de Jete’
‘Kempsey’
‘Libby’
‘Lisa’
‘Mossman’
‘Negrito’
‘Reretai’
‘White’
Australia
USA – California
USA – California
Chile
New Zealand
Spain
Chile
Spain
Ecuador
Spain
Australia
USA – California
USA – California
Australia
Spain
New Zealand
USA – California
41
Duarte, O. and Escobar, O. (1997) Mejora del cuajado de chirimoya
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Ellstrand, N.C. and Lee, J.M. (1987) Cultivar identification of
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George, A.P. and Nissen, R.J. (1986b) The effects of root temperature
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in subtropical Queensland. Australian Journal of Experimental
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Agriculture and Human Resources (HITAHR), College of
Tropical Agriculture and Human Resources, Hawaii.
Annona muricata
soursop
Soursop, Annona muricata L. (Annonaceae), is the most
tropical member of the genus and produces the largest fruit in
the family and is the only one lending itself to preserving and
processing. It was distributed very early to the warm lowlands
of eastern and western Africa, Asia and to south-east China.
It is called soursop in English and guanabana in most
Spanish-speaking countries. French-speaking areas of West
Indies and West Africa call it corossol, corossol epineux or
cachiman epineux. It is known as guayabano, guayabana (the
Philippines); catoche (Venezuela); nangka belanda, zuurzak,
sirsak (Indonesia); thurian-khaak, noina (Thailand); mundla
(India); fruta de conde, araticum do grande, graviola, jaca do
Para (Brazil); guanaba (El Salvador); nona sri kaya, durian
belanda, durian maki (Malaysia); zapote de viejas, cabeza de
negro (Mexico); huanaba (Guatemala); anona de puntitas,
anona de broquel (Argentina); sinini (Bolivia); and sorsaka,
zuurzak (the Netherlands).
World production and yield
This small, evergreen and quick-growing tree is commonly
found on subsistence farms in South-east Asia and the Pacific
islands. It is grown extensively in Mexico from Culiacan to
Chiapas, and from Veracruz to the Yucatan Penninsula in the
Gulf region. Orchards as large as 20 ha occur. In many other
areas in the world, soursop remains cultivated as a backyard
tree planted along the roads, in house yards, in mixed planting
and wastelands.
It is a highly nutritious fruit crop. The harvesting season is
similar in most areas, differing only in range. Mean monthly
soursop production in Hawaii shows a year-round production
with two peaks. Soursop yields in Hawaii from trees grown in
a marginal field have shown approximately 43 kg/tree on 4year-old trees increasing up to 83 kg/tree on 6-year-old trees.
At Paramaribo, Surinam, soursop yields of 54 kg/tree at 278
trees/ha are reported. No world production, export and
import figures are available. Commercial plantings are mostly
limited to the Philippines, the Caribbean and South America.
Soursop pulp is available commercially from South America.
Uses and nutritional composition
Fruit are harvested when fully mature, firm, yellowish green
with spines set apart. The ripe fruit are usually consumed
fresh as a dessert or snack item. This fruit has the greatest
processing potential of the annonas because of the excellent
flavour of the pulp and high recovery from the large fruit. At
processing plants soursop fruit is stored on racks in the shade
and inspected daily. All fruit found to yield to finger pressure
are removed for processing. Slightly immature fruit will ripen
but such fruit lack the full flavour and aroma, and nectars
prepared from such purée have a flat taste. The fragile skin of
the fruit, irregular shape and softness limit machine
processing and the fruit has to be hand peeled and cored. Pulp
recovery percentages ranged from 62 to 85.5% (Paull, 1982).
Differences in recovery percentages are due to differences in
equipment, extraction methods, cultivar and cultural
practices, including environmental influences. The number of
seeds per fruit also influences pulp recovery.
Soursop pulp is viscous and requires dilution, the pH is
adjusted to 3.7 by addition of citric acid and sugar to 15% to
create a desirable balance between acidity, sweetness and
flavour in the diluted nectar. Unsweetened and sweetened
soursop pulp processed below 93°C show no changes in
organoleptic properties, though freeze preservation produces a
higher quality product. Enriched pulp, sweetened or
unsweetened, can be processed and stored frozen for use in
various products or reconstituted directly by the consumer.
Purée and juice concentrates can be used to prepare iced
soursop drink or mixed with other juices, or made into
sherbets and gelatin dishes. The juice, with the addition of
sugar, makes an excellent ice cream or sherbet for making a
refreshing drink. ‘Champola’ made with strained pulp, milk
and sugar is a famous fruit drink in Havana, Cuba. In the
Dutch East Indies, the juice mixed with wine or brandy is a
popular drink. The fruit also makes excellent preserve, jam or
jelly. In Malaysia, the delicate flavour of soursop is popular for
flavouring ice cream and puddings. In the Philippines, young
soursop fruit with seeds that are still soft may be used as a
vegetable and cooked with coconut milk. Mature but hard
fruit may be made into sweets.
The edible portion of the soursop fruit is an excellent
source of vitamins B and C, and a fair to poor source of
calcium and phosphorus (Table A.21). The predominant
organic acids in the fruit are acetic, lactic and malic acids with
some quantities of citric and oxalic acids. Soursop is a good
source of potassium, riboflavin and niacin.
The soursop has some medicinal uses. A decoction of young
shoots and leaves is a remedy for gall bladder infection,
coughs, diarrhoea, dysentery and indigestion. Mashed leaves
are used as a poultice to alleviate eczema and rheumatism. The
seeds, like those of other members of the genus, contain a
toxic alkaloid. The flowers are antispasmodic. The ripe fruit
prevents scurvy while the unripe fruit may be used for
dysentery and has astringent properties. The tree can also be
used in landscapes.
Botany
The soursop is a small, evergreen, slender and
upright or low-branching and bushy tree, growing to heights
of 4.5–9 m and becomes straggly and untidy with age. The
leaves are alternate, smooth, dark green and glossy above, dull
and paler below, obovate to elliptic, 7–20 cm long. When
crushed, the leaves will emit a strong odour.
Flowers are solitary, 2.5–4 cm long with three thick, fleshy
DESCRIPTION
Annona
43
Table A.21. Proximate fruit composition of soursop in 100 g edible portion
(Source: Wenkam, 1990).
Proximate
Edible flesh
Seed/skin
Water
Energy (kcal)
Energy (kJ)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
%
34
34
80.1
71
247
0.69
0.39
18.23
0.95
0.58
mg
9
0.82
22
29
20
22
mg
16.4
0.07
0.12
1.52
0
petals and three close-set, pale yellow inner petals alternating
with three fleshy, slightly spreading, yellow-green outer petals.
The flower has a peculiar smell. The flowers are
hermaphrodite and are often produced singly or in small
clusters on old wood.
The fruit, a syncarp, is broadly ovoid or ellipsoid and
usually irregularly shaped or curved due to improper carpel
development (Fig. A.5). Fruit are nearly always longer than
they are wide. Fruit size varies from < 0.45 kg to > 4.5 kg,
largely dependent upon the extent of pollination and
fertilization. A normal fruit is generally heart-shaped to oval,
but if there is poor pollination, unfertilized ovules fail to
develop and the resulting fruit assumes distorted irregular
shapes and is usually undersized. The skin is dark green with
many recurved, soft spines 0.5–1.3 cm apart. There is often a
constriction like a fault on the side of the fruit, where the skin
has not swollen and the spines are much closer together. The
fruit stalk is about 3–8 cm long and woody. The ripe pulp,
which adheres to the skin but is easily separated into segments
(which were the separate ovaries), is juicy, creamy white with a
cottony texture and contains many black seeds about 2 cm
long. The pulp has an agreeable subacid flavour with a distinct
aroma.
ECOLOGY AND CLIMATIC REQUIREMENTS Soursops are
capable of growing in a wide range of soil types from sandy to
clay loams provided that the soil has good drainage. The tree is
commonly grown on slightly acid soils with optimum pH at
5–6.5. It also grows on the porous, oolitic limestone of south
Fig. A.5. Annona fruit – soursop (top), sweetsop (bottom left) and
cherimoya (bottom right) (Source: Nakasone and Paull, 1998).
Florida and the Bahamas. Higher and more consistent yields
are obtained on trees grown on well-drained sandy to sandy
loam soils. Waterlogging is a major cause of floral abscission
and root rot such as bacterial wilt caused by Pseudomonas spp.
Soursop cannot tolerate standing water for any length of time
but can tolerate dry soil conditions.
The tree thrives in the warm and humid tropics below
1000 m and is very susceptible to cold. In mountainous areas
(15°–25°C mean minimum) the tree produces very few fruit.
Frost kills young trees, but older trees show some tolerance.
Defoliation and an interruption of fruiting occur when the
temperature drops to near freezing. Poor pollination is a
frequent problem and occurs at high temperature (30°C) and
low humidity (30% relative humidity (RH)), even with hand
pollination. Lower temperature (25°C) and high humidity
(80% RH) greatly improves pollination.
The softwood of the trees makes them susceptible to wind
damage and limb breakage. Fruit productivity is improved by
the provision of windbreaks.
REPRODUCTIVE BIOLOGY Annona species generally require
27–35 days for flower bud development from initiation to
anthesis. Flowering can extend from 3 to 6 months or even
longer, with heavy peaks. The soursop produces fruit
44
Annonaceae
throughout the year but peak production in most areas comes
during summer and early autumn, sometimes with a
secondary peak during early spring. Two major flowering
periods occur after periods of vegetative flushes with the
second peak coinciding with the onset of monsoon in India
(Kumar et al., 1977). Defoliation of A. muricata manually or
by using ethephon spray promotes lateral branch growth and
induces additional flower formation near the apex of the
branches. No photoperiod responses have been reported.
The flowers exhibit both dichogomy and a protogynous
nature. Soursop floral anthesis takes place mostly between
noon and 8 p.m. and 4 a.m.–8 a.m., with pollen release
occurring between 4 a.m. and 8 a.m. Flower opening and
pollen dehiscence are not synchronous and thus very low selfpollination occurs. Cross-pollination may take place early in
the morning because at anthesis the flower usually emits a
fragrance that attracts insects. In order to increase yield, hand
pollination has become an important aspect of cultivation
practices in some areas.
Nitidulid beetles (Carpophilus and Uroporus spp.) are the
important pollinators of Annona flowers with wind and selfpollination being low (1.5%). Also studies have shown that
these beetles breed rapidly in rotting fruit media and that
populations of these beetles are increased by maintaining the
rotting fruit attractant (Thakur and Singh, 1964).
FRUIT DEVELOPMENT Fruit growth shows the typical
sigmoidal curve with maturation occurring in 16–24 weeks.
Low humidity (< 60% RH) and temperature (< 13°C) near
fruit maturity can increase the severity of fruit skin russetting
as well as delaying fruit maturation.
Horticulture
The soursop are usually propagated by seed.
There is a rapid loss of seed viability (6 months) after harvest
and seeds should be planted as soon as possible after removal
from the fruit. Fresh seeds can take up to 30 days to germinate
with about 85–90% germination. Propagation by cuttings or
air layering has not been very successful. Hypocotyl explants
of soursop are suitable for in vitro culture with good rooting
but poor shoot induction (Rasai et al., 1995).
Transplanting should be done at the beginning of the wet
season if there are seasonal dry periods and no irrigation
facilities. Plants should have attained a height of 30–46 cm at
transplanting time with the union of grafted or budded plants
placed 15 cm or so above the ground. Trees should be irrigated
as soon as possible after transplanting. Soursop trials in
Hawaii showed that the spacing should be 4.6 ⫻ 6 m without
affecting growth or interfering with cultural practices.
PROPAGATION
ROOTSTOCKS Atemoya is not compatible with soursop as
rootstock (Sanewski, 1991). In Florida, soursop has been
grafted on 1-year-old seedlings of sugar apple and pond apple,
the latter rootstock being tolerant to flooding.
The annonas are grown in many areas without
irrigation when rainfall is well distributed. Except for pond
apple (Annona glabra), soursop can stand periods of drought
and prefer rather dry conditions. Water stress should be
IRRIGATION
prevented during flowering, fruit set and fruit development as
fruit are more sensitive than leaves. Soursop has a shallow
fibrous root system and may benefit from mulching.
The soursop usually produces a
symmetrically conical tree and is well adapted to the central
leader system. An alternative is to develop a mushroomshaped tree that is topped at 1.8–2.4 m. The fruit in this
system is borne on the lateral branches and hangs down for
ease of harvesting. When properly trained, little pruning is
required except to thin out poorly placed and weak branches.
To contain trees within a certain space allocation and height
limitation the longest branches extending horizontally and
vertically may be pruned annually, preferably immediately
after harvest. Very severe pruning reduces subsequent fruiting.
PRUNING AND TRAINING
Observations in Hawaii and
Mexico indicate that it is desirable to provide 1.3 kg of a
triple-15 fertilizer formulation during the first year of
production, split into two applications. In Hawaii, the first
increment should be given around February for the primary
crop in July and the second increment applied in August for
the December–January secondary crop. Each year thereafter,
up to approximately the sixth-bearing year, the total amount
can be increased by approximately 0.45 kg/tree/year. In the
Philippines, the application of 100–150 g ammonium sulphate
a month after planting and an equal amount 6 months after or
at the end of the rainy season is recommended. The quantity is
increased every year until the trees start to bear fruit, at which
time, about 250–300 g complete fertilizer, high in nitrogen
and potassium, is used. A full-grown tree may need at least
500 g complete fertilizer per application.
NUTRITION AND FERTILIZATION
POSTHARVEST HANDLING AND STORAGE High temperature
can cause premature fruit ripening and fermentation of the
fruit. Fruit is harvested when fully mature and firm. The skin
colour changes as the fruit approaches maturity. The
immature soursop is dark green and shiny, losing its sheen and
becoming slightly yellowish green with spines set apart at
maturity. Soursop respiration begins to increase within a day
after harvest and reaches its peak at the sixth to eighth day.
Ethylene production is initiated approximately 48 h after
initiation of respiration rise and reaches its peak at about the
same time as the respiration peak reaches a plateau (Paull,
1982). Total soluble solids increases from around 10–16%
during the first 3 days of ripening. The major titratable acids
are malic and citric acids. After day 5 to 6, titratable acidity,
ethylene production and total phenols decline, changes that
produce a bland flavour and even a slightly objectionable
odour. The optimum edible stage is at days 6 and 7, which
coincide with ethylene production (Paull, 1982).
Fruit is hand harvested and put into lug boxes or baskets. In
large soursop orchards mechanical harvesting aids are feasible
and accelerate handling. Harvested fruit should be handled
with care to prevent bruising of the skin. Firm fruit are held
after harvest for 4–7 days at room temperature before
softening begins; optimum quality processing occurring 5 and
6 days later (Paull et al., 1983). The skin of ripening soursop
gradually turns dark brown to black, but the flesh is unspoiled.
Storage below 15°C causes chilling injuries and a failure to
Annona
develop full flavour. At lower temperatures, skin discoloration
rapidly occurs.
DISEASES, PESTS AND WEEDS A number of diseases have been
reported in the literature (Table A.22). Anthracnose caused by
Colletotrichum gloeosporioides (Glomerella cingulata) is the most
serious on soursop, particularly in areas of high rainfall and
atmospheric humidity and during the wet season in dry areas
(Alvarez-Garcia, 1949; Dhingra et al., 1980). This disease
causes twig dieback, defoliation, dropping of flowers and fruit.
On mature fruit the infection causes black lesions.
Black canker (Phomopsis anonacearum) and diplodia rot
(Botryodiplodia theobromae) occur mostly on neglected trees
and cause similar symptoms of purplish to black lesions
resulting in mummified fruit. Marginal leaf scorch is also
caused by P. anonacearum and B. theobromae and causes twig
dieback. Diplodia rot has darker internal discoloration and
deeper, more extensive corky rot in fruit. Cylindrocladium
fruit and leaf spot is caused by a soil-borne fungus,
Cylindrocladium colhounii. It can cause almost total loss of fruit
during years of persistent heavy rains. Symptoms begin with
small dark spots on the shoulders of the fruit that spread along
the sides, enlarge, become dry and crack. Infection is skindeep but fruit becomes unmarketable.
Some insect pests occur in numerous growing areas (Table
A.23). One of the most serious insects in Trinidad is the
45
cerconota moth (Cerconota anonella) that lays its eggs on
young fruit. The emerging larvae tunnel into the pulp, causing
blackened, necrotic areas. It is not uncommon to find every
fruit larger than 7.5 cm infested. Bagging the fruit is
sometimes done. This moth has been reported in the
American tropics as far south as Brazil and is a major limiting
factor in Surinam.
The bephrata wasp (Bephrata maculicollis) is widely
distributed throughout the Caribbean and Mexico, Central
and northern South America. This wasp is considered to be
the most important pest in Florida (Campbell, 1985). The
larvae infest the seeds and cause damage to the pulp as they
bore through the flesh to emerge when the fruit matures. The
thecla moth (Thecla ortygnus) is widespread through parts of
the Caribbean and in the American tropics but it is not
considered to be as serious as the cerconota moth and the
bephrata wasp. Primary damage is done to the flowers. The
larvae feed on flower parts such as the perianth, stamen and
stigmas with the flowers failing to set fruit.
Mature green annonaceous fruit are rarely infested by the
Mediterranean fruit fly (Ceratitis capitata) and oriental fruit fly
(Dacus dorsalis), but these are found on occasion in treeripened fruit.
Mealy bugs and various species of scale insects are found
universally and usually become a serious pest on neglected
trees. Red spider mites can become a serious problem in dry
Table A.22. Major diseases of soursop.
Common name
Organism
Parts affectedand symptoms
Country/region
Anthracnose
Colletotrichum gloeosporioides
(Glomerella)
Armillaria leuteobubalina
Pseudomonas solanacearum
Botryodiplodia theobromae
Phomopsis anonacearum
Phytophthora palmivora
Phakospora cherimoliae
Gliocladium roseum
Rhizopus stolonifer
Cercospora sp.
Flowers, fruit, leaves, dieback, seedling damping off
Universal
Roots, base of trees, decline
Tree wilt
Leaf scorch, twig dieback
Leaf scorch, twig dieback
Spots on immature fruit, fruit drop, twig dieback
Leaves
Fruit
Fruit
Leaves
Australia
Australia
Australia, Brazil
Australia
Australia
Florida
India
Brazil
Colombia
Armillaria root rot
Bacterial wilt
Black canker (diplodia rot)
Black canker
Purple blotch
Rust fungus
Fruit rot
Rhizopus rot
White spot
Table A.23. Major insect pests of soursop.
Common name
Organism
Parts affected
Country/region
Bephrata wasp (soursop wasp)
Wasp
Cerconota moth (soursop moth)
Thecla moth
Banana spotting
Mealy bug
Citrus mealy bug
Southern stink bug
Caribbean fruit fly
Queensland fruit fly
Potato leaf hopper
Red spider mite
Scale insects
Coconut scale
Bephrata maculicollis
Bephratelloides paraguayensis
Cerconota anonella
Thecla ortygnus
Amblypelta lutescens
Dysmicoccus
Planococcus citri
Nezara viridula
Anastrepha suspensa
Dacus tryoni
Empoasca fabae
Several genera, species
Saissetia coffeae
Aspidiotus destructor, other genera and species
Fruit
Fruit
Fruit
Flower, young fruit
Young fruit
Stem, leaves
Fruit
Fruit
Fruit
Fruit
Leaves
Leaves, flowers
Leaves, stem
Leaves, stem
Mexico, Americas, Trinidad, Surinam
Americas, Barbados
Americas, Trinidad, Surinam
Americas, Caribbean
Queensland
Universal
Queensland
Caribbean
Caribbean, Mexico
Australia
Caribbean
American tropics
Universal
Caribbean
46
Annonaceae
areas or during dry seasons. Heavy infestations have been
observed on soursop flowers and leaves in the Tecoman area of
Mexico during the prevailing dry period with trees showing
heavy flower drop.
Problem weeds especially grasses and twining weeds should
be controlled before planting. The shallow root system limits
the use of cultivation under the tree.
A desirable hybrid would be
between the cherimoya and soursop. This would combine the
larger fruit size and acidity of the soursop and the sweetness,
flavour and texture of cherimoya. Attempts to cross the
soursop with cherimoya, ilama, bullock’s heart or sweetsop
have not been successful and may reflect considerable genetic
distance of soursop from the other species (Samuel et al.,
1991). Breeding programmes have focused on selections from
seedling populations. Early maturity, greater yield, better fruit
appearance, and in the subtropics greater cold tolerance are
the most frequent breeding objectives. In Indonesia and the
Philippines, two distinct soursop forms exist, that of the sourand the sweet-fruited with the sweet-fruited having fewer
seeds than the sour-fruited. The sweet ‘Sirsak Ratu’ from Java
occurs in a few regions only, whereas the sour type is more
common (Yaacob and Subhadrabandhu, 1995).
Emma Ruth V. Bayogan and Robert E. Paull
MAIN CULTIVARS AND BREEDING
Literature cited and further reading
Alexander, D.E. (1982) Some Tree Fruits for Tropical Australia.
Commonwealth Scientific and Industrial Research Organization,
Canberra, Australia, pp. 44–47.
Allen, B.M. (1967) Malayan Fruit – An Introduction to Cultivated
Species. Donald Moor Press Ltd, Singapore.
Alvarez-Garcia, L.A. (1949) Anthracnose of the Annonaceae in Puerto
Rico. University of Puerto Rico Journal of Agriculture 33, 27–43.
Braga Sobrinno, R., Bandeira, C.T. and Mesquita, A.L. (1999)
Occurrence and damage of soursop pests in Northeast Brazil.
Crop Protection 18, 539–541.
Campbell, C.W. (1985) Cultivation of fruits of the Annonaceae in
Florida. Proceedings of the American Society for Horticultural
Science Tropical Region 29, 68–70.
Coronel, R.E. (1998) Promising Fruits of the Philippines. College of
Agriculture University of the Philippines at Los Baños.
Dhingra, J., Mehrotra, R.S. and Aneja, I.R. (1980) A new postharvest
disease of Annona squamosa L. Current Science 49, 477–478.
George, A.P. and Nissen, R.J. (1986) The effects of root temperature
on growth and dry matter production of Annona species. Scientia
Horticulturae 31, 95–99.
George, A.P. and Nissen, R.J. (1987) Propagation of Annona species:
a review. Scientia Horticulturae 33, 75–85.
Hoyos, P.P. and Zarate, R.R.D. (1985) Etiological and epidemiological
study of white spot on the soursop (Annona muricata) in Valle del
Cauca, Colombia. Acta Agronomy Palmira 35 (1), 81–92.
Ithnin, B. and Mohd Khalid, M.Z. (1996) Flowering and yield
patterns of soursop, Annona muricata. In: Abstracts of the
International Conference on Tropical Fruits, 23–26 July, Malaysia,
p. 64.
Kumar, R., Hoda, M.N. and Singh, D.K. (1977) Studies on the floral
biology of custard apple (Annona squamosa Linn.). Indian Journal
of Horticulture 34, 252–256.
Marler, J.E., George, A.P., Nissen, R.J. and Andersen, P.J. (1994)
Miscellaneous tropical fruits – annonas. In: Schaffer, B.C. and
Andersen, P.C. (eds) Handbook of Environmental Physiology of
Fruit Crops. Vol. II. Subtropical and Tropical Crops. CRC Press,
Boca Raton, Florida, pp. 200–206.
Morton, J.F. (1987) Fruits of Warm Climates. J.F. Morton, Florida,
pp. 65–90.
Nakasone, H.Y. (1972) Production feasibility for soursop. University
of Hawaii, Hawaii Farm Science 21, 10–11.
Nakasone, H.Y. and Paull, R.E. (1998) Tropical Fruits. CAB
International, Wallingford, Oxon, UK.
Paull, R.E. (1982) Postharvest variation in composition of soursop
(Annona muricata L.) fruit in relation to respiration and ethylene
production. Journal of the American Society for Horticultural
Science 107, 582–585.
Paull, R.E. (1998) Soursop. In: Shaw, P.E., Chan, H.T., Jr and Nagy,
S. (eds) Tropical and Subtropical Fruits. AgScience Inc.,
Auburndale, Florida, pp. 386–400.
Paull, R.E., Deputy, J. and Chen, N.J. (1983) Changes in organic
acids, sugars, and headspace volatiles during fruit ripening of
soursop (Annona muricata L.). Journal of the American Society for
Horticultural Science 108, 931–934.
Quisumbing, E. (1951) Medicinal Plants of the Philippines. Bureau of
Printing, Manila, 1234 pp.
Rasai, S., George, A.P. and Kantharajah, A.S. (1995) Tissue culture
of Annona spp. (cherimoya, atemoya, sugar apple and soursop): a
review. Scientia Horticulturae 62, 1–14.
Samuel, R., Pineker, W., Balasubramaman, S. and Morawetz, W.
(1991) Allozyme diversity and systematics in Annonaceae – a pilot
project. Plant Systematics and Evolution 178, 125–134.
Sanewski, G.M. (ed.) (1991) Custard Apples – Cultivation and Crop
Protection. Information Series QI90031, Queensland Dept.
Primary Industry, Brisbane, Australia.
Smith, D. (1991) Insect pests. In: Sanewski, G.M. (ed.) Custard
Apples – Cultivation and Crop Protection. Information Series
QI90031, Queensland Dept. Primary Industry, Brisbane,
Australia, pp. 73–79.
Sturrock, D. (1940) Tropical Fruits for Southern Florida and Cuba and
their Uses. The Arnold Arboretum of Harvard University,
Cambridge, Massachusetts, 131 pp.
Thakur, D.R. and Singh, R.N. (1964) Studies on pollen morphology,
pollination and fruit set in some annonas. Indian Journal of
Horticulture 22, 10–17.
Wenkam, N.S. (1990) Foods of Hawaii and the Pacific Basin. Fruits and
Fruit Products: Raw, Processed, and Prepared. Vol. 4: Composition.
Research Extension Series 110. Hawaii Institute of Tropical
Agriculture and Human Resources (HITAHR), College of
Tropical Agriculture and Human Resources, Hawaii.
Worrell, D.B., Carrington, C.M.S. and Huber, D.J. (1994) Growth,
maturation, and ripening of soursop (Annona muricata L.) fruit.
Scientia Horticulturae 57, 7–15.
Yaacob, O. and Subhadrabandhu, S. (1995) The Production of
Economic Fruits in Southeast Asia. Oxford University Press, New
York, 419 pp.
Annona senegalensis
wild soursop
This very popular African Annona (Annona senegalensis Pers.,
Annonaceae) is widespread in tropical semiarid and sub-humid
Annona
areas, often growing in savannah regions. It occurs from the
Nile River area to the Transvaal and Zululand in South Africa.
It is common as a single understorey shrub in wooded
savannah areas. Annona senegalensis is among the popular
shrubs and trees browsed by Fulani pastoralists’ cattle in
central Nigeria (Bayer, 1990). The common names include
wildesuikerappel in Afrikaans, giishta and yebere lib in
Amharic, gishta and gishta gaba in Arabic, wild custard apple
and wild soursop in English, annone and pomme cannelle du
senegal in French, and numerous local African names. Morton
(1987) gives mavulu, mugosa, mbokwe, makulo and mlamote
in Kenya; mtopetope and mchekwa in Zanzibar and Pemba;
mabengeya, elipo, obwolo and ovolo in Uganda; aboboma,
batanz, bangoora and bullimbuga in Ghana; mposa, muroro
and mponjela in Malawi; dilolo, iolo and malolo in Angola;
sougni, mete, dangan, sounsoun, tangasou, dougour, ianouri,
ndong and anigli in West Africa.
Uses and nutritional composition
The ripe fruit’s edible white pulp has a pleasant, pineapplelike taste. The leaves are sometimes used as vegetables, while
the flowers serve as a spice for various meals. A yellow or
brown dye and an insecticide are obtained from the bark. The
bark is also used for treating guinea worms and other worms,
diarrhoea, gastroenteritis, snakebite, toothache and respiratory
infections (Fatope et al., 1996; Alawa et al., 2003). The leaves
are used for treating pneumonia and as a tonic to promote
general well-being. The roots are used for stomach ache,
venereal diseases, chest colds and dizziness. Various plant parts
are combined for treating dermatological diseases and
ophthalmic disorders. Annona senegalensis is shown to be
therapeutically effective against Trypanosoma brucei brucei in
mice, which agrees with the claims of some practitioners of
traditional medicine that it is effective against trypanosomiasis
in man (Igweh and Onabanjo, 1989; Sahpaz et al., 1994;
Freiburghaus et al., 1996).
The edible fleshy pulp contains many seeds, which are
particularly eaten in northern Nigeria. About 11 fatty acids
have been identified in the seeds of A. senegalensis of which
oleic, gondoic, palmitic and stearic acids predominate (Wélé et
al., 2004). Terpinen-4-ol is the major volatile component.
47
2.5–4 cm) is formed from many fused carpels. The unripe
fruit is green, turning yellow to orange on ripening, with the
outline of the carpels forming a coloured network. The edible
fleshy pulp contains numerous oblong orange-brown seeds.
TAXONOMY AND NOMENCLATURE There are many synonyms
for this species including Annona arenaria Thonn. ex Schum.,
Annona chrysophylla Boj., Annona porpetac Bail., Annona
senegalensis Pers. var. chrysophylla Boj., Annona senegalensis
Pers. var. latifolia Olive, and Annona senegalensis var. porpetac
(Bail.) Diels.
ECOLOGY AND CLIMATIC REQUIREMENTS Richardson et al.
(2004) described the Annonaceae as a pantropically distributed
family found predominantly in rainforests, so they are
megathermal taxa. However, A. senegalensis grows best in low
and mid-elevation tropical climates as a deciduous shrub. It
may grow in drier areas as long as the roots have continuous
access to water. The species occurs along riverbanks, on fallow
land, in swamp forests and at the coast, often as a single plant
in the understorey of savannah woodlands. It is found from
coastal areas to 2400 m. The major habitat, according to
Isawumi (1993), is the savannah, although it is identified by
Salami (2001) as one of the common tree species now found in
the tropical rainforest belt of Nigeria. The mean annual
temperature is from 17–30°C and mean annual rainfall of
700–2500 mm. It is very sensitive to frost. The tree is found
on diverse soil types and does well on sandy loam soils.
REPRODUCTIVE BIOLOGY The tree starts bearing fruit in 3
years. Along the coast of Tanzania, it flowers from December
through to February (Mbuya et al., 1994). Flowering occurs
earlier from October to December elsewhere. The period of
defoliation is generally brief, and leaf flushing, flowering and
fruiting occur mostly in the dry season (Devineau, 1999). The
fruit matures during January to March, sometimes extending
to April. Poor pollination leads to misshaped fruit. Hand
pollination can improve the shape and yield. As the fruit
ripens, there is a tendency for cracking to occur.
Horticulture
Propagation is normally by seed and root suckers.
Scarification of the seed improves germination. The trees
coppice well after felling and are readily pruned to a desired
shape. Young plants do not compete well with weeds and need
to be protected from fire and browsing animals.
CULTURE
Botany
This shrub or small tree is 2–6 m tall, taller
under favourable conditions. The bark is smooth to rough,
silvery grey or grey-brown, with leaf scars; the slash is pale
pink (Isawumi, 1993). Young branches have dense, brown,
yellow or grey hairs that are lost later. The leaves (6–18.5 ⫻
2.5–11.5 cm) are simple, alternate, oblong to ovate or elliptical,
with eight to ten pairs of prominent lateral nerves. They are
green to bluish green with almost no hairs on top, but often
with brownish hairs on the underside. The solitary or groups
of two to four flowers are up to 3 cm in diameter, on 2 cm long
stalks. The flowers arise in leaf axils. The three ovate sepals are
free and smaller than the petals. There are six fleshy cream to
yellow petals in two whorls that are greenish outside, creamy
or crimson inside. The inner whorl of the petals curve over the
stamens and ovary. The stamens are 1.7–2.5 mm long. As with
other Annona spp., the fleshy ovoid to globose fruit (2.5–5 ⫻
DESCRIPTION
DISEASES, PESTS AND WEEDS As with other annonas,
anthracnose caused by Colletotrichum gloeosporioides is a major
problem on leaves, flowers and fruit. On the leaf, it causes
small, light green spots, dark spots on the flowers and
shedding and fruit mummification.
Victor N. Enujiugha
Literature cited and further reading
Alawa, C.B.I., Adamu, A.M., Gefu, J.O., Ajanusi, O.J., Abidu, P.A.,
Chiezey, N.P., Alawa, J.N. and Bowman, D.D. (2003) In vitro
screening of two Nigerian medicinal plants (Vernonia amygdalina
and Annona senegalensis) for anthelmintic activity. Veterinary
Parasitology 113, 73–81.
48
Annonaceae
Bayer, W. (1990) Use of native browsing by Fulani cattle in central
Nigeria. Agroforestry Systems (Historical Archive) 12, 217–228.
Devineau, J.L. (1999) Seasonal rhythms and phonological plasticity
of savanna woody species in a fallow farming system (South-West
Burkina Faso). Journal of Tropical Ecology 15, 497–513.
Fatope, M.O., Audu, O.T., Takeda, Y., Zeng, L., Shi, G., Shimada, H.
and McLaughlin, J.L. (1996) Bioactive ent-kaurene diterpenoids
from Annona senegalensis. Journal of Natural Products 59, 301–303.
Food and Agriculture Organization (FAO) (1983) Food and fruit
bearing forest species. 1: Examples from eastern Africa. Forestry
Paper #44/1, Food and Agricultural Organization, Rome.
Freiburghaus, F., Kaminsky, R., Nkunya, M.H. and Brun, R. (1996)
Evaluation of African medicinal plants for their in vitro
trypanocidal activity. Journal of Ethnopharmacology 55, 1–11.
Igweh, A.C. and Onabanjo, A.O. (1989) Chemotherapeutic effects of
Annona senegalensis in Trypanosoma brucei brucei. Annals of
Tropical Medicine and Parasitology 83, 527–534.
Isawumi, M.A. (1993) The common edible fruits of Nigeria – Part I.
The Nigerian Field 58, 27–44.
Mbuya, L.P., Msango, H.P., Ruffo, C.K., Birnie, A. and Tengnas, B.
(1994) Useful trees and shrubs for Tanzania. Regional Soil
Conservation Unit (RSCU) and Swedish International
Development Authority (SIDA), Nairobi, Kenya, 542 pp.
Morton, J. (1987) Wild custard apple. In: Morton, J.F. (ed.) Fruits of
Warm Climates. Creative Resources Inc., Miami, Florida,
pp. 86–88.
Richardson, J.E., Chatrou, L.W., Mols, J.B., Erkens, R.H.J. and Pirie,
M.D. (2004) Historical biogeography of two cosmopolitan
families of flowering plants: Annonaceae and Rhamnaceae.
Philosophical Transactions: Biological Sciences 359, 1495–1508.
Sahpaz, S., Bories, C., Loiseau, P.M., Cortes, D., Hocquemiller, R.,
Laurens, A. and Cave, A. (1994) Cytotoxic and antiparasitic activity
from Annona senegalensis seeds. Planta Medica 60, 538–540.
Salami, A.T. (2001) Agricultural colonization and floristic degradation
in Nigeria’s rainforest ecosystem. The Environmentalist 21,
221–229.
Wélé, A., Ndoye, I. and Badiane, M. (2004) Fatty acid and essential
oil compositions of the seed oil of five Annona species. Nigerian
Journal of Natural Products and Medicine 8, 62–65.
Annona squamosa
sweetsop
Sweetsop (Annona squamosa L., Annonaceae) is a small tropical
tree originating in the New World tropics, probably in the
Caribbean region. This species is the most widely grown
Annona spp. in the tropical regions of the Americas, Africa,
Asia and the Pacific. Sweetsop is also named sugar apple and
has many other regional names, such as custard apple (India),
anon (Spanish), ata (Portuguese), noi-na (Thailand), atis
(Philippines) and fan-li-chi (Taiwan).
annual production of sweetsop in Taiwan was 50,005 t from a
total harvested area of 4978 ha. More than 80% of the
production is in Taitung county of southern Taiwan.
Fruit is usually harvested from July to October, and can be
extended to March if summer pruning is conducted. Taiwan’s
peak of production occurs between July and March. In India
(Poona), the peak of production occurs later between August
to November, and in Thailand, Florida and the Caribbean
between July and September.
Uses and nutritional composition
Sweetsop is a good source of carbohydrate, potassium,
calcium, phosphorus and ascorbic acid (Table A.24). The fruit
is usually consumed fresh and also can be used to make juices,
shakes and ice creams. There are folk medicinal applications of
sweetsop. The stems, leaves and seeds of sweetsop contain the
toxic aporphine alkaloids (anonaine) (Bhakuni et al., 1972). In
tropical America, a leaf decoction is used as a cold remedy
while a bark decoction is used to stop diarrhoea. Sweetsop
juice is also able to resolve diarrhoea in children (Enweani et
al., 1998). The root is used in the treatment of dysentery and
the ground seed powder has insecticidal properties. The bark
extracts have been shown to have an anti-tumour effect in the
laboratory (Hopp et al., 1996).
Botany
TAXONOMY AND NOMENCLATURE The Annonaceae, or
custard apple family, comprises about 120 genera and more
than 2000 species (Leboeuf et al., 1982). The genus Annona is
the most economically important one, containing c.120
Table A.24. Proximate sweetsop fruit compositions per 100 g (USDA, 2002).
Proximate
Water
Energy (kcal)
(kJ)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
World production and yield
Sweetsop is the most widely grown Annona species. The fruit
is frequently found in village markets but has not shown much
potential for large commercial cultivation due to the small
fruit size, frequent cracking at maturity and poor shelf life.
The perishable nature and supply shortages make marketing
localized or air shipment essential. However, sweetsop is
intensively cultivated in Taiwan. In the last 5 years, the average
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin E
Panthothenic acid
Vitamin B6
Vitamin A
%
73.32
94
393
2.06
0.29
23.64
4.40
0.78
mg
24
0.6
21
32
247
9
mg
363
0.111
0.113
0.883
0.590
0.226
0.200
6 IU
Annona
species. Three major commercial species throughout the world
are: the cherimoya (Annona cherimola), sweetsop (A. squamosa)
and atemoya (a hybrid of A. cherimola and A. squamosa). The
sweetsop species name ‘squamosa’ refers to the knobbly
appearance of the fruit. The chromosome number of A.
squamosa is 2n = 14.
The sweetsop tree is normally smaller than the
cherimoya, attaining heights of 3–6 m with slender branches.
The leaves are oblong-lanceolate, 10–15 cm long and 3–5 cm
wide, alternately arranged on short petioles and narrower than
those of the cherimoya (Fig. A.6). Young leaves are slightly
hairy. The sweetsop is semi-deciduous in growth habit and
most leaves are shed before new shoots appear. The flowers of
sweetsop are hermaphrodite and exhibit a protogynous
dichogamy nature. Flowers are axillary, pendant, single or in
clusters of two to five on leafy shoots. The flower is 2–4 cm
long and contains three degenerated sepals and six petals. The
six petals are arranged into two whorls with three each and the
petals of the inner whorl are degenerated into small scales or
completely disappear. The multiple pistils grow on the conical
receptacle, in the centre of the flower with a number of
stamens at the periphery. The compound fruit is nearly heartshaped, 6–10 cm in diameter (see Fig. A.5 in entry for Annona
muricata). It is yellowish green in colour but a purple-fruited
variant is also known. The exterior parts of adjacent carpels
(the conical segments) are not completely fused and these
rounded protuberances separate frequently, exposing the
white flesh upon ripening. Many of the conical segments
contain a single black or dark-brown seed. There are 30–40
seeds in an average fruit.
DESCRIPTION
The sweetsop is
probably the most drought tolerant Annona species as it grows
and produces poorly where rains are frequent. Sweetsop does
very well in the drier north of Malaysia than in the south
which has year-round high moisture.
Temperature is a limiting factor, with frost killing young
trees, but older trees show some tolerance. Sweetsop does not
require a period of low temperature to flower and does well
under lowland conditions and dry periods. Seedlings have
49
high photosynthesis activity at 30°C and show vigorous shoot
growth (Higuchi et al., 1998). Poor pollination is a frequent
problem under high temperatures (> 30°C) and low humidity
(< 60% relative humidity (RH)), even with hand pollination.
Lower temperature (25°C) and higher humidity (80% RH)
greatly improves pollination. However, very high RH (> 95%)
may reduce the stigma receptivity (Marler et al., 1994) and
affect anther dehiscence. Hence, high humidity but no rain
during blooming is advantageous.
Light penetration to the base of vigorous trees with a dense
canopy in a closely spaced orchard can be extremely low and
can reduce fruit set. Pruning practices and spacing need to be
adjusted to increase light penetration. No photoperiod
responses have been reported.
The softwood makes the trees susceptible to wind damage
and limb breakage. Tree shaking may also be partially
responsible for penetration by collar rot organisms. The fruit
skin is easily damaged by rubbing and exposure to drying
winds (Marler et al., 1994). Productivity is improved by
windbreaks and overhead misting to raise the RH above 60%.
Under-tree sprinklers and efficient irrigation scheduling are
also helpful. The tree is sensitive to salinity stress (Marler and
Zozor, 1996). Trees grown in tropical coastal areas may be
damaged by wind-borne salt that can be alleviated by the use
of windbreaks and overhead sprinklers.
Sweetsop is capable of growing in a wide range of soil types,
from sandy soil to clay loams. However, the tree is shallowrooted and waterlogging can lead to root rot. Flooding of
sweetsop seedlings and rootstocks greatly reduces growth and
photosynthetic rate (Núñez-Elisea et al., 1999). High yields
occur on well-drained sandy to sandy loam soils. The optimal
soil pH for sweetsop is 6.0–6.5.
ECOLOGY AND CLIMATIC REQUIREMENTS
Fig. A.6. Leaf and flower of Annona squamosa also showing buried
axillary bud (Source: Nakasone and Paull, 1998).
REPRODUCTIVE BIOLOGY The flowers of sweetsop are
hermaphrodite and are produced singly or in small clusters on
the current season’s growth. Flower initiation begins at the
basal end of the growing branch (Lo, 1987). New flowers
continue to appear towards the apex of the shoot as flowers
produced earlier at the basal portion mature.
The period from floral bud initiation to anthesis is highly
variable. Differences in floral behaviour can be attributed to
both genetic variability and climatic differences. Flowering can
be extended from 3 to 6 months or longer, with heavy peaks.
Two major flowering periods occur after periods of vegetative
flush with the second peak coinciding with the onset of
monsoon in India (Kumar et al., 1977).
The flowers exhibit a protogynous dichogamy nature that
limits self-pollination and cross-pollination can increase fruit
set. Nitidulid beetles (Carpophilus spp. and some others) are
the important pollinators (George et al., 1992) with wind and
self-pollination being low. Pollen grains appearing early in a
flowering season have lower germination rates than pollen
from late flowers (Yang, 1988). The pollen is usually shed in
loosely bound tetrads; individual grains are also observed.
Pollen grains deposited on the stigma surface only germinate
during the female phase and are inhibited at or after anther
dehiscence. Once the pollen germinates, there appears to be
no apparent barrier to the growth of compatible pollen tubes
towards the embryo sac (Vithanage, 1984). Artificial
pollination is frequently practised to ensure pollination and
50
Annonaceae
good fruit shape. Hand pollination is normally carried out
before 8 a.m. every week using a small brush. Pollen can be
collected in the morning between 5 and 8 a.m. from fully open
flowers, when the sacs have turned from white to cream. The
collected pollen is used to pollinate half-open flowers whose
pistils are already receptive. Hand pollination has shown some
variable results and is less successful under non-favourable
climatic conditions and on young vigorous trees.
Hand pollination in commercial orchards is tedious, time
consuming and a costly practice. Attempts have been made to
use growth regulators, with considerable variation in the
results obtained. Treatments with naphthalene acetic acid
(NAA), benzyladenine (BA) and gibberellic acid-3 (GA3) at
25–250 ppm can induce fruit set with high variation among
treatments. Most of the fruit induced are seedless and are
smaller, with lower flavour and less fruit splitting than occurs
in seedy fruit that result from pollination (Yang, 1988).
FRUIT DEVELOPMENT Fruit growth shows a double sigmoidal
curve with maturation occurring in 14–18 weeks depending upon
cultivars and growing conditions. Fruit is harvested when fully
mature and firm. Low humidity (< 60% RH) and temperature
(< 13°C) near fruit maturity can increase the severity of fruit skin
russetting as well as delaying fruit maturation.
Horticulture
PROPAGATION Sweetsop is usually propagated by seeds. The
recalcitrant seeds should be planted as soon as possible after
removal from the fruit. Seeds from fruit having a rest of about 1
week after harvest are better than those directly taken from the
harvested fruit. Seeds can take up to 20–30 days to germinate
and germination is increased by soaking seeds for 3–4 days.
Inarching of A. squamosa to Annona reticulata rootstock has
had up to 70% success rate. Nevertheless, inarching is time
consuming and costly for large-scale propagation (George and
Nissen, 1987). Grafting and budding have a similar success
rate. The branches should be defoliated 1–2 weeks before
scion wood is cut to induce bud swelling. Rooting of tip and
stem cuttings has been used for some cultivars, while good
success has been achieved with micropropagation. Air layering
has been less successful (< 10% success rate).
Transplanting should be done at the beginning of the wet
season if there are seasonal dry periods and no irrigation. In
the subtropics, planting should not occur if there is a risk of
frost. Plants should have attained a height of 30–45 cm at
transplanting time with the union of grafted or budded plants
placed 15 cm or so above the ground. Trees should be irrigated
as soon as possible after transplanting, with wind and sun
guards sometimes required.
The periodically pruned small sweetsop can be spaced at
3.5–4.0 ⫻ 4.5–5.0 m. In dry areas with less luxuriant growth,
closer within-row spacing can be considered. A closer spacing
would increase humidity and benefit the longevity of stigma
receptivity. However, for the convenience of field machine
operations and consistent yields, spacing can be increased to
5.0–5.5 ⫻ 6.0–6.5 m.
ROOTSTOCKS Although sweetsop is often propagated by
seeds, superior cultivars can be propagated via inarching,
budding and grafting to sweetsop or other Annona rootstocks,
such as A. cherimola and A. reticulata (Sanewski, 1991).
Annona glabra is compatible but less hardy. Annona muricata
and Annona palustris are not compatible rootstocks for A.
squamosa.
PRUNING AND TRAINING Training of trees should begin in the
nursery and pruning should continue after transplanting. It is
desirable to train the tree to a single trunk up to a height of
about 80–90 cm and then headed back to produce lateral
branches. The scaffolds should be trimmed to about 60 cm
long to induce lateral branching. The tree height should be
maintained at c.2–2.5 m with a canopy of c.2.5–3 m in diameter
to prevent wind damage and to make it convenient for field
practices. In general, pruning is carried out when the trees are
dormant and involves removal of lower limbs touching the
ground and branches in the centre that may be rubbing against
each other. One-year-old branches are cut back to 10–15 cm to
induce new growth that will be the fruiting shoots, and the
pruning leaves 120–150 branches/tree.
The lateral buds of sweetsop are subpetiolar, in the base of
the swollen leaf petiole (Fig. A.6). Leaf shed must occur prior
to elongation of the ‘buried’ buds. Removal of leaves
mechanically by stripping or chemically with urea or ethephon
releases these buds. In Taiwan, normal pruning occurs in
February/March with fruit harvest from July through to
October. Pruning in January can induce flowering but it takes a
longer time to induce floral development and the total flower
number is less than pruning in February/March. Summer
pruning of selected shoots (June–September) can lead to
harvesting fruit from November to the next March (Yang,
1987). The shoots are pruned back to c.10 cm with two or
three buds left and the leaves on the pruned shoots must be
removed to release the buds.
AND FERTILIZATION The annonas have an
indeterminate growth habit (axillary flowering) and applying
nitrogen does not greatly interfere with floral initiation.
However, excessive tree vigour is usually associated with
reduced flowering and yields in many tree crops. In Taiwan,
continued research and field observation of sweetsop nutrition
(Chang, 2000) has led to greater refinements in terms of
fertilizer quantity and tree age applications (Table A.25). After
8 years of age, the annual amounts of NPK (nitrogen,
phosphorus, potassium) fertilizer remain the same, as the tree
size is kept constant by annual pruning and by competition
from adjacent trees. The annual requirements of nitrogen,
phosphorus and potassium are split into three increments.
The early spring application includes all the annual
phosphorous application and 20% of the nitrogen and
NUTRITION
Table A.25. A guide to annual application of nitrogen, phosphorus and
potassium (NPK) for sweetsop trees of different ages in Taiwan (Source:
Chang, 2000).
Tree age (years)
2
4
6
8
N (g/tree/year)
P (g/tree/year)
K (g/tree/year)
100–150
350–500
450–650
700–1000
44–220
88–220
132–220
175–220
83–125
208–415
374–415
580–830
Annona
potassium. In the summer application, 70% of the nitrogen is
applied to stimulate vegetative growth with 40% of the
potassium after summer pruning. The remainder is applied in
the autumn (Chang, 2000). The use of foliar nutrient analysis
has become a useful management tool in determining
sweetsop fertilizer programmes (Table A.26). The most
recently matured leaf, the third or fourth leaf from the apex on
a non-fruiting shoot without a leaf flush is used. The best
sampling time is in early December.
Black speck is a physiological disorder possibly caused by
calcium deficiency. Symptoms begin with small dark spots
primarily on the shoulders and waist of the fruit skin. In
serious cases, the number of dark spots increases and covers
the whole fruit. The damage is only limited to a thin layer of
flesh tissue right under the skin but fruit has reduced market
values. Spraying Ca(NO3)2 or CaCl2 (0.3–0.5%) two to three
times over 5–7 days when early symptoms of black speck
appears, controls the symptoms (Lin, 2000).
POSTHARVEST HANDLING AND STORAGE The skin colour
changes from greyish green to yellow-green as the fruit
approaches maturity. Adjacent carpels near the peduncle
Table A.26. Tentative leaf nutrient standards for sweetsop in Taiwan,
presented as a guide (Source: Anon., 1995).
Acceptable range
Nutrient
Sampling in
May–June
N (%)
P (%)
K (%)
Ca (%)
Mg (%)
Mn (ppm)
Fe (ppm)
Zn (ppm)
Cu (ppm)
B (ppm)
2.75–3.25
0.15–0.20
1.30–1.80
0.40–0.90
0.30–0.50
200–350
40–80
8–20
5–40
30–50
Sampling in
September–December
2.60–3.10
0.11–0.15
0.80–1.20
0.40–1.50
0.30–0.50
200–350
40–80
8–20
5–40
30–50
51
commonly separate and radiate out at maturity, exposing the
white pulp. Harvested fruit should be handled with care to
prevent bruising of the skin. Harvesting immature fruit results
in poor quality and a failure to ripen. Mature fruit on the tree
tend to split.
The fruit is climacteric and rapidly ripens within 3–6 days
after harvest. The onset of increased rates of carbon dioxide
and ethylene production occur about 3 days after harvest, but
the respiratory peak appears before the ethylene climacteric or
both occur simultaneously. The optima storage temperature of
sweetsop is between 15 and 20°C (Broughton and Tan, 1979;
Vishnu Prasanna et al., 2000). Fruit stored under low RH
conditions ripen faster and have better taste and appearance
than those stored under high RH.
DISEASES, PESTS AND WEEDS A number of diseases have been
reported (Table A.27). Anthracnose caused by Colletotrichum
gloeosporioides (Glomerella cingulata) is the most serious in
areas of high rainfall and atmospheric humidity and during
the wet season in dry areas (Dhingra et al., 1980). This disease
causes twig dieback, defoliation, dropping of flowers and fruit.
On mature fruit the infection causes black lesions. Another
severe fruit rot disease is attributed to Gliocladium roseum and
affects 20–90% of the fruit in India. Symptoms consist of
water-soaked spots that turn soft and brown.
Black canker (Phomopsis anonacearum) and diplodia rot
(Botryodiplodia theobromae) occur mostly on neglected trees
and cause similar symptoms of purplish to black lesions
resulting in mummified fruit. Marginal leaf scorch and twig
dieback can be also caused by P. anonacearum and B.
theobromae. Diplodia rot has darker internal discoloration and
deeper, more extensive corky rot in fruit. Phytophthora blight
(purple blotch) caused by Phytophthora citrophthora and
Phytophthora nicotianae that mostly infest fruit and leaves
occurs during the wet season (Huang et al., 1991). Fruit may
be mummified and stay on the tree or drop, and the infected
leaves show water-soaking spots that turn brown-black. Pink
disease caused by Corticium salmonicolor mainly infests stems
and causes twig dieback.
Table A.27. Major diseases of sweetsop.
Common name
Organism
Parts affected and symptoms
Country/region
Anthracnose
Colletotrichum gloeosporioides
(Glomerella cingulata)
Fruit, leaves, twigs
Universal
Armillaria root rot
Armillaria leuteobubalina
Roots, base of tree, decline
Australia
Bacterial wilt
Pseudomonas solanacearum
Tree wilt
Universal
Black canker
Phomopsis anonacearum
Fruit, leaves
Universal
Black canker (diplodia rot)
Botryodiplodia theobromae
Fruit, leaves
Universal
Brown root rot
Phellinus noxius
Root, base of tree, decline
Taiwan
Fruit rot
Gliocladium roseum
Fruit
India
Phytophthora blight (purple blotch)
Phytophthora citrophthora and
Phytophthora nicotianae (or
Phytophthora parasitica)
Phytophthora palmivora
P. parasitica
Fruit, leaves; immature fruit may be
mummified and stay on the tree
or drop
Same effects
Same effects
Taiwan
Australia
India
Pink disease
Corticium salmonicolor
Trunk, stem, twig dieback
Taiwan
Rust fungus
Phakopsora cherimoliae
Leaves
Florida
52
Annonaceae
Bacterial wilt is caused by Pseudomonas solanacearum and is
characterized by rapid wilting and death of young trees and
slow decline of old trees. There is a general decline of vigour
and defoliation on affected limbs. Vascular discoloration of
woody tissues occurs in the roots and up to the trunk at
ground level. Brown root rot is caused by Phellinus noxius
mainly infesting the roots and base of the tree and is
characterized by a slow wilting and eventually results in death
of the whole tree.
Some insect pests of sweetsop occur in most growing areas
(Table A.28). The most serious pest is the annona seed borer
(Bephratelloides cubensis) that is widely distributed throughout
the Caribbean and Mexico, Central and northern South
America and is found in Florida. This chalcidoid wasp lays its
eggs on young fruit and the emerging larvae tunnel into the
pulp, causing blackened, necrotic areas. Bagging the fruit is
sometimes done. The annona fruit borer (Cerconota anonella)
is another important pest in the American tropics. The atis
moth borer (Anonaepestis bengalella) is the most serious pest of
sweetsop in Taiwan.
Mealy bugs and various species of scale insects are found
universally and usually become a serious pest on neglected
trees. Red spider mites can become a serious problem in dry
areas or during dry seasons. Yellow tea thrips (Scirtothrips
dorsalis) can damage young shoots, flowers and fruit.
Mature green annona fruit are rarely infested by fruit flies,
but are found occasionally in tree-ripened fruit. Use of bait
sprays and field sanitation are recommended measures to
minimize fruit fly infestation (Smith, 1991). Fruit bagging also
provides protection.
Problem weeds, especially grasses and twining weeds,
should be controlled before planting by cultivation and
herbicides. Young trees should be protected from weed
competition by hand weeding, mulching or contact herbicides.
The shallow root system limits the use of cultivation under the
tree. A translocated herbicide may be needed for perennial
weeds and is applied as a spot spray. Nevertheless, to reduce
the use of herbicides, grass cover is recommended, in which
the whole field is covered with grass except the ground under
canopies.
The origin of most cultivars
is unknown. Existing commercial cultivars show considerable
variation in growth, fruit set, fruit size and quality. No single
cultivar has all the desirable characteristics. The length of the
juvenile period varies with earliest production occurring in 2–3
years and full production in 5–6 years. This juvenile period is
extremely variable with scions on seedling rootstocks. India and
Taiwan have produced a few named cultivars (Table A.29). The
major named cultivars of A. squamosa in Taiwan are ‘Ruan-zhi’,
‘Cu-lin’, ‘Da-mu’, ‘Xi-lin’ and ‘Tai-nong no. 1’. In India,
commercial varieties include ‘Balanagar’, ‘Mammoth’, ‘Arka’,
‘Arka Sahan’, ‘Barbadose Seedling’, ‘Washington’, ‘Red
Sitaphal’ and ‘Purandhar’. ‘Cuban Seedless’ is a seedless
cultivar with medium-sized fruit developed in Cuba; another
Cuban cultivar is low in fibre content. Seedling populations
have been established in Taiwan to select superior lines with
increased yield and improved quality. Early maturity, better
fruit appearance, higher edible flesh ratio, postharvest and
shipping quality and in the subtropics greater cold tolerance
are the most frequent objectives.
Chung Cheng Chen and
Robert E. Paull
MAIN CULTIVARS AND BREEDING
Literature cited and further reading
Anon. (1995) Production and Marketing of Sweetsop. Taitung District
Agricultural Improvement Station, Special Publication No. 1.
Taiwan, 48 pp. (In Chinese)
Bhakuni, D.S., Tewari, S. and Dahr, M.M. (1972) Aporphine
alkaloids of Annona squamosa. Phytochemistry 11, 1819–1822.
Bhaumik, P.K., Mukherjee, B., Juneau, J.P., Bhacca, N.S. and
Mukherjee, R. (1979) Alkaloids from leaves of Annona squamosa.
Phytochemistry 18, 1584–1586.
Broughton, W.J. and Tan, G. (1979) Storage conditions and ripening
of the custard apple Annona squamosa L. Scientia Horticulturae 10,
73–82.
Campbell, C.W. (1985) Cultivation of fruits of the Annonaceae in
Florida. Proceedings of the American Society for Horticultural
Science Tropical Region 29, 68–70.
Chang, M.S. (2000) Orchard soil and fertilization management. In:
Sweetsop Cultivation Manual. Taitung District Agricultural
Improvement Station, Taiwan, pp. 33–48. (In Chinese)
Table A.28. Major insect pests of sweetsop.
Common name
Organism
Parts affected
Country/region
Ambrosia beetles
Many species in the Scolytidae family
Twigs, branches, trunk
Florida
Annona fruit borer
Cerconota anonella
Fruit
Florida, Caribbean, American tropics
Annona seed borer
Bephratelloides cubensis
Seeds (fruit)
Florida, Caribbean, American tropics
Atis moth borer
Anonaepestis bengalella
Fruit
Taiwan, Philippines, India
Fruit fly
Dacus zonatus Saunders
Fruit
India
Mealy bugs
Pseudococcus maritimus and Pseudococcus
calceolariae, Planococcus pacificus, Planococcus citri,
Ferrisia virgata, Pesudococcus chiponensis,
Pseudococcus virgatis, Pseudococcus lilacinus
Stems, leaves, fruit
Florida, Taiwan, Australia, India
Red spider mite
Oligonychus coffeae, Oligonychus mangiferus and
several other genera and species
Leaves, flowers
American tropics, Taiwan
Scale insects
Aspidiotus destructor, Chrysomphalus ficus,
Philephedra tuberculosa, Ceroplastes floridensis
Leaves, twigs
Caribbean, Taiwan, Florida, India
Thrips
Scirtothrips dorsalis
Twigs, flowers, fruit
Taiwan
Annona
Table A.29. Some selected cultivars of sweetsop.
Country/region
Name
Taiwan
‘Ruan-zhi’, ‘Cu-lin’, ‘Da-mu’, ‘Xi-lin’, ‘Tai-nong no. 1’
India
‘Balanagar’, ‘Mammoth’, ‘Arka’, ‘Arka Sahan’, ‘Barbadose
Seedling’, ‘Washington’, ‘Red Sitaphal’, ‘Purandhar’
Thailand
‘Fai Kaew’, ‘Fai Krung’, ‘Nang Kaew’, ‘Nang Sir Krung’,
‘Nang Thong’
Florida
‘Lessard’, ‘Kampong Mauve’, ‘Red Sugar’, ‘Cuban Seedless’
Egypt
‘Abd El-Razik’
Dhingra, J., Mehrotra, R.S. and Aneja, I.R. (1980) A new postharvest
disease of Annona squamosa L. Current Science 49, 477–478.
Enweani, I.B., Esebelahie, N.O., Obroku, J. and Obi, L.C. (1998) Use
of soursop and sweetsop juice in the management of diarrhoea in
children. Journal of Diarrhoeal Disease Research 16, 252–253.
George, A.P. and Nissen, R.J. (1987) Propagation of Annona species:
a review. Scientia Horticulturae 33, 75–85.
George, A.P., Nissen, R.J. and Campbell, J.A. (1992) Pollination and
selection in Annona species (cherimoya, atemoya, and sugar
apple). Acta Horticulturae 321, 178–185.
Higuchi, H., Utsunomiya, N. and Sakuratani, T. (1998) Effects of
temperature on growth, dry matter production and CO2
assimilation in cherimoya (Annona cherimola Mill.) and sugar
apple (Annona squamosa L.) seedlings. Scientia Horticulturae 73,
89–97.
Hopp, D.C., Zeng, L., Gu, Z. and McLaughlin, J.L. (1996)
Squamotacin: an annonaceous acetogenin with cytotoxic
selectivity for the human prostate tumor cell line (PC-3). Journal
of Natural Products 59, 97–99.
Huang, T.C., Liu, L.H. and Lee, H.L. (1991) Phytophthora fruit rot
of Annona squamosa L. caused by Phytophthora citrophthora and P.
nicotianae. Plant Protection Bulletin 33, 103–112.
Huang, T.C., Shieh, J.L. and Lee, H.L. (2000) Ecology and control
of major diseases and insect pests. In: Sweetsop Cultivation
Manual. Taitung District Agricultural Improvement Station,
Taiwan, pp. 59–77. (In Chinese)
Kumar, R., Hoda, M.N. and Singh, D.K. (1977) Studies on the floral
biology of custard apple (Annona squamosa Linn.). Indian Journal
of Horticulture 34, 252–256.
Leboeuf, M., Cavé, A., Bhaumik, P.K., Mukherjee, B. and
Mukherjee, R. (1982) The phytochemistry of the Annonaceae.
Phytochemistry 21, 2783–2813.
Lemos, E.E.P. and Blake, J. (1996) Micropropagation of juvenile and
adult Annona squamosa. Plant Cell, Tissue and Organ Culture 46,
77–79.
Lin, C.H. (2000) Common nutrient disorders. In: Sweetsop
Cultivation Manual. Taitung District Agricultural Improvement
Station, Taiwan, pp. 49–58. (In Chinese)
Lo, S.S. (1987) Pruning technique, use of sprouting chemical and
flower initiation in sugar apple (Annona squamosa). In: Chang,
L.R. (ed.) Proceedings of a Symposium on Forcing Culture of
Horticultural Crops. Taichung District Agricultural Improvement
Station, Special Publication No. 10, Taiwan, pp. 147–150. (In
Chinese, English summary)
Marler, T.E. and Zozor, Y. (1996) Salinity influences photosynthetic
characteristics, water relations, and foliar mineral composition of
Annona squamosa L. Journal of the American Society for
Horticultural Science 121, 243–248.
53
Marler, J.E., George, A.P., Nissen, R.J. and Andersen, P.J. (1994)
Miscellaneous tropical fruits – annonas. In: Schaffer, B.C. and
Andersen, P.C. (eds) Handbook of Environmental Physiology of
Fruit Crops. Vol. II. Subtropical and Tropical Crops. CRC Press,
Boca Raton, Florida, pp. 200–206.
Nadel, H. and Peña, J.E. (1994) Identity, behaviour, and efficacy of
nitidulid beetles (Coleoptera: Nitidulidae) pollinating commercial
Annona species in Florida. Environmental Entomology 23,
878–886.
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J.H. (1999) Influence of flooding on net CO2 assimilation, growth
and stem anatomy of Annona species. Annals of Botany 84,
771–780.
Pal, D.K. and Sampath Kumar, P. (1995) Changes in the physicochemical and biochemical compositions of custard apple (Annona
squamosa L.) fruits during growth, development and ripening.
Journal of Horticulture Science 70, 569–572.
Saavedra, E. (1977) Influence of pollen grain stage at the time of
hand pollination as a factor on fruit set of cherimoya. HortScience
12, 117–118.
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Annona squamosa L. (Annonaceae). Annals of Botany 54, 153–167.
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54
Annonaceae
Annona squamosa ⫻ Annona cherimola
atemoya
Table A.30. Proximate fruit composition of atemoya per 100 g
(Source: Leung and Flores, 1961).
Atemoya (A. squamosa ⫻ A. cherimola, Annonaceae) is a hybrid
in the custard apple family, consisting of about 120 genera,
widely distributed. All are native to the American tropics and
subtropics. Some are grown as ornamentals while others are
known for their edible fruit and perfume. Three major species
of Annona are grown commercially: the cherimoya (Annona
cherimola Miller), sugar apple (Annona squamosa L.) and
atemoya or custard apple (Annona spp. hybrids) (George et al.,
1987). The cherimoya is indigenous to the tropical highlands
of Peru and Ecuador, while the sugar apple is widely
distributed throughout tropical South America (George and
Nissen, 1985a, b). Most atemoya hybrids resemble cherimoya
in vigour and tree habit, but exhibit flowering and fruiting
characteristics intermediate to both parents.
Proximate
World production
In the late 1990s, atemoya production was significant in
Australia (3000 t), Israel (500 t), Florida (200 t) and Hawaii
(50 t). In comparison, cherimoya production was significant in
Spain (15,000 t), Bolivia (6000 t), Chile (2300 t) and Peru (400
t) while sugar apple production was significant in Thailand
(75,000 t) and the Philippines (6000 t) (George and Nissen,
1992a; George et al., 1997). Chile exports 1000 t of cherimoya
annually. Yields of 80–150 kg/tree (15–25 t/ha) have been
recorded for mature cherimoya and custard apple trees
(George and Nissen, 1986a, 1992a). In Australia, the annual
production of atemoya is expected to double in the next 10
years due to the planting of new, higher yielding cultivars with
exceptional eating quality (George et al., 1999).
Uses and nutritional composition
Atemoyas are usually consumed as dessert fruit. Because seeds
of the fruit are interspersed throughout the flesh, cultivars
containing few seeds are the most desirable. The perishable
nature and supply shortage limits the market area or makes air
shipment essential. The fruit is rich in starch when firm but
increases markedly in sugar as it softens. The main sugars are
glucose and fructose (80–90%). Compared with other fruit,
atemoya fruit contain significant quantities of vitamin C,
thiamine, potassium, magnesium and dietary fibre (Table A.30).
The calorific value is high (> 300 kJ/100 g) and is almost double
that of peach, orange and apple (George and Nissen, 1993).
Botany
AND NOMENCLATURE The Annonaceae (120
genera) is considered a ‘primitive’ member of the Magnoliidae.
The genus Annona is the most important in the Annonaceae,
since among its 100 or more species, seven species and one
hybrid are grown commercially. Atemoya, commonly called
custard apple in Australia, is a hybrid between A. squamosa and
A. cherimola. P.J. Wester of Florida produced the first hybrids
in 1908 and called it the ‘atemoya’, by using the Brazilian
name ‘ate’ for sweetsop and ‘moya’ from cherimoya. In 1927,
hybrids were also developed in Poona, India. It should be
noted that common names, such as custard apple, have been
used with reference to many different species of Annona.
TAXONOMY
Edible portion
Water
Energy (kJ)
Protein
Lipid
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
%
70
71.5–78.7
310–394
1.1–1.4
0.4–0.6
18.1
0.05–2.5
0.4–0.75
mg
17
0.3
32
40
250
4.5
mg
50
0.05
0.07
0.8
–
Cherimoya trees are 3–10 m tall whereas sugar
apple is a shrub of 3–6 m. Atemoya trees are hybrids,
morphologically intermediate between the above two species
and vary in height from 3 to 10 m. All three species are semideciduous in growth habit. The tropical species, sugar apple
sheds its leaves in the dry season, whereas cherimoya and
custard apple from the subtropics shed their leaves in spring.
Leaves are alternate, simple and entire. Vegetative buds are
often subpetiolar (Fig. A.6), consequently natural leaf
abscission or artificial defoliation is necessary prior to
emergence of new growth flushes (George and Nissen, 1987a).
The Annona flower is hermaphrodite and exhibits
protogynous dichogamy (George and Nissen, 1992a). The
flowers are two series of three petals, and the outer series are
thick and fleshy. With cherimoya and atemoya, floral buds can
be initiated on 1-year and older wood but most are produced
in synchronization with the current season’s shoot growth
(Moncur, 1988; Higuchi and Utsunomiya, 1999). Annona spp.
generally require 27–35 days for flower bud development from
initiation to anthesis. Differences in floral behaviour in the
various areas may be attributed to both genetic variability and
climatic differences (Kshirsagar et al., 1976). Flowering can
extend from 3 to 6 months, with normally between two and
three peaks. For example, in Australia, atemoyas produce the
first flowers during early summer with a second flowering
during the late summer or early autumn.
Fruit are a pseudocarp formed by the fusion of the carpels
and the receptacle into a fleshy mass. Fruit shape is highly
variable ranging from spheroid to ovoid with the fruit surface
covered with U-shaped areoles, which can be smooth or
pointed. The white pulp is easily separable from the seeds and
is very popular, especially in the Americas.
DESCRIPTION
Annona
ECOLOGY AND CLIMATIC REQUIREMENTS The cherimoya is
indigenous to the tropical highlands of Peru and Ecuador,
while the sugar apple is widely distributed throughout tropical
South America. Most hybrids resemble cherimoya in vigour
and tree habit, but exhibit flowering and fruiting
characteristics intermediate to both parents. All three species
are now widely distributed throughout the world.
Both atemoya and cherimoya are best grown in frost-free
locations as young trees are killed at ⫺1°C and mature trees at
⫺3°C (George and Nissen, 1985a, 1992a). Cherimoya is more
cold tolerant than custard apple and can withstand a longer
duration at ⫺3°C. Sugar apple, on the other hand, is frost
sensitive. In the cool subtropics, cherimoya rootstocks, because
of their ability to grow at lower soil and air temperatures than
sugar apple, may increase tree growth and productivity.
Conversely, in warm subtropical areas, trees may be vigorous
but unproductive.
Trees of atemoya are semi-deciduous and are dormant in
late winter and early spring. This dormancy or rest period
enables the tree to avoid frost or drought. Cherimoya appears
to initiate growth at 7°C compared with 10°C for custard
apple (George and Nissen, 1985a, 1987a). Budbreak of most
atemoya normally occurs in late spring, after a winter
dormancy. In warm areas, chemical defoliation can be used to
advance budbreak by 4–6 weeks (George and Nissen, 1987a).
This treatment does not work in atemoya at < 12°C because
shoot growth is erratic (George and Nissen, 1986a, b).
Provided insect pollinators are present and environmental
conditions are conducive for set, defoliation has the potential
to increase early fruit production in warm regions.
In glasshouse studies, 32°C days/27°C nights, used to
simulate tropical conditions, induced strong vegetative
flushing and reduced floral production in atemoya (George
and Nissen, 1987b). In warm areas such as Florida, Hawaii
and Queensland, strong vegetative flushing occurs in
midsummer during flowering and may reduce fruit set unless
trees are propagated on dwarfing rootstocks or interstocks
(George and Nissen, 2002a). Under cooler conditions in
Chile, California and New Zealand, trees are less vigorous.
Temperatures between 22–28°C during flowering period are
ideal for fruit set in atemoya (George and Nissen, 1985a)
whereas 18–28°C is more suitable for cherimoya (Saavedra,
1977). Flower opening and anther dehiscence are both
advanced with increasing temperature, however at > 28°C the
stigmas desiccate before the anthers dehisce (George and
Nissen, 1988). In controlled-environment studies, relative
humidity (RH) of > 95% and < 60% severely reduced fruit
set (George and Nissen, 1988). The adverse effects of high RH
appear to be due to changes in stigmatic secretions, which
restrict pollen germination and pollen tube growth. At low
RH, desiccation of the stigma may occur before anthers
dehisce as most flowers are at the female stage in the early
morning and the anthers only release pollen in the afternoon
(Saavedra, 1977). Studies in Chile (Saavedra, 1977) and Egypt
(Ahmed, 1936) showed that spraying flowers with water or
inserting a drop of water into the flower at anthesis could
increase fruit set about sixfold. Although stigmas appear to
require constant, moderately high RH, a diurnal change
appears to be necessary for anther sacs to split (Kshirsagar et
al., 1976). As flowering of Annona spp. occurs during summer
55
in the subtropics, RH levels below 30% are common. Late
afternoon rain or under-tree irrigation can increase fruit set
perhaps by causing condensation of water droplets within the
fleshy petals of the flower (George and Nissen, 1985b). In
contrast, light rain or continuous daily sprinkling are
detrimental possibly because they dilute the floral scents
which attract pollinators.
Excessively low temperatures may retard fruit maturation
while excessively high temperatures may cause premature
ripening and fermentation (George and Nissen, 1985a, 1992a).
As expected, cultivars appear to vary in this regard.
Physiological disorders such as russetting also appear to be
more prevalent when temperatures fall below 13°C (George
and Nissen, 1985a).
Atemoya is capable of growing in a wide range of soil types
from sandy soil to clay loams. Higher yields occur on welldrained sandy to sandy loam soils. Drainage is essential to
avoid root rot diseases.
Moderate drought (L of ⫺1.5 Mpa) has been shown to
reduce shoot growth by about 20–30% and increase the
number of flowers per lateral by about 40% compared with
well-watered controls, due to reduced apical dominance and
increased lateral branching (George and Nissen, 2002a).
However, severe drought (L of ⫺2.0 MPa) reduced flowering
and fruit set by 30% in trees growing at 28°C and moderately
high vapour pressure deficity (VPD) of 1.2 kPa (George and
Nissen, 1988). Drought has been shown to reduce fruit size by
about 10% due to reduced stomatal conductance (gS) and
carbon dioxide assimilation (George and Nissen, 1992b,
2002a). In controlled-environment glasshouses there was a
continuous but decreasing decline in gS over a range of L
from ⫺0.8 MPa to ⫺4.0 MPa at high RH of 90% (George et
al., 1990), whereas at 60% RH, gS was extremely low,
irrespective of the tree water status. The stomata of atemoya
are thus more sensitive to RH than L.
Overall, atemoya benefits from uniform soil moisture
during the fruit development period with extremes of
moisture lowering production. The adverse effects of drought
on productivity can be minimized by maintaining soil water
potentials (S) > ⫺20 kPa. Bearing atemoya trees may need up
to 1440 l/tree every 4 weeks during slow growth phase and
from 500 to 750 l/tree every 3–5 days during flowering fruit
set and fruit growth (Sanewski, 1991). Reducing irrigation in
late winter to force atemoya trees into dormancy for 1 or 2
months in spring is recommended in Australia and California,
respectively. The amount and frequency of irrigation must be
determined by experience in any particular location and soil
type. Water stress should be prevented during flowering, fruit
set and fruit development, as fruit are more sensitive than
leaves.
Heavy shading of vigorous trees can reduce fruit set in
atemoya (Marler et al., 1994). Light penetration to the base of
vigorous trees with a dense canopy in a close spacing can be
2% of full sunlight and there is very little fruit set. Pruning
practices and spacing need to be adjusted for this growth
aspect. No photoperiod responses have been reported.
Trees are susceptible to wind damage and limb breakage.
Tree shaking may also be partially responsible for penetration
and infection by collar rot organisms. The fruit skin is easily
damaged by rubbing and exposure to drying winds (Marler et
56
Annonaceae
al., 1994). Productivity can be improved by windbreaks and
under-tree sprinklers to raise the RH above 60%.
REPRODUCTIVE BIOLOGY The flowers exhibit protogynous
dichogomy. Dichogamy appears to be the main factor limiting
self-pollination and fruit set in Annona and poses a serious
problem in obtaining high yields. The atemoya female parts
are receptive between 4 and 8 a.m. and appear moist and
sticky (Thakur and Singh, 1964). The pollen is discharged in
the afternoon of the same day from 3 to 6 p.m. if the RH is
above 80% and temperature > 22°C. At lower temperatures
pollen is released on the afternoon of the second day. The
pollen sacs turn a greyish colour as pollen is discharged. Upon
opening, flowers are receptive for about 24 h. The flowering
seasons of A. squamosa and A. cherimola coincide. When
sweetsop pollen is shed at about 2 a.m., cherimoya flowers are
receptive, opening around 7 to 9 a.m. and when cherimoya
pollen is shed at 3 to 4 p.m., sweetsop flowers are receptive.
This flower synchrony together with complementary
functional sexes favours cross-pollination leading to natural
hybridization. This is attested to by the frequent appearance of
hybrid seedlings under the trees of sweetsop and cherimoya
when grown in close proximity.
Beetles of the Nitidulidae family are the main insect
pollinators of Annona flowers. Nitidulid beetles (Carpophilus
and Uroporus spp.) are the important pollinators of Annona
flowers with wind and self-pollination being low (1.5%). Fruit
set of ‘African Pride’ atemoya increases linearly with
increasing numbers of nitidulid beetles per flower (George et
al., 1992). Three or more beetles per flower increased fruit set
to nearly 25%. Studies also showed that these beetles breed
rapidly in rotting fruit media and that populations of these
beetles can be increased by maintaining the rotting fruit
attractant. Alternatively, fruit set may be increased by using
pheromone bait stations (Pena, 2002).
Pollen early in a flowering season has thick walls, is high in
starch, germinates poorly and gives poor fruit set. Pollen of
later flowers shows a high proportion of individual pollen
grains without starch grains that germinate well. Hand
pollination can increase fruit set in excess of 40% (Schroeder,
1943) and is frequently practised, improving fruit set and fruit
shape. Pollen must be collected in the evening from fully open
flowers, when the sacs have turned from white to cream. The
flowers are held in a paper bag, not a closed container, and
should discharge that afternoon. The flowers are shaken over a
shallow tray or paper to collect the pollen that is transferred to
a small container and held in the refrigerator for use the next
morning. Enough pollen to pollinate 50 to 60 flowers can be
obtained from 20–30 flowers. Pollination is done before 7 a.m.
every few days during the flowering period using a small brush
or puffer. Hand pollination has shown some variable results
and is less successful on very humid, overcast days and on
young vigorous trees. About 150 flowers can be pollinated in
an hour and a success rate of 80–100% can be achieved.
Hand pollination in commercial orchards is tedious, time
consuming and a costly practice. Attempts have been made to
use growth regulators, with considerable variation in the
results obtained. Auxin (indole acetic acid (IAA), napththalene
acetic acid (NAA)) induces growth very slowly with less fruit
drop, while gibberellic acid-3 (GA3) promotes fruit set and
growth rate, however, the effects are short lived and repeated
applications are needed (Yang, 1988). Applications of the two
substances separately at appropriate times have produced
seedless fruit between 200 and 300 g in size (Saavedra, 1977).
In Japan, four sequential applications of gibberellin (GA3,
1500 ppm) produced near seedless fruit of 300–400 g in
selected cherimoya cultivars (Yonemoto et al., 2000a, b).
Repeated spraying is necessary to prevent fruit abscission
during the first 2 months. Seedless fruit are generally smaller,
with less flavour and less fruit splitting than occurs in seedy
fruit that result from pollination (Yang, 1988). Spraying with
GA3 is not recommended as a general management practice
for atemoya because of variable results, though it could be
used in areas with poor natural pollination.
FRUIT DEVELOPMENT Fruit exhibits a two-stage growth
pattern with maturation occurring 16 to 26 weeks after fruit
set, depending upon species and growing conditions (George,
2000). Low humidity (< 60% RH) and temperature (< 13°C)
near fruit maturity can delay fruit maturation whereas very
high temperatures (> 35°C) can cause premature fruit
ripening and fermentation of the fruit. In Australia, due to
climatic differences between regions within the country, fruit
may be harvested over a period of 8 months.
Horticulture
PROPAGATION Atemoya is normally propagated by grafting
(Table A.31). Varieties are very difficult to propagate by
cuttings or layering. There are cultivar differences in rooting
ability of atemoya, with ‘African Pride’ having a higher rooting
response (15%) than ‘Pink’s Mammoth’ and cherimoya
(<5%). Time of cutting removal is crucial for success; cuttings
taken at the end of the cool season have a higher rooting rate.
Roots should occur in 8–12 weeks and are ready to pot in
16–20 weeks. Recent research has shown softwood cuttings to
give a higher strike than hardwood cuttings (Nissen, personal
communication, 2003).
Air layering can be used with some cultivars, though
cherimoya are not propagated easily by this method. A
modification where the new shoot is clamped and only the
shoot tip is exposed is successful. Inarching of atemoya to
Annona reticulata rootstock has been successful. Although
Table A.31. Rootstock and scion compatibility of atemoya (Source:
Sanewski, 1991).
Sciona
Rootstock
Atemoya
Atemoya
Gefner
Pink’s Mammoth
Page/Bradley
Annona cherimola
Annona glabra
Annona muricata
Annona palustris
Annona reticulata
Annona squamosa
C
aC,
Gefner
Pink’s Mammoth
Page/Bradley
C
C
C
N
N
–
N
C
P
C
C
C
C?
P
C
C?
compatible; P, partially compatible; N, not compatible; –, unknown.
Annona
inarching has given good results, it is time consuming and
costly for large-scale propagation.
Grafting is superior to budding in percentage takes and
subsequent growth, with whip graft and cleft graft techniques
giving the best results (Duarte et al., 1999). The branches
should be defoliated 1–2 weeks before scionwood is cut to
induce bud swelling. T-budding and chip budding methods
are sometimes successful. There are considerable graft
incompatibilities among Annona and Rollinia spp. and types.
Transplanting is best done when the trees are dormant. In
the subtropics, planting should not occur if there is a risk of
frost. Plants should have attained a height of 30–46 cm at
transplanting time with the union of grafted or budded plants
placed 15 cm or so above the ground. Trees should be irrigated
as soon as possible after transplanting, with wind and sun
guards sometimes required.
In Australia and Florida, depending on rootstock, atemoya
trees are planted 4–6 m apart within the row with 6–8 m
between rows (Campbell, 1985). Narrower tree spacing is used
for ‘African Pride’ on A. squamosa rootstock or interstock and
the widest spacing is used for ‘Pink’s Mammoth’ on A.
cherimola. More recently, with new, exceptionally highyielding cultivars such as ‘KJ Pinks’ trained onto the open
Tatura system, higher planting densities have been made
possible.
Cherimoya has been found to be a vigorous
rootstock for atemoya. Atemoya is not compatible with Annona
glabra, Annona muricata and A. reticulata as rootstocks
(Sanewski, 1991) (Table A.31). This is complicated by cultivar
differences in compatibility with common rootstocks. Atemoya
cultivars ‘Bradley’ and ‘Page’ are compatible with custard
apple rootstocks but ‘Gefner’ shows partial incompatibility
with the same rootstock. With atemoya, George and Nissen
(2002b) showed that tree size could be reduced by half using
sugar apple rootstocks, but sugar apple was found to be
susceptible to bacterial wilt, caused by Pseudomonas
solanacearum, with up to 30% of trees dying within 6 years
after planting. Consequently, cherimoya, which is only mildly
susceptible to bacterial wilt, has been the preferred rootstock
of choice in Australia.
ROOTSTOCKS
PRUNING AND TRAINING Training of trees begins in the
nursery with pruning continuing after transplanting. It is
desirable to train the tree to a single trunk up to a height of
about 90 cm and then headed back to produce four to six main
branches or sub-leaders. These branches should be spaced
15–25 cm above each other and be allowed to grow in different
directions to develop a good scaffold. After about 2 m, they
could be left to natural growth. Pruning is carried out when
the trees are dormant and in heavy trees involves removal of
lower limbs touching the ground and branches in the centre
where branches may be rubbing against each other. The
objective is to allow sunlight access to the centre of the tree.
More recently, with the development of exceptionally highyielding varieties with lower tree vigour, trees are now being
trained onto an open Tatura or Y trellis system. These systems
allow for greater light interception and overcome the problems
of deep shading with goblet-trained trees.
With atemoya, all lateral buds can have up to two vegetative
57
buds and three flower buds. The lateral buds are normally
‘buried’ (subpetiolar) in the base of the swollen leaf petiole
(Fig. A.6). Leaf shed must occur prior to the elongation of
‘buried’ buds (George and Nissen, 1987a). During spring, just
prior to budbreak, removal of leaves mechanically by stripping
or chemically with urea or ethephon, followed by the
application of rest-breaking chemicals, can help to release
these buds from dormancy (Sanewski, 1991; George et al.,
2002b). Dormant pruning is also normally carried out in late
winter or early spring, just before budbreak. At this time,
laterals are moderately pruned as George et al. (2001) have
shown that severe stub pruning of laterals to < 20 cm long is
detrimental to yield and fruit quality. This pruning strategy
aims to produce one or two fruit on new season laterals. The
ideal seasonal growth of laterals producing fruit is about 60 cm
long and about 10 mm in diameter at the base. Annual lateral
growth more than 60 cm is considered excessive for mature
fruiting trees, and increases the severity of internal fruit
disorders (George et al., 2002b). Vigorous trees are also
summer pruned, which involves shoot tipping and usually leaf
stripping to force the subpetiolar buds into lateral growth
concomitantly with late flowering.
NUTRITION AND FERTILIZATION The Annonas spp. have an
indeterminate growth habit (axillary flowering) and applying
nitrogen in somewhat excessive amounts does not interfere
greatly with floral initiation, as is the case with plants having a
determinate growth habit. However, excessive tree vigour is
usually associated with reduced flowering, yield and fruit
quality in many tree crops and the atemoyas are no exception
(George et al., 1989).
In Australia, continued research and field observations of
atemoya nutrition (Sanewski, 1991; George et al., 2002c) have
led to greater refinements in terms of quantity of fertilizer and
times of incremental applications during the annual growth
and fruiting cycles. After 10 years of age, the annual amounts
of nitrogen, phosphorus and potassium (NPK) remain the
same, as tree size is kept relatively constant by annual pruning
and competition from adjacent trees. The annual requirements
of nitrogen and potassium are split into four increments
(Table A.32). In the cool subtropical areas, greatest vegetative
growth takes place during the warmer months from spring to
autumn. Reduction in nitrogen during the winter minimizes
new vegetative growth in young trees that are vulnerable to
cold temperatures. This adjustment is not necessary in the
warm tropics. There is one application of phosphorus per year,
during the early autumn.
The use of foliar nutrient analysis has become a useful
management tool in determining atemoya fertilizer
programmes (Table A.33). Sampling for foliar analysis consists
of obtaining the most recently matured leaf; the fourth or fifth
leaf below the growing point. Sample leaves are selected from
non-bearing shoots without a leaf flush, during late summer or
early autumn.
The primary sink for potassium in the atemoya is the fruit,
rather than the leaves and thus there is a high requirement,
with deficiency likely. About 60% of the potassium
requirement is applied during the fruit development period.
Atemoyas also have a fairly high requirement for magnesium
and calcium. Heavy vegetative growth during the fruit
58
Annonaceae
Table A.32. A guide to annual application of nitrogen, phosphorus and potassium (NPK) for ‘Pink’s Mammoth’ atemoya trees of
different ages using straight fertilizers (g/tree/year) and percentage distribution of application of annual amounts (Source: Sanewski,
1991).
Tree age (years)
2
4
6
8
10
Fertilizer
Urea (%)
Superphosphate (%)
Potassium chloride (%)
Urea (g/tree/year)
Superphosphate (g/tree/year)
Potassium chloride (g/tree/year)
400
860
1300
1600
1750
500
550
780
880
880
360
930
1170
1500
1650
Early spring
Early summer
Early autumn
Late autumn
20
30
10
10
30
40
100
40
Table A.33. Tentative leaf nutrient standards for atemoya in Queensland,
Australia presented as a guide (Source: George et al., 2002).
Nutrient
Nitrogen (%)
Phosphorus (%)
Potassium (%)
Calcium (%)
Magnesium (%)
Sodium (%)
Chloride (%)
Manganese (ppm)
Copper (ppm)
Zinc (ppm)
Iron (ppm)
Boron (ppm)
Acceptable range
2.4–3.2
0.15–0.21
1.0–1.5
1.0–1.6
0.35–0.4
< 0.02
< 0.3
60–140
10–20
40–70
50–110
18–30
development period competes for nutrients such as calcium
and boron. Calcium-deficient fruit develop hard, brown,
lumpy tissues around the central core. Deficiency in calcium
and boron are considered as causal factors for these lumps
(Cresswell and Sanewski, 1991; George et al., 2002d).
Applying excessive boron can be phytotoxic, especially in
sandy soils. A desirable practice is to use organic fertilizers
with inorganic fertilizer as a supplement to maintain a balance
and to control cropping (Sanewski, 1991).
HANDLING AND STORAGE Atemoyas are
harvested every 3–7 days with experienced pickers harvesting
from 150 to 180 kg of fruit/h. Heart-shaped fruit are
preferred with a smooth cherimoya-like skin, instead of the
bumpy sweetsop skin type. Besides shape, size (200–500 g)
and skin texture, the fruit should be free of blemishes and
mechanical injury that can lead to skin blackening. The fruit
skin colour changes from darker to lighter green and can be
greenish yellow at harvest. During ripening of this climacteric
fruit, the skin darkens further and splitting occurs. Harvested
fruit should be handled with care to prevent bruising of the
skin. This is especially important for fruit that is marketed for
fresh consumption. Australian standards specify mature
atemoya fruit 75 mm in diameter, firm with ‘creaming’
between segments on the skin. Containers are about 7 kg in
size, fibreboard or polystyrene (450 ⫻ 215 ⫻ 180 mm), well
ventilated and marked with ‘custard apple’ and the number of
POSTHARVEST
20
fruit. Foam sleeves or paper wrapping are used to minimize
damage. The presence of soft fruit and even one fruit-flydamaged fruit can lead to rejection of the consignment.
During ripening, skin splitting occurs and the skin darkens.
Fruit are stored at 10–13°C and 90–95% RH. Atemoya is
sensitive to chilling injury and shows skin darkening and loss
of aroma and flavour. Ethylene production is high (100–300
l/kg/h at 20°C) and ripening is accelerated by exposure to
100 ppm ethylene for 24 h. Respiration rate at 20°C is 40–460
mg carbon dioxide/kg/h.
DISEASES,
PESTS AND WEEDS Black canker (Phomopsis
anonacearum) and diplodia rot (Botryodiplodia theobromae)
occur mostly on neglected trees and cause similar symptoms
of purplish to black lesions resulting in mummified fruit
(Table A.34). Marginal leaf scorch is also caused by P.
anonacearum and B. theobromae and causes twig dieback.
Diplodia rot has darker internal discoloration and deeper,
more extensive corky rot in fruit. Fruit and leaf spot may also
be caused by a soil-borne fungus, Cylindrocladium colhounii
(Hutton, 1999) which can cause almost total loss of fruit
during years of persistent heavy rains. Symptoms begin with
small dark spots primarily on the shoulders of the fruit that
spread along the sides, enlarge, become dry and crack.
Infection is skin-deep but fruit becomes unmarketable. The
control measures recommended are good orchard maintenance
with heavy mulching and lower branch pruning to prevent
splashing of soil during heavy rainfall (Sanewski, 1991). A new
fruit spotting disease (Pseudocercospora spp.) has been
identified in recent years (Hutton, 1999). Symptoms are small
grey spots 1–5 mm in diameter. It develops under wet
conditions and is spread by the wind.
Bacterial wilt of atemoya is caused by Pseudomonas
solanacearum and is characterized by rapid wilting and death
of young trees and slow decline of old trees. There is a general
decline of vigour and defoliation on affected limbs. Vascular
discoloration of woody tissues occurs in the roots and up to
the trunk at ground level. It has caused up to 70% of tree
deaths in 12 years in orchards using A. squamosa rootstocks in
Queensland.
One of the most serious insect pests in Trinidad is the
cerconota moth (Cerconota anonella) that lays its eggs on
young fruit (Table A.35). The emerging larvae tunnel into the
pulp, causing blackened, necrotic areas. It is common to find
Annona
59
Table A.34. Major diseases of atemoya.
Common name
Organism
Parts affected and symptoms
Country/region
Anthracnose
Colletrotrichum gloeosporioides
(Glomerella)
Flowers, fruit, leaves, dieback,
seedling damping off
Universal
Armillaria root rot
Armillaria leuteobubalina
Roots, base of trees, decline
Australia
Bacterial wilt
Pseudomonas solanacearum
Tree wilt
Australia
Black canker (diplodia rot)
Botryodiplodia theobromae
Leaf scorch, hard black lumps on surface
of fruit, twig dieback
Australia
Black canker
Phomopsis anonacearum
Same effects
Australia
Purple blotch
Phytophthora palmivora
Spots on immature fruit, fruit drop, twig dieback
Australia
Cylindrocladium fruit rot
Cylindrocladium colhounii
Small spots and blotches on the fruit
Australia
Pseudocercospora fruit spot
Pseudocercospora spp.
Purplish-grey spots on the fruit
Australia
Rust fungus
Phakopsora cherimoliae
Leaves
Florida
Fruit rot
Glioclacium roseum
Fruit
India
Table A.35. Major insect pests of atemoya.
Common name
Organism
Parts affected
Country/region
Bephrata wasp (soursop wasp)
Bephrata maculicollis
Fruit
Mexico, Americas, Trinidad, Surinam
Wasp
Bephratelloides paraguayensis
Fruit
Americas, Barbados
Cerconota moth (soursop moth)
Cerconota anonella
Fruit
Americans, Trinidad, Surinam
Thecla moth
Thecla ortygnus
Flower, young fruit
Americas, Caribbean
Banana spotting
Amblypelta lutescens
Young fruit
Queensland
Mealy bug
Dysmicoccus
Stem, leaves
Universal
Citrus mealy bug
Planocuccus citri
Fruit
Queensland
Southern stink bug
Nezara viridula
Fruit
Caribbean
Caribbean fruit fly
Anastrepha suspensa
Fruit
Caribbean, Mexico
Queensland fruit fly
Dacus tryoni
Fruit
Australia
Potato leaf hopper
Empoasca fabae
Leaves
Caribbean
Red spider mite
Several genera, species
Leaves, flowers
American tropics
Scale insects
Saissetia coffeae
Leaves, stem
Universal
Coconut scale
Aspidiotus destructor, other genera and species
Leaves, stem
Caribbean
every fruit larger than 7.5 cm infested. Bagging the fruit is
sometimes done. This moth has been reported in the
American tropics as far south as Brazil and is a major limiting
factor in Surinam. The bephrata wasp (Bephrata maculicollis)
is widely distributed throughout the Caribbean and Mexico,
Central and northern South America. This wasp is considered
to be the most important pest in Florida (Campbell, 1985).
The larvae infest the seeds and cause damage to the pulp as
they bore through the flesh to emerge when the fruit matures.
The thecla moth (Thecla ortygnus) is widespread through parts
of the Caribbean and in the American tropics but it is not
considered to be as serious as the cerconota moth and
bephrata wasp. Primary damage is done to the flowers. The
larvae feed on flower parts such as the perianth, stamen and
stigmas with the flowers failing to set fruit. The banana
spotting bug (Amblypelta lutescens) and the fruit spotting bug
(Amblypelta nitida) are considered to be serious atemoya pests
(Waite and Huwer, 1998). The banana spotting bug is reported
to be confined to northern Queensland with both being found
in southern Queensland. The bugs cause small black 2–10 mm
spots on the shoulders of young fruit and penetrate about
1.0 cm into the fruit. The damage resembles the symptoms of
diplodia rot (black canker) (Sanewski, 1991).
Mealy bugs and various species of scale insects are found
universally and usually become a serious pest on neglected
trees. The former is reported to be a major pest on marketable
fruit in some areas of Australia (Sanewski, 1991; Smith, 1999).
Red spider mites can become a serious problem in dry areas or
during dry seasons.
Mature green annonaceous fruit have been shown to be
rarely infested by the Mediterranean fruit fly (Ceratitis
capitata) and oriental fruit fly (Dacus dorsalis), but are found
on occasion in tree-ripened fruit. In Australia, the Queensland
fruit fly (Dacus tryoni) infests ripening atemoya fruit. ‘African
Pride’ appears more susceptible than ‘Pink’s Mammoth’. Use
of bait sprays and field sanitation are recommended measures
to minimize fruit fly infestation (Smith, 1991; Lloyd, 1999).
Fruit bagging also provides protection.
Problem weeds especially grasses and twining weeds should
be controlled before planting by cultivation and herbicides.
60
Annonaceae
Young trees should to be protected from weed competition by
hand weeding, mulching or contact herbicides. The shallow
root system limits the use of cultivation under the tree. A
translocated herbicide may be needed for perennial weeds and
are applied as a spot spray.
In Australia, the earliest
cultivar that was grown was ‘Pink’s Mammoth’ which was
introduced from British Guiana in the 1890s. This cultivar
takes 6–7 years to begin producing commercial-size yields of
fruit that are large, weighing 800 g to as large as 2 kg and is a
less precocious bearer than ‘African Pride’, which was
probably introduced into Australia from South Africa,
although its origin may have been Israel. Both these cultivars
have now been superseded by two new cultivars, ‘Maroochy
Gold’, originating out of the Queensland Department of
Primary Industries breeding programme, and ‘KJ Pinks’, a
budsport of ‘Pink’s Mammoth’ with exceptionally high fruit
setting. Both these cultivars produce medium to large fruit
with excellent eating characteristics. The main cultivar grown
in Florida and Hawaii is ‘Gefner’, an Israeli cultivar; this
cultivar prefers semi-tropical conditions. It produces small- to
medium-sized fruit. Most minor cultivars listed in the
literature have now been discarded.
Breeding and selection in atemoya (Annona spp. hybrids)
and cherimoya (A. cherimola) has been much neglected. Most
atemoya cultivars are hybrids of cherimoya (A. cherimola) and
sugar apple (A. squamosa). However, other atemoya cultivars
are of unknown genetic origin. Few new cultivars of atemoya
and cherimoya have been selected in the past 20 years due to
the small population of naturally occurring seedlings, and lack
of a breeding programmes/strategies for these fruit. In
contrast, other subtropical and tropical fruit such as mango
have been intensively selected from over several hundred
thousand seedlings for more than 100 years.
Wester (1913, 1915) was the first person to realize the
possibilities of genetic improvement of Annona and initiated a
breeding programme in Florida and the Philippines. However,
because of the small number of progeny he evaluated, no new
varieties were selected. No further breeding of atemoyas was
carried out until the late 1980s when the authors initiated a
breeding programme in atemoya in Australia and coincidently
Zill and Medeem (personal communication, 1997) were
undertaking introductions and breeding for atemoya in
Florida. In Florida approximately 3000 seedling progeny,
mainly interspecific crosses, have been planted out in Zill
orchards near Boynton Beach. To date, none of the progeny of
these crosses has produced commercial cultivars.
Many interesting species and selections have been introduced
into Florida from Central America over several decades
(Whitman, 1972; Popenoe, 1974; Zill, personal communication,
1997). Many of these have been used in the Florida breeding
programme, and more recently in the Australian breeding
programme since their recent introduction in 1998. In Spain,
over the past 20 years, Dr Jose Farre (personal communication,
1997) has conducted a major cherimoya varietal introduction
programme from South America.
The main characteristics being selected in atemoya are fruit
symmetry, smooth skin, chilling tolerance and low seed
number or seedlessness. Seedlessness is a desirable
MAIN CULTIVARS AND BREEDING
characteristic being sought in atemoya and may be achieved
through various strategies. The most commonly used
approach in other crops is to double the chromosomes of
diploids to produce tetraploids which, in turn, are crossed
back to a diploid to produce a triploid plant which is often
sterile and produces seedless fruit. This technique should also
work with atemoya and other Annona spp. which are diploids
(Thakur and Singh, 1964, 1965).
In Queensland, considerable recent progress has been made
in selecting new types, identifying appropriate parents and
gaining an understanding of the inheritance of desirable traits
in atemoya (George et al., 2002c). Inter-varietal crosses have
been made between the best selections of atemoya and the
main commercial cultivars such as ‘African Pride’, ‘Pink’s
Mammoth’ and ‘Hillary White’ (Table A.36). Interspecific
crosses have also been made between four different species: A.
cherimola (cherimoya), A. squamosa (sugar apple), A. reticulata
(bullock’s heart) and Annona diversifolia (ilama). Some 10,000
breeding lines have been field planted since 1992.
Several cultivars have been selected in Israel (Gazit and
Eistenstein, 1985). More recently, active breeding and
selection programmes have been conducted in Florida and
Australia (George et al., 2002a). The latter programme has
produced four named cultivars to date and has successfully
hybridized atemoya with A. diversifolia and A. reticulata to
produce red and pink skin types. There is considerable
variation among seedlings.
An alternative approach to conventional breeding using
mutation techniques is also being evaluated in Australia. Early
results indicate that it may be feasible to produce tetraploids
using colchicine applications to juvenile buds. Ten advanced
selections are being trialed at six evaluation sites throughout
Queensland and northern New South Wales.
The Queensland and Florida programmes have successfully
developed hybrids with red skin colour and pink internal flesh.
Red skin colour may be carried by either a single or double
recessive gene. Fruit symmetry, flesh recovery and flavour
characteristics of some crosses are excellent. To date, one
advanced selection, ‘Maroochy Gold’, has been named.
Alan George and Robert E. Paull
Literature cited and further reading
Ahmed, M.S. (1936) Pollination and selection in Annona squamosa
and Annona cherimola. Bulletin of Egypt Ministry of Agricultural
Science Services, Horticulture Section 157, 1–23.
Brown, B.I., Wong, L.S., George, A.P. and Nissen, R.J. (1988)
Comparative studies on the postharvest physiology of fruit from
Table A.36. Selected cultivars of atemoya.
Name
Origin
‘Maroochy Gold’
‘KJ Pinks’
‘Pink’s Mammoth’
‘African Pride’
‘Bradley’
‘Page’
‘Gefner’
‘Kabri’
‘Malalai’
Australia
Australia
Australia
Southern Africa
USA – Florida
USA – Florida
Israel
Israel
Israel
Annona
different species of Annona (custard apple). Journal of
Horticultural Science 63, 521–528.
Campbell, C.W. (1985) Cultivation of fruits of the Annonaceae in
Florida. Proceedings of the American Society for Horticultural
Science Tropical Region 29, 68–70.
Cresswell, G. and Sanewski, G. (1991) Diagnosing nutrient disorders.
In: Sanewski, G. (ed.) Custard Apples – Cultivation and Crop
Protection. Information Series QI90031, Queensland Dept.
Primary Industry, Brisbane, Australia.
Dhingra, J., Mehrotra, R.S. and Aneja, I.R. (1980) A new postharvest
disease of Annona squamosa L. Current Science 49, 477–478.
Duarte, O., Pineda, A. and Rodríguez, yP. (1999) Mejora al cuajado
de atemoya ‘Gefner’ (Annona cherimola ⫻ Annona squamosa) con
diversos tratamientos de polinización manual. Proceedings of
InterAmerican Society for Tropical Horticulture 45, 74–76.
Duckworth, R.B. (1996) The Composition of Fruit and Vegetables.
[Fruit and vegetables, Appendix A.] Permagon Press, London.
Gazit, S. and Eistenstein, D. (1985) Floral biology of Annona
squamosa and Annona cherimola in relation to spontaneous
appearance of atemoya in Israel. Proceedings of the American
Society for Horticultural Science Tropical Region 29, 66–67.
Gazit, S., Galon, I. and Podoler, H. (1982) The role of nitidulid
beetles in pollination of Annona in Israel. Journal of American
Society of Horticultural Science 107, 849–852.
George, A.P. (2000) Improving productivity of custard apple (Annona
spp. hybrids) in subtropical Australia. PhD dissertation,
University of Queensland, Australia.
George, A.P. and Nissen, R.J. (1985a) The custard apple. Part 1.
Species, varieties and rootstock selection. Australian Horticulture
83, 100–106.
George, A.P. and Nissen, R.J. (1985b) The custard apple. Part 2.
Australian Horticulture 83, 105–110.
George, A.P. and Nissen, R.J. (1986a) Effect of pruning and
defoliation on precocity of bearing of custard apple (Annona
atemoya Hort.) var. African Pride. Acta Horticulturae 175,
237–241.
George, A.P. and Nissen, R.J. (1986b) The effects of root temperature
on growth and dry matter production of Annona species. Scientia
Horticulturae 31, 95–99.
George, A.P. and Nissen, R.J. (1987a) Effects of cincturing,
defoliation and summer pruning of vegetative growth and
flowering of custard apple (Annona cherimola ⫻ Annona
squasmosa) in subtropical Queensland. Australian Journal of
Experimental Agriculture 27, 915–918.
George, A.P. and Nissen, R.J. (1987b) The effects of day/night
temperatures on growth and dry matter production of custard
apple. Scientia Horticulturae 31, 269–274.
George, A.P. and Nissen, R.J. (1987c) Propagation of Annona species:
a review. Scientia Horticulturae 33, 75–85.
George, A.P. and Nissen, R.J. (1988) The effects of temperature,
vapour pressure deficit and soil moisture stress on growth,
flowering and fruit set of custard apple (atemoya) ‘African Pride’.
Scientia Horticulturae 34, 183–191.
George, A.P. and Nissen, R.J. (1992a) Annonaceae. In: Verheij,
E.W.M. and Coronel, R.E. (eds) Edible Fruits and Nuts. Plant
Resources of South East Asia. PROSEA Foundation, Bogor,
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George, A.P. and Nissen, R.J. (1992b) Effects of environmental
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Science 67, 445–455.
61
George, A.P. and Nissen, R.J. (1993) Fruits of tropical climates.
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George, A.P. and Nissen, R.J. (2002a) Effects of drought on fruit set,
yield and fruit quality of custard apple (Annona spp. hybrids)
‘African Pride’ plants. Journal of Horticultural Science and
Biotechnology 77, 418–427.
George, A.P. and Nissen, R.J. (2002b) Control of tree size and vigour
in custard apple (Annona spp. hybrid) cv. African Pride in
subtropical Australia. Australian Journal of Experimental
Agriculture 42, 503–512.
George, A.P., Nissen, R.J. and Brown, B.T. (1987) The custard apple.
Queensland Agriculture Journal 113, 287–297.
George, A.P., Nissen, R.J. and Carseldine, M.L. (1989) Effect of
season (vegetative flushing) and leaf position on the leaf nutrient
composition of Annona spp. hybrid cv. Pink’s Mammoth in southeastern Queensland. Australian Journal of Experimental
Agriculture 29, 587–595.
George, A.P., Nissen, R.J. and Howitt, C. (1990) Effects of
environmental variables and cropping on leaf conductance of
custard apple (Annona cherimola ⫻ Annona squamosa) ‘African
Pride’. Scientia Horticulturae 45, 137–147.
George, A.P., Nissen, R.J. and Campbell, J.A. (1992) Pollination and
selection in Annona species (cherimoya, atemoya, and sugar
apple). Acta Horticulturae 32, 178–185.
George, A.P., Nissen, R.J. and Broadley, R. (1997) An overview of the
Australian custard apple industry. In: Proceedings of the First
International Symposium on Annonaceae, Chapingo, Mexico,
pp. 51–57.
George, A.P., Nissen, R.J. and Broadley, R. (1999) Past, present and
future of the Australian custard apple industry. In: Broadley, R.
(ed.) Proceedings of the Second Australian Custard Apple
Conference. Queensland Depart. Primary Ind. Publication, Twin
Water, Queensland, Australia, pp. 4–13.
George, A.P., Subhadrabandhu, S. and Nissen, R.J. (2001) Effects of
pruning, girdling and paclobutrazol on shoot growth, yield and
quality of atemoya (Annona spp. hybrids) cv. African Pride in
subtropical Australia. Thai Journal of Agricultural Science 34,
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Horticulturae 575, 323–328.
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Effects of new rest-breaking chemicals on flowering, shoot
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575, 835–840.
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Ind. Publication, Twin Water, Queensland, Australia, pp. 54–56.
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Annonaceae
Kshirsagar, S.V., Shinde, N.N., Rane, D.A. and Borikar, S.T. (1976)
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South Indian Horticulture 24, 6–10.
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Central America and Panama, Guatemala, and National Institute
of Health, Bethesda, Maryland.
Lloyd, A. (1999) A fly in the ointment – fruit flies and custard apples.
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Twin Water, Queensland, Australia, pp. 63–68.
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Asimina triloba
pawpaw
The North American pawpaw, Asimina triloba (L.) Dunal,
grows wild as an understorey tree or thicket-shrub in mesic
hardwood forests ranging from northern Florida to southern
Ontario (Canada) and as far west as eastern Nebraska. Fruit
(100–1000 g) may be borne singly or in clusters, are highly
nutritious, have a strong aroma and a unique flavour that
resembles a combination of banana, mango and pineapple.
This oblong-shaped fruit has both fresh market and
processing potential.
Pawpaws have a well-established place in folklore and
American history. The traditional American folk song, ‘Way
down, yonder in the pawpaw patch’ was quite popular at one
time and autumn hunting for pawpaw in the woods is a
cherished tradition for many rural families in the eastern
USA. The first report of pawpaw dates back to 1541 when
followers of the Spanish explorer Hernando de Soto found
Native Americans growing and eating pawpaws in the valley of
the Mississippi. The Native Americans used the bark of
pawpaw trees to make fishing nets. Daniel Boone and Mark
Twain were reported to have been pawpaw fans. Lewis and
Clark, the explorers who between 1804 and 1806 successfully
crossed America from St Louis to the mouth of the Colombia
River, recorded in their journal (18 September 1806) how
pawpaws helped save them from starvation. Several American
towns, townships, creeks and rivers were named after the
pawpaw during the 19th century. Interest in pawpaw as a fruit
crop was evident in the early 1900s, however, the rapid
perishability of the fruit is likely to have decreased interest in
pawpaw. Interest in pawpaw did grow in the years between
1950 and 1985. Recently, there has been developing interest in
pawpaw as a gourmet food.
Asimina
World production
Although pawpaw has great potential for commercial
production, orchard plantings remain limited. Currently, most
pawpaw fruit for sale are collected from wild stands in the
forest. However, in a number of states in the USA small
private orchards, usually less than 1 ha in size, have been
planted. There are also pawpaw plantings in Italy, China,
Israel, Japan, Romania, Belgium and Portugal. In the USA,
pawpaw fruit and products are mainly sold at farmers’
markets, directly to restaurants and via entrepreneurs on the
Internet. However, at present the grower base is insufficient to
establish a commercial processing industry.
Uses and nutritional composition
Pulp from fruit can be eaten fresh or processed. The flavour of
the fruit can intensify as it over-ripens, as with banana,
resulting in pulp that is excellent for use in cooking. The seeds
and skin are generally not eaten. Local delicacies made from
fruit pulp include ice cream, compote, jam, pies, custards and
wine. The pawpaw fruit has a high nutritional value (Table
A.37). Pawpaw and banana are similar in dietary fibre content
and overall nutritive composition. The highly perishable
nature of the fruit can limit the available supply.
Botany
The pawpaw, A. triloba, is a
member of the mostly tropical custard apple family,
Annonaceae, which is the largest primitive family of flowering
plants. This family includes several delicious tropical fruit
such as the custard apple (Annona reticulata L.), cherimoya
(Annona cherimola Mill.), sweetsop or sugar apple (Annona
squamosa L.), atemoya (A. squamosa ⫻ A. cherimola) and
soursop (Annona muricata L.). The genus Asimina is the only
temperate-zone representative of the tropical Annonaceae, and
includes nine species, most of which are native to the extreme
south-eastern regions of Florida and Georgia. The North
American native pawpaw (A. triloba) produces the largest
fruit, has the most northerly and largest native range and the
greatest commercial potential of the Asimina genus. Pawpaw is
also a common name for papaya (Carica papaya), a tropical
fruit in the family Caricaceae. The two fruits are genetically
unrelated, but some pawpaws do have a papaya-like flavour.
TAXONOMY AND NOMENCLATURE
Pawpaw is a moderately small, deciduous tree or
shrub that flourishes in the deep, rich fertile soils of riverbottom lands of the forest understorey. Trees may attain
5–10 m in height and are usually found in patches, due to root
suckering. In sunny locations, trees typically assume a
pyramidal habit, with a straight trunk and lush, dark green,
long, drooping leaves (Fig. A.7 E). Leaves occur alternately,
are obovate-oblong in shape, glabrous, with a cuneate base,
acute midrib, and may be 15–30 cm long and 10–15 cm wide.
Vegetative and flower buds occur at different nodes on the
stem, the flower buds being basipetal. Vegetative buds are
narrow and pointed, and the flower buds are round and
covered with a dark-brown pubescence.
The dark maroon-coloured flowers of the pawpaw are
hypogynous and strongly protogynous. Flowers are pendant or
63
Table A.37. Nutritional composition of pawpawa per 100 g (Source: Peterson
et al., 1982; Jones and Layne, 1997).
Proximal analysis
g
Food energy (kcal)
Protein
Total fat
Carbohydrate
Dietary fibre
80
1.2
1.2
18.8
2.6
Vitamins
Vitamin A REb
Vitamin A Ius
Vitamin C
Thiamine
Riboflavin
Niacin
Minerals
Potassium
Calcium
Phosphorus
Magnesium
Iron
Zinc
Copper
Manganese
Essential amino acids
Histidine
Isoleucine
Leucine
Lysine
Methionine
Cystine
Phenylalanine
Tyrosine
Threonine
Tryptophan
Valine
mg
8.6
87
18.3
0.01
0.09
1.1
mg
345
63
47
113
7
0.9
0.5
2.6
mg
21
70
81
60
15
4
51
25
46
9
58
a Pawpaw analysis was done on pulp with skin, although the skin is not
considered edible. Probably much of the dietary fibre, and possibly some of
the fat, would be thrown away with the skin.
b RE, Retinol equivalents – these units are used in the Recommended Dietary
Allowances table (National Research Council, 1989).
c IU, International units.
DESCRIPTION
nodding, with sturdy pubescent peduncles up to 4 cm long
(Fig. A.7 B). The mature flowers have an outer and inner
whorl of three, maroon-coloured, three-lobed petals reaching
up to 5 cm in diameter (Fig. A.7 A). The inner petals are
smaller and fleshier, with a nectary band at the base. The
flower has a fetid aroma. Flowers have a globular androecium
and a gynoecium usually composed of three to seven carpels
resulting in three to seven fruited clusters; up to nine-fruited
clusters have been noted (Fig. A.7 C). Flowers emerge before
leaves in spring (about April in Kentucky). Pawpaw blossoms
occur singly on the previous year’s wood.
Pawpaw’s custard apple-like fruit are berries. The fruit have
an oblong shape, green skin, a pleasant but strong aroma when
64
Annonaceae
Fig. A.7. (A) Mature flower with an outer and inner whorl of three maroon-coloured, three-lobed petals; (B) a mature pawpaw flower and
developing cluster from an earlier flower; (C) pawpaw cluster with ripe fruit; (D) a fruit cut open lengthwise and seeds removed; and (E) a
pawpaw tree showing natural pyramidal growth habit in a full sun exposure.
ripe, and intense flavour (Fig. A.7 C, D). However, flavour
varies among cultivars, with some fruit displaying complex
flavour profiles. Fruit from poor quality pawpaw genotypes can
have a mushy texture, lack sweetness and have an overly rich
flavour with turpentine or bittersweet aftertaste; many wild
pawpaws have poor eating quality. Fruit from superior
genotypes have a firm texture, a delicate blend of flavours, are
rich but not cloying, and have no bitter aftertaste. The flavour of
a pawpaw fruit can intensify when it over-ripens, as with
banana, resulting in pulp that is excellent for use in cooking.
The fruit are oblong-cylindrical, typically 3–15 cm long,
3–10 cm wide and weigh from 100 to 1000 g. They may be
borne singly or in clusters which resemble the ‘hands’ of a
banana plant. In the fruit, there are two rows of seeds (12–20
seeds) that are brown and bean shaped and that may be up to
3 cm long. The seeds and skin of the fruit are generally not
eaten. The endosperm of the seeds contain alkaloids that are
emetic and if chewed may impair mammalian digestion.
Pawpaw fruit allergies have been reported in some people.
Flowers are strongly protogynous
and are predominantly self-incompatible, although the
pawpaw cultivar ‘Sunflower’ may be self-fruitful. Pollination is
by flies (Diptera) and beetles (Nitidulidae), and possibly other
nocturnal insects. Seedlings normally begin to flower upon
reaching about 1.8 m in height, but may not set fruit; cropping
is achieved at 5–8 years of age. Grafted pawpaw trees often
flower within 3 years of planting, but often fail to set fruit at
that time. This may be due to inadequate pollination or
inadequate canopy to support fruit development. Grafted
trees usually begin reliable fruit production at 5–6 years of
age.
For an individual tree, the bloom and pollination period
may last from 3 to 4 weeks. There is also cultivar variation for
bloom date that may be related to chilling hour requirement.
As a result, harvest for an individual tree may be extended
over a 3–4 week period. Thus, multiple harvests are necessary
depending on fruit ripeness. Each fruit cluster develops from
an individual flower, and fruit within a cluster develop and
REPRODUCTIVE BIOLOGY
Asimina
often ripen at different times (Fig. A.7 C). In cultivation,
pawpaw yields per tree are often low. Yields for mature grafted
trees in the seventh year can average between 2.0 and 6.5
kg/tree, depending on the cultivar. The tropical Annonaceae
relatives of the pawpaw, such as cherimoya, sweetsop (sugar
apple), soursop and atemoya also have low yields, due to low
rates of natural pollination. Pawpaws in the wild often have
poor fruit set due to low light levels in the understorey and
pollinator limitation. Pawpaws in the wild often produce many
root suckers that could potentially result in large clonal
patches contributing to poor fruit set because of selfincompatibility. Fruit set can be improved by hand crosspollination and it is likely this could be used to improve yields.
Fruit increase in size
during the course of spring and summer, ripening in late
summer or early autumn. Pawpaw fruit ripening is
characterized by an increase in soluble solids concentration
(up to 20%), flesh softening, increased volatile production
and, in some genotypes, a decline in green colour intensity of
the skin. Within 3 days after harvest, ethylene and respiratory
climacteric peaks are clearly evident as pawpaw fruit rapidly
soften. A common practice to determine maturity is to touch
each fruit to determine if it is ready to harvest; ripe softening
pawpaw fruit yield to slight pressure, as ripe peaches do, and
can be picked easily with a gentle tug. Thus, fruit are
harvested when they have already begun ripening and have
lost some firmness.
FRUIT GROWTH AND DEVELOPMENT
Horticulture
The pawpaw produces a relatively large, flat
seed with a dark brown fibrous seedcoat. Seed can be collected
from fruit when the flesh is soft or over-ripe. Pawpaw has
moderately recalcitrant seed that does not tolerate desiccation,
and it only has a relatively short period of viability at room
temperature. As little as 5 days under open-air conditions can
reduce the moisture content of pawpaw seeds to 5% and result
in total loss of viability. Pawpaw seed requires stratification for
optimal germination. Pawpaw seeds must be stored moist at
chilling temperature (5°C) to overcome embryo dormancy.
Seed can be stored in moist peat moss in ziplock bags for 2–3
years at 5°C and maintain a high germination percentage.
Storing pawpaw seed in a freezer (-15°C) will kill the embryo
and make the seed not viable.
Stratified seed can be sown in a well-aerated potting
substrate with a high sphagnum peat moss component (> 75%
by volume), cation exchange capacity and water-holding
capacity. Tall containers should be used to accommodate the
developing taproot of seedlings. Because pawpaw has a coarse
fibrous root system that is quite fragile, most commercial
nurseries propagate pawpaw in containers rather than in a
nursery bed. Although some commercial nurseries sell bareroot trees grown in nursery beds, we do not recommend this
practice. Transplant shock is common with bare-root trees and
field establishment is usually poor. Young pawpaw seedlings
are sensitive to excessive ultraviolet (UV) light and can be
damaged under full sun conditions. If growing seedlings
outside, the plants should be kept in moderate shade their first
year (we use 55% shade cloth) for maximum growth of the
PROPAGATION
65
plant. Seedlings will grow well in whitewashed or even
unshaded greenhouses. Plants in their second year of growth
outside do not require shading and will grow nicely in full sun
provided water is not limiting.
Chip-budding and whip-and-tongue grafting are the two
most reliable means to clonally propagate pawpaw. Winter
collected, dormant budwood should have its chilling
requirement fulfilled. Chip budding and grafting are most
successful when the seedling rootstock is at least 0.5 cm
diameter and actively growing. Bud take exceeding 90% can
be obtained. Clonal propagation of pawpaw by other methods
such as root cuttings or softwood cuttings has been
unsuccessful. Clonal propagation of pawpaw by tissue culture
has been attempted using various explant sources of different
physiological ages. The primary limitation in tissue culture is
the inability of explants to form roots.
Currently, pawpaw cultivars with superior fruit
characteristics are propagated by grafting and budding onto
seedling rootstocks. No clonal rootstocks are available for
pawpaw.
ROOTSTOCKS
AND PRUNING Present recommendations for
pawpaw plantings are 2.4 m within rows and 3.7–4.6 m
between rows. Row orientation should be north–south if
possible. Shading of pawpaw in the field the first year is
recommended and can be accomplished by installing
translucent double-walled polyethylene ‘tree-tubes’ around
each tree, securing them with bamboo stakes. However, trees
taller than 45 cm at planting do not require shading. During
warm summer temperatures (> 35°C), the tubes should be
removed from the trees, otherwise foliage within tubes can
became heat-stressed and desiccated. Weed control is
important to limit competition and improve establishment,
but there are no herbicides currently recommended for use on
pawpaw. Mulching with straw or other organic material can be
used to limit weed growth in the tree row. When natural
rainfall is inadequate, supplemental irrigation can substantially
improve tree survival rates.
Most pawpaw genotypes naturally develop a strong central
leader. The growth habit is similar to that of ‘Bradford’ pear, a
popular ornamental tree in the USA. Trees should not be
headed at planting and no pruning is required in the first year.
Branches can often develop narrow crotch angles in relation to
the trunk. Training to more horizontal scaffold limbs increases
scaffold strength and reduces limb breakage that may occur
under heavy crops or during ice storms. Pruning is conducted
in late winter–early spring and consists of removing low
branches to a height 60–90 cm on the trunk.
TRAINING
THINNING Pawpaw fruit set can often be low, but some
growers do practise hand thinning of fruit to increase fruit
size.
FERTILIZATION Fertilization requirements have not been
determined for bearing pawpaw trees. However, trees provided
with water-soluble fertilizer (20N-8.6P-16.6K) plus soluble
trace elements once in May, June and July during active
growth have achieved 30–45 cm of shoot extension each year
in Kentucky. Excellent growth has been achieved with
66
Annonaceae
granular ammonium nitrate fertilizer (34N-0P-0K) broadcast
under pawpaw trees in early spring at 30–60 g N/tree applied
before budbreak.
DISEASES AND PESTS Pawpaws have few disease problems;
however, leaves can exhibit leaf spot, principally a complex of
Mycocentrospora aiminae, Rhopaloconidium asiminae Ellis and
Morg. and Phyllosticta asiminae Ellis and Kellerm. At orchards
in Oregon (outside pawpaw’s native range), vascular wilt-like
symptoms have been observed in the spring after pawpaw trees
have leafed out. The pawpaw peduncle borer (Talponia
plummeriana Busck) is a small moth whose 5 mm-long larva
burrows into the fleshy tissues of the flower causing the flower
to wither and drop. The zebra swallowtail butterfly (Eurytides
marcellus), whose larvae feed exclusively on young pawpaw
foliage, will damage leaves, but this damage has been negligible
in plantings. The larvae of a leafroller (Choristoneura parallela
Robinson) may also damage flowers and leaves. Deer will not
generally eat the leaves or twigs, but they will eat fruit that has
dropped on the ground. Occasionally, male deer will rub their
antlers on young trees, scraping off bark and occasionally
breaking off branches. Japanese beetles (Popillia japonica
Newman) occasionally feed on young foliage and can damage
pawpaw trees.
Biologically active compounds known as annonaceous
acetogenins have been extracted from pawpaw twigs and have
potential as human medications and botanical pesticides.
About 250 of these compounds have been isolated and
characterized. Three of these compounds, bullatacin, bulletin
and bullanin have high potencies against human cancer cells in
vitro. Dr Jerry McLaughlin of Nature’s Sunshine Products
(Spanish Fork, Utah) has developed a commercial head-liceremoval shampoo from pawpaw. Botanically derived pesticides
that are environmentally compatible and biologically
degradable may also be obtained from pawpaw because the
annonaceous acetogenins are toxic to several economically
important insect species.
HANDLING AND POSTHARVEST STORAGE Pawpaw fruit soften
rapidly at room temperature after harvest. At room
temperature, very soft ripe fruit have a shelf life of 2–3 days
while those fruit that are just beginning to soften have a shelf
life of 5–7 days. Fruit that have just begun to soften can be
stored for 1 month at 4°C with little change in fruit firmness
and they will ripen normally when returned to room
temperature. Hard immature fruit will not ripen, even if
treated with ethephon. Because fruit are non-uniform in size
and shape, packaging that minimizes bruising during shipping
needs to be developed.
Efforts to domesticate the
pawpaw began early in the 20th century. In 1916, a contest to
find the best pawpaw was sponsored by the American Genetics
Association. This contest generated much interest and the
sponsors thought that with time and ‘intelligent breeding’
commercial quality varieties could be developed and an
industry begun. However, an industry did not develop. Pawpaw
enthusiasts noted that the rapid perishability of pawpaw fruit
was the major factor inhibiting commercialization.
Beginning in the 20th century, elite pawpaw selections from
the wild were assembled in extensive collections by various
enthusiasts and scientists, including Benjamin Buckman
(Farmington Illinois, c. 1900–1920), George Zimmerman
(Linglestown, Pennsylvania, 1918–1941), and Orland White
(Blandy Experimental Farm, Boyce, Virginia, 1926–1955).
From about 1900–1960, at least 56 clones of pawpaw were
selected and named. Fewer than 20 of these selections remain,
with many being lost from cultivation through neglect,
abandonment of collections, and loss of records necessary for
identification. Since 1960, additional pawpaw cultivars have
been selected from the wild or developed as a result of
breeding efforts of hobbyists. More than 40 clones are
currently available (Table A.38). From 1995 to 1999, Kentucky
State University (KSU) and the PawPaw Foundation (PPF)
established a Pawpaw Regional Variety Trial (PRVT) with 28
MAIN CULTIVARS AND BREEDING
Table A.38. Commercially available pawpaw cultivars (Source: descriptions derived from Jones and Layne, 1997; Jones et al., 1998; and unpublished data of K.
Pomper)a.
Cultivar
Descriptionb
‘Adam’s Secret’
From Pennsylvania, large fruit, few seeds, skin remains green when ripe
‘Blue Ridge’
Selected in Kentucky by Johnny Johnson; has white-fleshed fruit
‘Collins’
Selected in Georgia
‘Convis’
Selected from Corwin Davis orchard. Large-size fruit, yellow flesh; ripens first week of October in Michigan
‘Davis’
Selected from the wild in Michigan by Corwin Davis in 1959. Introduced in 1961 from Bellevue, Michigan. Medium-size fruit, up to
12 cm long; green skin; yellow flesh; large seed; ripens first week of October in Michigan; keeps well in cold storage.
‘Duckworth A’
Low-chill cultivar selected in San Mateo, Florida by Eric Duckworth, seedling of Louisiana native parent; tree with pyramidal shape
‘Duckworth B’
Low-chill cultivar selected in San Mateo, Florida by Eric Duckworth, seedling of Louisiana native parent; grows no larger than a
shrub
‘Estil’
Selected by Nettie Estil in Frankfort, Kentucky. Large fruit, smooth-textured flesh
‘Ford Amend’
Selected from wild seedling of unknown parentage by Ford Amend around 1950. Introduced from Portland, Oregon. Medium-size
fruit and earlier than ‘Sunflower’; ripens late September in Oregon; greenish-yellow skin; orange flesh
‘G-2’
Selected from G.A. Zimmerman seed by John W. McKay, College Park, Maryland, in 1942
‘Glaser’
Selected by P. Glaser of Evansville, Indiana. Medium-size fruit
‘IXL’
Hybrid of ‘Overleese’ and ‘Davis’; large fruit, yellow flesh; ripens second week of October in Michigan
‘Jack’s Jumbo’
Selected in California from Corwin Davis seed; large fruit
Asimina
Cultivar
Descriptionb
‘Kirsten’
Hybrid seedling of ‘Taytwo’ ⳯ ‘Overleese’; selected by Tom Mansell, Aliquippa, Pennsylvania
‘LA Native’
From LA, blooms late in Tennessee, small fruit, somewhat frost hardy
‘Little Rosie’
Selected by P. Glaser of Evansville, Indiana. Has small fruit. Reported to be an excellent pollinator
67
‘Lynn’s Favorite’
Selected from Corwin Davis orchard. Yellow fleshed, large fruit; ripens second week of October in Michigan
‘M-1’
Selected from ‘G-2’ seed by John W. McKay, College Park, Maryland, in 1948
‘Mango’
Selected from the wild in Tifton, Georgia, by Major C. Collins in 1970. Vigorous growth
‘Mary Foos Johnson’
Selected from the wild in Kansas by Milo Gibson. Seedling donated to North Willamette Experimental Station, Aurora, Oregon, by
Mary Foos Johnson. Large fruit; yellow skin; butter-coloured flesh; few seeds; ripens first week of October in Michigan
‘Mason/WLW’
Selected from the wild in Mason, Ohio, by Ernest J. Downing in 1938
‘Middletown’
Selected from the wild in Middletown, Ohio, by Ernest J. Downing in 1915. Small-size fruit
‘Mitchell’
Selected from the wild in Jefferson Co., Illinois, by Joseph W. Hickman in 1979. Medium-size fruit, slightly yellow skin, golden flesh,
few seeds
‘NC-1’
Hybrid seedling of ‘Davis’ ⳯ ‘Overleese’; selected by R. Douglas Campbell, Ontario, Canada, in 1976. Large fruit; few seeds; yellow
skin and flesh; thin skin; early ripening, 15 September in Ontario and early September in Kentucky
‘Overleese’
Selected from the wild in Rushville, Indiana, by W.B. Ward in 1950. Large fruit; few seeds; bears in clusters of three to five; ripens
first week of October in Michigan and early September in Kentucky
‘PA-Golden 1’
Selected as seedling from seed originating from George Slate collection by John Gordon, Amherst, New York. Early cropping.
Medium-size fruit, yellow skin, golden flesh; matures late August in Kentucky and mid-September in New York
‘PA-Golden 2’
Selected as seedling from seed originating from George Slate collection by John Gordon, Amherst, New York. Fruit: yellow skin,
golden flesh; matures mid-September in New York
‘PA-Golden 3’
Selected as seedling from seed originating from George Slate collection by John Gordon, Amherst, New York. Fruit: yellow skin,
golden flesh; matures mid-September in New York
‘PA-Golden 4’
Selected as seedling from seed originating from George Slate collection by John Gordon, Amherst, New York. Fruit: yellow skin,
golden flesh; matures mid-September in New York
‘Prolific’
Selected by Corwin Davis, Bellevue, Michigan, in mid-1980s. Large fruit; yellow flesh; ripens first week of October in Michigan
‘Rebecca’s Gold’
Selected from Corwin Davis seed, Bellevue, Michigan, by J.M. Riley in 1974. Medium-size fruit; kidney-shaped; yellow flesh
‘Ruby Keenan’
Medium-size fruit with excellent flavour
‘SAA-Overleese’
Selected from ‘Overleese’ seed by John Gordon, Amherst, New York, in 1982. Large fruit; rounded shape; green skin; yellow flesh;
few seeds; matures in mid-October in New York
‘SAA-Zimmerman’
Selected as seedling from seed originating from G.A. Zimmerman collection by John Gordon, Amherst, New York, in 1982. Large
fruit; yellow skin and flesh; few seeds
‘Silver Creek’
Selected from the wild in Millstedt, Illinois, by K. Schubert. Medium-size fruit
‘Sue’
Selected in southern Indiana. Medium-size fruit, yellow flesh, skin yellow when ripe
‘unflower’
Selected from the wild in Chanute, Kansas, by Milo Gibson in 1970. Tree reported to be self-fertile. Large fruit; yellow skin; buttercoloured flesh; few seeds; ripens early to mid-September in Kentucky and the first week of October in Michigan
‘Sunglo’
Yellow skin, yellow flesh, large fruit that ripens first week of October in Michigan
‘Sweet Alice’
Selected from the wild in West Virginia by Homer Jacobs of the Holden Arboretum, Mentor, Ohio, in 1934
‘Taylor’
Selected from the wild in Eaton Rapids, Michigan, by Corwin Davis in 1968. Small fruit; bears up to seven fruit in a cluster; green
skin; yellow flesh; ripens first week of October in Michigan
‘Taytwo’
Selected from the wild in Eaton Rapids, Michigan, by Corwin Davis in 1968. Sometimes spelled ‘Taytoo’. Small fruit; light-green skin;
yellow flesh; ripens first week of October in Michigan
‘Tollgate’
Yellow fleshed, large fruit that ripens first week of October in Michigan
‘Wells’
Selected from the wild in Salem, Indiana, by David Wells in 1990. Small- to medium-size fruit; green skin; orange flesh. Ripens midto late September in Kentucky
‘White’
Selected in Kentucky by Johnny Johnson; has white-fleshed fruit
‘Wilson’
Selected from the wild on Black Mountain, Harlan Co., Kentucky, by John V. Creech in 1985. Small fruit; yellow skin; golden flesh
‘Zimmerman’
Selected in New York from G.A. Zimmerman seed by George Slate
aDescriptions
bFruit
come from a wide variety of sources and most of the cultivars have not been compared for performance side by side at one geographic site.
size categories of small, medium and large are < 100 g, 100–150 g and > 150 g, respectively.
68
Annonaceae
clones at 13 sites across the USA. Cultivars being tested
include ‘Middletown’, ‘Mitchell’, ‘NC-1’, ‘Overleese’, ‘PAGolden 1’, ‘Rappahannock’, ‘Shenandoah’, ‘Sunflower’,
‘Susquehanna’, ‘Taylor’, ‘Taytwo’, ‘Wells’ and ‘Wilson’. The
other 15 clones were selections from the PPF breeding effort.
Tree survival, trunk cross-sectional area, fruit size and taste,
flesh-to-seed ratio, resistance to pests and diseases, and overall
productivity on a year-to-year basis are among the attributes
being evaluated. The pawpaw cultivars ‘PA-Golden 1’,
‘Overleese’, ‘NC-1’, ‘Sunflower’, ‘Shenandoah’ and
‘Susquehanna’ have performed well in Kentucky, have
excellent fruit size and flavour, and are recommended for
planting in the south-eastern USA. Complete results from the
PRVT and additional regional recommendations will be
available in a few years.
Kirk W. Pomper and Desmond R. Layne
Literature cited and further reading
Archbold, D.D. and Pomper, K.W. (2003) Ripening pawpaw fruit
exhibit respiratory and ethylene climacterics. Postharvest Biology
and Technology 30, 99–103.
Callaway, M.B. (1990) The Pawpaw (Asimina triloba). CRSHORT1–901T, Kentucky State University Publications, Kentucky.
Darrow, G.M. (1975) Minor temperate fruits. In: Janick, J. and
Moore, J.N. (eds) Advances in Fruit Breeding. Purdue University
Press, West Lafayette, Indiana, pp. 276–277.
Geneve, R.L., Pomper, K.W., Kester, S.T., Egilla, J.N., Finneseth,
C.L.H., Crabtree, S.B. and Layne, D.R. (2003) Propagation of
pawpaw – a review. HortTechnology 13, 428–433.
Huang, H., Layne, D.R. and Kubisiak, T.L. (2000) RAPD
inheritance and diversity in pawpaw [Asimina triloba (L.) Dunal].
Journal of American Society of Horticultural Science 125, 454–459.
Huang, H., Layne, D.R. and Kubisiak, T.L. (2003) Molecular
characterization of cultivated pawpaw [Asimina triloba (L.) Dunal]
using RAPD markers. Journal of American Society of Horticultural
Science 128, 85–93.
Jones, S.C. and Layne, D.R. (1997) Cooking with pawpaws. Bulletin
#PIB-001, Kentucky State University Cooperative Extension
Program, Kentucky.
Jones, S.C., Peterson, R.N., Turner, T., Pomper, K.W. and Layne,
D.R. (1998) Pawpaw planting guide: cultivars and nursery sources.
Bulletin #PIB-002, Kentucky State University Cooperative
Extension Program, Kentucky.
Kral, R. (1960) A revision of Asimina and Deeringothamnus
(Annonaceae). Brittonia 12, 233–278.
Layne, D.R. (1996) The pawpaw [Asimina triloba (L.) Dunal]: a new
fruit crop for Kentucky and the USA. HortScience 31, 777–784.
McLaughlin, J.L. (1997) Anticancer and pesticidal components of
pawpaw (Asimina triloba). Annual Report of the Northern Nut
Growers Association 88, 97–106.
National Research Council (1989) Recommended Dietary Allowances,
10th edn. National Academy Press, Washington, DC.
Peterson, R.N. (1991) Pawpaw (Asimina). Acta Horticulturae 290,
567–600.
Peterson, R.N. (2003) Pawpaw variety development: a history and
future prospects. HortTechnology 13, 449–454.
Peterson, R.N., Cherry, J.P. and Simmons, J.G. (1982) Composition
of pawpaw (Asimina triloba) fruit. Annual Report of the Northern
Nut Growers Association 73, 97–107.
Pomper, K.W., Crabtree, S.B., Brown, S.P., Jones, S.C., Bonney,
T.M. and Layne, D.R. (2003) Assessment of genetic diversity of
pawpaw varieties with inter-simple sequence repeat markers.
Journal of American Society of Horticultural Science 128, 521–525.
Pomper, K.W., Layne, D.R. and Jones, S.C. (2003) Container
production of pawpaw seedlings. HortTechnology 13, 434–438.
Pomper, K.W., Layne, D.R., Peterson, R.N. and Wolfe, D. (2003) The
pawpaw regional variety trial: background and early data.
HortTechnology 13, 412–417.
Reich, L. (1991) Uncommon Fruits Worthy of Attention: a Gardener’s
Guide. Addison-Wesley, New York.
Templeton, S.B., Marlette, M., Pomper, K.W. and Jones, S.C. (2003)
Favorable taste ratings for several pawpaw products.
HortTechnology 13, 445–448.
Wiese, T.D. and Duffrin, M.W. (2003) Effects of substituting pawpaw
fruit puree for fat on the sensory properties of a plain shortened
cake. HortTechnology 13, 442–444.
Willson, M.F. and Schemske, D.W. (1980) Pollinator limitation, fruit
production, and floral display in pawpaw (Asimina triloba).
Bulletin of the Torrey Botanical Club 107, 401–408.
Rollinia mucosa
biriba
Biriba tree, Rollinia mucosa (Jacq.) Baill. (Annonaceae), is
considered to have been cultivated in pre-Columbian times and
is now widely grown throughout the Amazon region. It is also
found in north-eastern Brazil, the Antilles and other parts of the
Caribbean. The fruit is appropriate for the fresh fruit market,
with prices varying according to size. The species is an excellent
alternative to diversify fruit gardens and to supply the demand
motivated mainly by originality of the native fruit. Moreover it is
an important genetic resource for its natural genetic variability,
which can be used in domestication studies, for selection of
superior genotypes or as a source of genes for related species.
The species is most commonly known as biriba. Other
names include biribazeiro, biriba-de-pernambuco or fruta-decondessa (Brazil); anon (Peru); chirimoya (Ecuador); mulato
(Colombia); sinon (Venezuela); anona babosa or zambo
(Mexico); cachiman morveusc, cachiman coehon or cachiman
montagne (Guadalupe); cachiman or anon cimamon (Puerto
Rico); anonillo (Panama); candongo or anona (Dominican
Republic); and wild sugar apple (English).
World production and yield
In Central Amazon, the fruit is sold per unit at open markets
or by street vendors. The species is not commercially
exploited, even considering its value as a food source and for
income for small farmers. It is most often found in home
gardens or urban yards and small farms. Fifteen-year-old trees
can produce over 150 fruit/year (Souza et al., 1997). During
the first three harvests of trees evaluated at EMBRAPA
Western Amazon Experimental Station, State of Amazon, the
average production reported was about 45 fruit/tree, with a
range from 35 to 55 (Sousa, 1998).
Uses and nutritional composition
Biriba has become a popular fruit due to its delicate flavour,
pulp yield and fruit size which is enough for one individual.
Rollinia
Fruit is consumed fresh, although some in the Amazon prefer
to blend it into a juice with or without milk. Fruit is 52% pulp,
42% peel and 6% seeds (Costa and Müller, 1995). The pulp is
creamy, slightly acid to sweet, with total soluble solids ranging
from 10 to 20% (Sousa, 1998). The fruit is regarded as being
rich in vitamin C (Table A.39). The fruit and seeds contain
acetogenens and alkaloids that may be anti-tumour agents and
can inhibit platelet aggregations (Liaw et al., 2003). The wood
is hard and heavy and is used for boats, masks and boxes.
Botany
TAXONOMY AND NOMENCLATURE There are approximately
65 species in the genus Rollinia, but only R. mucosa is
cultivated for its fruit. Synonyms include Rollinia deliciosa
Saff., Rollinia orthopetala A. DC., Rollinia pulchrinervis A.
DC., Rollinia sieberi A. DC. and Annona mucosa Jacq.
The tree is leafy with a round or conic canopy,
requiring sufficient space for satisfactory development.
During a study carried out on 80 plants conserved in a
diversified collection of Amazon indigenous species genetic
resources at EMBRAPA Western Amazon Experimental
Station, this species was considered to have a fast initial
growth, reaching a height of 3.7 m with a trunk 6.1 cm in
diameter at 50 cm in its first year (Sousa and Paiva, 2000). The
leaves are alternate, simple, oblong- or elliptical-oblongshaped, 15–25 cm long and 8–11 cm wide, coriaceous,
deciduous, without stipules, with a petiole 5–10 mm long.
Solitary flowers are hermaphrodite and arranged on large
pedicels. The fruit is a syncarp consisting of many joined
carpels of pyramidal-shaped fruitlets. The ripened fruit is
yellowish coloured and varies in shape, size, consistency and
pericarp. The black seeds are about 1–1.5 cm long.
DESCRIPTION
69
Municipality of Manaus, State of Amazonas, Brazil, where the
species is under study, the climate is characterized by a short
dry season, with an average rainfall above 60 mm during the
driest months (July and August) and above 300 mm during the
rainy season (January–March), annual rainfall averaging
around 2700 mm. The annual average temperature for the last
10 years has been around 26°C, with temperatures during the
coldest month never being below 18°C.
The tree grows better in deep soil with a high content of
organic matter and good drainage, even though it tolerates
poor, acid and heavy-textured soils. In the Brazilian Amazon,
the tree has been planted in upland xanthic ferralsols (yellow
oxisols) that have a clay texture, high acidity and high content
of exchangeable aluminum. In that region, the tree also grows
in lowlands subject to periodic flooding.
REPRODUCTIVE DEVELOPMENT The juvenile period from
seed is about 3 years. The biriba tree is among the most
tropical species and flowers just once a year after leaf fall,
which occurs in the Central Amazon during the low rainfall
season, between June and September. In Costa Rica, flowers
are observed between February and July.
FRUIT DEVELOPMENT In the state of Amazonas, fruit usually
mature between October and May, and between June and
November in Costa Rica. The fruit mature about 55 days from
anthesis (Falcão, 1993). Fruit may be spherical to oblongshaped, attaining 10–20 cm in length, 7–20 cm in diameter
and an average weight from 200 to 1000 g. Special care must
be taken in harvesting and handling to avoid fruit darkening
due to mechanical injury (Fig. A.8).
Horticulture
Trees are propagated either vegetatively or by
seeds. Seeds should be sown as soon as they are collected from
PROPAGATION
The biriba tree is
typically found growing in hot and humid climates. In the
ECOLOGY AND CLIMATIC REQUIREMENTS
Table A.39. Composition of edible flesh of biriba per 100 g (Source: Morton,
1987).a
Proximate
%
Water
Energy (kcal)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
77.2
80
2.8
0.2
19.1
1.3
0.7
Minerals
mg
Calcium
Iron
Phosphorus
24
1.2
26
Vitamins
mg
Ascorbic acid
Niacin
33
0.5
aThe
pulp is 52% of the total fruit weight and the total soluble solids/acidity
ratio is 28.
Fig. A.8. Biriba, Rollinia mucosa, is a syncarp of many joined
pyramidal-shaped fruitlets with soft spines that are easily damaged
during harvesting and handling.
70
Annonaceae
the fruit, preferably from high-yielding plants with highquality fruit. Germination takes about 30 days with about
80% germination. The most common vegetative method is
grafting. Rooted plantlets can be produced in vitro (Figueiredo
et al., 2000).
Cerconota anonnela (Lepidopterene)
larvae attack maturing fruit and cause considerable damage. A
borer (Cratosomus bombina) burrows into the bark and trunk
leading to secondary infections and branch death. White flies
(Aleurodicus cocois) and mealy bugs (Pseudococcus brevipes and
Aspidiotus destructor) are common on the leaves. Cercospora
leaf spot occurs and Glomerella cingulata causes stem dieback
and fruit rot.
DISEASES AND PESTS
Sousa, N.R. and Paiva, J.R. (2000) Conservação de recursos genéticos
de espécies autóctones da Amazônia em sistemas diversificados.
In: Actas del III Congreso Latinoamericano de Ecologia.
Universidad de Los Andes, Mérida, pp. 347–353.
Souza, A. das G.C. de, Sousa, N.R., Souza, S.E.L. da, Nunes,
C.D.M., Canto, A. do C., Cruz, L.A. de A. (1997) Fruit Trees of
the Amazon Region. Collection Biblioteca Botânica Brasileira 1,
EMBRAPA-SPI/Manaus, EMBRAPA-CPAA, Brasília, 204 pp.
APOCYNACEAE
Carissa congesta
karanda
No breeding programmes
have been described. The existing variation in the trees
planted from seeds offer excellent opportunities for selection
for yield, size, weight, consistency and total soluble solids
content. Some selections may have been made by Indians in
the Upper Solimôes River region, where fruit weighing above
4 kg with a smooth pericarp are found (Clement et al., 1982;
Lima and Costa, 1997). These selections have been known as
biriba do Alto Solimões.
Nelcimar Reis Sousa
The English names of Carissa congesta Wight. (Apocynaceae),
in addition to karanda, include Bengal currant and karaunda.
It is known as karandan and senggaritan in Indonesia; kerenda,
kerandang and berenda in Malaysia; caramba and pekunkila in
the Philippines; naam daeng, manaao ho and naam khee haet
in Thailand; and cay siro in Vietnam. The species is common
throughout its native range of India, Sri Lanka, Myanmar and
Malacca and it grows often in Thailand, Cambodia, South
Vietnam and East Africa. Introduced as a hedge, it is now wild
around Djakarta. The synonym is Carissa carandas Auct.
Literature cited and further reading
Uses and nutrient composition
Clement, C.R., Müller, C.H. and Chavez Flores, W.B. (1982)
Recursos genéticos de espécies frutíferas nativas da Amazônia
Brasileira. Acta Amazônica 12, 677–695.
Costa, J.P.C. da and Müller, C.H. (1995) Fruticultura Tropical: o
Biribazeiro Rollinia mucosa (Jacq.) Baill. Documentos 84,
EMBRAPA-CPATU, Belém, Brazil, 35 pp.
Falcão, M. de A. (1993) Aspectos Fenológicos, Ecológicos e de
Produtividade de Algumas Fruteiras Cultivadas na Amazônia
Brasileira: Araçá-boi (Eugenia stiptata McVaugh), Biribá (Rollinia
mucosa Jacq.), Camu-camu (Myrciaria dúbia (H.B.K.) McVaugh),
Cupuaçu (Theobroma grandiflorum (Wild. ex. Spreng) Schum.) e
Graviola (Annona muricata L.). 2nd edn rev. FUA, V.2, Manaus,
Brazil, 97 pp.
Figueiredo, S.F.L., Alarello, N. and Viana, V.R. (2000) Minor
propagation of Rollinia mucosa (Jacq.) Baill. In Vitro Cellular and
Development Biology 37, 471–475.
Kessler, P.J.A. (1993) Annonaceae. In: Kubitzki, K., Rohwer, J.G. and
Bittrich, V. (eds) The Families and Genera of Vascular Plants. Vol.
2. Flowering Plants. Dicotyledons. Magnoliid, Hamamelid and
Caryophyllid Families. Springer Verlag, Berlin, pp. 93–129.
Liaw, C.C., Chang, F.R., Wu, M.J. and Wu, Y.C. (2003) A novel
constituent from Rollinia mucosa, rollicosin, and a new approach
to develop Annonaceaus acetogenius as potencial anti-tumor
agents. Journal of Natural Products 66, 279–281.
Lima, R.R. and Costa, J.P.C. da (1997) Coleta de Plantas de Cultura
Pré-colombiana na Amazônia Brasileira – Metodologia e Expedições
Realizadas para Coleta de Germoplasma. Documentos 99,
EMBRAPA-CPATU, Belém, Brazil, 148 pp.
Morton, J. (1987) Fruits of Warm Climates. Creative Resource
Systems Inc., Winterville, North Carolina, pp. 88–90.
Sousa, N.R. (1998) Biribazeiro fruits in a diversified collection of
Amazon indegineous species genetic resources. Proceedings
InterAmerican Society for Tropical Horticulture 42, 140–142.
The sourish-sweet fruit is consumed fresh when ripe and the
more acid fruit is stewed with sugar (see Table A.40 for
proximate fruit composition). It is also used to make
beverages, pickles, curries, tarts, jellies and chutneys. The
fruit exudes a gummy latex when cooked, but the red juice is
clear and consumed as a cold beverage. Green fruit can be
pickled.
The fruit can be used for tanning and dyeing while a paste
of the pounded root serves as a fly repellent. The white or
yellow wood is hard, smooth, and useful for handicrafts and
utensils.
Unripe fruit are used medicinally as an astringent and ripe
fruit as an antiscorbutic, and as a traditional remedy for
biliousness. A leaf decoction is used in cases of intermittent
fever, diarrhoea, oral inflammation and earache. The bitter
root is a stomachic and vermifuge, and contains salicylic acid
and cardiac glycosides. Bark, leaves and fruit contain an
unnamed alkaloid.
MAIN CULTIVARS AND BREEDING
Table A.40. Proximate fruit composition of karanda per 100 g (Source:
Morton, 1987).
Proximate
Water
Energy (kcal)
Protein
Lipid
Carbohydrate
Fibre
Ash
Vitamins
Ascorbic acid
g
83
75
0.39–0.66
2.57–4.63
7.9–12.5
0.62–1.8
0.66–0.78
mg
9–11
Carissa
Botany
This is a woody, straggly, climbing shrub growing 3–5 m high.
It can reach the top of tall trees and numerous spreading
dense branches are set with sharp, simple or forked thorns
(5 cm long) that occur in pairs in the leaf axils. The evergreen
leaves are opposite, oval or elliptic, 2.5–7.5 cm long, dark
green, glossy and leathery on the upper surface and dull light
green underneath. The tubular white, sometimes tinged with
pink, fragrant flowers have five lobes that are twisted to the
left. The flowers occur in terminal clusters of two to 12.
The shrub requires full sun in non-humid tropical regions.
The species has some cold tolerances and can be grown up to
1800 m in the Himalayas. It can grow well on poor, rocky soils
though does better on well-drained fertile soils.
It can bloom and fruit throughout the year. Trimming
encourages new growth and profuse flowering. The fruit are in
clusters of three to ten and can be oblong, ovoid to round,
1.25–2.5 cm long. The purplish-red, smooth skin is thin and
glossy, but tough and turns dark purple to black when ripe.
The juicy red to pink pulp is acid to sweet sometimes bitter
with specks of latex. The flesh may enclose two to eight small
flat, brown seeds.
Horticulture
Karanda is normally propagated by seed, as cuttings are not
easily rooted. Young shoot cuttings can be rooted under
constant mist. Grafting is possible onto seedlings. Karanda is
a good rootstock for carissa (Carissa macrocarpa).
Karanda is often grown as a hedge and can be trimmed to
encourage new growth and profuse flowering. When young the
plant grows slowly, though once established, it grows more
vigorously and becomes difficult to control. When kept
trimmed new shoots are encouraged and it will bloom and
fruit profusely. Flowers occur in terminal clusters on this new
growth.
The pests and diseases of karanda are similar to those of
other carissas. Twig dieback due to Diplodia, stem canker
(Dothiorella spp.), green scurf (Cephaleuros virescens) and algal
leaf spot have been noted.
Some sweet selections have been made from the
considerable wide variation in seedling population. This
variation in fruit quality ranges from oval, dark-purple skin,
red flesh, acid fruit to round, maroon, pink-fleshed, seed
subacid.
Robert E. Paull
Literature cited and further reading
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resource
Systems Inc., Winterville, North Carolina, pp. 221–237.
Verheij, E.W.M. and Coronel, R.E. (1992) Carissa carandas L. In:
Verheij, E.W.M. and Coronel, R.E. (eds) Edible Fruits and Nuts.
Plant Resources of South East Asia No. 2. PROSEA Foundation,
Bogor, Indonesia, pp. 323–324.
Carissa macrocarpa
Natal plum
Carissa (Carissa macrocarpa (Ecklon) A. DC., Apocynaceae) is
known widely as Natal plum and amantungula in English.
Other names include akamba, agamita, agam, amatungulu,
71
agamssa and adishawel. It is native to Natal, South Africa and
is now cultivated worldwide in the tropics and subtropics,
although not in South-east Asia.
World production
As a native to the coastal region of Natal, it is also cultivated in
the Transvaal of South Africa. It was first introduced into the
USA in 1886 by the horticulturist Theodore L. Meade and
again in 1903 by Dr David Fairchild. The latter seed
introduction was distributed for evaluation in different
climatic zones of Florida, the Gulf States and California. It
was introduced into Hawaii in 1905, the Bahamas in 1913 and
the Philippines in 1924. It was widely planted in Israel but
rarely set fruit. It is valued as a protective hedge and the fruit
is a by-product.
Uses and nutritional composition
The fruit is dark red when fully ripe and is eaten fresh when
slightly soft to the touch. It has a sour taste and is eaten whole
without peeling or seeding (see Table A.41 for proximate fruit
composition). If halved or quartered and seeded it may be
used in fruit salads, and as a topping for cakes, puddings and
ice cream if cooked as a sauce or used in pies and tarts.
Stewing or boiling causes the latex to leave the fruit and
adhere to the pot. It can be pickled.
Botany
TAXONOMY
The synonym is Carissa grandiflora A. DC.
This vigorous, spreading woody shrub is up to
5.5 m in height and width. All parts have a gummy white sap.
The thorns (5 cm) are double-pronged and appear on all the
branches. The leathery, evergreen leaves are opposite, ovate,
glossy and 2.5–5 cm long. Shoots grow from the axil in a pair
of small scale leaves.
DESCRIPTION
Table A.41. Proximate fruit composition for Natal plum per 100 g.
Proximate
Water
Energy (kcal)
Protein
Lipid
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Phosphorus
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
g
82
68
0.4
0.9
16.4
0.8
0.4
mg
11
1.3
7
mg
56
0.04
0.06
0.2
25 IU
72
Apocynaceae
The white tubular flowers are 5 cm across, five-lobed, and
borne singly or a few occur at the tips of the branchlets. The
flowers have a sweet fragrance. Some plants bear functionally
male flowers with stamens longer than the style. The stamens
in functionally female flowers are the same length as the style
and have no pollen.
The round, oval or oblong fruit (6.25 cm long, 4 cm across)
has a gummy latex when unripe. The berries’ tender, smooth
skin changes from green to a bright magenta-red crimson
coated with a thin, whitish wax bloom when ripe. The flesh is
juicy, strawberry-coloured with a milky sap enclosing six to 16
small, thin, flat brown seeds that can be eaten.
Cooper, A., Poirier, S., Murphy, M., Oswald, M.J. and Procise, C.
(1998) South Florida Tropicals: Carissa (Natal Plum). Fact Sheet
#FCS 8522. University of Florida, Cooperative Extension
Service, Gainesville, Florida, 3 pp.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resource
Systems Inc., Winterville, North Carolina, pp. 420–422.
Singh, R., Chopra, D.P. and Gupta, A.K. (1982) Some carissas for
western Rajasthan. Indian Horticulture 27, 10–11.
This subtropical to
near tropical species can withstand light frost to ⫺4°C when
well established. Best growth occurs in the full sun. It grows
well in many soil types and has moderate drought and salt
tolerance. Well-drained soils are required, as the tree is
intolerant of waterlogging.
Couma utilis Muell. Arg., Couma macrocarpa Barb. Rodr. and
Couma guianensis Aubl. (Apocynaceae) are Amazonian fruit and
latex species of limited economic value. The species produce a
non-elastic latex that was important in the manufacture of
chewing gum and is still in some demand. The origin of these
species is not known with precision, though the centres of
diversity are in the forests of Central and Western Amazonia
(Cavalcante, 1991).
The common names for C. utilis Muell. Arg. are sorva or
sorvinha (Brazilian Portuguese), couma (Spanish) and milk tree
(English). Couma macrocarpa Barb. Rodr. is called cumã-açu or
leche caspi (Peru), juansoco (Colombia), capirona (Spanish)
and sorva grande (Portuguese). Couma guianensis Aubl. is
commonly known as Poirier de la Guiana (French Guiana).
ECOLOGY AND CLIMATIC REQUIREMENTS
The plant flowers and sets fruit all
year. The flowering peaks in Florida occur from May through
to September. A vegetative growth phase is followed by
flowering and thorn formation. The pollinators are small
beetles and hawkmoths and other night-flying insects.
Unproductive plants that set fruit are apparently self-infertile
as they bear fruit after hand cross-pollination.
REPRODUCTIVE BIOLOGY
Horticulture
Seeds germinate in 2 weeks and seedlings grow
slowly. Air laying, ground layering and shield budding are the
preferred propagation methods. Treated softwood cuttings are
also used. Karanda (Carissa congesta Wight.) is a desirable
rootstock and gives increased yields. Trimming allows shape to
be maintained and a new flowering site to develop.
Seedlings begin to bear fruit in 2 years and cuttings sooner. A
standard, well-balanced fertilizer is adequate. Micronutrients
maybe needed on limestone soils.
PROPAGATION
DISEASES, PESTS AND WEEDS Spider mites, thrips, white flies
and scale attack the young plants. A number of fungal diseases
have been reported in Florida including leaf spot and green
scurf caused by Cephaleuros virescens; leaf spot caused by
Alternaria sp., Botryosphaeria querquum, Fusarium sp.,
Gloeosporium sp., Phyllosticta sp. and Colletotrichum
gloeosporioides. Anthracnose on the fruit is also caused by C.
gloeosporioides. Dieback caused by Diplodia natalensis and
Rhizoctonia solani, and root rot from Phytophthora parasitica
and Pythium sp. also occur.
MAIN CULTIVARS AND BREEDING Selections have been made
in South Africa, California and Florida. These include
‘Fancy’, ‘Torrey Pines’, ‘Gifford’, ‘Extra Sweet’ and ‘Frank’.
Numerous landscape cultivars have been released which are
compact, dwarf and less thorny.
Robert E. Paull
Literature cited and further reading
Cohen, L. and Arzee, T. (1980) Twofold pathway of apical
determination in the thorn system of Carissa grandiflora. Botanical
Gazette 141, 258–263.
Couma guianensis, Couma macrocarpa,
Couma utilis
World production and yield
Couma latex production in Amazonia reached 66,000 t/year
just after World War II, probably from the two central
Amazonian Couma species combined, but currently
production is only about 1000 t/year. This fall in the
production is due in part to the predatory exploitation
practices used, since the trees are cut to maximize latex
extraction. A greater cause is the gradual substitution of
Couma latex by synthetics in the chewing gum industry.
No plantations for latex or fruit production are known to
exist, hence the Couma species are not accounted for in
agricultural statistics for the Amazon basin. Fruit yield of C.
utilis is about 30 kg/plant from 6-year-old trees.
Uses and nutritional composition
The fruit are edible and the trees are beautiful ornamentals
whose potential has been unexploited to date. Occasionally the
pulp of C. utilis is used to make ice creams, juices, sweets and
creams. The composition of 100 g of pulp contains 68.7%
water, 3.6% proteins, 6.9% lipids and 15.8% of other
carbohydrates (Villachica et al., 1996).
Couma utilis and C. macrocarpa produce the raw latex
material for chewing gum that is mixed with latex from species
of the Sapotaceae family, principally Manilkara zapota (L.) P.
van Royen, the chicle. The latex of C. utilis is preferred, but
extractors don’t always maintain purity. The latex of C.
guianensis is not used because it has a bitter flavour.
The latex is extracted from sorva trees by cutting the bark
in helical grooves (similar to the technique used in rubber
Hevea brasiliensis) and the latex exudes in abundance. After
coagulation and moulding into a compact block, it is
Couma
commercialized under the name ‘sorva’. Cavalcante (1991)
reported that the latex is used as a milk substitute to mix with
coffee (which explains the common name in English), and to
mix with cooked maize or banana in a local dish called
‘mingau’. Some indigenous Amazonian peoples mix Couma
latex with banana pulp to cure diarrhoea; others mix the latex
with castor bean oil (Ricinus communis L.) as an antihelminthic remedy. It is used by both native Amazonians and
mixed-blood peasants to caulk canoes and boats, as well as for
whitewashing wooden houses.
Botany
TAXONOMY AND
NOMENCLATURE The
genus Couma
(Apocynaceae) contains 15 species in the neotropics, of which
three are the most important in northern South America: C.
utilis Muell. Arg. (synonym Couma rigila Muell.), C.
macrocarpa Barb. Rodr. and C. guianensis Aubl.
73
FRUIT GROWTH AND DEVELOPMENT Fruit growth starts
immediately after flowering, unless pollination is inadequate,
then the flowers are immediately shed. Ripe fruit are harvested
(or fall) about 103 days after fruit set. Although there are
frequently two flowering peaks, there is normally only one
fruit harvest per year, with peak yield in August (the mid-dry
season in Manaus).
ECOLOGY The three Couma species are native to humid
tropical climate, and are found naturally in the upland forests
of Amazonia, with altitudes less than 500 m. They grow well
in the acid soils that predominate in these regions. Couma utilis
is often found in fields and second growth, but C. guianensis
and C. macrocarpa are generally only found in undisturbed
forests.
Horticulture
Seed propagation is the only method
commonly used, although side veneer grafting has been
reported. The seeds are extracted from completely ripe fruit,
which are recognized by the dark green coloration of the
epicarp and by the soft pulp consistency. The number of seeds
per fruit varies from two to 42, with an average of 12. The
seeds are orthodox, tolerating drying and freezing, and can be
maintained in storage by conventional methods.
Germination is fast and uniform, with emergence starting
22 days after sowing and stabilizing 10 days later, when it
reaches 90%. Initially the seedlings must be protected from
direct sunlight, and after 6–8 months they can be hardened off
and transplanted to the field.
PROPAGATION
In open areas, C. utilis grows to 12 m and in
the forest it may reach 20 m.The bark is smooth, thick and
decorated with big white to brown spots caused by lichens.
When the bark is cut it exudes large amounts of white latex.
The canopy is wide, with dense, dark green foliage, with
abundant low branching in open areas. The inflorescence is
axillary corymbose. The small flowers are hermaphrodite,
gamopetalous and the corolla is pink or light purple, with five
petals. The fruit are small glabrous berries (10–20 g),
succulent, generally green, sometimes brown when ripe,
containing many small seeds. The fruit pulp is edible, with a
pleasant flavour, often reminiscent of sun-dried raisins. The
fruit are harvested from the tree when they are almost ripe,
and often forced to ripen with chemical treatments, and then
tied into clusters of 20–25 fruit to be marketed.
Couma macrocarpa is a large tree, reaching 30–40 m in
height. The trunk has thick, spongy, dark-coloured bark,
decorated with clear spots created by lichens, and exudes a
thick, white, viscous latex with a sweet flavour. Leaves are
simple, up to 20 cm in length by 13 cm in width, the lateral
nerves of up to 20 pairs are regularly parallel while the basal
ones are perpendicular to the central nerve. The fruit is a
round berry, 5–7 cm in diameter, with a green to brown
exocarp, a fibrous, juicy pulp and several seeds.
The tree and fruit morphologies of C. guianensis are quite
similar to C. utilis and C. macrocarpa, however the size of the
C. guianensis tree is smaller than C. utilis. The leaves are equal
in size to those of C. macrocarpa but distinct in secondary
nerve patterns. The fruit of C. guianensis are smaller than C.
macrocarpa and larger than C. utilis.
DESCRIPTION
Couma utilis starts flowering in 3
years in Manaus and 6 years in Belém, Brazil. In the first
years, yields are low and variable. Flowering occurs in the first
half of the rainy season (December–March in Manaus),
followed by a shorter (or equivalent) flowering peak at the end
of the rainy season (May–June/July) (Falcão et al., 2003). The
main pollinators are probably bees, including Eulaema
mocseryi Friese, Eulaema nigrita Lepetier, Xylocopa frontalis
Olivier, Epicharis sp. and Tetrapedia sp. Similar information
does not exist for C. guianensis and C. macrocarpa.
REPRODUCTIVE BIOLOGY
CULTIVATION Although tolerant of poor acid soils, an
orchard should be established on good soils after ploughing,
disking and correcting acidity. Planting pits should measure 50
⫻ 50 ⫻ 50 cm, be fertilized with 5–10 kg of mature manure
and 200 g superphosphate, well mixed. Spacing of 7 ⫻ 7 m for
C. utilis and at least 8 ⫻ 8 m for C. macrocarpa is
recommended. Weeding is essential at least in the first 2 years.
Manure (5–10 kg) should be applied yearly in orchards for
fruit.
MAIN PESTS AND DISEASES The only pest observed attacking
sorva in Brazil is a Coleoptera (Protopulvinasia sp.). The
intensity of infestation is normally low and control measures
are generally unnecessary. The fungus Meliola sp. (called
fumagina) is the only pathogen that may limit the
photosynthetic capacity of leaves, as it covers them with a
powdery black fungal layer.
Danival Vieira de Freitas
Literature cited and further reading
Cavalcante, P.B. (1991) Frutos Comestíveis da Amazônia. Edições
CEJUP, Belém, Pará, Brazil, 279 pp.
Falcão, M. de A., Clement, C.R. and Gomes, J.B.M. (2003) Fenologia
e produtividade da sorva (Couma utilis (Mart.) Muell. Arg.) na
Amazônia Central. Acta Botanica Brasilica 17, 541–547.
Villachica, H., Carvalho, J.E.U., Müller, H.C., Díaz, C.S. and
Almanza, M. (1996) Frutales y Hortalizas Promisorios de la
Amazonia. SPT-TCA No 44. Tratado de Cooperacion Amazonica,
Lima, Peru, 367 pp.
74
Apocynaceae
Hancornia speciosa
mangaba
Mangaba, Hancornia speciosa Gomes (Apocynaceae), a
Brazilian indigenous fruit, is very much appreciated in the
north-east region of the country. The common names in
Brazil are magabeira, magabiba, mangaiba-uva and mangaba
tree-de-minas. The English name is mangaba tree and in the
French caoutchouc de Pernambouc. The mangaba tree
predominates in the Amazon and north-east region of Brazil.
It is also found in Venezuela, Colombia and Peru. The name of
this fruit is derived from the tupi-guarani language, which
means ‘good fruit for eating’. The production of mangaba is
largely from collected fruit and the tree is still not cultivated.
is used to produce rubber and utilized as a popular medicine
for treatment of ulcers and tuberculosis.
Botany
TAXONOMY AND NOMENCLATURE The genus Hancornia
(Apocynaceae) has a single species with six varieties
(Monachino, 1945). The varieties are H. speciosa var. speciosa,
H. speciosa var. maximilliani A. DC., H. speciosa var. cuyabensis
Malme, H. speciosa var. lundii A. DC., H. speciosa var. pubescens
(Nees. et Martius) Muell. Arg. and H. speciosa var. gardneri (A.
DC.) Muell. Arg.
The mangaba tree is a xerophytic plant that has
a semi-deciduous life cycle. The latex-producing tree is
evergreen and grows 2–10 m high, rarely to 15 m in height.
Canopy diameter varies from 4 to 5 m. The reddish bark is
wrinkled. The branches occur as whorls on the main stem.
The simple green leaves are leathery. The main roots grow
deep in the soil. The inflorescence consists of one to seven
white, hermaphrodite double flowers, about 6 cm long that
occur terminally on the latest growth. The fruit is an
ellipsoidal or round berry varying from 2.5 to 6 cm in
diameter (3–51 g). The yellow exocarp is often stained with
red or has red grooves. The soft, fibrous, white fleshy pulp is
often both sweet and acidic, containing 2–15 or as many as 30
seeds. Seeds are 7–8 mm in diameter, disc-shaped, wrinkled
and a clear chestnut brown. The poisonous latex is white and
produced mainly in stem and leaves.
DESCRIPTION
World production and yield
Current studies are underway in the north-east region of Brazil
to quantify production and determine fruit characteristics
suitable for commercial plantings. Species conservation is
threatened by deforestation and the number of wild plants has
declined. Brazilian annual fruit production was estimated to be
1500 t in 2002. About 80% of the production occurs in the first
half of the year, with 55% concentrated between January and
March, lower production occurs during August to November.
Uses and nutritional composition
The fruit is consumed fresh or as juice, compotes, jellies,
liqueurs and vinegars. The mangaba fruit is industrialized in
the form of frozen pulp and is used as juices and in ice cream.
The pulp yield is about 86% of whole fruit. The gummy pulp
is a good source of iron and vitamin C (Table A.42). The latex
Table A.42. Pulp characteristics of the ripe mangaba based on 100 g of fresh
pulp (Source: Alves et al., 2000).
Constituent
Weight total (g)
Seeds (%)
Shell + pulp (%)
Length (mm)
Diameter (mm)
Total soluble solids (%)
Total titratable acidity (%)
Soluble solids per unit acidity
pH
Total soluble sugars (%)
Reducing sugars (%)
Starch (%)
Total pectin (%)
Soluble pectin (%)
Fractional pectin (% – in relation to ISAa)
High methoxlyation
Low methoxylation
Insoluble protopectin
Pectinmethylesterase (UEAb)
Polygalacturonase (UEA)
Total vitamin C (mg/100 g)
Soluble phenolics in water (%)
Soluble phenolics in 100% methanol (%)
Soluble phenolics in methanol 50% (%)
aISA,
insoluble solids in alcohol.
units of enzymatic activity.
bUEA,
Average
19.8
13.2
86.5
33.4
30.1
16.7
1.77
9.51
3.29
13.0
7.72
0.52
0.54
0.24
10.35
1.10
0.29
498.3
17.3
139
0.29
0.33
0.31
ECOLOGY AND CLIMATIC REQUIREMENTS For optimum
development, the mangaba tree requires areas with an annual
mean temperature of about 25°C and annual rainfall of
750–1500 mm. However, the plant tolerates drought and
grows in the hottest periods. The mangaba tree grows better in
sandy acidic soils that are poor in nutrients and organic matter,
and in soils with little water retention. However, the mangaba
tree can develop in deep, well-drained and sandy-loamy soil.
REPRODUCTIVE BIOLOGY Fruit production begins 5–6 years
after planting, although there are reports of fruit production
after 1 year. The tree has two main flowering periods. In the
state of Sergipe in Brazil, flowers occur in August and
February. The flowering period varies from 90 to 120 days. The
flowers produce a faint odour and are open from about 4:30
p.m. to 10:00 a.m. the next day. Bees (Euglossini), hawkmoths
(Sphingidae) and Ninphaelidae (Heliconius) are the most
frequent visitors and are thought to be the main pollinators.
FRUIT GROWTH AND DEVELOPMENT
The fruit begins to ripen
about 112 days from anthesis.
Horticulture
Propagation is from recalcitrant seeds obtained
from ripe fruit. These fruit should be from selected healthy
plants, with good organoleptic characteristics. Immediately after
extraction, the seeds should be washed to totally eliminate the
pulp, and dried under shade for 24 h and usually sown within 4
days. Care must be taken to avoid calcareous substrates, excess
organic matter or overly wet conditions to avoid seedling
PROPAGATION
Saba
disease. Vegetative propagation by grafting can be done on
plants 50–80 cm tall. However, grafting is commercially
difficult due to the very thin stems. Shoot cuttings show poor
rooting response and micropropagation is being tried.
PRUNING The slow growing tree usually produces a broad
canopy and many branches droop down and touch the ground
during wind. The first pruning should be performed when the
plant reaches 1 m in height. After fruiting, it is important to
prune to remove dry, broken and diseased and insect-damaged
branches.
DISEASES AND PESTS The main pest is green aphid (Aphis
gossypii) that attacks the branch terminal leaves causing leaf
rolling. Control can be achieved by bi-weekly spraying of
commercial insecticides. Green cochineal (Coccus viridis) attacks
the new branches and the inferior part of the leaves along with
the main rib. Caterpillars occasionally attack and defoliate young
plants. This pest is controlled by commercial pesticides. Other
insects such as leaf-cutting ant (Atta spp.), arapuá (Trigona
spinipes) and bugs (Theogonis stigma) also attack the plants.
The most serious disease is anthracnose caused by
Colletrotichum gloeosporioides that can cause complete
defoliation of young plants and dark stains on the fruit when
infection occurs at flowering and fruiting. Other diseases
include root rot (caused by Cylindrocladium clavatum and
Fusarium solani), wet rot, damping-off or wilt (Sclerotium
rolfsii Sacc.) and brown spot or leaf spot (Mycosphaerella
discophora Syd. var. macrospora).
Since the ripe fruit
is very perishable it should be harvested at the half-ripe stage.
The optimum harvest is based on the change of colour from
green to clear yellow. Mature green fruit are physiologically
mature and are able to reach the peak of edible quality in 2–4
days. This very short postharvest life limits transport to
markets and commercialization. Abscised fruit, ‘natural fall’,
ripen in 12–24 h and must be consumed quickly and are
regarded as being higher in quality.
Fruit are packed in plastic boxes immediately after harvest
and shipped. Fruit have a high rate of respiration which at
higher temperatures further reduces shelf life and limits
marketing to 3 or 4 days. Storage temperatures of 6–9°C
extend the useful postharvest life to 7–10 days when held in
polyethylene films. At 6°C or lower, the fruit develop chilling
injury. Storage at 8°C is recommended.
HANDLING AND POSTHARVEST STORAGE
GERMPLASM, CULTIVARS AND BREEDING
No select cultivars
are available, though research in the area of genetic resources
and improvement is promising. The Farming Research
Organization at João Pessoa city in the Paraíba State (EMEPAPB) Brazil, has 324 accessions collected from Paraíba,
Pernambuco and Rio Grande do Norte states. The Federal
University of Alagoas (UFAL), in partnership with the
Secretariat of Agriculture, Supplies and Fishes of Alagoas
(SEAP-AL), Rio Largo, has a collection with accessions
collected along the coast of Alagoas state. The Agricultural
Research Company (IPA) from Pernambuco State maintains
on experimental station at Porto de Galinhas that has 125
accessions on 1.3 ha.
Marcelo A.G. Carnelossi and
Narendra Narain
75
Literature cited and further reading
Aguiar Filho, S.P. de, Bosco, J. and Araujo, I.A de (1998) A Mangaba
Tree (Hancornia speciosa): Domesticação e Técnicas de Cultivo.
EMEPA-PB Documentos 24. EMEPA-PB, João Pessoa, Brazil.
Alves, R.E., Filgueiras, H.A.C. and Moura, C.F.H. (2000)
Caracterização de Frutas Nativas da América Latina. Série Frutas
Nativas 9. Funep, Jaboticabal, Brazil.
Anon. (2000) Mangaba. Globo rural, São Paulo. Globo 15 (179), 83–85.
Carnelossi, M.A.G., Toledo, W.F.F., Souza, D.C.L., Lira, M.L.,
Silva, G.F., Jalalli, V.R.R. and Viegas, P.R.A. (2004) Conservação
pós-colheita de mangaba (Hancornia speciosa Gomes). Ciência e
Agrotecnologia 28, 1119–1125.
Epstain, L. (2004) Mangaba: coisa boa de comer. Bahia Agriculture 6,
19–22.
Lederman, I.E., Silva Junior, J.F. da, Bezerra, J.E.F. and Espíndola,
A.C. de M. (2000) Mangaba (Hancornia speciosa Gomes). Série
Frutas Nativas 2. Funep, Jaboticabal, Brazil.
Monachino, J. (1945) A Revision of Hancornia (Apocynaceae). Lilloa,
Tucumán, Argentina, pp. 19–48.
Moura, C.F.H, Alves, R.E, Filgueiras, H.A.C., Araújo N.C.C. and
Almeida, A.S. (2002) Quality of fruits native to Latin America for
processing: mangaba (Hancornia speciosa Gomes). Acta
Horticulturae 2, 549–554.
Narain, N. (1990) Mangaba. In: Nagy, S., Shaw, P.E. and Wardowski,
W.F. (eds) Fruits of Tropical and Subtropical Origin: Composition,
Properties and Uses. Florida Science Source, Lake Alfred, Florida,
pp. 159–165.
Pereira-Netto, A.B. de and McCown, B.H. (1999) Thermally induced
changes in the shoot morphology of Hancornia specioza
microcultures: evidence of mediation by ethylene. Tree Physiology
19, 733–740.
Severino, P.A.F., Bosco, J. and Araujo, I.A. (1998) A Mangaba Tree
(Hancornia speciosa): Domesticação e Técnicas de Cultivo. EMEPA,
João Pessoa, Brazil.
Vieira Neto, R.D. (1994) Cultura da Mangaba Tree. Circular técnica 2.
EMBRAPA-CPATC, Aracaju, Brazil.
Vieira Neto, R.D. (1997) Caracterização física de frutos de uma
população de mangaba trees (Hancornia speciosa Gomes). Revista
Brasileira de Fruticultura 19, 247–250.
Vieira Neto, R.D. (2002) Mangaba. In: Frutíferas Potenciais Para os
Tabuleiros Costeiros e Baixadas Litorâneas. EMBRAPA Tabuleiros
Costeiros/Empresa de Desenvolvimento Agropecuário de Sergipe
– Emdagro, Aracajú, Brazil, pp. 115–140.
Villachica, H., Carvalho, J.E.U., Muller, C.H., Diaz, S.C. and
Almanza, M. (1996) Frutales y Hortalizas Promisorios de la
Amazônia. SPT – TCA 44. Tratado de Cooperacción Amazônica,
Lima, Peru, pp. 227–231.
Wisniewski, A. and Melo, C.F.M. de (1982) Borrachas Naturais
Brasileiras. III. Borracha de Mangaba Tree. EMBRAPA-CPATU
Documentos 8. EMBRAPA-CPATU, Belém, Brazil.
Saba comorensis
rubber vine
This forest liana, Saba comorensis (Boj.) Pichon (Apocynaceae),
grows on other trees in the riparian equatorial rainforest of
Africa. It is native to the Comoros, Ghana, Kenya, Malawi,
Mozambique, Tanzania and Uganda. The common names are
rubber vine in English and mbungo and mpira in Swahili. The
fruit is commonly found in the local markets. It can be used as
an ornamental for its flowers and their fragrance.
76
Apocynaceae
Uses and nutritional composition
The fruit pulp is edible and it makes a refreshing sour drink.
The fruit does not abscise and must be harvested when it
turns yellow. It has a relatively long postharvest life. The stem
yields latex that is an inferior rubber. Bark decoctions are used
to treat rheumatism.
Botany
TAXONOMY AND
NOMENCLATURE The
synonyms are
Landolphia comorensis (Boj.) K. Schum. var. florida (Benth.) K.
Schum. and Saba florida (Benth.) Bullock.
This strong forest liana grows up to 20 m long
on other trees. The stem is lenticillate and exudes a white,
sticky latex when cut. The ovate or elliptical leaves are 7–16 ⫻
4–8.5 cm. The fragrant flowers are borne on many shortstalked terminal or axillary corymbs. The corolla is tubular
with a yellow throat and white petals. The fruit is subglobose
4–8 cm long and 3.5–6 cm wide, greenish when young,
turning orange-yellow when ripe. The yellow flesh contains
numerous brown-black seeds.
DESCRIPTION
The vine is very
abundant in undisturbed forests, coastal areas and around the
Great Lakes region of Africa from sea level to 1250 m. It is
rare in open areas. The area has a mean annual temperature
around 20°C and a mean annual rainfall of 900–2000 mm. The
liana grows on a variety of soil types.
ECOLOGY AND CLIMATIC REQUIREMENTS
REPRODUCTIVE BIOLOGY A vine will not flower every year
and flowering time is not regular among populations. In
Tanzania, flowers occur between February and November with
fruit maturing 10 months later from December to May. The
seeds are dispersed by birds and monkeys.
Horticulture
PROPAGATION The liana regenerates naturally by seed on
fertile moist soils under partial or full shade. It can be
propagated by cuttings and the vine can be coppiced. The
seeds germinate in about 12 days with a high germination rate
in excess of 90%.
Robert E. Paull
Literature cited and further reading
Agroforestree Database. Available at: http://www.worldagro forestry.
org/Sites/TreeDBS/AFT/SpeciesInfo.cfm?SpID=18051
(accessed 20 November 2006).
Food and Agriculture Organization (FAO) (1983) Food and fruit
bearing forest species. 1: Examples from eastern Africa. FAO Forestry
Paper #44/1, FAO, Rome.
Maundu, P.M., Ngugi, G.W. and Kabuye, C.H.S. (1999) Traditional
food plants of Kenya. KENRIK (Kenya Resource Centre for
Indigenous Knowledge) National Museums of Kenya, Nairobi.
Niger, Senegal and Tanzania. The common names are liane
saba or saba in French and mandinka and madd in Wolof.
Uses and nutritional composition
The fruit has a sweet-sour taste and often appears in local
markets during the fruiting season (see Table A.43 for
proximate fruit composition). The leaves are used in Senegal
to prepare sauces and condiments as a salty appetizer. In Côte
d’Ivoire, the latex is used as an adhesive for poison
preparations for arrows as it hardens upon exposure. The
inferior rubber produced from the latex is sometimes used to
adulterate genuine rubber. The leaves are eaten to stop
vomiting and bark decoctions are taken for diarrhoea and food
poisoning.
Botany
AND
NOMENCLATURE The
synonyms are
Landolphia florida var. senegalensis (A. DC.) Hall. f.,
Landolphia senegalensis (A. DC.) Kotschy and Peyr.,
Landolphia senegalensis var. glabrifolia Hua, Saba senegalensis
var. glabrifolia (Hua) Pichon, and Vahea senegalensis (A. DC.)
Pichon.
TAXONOMY
This liana grows up to 40 m long and can often
be shrub-like with a trunk up to 20 cm in diameter. The bark
is rough or scaly. The opposite leaves are one-and-a-half to
three times as long as wide and have a 4–14.5 mm long petiole.
The flowers (3–30) occur in a cyme on a 2.5–6 cm peduncle
and each flower is on a 2.5–8 mm pedicel. The corolla has a
yellow throat with the tube being five to nine times as long as
the calyx. The stamens are inserted 3.5–6 mm above the
corolla base with 0.4–1 ⫻ 0.1 mm stamens and anthers 1–2 ⫻
0.2–0.5 mm. The ovary is often ribbed and has about 30 ovules
and a style that is 1.5–3 mm long. The fruit is 5–15 ⫻
4–10 cm wide with a 1 mm thick wall.
DESCRIPTION
Table A.43. Proximate fruit composition of saba per 100 g.
Proximate
Water
Energy (kcal)
Protein
Lipid
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Phosphorus
Vitamins
Saba senegalensis
saba
This riverine liana, Saba senegalensis (A. DC.) Pichon
(Apocynaceae), from western Africa is found in Burkina Faso,
Côte d’Ivoire, Gambia, Ghana, Guinea, Guinea-Bissau, Mali,
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
%
80
71
0.8
0.2
18.5
1.3
0.5
mg
58
1.0
28
mg
48
15
0.03
5
Trace
Monstera
77
AND CLIMATIC REQUIREMENTS The vine is
commonly found in riverine areas and open woodland from
sea level to 800 m.
ECOLOGY
REPRODUCTIVE BIOLOGY
Flowers are produced all year-
round.
Horticulture
The species grows directly from seed and can regenerate
naturally.
Robert E. Paull
Literature cited and further reading
Agroforestree Database. Available from: http://www.worldagro
forestry.org/Sites/TreeDBS/AFT/SpeciesInfo.cfm?SpID=1799
9 (accessed 20 November 2006).
Burkill, H.M. (1994) Useful Plants of West Tropical Africa. Vol. 2.
Families E–I. Royal Botanical Gardens, Kew, UK.
Leeuwenberg, A.J.M. and van Dilst, F.J.H. (1989) Saba (Pichon)
Pichon, series of revisions of Apocynaceae 27. Bulletin Jardine
National Belgium 59, 189–206.
ARACEAE
Monstera deliciosa
ceriman
The ceriman, Monstera deliciosa Liebm. (Araceae), is the only
aroid grown for its compound fruit. Another English name for
this vine is monstera. In Mexico and Latin America it is
known as piñanona or piña anona, ceriman de México or
balazo. Other names include ojal or huracán in Venezuela;
hojadello in Colombia; Costilla de Adán in Peru; harpón or
arpón común in Guatemala; caroal, liane percee or liane
franche in Guadeloupe; siguine couleuvre in Martinique;
arum du pays or arum troud in French Guiana; and banana de
brejo, banana do mato or fruta de México in Portuguese. As an
ornamental and as a foliage house plant, it is known as
‘Mexican breadfruit’, ‘hurricane plant’, ‘Swiss-cheese plant’,
‘split leaf philodendron’ and ‘windowleaf ’.
World production and yield
The species is native to the wet forest of southern Mexico,
Guatemala, and parts of Costa Rica and Panama. In 1908, it
was reported to be cultivated in Florida, Portugal and Algeria
(Labroy, 1908). Though no longer cultivated on any scale for
its fruit, it is found for sale at roadside markets in southern
Florida (Fig. A.9). It has been spread around the world as an
ornamental foliage plant that can be used indoors or outdoors
generally climbing on some structure or tree.
Fig. A.9. Monstera deliciosa fruit on sale at a roadside stall in Florida.
component (Peters and Lee, 1977). Table A.44 shows the
proximate fruit composition of ceriman.
Acridity associated with the oxalic acid raphides and
associated proteins occurs in the growing fruit and floral
remnants and all other parts of the plant. Aerial roots have
more raphides than soil-borne roots, suggesting tissue
variation. Sensitive individuals suffer throat irritation,
urticaria and anaphylaxis due to the acridity of the raphides.
Ripe fruit lack acridity, although there are individuals that will
suffer from diarrhoea or intestinal gases when eating the ripe
fruit, so that small amounts should be eaten the first time. The
fruit can be consumed when the rind has loosened the entire
length of the fruit, otherwise only the part where the rind is
loose should be eaten.
The aerial roots are used as ropes and to fashion baskets. A
root and leaf infusion is taken for arthritis. A preparation of
the roots is used in some places as a remedy for snake bites.
The inflorescences have a high concentration of the aromatic
amines, tyramine and dopamine, though these are absent from
reproductive organs.
Botany
This fast-growing stout herbaceous vine up to
24 m long spreads over the ground forming mats and can
climb trees. The rarely branched stems are cylindrical
(6.25–7 cm thick) and rough with leaf scars producing
numerous tough aerial roots normally at a node. The oval
leaves are leathery on stiff, flattened petioles (up to 105 cm
long). Mature leaves are up to 90 cm long and 80 cm wide,
deeply cut at the margins to 23 cm strips and perforated on
each side of the midrib with oblong holes of various sizes.
Young leaves are heart-shaped and without holes and these are
the ones used for smaller pots as indoor plants.
DESCRIPTION
Table A.44. Proximate fruit composition of ceriman per 100 g
(Source: Morton 1987).
Uses and nutritional composition
Proximate
The fully ripe fruit pulp is served as a dessert with cream, in
fruit salads or ice creams. The flavour is between pineapple
and banana with sweet, lactone and coconut overtones,
somewhat like piña colada. The flavour is due mainly to ethyl
esters with the only terpene being linalool (Peppard, 1992). It
can be stewed. Ripe fruit soluble solids are about 19% with
7–8 meq/100 g titratable acidity, oxalic acid being a major acid
Water
Energy (kcal)
Proteins
Lipids
Carbohydrates
Fibre
Ash
%
77.9
73.7
1.8
0.2
16.2
0.57
0.85
78
Araceae
Multiple inflorescences arise from the leaf axils on tough
cylindrical stalks. The spadix is cream to tan, surrounded
initially by a waxy white spathe with a pointed apex. This
spadix develops into a green compound fruit (20–30 cm long
by 5–8 cm wide) with a cylindrical form resembling corn on
the cob. The sessile flowers have two-carpellate and twolocular gynoecia. A viscous gum-like nectar is produced near
the stigma that attracts bees. The thick, hard rind of hexagonal
plates (scales) cover the individual segments made up of a
juicy ivory pulp. Between the individual segments are thin,
black membranes. Occasionally, a pale green, pea-size seed
occurs in some of the individual segments of this compound
fruit. The individual fruit of the inflorescence are berries 1 cm
long by 5–6 mm in diameter.
Monstera deliciosa is a
member of the Araceae whose synonym is Philodendron
pertusum Kunth & Bouche, a name which is still used to some
degree in the ornamental foliage plant business as it is closely
related to Philodendron. It has a close relative Monstera
adansonii from Surinam called five-hole plant.
TAXONOMY AND NOMENCLATURE
ECOLOGY AND CLIMATIC REQUIREMENTS This is a tropical
species found all over tropical America in the hot humid
forests, under 600 m altitude with a yearly rainfall above
1000 mm, where it lives under the shade of the large trees on
which it climbs as an epiphytic plant. It does best in semishade under high moisture conditions. Under subtropical
conditions, it can be grown especially as an indoor plant. The
plant grows well in almost any soil that is well drained and rich
in organic matter. It does not stand saline conditions.
REPRODUCTIVE BIOLOGY Suckers begin to produce fruit in
2–4 years while cuttings take 4–6 years to fruiting. Flowering
and fruiting overlap as it requires 12–14 months from the
opening of inflorescence to fruit maturity. Cross fertilization is
required for fertilization and seed initiation.
FRUIT DEVELOPMENT Fruit development can take 12–14
months and it does not require fertilization. As the fruit
matures, the rind takes on a lighter shade and progressively
ripens toward the apex over 5–6 days. The portion eaten is
only that for which the rind can be easily removed.
Horticulture
PROPAGATION It can be raised from seed or by tissue culture,
though generally it is by stem cuttings that take very easily;
these cuttings are stem pieces, 15–30 cm long with two nodes
or stem tips when there is no new leaf unfolding. Suckers with
or without roots can also be used. Seeds are used occasionally
and they should be sown as soon as possible after removal
from the fruit since they are short lived. The plant should be
transplanted close to a tree trunk or a wall so that it can climb.
The harvested fruit kept at room temperature will finish
their ripening in about 5–6 days and the ripe fruit can be kept
in the refrigerator for a week. A pronounced ethylene and
respiratory climacteric occurs during ripening concurrent to a
rapid conversion of starch to sugars. The whole fruit can be
ripened in a plastic wrap, paper or aluminium foil.
DISEASES, PESTS AND WEEDS
Plants used as indoor ornamentals
have usually pest problems while those outside normally do not.
Scale insects, mites, mealy bugs and caterpillars do attack the
plant. In Florida, a grasshopper can cause severe leaf losses in
certain years. It is not regarded as a host to Caribbean fruit fly.
Leaf spots caused by Botriodiplodia theobromae or Acrosperia
fluctuata are frequent when ventilation is poor. Anthracnose,
bacterial soft rot and root rot have also been reported.
MAJOR CULTIVARS AND BREEDING
forms and regional selections exist.
Ornamental variegated
Robert E. Paull and
Odilo Duarte
Literature cited and further reading
Barabe, D. and Chretien, L. (1985) Anatomie florale de Monstera
deliciosa (Araceae). Canadian Journal of Botany 63, 1423–1428.
Eyde, R.H., Nicolson, D.H. and Sherwin, P. (1967) A survey of floral
anatomy in Araceae. American Journal of Botany 54, 478–497.
Geilfus, F. (1989) El Árbol al Servicio del Agricultor Vol. 2. Guía de
Especies. ENDA-Caribe y CATIE, Santo Domingo, Dominican
Republic, 778 pp.
Gould, W.P. and Hallman, G.J. (2001) Laboratory and field
infestation studies on Monstera to determine its host status in
relation to the Caribbean fruit fly (Dipteria: Tephritidae). Florida
Entomologist 84, 437–438.
Hinchee, M.A.W. (1981) Morphogenesis of aerial and subterranean
roots of Monstera deliciosa. Botanical Gazette 142, 347–359.
Labroy, M.O. (1908) Le ceriman de Mexique (Monstera deliciosa
Liebm.) espece fruitiere. Journal of Tropical Agriculture 8,
169–170.
Martin, F.W., Campbell, C.W. and Ruberté, R.M. (1987) Perennial
Edible Fruits of the Tropics – an Inventory. Agriculture Handbook
No. 642. US Department of Agriculture (USDA) – Agricultural
Research Service (ARS), Washington, DC.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resource
Systems Inc., Winterville, North Carolina, pp. 15–17.
Peppard, T.L. (1992) Volatile flavour constituents of Monstera
deliciosa. Journal of Agriculture and Food Chemistry 40, 257–262.
Peters, R.E. and Lee, T.H. (1977) Composition and physiology of
Monstera deliciosa fruit and juice. Journal of Food Science 42,
1132–1133.
Pochet, M., Martin-Tanguy, J., Marais, A. and Martin, C. (1982)
Hydroxycinnamoyl acid amines and aromatic amines in the
inflorescence of some Araceae species. Phytochemistry 21,
2865–2869.
Vásquez, R. and Coimbra, S. (2002) Frutos Silvestres Comestibles de
Santa Cruz, 2nd edn. Editorial FAN, Santa Cruz de la Sierra,
Bolivia, 265 pp.
ARECACEAE – the Palm Family
The palm family, consisting of over 2500 species arrayed
among 200 genera, is known to botanists as the Arecaceae,
though the old name Palmae is still sometimes used in
horticulture. Over 90% of the diversity within the family is
contained within the world’s tropics, and the utility of many
palms in human industry at both subsistence and worldmarket levels makes the Arecaceae the third most economically
important family of plants after the grasses and legumes.
Arecaceae
Morphology, growth and development
Palms, despite their ability to reach tree-like dimensions, have
more in common with lawn grasses, maize and rice than with
oak trees, maples or tropical hardwoods when it comes to their
basic structure and growth processes. Like the former three
familiar plants, palms are monocotyledons and, as monocots,
lack a vascular cambium. Palms therefore have little to no
capacity for secondary growth. Once a palm stem achieves its
maximum girth at a given point on the stem, it is largely
incapable of increasing its stem diameter. Furthermore, the
bundles of conducting tissue within the palm stem, formed
during the earliest stages of stem development, must last the
entire life of the palm. Palms are also not able to repair their
vascular tissue if damaged. It is thus impossible to graft one
part of a palm to another. However, the transport of water and
nutrients throughout the leaf canopy is efficient due to
numerous vascular bundles throughout the trunk. Most
importantly of all, the future of a palm stem rides upon the
continued health of a single actively growing apical meristem
within the bud with little or no ability to regenerate itself. Very
few palms have the ability to branch on their aerial stems in
the normal course of their growth (Tomlinson, 1973) although
occasionally an aberrant individual of an otherwise nonbranching species will produce a branched head. Thus if the
meristem is killed, the entire palm (if solitary) or an individual
palm stem (if clustering) is doomed to eventual death.
Tomlinson (1990) describes the life of a palm as successive
series of semi-discrete, but interdependent episodes or phases:
seed, embryo, seedling, establishment, mature vegetative and
mature reproductive. A palm requires varying horticultural
treatment depending on its phase of development, and may
express more or less tolerance for certain environmental
variables at one given stage versus another. Failure to understand these sometimes subtle but crucial phase differences can
result in damage or even death after various horticultural
operations such as transplanting.
Unlike broadleaved, dicotyledonous trees, palms complete
their increase in stem diameter before elongating. During this
‘establishment phase’, the palm is particularly sensitive to
growth checks or less than optimal environmental conditions.
Since palms do not produce growth rings in their trunks,
there is no absolutely reliable way to determine the age of a
palm without being witness to its entire history. A palm’s age
can be crudely determined by counting the number of:
(i) leaves in the canopy; (ii) adhering leaf bases; and (iii) leaf
scars on the trunk, then observing the rate of leaf production.
If the total number of leaves, bases and scars observed is
divided by the estimated number of leaves produced annually,
a reasonable estimate of the palm’s age can be derived.
Understorey palms are estimated to live for 60–100 years.
Canopy palms have longer lifespans, 100 to 700 or more years
(Morici, 1999).
Palm roots
Shortly after seed germination, the seedling root or radicle of
a palm ceases to function and is replaced by roots produced
from a specialized area of the stem called the root initiation
zone. It is during the establishment phase of its growth that a
young palm fully develops this initiation zone at the base of
79
the stem. Such roots, originating from the stem, are called
adventitious, in contrast to the underground root system of
many dicots, which develop sequentially from a perennial
seedling root. Again, unlike dicots, palm roots emerge from
the stem at maximum thickness; they are incapable of
secondary growth. However, they can branch to four orders,
with third and fourth order roots the primary absorbing
organs. Normally, root development is restricted to the
subterranean portion of the trunk, but on some palms the root
initiation zone extends for some distance above ground level.
Most extreme in this regard are the ‘stilt-root’ palms of wet
tropical rainforests that produce long, thick, support roots
from as high as 2–3 m above the ground. A positive
consequence of the adventitious root system in palms is that
transplanting of mature specimens is relatively easy.
Palm stems
The stems or trunks of palms are as diverse as the palms
themselves, varying in thickness, shape, surface features and
habit. A sizable group of palms (the rattans) even grow as
high-climbing vines into the canopies of rainforest trees.
Many palm stems remain covered with the remains of old leaf
bases for many years; others readily shed their dead leaves. For
the first years of a palm’s life, the stem consists of little more
than overlapping leaf bases shielding the all-important
meristem. Some palm trunks swell noticeably at the base as
they develop with age; others develop conspicuous bulges
further up on the stem. Most tall-growing palms eventually
produce a clear trunk, usually grey or brown, sometimes
green. The trunks of some palms are conspicuously spiny;
these spines are sometimes the remains of fibres that occurred
within the tissue of the leaf bases. The scars left behind by
fallen leaves frequently create a distinctive pattern on the
trunk. This may appears as rings, or, if the leaves incompletely
sheath the trunk, variously shaped scars.
Palm stems vary from about 1 cm to more than 1 m in
diameter. All growth in thickness is primary, that is increase in
diameter precedes the completion of extension growth. Below
the leaf-generating meristem, within the first one to several
nodes of the apex, is a ‘primary thickening meristem’ that
facilitates the expansion growth of the stem across those first
few nodes. All maturation of the stem tissue is basipetal
(downward from the apex). Once expansion growth is
complete within a given segment on the developing stem, little
or no additional cell division takes place in that region.
The outer surface layer on the stem may become thick
walled and sclerotic due to deposition of lignin. A number of
palms do produce cork cells in the outer cortex but there is no
uniform, single-layered, cork cambium. Instead, the phellogen
consists of tiers of meristematic cells. No lenticels or pores
appear on the outer surface, but occasional passive vertical
splits occur on some palm trunks.
The degree of heterogeneity in the anatomy of the palm
stem figures importantly in the use to which palm trunks can
be put. The densely fibrous peripheral ‘sheath’ external to the
vascular bundles, which forms the main mechanical support of
the palm stem, can be so hard as to dull cutting blades (which
is why palms are often left standing when tropical forest is
cleared by hand). Within the palm stem there may occur
80
Arecaceae
stegmata, small cells with uneven walls that contain a single
silica body. These are found in association with fibres and are,
in part, responsible for the hardness of this tissue.
Palms may either be single-stemmed (solitary) or multistemmed (clustering). The basal suckers on a clustering palm
may originate very close to the parent stem or some distance
from it. Both solitary and clustering forms may occur in the
same genus, or even (albeit rarely) in the same species.
Dichotomous branching of the aerial stems, in which the
apical meristem actually divides, is relatively rare in palms,
occurring only in the genera Hyphaene and Nypa. Axillary
branching from lateral meristems is typical of virtually all
clustering or multi-stemmed palms.
Palm leaves
Palm leaves are amazing feats of organic engineering; they are
the largest such organs in the plant kingdom. Leaf development
in the palms is unique in the plant kingdom. Segments (palmate
and costapalmate leaves) or pinnae (pinnately compound)
originate within a continuous tissue mass rather than from
separate primordial units. Two processes are involved: plication
(folding) via differential growth, and segmentation (Uhl and
Dransfield, 1987; Tomlinson, 1990). Many of the details of the
leaf development process in palms are still not well understood.
The way in which the leaf segments (fan palms) or leaflets
(feather palms) are folded around the main vein or midrib is
an important feature of palm leaves that has significance in the
taxonomy and identification of the major groups within the
family. Palms in which the leaflets or segments are folded
upward, forming a ‘V’, are called induplicate. Palms in which
the leaflets or segments are folded downward, forming an
inverted ‘V’, are termed reduplicate. Most of the fan palms
have induplicate leaves, while the majority of the feather palms
have reduplicate leaves. The best place to look to determine
which type of folding characterizes a particular species is right
at the point where the leaflet attaches to the rachis (feather
palms) or, on fan palms, the point where the segments first
split from the rest of the leaf.
All palm leaves consist of three main parts: the blade, the
petiole and the leaf base. Each part has a constrained
mechanical function that lends these large structures the
integrity required to resist wind and other stresses.
The petiole or leaf stem functions mechanically like a
tapered beam or cantilever. The extension of the petiole
through the lamina of a pinnate leaf is called the rachis; in
palmate leaves, the costa. The petiole can be short or long; in a
few species it is apparently obsolete. The petiole of a number
of palm species is toothed along the margins, ferociously so in
some. Generally, the rachis continues through the central
leaflet of induplicate leaves, while the blade of reduplicately
folded leaves is bifid at its apex.
The leaf base is that part of the petiole that sheathes the
stem. It functions mechanically as a stressed cylinder
(analogous to a barrel), and supports almost all of the
mechanical stresses to which the leaf is subject. It initially
develops as a closed tube, but goes through considerable
modification throughout the life of the palm. On many palms,
the base remains attached to the trunk or stem for some time
after the blade and the petiole drop off. In some cases, the
pattern of leaf-base stubs is a distinctive feature of the palm’s
appearance. On other palms, the sheath splits near its base or
disintegrates but leaves behind a mass of fibre of varying weave
and consistency. The tubular leaf bases of some feather-leaved
palms sheath each other so tightly around the stem that they
form a conspicuous neck-like structure called a crown shaft.
Often waxy and smooth, and sometimes strikingly coloured,
the crown shaft can be a structure of singular beauty. Crown
shaft palms are ‘self-cleaning’ in that the tubular leaf base
forms two abscission zones, one at its base and one along the
dorsal surface, that allow a leaf to fall freely after it senesces.
Palm leaf blades basically fall into three main classes: the
fan palms (palmate or costapalmate leaves); the feather palms
(pinnate or bipinnate leaves); and entire or bifid leaves. The
fan palms are described as either palmate or costapalmate. Fan
palm leaves are circular or shaped like an out-stretched hand.
They are divided shallowly or deeply into a variable number of
segments which are often split at the tips themselves. Palmate
and costapalmate leaves are similar in appearance except for
the extension of the petiole into the blade of the costapalmate
leaf. This extension is sometimes referred to as the costa.
Costapalmate leaves are often twisted or folded sharply along
or at the tip of the costa.
Feather palm leaves consist of a series of individual leaflets
arrayed along an extension of the petiole called the rachis.
Pinnately-compound palm leaves are feather leaves that are
only once-compound; that is, there is only a single series of
leaflets. The leaflets may be numerous or few, narrow or broad,
pointed at the tip or blunt and toothed. They can be regularly
arranged along the rachis or attached in groups of several.
Bipinnately-compound palm leaves are twice-compound; that
is, the primary leaflets themselves consist of a system of
smaller secondary leaflets. Bipinnately-compound leaves are
very rare in the palm family, occurring in only a single tribe
(Caryoteae) of the family.
Entire-leaved palms have neither segments nor leaflets.
Instead, the leaf consists of an unsplit blade longer than it is
wide. Bifid leaves are similar in general appearance to entire
leaves but have two lobes in their apical portions. Bifid leaves
are related structurally to pinnate leaves (and essentially
represent an early development stage of a pinnately compound
leaf), while entire leaves are developmentally related to
palmate or costapalmate leaves. The first leaves of many palm
seedlings are entire or bifid, regardless of what type of mature
leaf occurs on the palm. Palms that have either entire or bifid
leaves throughout their life are thus generally assumed to have
retained juvenile characteristics (neotony).
Palms are apparently largely devoid of chemical defences
against arthropod predators and other herbivores. Only a
single palm genus (Orania) is known to contain compounds
that are poisonous to humans. An important defence
mechanism in the palm arsenal appears instead to be the
impressive diversity of spiny structures that may occur on a
variety of vegetative or even reproductive parts.
Reproductive morphology
The duration of time that a palm spends in its mature
vegetative (sterile) phase may be a few years or many. Palms
enter the mature reproductive phase of their lives at the time
Arecaceae
they produce their first inflorescence. Palms are categorized
into two groups based on their reproductive behaviour. A
hapaxanthic palm stem exhibits determinate growth. It grows
vegetatively for a varying period of time, flowers and fruits
after ceasing vegetative growth, then dies. Hapaxanthic palm
stems have two possible phenological patterns: (i) a short
reproductive phase during which lateral inflorescences are
produced in axils of upper bract-like leaves; and (ii) a longer
reproductive phase during which lateral inflorescence buds are
initiated in the leaf axils but are suppressed until vegetative
growth ceases, at which time they mature basipetally. In most
palms, flowering axes mature acropetally (older inflorescences
are lower on the plant). This type of hapaxanthy is known only
in the fishtail palm tribe (Caryoteae). A solitary-stemmed palm
that exhibits hapaxanthy is also referred to as monocarpic
since the palm ceases to live after its fruit mature.
Far more common is the condition known as pleonanthy.
Pleonanthic palm stems grow indeterminately; each shoot is of
potentially unlimited vegetative growth and flowers are
produced on specialized axillary branch systems year after
year throughout the reproductive life of the palm.
Palm inflorescences are classified on the basis of where they
originate relative to the crown of the palm. Suprafoliar
inflorescences originate and stand above the canopy. An
interfoliar inflorescence is produced from leaf axils within the
canopy. An intrafoliar inflorescence is produced below the
canopy of leaves (typical of most palms with crown shafts).
The individual flowers of a palm are generally quite small and
inconspicuous, but are usually borne in such numbers on the
inflorescence that they may be collectively showy. The
inflorescences are palms are frequently quite long and much
branched, but on some species they are short and spike-like. The
palm inflorescence is a branch complex, made up of repeating
units and their associated bracts, with up to five branch orders.
The flowers themselves are borne on terminal segments of
all branch orders called rachillae (singular: rachilla). These are
often very short. Each flower is subtended by a minute scalelike bract. The pedicel (flower stalk) may also bear a small
bracteole. Flowers may be solitary but are usually clustered
with varying complexity.
Bisexual flowers (with both stamens and carpels) are the
exception. Most palms have unisexual flowers, and the plants are
either monoecious (same plant) or dioecious (different plants).
On monoecious palms, flowers of both sexes may be spatially
segregated on the same inflorescence, but only rarely are they
restricted to a single inflorescence of one sex or the other.
Flowers of each sex are often functional at different times.
The simplest palm flowers are trimerous (parts in threes)
with three slightly imbricate (overlapping) sepals, three slightly
imbricate petals, six stamens in two whorls, and three distinct,
uniovulate carpels in a superior ovary. However, there is an
enormous amount of variation in flower structure throughout
the family. The basic structure is actually very rare and is found
in nine genera of the Coryphoideae (the subfamily considered to
be closest to the ancestral palm group (Moore, 1973; Moore
and Uhl, 1982)). The basic plan is modified in many ways via
connation and/or adnation of parts, increase, reduction or loss,
and by differential elongation of the receptacle (Moore and
Uhl, 1973; Uhl, 1988). In many genera, the female flowers do
not have very well developed sepals and petals.
81
The palms exhibit extraordinary diversity in pollen
morphology, even within genera, and may be an indication of
great antiquity for the family, since pollen morphology is often
considered a fairly conservative character in plant evolution.
The basic monocot morphology, that is elliptic shape,
monosulcate (single germination pore), reticulate (net) exine
ornamentation, occurs in at least some members of each
subfamily and is very common in the Coryphoideae
(considered the most primitive subfamily).
In contrast to the often diminutive flowers, the fruit (and
seeds as well) of many palm species are fairly large and
conspicuous. In fact, the largest seed of any plant known on
Earth belongs to a palm, Lodoicea maldivica. A palm fruit
consists of three layers: a thin, superficial exocarp or epicarp
(outer surface); a thick and fleshy or fibrous mesocarp; and a
thick and bony (or thin) endocarp (the innermost layer). The
majority of palm fruit are described as drupes. A drupe is
defined as a fleshy, one-seeded fruit with a thick and sclerotic
endocarp that does not open or split at maturity. Palm fruit
with thin endocarps qualify as berries. Palm fruit with fleshy
mesocarps are generally dispersed by animals; those with a
fibrous mesocarp are sometimes dispersed and may float (e.g. a
coconut). A husked coconut in the supermarket has been
cleaned down to the endocarp. Structural variation in palm
fruit is found in size, shape, surface texture, mesocarp
composition, extent of the endocarp and the number of seeds
they contain. Most palm fruit are smooth, but some can be
scaly, hairy, warty or prickly. For more than a few species, the
display afforded by the ripe fruit is much more conspicuous
than that of the flowers.
The innermost layer of the fruit wall, the endocarp, remains
adherent to the seedcoat in many palm species. In particular,
the endocarp of cocosoid palms (subfamily Arecoideae, tribe
Cocoeae) is fused to the seedcoat. When seeds of many palm
species are cleaned before sowing, usually the endocarp is at
least partially retained.
The seedcoat of some species bears interesting patterns of
ornamentation or sculpturing on its surface. In subfamily
Calamoideae, the seedcoat is fleshy (sarcotesta). Most of the
volume of the seed is taken up by the nutritive tissue called the
endosperm that feeds the developing seedling. In most palm
seeds, the endosperm is liquid early in development but
becomes solid at maturity. Seeds with hollow centres, such as
the coconut, are very rare. The actual embryo of a palm is quite
small, and is located in a small chamber at one end of the seed.
Seed germination
The way palm seeds germinate falls into one of two categories.
In palms with remote germination, the seedling axis develops at
some distance from the actual seed. The first structure to emerge
from the seed is called the cotyledonary petiole. It resembles,
and many people mistake it for, the first seedling root. The
cotyledonary petiole grows downward into the soil (sometimes
very deeply) and swells at its base. From this swelling emerges
the first seedling root (radicle) and seedling shoot (plumule).
The actual cotyledon or seed leaf remains inside the seed
functioning as an absorptive organ called the haustorium. The
haustorium transfers nutrients from the endosperm to the young
seedling. In palm seeds with remote germination, the radicle
82
Arecaceae
persists for some time and produces lateral roots. The seeds of
date palms (Phoenix spp.) have remote germination. A number of
palm species with remote germination (Borassus, for example)
bury the seedling axis deep in the soil.
The other main class of palm seed germination is called
adjacent germination. In these seeds, only a small portion of
the cotyledon emerges from the seed. It appears as a swollen
body abutting the seed surface and is called the ‘button’. The
radicle and plumule emerge from the bottom and top of the
button. In palms with adjacent germination, the first seedling
root or radicle is usually narrow, very short lived, and is quickly
replaced by roots formed at the seedling stem base
(adventitious roots). As with remote germination, a haustorium
remains inside the seed absorbing food from the endosperm.
Some common palms with adjacent germination include
coconut (Cocos nucifera). In coconut, however, the first stages of
germination occur in the fibrous fruit wall that adheres to the
seed and cannot be observed without dehusking the nut.
Taxonomy and classification
Modern classification of the palm family began with the work
of Harold E. Moore (1973), who recognized 25 natural groups
of genera but without any formal taxonomic rank. Uhl and
Dransfield (1987) formalized Moore’s classification and
summarized the wealth of information that had accumulated
on each group of the palms, much of it as a result of Moore’s
extensive fieldwork.
Uhl and Dransfield recognized six subfamilies in the palms,
each in turn divided further into tribes and, in some cases,
subtribes. These are: Coryphoideae (three tribes), Calamoideae
(two tribes), Nypoideae (only one species), Ceroxyloideae (three
tribes), Arecoideae (six tribes) and Phytelephantoideae. These
six subfamilies basically represent six major lines of evolution.
Four characters are most useful in delimiting the
subfamilies: (i) whether the leaf is palmate, costapalmate or
pinnate and either induplicate or reduplicate; (ii) the number
of peduncular bracts on the inflorescence; (iii) the
arrangement of the flowers on the rachillae; and (iv) the
structure of the gynoecium (female reproductive parts).
number of rattan species in the Asian tropics. Only four
genera occur in the New World (Raphia occurs in both
hemispheres); the subfamily is most abundant in eastern Old
World tropics, often in high rainfall or swampy areas. Many of
the species are conspicuously spiny. Calamoideae contains the
only palmate-leaved palms outside of the Coryphoideae, but all
are reduplicate, and the group includes many climbing species
(the rattans). Plants are monoecious, dioecious or polygamous
(bearing both unisexual and bisexual flowers) and are
characterized by tubular inflorescence bracts, a dyad of
bisexual or unisexual flowers as the basic flower cluster, and by
the closely overlapping scales covering the ovaries and fruit. A
number of genera have fleshy seedcoats (sarcotesta).
Nypoideae
This tribe consist of a single monotypic genus, Nypa. Nypa
fruticans is a mangrove palm of Asia and the west Pacific and is
very different from other palms. It grows from a prostrate,
dichotomously branched stem and bears reduplicate, pinnate
leaves and erect inflorescences with a terminal head of female
flowers and branched, lateral spikes of staminate flowers. The
flowers are solitary, both sexes with sepals and petals and the
females are apocarpous. Nypa is one of the earliest
recognizable palms in the fossil record, with occurrences
throughout both hemispheres.
Ceroxyloideae
The Ceroxyloideae are mostly New World palms, but two
genera occur in Madagascar, one in the Mascarene Islands and
one in Australia. The subfamily is distinguished by reduplicate
leaves that are regularly or irregularly pinnate, bifid or entire
(the latter two only occur in Chamaedorea), numerous
peduncular (empty) bracts on the inflorescence, and flowers
arranged singly in rows (acervuli). A crown shaft is sometimes
formed by the leaf sheaths. The species are bisexual,
monoecious or dioecious. New evidence from DNA sequences
indicates that this subfamily may be an artificial one, as has
been suggested on the basis of morphological evidence.
Arecoideae
Coryphoideae
The Coryphoideae is the most diverse as well as the most
unspecialized (‘primitive’) subfamily of the palms. The leaves
of Coryphoideae are palmate, costapalmate, rarely entire or
pinnate, and induplicate. With the exception of the Nypoideae,
all apocarpous palms (with free carpels) are in this subfamily.
The flowers of Coryphoideae are solitary or clustered, but
never in triads. Palms in this subfamily are never strictly
monoecious (separate male and female flowers on the same
plant). Many coryphoid palms are valued ornamentals, and the
hardiest palms in the world belong to this subfamily. The date
palms (Phoenix) belong to this subfamily as well, and the
induplicate-leaved, pinnate or bipinnate tribe Caryoteae (the
fishtail palms) is also allied with the group.
Calamoideae
The Calamoideae, the rattans, contain only 22 genera, but this
represents a quarter of all palm species because of the sizable
This pantropical subfamily is the largest in the palm family
and contains the greatest number of horticulturally and
agronomically significant species such as coconut and African
oil palm. The Arecoideae consists of approximately 113 genera
in six tribes. All are reduplicate and monoecious. The basic
flower cluster of the Arecoideae is the triad, but it is reduced to
a single flower along some portions of the rachillae in many
genera.
Phytelephantoideae
This morphologically isolated subfamily comprises three genera
(Ammandra, Phytelephus and Palandra) in north-west South
America and Panama. Male flowers contain a large number of
stamens and female flowers are clustered in a head-like
structure. The fruit are many seeded. The hardened endosperm
of the phytelephantoid palms is called tagua or vegetable ivory
and is carved into figurines or fashioned into buttons. A sizable
industry for the latter flourished in the early part of this century
Acrocomia
but was rendered obsolete by the plastics industry. The desire to
promulgate sustainable development of rainforest products
coupled with the worldwide ivory trade ban has stimulated
renewed interest in tagua as a marketable commodity.
The palms are a very ancient branch of the monocotyledons,
and differentiated when the continents were closer together.
The present geographic distribution of the family was strongly
influenced by the break up of the ancient continents, Laurasia
and Gondwanaland. The earliest unequivocal occurrence of
fossil palms is from the late Cretaceous (c.80 to 90 million years
ago). The earliest fossil palm leaf remains are costapalmate,
followed by pinnate, then strictly palmate. Fossil evidence
suggests that the subfamilies Coryphoideae, Arecoideae,
Nypoideae and Calamoideae have been distinct for more than 50
million years.
Pollination biology
The small size and fairly bland coloration of most palm
flowers, as well as the copious amount of pollen produced, led
early botanists to conclude that most palms were pollinated by
wind. It is now known that, in fact, most palms are insect
pollinated (Henderson, 1986). The large amount of pollen
produced may offset predation by pollen-feeding insects
(often the very same ones responsible for pollination). Beetle
(particularly weevil) pollination is very common among the
palms. Palms pollinated by weevils tend to mature their female
flowers first (or, if the flowers are bisexual, the stigma will be
receptive before the pollen is shed), have a short flowering
period and tightly spaced flowers. Bee-pollinated palms
typically are protandrous, have a longer period of flowering
and flowers considerably separated on the rachillae. Thrips,
flies and ants have also been reported to pollinate palms. Other
types of animal pollination are rare, but include bats
(Calyptrogyne) and birds (Pritchardia).
The ecological role of palms
Our understanding of the role that palms play in the complex
ecology of tropical forests is probably very incomplete. Fleshy
palm fruit are important sources of food for numerous birds,
rodents and primates in the tropics. Other animals feed on the
carbohydrate- and oil-rich endosperm of the seeds. Some
species of parrots are completely dependent on certain palm
seeds as a high-energy food source during their breeding season
and may fail to breed if the supply of seed is somehow
restricted. Palms provide roosting places for birds and bats, and
the large leaves serve as shelter for numerous smaller animals.
Ethnobotany and economic botany of palms
Many people are surprised to learn that the palm family is
third only to the grass and bean families in economic
importance worldwide. Cocos nucifera and Elaeis guineensis are
major cash crops throughout the world’s tropics, as is the date
palm (Phoenix dactylifera) in subtropical arid zones. Equally
significant are the myriad uses to which local palms are put by
indigenous human cultures wherever palms are found
naturally (Balick and Beck, 1990; Schultes and Raffauf, 1990;
Johnson, 1998). These include exploitation for food, oil, fibre
and construction, as well as medicinal and ceremonial use.
Alan W. Meerow
83
Literature cited and further reading
Balick, M.J. and Beck, H.T. (1990) Useful Palms of the World, a
Synoptic Bibliography. Columbia University Press, New York,
724 pp.
Henderson, A. (1986) A review of pollination studies in the Palmae.
Botanical Review 52, 221–259.
Janos, D.P. (1977) Vesicular-arbuscular mycorrhizae affect the growth
of Bactris gasipaes. Principes 21, 12–18.
Johnson, D. (1998) Tropical palms. Food and Agriculture
Organization (FAO) Forestry Report, Non-Wood Forests
Products No. 10. FAO, Rome.
Moore, H.E. Jr (1973) The major groups of palms and their
distribution. Gentes Herbarium 11, 27–141.
Moore, H.E. Jr and Uhl, N.W. (1973) The monocotyledons: their
evolution and comparative biology. VI. Palms and the origin and
evolution of monocotyledons. Quarterly Review of Biology 48,
414–436.
Moore, H.E. Jr and Uhl, N.W. (1982) Major trends of evolution in
palms. Botanical Review 48, 1–69.
Morici, C. (1999) Death and longevity of palms. Palms 43, 20–24.
Schultes, R.E. and Raffauf, R.F. (1990) The Healing Forest.
Dioscorides Press, Portland, Oregon, 500 pp.
Tomlinson, P.B. (1973) Branching in monocotyledons. The
monocotyledons, their evolution and comparative biology VIII.
Quarterly Review of Biology 48, 458–466.
Tomlinson, P.B. (1990) The Structural Biology of Palms. Clarendon
Press, Oxford, UK, pp. 492.
Uhl, N.W. (1988) Floral organogenesis in palms. In: Leins, P.,
Tucker, S.C. and Endress, P.K. (eds) Aspects of Floral
Development. J. Cramer, Berlin, Germany, pp. 25–44.
Uhl, N.W. and Dransfield, J. (1987) Genera Palmarum. Allen Press,
Lawrence, Kansas, 610 pp.
Acrocomia aculeata
gru-gru palm
Gru-gru palm, macauba or mucaja Acrocomia aculeata (Jacq.)
Lodd. (Arecaceae), is a single-stemmed spiny palm widely
distributed in (sub)tropical America. The fruit mesocarp and
seed both yield useful oils.
World production and yield
Most oil extraction from A. aculeata occurs at the subsistence
level, but some commercial exploitation has taken place in
Brazil and Paraguay. In 1980, 190 t of oil was produced in the
three Brazilian states of Maranho, Ceara and Minas Gerais.
One 18 g fruit yields about 2.4 g of fruit oil and 1 g of seed oil
(Pesce, 1985). In 1971, Paraguay exported 7400 t of kernel oil.
Uses and nutritional composition
The oil pressed from the fruit can be utilized for cooking
without refinement if extracted from very fresh fruit. It is also
processed into soap. The seed (kernel) oil is sweet to the taste
and has been used in the manufacture of margarine. The seed
is 60% fat, 17% saturated (75% oleic and 8% linoleic acids).
The fruit contains 4.6 mg of carotene/100 g and is edible.
Dried seeds yield about 65% oil, and the fruit pulp, 64%
(Balick, 1979). The yellow oil from the mesocarp has higher
84
Arecaceae
iodine content than African oil palm fruit, but the fruit
hydrolizes rapidly after harvest if not extracted quickly,
especially if damaged (Duke, 2001). The seed meal is used as
livestock food. More details of the nutrient composition of the
fruit and seed are listed in Table A.45.
REPRODUCTIVE BIOLOGY Scariot et al. (1991) studied the
reproductive biology of this species near the Brazilian capital
of Brasilia. Plants flowered between August and December
and ripened their fruit between March and June. The flowers
are pollinated by various beetle species.
Botany
Horticulture
TAXONOMY AND NOMENCLATURE Acrocomia at one time
contained many more species, all of which have been reduced
to synonyms of A. aculeata (Henderson, 1994). There is one
other species in the genus, Acrocomia hassleri (Barb. Rodr.)
Hahn, a dwarf palm of southern Brazil and adjacent Paraguay.
The genus is classified in the tribe Cocoeae of the subfamily
Arecoideae. Synonyms for A. aculeata include Acrocomia
sclerocarpa Mart., Acrocomia totai Mart. and numerous others
(Henderson et al., 1995).
PROPAGATION The seed of A. aculeata is surrounded by a
bony endocarp after the fleshy mesocarp is removed. If this is
cracked or otherwise scarified, seeds germinate in 4–6 months.
They are thought to be recalcitrant, and should thus be
planted soon after falling from the tree.
Acrocomia aculeata is a robust, tall-growing
palm, 4–11 m in height. The grey trunk is 10–35 cm in
diameter, often swollen, and armed with spines. A skirt of
dead leaves sometimes persists below the crown of 10–30
greyish-green, arching leaves. The pinnate leaves are several
metres long, with numerous leaflets arranged irregularly
around the rachis in various planes, thus giving the leaf a
plume-like appearance. The leaflets are coated with white
wax on their lower surface. The once-branched spiny
inflorescences emerge from the axils of the leaves. Male and
female flowers are borne on the same inflorescence, the males
at the tip of the rachilla and the females near the base. The
one-seeded fruit are globose, 2.5–5 cm in diameter and ripen
to yellowish orange.
DESCRIPTION
ECOLOGY AND CLIMATIC REQUIREMENTS Gru-gru palm is
widely distributed from Mexico to Argentina, Bolivia,
Paraguay and the West Indies, but is not found in Peru and
Ecuador. It typically occupies open woodlands and savannahs
in areas with seasonal rainfall, mostly at low elevation but
reaching 1200 m in the Andes of Colombia. It is believed to
have been introduced to some portions of its range by human
activity (Henderson et al., 1995). In some areas it forms large
populations. It is quite drought tolerant, and can also
withstand several degrees of frost (at least some of its
ecotypes). It tolerates slightly acid to slightly alkaline soils.
DISEASES, PESTS AND WEEDS In some areas of its range, grugru nut is attacked by a stem-boring weevil, Ryna barbirostris,
which can completely destroy the trunk. The fungus
Phaeophora acrocomiae causes a leaf spot disease.
Alan W. Meerow
Literature cited and further reading
Balick, M.J. (1979) Amazonian oil palms of promise: a survey.
Economic Botany 33, 11–28.
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Henderson, A. (1994) The Palms of the Amazon. Oxford University
Press, New York, 388 pp.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
Markley, K.S. (1956) Mbocaya or Paraguay cocopalm – an important
source of oil. Economic Botany 10, 3–32.
Pesce, C. (1985) Oil Palms and Other Oil Seeds of the Amazon. Edited
and translated by D.V. Johnson. Reference Publications, Algonac,
Michigan.
Scariot, A., Lleras, E. and Hay, J. (1991) Reproductive biology of the
palm Acrocomia aculeata in Central Brazil. Biotropica 23, 12–22.
Aiphanes aculeata
mararay
Mararay, Aiphanes aculeata Willd. (Arecaceae), is a widespread
and very spiny, Neotropical palm with edible fruit and seeds.
The fruit are offered in Colombian markets, but no summary
data are available.
Table A.45. Percentage composition of various fruit and seed products of Acrocomia aculeata (Source: Markley, 1956).
Constituent
Moisture (H2O)
Lipids (oil)
Nitrogen
Protein
Crude fibre
Sugars
Ash
Potassium
Phosphorus
Calcium
Outer fruit
wall (epicarp)
Pulp
(mesocarp)
Pulp, expeller
cake
Shell
(endocarp)
Kernel
(endosperm)
Kernel,
expeller cake
6.65
3.88
0.74
4.62
36.00
–
5.82
2.18
0.10
0.07
4.31
27.94
0.67
4.18
8.82
4.85
10.32
2.18
0.12
0.09
5.26
6.26
0.98
6.12
6.83
5.16
9.16
2.75
0.16
0.10
6.84
2.46
0.31
1.94
49.69
–
3.26
1.02
0.04
0.04
3.17
66.75
2.02
12.62
8.60
1.28
1.98
1.36
0.42
0.08
7.44
7.22
5.50
34.38
11.65
2.80
5.37
1.55
1.14
0.27
Allagoptera
85
Uses and nutritional composition
Uses and nutritional composition
The epicarp and mesocarp of the fruit is rich in carotene
(Balick and Gershoff, 1990). The seed is used in candies
(Bernal, 1992).
The fruit and the seeds of most Allagoptera spp. are edible. No
information on their nutritional value is available.
Botany
Botany
TAXONOMY AND NOMENCLATURE There are about 22 species
of Aiphanes, distributed mostly in the Andean region of South
America, especially Colombia and Ecuador. Most are
rainforest palms, occurring from sea level to 2800 m in
elevation. The genus is classified in the tribe Cocoeae of
subfamily Arecoideae. Aiphanes aculeata was long known as
Aiphanes caryotifolia (Kunth) H. Wendl. Henderson et al.
(1995) list many others. Other common names include cocos
rura (Bolivia), corozo (Colombia, Ecuador) and macaguita
(Venezuela).
Mararay is a solitary-stemmed, feather-leafed
palm growing 4–12 m tall, with stems about 10 cm in diameter
that are armed with sharp fibre spines. The 10–15 pinnate
leaves bear 25–40 pairs of leaflets arranged irregularly in
clusters of four to six that spread in different planes. The
leaflets characteristically widen abruptly towards their tip.
Petioles and leaflets bear spines similar to those of the stem.
The nodding inflorescences emerge from among the leaves,
subtended by a woody spathe, also armed with spines. The
one-seeded fruit are globular, 1.5–2.3 cm in diameter, usually
bright red, but sometime orange or white, with a juicy orange
mesocarp. The bony endocarp surrounding the seed has three
pores near the middle.
DESCRIPTION
ECOLOGY AND CLIMATIC REQUIREMENTS Unlike most of its
congeners, A. aculeata is a denizen of dry forest between 500
and 1500 m in Venezuela, Colombia, Peru, Bolivia and western
Brazil, but absent (at least in the wild) from Ecuador.
Bee and fly pollination has been
reported for many members of the genus.
Alan W. Meerow
REPRODUCTIVE BIOLOGY
Literature cited and further reading
Balick, M. and Gershoff, S. (1990) A nutritional study of Aiphanes
caryotifolia (Kunth) Wendl. (Palmae) fruit: an exceptional source
of vitamin A and high quality protein from tropical America.
Advances in Economic Botany 8, 35–40.
Bernal, R. (1992) Colombian palm products. In: Plotkin, M. and
Famolare, L. (eds) The Sustainable Harvest and Marketing of
Rainforest Products. Island Press, Covelo, California, pp. 158–172.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
TAXONOMY AND NOMENCLATURE Allagoptera consists of four
species of cocosoid palm distributed from eastern Brazil to
Bolivia, Paraguay and Argentina. It is classified in the tribe
Cocoeae of subfamily Arecoideae. Allagoptera arenaria (Gomes)
Kuntze (caxandó, coco da pria, seashore palm) is found along
the Atlantic coast of Brazil from Bahia in the north to Sao
Paulo in the south. It is short stemmed and mostly
underground. The leaves are densely waxy on the underside.
The sweet mesocarp of the fruit is eaten. Allagoptera
brevicalyx Moraes (buri da prioa, caxandó) is endemic to
Brazil. The leaflets of this species are slit at the tip and waxy
grey on both surfaces. Allagoptera campestris is found in
southern Brazil, northern Argentina and Paraguay, from
600 to 1500 m elevation. The immature fruit is edible.
Allagoptera leucocalyx (Mart.) Kuntze has a distribution
similar to that of A. campestris. Both the mesocarp and seeds
are edible.
The stems are mostly subterranenan and short,
solitary, but appearing clustered, often somewhat postrate.
The stem sometimes dichotomously branches and the growing
point is lower in the ground than the base of the stem. The
pinnate leaves usually number four to ten and are frequently
waxy, at least on the underside. Petioles are short. Narrow
leaflets are clustered irregularly and radiate in different planes,
and are often split at the tip. The inflorescences are spicate and
emerge from among the leaves. The flowers are densely packed
on a short, broad flowering branch. On the lower portion of
the rachilla, flowers occur in triads of two lateral staminate
flowers and a central pistillate flower. Only staminate flowers
occur on the upper portion of the rachilla. The yellowishgreen fruit bear one or two seeds, and are densely crowded
into a club-shaped fruiting stem.
DESCRIPTION
ECOLOGY AND CLIMATIC REQUIREMENTS All Allagoptera
spp. come from seasonally dry habitats and do best on sandy
soils. The more southerly occurring species are hardy to
ⳮ4°C. Allagoptera arenaria is extremely salt tolerant. It
grows on dunes or adjacent restinga vegetation on very sandy
soils, often forming large colonies, from 0 to 10 m elevation.
Allagoptera brevicalyx is restricted to Bahia and Sergipe
states on coastal dunes or dry woods near the ocean, from
0 to 20 m elevation. Allagoptera campestris occurs in the
inland cerrado vegetation from 600 to 1500 m elevation and
A. leucocalyx is found on dry, rocky, sandy soils from 200 to
700 m elevation.
Horticulture
Allagoptera spp.
Allagoptera (Arecaceae) consists of four species of cocosoid
palm distributed from eastern Brazil to Bolivia, Paraguay and
Argentina. These species are not produced commercially.
Allagoptera arenaria is extremely salt tolerant, and makes a fine
ornamental in tropical and subtropical coastal areas.
Propagation is by seed, which germinates erractically over 3–6
months. High temperature helps speed germination. All are
slow-growing palms.
Alan W. Meerow
86
Arecaceae
Areca catechu
betel nut
Betel nut, Areca catechu L. (Arecaceae), is a single-stemmed,
pinnate-leafed palm cultivated extensively throughout tropical
Asia and Africa for its seed, which is chewed as a breath
freshener. Also the nuts are said to have stimulant properties.
It is estimated that between 10 and 25% of the world’s
population chew betel nuts with some frequency.
World production and yield
A single betel nut palm produces two to three fruiting stems
annually, each containing 150–250 fruit; larger-fruited varieties
will produce fewer. One hundred fruit weigh 1.5–2.3 kg. A
hectare with 1000 palms yields 450,000–750,000 fruit, which is
processed down to 15–25 cwt of dried areca nuts (the seed). An
average annual yield of nuts is estimated at 17.5 cwt/ha (Duke,
2001). India and Pakistan are major producers, but almost all of
the production is locally consumed. In 1969 and 1970, there
were reportedly 1 million ha of betel nut in production in
Pakistan alone (Duke, 2001). It is also produced throughout
Malaysia and the Philippines, and the latter exports a large
quantity to India, as does Sri Lanka. An estimate of world
production is 184,000 ha, producing 191,000 t/year (Bavappa
et al., 1982).
Betel nut palm is a single-stemmed palm with
a slender trunk, conspicuously ringed by the scars of fallen
leaves, capable of reaching 30 m in height (Fig. A.10), but
often smaller, especially when grown in full sun. The
sheathing leaf bases form a smooth, greyish crown shaft. The
pinnate leaves are 1–1.5 m long, with several dozen obliquely
toothed leaflets. The apical pinnae are fused together to form a
fishtail-like shape. The branched flower stems emerge from
below the crown shaft, and are roughly 1 m long. The small
flowers are arranged in triad clusters of two staminate flowers
flanking a slightly larger pistillate flower. The orange or red
drupes are 5–6 cm long and 4–5 cm wide, varying from
spherical to somewhat flattened, and contain a single seed.
DESCRIPTION
ECOLOGY AND CLIMATIC REQUIREMENTS Areca catechu is
thought to be originally native to the Malaysian peninsula, but
its use has resulted in a long history of cultivation and
naturalization through south and South-east Asia. It requires a
warm, humid tropical climate to thrive, and is damaged at 0°C.
In the tropics, elevations below 1000 m suit the palm best. It
performs best with annual rainfall in excess of 500 mm, but
responds favourably to irrigation in drier climates. In the wild,
it is usually found at the margins of rainforest or below the
Uses and nutritional composition
Dried areca nuts are either ground into powder or sliced. Slices
are typically enclosed within a leaf of betel pepper (Piper betel),
mixed with various spices (especially cloves), some lime and
sometimes tobacco. Powdered betel may be carried about in a
pouch and taken like tobacco snuff. The nuts are chewed at social
occasions and after meals. They sweeten the breath, especially if
mixed with spices. Chewing the nuts stains saliva red and
sustained use will turn the gums and teeth black. A large number
of folk medicinal uses for the nuts are also recorded (Duke,
2001). Black and red dyes are also manufactured from the nuts.
The fibrous mesocarp is an important industrial product used
for insulation and particle board.
Almost 400 calories are contained within 100 g of fresh
betel nuts, along with 6.0 g protein, 11 g fat, 70 g
carbohydrates, 540 mg calcium, 63 mg phosphorus, 5.7 mg
iron, 76 mg sodium, 446 mg potassium, 0.17 mg thiamine,
0.69 mg riboflavin, 0.6 mg niacin and trace amounts of
vitamin C. The seeds also contain vitamin A. A number of
unique alkaloids are contained within the seed, and some
health authorities claim that frequent habitual use has negative
health effects (Van McCrary, 1998). Arecaine and arecoline are
two of these alkaloids, sometimes compared to nicotine in
their stimulating, mildly intoxicating and appetite-suppressing
effects. Epidemiologic studies have linked the use of betel nut
to various oral cancers, though this may be more a factor of
concurrent abuse of tobacco and alcohol.
Botany
TAXONOMY AND NOMENCLATURE Areca catechu L. is one of
about 60 species of mostly small or modest-sized monoecious
palms found in the understorey of Asian tropical forests. It is
classified in the palm subfamily Arecoideae, tribe Areceae.
Fig. A.10. Areca catechu palm and insert showing fruit (with permission
from Sitijati Sastrapradja from Palem Indonesia, Lembaga Biologi
Nasional, 1978; fruit photograph provided by Ken Love).
Arenga
canopy; in time it may penetrate the upper strata of vegetation.
It colonizes secondary forest readily. The palm grows on a
variety of soils, as long they have good water-holding capacity.
REPRODUCTIVE BIOLOGY Areca catechu is monoecious, with
separate staminate and pistillate flowers borne on the same
inflorescence. The palms are insect pollinated. Little is
recorded in detail about the species’ reproductive biology. The
palms are diverse in flowering and fruiting season depending
on the area in which they are grown. Palms usually begin to
flower and fruit after 7 years from seed.
FRUIT DEVELOPMENT Betel nuts mature in 6–8 months after
pollination of female flowers. The fruit are harvested typically
when their full colour (yellow, orange or red, depending on
source) is developed. Optimum fruit production usually takes
10–15 years, and the palms will generally bear heavily for
30–75 years of their life (60–100 years is the best estimate
available).
Horticulture
Seeds are the only means of propagation of
betel nut. Ripe fruit are gathered, sun-dried for several days or
in shade for a week. Seeds are sown either in rows or in groups
of several dozen in pits of prepared soil. Plantain leaves are
sometimes used as germination containers. Of course, the
seeds may be sown in trays or other nursery containers as well,
but this is seldom done in the main production areas. Seeds
germinate erratically in as little as 6 weeks and upward to a
year. Typically 1–2-year-old seedlings are planted into their
permanent sites at a density of 1000–1500 palms/ha. Some
degree of shade is generally provided by bananas or fruit trees.
Betel nut is thus an important component of agroforestry
systems in Asia.
PROPAGATION
Betel nut palms are tolerant
of many different soils types, except very sandy soils with low
water-holding capacity. A pH of 6.3–6.5 is probably ideal,
though the palm tolerates acidity to at least 5.0 and alkalinity
to 8.0. Year-round irrigation is essential where a marked dry
season occurs. Applications of manure or inorganic NPK
(nitrogen, phosphorus, potassium) fertilizers have proven
beneficial (Duke, 2001).
NUTRITION AND FERTILIZATION
DISEASES, PESTS AND WEEDS
Koleroga disease (Phytophthora
omnivorum var. arecae), which attacks the fruit, and foot rot
(Ganoderma lucidum), which infects the base of the stem, are the
two most serious diseases of betel nut palm. Innumerable other
fungal blights, mostly leaf spot diseases, have been reported on
A. catechu. Thielaviopsis paradoxa is an endophytic fungus that
causes the stems to split lengthwise. Bacterial decline caused by
Xanthomonas vasculorum has been reported. Several nematode
species are a problem in Thailand. Insect pests include
rhinoceros beetle (Orcytes rhinoceros), a caterpillar that feeds on
the leaves (Nephantis serinopa) and a borer (Arceerns fasiculatus).
MAIN CULTIVARS AND BREEDING In some of the areas
throughout its natural range and where it has been introduced,
betel nut populations have been selectively propagated for
87
various characteristics such as fruit shape and size, nut flavour
and, presumably, concentration of biologically active
constituents (Duke, 2001). Years of selection have resulted in
those forms coming true from seed for those specific
characteristics (i.e. they are likely to be homozygous for the
genes promoting these traits). These selections have at times
been given formal botanical rank, but are probably better
considered as cultivars. ‘Deliciosa’ has been applied to
varieties with very mildly flavoured nuts. ‘Batanensis’
produces shorter and thicker stems. ‘Communis’ has orangered fruit.
Alan W. Meerow
Literature cited and further reading
Bavappa, K.V.A., Nair, M.K. and Kumar, T.P. (1982) The Arecanut
Palm (Areca catechu Linn.). Central Plantation Crops Research
Institute, Kasaragod, India.
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Van McCrary, S. (1998) The Betel Nut: An Emerging Public Health
Threat? University of Houston Health Law Institute. Available
at: http://www.law.uh.edu/healthlawperspectives/HealthPolicy/
980908Betel.html (accessed 20 November 2006).
Arenga pinnata
sugar palm
Sugar palm, Arenga pinnata (Wurb) Merr. (Arecaceae), is a
solitary palm from Asia widely cultivated for its starchy pith
and sugary sap which contains much higher levels of sucrose
than most sugarcane varieties, as well as the industrially
valuable fibre extracted from the leaf sheaths. A minor use for
the palm is the extraction of the edible endosperm of partially
ripened seeds for preparation as sweetmeats.
Uses and nutritional composition
Half-ripe fruit are first peeled to remove the fruit wall which
contains high levels of irritating calcium oxalate crystals. The
seeds are washed and the seedcoat removed. The endosperm is
first soaked in lime water for a few days then boiled in sugar
solution, sometimes flavoured with various spices. The juice of
the ripe fruit is used as a fish poison and should never be
consumed. While the stem starch and the sugary sap of A.
pinnata has been analysed, no nutritional information on the
seed is available.
Botany
TAXONOMY AND NOMENCLATURE Arenga pinnata is one of
about 17 Asiatic species in the genus. It is classified in the tribe
Caryoteae, which was once placed in the subfamily Arecoideae,
but has been shown to be more closely related to the subfamily
Coryphoideae, which consists of mostly fan-leafed palms.
Arenga saccharifera Labill. is a synonym.
Sugar palm grows up to 15 m in height with a
solitary stem up to 0.5 m in diameter (Fig. A.11). The remains
of old leaf sheaths cover the stem with a dense coat of dark
brown fibres and spines. The induplicate, pinnate leaves are
erect, nearly 10 m long with 100 or more, dark-green leaflets
with a white waxy coating on their underside. The stout
DESCRIPTION
88
Arecaceae
long period of juvenility (6–12 years), and lives for about
another 15 years after flowering begins. Flowering and fruiting
occurs throughout the year in the lowland tropics.
FRUIT DEVELOPMENT
It takes 2 years for the fruit of A.
pinnata to mature.
Horticulture
PROPAGATION Sugar palm is propagated from seed, which
germinates in 3–12 months.
DISEASES, PESTS AND WEEDS Sugar palm is rarely troubled
by pests or disease. The stem-rotting fungus Ganoderma
pseudoferreum has been reported to affect A. pinnata. In Southeast Asia, the rhinoceros beetle, Orcytes rhinoceros, has been
known to feed on the foliage.
While some degree of
regional selection has been applied to sugar palm, mostly
towards reducing the juvenile period, no rigorous breeding or
cultivar evaluation programme has ever been conducted.
Alan W. Meerow
MAIN CULTIVARS AND BREEDING
Literature cited and further reading
Fig. A.11. Arenga pinnata palm showing multiple inflorescences
(with permission from Sitijati Sastrapradja from Palem Indonesia,
Lembaga Biologi Nasiona, 1978).
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Astrocaryum aculeatum
petioles are 1.5–2 m long and covered with fibre. The leaflets
are about 1 m long, 5–8 cm wide, and usually with jagged,
‘fishtail’ tips. Large pendent inflorescences emerge from the
leaf axils sequentially from bottom to top, consisting of many
long, slender rachillae drooping from a short peduncle.
Typically, the flowers are arranged in a triad of two staminate
flowers flanking a central pistillate flower, but unisexual
inflorescences are sometimes produced, especially those
developing in the upper axils. The flowers are stalkless and
purple, sometimes scented. The ovoid fruit is 5–6 cm in
diameter, yellow to yellowish brown with a fleshy white
mesocarp that is irritating to the skin. Each contains two to
three grey-brown seeds, 2.5–3.5 cm long and 2–2.5 cm wide.
ECOLOGY AND CLIMATIC REQUIREMENTS Arenga pinnata is
thought to have originated in Indonesia but is now widely
distributed through India, Sri Lanka, southern China, Southeast Asia, New Guinea and Guam, from sea level to 1200 m
elevation. It is a rainforest palm in its natural state, and thrives
on rich, moist soil in partial shade, but is adaptable to drier areas
and to full sun. Sugar palm is tolerant of temperatures below
freezing but is damaged at –2°C. Specimens have been known to
recover from temperatures as low as –4°C (Duke, 2001).
The flowers of sugar palm are insect
pollinated, though details are lacking in the literature. In its
natural forest habitat, the fruit are dispersed by fruit bats,
civets and probably other small mammals. The sugar palm is a
hapaxanthic palm, and begins to decline (and eventually dies)
after the uppermost inflorescences set fruit. The palm has a
REPRODUCTIVE BIOLOGY
chonta palm
Chonta palm, Astrocaryum aculeatum G.F. Mey (Arecaceae), is
a solitary palm widely distributed in the Amazon region, and
often associated with former or present human habitation. Its
edible fruit is valued locally. The fruit are sold in markets in
Amazonian Colombia and Brazil.
Uses and nutritional composition
The orange, fleshy mesocrap of the fruit is consumed. The
fruit contains 3.5% protein, 19.1% carbohydrate and 16.6%
fat. In addition, the level of vitamin A is reported to be 50,000
IU/100 g of pulp, three times that of carrots. The oil pressed
from the seed is chemically similar to coconut oil. Local
people steam the fruit and eat them, or crack open the seed
inside young fruit to drink the clear, sweet endosperm. Other
species of Astrocaryum with edible fruit or seeds are listed in
Table A.46. No information on their nutritional value is
available. Astrocaryum jauari is a waterside plant, and the fruit,
which drop into the watercourses along which the palms grow,
are an important food source for river fish. Use of the seeds
for fish food in aquaculture projects has been proposed
(Borgtoft Pedersen and Balslev, 1992).
Botany
There are perhaps 18
species of Astrocaryum (Henderson et al., 1995). Astrocaryum
princeps Barb. Rodr. and Astrocaryum tucuma Mart. are
common synonyms for A. aculeatum; Henderson et al. (1995)
list others (Table A.46). The genus is classified in the tribe
Cocoeae of subfamily Arecoideae.
TAXONOMY AND NOMENCLATURE
Table A.46. Other species of Astrocaryum with edible fruit.
Species
Common name
Astrocaryum acaule Mart.
Origin
Uses
Characteristics
References
Espina, corozo, palmeira lú, Guiana, Amazon
tucumai
Edible pulp and nut
Fouqué, 1973; Martin et al., 1987
Astrocaryum campestre Mart.
Jarivá, tucum
Brazil, Bolivia
Edible fruit
Astrocaryum chambira Burret
Chambira
Western Amazon
Edible fruit
Astrocaryum gynacanthum Mart.; Syn.:
Astrocaryum munbaca Mart.
Coco de puerco (Colombia), Amazon mostly east and
cubarro (Venezuela);
centre, on non-inundated
mumbaca
lowlands
Mesocarp occasionally
eaten
Astrocaryum huaimi Mart.
Chontilla
Pulp and nut oily, edible
Stem very short and underground; five
to nine leaves, leaflets 55–103 per side
in irregular clusters; fruit obovoid,
2.5–3 cm long, 1.5–2 cm wide,
yellow-green to orange
Stem short, underground; three to six
leaves, leaflets 17–43 per side, in
irregular clusters; fruit 3–3.5 cm long,
2–2.5 cm wide, orange or yellow-green
Stem to 30 m tall; leaves 9–16, erect,
to 5 m long; inflorescence solitary and
erect; large, light green fruit,
mesocarp fibrous
Stems clustered, to 12 m, 3–10 cm
wide; fruit obovoid 2.5–3 ⳯
1.2–1.5 cm, densely crowded, bright
orange; mesocarp orange, floury
Spiny stem to 10 m, clustered or
solitary; fruit obovoid 3–4.5 ⳯ 2–3cm,
yellow to orange
Astrocaryum jauari Mart.
Jauari
Astrocaryum macrocarpum Huber
Palmeira tucumã-assi
South-western periphery of
the Amazon: Peru, (Madre
de Dios), eastern Bolivia to
Mato Grosso in Brazil
North of South America, on
regularly flooded sandy soils
Brazil
Pulp has a flat taste;
edible nut (oil)
Pulp
Mexico to Central America
Inflorescence and
endosperm eaten; leaves
for thatching; trunks as
tool handles
Mesocarp and nut
occasionally eaten;
leaves and stems
in house construction
Astrocaryum murumuru Mart.
Chonta (Bolivia),
chuchana (Colombia,
Ecuador), huicongo
(Peru), murumuru,
muruí (Brazil)
Amazon, periodically
flooded areas, up to 900 m
in eastern Andes
Astrocaryum princeps Barb. Rodr.
Tucumã-açú
Brazil
Pulp
Astrocaryum vulgare Mart. Syn.:
Astrocaryum awarra de Vriese,
Astrocaryum guianense Splig. ex Mart.,
Astrocaryum segregatum Drude,
Astrocaryum tucuma of Wallace,
Astrocaryum tucumoides Drude
Tucuma; tucumã,
tucumã do Pará,
cumari, acquiere,
awarra, palmier
tucuman, aouara
Amazon
Juice, oil
Villachica, 1996
Henderson et al., 1995
Fouqué, 1973; Henderson et al.,
1995
Fouqué, 1973; Martin et al., 1987
Martin et al., 1987
Henderson et al., 1995
Henderson et al., 1995;
Silva, 1996
Martin et al., 1987
Fouqué, 1973; Cavalcante, 1991;
Henderson et al., 1995;
Villachica, 1996
Astrocaryum
Astrocaryum mexicanum Liebm. ex Mart. Lancetilla (Honduras),
chocho, chichón (Mexico)
Multiple spiny stems to 15 m; fruit ovoid
4–5 ⳯ 2.5–3 cm, greenish orange
Syn. of A. aculeatum according to
Henderson et al., 1995
Stem solitary, to 8 m, 2.5–8 cm wide;
fruit ellipsoid to obovoid,
4–6 cm, densely covered with
short black spinules, brownish
Spiny stem, solitary or clustered, to
15 m; fruit obovoid, 3.5–9 ⳯
2.5–4.5 cm, brown, tomentose to
scarcely to densely covered with
short black spinules; mesocarp fleshy
or fibrous
Syn. of A. aculeatum according to
Henderson et al., 1995
Usually clustering, stems 410 m tall;
leaves 8–16, erect, leaflets 73–120 per
side, irregularly clustered;
fruit globose, 4–5 cm and 3–3.7 cm
wide, orange
Fouqué, 1973
89
90
Arecaceae
Chonta palm is a solitary-stemmed feather
palm growing 20 m or more tall. The trunk may reach 25 cm in
diameter and is covered with long black spines. There are
between six and 15 erect leaves extending to 6 m in length.
There are 73–130 pairs of leaflets arranged in irregular clusters
that spread in various planes. The inflorescence emerges from
the leaves and is erect, subtended by a spiny bract. The
flowering branches have only two to four female flowers at their
base; the rest of the flowers are male. The globose fruit are 4–5
cm in diameter, yellow-orange or yellow-green.
DESCRIPTION
Chonta palm is
widespread throughout the central and eastern Amazon of
Colombia and Venezuela, Trinidad, the Guianas and Brazil,
below 1000 m. It is rarely found in forests of the Amazon, but
is much more common in deforested areas, and is thought to
have been introduced in some parts of its range. It has little
tolerance of frost.
ECOLOGY AND CLIMATIC REQUIREMENTS
Consiglio and Bourne (2001) found
that beetles were the most efficient pollinator of chonta palms.
REPRODUCTIVE BIOLOGY
World production and yield
No data are available. Yields are said to vary, which has
prevented sustainable large-scale processing. Fruit and/or nuts
are exported from Central America for soap manufacturing.
Uses and nutritional composition
The seed of the cohune palm yields a non-drying oil used in
food, for lighting and soap production. The fruit are sometimes
made into sweetmeats and also fed to livestock. Per 100 g, the
seed contains 6.8 g protein and 52.2 g fat (Duke, 2001).
Botany
The cohune palm was
formerly known as Orbignya cohune (Mart.) Dahlgren ex Stanl.
Palm species formerly recognized as the separate genera
Maximiliana, Orbignya and Scheelea have been combined with
Attalea (Henderson et al., 1995). It is classified in the tribe
Cocoeae of subfamily Arecoideae. Other Attalea species with
edible fruit or seeds are listed in Table A.47.
TAXONOMY AND NOMENCLATURE
The cohune palm grows 16–20 m tall,
producing a trunk up to 30 cm in diameter that is
conspicuously ringed with old leaf scars after the persistent old
leaf bases finally fall. The 15–30 leaves are as much 10 m long,
and fairly erect, arching ultimately at their tip. Each leaf bears
30–50 leaflet pairs, each up to 45 cm long, rigid and dark
green. The flower stems emerge from among the leaves,
contained by a woody bract in bud. The branches of the
staminate inflorescences are up to 15 cm long; staminate
flowers have as many as 24 stamens. The fruit are borne in
pendulous clusters; each fruit is 4–8 cm long, 3.3–4.5 cm wide,
brown or yellow-brown and containing one to three seeds.
DESCRIPTION
Horticulture
Seeds of chonta palm can take more than a year to germinate.
Alan W. Meerow
Literature cited and further reading
Borgtoft Pedersen, H. and Balslev, H. (1992) The economic botany of
Ecuadorean palms. In: Plotkin, M. and Famolare, L. (eds)
Sustainable Harvest and Marketing of Rain Forest Products. Island
Press, Washington, DC, pp. 173–191.
Cavalcante, P.B. (1991) Frutas Comestíveis da Amazônia, 5th edn.
Edições CEJUP, Belém, Brazil, 279 pp.
Consiglio, T.K. and Bourne, G.R. (2001) Pollination biology and
breeding system of the palm Astrocaryum vulgare in Guyana: a
test of predictability of syndromes. Journal of Tropical Ecology 17,
577–592.
Fouqué, A. (1973) Espèces fruitières d’Amérique tropicale. Fruits 28,
290–299.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
Martin, F.W., Campbell, C.W. and Ruberté, R.M. (1987) Perennial
Edible Fruits of the Tropics – an Inventory. Agriculture Handbook
No. 642. United States Department of Agriculture, Agricultural
Research Service, Washington, DC, 247 pp.
Silva, S. (1996) Frutas no Brasil. Empresa das Artes, São Pablo, Brazil.
Villachica, H. (1996) Frutales y Hortalizas Promisorios de la Amazonía.
Tratado de Cooperación Amazónica, Lima, Peru, 374 pp.
Attalea cohune
cohune
Cohune, corozo, Attalea cohune Mart. (Arecaceae), is a tallgrowing single-stemmed, pinnate-leafed palm native to the
wet Atlantic coast lowlands from Mexico to Honduras and
Belize, but cultivated as far south as northern South America.
The seeds are a source of a high-quality oil.
The cohune palm
occurs from sea level to 600 m on a variety of soil types from
southern Mexico to Belize, but also sparingly in northern
South America. It is most abundant in wet rainforest on rich
soils, but is found in disturbed, open areas. Some populations
are undoubtedly relics of cultivation. Little or no tolerance of
freezing temperatures is to be expected. Wide tolerance of pH
is reported (Duke, 2001).
ECOLOGY AND CLIMATIC REQUIREMENTS
Horticulture
PROPAGATION Cohune palm seeds remain viable for 6
months. They should be planted about 5 cm deep and kept
moist. A spacing of 100 trees/ha has been recommended.
DISEASES, PESTS AND WEEDS
Fruit are often parasitized by
the larvae of bruchid beetles. Fungal pathogens that have been
reported include Achorella attaleae, Gloeosporium palmigenum
and Poria ravenalae (Duke, 2001).
Alan W. Meerow
Literature cited and further reading
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
Table A.47. Some additional Attalea species with edible fruit or seeds (Source: Henderson et al., 1995).
Species
Synonym
Common names
Distribution and habitat
Descriptive notes
Edible product
Taparin, táparo, igua, mangué
Atlantic slopes of Panama and
NW Colombia; lowland rainforest
below 500 m
Subterranean stems; leaves 8–15,
leaflets in irregular clusters, apical
leaflets partially joined along margins;
fruit light brown, 6–8 cm long,
3.5–5 cm wide
Liquid endosperm is drunk; seeds
eaten
Táparo, almendrón
Colombia, dry to wet forested
ravines, 1000–1600 m
Subterranean stems; leaves 10–15,
leaflets two-ranked; fruit 6–9 cm long,
5 cm wide
Seeds edible; high oil content
Attalea butyracea
Scheelea butyracea
(Mutis ex L. f.) Wess. Boer (Mutis ex L. f.) H. Karst.
ex H. Wendl.; many
others listed in
Henderson et al. (1995)
Palla, jací, palama real, corozo,
canambo, coquito, coyol real,
shebon, yagua
Widespread from Mexico to
northern South America in
seasonal and wet forest as
well as savannah and
disturbed areas, usually
below 300 m
Stem to 20 m and 25–50 cm broad;
Fruit and seed edible
leaves 15–35, petiole and sheath spiny,
leaflets two-ranked; fruit 4.5–8.5 cm long,
3–4.5 cm wide, orange, yellow or brown
Attalea colenda (O. F.
Cook) Balslev &
Henderson
Palma real
SW Colombia and western
Ecuador, lowland rainforest or
deciduous forest, below 900 m
Stem to 30 m, leaves 15–20, petioles
long, leaflets two-ranked; fruit to 6 cm
long, 3.5 cm wide, orange-brown
Attalea exigua Drude
Catolé, indaia rasteira
Brazil, cerrado vegetation below
800 m
Subterranean stems; leaves 4–8, arched, Endosperm of seed used to make
leaflets irregularly arranged in clusters;
confections and as a sweetener
fruit 4–6 cm long, 3–5.5 cm wide, reddish
orange to dark purple
Attalea allenii H.E. Moore
Attalea amygdalina Kunth
Attalea maripa (Aubl.)
Mart.
Attalea victoriana Dug.
Maximiliana maripa
(Aubl.) Drude; many
others listed in
Henderson et al. (1995)
Cusi, anajá, inajai, guichire, inayo, Widespread in N South America
maripa, kukarit, inayuga, cucurito east of the Andes in primary and
secondary forest, open and
disturbed areas at low elevation
Stems 3.5–20 m tall, leaves 10–22 in five
distinct vertical rows on long petioles,
leaflets irregularly arranged in tight
clusters; fruit 4–6 cm long, 2.5–3 cm
wide, brown
Oil similar to coconut and African oil
palm extracted from seed
commercially
Seed edible
Attalea
91
92
Arecaceae
Attalea speciosa
babassu
Babassu, Attalea speciosa Mart. ex Spreng. (Arecaceae), is a
single-stemmed, pinnate-leafed palm native to the Amazon
region. The Brazilian common name babaçu or babassu is
from the Tupi-Guarani Indian language: ba = fruit; açu =
large); it is known as cusi in Bolivia (Anderson et al., 1991). It
is an important local resource in the southern Amazon,
especially in Brazil, yielding an oil comparable to coconut. Other
parts of the palm are also used by local people. There has been
some developmental interest in Brazil on expanding production
as well as increasing oil extraction efficiency.
World production and yield
It is estimated that several hundred thousand households
harvest the fruit of the babassu palm in the Brazilian state of
Maranhão alone (Balick and Pinheiro, 1993). The economic
value of the palm in that state was estimated at US$85 million
(May, 1986), but that included all uses to which this versatile
palm is put. However, most of the value resides in subsistence
economies, thus an exact figure is elusive. Oil extracted from
the seeds averages 90–150 kg/ha annually (it constitutes about
7% of the fresh fruit), and a minimum of 85,000 t of babassu
kernel oil were extracted annually during the 1970s in
Maranhão state (Pesce, 1985). During both World Wars, there
was significant export of babassu oil to Europe from Brazil, a
trade which all but disappeared by the 1960s (Anderson et al.,
1991). A shortage of coconut oil in the 1980s resurrected the
export industry briefly. Most production outside of
subsistence use feeds the Brazilian cosmetic industry, and
150,000 t of the oil was produced in 1985 (Balick and
Pinheiro, 1993).
Uses and nutritional composition
The seed (kernel) of babassu is rich in lauric acid (60–70%),
and is thus comparable to that of coconut (Cocos nucifera) or
African oil palm kernel (Elaeis guineensis). Over 80% of the oil
is saturated fat, about 11% monosaturated and the remainder
polyunsaturated. The oil contains 19 g of vitamin E/100 g. It
does not turn rancid as quickly as other palm oils. The seed
meal left over after oil extraction is often used as feed for
livestock. The hard fruit husks make an excellent charcoal
(Balick and Pineiro, 1993). The seeds are an important food
resource for the hyacinth macaw (Munn et al., 1988).
Botany
Babassu was long known as
Orbignya phalerata Mart. Other synonyms include Orbignya
martiana Barbosa Rodrigues, Orbignya barbosiana Burret and
Orbignya speciosa (Martius) Barbosa Rodrigues. Palm species
formerly recognized as the separate genera Maximiliana,
Orbignya and Scheelea have been combined with Attalea
(Henderson et al., 1995).
TAXONOMY AND NOMENCLATURE
The babassu is a single-stemmed palm,
reaching to 30 m in height and a girth of 20–50 cm. The
crown consists of 10–25 large, pinnate leaves that are at first
sub-erect but then become arching. The apical portion of the
leaf is often twisted. The petiole is short, 10–40 cm long,
DESCRIPTION
while the rachis extends from 5.5 to nearly 9 m in length,
supporting several hundred leaflets. Each pinna is 20–185 cm
long and 1–6 cm wide. The pendent inflorescences, arising
from the leaf axils, are either entirely male or bisexual, and are
up to 2 m long. In bud they are contained by a woody bract.
Staminate inflorescences are branched into as many as 400
flower-bearing rachillae, each with 15–100 flowers. Bisexual
inflorescences have slightly more branches, each with one or
two (sometimes three) pistillate flowers and one to several
staminate flowers that may not fully develop. The fruit is an
oblong drupe, 6–13 cm long and 4–10 cm wide. The outer and
middle layers are fibrous and mealy, respectively. A tough,
woody inner wall (endocarp) surrounds three to six (rarely
fewer, or even more rarely up to 11) ovoid seeds 3–6 cm long,
with oily white endosperm.
The babassu is
widely distributed along the southern edges of the Amazon
basin from the Atlantic Ocean to Bolivia, extending
throughout eastern and central Amazonas and northward to
Guyana and Surinam. Most populations are found south of
the Amazon River. There are areas in Maranhão and Piauí
states of Brazil where huge populations, as many as 10,000
palms/ha (Anderson et al., 1991) can be found. These socalled ‘babassu zones’, with high numbers of juvenile palms
(fruiting individuals are usually in the range of 100–200/ha),
may be in part artefacts of human activity, as babassu palm
colonizes disturbed sites very readily. These zones constitute
as much as 150,000 km2 in south-eastern Amazonas, often in
the transitional areas between forest and savannah. In primary
rainforest, mature reproductive palms are more scarce
(c.50/ha) because of light limitations. Babassu palms in the
forest may remain in the juvenile phase of growth for as much
as 50 years (Anderson, 1983).
Babassu has fairly broad ecological tolerance. Though the
babassu zones are usually on good soils with high annual
rainfall, A. speciosa also occupies the savannahs of the Brazilian
cerrado vegetation with as much as a half-year dry season
(though the larger populations are always along rivers). The
palm is not, however, tolerant of frequently inundated soils.
Annual average rainfall of 1500–2500 mm appears optimal,
and little or no tolerance of freezing temperatures is to be
expected. Wide tolerance of pH is reported (Duke, 2001).
ECOLOGY AND CLIMATIC REQUIREMENTS
Babassu has a consistent phenology
over a wide range. Leaf emergence and flowering occurs
during the local rainy season, followed approximately 9
months later by fruit ripening and leaf senescence and loss
(Anderson et al., 1991). In some stands, flowering occurs
throughout the year.
REPRODUCTIVE BIOLOGY
Babassu palms rarely begin to fruit
before 8 years under the best conditions. Fruit production
increases for the next dozen years, and trees bear for upwards
of 75 years.
FRUIT DEVELOPMENT
Horticulture
PROPAGATION Research on germination of babassu seed is
fairly extensive (Frazão and Pinheiro, 1985; Pinheiro, 1986;
Bactris
Pinheiro and Araujo Neto, 1987a, b). The babassu seed has a
type of germination known as ‘remote tubular’. It germinates
hypogeally (the cotyledon does not emerge from the seed).
The cotyledonary petiole emerges from the seed and grows
down to a depth of as much as 60 cm. The seedling stem and
root thus develop deeply underground while the rest of the
cotyledon (haustorium) absorbs the endosperm and enlarges
within the seed and occupies the space formerly filled by the
endosperm.
Fire and shade stimulate germination, and both conditions
are found in the forest–savannah transition zone where the
palm is often most common. An extensive adventitious root
system is formed early in the life of the palm. The apical
meristem remains underground for several to many years as
the stem expands in diameter before elongation commences.
The palm is thus able to re-grow successfully after injury.
Needless to say, such a system is difficult to adapt to nursery
production. Successful nursery seed germination of such
palms has been accomplished in very deep containers, or even
in long lengths of plastic pipe.
Babassu palms are most
productive on fertile soils, which suggests that the palms will
respond to fertilization. No information has been published on
mineral nutrition, however. The extensive root system of the
palms implies a fairly efficient mechanism of nutrient uptake.
NUTRITION AND FERTILIZATION
MANAGEMENT Most management strategies for babassu
involve enhancement of natural stands, rather than
establishment of plantations. The numerous seedlings and
stemless juveniles require thinning, which can only be
accomplished by harvest for palm heart (which kills the palm)
or with systemic herbicides. A combination of juvenile palms
and young reproductives are retained, and older palms as well
as those producing only male flowers are eliminated.
DISEASES, PESTS AND WEEDS Fruit are often parasitized by
the larvae of Pachymerus nucleorum, a bruchid beetle, after they
fall from the tree.
MAIN CULTIVARS AND BREEDING The wide ecological
tolerances of babassu suggest that seed-propagated lines with
specifically adapted features, as well as more productive
individuals, could be isolated, but no breeding or sustained
selection programmes have been initiated.
Alan W. Meerow
Literature cited and further reading
Anderson, A. (1983) The biology of Orbignya martiana (Palmae), a
tropical dry forest dominant in Brazil. PhD thesis, University of
Florida, Gainesville.
Anderson, A.B., May, P.H. and Balick, M.J. (1991) The Subsidy from
Nature – Palm Forests, Peasantry, and Development on an Amazon
Frontier. Columbia University Press, New York.
Balick, M.L. and Pinheiro, C.U.B. (1993) Babassu. In: Clay, J.W. and
Clement, C.R. (eds) Selected species and strategies to enhance income
generation from Amazonian forests. Food and Agriculture
Organization (FAO) Miscellaneous Working Paper 93/6, FAO,
Rome. Available at: http://www.fao.org/docrep/v0784e/v0784e0u.
htm#babassu (accessed 27 November 2005).
93
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Frazão, J.M.F. and Pinheiro, C.U.B. (1985) Métodos Para Acelerar e
Uniformizar a Germinação de Sementes de Palmeiras do Complexo
Babaçu (Palmee, Cocosoideae). Pesquisa em Andamento no. 38.
EMBRAPA-UEPAE, Teresina, Brazil, p. 2.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
May, P. (1986) A Modern Tragedy of the Non-Commons: AgroIndustrial Change Equity in Brazil’s Babaçu Palm Zone. Latin
American Programme Dissertation Series, Cornell University,
Ithaca, New York.
Munn, C., Thompson, J. and Yamashita, C. (1988) The hyacinth
macaw. In: Chandler, W.J., Labate, L.W. and Christopher, M.
(eds) Audubon Wildlife Report 1989–90. Elsevier Science and
Technology Books, San Diego, California, pp. 405–419.
Pesce, C. (1985) Oil Palms and Other Oil Seeds of the Amazon. Edited
and translated by D.V. Johnson. Reference Publications, Algonac,
Michigan.
Pinheiro, C.U.B. (1986) Germinação de Sementes de Palmeiras: Revisao
Bibliográfica. Documentos no. 5. EMBRAPA-UEPAE Teresina,
Brazil, 102 pp.
Pinheiro, C.U.B. and Araujo Neto, A. (1987a) Descrição do Processo
Germinativo de Sementes de Babaçu (Orbignya phalerata Martius).
Comunicado Técnico no. 14. EMAPA, Brazil, 7 pp.
Pinheiro, C.U.B. and Araujo Neto, A. (1987b) Teste Comparativo
Entre a Germinação de Frutos Inteiros e Amêndoas de Babaçu
(Orbignya phalerata Martius) em Vermiculita. Pesquisa em
Andamento no. 29. EMAPA, Brazil, 6 pp.
Bactris gasipaes
peach palm
The peach palm, Bactris gasipaes Kunth (Arecaceae), is known
as pupunha (Brazil), chontaduro (Columbia), pejibaye (Costa
Rica) or pijuayo (Peru) and is the only domesticated palm in
tropical America. It was probably first used for its wood, was
fully domesticated for its starchy-oily fruit and is now most
important for its heart-of-palm. The name in English and
other European countries is a misnomer, as the fruit is more
like a tree cassava (Manihot esculenta Crantz) than a juicy
peach (Prunus persica (L.) Batsch) (Clement et al., 2004).
During the last two decades of the 20th century, cultivated
peach palm supplanted wild harvested palms in most
producing countries (except Brazil, where Euterpe spp.
dominate) as the principal source of heart-of-palm in both
Latin American (the major production and consumption
region) and world markets (Mora Urpí and Gainza
Echeverria, 1999). While this is a modern success story, peach
palm is under-utilized for its originally important product –
the fruit.
The origin of peach palm has been debated extensively and
inconclusively for more than a century. Recent morphoanatomical evidence suggested that peach palm’s origin will
probably be found in south-western Amazonia (Ferreira,
1999), in what is now northern Bolivia, south-eastern Peru and
western Brazil. Recent allozyme evidence also pointed towards
that region (Rojas-Vargas et al., 1999), as does a DNA (RAPD)
analysis (Rodrigues et al., 2004). It now appears that two
94
Arecaceae
dispersions occurred from that region: one to the north-east,
resulting in the Pará microcarpa landrace and undescribed
intermediate populations; one to the north-west, resulting in
the complex of micro-, meso- and macrocarpa landraces and
undescribed populations that occupy central and western
Amazonia, the rest of north-western lowland South America,
and Central America up to Nicaragua (Fig. A.12). The full
number and distribution of peach palm landraces remain to be
determined.
Archaeological evidence on early distribution and possible
origin(s) and dispersion is still fragmentary (Morcote-Rios
and Bernal, 2001). The earliest records are of carbonized seeds
from the lowlands of Costa Rica, dated to 2250–1650 years
before present (BP) and 2190 ⫾ 60 BP, while the earliest
records in Amazonia are from Colombia, dated to 1080 ⫾ 40
BP. No records exist for the putative region of origin nor the
majority of peach palm’s Amazonian distribution. None the
less, archaeology will be important for confirming the
hypotheses based on modern plant morphology, anatomy and
genetics. A linguistic study of indigenous names of peach palm
is underway and will offer further clues.
During the century immediately following European
conquest, the fruit was reported to be used principally as a
cooked starchy staple, or fermented to make a drink, or
ground and dried into flour. The wood was important for tools
and weapons because of its straight grain and durability (the
modern Colombian name, chontaduro, means the ‘tough
palm’). Throughout western Amazonia and extending up to
Costa Rica the peach palm appears to have been a staple starch
crop, perhaps as important as maize (Zea mays L.) and cassava
in much of this region. The date palm (Phoenix dactylifera L.)
is the Old World’s dry tropical domesticate with similar
importance in subsistence, as was noted by the earliest
European conquerors who were familiar with it from southeastern Spain.
Peach palm’s pre-Columbian importance in Central
America was attested to by one of the first legal cases in the
Spanish colonies (1541–1546; Patiño, 1963: 121–122). In 1540,
a band of adventurers was authorized by the governor of
Panama to establish a settlement in the Sixaola River valley (in
what is now southern Costa Rica’s Atlantic coast), where they
reported that the local people depended on peach palm for
Fig. A.12. Approximate distribution of Bactris gasipaes var. gasipaes (light shading) in the lowland Neotropics, with the distribution of valid
(defined by molecular characterization and morphometric data) and still-to-be-validated landraces. Occidental (Central America and northwestern South America) landraces: 1. microcarpa Rama; 2. mesocarpa Utilis; 3. mesocarpa Cauca. Oriental (Amazonia) landraces: 4.
microcarpa Tembé; 5. microcarpa Juruá; 6. microcarpa Pará; 7. mesocarpa Pampa Hermosa; 8. mesocarpa Tigre; 9. mesocarpa Pastaza; 10.
mesocarpa Inirida; 11. macrocarpa Putumayo; 12. macrocarpa Vaupés (Source: Rodrigues et al., 2004).
Bactris
their subsistence. The governor of Nicaragua heard of this
settlement and, considering the Sixaola valley to be part of his
territory, sent an expedition to expel the Panamanian group.
This second expedition included a large group of Nicaraguan
natives, who relied upon maize for subsistence. When the
invaders did not find enough food in the Sixaola valley, they
started cutting peach palm for its heart-of-palm. This led to
expanded conflict, and between 30,000 and 50,000 palms were
cut in the valley to subjugate the local population and expel
the original band of adventurers. The Panamanian governor
prepared a detailed legal case against the Nicaraguan governor
about this conflict, which was sent to Madrid and provided an
enormous amount of detail about peach palm. The cut trees
would have furnished about 750 t of fresh fruit each year
(assuming 30,000 cut, with each yielding five or six bunches,
each with 50–70 fruit (details in the legal documents)), a
major subsistence contribution in a small river valley and a
strong argument for its role as a staple of the subsistence diet.
North-western Amazonia, the Colombian Pacific coast
(called the Chocó) and southern Central America (up to Costa
Rica) is where peach palm was most important (see distribution
of meso- and macrocarpa landraces in Fig. A.12). Although
maize and cassava were both present, peach palm seems to have
been a staple and was certainly the reason for harvest festivities.
The beginning of the harvest season was celebrated by
preparation of large amounts of cooked fruit, which was
consumed both directly and after being transformed into a
fermented drink (chicha). Depending upon the number of days
of fermentation, the chicha would have been similar in potency
to a low alcohol beer (3–5%) or as strong as wine (10–12%).
These harvest festivities were followed, 9 months later, by a
peak in births, probably due to the better nutritional status of
the mothers during the harvest season (Patiño, 1992) and may
be how peach palm earned its reputation as an aphrodisiac as
well. The 500 years since the European conquest has seen a
significant decline in peach palm’s importance, to the point
that it is currently an under-utilized minor crop.
95
120,000 t, of which only about 50% is commercialized as fresh
fruit, while the other 50% is used for subsistence, either
directly or as animal feed, or is wasted (Clement et al., 2004).
A typical fresh bunch weighs 2–5 kg, and is worth
US$0.50–1.00 at the farm gate and US$1.00–3.00 in the
market, so Neotropical farmers earn about US$11 million/
year from the commercialized fruit, while consumers pay
about US$30 million/year to enjoy it.
In contrast, the heart-of-palm is grown in high-density (>
5000 plants/ha), high input commercial monocultures and
some production statistics exist. Brazil had approximately
20,000 ha in production in 2002 and another 3000–5000 ha
recently planted. Bolivia had 3000 ha in the Cochabamba
region and Colombia had 1000 ha in its Chocó region. Costa
Rica had 8000 ha in production in 2002, down from 12,500 in
1998 due to severe competition from Ecuador, which had
10,000 ha in production in 2002. Panama, Peru and Venezuela
have minor production areas. Extrapolation from this data set
suggests that the total Neotropical production area is greater
than 43,000 ha. Assuming standard density (5000 plants/ha),
this area yields 322.5 million hearts-of-palm/year, each
weighing about 200 g, worth about US$0.25 per heart at the
farm gate, so farmers earn about US$80 million. An additional
200 g of edible stem tissue is also processed from each stem,
but farmers don’t get paid for this. The majority of consumers
buy 300 g net weight bottles or 500 g net weight cans of
processed heart-of-palm, paying US$2–5 per bottle in Brazil
(variation due to distances between production area and
market, and perceived quality of brand name) and more in the
USA and the European Union. Bottles are standard in Brazil,
as consumers feel more confident of quality when they can see
the hearts. The state of São Paulo, Brazil, is the largest world
consumer of heart-of-palm, followed by the rest of Brazil, the
European Union, the USA and other Latin American
countries. Ecuador and Costa Rica are the major exporting
countries of peach palm hearts; Brazil exports less than 10%
of its total heart-of-palm production, which includes many
more hearts of Euterpe oleracea Mart. than of peach palm.
World production and yield
Peach palm is still almost exclusively a Neotropical crop, with
only experimental areas in Africa, Asia and Oceania. As a fruit
crop, it is grown almost exclusively by smallholders in home
gardens and swiddens, with a few small orchards near major
consumption areas. Hence, all production data are estimates.
The estimated production of the State of Amazonas, Brazil, in
2000 was 13,600 t of fresh fruit bunches; Brazilian Amazonia
probably produced at least twice that amount. Colombian
production was estimated at 49,000 t in 2002, divided between
its Amazonian and Chocó (Pacific) lowlands. Costa Rican
production was estimated at 10,500 t in 2002, divided between
its Atlantic and humid Pacific lowlands, and there are probably
more orchards there than in other countries (Clement et al.,
2004). Bolivia, Ecuador, Peru and Venezuela also produce
moderate amounts of peach palm in their Amazonian
lowlands, while Venezuela also produces in its Orinoco river
basin lowlands and Ecuador in its Pacific coastal lowlands.
French Guiana, Guyana, Nicaragua, Panama and Surinam are
minor producers. Extrapolation from this scanty data set
suggests that total Neotropical production is probably about
Uses and nutritional composition
Today the fruit is used principally as a cooked snack, although
the major-producing countries all have cookbooks with varied
recipes. There is a niche market demand for dry flour, but the
benefit/cost ratio of supplying this demand is generally
negative, principally because of supermarket stocking strategies,
and Latin American research and development institutes have
not devoted enough attention to changing this ratio (Clement et
al., 2004). As fruit drinks gain continually greater market share
among health-conscious consumers, chicha could make a comeback. The name peach palm was coined by Alexander van
Humboldt and is derived from the aroma of the fermented
pulp, which recalls that of fresh tree-ripened peaches. The
flavour is distinctly fruity (though not peachy), rather than
starchy, and is very pleasant. More food technology research
and development is needed to exploit this opportunity,
especially in countries that already know peach palm’s flavour.
The fruit is energy rich, due both to starches and to oils,
while the heart-of-palm is essentially a dietary product (Table
A.48). The relative proportions of starch to oil vary inversely
96
Arecaceae
Table A.48. Mean chemical composition per 100 g of peach palm fruit
mesocarpa and of heart-of-palmb.
Fruit mesocarp
Heart-of-palm
g
g
273.5
3.3
6.0
2.2
3.3
0.5
34.9
2.0
0.8
47.6
1.5
1.3
0.73
0.35
0.22
5.2
0.9
nac
mg
mg
18.9
0.59
17.1
na
240.5
4.3
42.4
0.23
3.4
na
193.6
0.1
Vitamins
mg
mg
Vitamin A (carotene)
Vitamin C
Thiamine (vitamin B1)
Riboflavin (vitamin B2)
Niacin
1.1
18.7
0.045
0.135
0.81
Proximate
Energy (kcal)
Protein
Fat
Saturated
Monounsaturated
Polyunsaturated
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
part of the reason that peach palm offers an opportunity for
enhanced food security in humid tropical areas, an
opportunity that is unexploited in Africa and Asia.
Botany
Henderson (2000) revised
Bactris and provided a testable hypothesis about the
relationships among taxa closely related to peach palm.
Seventeen different species in three genera had been
considered close peach palm relatives since Ruiz and Pavon
described the first wild population in 1798 as Martinezia
ciliata; they are now all synonyms. Taxonomic controversy of
this type is common in species with domesticated populations
because morphological variation is enhanced during the
domestication process (Clement, 1999). In the new hypothesis,
all cultivated populations and landraces are now Bactris
gasipaes Kunth var. gasipaes and all wild populations are now
B. gasipaes var. chichagui (H. Karsten) Henderson.
A complex hierarchy of landraces was proposed on
morphometric grounds. As understood here, each landrace is a
meta-population composed of a variable number of closely
related domesticated and always cultivated subpopulations,
defined by a specific combination of morphological characteristics, a distinct geographic distribution and associated
ethnohistory. The original landrace hierarchy proposal had a
primary geographic division defined by the Andes, with an
Oriental group in lowland northern South America and an
Occidental group in lowland north-western South America
northwards into Central America (Fig. A.12). Within these
groups, fruit size is the primary factor to distinguish landraces
into microcarpa (10–30 g), mesocarpa (30–70 g) and macrocarpa
(> 70 g), since this trait was most modified by human selection.
In the Oriental group, three microcarpa, five mesocarpa and two
macrocarpa landraces had originally been mapped, while in the
Occidental group, one microcarpa and four mesocarpa landraces
had been mapped. The new genetic analysis (Rodrigues et al.,
2004) reduced this to three microcarpa, four mesocarpa and two
macrocarpa in the Oriental group, and one microcarpa and two
mesocarpa in the Occidental group (Fig. A.12). This new
analysis also suggests that the Andes was not a barrier to
dispersion (as originally proposed by Prance, 1984) and the
Oriental/ Occidental distinction may not be important.
TAXONOMY AND NOMENCLATURE
na
3.2
na
na
na
a
Mesocarp of three fruit.
A section 9 cm long, 2 cm in diameter.
c na = not available.
b
along the domestication continuum, with wild-type fruit being
rich in oils while the domesticates are rich in starches (Table
A.49). Fruit protein quality is not exceptionally high, but fruit
mesocarp oil is rich in monounsaturated oleic acid (Yuyama et
al., 2003). The fruit contain two antinutritional factors, a
trypsin inhibitor and calcium oxalate crystals, which are
denatured or eliminated by boiling, respectively. The heart-ofpalm contains calcium oxalate crystals. The fruit chemical
composition is similar to that of maize and better than that of
potato or cassava on a fresh weight basis (Table A.49), which is
Table A.49. Comparison of mean chemical composition (g/100 g)a of peach palm (Amazonian mean and three landraces), cassava, maize, sweet potato and a
set of 21 succulent Amazonian fruits.
Crop
palmb
Peach
Juruá landracec
Solimões landracec
Putumayo landracec
Cassavad
Maize (fresh)d
Sweet potatod
Succulent fruitsc
a
Water
Protein
Oil
Carbohydrate
Fibre
Energy (MJ)
45.0
54.4
42.7
52.6
65.2
63.5
67.2
82.8
3.5
3.1
4.1
1.9
1.0
4.1
0.9
0.9
27.0
13.8
12.0
3.5
0.4
1.3
0.2
0.8
19.8
19.6
31.2
38.0
32.8
30.3
29.6
11.9
3.8
8.4
9.3
3.2
1.0
1.0
1.1
2.9
1.47
1.04
1.20
0.85
0.55
0.54
0.53
0.26
Fresh weights; the difference between the sum of the means and 100 is due to ash content.
Mora Urpí et al. (1997).
c Clement et al. (2004) and references therein.
d Leung and Flores (1961).
b
Bactris
The peach palm is a multi-stemmed palm that
may attain 20 m in height (Plate 13). Stem diameter varies
from 15 to 30 cm and internode length from 2 to 30 cm, and
becomes reduced with age after 5 years. The internodes are
armed with numerous black, brittle spines, although spineless
mutants occur and have been selected for in several areas. The
stem is topped by a crown of 15–25 pinnate fronds, with the
leaflets inserted at different angles. The heart-of-palm is a
gourmet vegetable composed of the tender unexpanded leaves
in the palm’s crown. The inflorescences appear among the
axils of the senescent fronds. After pollination, the bunch
contains between 50 and 1000 fruit (there is a strong negative
correlation between fruit size and fruit number in a bunch)
and weighs 1–25 kg. Numerous factors cause premature fruit
drop: poor pollination, poor plant nutrition, drought,
crowding, insects and diseases. The fruit is a drupe with an
humid starchy/oily mesocarp, a fibrous red, orange or yellow
exocarp, and a single endocarp, with a fibrous/oily white
kernel. Individual fruit of var. gasipaes weigh between 10 and
250 g, with means varying according to landrace, and seeds
weigh between 1 and 4 g; fruit of var. chichagui weigh between
0.5 and 5 g and seeds weigh between 0.3 and 1 g (Plate 14).
DESCRIPTION
Wild peach palm
(var. chichagui) occurs in transitional natural ecosystems and
where natural disturbances are frequent, principally along
riverbeds and in primary forest gaps, while cultivated peach
palm (var. gasipaes) only occurs in ecosystems created by
humans. Extensive natural stands of wild peach palm have not
been reported (see population structure above), nor are wild
palms harvested today.
Cultivated peach palm is adapted to a wide range of
ecological conditions, reflecting its wide anthropogenic
geographical distribution in the humid Neotropics. It is most
productive on relatively deep, fertile, well-drained soils at low
to middle altitudes (< 1000 m above sea level), with abundant
and well-distributed rainfall (2000–5000 mm/year) and
average temperatures above 24°C. It produces relatively well
on low-fertility soils, including highly eroded laterites with
50% aluminium-saturated acid soils, following the slashingand-burning of primary or secondary forest, since the burn
releases calcium and magnesium that neutralize the acidity
and aluminium toxicity, but fruit production decreases in the
long term without additional lime and nutrient inputs. It does
not tolerate waterlogged soils. It can withstand relatively short
dry seasons (3–4 months) if soils are not excessively sandy, but
dry seasons significantly reduce growth and yield. Symbiotic
associations with vesicular-arbuscular mycorrhizae improve
growth and are often essential for normal development.
The peach palm is often reputed to be rustic and well
adapted to tropical soils with few nutrients (NAS, 1975). In
fact, like any domesticated crop, it requires good husbandry.
These contradictory observations are due to peach palm’s
domestication in traditional agroecosystems in the American
tropics. Wild peach palm produces abundant fruit with no
additional inputs on low fertility soils, but these fruit are very
small, so total yield is low and export beyond the system is
minimal. During the domestication process, peach palm
became well adapted to scavenging nutrients after the burning
of previous vegetation. In these agroecosystems, the palms are
ECOLOGY AND CLIMATIC REQUIREMENTS
97
generally transplanted, arriving with a partially developed root
system already inoculated with mycorrhizae. This initial
advantage is complemented by its fibrous root system and its
strong competitive ability to absorb nutrients (Fernandes and
Sanford, 1995). The result is fast initial growth, to get the
crown above competing cassava and low stature fruit trees. In
traditional agroecosystems, weedy regrowth is managed at
progressively lower rates until a managed fallow develops, with
nutrient recycling similar to natural forests. In the initial
stages of this sequence, peach palm does well, but is eventually
shaded out by taller fallow trees. This sequence of events
apparently supports the contention that peach palm is both
rustic and well adapted to low nutrient soils, but in fact it is
well adapted to compete for nutrients and light when these are
available in traditional systems, and dies out as these become
less available. In modern agroecosystems, peach palm requires
appropriate fertilization to remain productive.
Phenology varies both within and
among countries, and the environmental events that trigger
flowering are not yet clear. In central Brazilian Amazonia, the
main flowering season extends from the mid-dry season
(August–September) to the beginning of the rainy season
(November), and fruit ripen between late December and late
March. In Costa Rica’s Atlantic zone, the main flowering
season extends from May to July and the fruiting season from
August to October; this region has a much less pronounced
dry season than central Amazonia. When the dry season is less
pronounced, a second flowering period may occur in plants
with good nutritional status. When this happens, the harvest
seasons are separated by about 6 months. Costa Rican farmers
have taken advantage of micro-climatic, soil and altitudinal
variability to supply their major San José market year-round,
effectively exploiting phenological variability (Clement et al.,
2004).
Peach palm is predominantly allogamous, having separate
pistillate and staminate flowers, and protogynous development
(i.e. the female flowers are receptive before the male flowers
shed pollen). Self-pollination is thought to be regulated by a
genetic incompatibility mechanism, but there is considerable
variation in self-fertility. Self-pollination may occur: (i) within
the same inflorescence; (ii) between inflorescences of the same
stem; or (iii) between inflorescences on different stems of the
same plant. The latter event is probably much more common
than the other two events, although the first event may be
common at the beginning and end of the season when
sufficient cross-pollen is not available.
The pollination cycle lasts 3 days in a given inflorescence.
Female anthesis begins when the inflorescence bract opens in
the afternoon, and unfertilized female flowers may remain
receptive for up to 48 h. Late in the afternoon of the second
day, female flower anthesis normally ends and male flower
anthesis begins (protogyny). Male flowers release their pollen
in 15–30 minutes, showering the inflorescence and visiting
insects, and then the male flowers abscise. The insects then
leave and search for a recently opened inflorescence, attracted
by a musky scent produced by a gland on the male flowers.
The reproductive biology of peach palm suggests a tight coevolutionary history with very small curculionid beetles,
thousands of which are attracted by the musk to a single
REPRODUCTIVE BIOLOGY
98
Arecaceae
inflorescence. Where peach palm has been introduced recently,
the lack of these curculionid beetles may severely limit fruit
set.
Wild peach palm (var. chichagui) occurs in small subpopulations (three to 20 plants), normally quite distant from
the next such sub-population (500 m to several kilometres).
This meta-population structure favours genetic drift and selfpollination within sub-populations, with consequent
reduction in genetic diversity and random shifts in allele
frequencies, making the genetic incompatibility mechanism
very important as a way of counteracting this. However, to
assure population survival, the mechanism must also allow
some self-fertility. Population structure of cultivated peach
palm (var. gasipaes) is remarkably similar to that of wild peach
palm, as traditional farmers tend to have three to ten plants in
their home gardens and five to 30 in their agroforestry plots
(Clement et al., 2004), each of which is a distinct subpopulation. In these agroecosystems, human selection
pressure tends to reinforce self-pollination, as fewer moreclosely related plants are used to create each new subpopulation, resulting in a further reduction of genetic
diversity, as observed by Rodrigues et al. (2004). This
reduction is typical of domesticated species, but a human
behavioural characteristic – seed exchange – acts to enhance
genetic diversity and maintain genetic viability. Neighbours
exchange seed quite frequently within villages, and seeds are
often taken as gifts when visiting other villages or are
requested as gifts when a special tree is observed (Adin et al.,
2004). Seed exchange and buying seed in the market are
important mechanisms for maintaining genetic diversity in any
crop, but are especially important in under-utilized crops that
are more likely to be threatened by or suffering genetic
erosion.
GROWTH AND DEVELOPMENT Fruit develop to
maturity in 3–4 months. Inflorescences develop in sequence
on the stem, so one cannot harvest all bunches on the stem at
the same time. The fruiting season typically extends over a
2–4 month period (rarely more in exceptionally good years).
The first harvest of the season normally yields the largest and
best-quality fruit, after which fruit quality gradually
deteriorates due to increasing insect damage and fungal
infection, favoured both by the presence of fruit and by the
gradual depletion of physiological reserves.
During the first 2 months of growth, the fruit expands in
size and its composition is rich in starches and some protein.
During the third month, starches are metabolized to produce
oils and carotenes, and the mesocarp and exocarp gain colour.
When the exocarp has 50% of its final colour, the seed is
mature and the fruit can be harvested. Full flavour and colour
come a week or so later, and the full flavour may be too strong
for some consumers. Hence, fruit tend to be harvested and
commercialized before full ripeness. After harvest, fruit
deteriorate rapidly (3–7 days), so must be handled carefully
and expeditiously.
FRUIT
Horticulture
PROPAGATION The peach palm is propagated by seed, as
vegetative propagation of off-shoots is difficult and a
commercial tissue culture protocol has not yet been developed.
The seed is considered to be recalcitrant (i.e. it can not be
dried or frozen for storage). Seeds obtained from healthy
productive trees at the beginning of the harvest season have
greatest germination success, better than 80% with standard
practices. These practices include careful pulp removal and
seed cleaning, sowing in appropriate substrates with sufficient
(but not excessive) irrigation tailored to the substrate,
moderate shade (75% of full sun) and frequent inspection for
pests and diseases. Fresh seeds take 30–90 days to germinate,
depending upon temperature and humidity, and a field-ready
seedling takes another 4–6 months to produce in an organicmatter rich, well-fertilized substrate.
FIELD MANAGEMENT Planting density, plant management
and fertilization vary significantly between the fruit crop and
the heart-of-palm crop. The fruit crop is generally grown in
agroforestry systems of varying diversity and less frequently in
monoculture orchards. In the latter, spacing must be at least 5
⫻ 5 m (400 plants/ha) or different rectangular or triangular
arrangements to allow sufficient light into the orchard; wider
spacings allow more intercropping during the early years. Offshoot management is critical to reduce within-clump
competition for light and nutrients, while maintaining enough
off-shoots to replace the fruiting stem when this grows too tall
to harvest economically (above 10 m, which is often attained in
10 years or less).
Dolomitic lime is recommended for managing soil acidity in
tropical America’s typically acid Oxisols and Ultisols,
especially when aluminium concentration is high. During field
preparation 2 t/ha is recommended and 1 t/ha at 3-year
intervals for maintenance (Bovi, 1998) to provide sufficient
magnesium and calcium, while making phosphorus more
available. Juvenile plants require abundant nitrogen (N) and
moderate phosphorus (P) (90–120 kg/ha N; 45 kg/ha P;
80–90 kg/ha K), while fruiting plants require abundant
potassium (K) and nitrogen, with moderate phosphorus
(140–190 kg/ha N; 90 kg/ha P; 150–180 kg/ha K). In most
tropical soils micronutrient deficiencies are common, but
poorly studied with peach palm. Hence, animal manure is
strongly recommended and mineral fertilization can be
reduced proportionately. Leguminous groundcover crops are
also highly recommended, as they suppress weeds and provide
organic matter and nitrogen to the system.
The heart-of-palm crop is grown in high density (> 5000
plants/ha), high input stands (standard spacing is 1 ⫻ 2 m, but
numerous alternatives exist, especially when small tractors are
used for maintenance). After field planting, the first harvest is
obtained within 18–24 months and the clumps are maintained
by periodic harvesting of the larger off-shoots – the clumps are
essentially immortal if adequately fertilized and weeded. Most
researchers and growers consider off-shoot management
important, but the Pampa Hermosa landrace germplasm that is
used in most plantations produces fewer off-shoots than Utilis
landrace germplasm, for example, and thus requires less
management on average. Dolomitic lime should be applied as
above. The perennially juvenile plants require abundant nitrogen
(200–250 kg/ha N) and moderate levels of other macronutrients
(20–40 kg/ha P; 100–120 kg/ha K). Harvesting leaves 90% of
the biomass to mulch the field and many agroindustries compost
Bactris
the other 9% for use in the nursery or return it to the field, all of
which makes the crop very sustainable. Both organic and
conventional (mineral fertilizer, but almost no pesticides) heartof-palm are available, although little research has been published
on the organic alternative to date.
Erwinia bacteria can be a problem
where drainage is poor or shade intense in heart-of-palm
stands. Anthracnose (Colletotrichum) fungal attack indicates
inadequate
phosphorus
fertilization.
Poor
nursery
management allows damping-off (Fusarium) and other fungal
diseases to become important. Poor plant nutrition leads to
increasing fungal attacks on fruit during the season, but these
tend not to reach critical levels. Hence, as in other minor
crops, most peach palm diseases can be managed with
appropriate fertilization and field practices, and no pesticides
are normally used.
Beetles (Coleoptera) may be locally important fruit or seed
pests, but only in the Colombian Pacific have they seriously
affected fruit yields. A seed-boring beetle has occasionally
been reported in the south-west Amazon. Foliage mites
(Retracus spp.) are indicative of poor plant nutrition and
occasionally require chemical control on fruiting plants, but
not in heart-of-palm orchards, where rapid growth and harvest
keep their populations under control. Gophers (ground
squirrels), pigs, rabbits and rats are pests in some areas, and all
can be controlled with fencing or poison baits.
MAIN DISEASES AND PESTS
POSTHARVEST HANDLING AND STORAGE Fruit from spineless
peach palm is typically collected by climbing the stem and
lowering the fruit bunches to the ground with a rope or
dropping them into a net. Most peach palms have spiny stems,
however, and these are very difficult to climb. Hence, farmers
use poles with a hook or curved knife at the end, with which
they dislodge or cut the fruit bunch and catch it with a net or
foam cushion. Harvesting from the ground is faster and safer
than climbing the stem to collect fruit, but it causes more
damage to the fruit. Both methods are expensive in man-hours
and help explain why many bunches don’t get to market
during the peak harvest season, when bunch prices are
depressed by excess supply. In Brazilian Amazonia, a 10 m tall
tree requires three men (or adolescent boys) to harvest a
bunch: one man to manage the pole and dislodge the bunch;
two men to manage the net or cushion. At twice the national
minimum wage (the most common agricultural labour rate in
the region – the minimum is R$240 = US$81/month, January
2004), this can cost US$0.45, equivalent to 90% of the farm
gate price of a fresh bunch (Clement, 2000). Needless to say,
height growth is a major problem in growing peach palm for
fruit and the reason that tall stems are cut to make room for
younger off-shoots.
Fresh fruit are very perishable, due to its chemical
composition (Table A.48). A ripe fresh bunch can be
maintained in good condition without refrigeration for only
4–6 days if kept in the shade and well ventilated. With
refrigeration (20°C, 70% relative humidity) and waxing,
storage can be extended to 8 days, but few merchants have the
necessary capital and facilities. Shelf life of fresh fruit can be
extended by collecting well-developed fruit that are just
starting to change from green to their final colour, and this is
99
now standard practice throughout the region. Frozen, dried or
canned fruit can be conserved for months, but with the
consequent loss of final flavour and colour.
In practice, no commercial ventures have survived by
industrially processing the fruit. Only Costa Rica had
successfully developed the technology, and during the 1980s
several small processing and commercialization businesses had
fair success. During the late 1980s and 1990s, farmers
discovered the advantages of planting where phenology
offered them a market window (see ‘Fruit growth and
development’ above) and fresh fruit are now available yearround. This drove the processing businesses to ruin, since
their product was perceived to be of inferior quality and was
more expensive (Clement et al., 2004).
Fresh fruit are commonly sold by the bunch, or they are
minimally processed and packaged, especially in Costa Rica.
The processing involves only removal of fruit from the bunch,
washing, waxing (by the more capitalized communities),
sorting and classifying, and packaging in net bags of specified
weight. Cooked fruit is handled like fresh fruit. In all other
countries, fruit are sold at market in bunches or as cooked
fruit.
If fruit are destined for flour, they should be processed on
the day of harvest or the following day. Processing involves:
cooking the entire bunch to facilitate removal of the fruit,
denature potential toxins and improve starch quality; cutting
the whole fruit into small pieces; removing the seeds and
drying the pulp and peel (red peels give a golden colour to the
flour); and then grinding and packaging. Processing fruit of
low phytosanitary quality is more difficult and expensive,
requiring careful sorting and peeling. While a niche market
demand exists, modern supermarket logistics have made
commercialization too expensive for small processing
businesses, and none have survived the changes in
supermarket supply strategies (Clement et al., 2004).
Hearts-of-palm are an entirely different story. Off-shoots
are harvested when they reach commercial dimensions, which
depend on factory and market demands for heart-of-palm. For
the international market there is essentially one basic quality:
true or export heart-of-palm is a cylinder composed of a
tender petiole-sheath enveloping the developing leaves above
the apical meristem. This also explains the name, as the apical
meristem is truly the heart of the palm, as all leaves and stem
arise there. The true heart commands a higher price when it
has a narrow diameter, and is called ‘extra fine’. In Brazil,
there is a demand for three heart-of-palm dimensions: thin
(1.5–2.5 cm) hearts to be canned for the export markets;
medium (2–4 cm) hearts for both the bottled and the fresh
markets in Brazil; and thick (3–6 cm) hearts for the Brazilian
churrascaria market (restaurants that specialize in barbecued
meat with thick hearts-of-palm as garnish), either bottled or
fresh.
For all national markets (and a few minor international
markets) there is a second quality: the tender stem tissue
below the apical meristem. When fresh or processed, this stem
tissue has the same flavour as the heart, but a different texture,
as the stem is fibrous parenchymous tissue and the heart is leaf
sheath, petiole and blade tissue. The stem tissue commands a
much lower price, and is presented in different sizes and
shapes, but provides the industry with larger earnings because
100
Arecaceae
it comes as free material with the stems they buy from the
farmers. All industries also offer chunks and slices of true
heart or tender stem that doesn’t meet their other standards.
These are sold very cheaply to restaurants and pastry shops
that don’t demand higher quality materials.
For hearts-of-palm with 2–3 cm diameter, off-shoots are
harvested when they attain diameters of larger than 9 cm,
measured at 20–30 cm above the ground. Off-shoot diameter
and other morphological characteristics are correlated with
heart-of-palm yield, but in practice only diameter is measured
to determine if the off-shoot is ready for harvest because there
is a good correlation between yield and diameter under normal
nutritional conditions. In Brazil, the slightly larger-diameter
hearts are more easily evaluated by height than diameter
(Clement and Bovi, 2000), and this has become standard
practice for both the processed and the fresh markets.
When off-shoots are ready for harvest, they are cut and the
outer fibrous leaf sheaths are removed. Two non-commercial
leaf sheaths, surrounding the heart-of-palm, are normally left
to protect it from rapid moisture loss and mechanical damage
during transport. Ideally the heart-of-palm should be
transported to the processing plant on the day of harvest to
minimize moisture loss. If transport delays are anticipated,
more leaf sheaths should be left surrounding the heart-ofpalm, a paraffin/beeswax mixture should be applied to the cut
ends, and they should be stored in a shady place. These
postharvest treatments will normally conserve fresh heart-ofpalm for 4–6 days without significant moisture loss or fungal
infection.
All processing is done following Codex Alimentarius
regulations (FAO/WHO, 1985). The Codex is designed to
guarantee safe food for consumers. Unfortunately, flavour and
appearance are lost en route. This helps explain why the world
market for this product is essentially saturated and why Costa
Rica lost market share to Ecuador over the last decade – in a
saturated market, price determines who gets the sales. In
Brazil, Costa Rica and Hawaii (USA) a new market for fresh
heart-of-palm (and tender stem) is being developed by
numerous small entrepreneurs. This new market is still only a
niche, but appears to have great promise.
MAIN CULTIVARS AND BREEDING No named cultivars have
been brought to market. Rather, the breeding effort aims at
general population improvement to maintain the variability
that helps control pests and diseases. This strategy is perfect
for heart-of-palm production, which is now based principally
on the Pampa Hermosa (Yurimaguas, Peru) landrace because
of its spinelessness, rapid growth and good quality (Mora
Urpí et al., 1999). This strategy is not good for the fruit
market, since consumers desire uniformly high quality fruit,
which are not abundantly found in open-pollinated landrace
populations. A new selection strategy is being tried in Peru to
remedy this and will certainly be adopted in the rest of Latin
America.
Basic production parameters in one region may serve as a
reference for comparing production in other regions, and offer
targets for agronomic and genetic improvement programmes.
Production parameters (Mora-Urpí et al., 1999; based on 5000
plants/ha) for heart-of-palm in Costa Rica are:
● time from plantation establishment to first harvest of all
plants (9 cm off-shoot diameter) should be 18 months;
● number of harvested off-shoots should 8000/ha in the first
year of production (12–24 months) and 10,000/ha each
year thereafter;
● field-harvested shoots should contain 70% export quality
hearts and 30% tender stem quality;
● average yield of export-quality heart-of-palm after
processing should be 1.35 t/ha, beginning in the second
year of production (10,000 harvested off-shoots/ha, each
yielding 135 g of export quality and 50 g of stem quality).
In Brazil, with somewhat different internal market demands,
these production parameters vary a little, with slightly longer
time frames given that plants must grow for another few months
to attain Brazilian size specifications. Another difference is that
the longer time frame allows a larger tender stem section, giving
better returns to the processing operations. Charles R. Clement
Literature cited and further reading
Adin, A., Weber, J.C., Sotelo Montes, C., Vidaurre, H., Vosman, B.
and Smulders, M.J.M. (2004) Genetic differentiation and trade
among populations of peach palm (Bactris gasipaes Kunth) in the
Peruvian Amazon – implications for genetic resource
management. Theoretical and Applied Genetics 108, 1564–1573.
Bovi, M.L.A. (1998) Palmito Pupunha: Informações Básicas Para
Cultivo. Boletim Técnico 173, Instituto Agronômico, Campinas,
São Paulo, Brazil, 50 pp.
Clement, C.R. (1999) 1492 and the loss of Amazonian crop genetic
resources. I. The relation between domestication and human
population decline. Economic Botany 53, 188–202.
Clement, C.R. (2000) Pupunha (Bactris gasipaes Kunth, Palmae). Série
Frutas Nativas, 8, Fundep, Jaboticabal, São Paulo, Brazil, 48 pp.
Clement, C.R. and Bovi, M.L.A. (2000) Padronização de medidas de
crescimento e produção em experimentos com pupunheira para
palmito. Acta Amazonica 30, 349–362.
Clement, C.R., Weber, J.C., van Leeuwen, J., Domian, C.A., Cole,
D.M., Arévalo Lopez, L.A. and Argüello, H. (2004) Why
extensive research and development did not promote use of peach
palm fruit in Latin America. Agroforestry Systems 61, 195–206.
Fernandes, D.N. and Sanford, R.L. (1995) Effects of recent land use
practices on soil nutrients and succession under tropical wet
forest in Costa Rica. Conservation Biology 9, 915–922.
Ferreira, E. (1999) The phylogeny of pupunha (Bactris gasipaes
Kunth, Palmae) and allied species. In: Henderson, A. and
Borchsenius, F. (eds) Evolution, Variation and Classification of
Palms. Memoirs of the New York Botanical Garden 83. New York
Botanical Garden Press, Bronx, New York, pp. 225–236.
Food and Agriculture Organization/World Health Organization
(FAO/WHO) (1985) Codex Standard for Canned Palmito (Codex
Stan 144–1985). FAO/WHO, Rome.
Henderson, A. (2000) Bactris (Palmae). Flora Neotropica
Monograph 79, New York Botanical Garden, New York, 181 pp.
Leung, W.T.W. and Flores, M. (1961) Food Composition Table for Use
in Latin America. Guatemala City, Instituto de Nutrición de
Centro América y Panamá and Bethesda National Institute of
Health, Bethesda, Maryland.
Mora Urpí, J. and Gainza Echeverria, J. (eds) (1999) Palmito de
Pejibaye (Bactris gasipaes Kunth): Su Cultivo e Industrializacíon.
Editorial Universidad de Costa Rica, San José, Costa Rica, 260 pp.
Bactris
Mora Urpí, J., Weber, J.C. and Clement, C.R. (1997) Peach palm.
Bactris gasipaes Kunth. Promoting the Conservation and Use of
Underutilized and Neglected Crops 20. Institute of Plant Genetics
and Crop Plant Research (IPK), Gatersleben and International
Plant Genetic Resources Institute (IPGRI), Rome, 83 pp.
Mora Urpí, J., Bogantes Arias, A. and Arroyo Oquendo, C. (1999)
Cultivares de pejibaye para palmito. In: Mora Urpí, J. and Gainza
Echeverria, J. (eds) Palmito de Pejibaye (Bactris gasipaes Kunth):
Su Cultivo e Industrializacíon. Editorial Universidad de Costa
Rica, San José, Costa Rica, pp. 41–47.
Morcote-Rios, G. and Bernal, R. (2001) Remains of palms (Palmae)
at archaeological sites in the New World: a review. Botanical
Review 67, 309–350.
National Academy of Sciences (NAS) (1975) Underexploited Tropical
Plants with Promising Economic Value. NAS, Washington, DC,
188 pp.
Patiño, V.M. (1963) Plantas Cultivadas y Animales Domesticos en
America Equinoccial. Tomo 1. Frutales. Imprenta Departamental,
Cali, Colombia, 547 pp.
Patiño, V.M. (1992) An ethnobotanical sketch of the palm Bactris
(Guilielma) gasipaes. Principes 36, 143–147.
Prance, G.T. (1984) The pejibaye, Guilielma gasipaes (H.B.K.) Bailey,
and the papaya, Carica papaya L. In: Stone, D. (ed.) PreColumbian Plant Migration. Peabody Museum of Archeology and
Ethnology, Paper No. 76. Harvard University Press, Cambridge,
Massachusetts, pp. 85–104.
Rodrigues, D.P., Astolfi Filho, S. and Clement, C.R. (2004)
Molecular marker-mediated validation of morphologically defined
landraces of pejibaye (Bactris gasipaes) and their phylogenetic
relationships. Genetic Resources and Crop Evolution 51, 871–882.
Rojas-Vargas, S., Ramírez, P. and Mora Urpí, J. (1999) Polimorfismo
isoenzimático en cuatro razas y un híbrido de Bactris gasipaes
(Palmae). Revista de Biologia Tropical 47, 755–761.
Yuyama, L.K.O., Aguiar, J.P.L., Yuyama, K., Macedo, S.H.M.,
Fávaro, D.I.T., Afonso, C. and Vasconcellos, M.B.A. (1999)
Determinação de elementos essenciais e não essenciais em palmito
de pupunheira. Horticultura Brasileira 17, 91–95.
Yuyama, L.K.O., Aguiar, J.P.L., Yuyama, K., Clement, C.R., Macedo,
S.H.M., Fávaro, D.I.T., Afonso, C., Vasconcellos, M.B.A.,
Vannucchi, H., Pimentel, S.A. and Badolato, E.S.G. (2003)
Chemical composition of the fruit mesocarp of three peach palm
(Bactris gasipaes) populations grown in central Amazonia, Brazil.
International Journal of Food Sciences and Nutrition 54, 49–56.
Bactris spp.
Bactris Jacq. ex Scop. (Arecaceae) is one of the largest genera
of palms in the Americas and is notorious for the degree to
which most species are armed with black spines – on the
stems, leaves and inflorescences. In addition to the
economically important Bactris gasipaes (treated separately in
this volume, see previous entry) many other species produce
edible fruit (Table A.50). It is likely that fruit of species other
than those listed can be consumed.
World production and yield
The species of Bactris treated here are exploited chiefly from
wild populations or, at best, small plantings established by
subsistence agriculturists, thus no production data are available.
101
Uses and nutritional composition
No nutritional information is available on these species. The
mesocarp pulp of many of the fleshy-fruited species have an
agreeable flavour that has been likened by some to that of
grapes. They are eaten fresh or juiced to make a refreshing
beverage.
Botany
TAXONOMY AND NOMENCLATURE The taxonomy of Bactris is
still poorly understood despite the ubiquity of the genus in
tropical forests throughout the Americas. Henderson et al.
(1995) recognize 64 species. Several of these form large
complexes that may ultimately be split up into distinct species.
The genus is classified in the tribe Cocoeae of subfamily
Arecoideae.
DESCRIPTION In general, many Bactris species can be
described as medium-sized, spiny clustering palms, but the
genus includes non-spiny species, tall-growing species and
solitary-stemmed species as well. Leaves are most frequently
pinnate but can be simple, and the canopy holds between four
and 20. Leaflets are typically clustered in groups that radiate
in different planes, but some species have regularly arranged
pinnae. The leaf sheath of most species extends above the
point of insertion of the petiole. Spineless species are the
exception, but in a few species the spines are reduced in size
or restricted to the leaflet tips. The inflorescences are spikelike or branched to one order and are borne from among the
leaves. The rachillae are slender, and separate male and female
flowers are arranged on them in a triad of one central female
flanked by two males, but the female flower may be missing on
some branches or portions thereof. The one-seeded fruit vary
from globose to ovoid to ellipsoid, and are coloured green,
orange, red or dark purple. They are generally smooth. A
group of purple-fruited species have a distinctive cup-like
structure at the proximal end of the fruit. The mesocarp can
be either mealy or juicy. The seed is covered by a bony
endocarp with three pores near its middle.
ECOLOGY AND CLIMATIC REQUIREMENTS Bactris species are
distributed from Mexico and the West Indies through Central
America and throughout tropical South America. They are
chiefly denizens of wet tropical rainforest, usually at lower
elevations. They have little tolerance of frost and prosper with
year-round rainfall.
REPRODUCTIVE BIOLOGY Weevils have been implicated as
pollinators of some Bactris species (Essig, 1971; Urpi, 1982).
The typical Bactris phenological syndrome is short nocturnal
anthesis of the pistillate flowers followed by a brief period of
staminate anthesis (Henderson et al., 1995).
Horticulture
PROPAGATION Solitary-stemmed
Bactris
species
are
propagated by seed, which should be fresh. The pulpy or
mealy mesocarp must be cleaned from the seed before sowing.
Vegetative propagation of clustering species is possible by
division.
Alan W. Meerow
102
Arecaceae
Table A.50. Bactris species other than B. gasipaes with edible fruit.
Species
Synonyms
Bactris arundinaceae
Trail.
Bactris brongniartii
Common name
Origin
Characteristics
Fruit uses
References
Syn. of B. tomentosa
var. tomentosa
according to
Henderson et al., 1995
Palmeira lú-I (Brazil)
Brazil
Clustering, flattened, yellowish leaf
spines, spicate inflorescence,
fruit purple-black
Pulp
Martin et al., 1987
B. burretii Glassman,
B. marajaacu Barb.
Rodr., B. piscatorum
Wedd. ex Drude
Marajá (Brazil),
Amazon region
chacarra, cubarro
and adjacent areas
(Colombia), bango
palm (Guyana),
ñejilla (Peru),
caña negra (Venezuela)
Clustering, often rhizomatous;
Pulp
leaflets bifid at tip; spine flattened
and yellowish brown in middle; fruit
purple with cup-like structure
Henderson et al., 1995
Marajaú (Bolivia),
chontilla (Ecuador),
ñejilla (Peru),
marajá (Brazil)
Western Amazon
Stems clustered, to 8 m, 1.5–5 cm
wide; fruit congested, irregularly
and narrowly obovoid, 2–4.5 ⳯
1–2.5 cm, purple black
Fleshy fruit
marketed locally.
Pulp eaten fresh.
Also fed to animals
Fouqué, 1973; Henderson et al.,
1995
Sp: corozo, lata
(Colombia), biscoyol
(Costa Rica), coyolito
(Nicaragua), uvita
de monte (Panama),
piritu, uvita
(Venezuela), uiscoyol
Central America to Stems clustered, spiny, to 3 m,
north of Colombia 2.5–3 cm wide; fruit depressed
and Venezuela
globose, 1.5–2 cm, purple black
Pulp eaten fresh.
Refreshing drink
Fouqué, 1973; Henderson et al.,
1995
Sp: chontilla (Bolivia),
chinamato (Colombia),
pijuayo del monte
(Peru), macanilla
(Venezuela);
Po: pupunha brava
Venezuela (Barinas,
Cojedes, Zulia),
Colombia (north
and Valle), Peru
(Huánuco, Madre
de Dios), Brazil
(Acre, Rondônia)
and Bolivia (Santa
Cruz)
Edible fruit
Henderson et al., 1995
Bactris concinna Mart.
Bactris guineensis
(L.) H.E. Moore
Subspecies
var. concinna,
var. inundata,
var. sigmoidea
B. horrida Oerst.,
B. minor Jacq,
B. oraria L.H. Bailey,
B. rotunda Stokes
Bactris macana (Mart.) B. caribaea H. Karts.,
B. dahlgreniana
Pittier
Glassman
Stems solitary or clustered, spiny,
to 12 m, 10–20 cm wide; fruit
subglobose to obovoid, 1–1.6 cm,
orange. Possibly the wild ancestor
of B. gasipaes
Bactris major Jacq.
B. subglobosa
H. Wendl., numerous
others
var. major, var.
megalocarpa,
var. infesta,
var. socialis
Sp: marayáu (Bolivia),
corozo de gallina, lata
(Colombia), huiscoyol
(El Salvador,
Guatemala, Honduras,
Nicaragua), jahuacté
(Mexico), caña brava
(Panama), cubarro,
moporo (Venezuela),
viscoyol; Po: marajá;
En: beach palm,
beach spiny club
palm, hones (Belize);
Fr: zagrinette
Bactris maraja Mart.
B. monticola Barb.
var. chaetospatha, Sp: chontilla (Bolivia,
Rodr., B. piranga Trail., var. juruensis,
Colombia, Peru),
cacharra, espina
numerous others
var. maraja
(Colombia), uvita
(Panama), ñeja (Peru),
uva de montaña, piritu
(Venezuela), niejilla;
Po: marajá, marja açu,
tucum bravo
Lowlands from
Costa Rica to north
of South America
including central
and western
Amazon, the
Guianas and Bahia
in Brazil
Stems clustered, spiny; fruit
Pulp eaten fresh;
widely depressed, obovoid, 1–2 cm, seed sometimes
purple black, occasionally minutely eaten
spinulose; mesocarp juicy
Cavalcante 1991; Henderson et
al., 1995
Bactris setosa Mart.
B. cuyabensis Barb.
Rodr., B. polyclada
Burret
Atlantic coast of
Brazil
Stems clustered, to 6 m, 3–4 cm
wide, spiny; fruit depressedglobose, 1–1.5 ⫻ 1.5–2 cm, purple
black; mesocarp juicy
Martin et al., 1987
Po: tucum, jucum
Coasts of Central
Stem forming dense clumps, 10 m,
America and north 2.6 cm wide, with spines 5 cm long;
of South America fruit ellipsoid or obovoid, 2.5–4.5
⳯ 1.3–3.5, apiculate, purple black;
mesocarp yellowish, fibrous and
juicy
Pulp eaten fresh,
drinks; seed
sometimes eaten
as well
Juicy pulp
Fouqué, 1973; Henderson et al.,
1995
Borassus
103
104
Arecaceae
Literature cited and further reading
Cavalcante, P.B. (1991) Frutas Comestíveis da Amazônia. Edições
CEJUP, Brazil, 279 pp.
Essig, F.B. (1971) Observations of pollination in Bactris. Principes 15,
20–24.
Fouqué, A. (1973) Espèces fruitières d’Amérique tropicale. Fruits 28,
290–299.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
Martin, F.W., Campbell, C.W. and Ruberté, R.M. (1987) Perennial
Edible Fruits of the Tropics – An Inventory. Agriculture Handbook
No. 642. US Department of Agriculture, Agricultural Research
Service, Washington, DC.
Urpi, J.M. (1982) Polinización en Bactris gasipaes H.B.K. (Palmae):
nota adicional. Revista Biologia Tropiale 30, 174–176.
Borassus flabellifer
palmyra palm
Palmyra palm, Borassus flabellifer L. (Arecaceae), is a slowgrowing fan palm with a broad Old World distribution and a
venerable history of cultivation by man. Its sap is an important
source of sugar and the fruit and seeds have been consumed
by people for centuries.
or four will remain after detailed revision of the genus. Some
authors feel that only a single species, B. flabellifer, should be
recognized. It is member of the tribe Borasseae, subfamily
Coryphoideae, all dioecious costapalmate fan palms. Borassus
aethiopicum is very similar in all respects to B. flabellifer.
The palmyra palm is a large tree up to 30m high
and the trunk may have a circumference of 1.7 m at the base
(Fig. A.13). There may be 25–40 fresh leaves. The leaves are
leathery, grey-green, fan-shaped, 1–3 m wide, folded along the
midrib, and divided to the centre into 60–80 linear-lanceolate,
0.6–1.2 m long, marginally spiny segments. Their strong,
grooved petioles, 1–1.2 m long, black at the base and blackmargined when young, are edged with hard spines. The palms
are dioecious, and male and female inflorescences differ in their
order of branching (the males twice, the female unbranched or
branched but once). The staminate flowers are borne on catkinlike rachillae, the pistillate flowers on thicker branches. The
fruit are large drupes, 10–20 cm in diameter, brown, with a
fibrous mesocarp. They contain two or three large seeds.
DESCRIPTION
AND CLIMATIC REQUIREMENTS Palmyra palm
grows from the Persian Gulf to the Cambodian–Vietnamese
border and is commonly cultivated in India, South-east Asia,
ECOLOGY
World production and yield
Each palm may bear between six and 12 bunches of about 50
fruit/year. An average crop of B. flabellifer in Ceylon is 350
fruit.
Uses and nutritional composition
The chief product of the palmyra is the sweet sap (toddy)
obtained by tapping the tip of the inflorescence, as is done
with the other sugar palms and, to a lesser extent, with the
coconut. Small fruit are pickled in vinegar. In April and May
in India, the shell of the seed can be punctured with a finger
and the sweetish liquid sucked out for refreshment like
coconut water. Immature seeds are often sold in Indian
markets. The kernels of such young seeds are obtained by
roasting the seeds and then breaking them open. The halfgrown, soft-shelled seeds are sliced longitudinally to form
loops, or rings and these, as well as the whole kernels, are
canned in clear, mildly sweetened water, and exported. Tender
fruit that fall prematurely are fed to cattle. The pulp of mature
fruit is sucked directly from the wiry fibres of roasted, peeled
fruit. It is also extracted to prepare a product called punatoo in
Ceylon. It is eaten alone or with the starch from the palmyra
seedlings. The fresh pulp is reportedly rich in vitamins A and
C. The fruit contains, per 100 g, 43 calories, 87.6 g water, 0.8 g
protein, 0.1 g fat, 10.9 g total carbohydrate, 2.0 g fibre, 0.6 g
ash, 27 mg calcium, 30 mg phosphorus, 1.0 mg iron, 0.04 mg
thiamine, 0.02 mg riboflavin, 0.3 mg niacin and 5 mg ascorbic
acid.
Botany
While seven species have
been described in the genus Borassus, it is likely than ony three
TAXONOMY AND NOMENCLATURE
Fig. A.13. Borassus flabellifer palm (with permission from Sitijati
Sastrapradja from Palem Indonesia, Lembaga Biologi Nasional, 1978).
Brahea
Malaysia, tropical Africa and occasionally in other warm
regions including Hawaii and southern Florida. Its area of
origin is not known but undoubtedly man has influenced its
range, and large populations, in many areas covering
thousands of hectares, that appear ‘wild’ may be naturalized
from introductions made more than 2000 years before the
present. It is found in seasonally dry areas in the main, and is
hardy to about –4°C. Though well adapted to dry, tropical
climates, the palmyra grows better with regular irrigation, but
will not tolerate waterlogged soils.
Inflorescences begin to appear in
November–December, but anthesis does not occur until
March. Fruit mature in July and August.
REPRODUCTIVE BIOLOGY
FRUIT DEVELOPMENT The coconut-like fruit are three-sided
when young, becoming rounded or more or less oval,
12–15 cm wide, and capped at the base with overlapping
sepals. The outer covering is smooth, thin, leathery and
brown, turning nearly black after harvest. Inside is a juicy
mass of long, tough, coarse, white fibres coated with yellow or
orange pulp. Within the mature seed is a solid white kernel
(endosperm) which resembles coconut meat but is much
harder. When the fruit is very young, this kernel is hollow, soft
as jelly and translucent like ice, and is accompanied by a
watery liquid, sweetish and potable.
Horticulture
PROPAGATION The seeds of palmyra palms germinate remotely
and deeply, producing a very long cotyledonary petiole (‘sinker’
or ‘dropper’) that requires a deep container. The cotyledonary
petiole may bury itself as much as 0.5 m below ground before a
shoot forms. Seeds begin to germinate in 2–6 weeks.
MAINTENANCE AND TRAINING Palmyra palms are very slow
growing, and do not even show any aerial stem elongation for
the first 15–20 years of their life. Flowering may begin at
12–15 years of age, and will continue for about 50 years.
DISEASES, PESTS AND WEEDS
Bud rot fungi such as Pythium
palmivorum and Phytophthora palmivora are the most serious
pathogens of palmyra palms, but a number of other fungal
diseases cause foliar blights. Rhinoceros beetle (Oryctes
rhinoceros), black-head caterpillar (Nephantis serinopa) and red
palm weevil (Rhynchophorus ferrugineus) can seriously infest
the palms (Duke, 2001).
MAIN CULTIVARS AND BREEDING Some regional variation,
mostly in terms of ecological tolerances, has been selectively
developed, but no formal cultivars are recognized.
Alan W. Meerow
Literature cited and further reading
Davis, T.A. and Johnson, D.V. (1987) Current utilization and further
development of the palmyra palm (Borassus flabellifer L.,
Arecaceae) in Tamil Nadu State, India. Economic Botany 41,
247–266.
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
105
Morton, J.F. (1988) Notes on distribution, propagation, and products
of Borassus palms (Arecaceae). Economic Botany 42, 420–441.
Sambou, B., Lawesson, J.E. and Barfod, A.S. (1992) Borassus
aethiopum, a threatened multiple purpose palm in Senegal.
Principes 36, 148–155.
Brahea dulcis
palma de sombrero
Palma de sombrero, suyate, capulín and soyate, Brahea dulcis
(Mart.) Becc. (Arecaceae), is a widespread fan palm found in
Mexico and Central America, the edible fruit of which are
sweet and of an agreeable flavour.
Related species with edible fruit include Brahea armata S.
Watson, blue hesper palm, which is found in northern Baja
California and north-west Mexico. The Yuman Indians grind
the seeds of this palm into a meal, and also eat the fruit fresh
and use the juice for drinks. Guadalupe palm, Brahea edulis,
native to Guadalupe Island off the coast of Baja California,
also has edible fruit and it is said to taste similar to dates. It is
extremely rare in the wild, due to predation by feral goats
(Henderson et al., 1995). Also Brahea aculeata from western
Mexico is said to have edible fruit.
Uses and nutritional composition
Brahea dulcis is not produced commercially. The fruit are
eaten locally. The fruit are picked when ripe and can be eaten
fresh or made into preserves. With refrigeration they can be
stored for a month or longer.
Botany
TAXONOMY AND NOMENCLATURE The genus Brahea consists
of six to nine species of mostly Mexican fan palms, though
several species range south to Central America. They are
found in dry woodland to semi-desert. It is classified in the
tribe Corypheae of the subfamily Coryphoideae. Some
synonyms for B. edulis include Brahea bella L.H. Bailey,
Brahea calcarea Liebm., Brahea conzatti Bartlett and Brahea
salvadorensis H. Wendl. Ex Becc. Henderson et al. (1995) list a
number of others.
DESCRIPTION Brahea dulcis is a solitary or clustering fanleafed palm with stems reaching 2–7 m in height and
12–20 cm in diameter. The stems often lean. The canopy
consists of ten to 15 dull-green, sometimes waxy, palmate
leaves with toothed petioles. The blade is split down to its
middle into 30 to 50 stiff segments. The long, arching
branched inflorescences emerge from among the leaves and
are densely hairy. The one-seeded fruit are ovoid, 1–1.5 cm
long and brown or green.
ECOLOGY AND CLIMATIC REQUIREMENTS Brahea dulcis is
found in dry, oak woodlands or open areas on rocky, calcareous
soils from 300 to 1700 m. Though hardy to at least –5°C, it
grows poorly in humid, subtropical or tropical climates, but
thrives in Texas, Arizona and California and similar climates.
It ranges from eastern and southern Mexico up the Mexican
Pacific coast and south as far as Nicaragua.
106
Arecaceae
Horticulture
Propagation is from seed, which germinate in 2–4 months. The
seed stores well at room temperature for at least a year. It is
rather slow growing. It and other Brahea species are planted as
ornamentals in semiarid climates with mild winters.
Alan W. Meerow
Literature cited and further reading
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
ECOLOGY AND CLIMATIC REQUIREMENTS Butia capitata is a
palm of open savannah (the cerrado and restinga vegetation of
Brazil), growing on sandy soils from southern Brazil into
northern Argentina, Paraguay and Uruguay at low elevation. It
is hardy to at least –12°C.
Horticulture
PROPAGATION Pindo palm is easily propagated from seed,
which germinates unevenly over several months to a year
unless the bony endocarp surrounding the seed(s) is cracked.
DISEASES, PESTS AND WEEDS
the seed.
Butia capitata
pindo palm
Pindo palm, also known as jelly palm, butiá and cabeçudo,
Butia capitata (Mart.) Becc. (Arecaceae), is a modest-sized
single-stemmed palm widely grown as an ornamental in warm
temperate and subtropical areas of the world. The fleshy/
fibrous fruit is pleasantly sweet and flavourful.
Related species with edible fruit include Butia eriospatha
(Mart.) Becc., which is found in open areas and within Araucaria
forests of southern Brazil and is very similar in appearance to B.
capitata but the spathe bract of the inflorescence is densely
tomentose. The fruit of this species are soaked in alcohol to make
a beverage. Butia yatay (Mart.) Becc. is larger in all respects than
B. capitata, and is reportedly hardier. It ranges from southern
Brazil to northern Argentina and Uruguay.
Uses and nutritional composition
This palm is not produced commercially and the fruit of
B. capitata are eaten locally or made into preserves. No
information on their nutritional value is available.
Botany
TAXONOMY AND NOMENCLATURE Butia is a small genus of
about eight species closely related to Syagrus and capable of
hybridizing with species of that genus. Most of the species are
endemic to Brazil. Butia capitata occurs in two distinct and
widely separated population clusters, and some palm specialists
have advocated recognizing the southernmost populations as a
distinct species, Butia odorata (Henderson et al., 1995). The
genus is classified with the tribe Cocoeae of subfamily Arecoideae.
Pindo palm is a solitary-stemmed feather palm
growing 2–6 m in height. The trunk can reach about 0.5 m in
diameter and remains clothed in old leaf bases for many years.
The canopy consists of 18–32 arching leaves that vary from
yellowish green to greyish green. Each leaf is 2.5–3 m long.
The petiole is short, broad and armed with fibre spines.
Arranged regularly along the rachis are 44–48 pairs of stiffly
upright leaflets that form a distinct ‘V’. The inflorescence is
many-branched but fairly short, substended by a conspicuous
woody bract. Yellow male and female flowers are borne on the
same rachillae. The ovoid fruit are 1.8–3.5 cm long,
1.2–2.2 cm wide, vary from yellow to orange and contain one
to three seeds surrounded by a bony endocarp with three
pores near the middle.
DESCRIPTION
Bruchid weevils sometimes infest
Alan W. Meerow
Literature cited and further reading
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
Calamus rotang
rattan
Rattan, Calamus rotang L. (Arecaceae), is one of nearly 400
species of mostly climbing vines found throughout South-east
Asia, Australasia and tropical Africa. The genus is the largest
in the entire palm family. The most important economic use of
these vining palms is for their long, strong and flexible stems
(canes) that are a significant source of export income for the
countries where they grow. The seeds of many species are
surrounded by a sweet, pulpy aril that is locally consumed.
Calamus rotang is the type species for the genus and is used
herein as representative.
World production and yield
The collection of fruit for consumption can only be
considered an incidental use for rattan vines. The production
of cane for furniture manufacture is an important industry in
India, the Philippines, Malaysia and other areas rich in
species. In the Philippines, the industry provides jobs for
10,000 people, and the value of raw rattan has been estimated
at US$50 million, with finished products valued at US$1.2
billion (Duke, 2001). Cirebon, a city in West Java, Indonesia,
produces almost half of all Indonesian rattan products and
exports 1500 containers of furniture each month, worth
US$15 million. The Malaysian Timber Council estimated the
total value of rattan furniture exported from Malaysia at over
US$90 million.
Uses and nutritional composition
The fruit of Calamus ornatus contains, per 100 g, 79 calories,
0.6 g protein, 1.2 g fat, 18.6 g total carbohydrate, 0.5 g fibre, 19
mg calcium, 10 mg phosphorus, 1.7 mg iron, 0.06 mg
thiamine, 0.01 mg riboflavin, 0.9 mg niacin and 5 mg vitamin
C (Duke, 2001). The subacid pulp surrounding the seed is
thirst quenching. The fruit are sometimes pickled in brine as
well, and the seeds are also edible. These uses are minor ones
by comparison to the great economic importance of the long
flexible stems of the many species of rattan vines.
Cocos
Botany
TAXONOMY AND NOMENCLATURE The taxonomy of the
rattans is poorly understood, no doubt due to the difficulty in
collecting specimens of the spiny stems, but at least 400
species are estimated to be in the genus Calamus. Calamus is
one of several genera in the subfamily Calmoideae that are
called rattans, all of which consist primarily of high-climbing
vines.
DESCRIPTION The slender stems of C. rotang can be up to
200 m in length, and typically climb into the canopy of
rainforest trees. The leaf sheaths and petioles are armed with
straight or recurved spines. The leaves are 60–90 cm long,
pinnate with numerous, narrow leaflets, 20–23 cm long and
1.3–2 cm wide, sometimes with spines on one or both surfaces
along the midrib. The staminate inflorescence is slender, whiplike, branched to one order, and often spiny. The pistillate
inflorescence is slightly more robust, with catkin-like, recurved
branches. The apex of the inflorescence is modified as a
climbing grapnel by which the stems are able to ascend to the
canopy of forest trees over 100 m tall. The fruit are 1.5–2 cm
in diameter, pale yellow in colour and covered with
overlapping scales in vertical rows.
107
the shade, treated with fungicide, then planted directly or
stored in moist sawdust for several days. Seeds begin to
germinate in 65–100 days. Seedlings about 15 cm tall with
four or five leaves are planted directly in the field. Two
seedlings are planted in each hole, each hole spaced 2 m on
centre.
In order to grow properly,
rattan has to be planted under some sort of tree cover, such as
logged-over forest, secondary forest, fruit orchards, tree
plantations or in rubber plantations.
MAINTENANCE AND TRAINING
NUTRITION AND FERTILIZATION Fertilization at 6 g per plant
of 20:10:5 N:P:K is recommended shortly after planting, and
an organic mulch is beneficial during the first 2–3 years (Duke,
2001).
DISEASES, PESTS AND WEEDS
The fungi Catacaumella calamicola, Doratomyces tenuis and Sphaerodothis coimbatorica are
significant foliar pathogens of rattans (Duke, 2001). Various
wood-boring insects can infest the canes after harvest.
Alan W. Meerow
Literature cited and further reading
Rattans are forestdwelling plants in the wild, and are more abundant in primary
than secondary forest. They are exclusively tropical plants and
have no tolerance of frost, preferring acid soils with abundant
moisture and organic matter. Most rattan in commerce was
historically harvested from the wild, which has resulted in
significant reduction in their occurrence.
ECOLOGY AND CLIMATIC REQUIREMENTS
REPRODUCTIVE BIOLOGY All Calamus species are dioecious
palms. Alloysius (1999) observed no clear relationship between
flowering and climatic conditions in Calamus manan Miq. in
Malaysia. Alloysius (1999) reported that C. manan flowered
each year in October–December with fruit maturing 16–17
months later. Staminate plants flower for a longer period than
pistillate plants. Anthesis occurs at night, suggesting nocturnal
insects (moths) might be pollinators, with bees the main
diurnal flower visitors. Bogh (1996) studied phenology and
pollination of Calamus longisectus Griff., Calamus peregrinus
Furt., Calamus rudentum Lour. and an unidentified species in
Thailand. He found that staminate plants flowered almost
continuously for several months, while pistillate plants have
much shorter flowering periods. He also determined that
Trigona bees were the most important pollinators.
FRUIT DEVELOPMENT Abdullah (2000) reported that fruit
production took 8–9 months for Calamus palustris, and 12–13
months for Calamus scipionum and Calamus ornatus in
peninsular Malaysia.
Horticulture
Rattans are propagated chiefly by seed, but
vegetative propagation by division of clumps or by suckers is
possible. The pericarp is removed from the fruit, which is then
fermented in water for about a day, after which the seed is
cleanly squeezed from the pulp by hand. The seed is dried in
PROPAGATION
Abdullah, M.Z.B.H. (2000) Reproductive biology and phenological
observation of three Calamus species in peninsular Malaysia. PhD
thesis, Universiti Putra, Malaysia.
Alloysius, D. (1999) Reproductive biology of Calamus manan Miquel.
PhD thesis, Universiti Putra, Malaysia.
Bogh, A. (1996) The reproductive phenology and pollination biology
of four Calamus (Arecaceae) species in Thailand. Principes 40,
5–15.
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Cocos nucifera
coconut
The coconut (Cocos nucifera L., Arecaceae) is instantly
recognized and obviously different from any other fruit or nut.
The coconut palm is romantically associated with beautiful
tropical beaches and is the most widespread and most easily
recognizable of all palm trees. The closest botanical relatives
(Cocoidae) largely occur in South America yet the centre of
diversity for this important crop species is in the islands of
South-east Asia and the Pacific. From the mid-19th century
until the 1960s, the dried kernel of the coconut, copra, became
the most important source of vegetable oil in international
markets. The oil is used for making candles, soap and high
explosives.
Strictly speaking the coconut is not an achene but a drupe
(like a plum). Its characterization as a ‘nut’ has tended to hide
the significant value of the fibres in the husk of the mature
fruit and the watery endosperm that fills the cavity of the
immature fruit. It is these two characters, rather than the oil
content of the kernel, that account for the original natural
spread of the coconut, by floating, and its value to people
travelling long distances at sea. Air space between the fibres
and the volume of the nut cavity gave coconut its most
108
Arecaceae
distinguishing feature – the ability to float hundreds or
thousands of kilometres between tropical oceanic islands and
germinate, even on newly emerged volcanic islands. This
ability enabled its earliest, pre-human, dispersal. Subsequent
human dissemination to places that it could not float to made
the coconut the first truly pantropical plant and it quickly
became the most convenient source of vegetable oil (for more
than a century from the mid-1800s to the late 1900s) for high
lauric acid content. Although the demand led to considerable
colonial plantation developments during that time, today more
than 95% of coconut production is in the hands of small,
economically weak producers. Coconut palms grown by these
farmers are already less productive than African oil palm.
Neither of these labour-intensive tropical crops are able to
directly compete with mechanically harvested, genetically
modified, high lauric rapeseed (canola). The future of the
coconut is assured by its multitude of uses and its natural
beauty in the tropical landscape.
World distribution and yield
The coconut (cocoa-nut, cokernut (archaic); côco, coquos
(Portuguese); coco, cocotero (Spanish); cocotier (French);
kokosnuss (German); kokosnoot (Dutch); mnazi (Swahili);
narikal (Arabic); kerala, narial (Hindi); tennai (Tamil); pol
(Sinhala); kelapa (Malaysian and Indonesian); niyog (Tagalog);
and niu (Polynesian/Hawaiian)) is found throughout the
tropics. The greater number of coconut types in South-east
Asia–Melanesia that have numerous local names and uses and
unique insect and crab associations, attest to a long presence,
making this region the centre of diversity. This region is
assumed to be the centre of origin, with the coasts of America
at the ends of eastward and westward movement, possibly
from New Guinea and Polynesia with man or by drifting
(Ward and Brookfield, 1992). The coconut cultivars on the
Pacific coast of tropical America, from Mexico to Peru, are
indeed distinct from those in the coasts and islands of the
Caribbean and the Atlantic coasts of South America and West
Africa but this can be attributed to 16th-century Portuguese
and Spanish activities.
The closest botanical relatives of the coconut (once
classified as other Cocos species) occur in South America,
southern Africa or Madagascar raising the possibility that the
true centre of origin for Cocos was at the conjunction of South
America and southern Africa when those two continents were
part of the Gondwana super-continent. If that were so, then a
sea level, coastal distribution of a primordial coconut due
to natural dispersal by floating (Harries, 2002) occurred
synchronously with the movement of the Indian subcontinental tectonic plate from Gondwana across the
primordial Tethys Sea. Geologically, this is supported by
tropical marine sediments identified in Himalayan strata and
the presence of cocosoid fossil fruit and stems (Sahni, 1946)
on the Indian subcontinent. Further prehistoric spread into
the Pacific is indicated by cocos-like fruit in New Zealand and,
more contentiously, a fossil fruit in Australia and pollen cores
in the Cook Islands. Such natural dispersal of ancestral
coconut palm can account for the predominance of coconut
cultivars with ‘wild type’ attributes on some tropical coasts
and remote islands from the Indian Ocean to the mid-Pacific.
Subsequently, domestication in South-east Asia–south-west
Pacific, followed by introgression of wild and domestic types,
led to human dissemination inland, upland and by boat to
coastlines to which the wild type could not float.
The numerous uses of the coconut palm and its fruit are
sometimes thought to have led to its wide cultivation in the
tropics. The reverse is the case. It had a wide natural
dissemination before Polynesians, and then Europeans, took it
for one use in particular – as a source of pure, fresh, sweet
drinking water – to regions to where it could not float. The
other uses, such as coir fibre for ropes, became important for
the mercantile activities of sailing ships and then oil, from
copra (the dried endosperm of the nut) led to coconuts being
traded commercially everywhere along sea coasts, grown at
inland sites having an adequate water supply, and even planted
in agriculturally unsuitable places in the tropics.
Uses and nutritional composition
Humans have used this palm for thousands of years for food,
thatch, fibre and wood, and it entered commercial trade as oil for
lamps, candles and soap. Coconut fruit is used fresh at both the
immature and the mature stage. The kernel (solid endosperm)
from mature coconuts can be shredded and dehydrated to
become desiccated coconut, or coconut oil can be extracted and
processed as an edible oil from the fresh kernel or as an
industrial oil from the kernel that has been dried (in the sun,
over a fire, in a kiln) or fried (in hot oil) to copra. The outer husk
(mesocarp) of the fruit (‘nut’) is used as a fibre (coir), along with
a non-fibrous product, coir dust (cocopeat). It now has
widespread uses in horticulture as a replacement for peat moss.
There has always been local trade in immature fruit and
dehusked mature nuts for eating but modern technology is
making coconut water available internationally. Fresh coconut
water is an important adjunct in media for tissue culture. The
juicy, jelly-like endosperm in the young nuts is highly prized
for eating out of the shell, for the water and for use in cooking.
Immature (‘green’, ‘jelly’) coconuts are harvested 7–9 months
after pollination by a climber so that the entire bunch can be
lowered by rope to the ground to avoid damage. The
epidermis is uniformly fresh in colour (shades of green,
brown, red or yellow depending on variety) and smooth, while
the coir is white. The dehusked fruit is about 10 cm in
diameter weighing about 500 g, having 100 g endosperm, 120
g shell and 250 g liquid endosperm (water). The liquid
endosperm in fresh ‘green’ coconuts can have 130–620 ml
water and 48 g/kg sugar, depending on the stage of harvest,
and is at a maximum 7–8 months after anthesis. The water is
marketed in sterilized long-life packs in South-east Asia and
Brazil. Mechanical damage will cause the white coir to turn
brown and can cause nut cracking. Younger nuts rupture with
less force than mature nuts (108 versus 537 kg) (Tongdee,
1991). The nuts are held in a cool place until processed or sold
at market. In Thailand, the green nut is trimmed and shaped,
removing most of the husk. The final product has a flat
bottom, round body with a pyramid top and the eyes showing.
Alternatively, all the husk is removed before dipping in the
sulphite solution.
To enter the fresh market chain, mature nuts are dehusked
before shipment. The dry nuts should be brown, free from
Cocos
fibre, damage and cracks and of the required weight or size (35
to 45 cm in circumference). The nuts are shipped in sacks or
cardboard cartons. Postharvest stress cracks are directly
related to coconut weight loss (Burton, 1982). Waxing of the
nuts minimizes water loss and dramatically reduces fruit
cracking. The shelf life is 2–3 months at 12°C before the
residual liquid has evaporated or the shell has cracked because
of desiccation. Low humidity and high temperature must be
avoided.
Mature nuts for copra, coir or desiccated coconut are left on
the palm until 11 months or more from pollination when the
fresh skin colour shows dry, brown patches to being fully
brown and the coir is brown. Mature nuts of some
varieties/cultivars fall to the ground when ripe and can be
collected, but more commonly all nuts on one or more ripe
bunches are cut using a blade on a pole at regular intervals
every 4–6 weeks, throughout the year. Monkeys can also be
trained to harvest coconuts. Mature nuts are comparatively
light and bulky, while green nuts are three times more dense.
The oil is extracted from copra and used for soap, detergent
and margarine, while the desiccated coconut is used in
confectionery. The processing of copra and the use of the
husks has been extensively reviewed (cf. Woodroff, 1970; Ohler,
1999). The fresh kernel (28% whole nut) contains 25–44% oil,
35–62% water, 9–14% carbohydrate, about 5 mg/100 g
vitamin C, as well as vitamins B1, B2 and B3 (Table A.51).
Coconut oil becomes solid at temperatures below about 25°C
and was used as ‘vegetable butter’ until the hydrogenation
process for making margarine from other vegetable oils was
developed in the 1890s. However, the use of glycerin, a byproduct of soap manufacture, to make nitroglycerine expanded
the demand for copra at the time of World War I (Harries et al.,
2003) and also in World War II, when Japan took control of all
major coconut-producing countries.
Table A.51. Proximate fruit composition of coconut (Source: Leung et al.,
1972; Siong et al., 1988).
Amount in 100 g edible portion
Variable
Proximate (g)
Water
Calories (kcal)
Protein
Fat
Carbohydrate
Fibre
Ash
Minerals (mg)
Calcium
Phosphorus
Iron
Sodium
Potassium
Vitamins (mg)
Ascorbic acid
Carotene
Thiamine
Niacin
Riboflavin
Immature
Mature
Water
Toddy
81.4
122
1.9
11.9
4
0.7
0.8
55
296
35
27.2
13.7
3.8
1
94
22
0.2
0.4
4.5
–
0.5
86
43
0.3
0.4
10
Trace
11
42
1.1
51
257
13
83
1.8
16
340
24
18
0.3
5
130
Trace
–
Trace
34
109
7
Trace
0.05
0.8
0.03
5
0
0.04
0.6
0.03
3
0
Trace
0.1
Trace
29
Trace
0.02
Trace
0.01
109
Numerous coconut recipes exist for its use in breads,
waffles, cakes, sweets, cookies, pies, ice cream, soup and other
main-course cooked dishes and the health benefits of dietary
coconut oil are becoming better known (Fife, 2000). In the
Philippines, acetic acid bacteria produce a ‘cartilaginous’
material from coconut water called nata de coco that is used in
a number of desserts.
The fibre from the husk is used for ropes, mats and
geotextiles. More recently the residual coir dust (cocopeat) is
replacing peat moss for environmentally sustainable,
horticultural seed-sowing and potting mixtures. Unopened
coconut inflorescences are tapped and the exudate (toddy)
collected and fermented for an alcoholic brew (palm wine) of
up to 12–13% alcohol. It is also distilled to produce a ‘whisky’.
Alternatively, the toddy is boiled down to produce a treacle or
sugar. Freshly gathered toddy has about 8.6% total soluble
solids, pH 3.6, 0.23% crude protein, 0.6% sucrose, 5.7%
reducing sugars and can have about 5% alcohol depending
upon collection frequency and hence the time allowed for
fermentation to occur.
Botany
TAXONOMY AND NOMENCLATURE The genus Cocos is
monotypic, containing only the highly variable C. nucifera L.
and occurs within the Arecoideae subfamily, tribe Cocoeae, as
do other economic palms such as the peach palm (subtribe
Bactridinae), oil palm (subtribe Elaeidinae) and betel nut (Uhl
and Dransfield, 1988). Previously, the genus contained over 30
other species that occurred in Central and South America;
these are now assigned to several other genera. A classification
of C. nucifera by analysis of the proportions of husk, shell and
endosperm in the fruit (Harries, 1978) identified the
geographical and historical distribution of populations with
wild-type, domestic-type and introgressed-type characteristics. Recently, sequence-tagged microsatellites and amplified
fragment length polymorphism (AFLP) methods have been
used to analyse genetic diversity (Teulat et al., 2000). While no
truly ‘wild’ coconuts are known, wild-type coconuts have been
identified in the Philippines (Gruezo and Harries, 1984),
Australia (Buckley and Harries, 1984) and elsewhere (Harries,
1990a).
This palm can grow 20–30 m high and live
80–100 years. The stem has only one terminal growing point,
no axillary vegetative buds and only rarely suckers from the
underground portion of the stem. Loss of the terminal
meristematic growing point leads to death. Branching of the
stem has been reported but is rare. During the initial years of
growth the stem gradually increases in thickness and then this
diameter is maintained until about 10 m when there is a
gradual reduction in diameter. Poor nutrition can also reduce
stem diameter. Early rapid stem growth occurs until fruiting,
then the rate of stem growth declines. The absence of a lateral
cambium layer means that there is no secondary thickening
and no capacity to repair injury. Stem strength and flexibility
is due to the fibrous sheath surrounding the numerous
vascular bundles in the stem periphery and a large number of
smaller bundle fibres in the stem vascular bundles (Tomlinson,
1990).
DESCRIPTION
110
Arecaceae
As a monocotyledonous plant, the coconut has an
adventitious root system. The primary roots are uniformly
thick, as are the secondary and tertiary, which have
progressive smaller diameters. These roots are produced from
the base throughout the plant’s life. A 25-year-old palm can
have 1500–4000 roots arising at the base of the bole or stem.
The long-lived primary roots produce branch roots about the
same thickness and short-lived rootlets. The older part of the
main roots and branches becomes sclerotic. About 25% of the
roots grow vertically downwards while the majority spread
horizontally and can reach 20 m from the palm and about 2 m
below the surface. However, about 70% of the roots are found
within a 1 m radius of the stem to a depth between 0.1 and
0.5 m (Cintra et al., 1993). The initial roots of the germinating
seedling can grow 1 cm/day, although this rate slows after
about 3 months of growth. Pruning of roots induces root
branching and continued growth.
The crown of the plant has about 30–40, open, 3–6 m long
leaves (fronds), 10–14 of which subtend fruit bunches at
different stages of development (Fig. A.14). There are also
30–50 unopened fronds in the crown. Leaves are produced in
succession, 8–20/year and take 4–5 months to emerge from
the sheath. The phyllotaxy is 5/2 or about 144° (Davis, 1970).
The rate of leaf production is higher in dwarf palms (17/year)
than tall palms (12/year). Leaves survive 3–3.5 years after they
are fully opened. After about 30 years, there is a gradual
reduction in the rate of emergence, leaf life and length and
Fig. A.14. Leaf, flower and fruit of Coco nucifera (Source: Vozzo,
2002).
therefore nut yield (Foale et al., 1994). Each leaf has 200–250
parallel-veined leaflets that are 70–80 cm long and 2.5 cm
wide at the base of the leaf, and about 45 cm long and 1.3 cm
wide at the apex.
The leaves intercept about 44% of the incident light
depending upon season and plant density (Nair and
Balakrishnan, 1976). This light interception makes coconut
suitable for mixed cropping. The functional leaf area in a
plantation is dependent more upon the intensity and duration
of seasonal drought and less on plant density. Damage to the
leaves by insects, storms or leaf pruning or clipping can lead to
significant nut yield reduction if more than 40% of the leaf
area is lost. The reduction in nut yield is due to nut shedding
and premature nut fall (Bailey et al., 1977). The impact of
50% leaf loss on yield can continue for 5 months and the loss
of 70% of the leaves for 17 months after defoliation. The
proportion of dry matter partitioned to the fruit can be as high
as 62%, in this C-3 plant (Corley, 1983).
This monoecious palm carries both staminate and pistillate
flowers borne on an axillary inflorescence (spadix) at each leaf.
Dwarf forms may begin flowering in 3 years, tall forms in 5–7
years. The first inflorescence may be all male, later
inflorescences will also produce female flowers. The immature
spadix is enclosed by a prophyll and peduncular bracts
(spathes). The spadix when mature and after emergence from
the bract is 1.2–1.8 m long, straw to orange coloured and a
simply branched rachis. Each branch (rachilla) bears one or
more female flowers near the base and numerous male flowers
above. There are generally up to about 50 female flowers per
bunch and possibly thousands of small male flowers.
The globose pistillate flowers are about 2.5 cm long and
3 cm in diameter with a reduced round perianth surrounding
the base. There is a short style with three stigmas, three ovules
are produced though normally only one is fertile. The
staminate flowers are small (3 mm) non-symmetrical with
small sepals and three longer petals and six stamens.
Pollinating insects are attracted by a small drop of nectar in
each newly opened male flower and by a prolonged nectar
supply for 2–3 days during the period of stigma receptivity of
each female flower.
ECOLOGY AND CLIMATIC REQUIREMENTS The palm grows
between latitudes 23°N and 23°S, with favourable
temperatures (27°C ± 7°C) and ground water or high, evenly
distributed rainfall, at elevations up to about 1000 m above sea
level. The canopy of coconut intercepts only about 40–50% of
the incident light. There is also little change in the canopy
spread with age and the limited root spread makes this an ideal
crop for intercropping. Numerous crops are grown under
coconut: fodder grass and other pastures for grazing, coffee,
cacao, long-kong, duku, pineapple, banana, maize, mango,
citrus, ginger, medicinal, aromatic and spice crops (black
pepper), yams, sweet potatoes, beans and groundnuts. Intercropping increases the income of small growers. Cash crops
are used among young coconut planting, and more long-term
crops such as bananas can be planted in alternate rows (Ohler,
1999).
Though the palm is grown on a wide range of soil types, it
yields best on rich river alluvial deposits with good drainage.
In most tropical countries, it grows on beach sands having low
Cocos
nutrient levels. In these sandy conditions, it requires higher
land or freshwater swamps to carry nutrients via percolation
towards the beaches. Management of coconuts on clay soil is
difficult, good drainage being essential. Soil pH of acid clays
(pH 5.0) to coral-derived sands (pH 8.0) are tolerated
(Murray, 1977).
Total rainfall of between 1300 mm and 2300 mm/year is
required for good production and the pattern of rainfall is
more important than the total amount. Irrigation is needed in
a new planting until the root system reaches the dry season
water table. In the absence of a water table, trickle irrigation is
recommended. Large plantations are not normally irrigated,
though nut and copra yield are reduced by drought. The rate
of application varies with soil type and weekly applications of
40–50 l/plant on an oxisol have been recommended in India
during the dry months (Salam and Mammen, 1990). The
mean transpiration rate of mature coconut palm is about 7.5
g/cm2/s and leads to a loss of about 90 l/day in each palm
(Kulandaivelu, 1990). Lower application rates mean that the
palm is still drawing some water from the soil.
It takes about 44 months from flower primordium initiation
to fruit maturity, including the 12 months from anthesis to
fruit maturity. Drought (or an extended dry season of 3
consecutive months) leads to inflorescence abortion, button
shedding, premature nut fall and low final nut yield. Hence,
rainfall in the first 3 months of nut development determines
crop size 12 months later. The effects of prolonged drought
can persist for up to 30 months.
Lightning can cause significant damage and leads to disease
at the damage site. Malaysia and Sri Lanka both report
lightning as a major contributing factor to disease.
A mean temperature of 27°C and diurnal variation of
6–7°C is considered optimum. These are found on tropical sea
coasts where the sea acts as a buffer against rapid temperature
changes. Mature coconut palms have survived frost and snow
in Florida but inflorescence abortion occurs at temperatures
below 15°C. Temperature is the deciding variable in altitude of
cultivation, a 20°C mean monthly temperature being the
minimum. At the equator, the limit for commercial production
is up to 1000 m and at 18°N (Jamaica) to 150 m. High
maximum temperatures can also decrease yield.
Too much shade or very cloudy conditions lead to poor
palm growth. The influence on yield has not been studied due
to the long nut development period. A minimum requirement
of 2000 h/year of sunshine has been suggested (Fremond et
al., 1966).
If adequate soil moisture is available, coconuts can tolerate
high winds. The windward palms may show some yield
reduction. Strong winds from hurricanes or cyclones can lead
to palm death by the crown being broken off, but palms that
are blown over may survive and root along the stem.
REPRODUCTIVE BIOLOGY Early flowering can be encouraged
by full sunlight, unrestricted water supply and additional
fertilizer applications. Reductions in these factors delay
flowering and constant shading can delay flowering
indefinitely. The inflorescence primordium is formed very
soon after the leaf. Little primordium growth takes place until
the subtending leaf growth has finished its expansion. The
inflorescence then begins to rapidly differentiate, followed by a
111
phase of elongation with the inflorescence opening about
12–13 months later. The number of spadices produced
depends on the rate of leaf production and amount of spadices
aborted. The number of pistillate flowers per spadix varies
with cultivar, palm age, season and palm management. Spadix
abortion is more common in young palms and during drought
(Menon and Pandalai, 1957).
The staminate flowers begin opening at the distal end of the
spadix during the morning, falling in the afternoon and the
whole process continues for upwards of a month on the same
spadix. There is a sweet scent at anthesis and nectar
production from the pistillodes in the base of the staminate
flowers. The female flowers produce nectar as they open from
the distal end, and anthesis lasts 6–15 days. Normally
staminate flower anthesis has finished before pistillate flowers
are receptive on the same spadix forcing cross-pollination in
most cultivars. However, during the warmer months, interspadix pollination on the same palm is possible in 40–50% of
the palms (Patel, 1938). Some dwarf cultivars have male and
female phases coinciding, leading to variable out-crossing
(Ashburner, 1995a). The presence of nectar and sweet scent
leads to considerable bee activity on both male and female
flowers. Flies and other insects may also be involved in
pollination and wind pollination may occur to a limited extent.
After anthesis, there is a period of immature nut (button)
shedding, most occurring in the first month. This shedding can
vary from 55 to 95% of the pistillate flowers and is due to a
number of causes: defective pollination, drought, disease, insects
or poor palm condition. There is a further shedding of more
mature nuts just before they are full grown and before the
endosperm has begun to form. This shedding is often greater
following a drought and accentuated by heavy rainfall following
drought. ‘Barren’ nuts can also develop, having an aborted
embryo. A higher number of female flower and nut set is
sometimes observed after tapping for toddy. In addition, dwarf
and hybrid palms carry a heavy load of nuts when they first
come into production. This last observation and biennial bearing
suggests a role for assimilate supply and demand in button nut
shed and immature nut fall (Foale, 1993). Several hormone
sprays particularly 2–4D and coconut water can double button
set and increase nut yield when applied after fertilization is
completed (Gangolly et al., 1956). Nut yield has been correlated
with a number of palm characters: total number of open leaves;
rate of leaf production; trunk length; trunk girth; and number of
pistillate flowers (Menon and Pandalai, 1957).
FRUIT DEVELOPMENT The coconut fruit is a fibrous drupe,
containing a hard-shelled ‘nut’. The mature fruit is either
ovoid and angular or spherical, depending on cultivar,
20–30 cm long, weighing from 1–2 kg. It has a thin epidermis,
covering a thick fibrous mesocarp (coir), within which is a
hard lignified endocarp (shell) that is brown when mature. The
‘wild type’ has a long, angular fruit, containing an ovoid nut
whereas in the ‘domestic type’ both the fruit and the nut are
more spherical or oblate because the proportions of husk, shell
and endosperm differ due to natural or domestic selection
(Harries, 1978). Inside the endocarp, at maturity, the white
flesh of the kernel (endosperm) is about 12–15 mm thick with
a large central cavity. The pea-sized embryo lies in the flesh
under the ‘soft eye’, one of the three generative pores at the
112
Arecaceae
basal end of the nut. The embryo weighs c.1/1000 of the fruit
weight. Occasionally, there will be two or three viable
embryos, one under each generative pore.
The fruit take approximately 12 months to reach maturity,
depending on cultivar. About 32% of the endosperm is
deposited in the first 8 months and 94% by the 11th month of
development. When young, the mesocarp comprises the major
portion of the nut and increases in thickness and number of
fibres up to maturity (Shivashankar, 1991). The shell is already
differentiated before fertilization and further development
occurs after the mesocarp has differentiated, about 4 months
after fertilization. The endosperm is the last to develop and
begins as a liquid containing free nuclei and some cells. These
cells begin to coalesce towards the periphery of the embryo sac
on the endocarp about 7 months after fertilization. Additional
cells are formed and adhere to the endocarp, resulting in the
cellular peripheral endosperm that is initially translucent and
jelly-like, hardening to a white flesh at 11 months. Oil content
in the endosperm parallels its development. Coconut water
begins to form about 3 months after fertilization and reaches a
maximum at 8 months after fertilization then declines. The
coconut water is of cytoplasmic origin but in mature coconut
there are no free cells.
As maturity approaches, the fibrous mesocarp begins to dry,
becoming reddish brown. This dehydration and the shrinkage
of the amount of liquid endosperm reduces the nut mass from
3–4 kg at 9 months to 1.5–2 kg at 12 months. For fresh
consumption, the fruit is harvested immature (7–8 months)
when the endosperm has begun to form and is jelly-like
(Tongdee, 1991) before it gradually thickens and hardens. The
coconut water has a maximum of 6% soluble solids, 8–9
months after fertilization. Mature coconuts (10–11 months)
are also used after the hardening of the endosperm. The
endosperm is shredded and squeezed to produce ‘coconut
milk’ and ‘coconut cream’.
Horticulture
PROPAGATION The coconut is only propagated by seed.
Mother palms should be selected on the basis of performance
(a high number of good-sized fruit) over 3 years or longer.
Fruit (seed nuts) are best reaped when the fresh skin colour is
just starting to turn brown and while they contain adequate
liquid and can be heard to ‘splash’ when shaken. Immature
fruit (full of water and very heavy) or over-mature fruit (with
no water and very light) should be rejected. The selected seed
nuts can withstand normal harvesting but careful handling
may avoid damage and consequential loss of viability. Setting
in a nursery should take place immediately since germination
can occur during storage, resulting in twisted and deformed
shoots. Speed of germination is a taxonomic characteristic
(Harries, 1981). Plantlets can also be obtained from excised
embryos after 5–6 months of culturing, avoiding the need to
handle large and heavy nuts (Assy Bah, 1986).
The nuts are normally germinated in nurseries using a
sandy soil, often now in polythene bags (450 ⫻ 450 mm). The
use of polythene bags minimizes root disturbance during
transplanting and reduces ‘transplanting shock’. Initially, the
nuts need to be watered daily to assure uniform germination
and development. Sometimes the epidermis is trimmed from
the germ end of the nut to facilitate water penetration and
germination. The shoot appears after several weeks. There is a
relationship between the speed of germination and vigour, and
the productivity of the palm grown from it, hence the slow
germinating and less vigorous seedlings are culled.
As germination commences, the embryo develops a spongy
haustorium ‘apple’ inside the seed cavity for nutrient
absorption and a shoot that emerges through the soft eye.
When coconuts are processed to copra many may have started
to germinate and, in Sri Lanka and elsewhere, the spongy
haustorium is eaten as a vegetable delicacy. The endosperm is
not completely absorbed in 5 months, having about 60% of
the solid endosperm still present after 9–18 months (Menon
and Pandalai, 1957). Coconut seedlings may therefore have
adequate amount of stored nutrients in the nut till
transplanting but fertilizer application can stimulate growth.
Transplanting can take place soon after the nuts have
sprouted (4–6 months), sometimes up to 9 months. This
should be scheduled so that germination occurs in time to
transplant at the start of the wet season. The cleared land is
planted commonly on a triangular pattern, rows running
north–south, with 6–10 m between plants to obtain optimum
number of palms/ha. A wider spacing is used for the tall
varieties (8–10 m) than the dwarf varieties (6–8 m). A spacing
of 9 m gives 160 palms/ha. Organic manure is frequently
added to the planting hole that is big enough to accept the
plant without significant root disturbance. Mulch may then be
put around the plant after the hole is refilled. If irrigation is
available, the seedling should be watered as soon as possible to
stimulate root development.
The coconut is difficult to vegetatively
propagate. Attempts to generate coconut palms by tissue
culture have achieved only a handful of plants since the 1970s,
despite the fact that coconut water (from the immature fruit)
is a vital ingredient in tissue-culture media for many other
plant species.
ROOTSTOCKS
The coconut palm, unlike the date
palm and the oil palm, is ‘self-pruning’ throughout its life,
dead leaves and over-mature fruit hang for a few months
before falling to leave a clean stem (some exceptions may be
due to growing conditions or unspecified abnormality). Since
all parts of the palm can be used, leaf trimming is sometimes
practised. Trimming can reduce yield.
PRUNING AND TRAINING
During seedling growth,
fertilizer application is recommended, such as 50 g/plant of
equal parts of calcium phosphate, potassium chloride and
magnesium sulphate on two occasions (Fremond et al., 1966).
The lack of uniformity and variability in response of planting
material, the large areas required for trials, difficulty in
recording data on tall palms, seasonal effects and the long
period before fertilizer effects are seen, are considered to be
major difficulties in fertilization experiments. Nitrogen,
potassium and magnesium have been shown to significantly
influence production. The amount of minerals exported in the
harvested product is greatest in the husk: 67% of the
potassium (K) and 85% chloride (Ochs et al., 1993). Leaf
NUTRITION AND FERTILIZATION
Cocos
analysis data have suggested critical levels for the leaflets of the
14th leaf: nitrogen (N) 1.8–2.0% dry mass, phosphorus (P)
0.12%, K 0.8–1.0%, calcium (Ca) 0.3–0.4% and magnesium
(Mg) 0.3% for talls (Magat, 1978, 1979; Rognon, 1987). For
hybrids, 2.2% N, 0.12% P, 1.4% K and 0.2% Mg are
recommended. Chloride has been shown to significantly effect
growth with critical concentration of 0.5–0.6% (Magat et al.,
1988). Nut water analysis is useful for K but not other
nutrients.
HANDLING AND STORAGE Maturity, size,
freedom from blemishes, cracking and free from fibre of
husked coconuts, and wet or mouldy eyes are the main quality
characteristics. Coconut milk is obtained by removing and
grating the hard, white flesh and squeezing out the milky
juice. Young coconuts are harvested 6–9 months after
flowering, as the nut approaches full size and the skin is still
green and the short stem (rachilla) on the top of individual
coconuts that originally held the male flowers (in Thai called
‘rat-tail’) becomes half green and brown. In immature nuts,
the skin surface around the calyx (cap) on the top of coconuts
is creamy-white or a whitish yellow. When the area
surrounding the cap is green the coconut is regarded as
mature and is 10–12 months old. At maturity the skin begins
to change from green to yellow then brown and the ‘rat-tail’ is
entirely brown.
There are no specific grades, but informal grades are
usually based on size and weight. Mature US dehusked
coconuts are sold in 34–36 kg woven plastic or burlap sacks
containing 40–50 coconuts, plastic mesh bags of 12 coconuts
or cartons with 20–25 film-wrapped coconuts, 17–18 kg. After
the husk is removed from immature coconuts in South-east
Asia, they are shaped, dipped in bisulphite, film wrapped, and
sold in single-piece cartons containing 10–16 nuts.
Alternatively, all the husk is removed and then dipped in
sodium bisulphite before packing. Bisulphite is not approved
in the USA for this purpose.
Room cooling is most often used for mature husked nuts,
though the nuts can be forced-air or hydro-cooled. Rapid
temperature changes of 8°C can cause cracking of mature
coconuts. Mature coconuts with husk can be kept at ambient
conditions for 3–5 months before the coconut water has
evaporated, the shell has cracked because of desiccation or
sprouting has occurred. Storage at 0–1.5°C and 75–85%
relative humidity (RH) is possible for up to 60 days for mature
de-husked coconut and 13–16°C and 80–85% RH for 2–4
weeks. Low humidity and high temperature are to be avoided.
Immature nuts have green skins that turn brown after 7 days at
0°C.
Young coconuts are normally held at 3–6°C and 90–95%
RH, and husked-wrapped, shaped fruit can be held for 3–4
weeks. Shaped young coconuts not treated with bisulphite
brown in 12 h, but when treated with 0.5–1.0% sodium metabisulphite the nuts can be held at ambient temperature for 2
days before browning occurs, and if treated with 2% sodium
meta-bisulphite can be held at ambient temperature for 2–7
days. Young coconuts that are not de-husked can be stored for
a longer period (28 days at 17°C) as the husk acts as an
insulator and appears to increase the storage life of young
coconuts.
POSTHARVEST
113
DISEASES, PESTS AND WEEDS Bud rot, basal stem rot and grey
leaf blight can be important diseases (Table A.52). Other
fungal diseases of coconut such as leaf rot (Helminthosporium
halodes) can cause losses (Nambiar and Rawther, 1993). The
viroid diseases caused by single-stranded RNA such as
cadang-cadang (Philippines) and tinangaya (Guam) can be
tested for using molecular probes (Hanold and Randles, 1994),
as can the virus causing foliar decay disease in Vanuatu
(Randles et al., 1986), thus avoiding transmission of infected
material. Lethal yellowing disease is epidemic in parts of the
Caribbean and West Africa (Arellano and Oropeza, 1995) and
related diseases have been identified in Indonesia (Allorerung
et al., 1999). The disease is caused by a phytoplasma spread by
a plant hopper (Jones et al., 1995) and is a major threat of
quarantine concern as it also infects more than 30 other palm
species. The Malayan Dwarf and other varieties from Southeast Asia show high to medium levels of resistance to lethal
yellowing disease, as does the F1 hybrid, Maypan. The Centre
for Information on Coconut Lethal Yellowing (CICLY) at
http://groups.yahoo.com/group/CICLY makes current
information on this disease available online.
Over 700 species of insects have been recorded to associate
with coconut, of which 165 are peculiar to coconut. Only a few
are serious pests (Fremond et al., 1966; Gallego, 1985). Oryctes
spp., especially the rhinoceros beetle (Oryctes rhinoceros),
burrow into the petiole and penetrate the young immature
leaves sometimes causing death of the palm through secondary
bacterial or fungal infection. The palm weevil Rhynchophorus
palmarum is the vector for the red ring disease nematode
(Table A.53) and other Rhynchophorus spp. cause damage by
laying eggs in the petiole and foliage. Scale insects (Aspidiotus
destructor) attack the leaflets and cause loss of palm vigour.
Leaf-eating and leaf-mining beetles (Brontispa and
Promecotheca) and caterpillars of various moths and butterflies
can cause severe damage to leaves if not controlled. Total loss
of fruit production in otherwise healthy palms can be caused
by infestation of fruit sucking bugs, Promecotheca (in Africa)
and Amblypelta spp. (in the Solomon Islands). The coconut
eriophyid mite, Aceria guerreronis, which seriously reduces
fruit size, was first recognized in Mexico in the 1960s and has
subsequently been reported elsewhere in Latin America, all
over the Caribbean and West Africa and is currently active in
India and Sri Lanka. Monkeys, squirrels and rats cause
problems in localized areas to nursery seedlings, young palms
and immature nuts that are chewed open (5-month-old nuts
are particularly preferred) for their water content. Despite its
close association with coconut, in Melanesia and Polynesia, the
robber crab (Birgus latro) is not a pest. It can climb the trunk
of the coconut palm but it does so to escape danger, and not to
feed off the nuts that it, reputedly, cuts off after climbing into
the crown. The real relationship between this land-living crab
and the coconut is to enable its short-lived aquatic larval stage
to disperse long distances to other islands, by floating
(Harries, 1983).
Weed control is essential during plantation establishment.
Failure to control weeds in the 2–4 m area around each palm
can lead to shorter leaf length and fewer new leaves produced
per year, fewer open bunches and nuts set per palm (Romney,
1988). The larger diameter (4 m) circle is more beneficial for
the 39–50-month-old palms.
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Table A.52. Some important diseases and nematodes of coconut (Source: Nambiar and Rawther, 1993; Ploetz, 1994).
Common name
Organism
Parts affected
Distribution
Bud rot and nut fall
Phytophthora palmivora, other Phytophthora spp.
Heart leaves chlorotic, wilted and easily removed
extending to stem apex. Affected nuts shed
All coconut growing areas
Basal stem rot
Ganoderma spp.
Similar to drought; gradual wilting leaf collapse
and drooping; growth slows
All coconut growing areas
Grey leaf blight
Pestalotiopsis palmarum
Small yellow to brown spots on the leaflets and
rachises coalescing into irregular necrotic areas
All coconut growing areas
Stem bleeding
Chalara paradoxa (Thielaviopsis paradoxa)
Infected roots, decay of pith leads to slow palm
All coconut growing areas
decline. Trunk pith decays when infected, reddish
brown exudate from trunk infection site
Cadang-cadang
Viroid (RNA), lethal disease (CCCVd)
Slow infection, nuts become rounded, leaflets
fine non-necrotic translucent yellow spots,
inflorescence becomes necrotic. Related viroid
found outside the Philippines but cadang-cadang
only recognized in the Philippines
Philippines (specific locations)
Foliar decay
Virus (single-stranded; circular DNA)
Progressive yellowing, necrosis and death.
Transmitted by plant hopper to imported
varieties; local varieties are resistant
Vanuatu
Lethal yellowing
Phytoplasma (previously mycoplasma-like
organism (MLO)), plant hopper as vector
First reported from Cuba and Jamaica in 1800s.
Rapid spread, palm death 3–6 months after
first symptoms: nut drop after latent phase,
new inflorescence black. Yellowing of foliage
from the oldest leaves on very susceptible tall
types, brown in more resistant tall and dwarf
types. Bud death, no control, resistant varieties
and hybrids
Caribbean (islands): Cuba,
Cayman, Dominican Republic,
Haiti, Jamaica; (Gulf coast):
Florida, Texas, Mexico, Belize,
Guatemala, Honduras. West
Africa: Cameroon, Ghana,
Nigeria, Togo. East Africa:
Kenya, Mozambique, Tanzania
Red ring disease
Burasaphelenchus cocophilus (obligate parasite
of many palms). The vector is a weevil
Severe insect damage to crown and infects the
palm resulting in leaf yellowing and internal red
ring symptom about 70 cm around the
inoculation point of the stem. Sanitation reduces
vector population
Latin America and Trinidad
Table A.53. Major insect pests of coconut.
Common name
Organism
Parts affected
Country/region
Rhinoceros beetle
Palm weevil
Scale insect
Leaf-eating and leaf-mining beetles
Oryctes spp.
Rhynchophorus spp.
Aspidiotus destructor
Brontispa and Promecotheca
Foliage and growing point
Stem and growing point (vector of red ring)
Foliage
Foliage; serious on seedlings
Moths and butterflies
Locusts and grasshoppers
Coconut fruit bugs
Various
Various
Pseudotheraptus wayii
Amblypelta spp.
Eriophyes guerreronis
Myndus spp.
Foliage and inflorecences
Foliage
Developing fruit
Asia and Pacific; Africa
Asia, Latin America
Widespread
Pacific but now reaching parts of South-east
Asia
Locally important
Locally important
Africa
Solomon Islands
America, Africa, South Asia
America (leaf yellowing), West Africa (blast)
and Vanuatu (foliar decay)
All coconut growing areas
Coconut fruit mite
Plant hopper
Ganoderma spp.
Developing fruit
Vector of lethal yellowing, blast and foliar
decay
Similar to drought; gradual wilting leaf
collapse and drooping; growth slows
This very variable species
has a chromosome number of 2n = 32. The terms variety,
cultivar, ecotype, population are often used interchangeably
but Latinized distinctions between typica for tall cultivars and
nana for dwarf cultivars are less useful than the system of
naming types according to their origin and habit (e.g. ‘Jamaica
Tall’) sometimes with fruit colour (e.g. ‘Malayan Yellow
Dwarf ’). Tall cultivars tend to be slower to begin flowering
MAIN CULTIVARS AND BREEDING
and are generally (but not always) cross-pollinated, while
dwarf cultivars are generally (but not always) self-pollinated
(Table A.54). Dwarf cultivars are more precocious and the first
fruit may be at or close to ground level – hence the
terminology ‘dwarf ’. Although they are never as high or as
vigorous as tall cultivars, they can reach 30 m high in 60 years.
The colours green, red and yellow seen in the petioles and
fruit serve to differentiate between blocks of self-pollinated
Cocos
115
Table A.54. The characteristics of tall and dwarf types of coconut and example of populations.
Characteristics
Dwarf type
Tall type
Height
Leaf production per year
Juvenile period
Pollination
Fruit
Disease resistance
Undesirable
8m
16–18
1.5–3 years
Self-pollinate
Smaller
Some resistance to lethal yellowing
Periodicity in bearing
Less adaptable
Weaker leaf and bunch attachment
Smaller nuts
Lower copra production
More water
Green Catigan-Davao Philippines
Tacunan-Davao Philippines
Aromatic Thailand
Yellow Malayan-Malaysia/Ivory Coast
Red Malayan-Malaysia/Ivory Coast
Cameron-Cameron, West Africa
25 m
About 12
6–8 years
Out-crossing
Large
Susceptible
Height for harvesting
Populations
dwarf populations but are less obvious in cross-pollinated tall
populations that tend to be mixtures of green- and bronzecoloured individuals. However, colours help to identify offtypes when hybrids are produced between yellow dwarf seed
parents and tall pollen parents. Commercially, dwarf cultivars
are considered to be less productive, more sensitive to poor
conditions and have lower quality copra. Some dwarf cultivars
are more resistant to serious virus and phytoplasma diseases
than tall cultivars. Morphological markers, disease, isozyme
and molecular markers are becoming available (Ashburner,
1995b; Ashburner et al., 1997; Teulat et al., 2000).
Of the population types, one of the most interesting for
fresh or green coconut use is the macapuno type. Whereas the
mature normal endosperm is hard and compact around the
periphery of the cavity, in the macapuno type it is soft and
curd-like, filling the entire cavity (Nunez and de Paz, 1990).
The endosperm cells of macapuno are large and multinucleate,
having low intercellular adhesion (Sebastian et al., 1987) and
have a higher cytokinin activity than non-macapuno nuts.
Macapuno endosperm has reduced hemicellulose content in
the cell walls and a different cell-wall sugar composition. A
type similar to the Philippine macapuno has been reported in
India as ‘Thairu thengai’ (Menon and Pandalai, 1957), in Sri
Lanka as ‘Dikiri’, in Thailand as ‘Maphrao Kathi’, in
Indonesia as ‘Korpyor’ and ‘Dua Dac Ruot’ in Vietnam. The
macapuno character is probably a single recessive gene hence
the homozygous palms need to be grown in isolation to ensure
100% yield of macapuno. Dwarf types have been selected
from this normally tall type. However, the nuts do not sprout,
although the embryo, once removed from the nut and washed
free of endosperm, can be grown aseptically in vitro. Another
variant is a fragrant cultivar, ‘Nam Hom’, a dwarf type that
smells like pandanus leaf, that has been produced in Thailand
and is now used for export. The water in ‘Nam Hom’ is not as
sweet as another Thai dwarf variety ‘Nam Wan’ that has
6.5–7% total soluble solids at the green stage. In Sri Lanka, a
cultivar ‘Tembili’ (King Coconut) has sweeter water and is
grown for drinking only.
Laguna-Philippine
Macapuno-Philippine
Tahiti-Tahiti
West African Tall-Ivory Coast
Rennel-Solomon Islands
Increased yield of copra per unit area, was the aim of most
breeding programmes (Santos, 1986), except where diseases or
selection for the fresh market criteria take precedence
(Harries, 1990b). Yield increase can be achieved by increasing
the number of nuts per palm, rather than the amount of copra
per nut, earliness of bearing (possibly linked to the annual rate
of leaf production), palm vigour and disease resistance.
Approaches have generally involved germplasm collection and
evaluation, selection and progeny testing, crossing and
evaluation in different growing regions for adaptability. The
rate of improvement is limited by the long juvenile phase, low
rate of plant multiplication, difficulty of making controlled
crosses with some types, large seed size and its poor
storability, high outcrossing of tall types leading to highly
heterozygous offspring and the diverse environments used for
coconut production. Successful hybrids between dwarf and
tall types include ‘Maypan’ (‘Malayan Dwarf ’ ⫻ ‘Panama
Tall’), ‘Mawa’ (‘Malayan Yellow Dwarf ’ ⫻ ‘West African
Tall’) and ‘Maren’ (‘Malayan Red Dwarf ’ ⫻ ‘Rennel Tall’).
‘Maypan’ was produced for planting in areas subjected to
lethal yellowing disease (Harries and Romney, 1974) and, like
‘Mawa’, achieves higher yields by early bearing and increasing
the number of nuts (Ooi and Chew, 1985; Bourdeix et al.,
1990) and, like ‘Maren’, has satisfactory fruit size (Illingworth,
1991).
Hugh C. Harries and Robert E. Paull
Literature cited and further reading
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through button shedding and immature nut fall. Ceylon Coconut
Planters Review 6, 97–106.
Allorerung, D., Harries, H.C., Jones, P. and Warokka, S. (eds) (1999)
Proceedings of the Workshop on Lethal Diseases of Coconut Caused by
Phytoplasma and their Importance in Southeast Asia. Asian and
Pacific Coconut Community (APCC), Jakarta, Indonesia.
Anon. (1980) Coconut (niu) Uses and Recipes. Na Lima Kokua Pacific
Tropical Botanical Garden, Lawai, Hawaii.
Arellano, J. and Oropeza, C. (1995) Lethal yellowing. In: Oropeza,
116
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C., Howard, F.W. and Ashburner, G.R. (eds) Lethal Yellowing:
Research and Practical Aspects. Kluwer Academic Publishers,
Dordrecht, the Netherlands, pp. 1–15.
Ashburner, G.R. (1995a) Reproductive biology of coconut palms. In:
Oropeza, C., Howard, F.W. and Ashburner, G.R. (eds) Lethal
Yellowing: Research and Practical Aspects. Kluwer Academic
Publishers, Dordrecht, the Netherlands, pp. 111–121.
Ashburner, G.R. (1995b) Genetic markers for coconut palms. In:
Oropeza, C., Howard, F.W. and Ashburner, G.R. (eds) Lethal
Yellowing: Research and Practical Aspects. Kluwer Academic
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Investigations on the shedding of buttons in the coconut (Cocos
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others (Colombia, Ivory Coast, Nigeria, Papua New Guinea)
the remainder. As the oil palm is a very productive (3–5
t/ha/year of oil) and profitable crop, its areas are expected to
increase further, particularly in Indonesia and Colombia.
Uses and nutritional composition
Elaeis guineensis
oil palm
The oil palm, Elaeis guineensis Jacq. (Arecaceae), is an
economically important crop to many third world countries in
the humid tropics. It is the highest yielding and a highly
profitable oil crop and is relatively easy to grow by large
plantations and small farmers alike. It is the leading export oil
crop providing a major source of external revenue to a number
of countries and its versatile uses spawn a host of domestic
industries. Oil palm cultivation has most of the elements of an
environmentally sustainable crop suitable for tropical
countries: mature oil palms replace the climax vegetation lost
in degraded forests; use of a leguminous cover crop in the
immature phase protects and replenishes organic matter and
nutrients in the soil; efficient recycling of the crop waste (e.g.
pruned fronds, bunch stalks, mill-waste fibres, sludge, shells)
to the fields; and contribution towards rural-community
development. The industry is well supported by very
organized and sophisticated research and development and
marketing programmes.
Historical origins
The African oil palm, E. guineensis, the oil palm of commerce,
is indigenous to tropical Africa, concentrated in the rainforest
areas of the West and Central African countries of Angola,
Cameroon, Congo, Ghana, Ivory Coast, Nigeria and Zaire.
The species occurs as semi-wild groves usually close to
settlements of indigenous peoples as the oil palm fruit is an
important dietary source of fat and vitamins to these people.
Crude palm oil obtained from the pulp or mesocarp of the
fruit is the primary internationally traded commodity. Oil palm
plantations were started in South-east Asia by the Europeans
very early in the 20th century to obtain a substitute for animal
fat in the production of wax, candles and margarine. They
started with seedlings that originated from the four thickshelled dura palms in Bogor Botanic Gardens, Java. The four
palms presumably originated from the same fruit bunch in
Africa and came to Indonesia via Amsterdam and Mauritius.
With increasing demand for the oil as a food as well as other
non-food uses and the availability of improved cultivars,
especially the thin-shelled tenera hybrid, oil palm plantations
have expanded tremendously since 1911 and continue to do so.
World production and yield
In terms of the world’s total oils and fats production of about
121 million t, palm oil (21%) occupies second place next to
soybean oil (25%). This will soon change. Palm oil, with 19.1
million t, worth about US$7.5 billion annually, dominates
(47%) the international edible oil trade (Oil World, 2002). It is
produced from about 7.4 million ha of plantations. Malaysia
accounts for 48% of the production, Indonesia 35%, and
The palm fruit produces palm oil from the mesocarp and
kernel oil from the nut. Typically an oil palm fruit bunch will
produce 20–25% palm oil and 3–5% kernel oil.
Crude palm oil, the main commodity, obtained from the
sterilized mesocarp is refined, bleached and deodorized to give
refined palm oil. Refined palm oil can be used directly or
blended with other oils as cooking oil, salad oil, shortenings,
margarines and spreads. Refined palm oil can be fractionated
to give olein (liquid) and stearin (semi-solid) and with further
fractionation gives fatty acids and alcohols, intermediate
commodities traded and used in food and oleochemical (e.g.
detergent, lubricant, plastics, pharmaceuticals, cosmetics)
industries. Stearic acid is a cocoa butter substitute. Palm oil or
as methyl ester (palm diesel) can be used as biofuel. However,
80% of palm oil is used as food but its other uses are
increasing (Fig. A.15). Palm kernel oil competes with coconut
oil in the food and oleochemical uses.
Prompted by health and environmental concerns besides
profit motive, added value products from secondary and waste
products from the palm oil industry are also gaining
importance. Carotene in crude palm oil has been processed
into a vitamim A supplement and a food dye. Tocols extracted
from palm oil have been encapsulated and marketed as palm
vitamin E. Sterols, squalene, co-enzyme Q and phospholipids,
components of palm oil, have applications in pharmaceutical,
nutraceutical, food and cosmetic industries. Kernel cake and
sludge cake from the mill waste have use as animal feed while
the bunch-fibre waste can be used directly in the plantation as
organic mulch or processed into compost, or burnt together
with the nut shells to provide power to the mill and the
plantation residential community. Surplus power can be
supplied to the national grid.
Palm oil contains c.50% saturated fatty acid and 50%
unsaturated fatty acid (Table A.55) but some doctors,
nutritionists and traders have equated it with coconut oil as
saturated oils from tropical palms. Consumption of high
amounts of such oils would predispose the individual to heart
disease. Proponents of palm oil counter-argue that the
saturated component is mainly palmitic acid that does not
cause a rise in blood cholesterol. It has also a large component
of monounsaturates (oleic acid, 40%) and substantial amounts
of polyunsaturates (linoleic acid, 10%). Palm oil also contains
antioxidants (carotene, tocols) which, besides conferring deepfrying stability, have anti-cancer and anti-atherosclerosis
properties. It does not require hydrogenation in the
production of margarines. Hydrogenation leads to the
production of the trans fatty acids.
Nevertheless, improvement in the degree of unsaturation of
palm oil is being sought through breeding and genetic
transformation means. The target value is an iodine value of
70 (c. 70% unsaturation) to put it on a competitive basis with
olive oil. Besides appeasing consumer health concerns, a more
liquid oil from unsaturation allows it to penetrate the salad-
119
Elaeis
FRESH FRUIT BUNCHES
MILL
PROCESS
Fruit residues
KERNEL
CRUDE PALM OIL
Crushing
Extraction
Refining (R)
Bleaching (B)
Deodorizing (D)
Palm kernel meal
Other uses
being researched
Fractionation
and refining
Technical uses
Soap etc.
Palm kernel oil
(PKO)
Blending
Fuel
RBD
Palm oil
RBD Olein
RBD Stearin
Splitting
Animal
feed
Fractionation
and refining
Soaps
Stearin
Olein
Refining
Hydrogenation
(H)
H. PKO
H. PK. Olein
Confectionery
Fats
Margarines
Shortenings
Vanaspati
Frying fats
Ice cream
Splitting
Margarine
Confectioneries
Filled milk
Ice cream
Biscuit cream
Frying
Cooking
Shortenings
Margarines
Shortenings
Margarines
Vanaspati
Fatty
acids
Soaps
2nd
Fractionation
Soaps
Food emulsifiers
etc.
Margarines
Confectioneries
Coffee whitener
Filled milk
Coating fats
Fatty Acid
Glycerol
Fatty
Alcohols
Amines
Amides
Emulsifiers
Humectants
Explosives
Double
fractionated
olein
Palm midfraction
Blending
Cocoa butter
equivalent
Fig. A.15. Uses of palm oil (Source: Pantzaris, 1997).
Table A.55. Variability of fatty acid composition in oil palm populations (Source: Arasu 1985; Rajanaidu, 1990; Rajanaidu et al., 2000).
Content (%)
Fatty acid
Nigerian Elaeis
guineensis
E. guineensis
IRHOb
E. guineensis
Elaeis oleifera
E. oleifera ⫻
E. guineensis
0.3–3.1
37.4–46.6
3.8–14.7
33.0–55.9
5.4–15.8
43.8–69.8
0.9–1.5
41.8–6.8
4.2–5.1
37.3–40.8
9.1–11.0
51.0–55.3
0.3–1.6
34.7–50.1
3.1–8.8
32.0–46.0
10.0–16.0
–
0.1–0.3
14.4–24.2
0.6–2.2
55.8–67.0
6.0–22.5
67.4–91.9
0.1–0.5
22.4–44.7
1.4–4.9
36.9–60.1
8.3–16.8
–
C14:0 (myristic)
C16:0 (palmitic)
C18:0 (stearic)
C18:1 (oleic)
C18:2 (linoleic)
Iodine value
a
b
PORIMa
PORIM, Palm Oil Research Institute of Malaysia.
IRHO, Institut de Recherche pour les Huiles et Oleaginuex.
and cooking-oil markets of the temperate countries. Genetic
variability for unsaturated fatty acid and other oil quality traits
are deficient in advanced oil palm breeding populations.
Botany
The genus Elaeis belongs
to the Palmae or Arecaceae family and subfamily Cocoideae.
Jacquin in 1763 first described the African oil palm, E.
guineensis Jacq. The American oil palm, Elaeis oleifera Cortes,
which has a slower-growing procumbent habit with leaflets in
TAXONOMY AND NOMENCLATURE
one plane and different fruit traits, was known earlier as Elaeis
melanococca and Corozo oleifera. Both species have 2n = 32
chromosomes and hybridize readily but the hybrids exhibit
varying degrees of sterility. The third species, Elaeis
madagascariensis, is thought to be a variant of E. guineensis.
There are also named vegetative and fruit variant forms of E.
guineensis: leaf form – idolatrica (fused leaflets); fruit form –
mantled/poissoni (fleshy mantle), dura (thick shell), tenera
(thin shell), pisifera (shell-less); immature fruit colour –
nigrescens (black/dark purple), virescens (green) and
albescens (white).
120
Arecaceae
The oil palm tree is mainly grown from seed,
which takes about 100–120 days to germinate in nature after
accumulating sufficient heat and moisture to break the
dormancy. In commercial seed production, controlled seed
germination is achieved by heating (37–39°C) the seeds at
18–20% moisture for 40–60 days and then setting for
germination at 22–23% moisture under ambient conditions.
The oil palm has only one terminal bud. Stem growth in
the first 3 years in the field is mainly in base enlargement. It
can grow to a height of 25–30 m but in commercial fields it is
replanted when it becomes too tall (12–15 m) for harvesting.
The oil palm stem anatomy, typical of monocotyledons,
consists of the cortex formed by the extension of the leaf
bases; the pericycle comprising numerous vascular bundles
with fibrous phloem sheaths embedded in sclerotic ground
tissue, providing mechanical support for the stem; and the
central core of less dense vascular bundles embedded in
parenchyma ground tissue. The apical bud or meristem
produces the leaf, which comprises the petiole, rachis and
leaflets. The petiole is 1.3–2.5 m long, triangular in crosssection with spiny upper edges. The rachis in a mature palm
measures 5.5–7.0 m long bearing 300–400 leaflets. The leaflets
vary in length according to their position along the rachis with
the middle being the longest at 1.2 m. Leaflet width ranges
from 4 to 6 cm. The leaves are arranged spirally, seen as two
sets of eight and 13 leaves running in opposite directions. A
well-grown mature oil palm has 41–50 leaves, each having a
dry weight of 4.5–5.5 kg and a leaf area of about 10 m2.
The oil palm has a typical fibrous root system The
hemispherical meristematic bole (c.80 cm diameter) grows
downwards 40–50 cm into the soil. About 6000–10,000
primary roots extend > 20 m horizontally from it. Secondary
roots branch out at right angles, upwards and downwards from
the primary roots followed by tertiary roots from the
secondary and quaternary from the tertiary, decreasing in size
and length accordingly. The structure of the different roots is
essentially similar with an outer epidermis, lignified
hypodermis surrounding a wide cortex with large lacunae for
root respiration and non-lignified root tips. The primary roots
presumably have palm anchorage and conduit functions while
the others are for water and nutrient absorption. Total root
length may exceed 60 km.
The oil palm is monoecious bearing male and female
flowers in separate clusters (Plate 15A, B). The inflorescence is
a raceme with flowers occurring on spikelets arranged on a
large spadix enveloped by two fibrous spathes. The male
inflorescence has a stalk about 40 cm long bearing 100–300
spikelets, each 10–30 cm long. Each spikelet holds 400–1500
sessile, male flowers with six stamens each having bilobed
anthers. A male inflorescence produces 10–50 g of pollen with
viability lasting 3–8 days. The female inflorescence with a
shorter and stouter stalk carries around 150 fibrous spikelets
bearing five to 30 flowers each. The female flower has a
protective spiny bract, two floral bracts and two whorls of
three perianths each. At receptivity, the trifid stigma curves
outward to receive the pollen. The stigmatic lobes are creamy
white with sticky glandular central tissue. They turn pink after
the first day, light brown on the second and violet on the third.
The receptive period lasts 2–4 days and after pollination and
fertilization, the stigma turns black and woody. Both male and
DESCRIPTION
female flowers develop and mature from the base of the
spikelets upwards. They emit a characteristic aniseed smell.
Upon fertilization, the female inflorescence develops into an
ovoid fruit bunch bearing between 500 and more than 4000
fruit and weighing 10–50 kg in a fully mature palm. Each fruit
is a sessile drupe, ovoid-oblong shape, 2–5 cm long, about
2 cm wide and weighing 3–30 g (Plate 15D). It contains a
kernel enclosed by an endocarp (shell), an oil rich mesocarp
and a coloured exocarp. In the commonly grown nigrescens
type, the fruit exocarp is black or dark purple turning to
orange red at maturity.
The oil palm flowers continuously
in successive cycles of male and female inflorescences ensuring
cross-pollination. Although the pollen is small and light (22 ⫻
33 m) and can be wind dispersed, in nature, pollination is
mainly by weevils (Elaeidobius kamerunicus) and to a lesser
extent by Thrips hawaiiensis. The insects are attracted to the
flowers by the strong aniseed smell during anthesis/
receptivity in search of food. Between 60 and 70% of the
flowers develop into fruit. Inflorescence initiation commences
at 37–39 months before fruit ripening and sex differentiation
at 19–22 months. Good nutrition, high light intensity or low
moisture deficit favours the development of female
inflorescences while stress conditions favour male
inflorescence production. Female inflorescences may abort a
few months before receptivity and bunch abortion may occur a
few months after pollination under severe water or other stress
conditions.
REPRODUCTIVE BIOLOGY
FRUIT DEVELOPMENT Normally, only one of the three ovules
is fertilized to form a seed within the fruit. The ovary develops
rapidly from 16 to 90 days after pollination when maximum
size is reached and the endosperm (kernel) completely fills the
endocarp (shell). The mesocarp develops simultaneously
attaining its maximum weight at about 130 days after
pollination. The next phase involves the solidification of the
kernel, which takes about 55 days, and the simultaneous oil
synthesis in the kernel and the mesocarp. The latter process
commences at 70 days from pollination, increases
exponentially from 100 days and reaches the peak at 155–188
days. Maximum oil is attained in the mesocarp when at least
6% of the fruit have abscised.
In the typical nigrescens fruit the external ovary wall
changes its colour from white to pale yellow prior to
pollination, the fruit becomes dark pink at 60 days, black at
140 days and then orange-red at ripening. In the less common
virescens type, the colour is greenish before turning orange at
maturity, while in the albescens it remains pale yellow. At this
stage, the fruit start to abscise and oil synthesis stops. Fruit
ripening occurs at 120–200 days after pollination, depending
on environmental conditions and genotype.
ECOLOGY AND CLIMATIC REQUIREMENTS The natural habitat
of oil palm is riverine forests or freshwater swamps. It cannot
thrive in primaeval forests and does not regenerate in tall
secondary forests. It has adapted to diverse ecological zones in
the tropical belt (longitudes c.15°N–15°S), from savannah to
rainforests and highlands (< 1500 m) although it is widely
grown on lowlands. It can grow on most tropical soils with
Elaeis
adequate water supply. The best oil palm growth and yields are
obtained from coastal and riverine alluvium without an acid
sulphate (jarosite) layer within 75 cm, and deep (> 100 cm)
soils of volcanic or sedimentary origin, and acid sands. Soils
with pH 4–6 and slopes of < 20° are suitable. It can tolerate
temporary flooding or a fluctuating water table, as found along
rivers. Waterlogged, shallow lateritic, loamy sand to sand,
stony or woody, peaty soils would confer low yields.
The ideal rainfall is 2000–2500 mm/year without any dry
season (monthly rainfall < 100 mm). However, the oil palm can
withstand annual rainfall between 650 and 4500 mm with 2–4
months of dry period. Severe moisture deficits of
> 200 mm/year may induce male inflorescence and floral and
bunch abortions. The suitable temperature for oil palm is a mean
maximum temperature of 28–34°C and mean minimum
temperature of 21–24°C. Oil palm growth is arrested below
15°C while floral abortion and delayed fruit maturity occur
below 18°C. The oil palm being a C3 plant requires full sunlight
for maximum photosynthesis. The desirable sunshine hours
should exceed 5 h/day or 1825 h/year provided the high
sunshine hours are not accompanied by drought, high water
vapour pressure deficit (> 1.8 kPa) and/or extreme temperatures
(> 38°C). The solar radiation should be at least 12 MJ/m2/day.
Horticulture
The oil palm is an out-breeding species.
Commercial propagation is through hybrid seeds. Typically,
tenera hybrids (thin shell, thick-mesocarp fruited palms) are
obtained by controlled pollination of the thick-shell fruited
dura female parent with the shell-less fruited female-sterile
pisifera male parent (Plate 15C, D). Commercial seeds are
mixtures of hybrids derived from parents which are non-true
inbreds and the parents on each side may or may not be
related. Consequently, considerable genetic variability exists
among commercial palms.
PROPAGATION
MAIN CULTIVARS AND GENETIC IMPROVEMENT The main oil
palm cultivars are designated by the parental populations, for
example ‘Deli’ ⫻ ‘AVROS’, ‘Deli’ ⫻ ‘Yangambi’, ‘Deli’ ⫻ ‘La
Me’. Here is a brief description of the major parental
populations.
● ‘Deli’ – This is the thick-shelled dura variety derived from
the four Bogor palms. Subsequent distribution and
selections in other countries led to the development of subpopulations, for example ‘Ulu Remis’, ‘Elmina’, ‘Serdang’,
‘Dabou’. The ‘Dumpy’ and ‘Gunung Melayu’ are short
variants. The ‘Deli’ is known for its bigger bunch and fruit.
It is the source of dura mother palms in all major
commercial hybrid seed-production programmes.
● ‘AVROS’ – This originated from the Djongo (best) palm in
Zaire’s Eala Botanical Garden and further bred in Sumatra,
Indonesia to give rise to the well-known progenitor, SP540.
Descendants of this palm have been widely distributed to
become the source of pisifera parents for major seed
producers worldwide, for example Colombia, Costa Rica,
Indonesia, Malaysia, Papua New Guinea. ‘AVROS’
pisiferas are noted for their high oil-yielding and vigorous
growth-conferring attributes.
121
● ‘Yangambi’ – This originated from the same source as the
Djongo palm and thus has similar attributes to ‘AVROS’.
● ‘La Me’ – This resulted from the breeding programme of
CIRAD (Centre de Cooperation Internationale en
Recherche Agronomique pour le Developpement) from the
‘Bret 10’ palm in La Me, Ivory Coast. Descendents of the
progenitor, ‘L2T’, are the pisifera parents of major seedproduction programmes in Ivory Coast and Indonesia. ‘La
Me’ progenies typically exhibit high bunch number,
smaller bunch and fruit and smaller stature.
Other known but less widely used pisifera sources are
‘Binga’ (Palm Bg 312/3), derived from ‘Yangambi’, ‘Ekona’
(Palm Cam. 2/2311) from Cameroon and ‘Calabar’ (Palm NF
32.3005) from Nigeria.
Genetic improvement objectives High oil yield is the primary
objective of genetic improvement. Improvement in oil content
and smaller sized, high harvest-indexed palms for higher
density plantings are aimed towards this. Ease of harvesting is
important in labour scarce countries such as Malaysia. Short
palms with long-stalked fruit bunches having non-abscising
ripe fruit with a distinct colour change (e.g. virescens) will
facilitate harvesting. Improvement in oil quality, particularly
unsaturation, has received much attention in Asian
programmes. High carotene oil palms are also being sought.
These quality traits are in the domains of the Oleifera genome
and wild Guineensis accessions and have to be introgressed into
advanced Guineensis materials by backcross breeding or by
genetic engineering because of hydrid sterility and the poor
agronomic traits from the donor species.
With the development of the genetically more uniform
hybrids and clones, breeding for adaptability and tolerance to
abiotic (e.g. drought, mineral deficiency) and biotic (e.g.
disease and pest) stresses would assume importance. Fusarium
wilt resistance and drought tolerance are important objectives
in African and American programmes. Breeding for
Ganoderma (basal stem rot) resistance appears to become
mandatory in many Asian programmes. Tolerance to
magnesium deficiency is an important breeding objective in
Indonesia and Papua New Guinea.
Breeding The shell trait exhibits monogenic inheritance. The
thick-shelled dura is homozygous dominant, the shell-less
pisifera is homozygous recessive and the thin-shelled tenera is
the heterozygote exhibiting incomplete dominance. In
breeding, ‘Deli’ dura mother palms are regenerated (100%)
from dura ⫻ dura crosses of selected parents. The pisiferas
being female sterile are regenerated from tenera ⫻ tenera and
tenera ⫻ pisifera crosses of selected parents. The former cross
gives 25% dura, 50% tenera and 25% pisifera progenies,
while the latter cross gives 50% tenera and 50% pisifera
progenies.
Oil palm hybrid seeds are obtained from recurrent selection
progammes. In the modified recurrent method, the dura
parents are selected based on the individual’s performance for
commercial seed production and for further breeding. The
female sterile pisiferas are selected as male parents for
commercial seed production based on their dura ⫻ pisifera
progeny tests. Pisifera improvement is effected using
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phenotypically selected teneras. In the other modified reciprocal
recurrent selection method, dura ⫻ tenera progeny tests are
done instead. Selfs of the dura and tenera parents are planted at
the same time as the progeny test. Based on the progeny test,
the superior progenies are reproduced as commercial dura ⫻
pisifera hybrids using the parental selfs. Further breeding
involves the progeny tested dura and tenera parents. This
scheme produces more genetically uniform hybrids.
In vitro propagation Commercial mixed hybrids exhibit
considerable variability in the field with individuals varying
more than 30% from the group mean. The differences are not
entirely genetic. Nevertheless, it stimulated the successful
development of the in vitro propagation technique in the oil
palm that cannot be vegetatively propagated and had been
considered a recalcitrant species for in vitro propagation. In
vitro propagation of oil palm is achieved via somatic
embryogenesis from callus cultures using young leaf explants
(Plate 15E). Large-scale commercial clonal propagation has
been hampered by the spectre of somaclonal variation,
manifested as mantled parthenocarpic fruit leading to fruit
bunch abortion and consequent partial or full sterility (Plate
15F); the inefficiency of the cloning process; and the
inefficiency of palm selection for cloning requiring rigorous
clonal field tests. With the demonstration of the feasibility
(improved cloning efficiency, low somaclonal variation) of
recloning field-tested clones and the liquid suspension culture
technique, large-scale commercial propagation of proven
clones appears assured.
Cloning the parents of superior hybrids to produce biclonal
(both clonal parents) or semi-clonal (one clonal parent) hybrid
seeds is an alternative approach albeit with lower genetic
improvement expected.
Genetic modification Transgenic oil palms carrying the Basta
resistance gene obtained through the biolistics approach are at
the field-testing stage. High oleic transgenic oil palm are in the
pipeline. The production of polyhydroxy butyrate (PHB) by
oil palm for biodegradable thermo-plastic production is a
recent research initiative. There is a host of other possibilities.
Transgenic palms can be commercially propagated as clones or
used as breeding parents for hybrid seed production.
The development of molecular markers (e.g. restriction
fragment length polymorphism (RFLP), amplified fragment
length polymorphism (AFLP), microsatellites) has allowed
breeding parents, families and populations to be accurately
segregated pedigree-wise for population genetic study, breeding strategy and intellectual property ownership applications.
Molecular markers linked to quantitative trait loci of desirable
traits (e.g. shell thickness, virescens, embryogenesis) are being
sought to develop marker-assisted selection to improve
breeding efficiency.
Stringent field-testing protocols are being drawn up to
ascertain the agronomic, environmental and consumer health
acceptability of genetically modified (GM) oil palm.
CULTIVATION Land preparation This operation involves
land clearing, drainage of wet areas, terracing on hills and
construction of a network of roads for transport. The existing
vegetation may be secondary logged jungle or a perennial tree
crop such as rubber, cocoa or oil palm. Careful timing of land
preparation is important to ensure the field is ready for
planting at the onset of the rainy season.
Clearing Burning is the traditional method of land clearing
for oil palm cultivation. The environmentally friendly zeroburn technique is now widely adopted in both new clearing
and replanting of oil palm. The existing vegetation is usually
mechanically cleared and stacked in windrows parallel to the
planting rows. In an oil palm replant, the old stand is
mechanically felled and the trunk chipped and stacked (using
an excavator fitted with a sharpened-lipped bucket) to
accelerate decomposition. Where pests (e.g. termites) and
diseases (e.g. Ganoderma spp.) are prevalent, burning may be
allowed by the relevant authorities. To control the Oryctes
rhinoceros beetle, the standing palm or chipped palm residues
are increasingly being mechanically pulverized to destroy
potential beetle breeding sites.
Drainage and terracing Drainage is often required for the
very productive flat coastal and alluvial plains to ensure a wellaerated rooting zone and to remove excess water during the
wet season. The water table should ideally be > 60 cm below
the soil surface.
Terracing is not necessary on rolling terrain of < 10° slopes,
although conservation terraces spaced 25–30 m apart are
important. On slopes > 10°, contour terracing is critical not
only for soil and water conservation but also to provide access
for field upkeep, fertilization and harvesting. Terraces are
usually 3–4 m wide and 10 m apart with a back slope of 3–5°.
Palms are planted along the terraces at varying distances apart
(depending on the slope) to achieve a density close to 138–149
palms/ha. Wide terraces allow mechanization of fieldwork and
must be incorporated in the land preparation process.
Layout of road system A well-planned road system is
fundamental to efficient crop production and an essential
component of land preparation. On flat to undulating terrain,
field roads are usually spaced at 200 m apart with connecting
roads at every 1 km. On hilly terrain roads should ideally have
< 8° slopes and run diagonally across the slope. Road density
depends on terrain and typically about 100–150 m/ha is
required. Requirements will be reduced with mechanized infield collection of harvested bunches.
Planting materials Tenera hybrid seedlings are
currently being planted in commercial fields. Commercial
plantings of clones are constrained by the high cost and
limited supply of proven clones. Seeds are usually obtained
only from specialist seed producers to ensure legitimate and
proven hybrids. Typically young oil palm seedlings are raised
in a nursery for 12 months before field planting. Optimum
conditions in the nursery ensure only uniform, vigorous and
healthy seedlings are planted out in the field.
NURSERY
Prenursery In a two-stage nursery, germinated seedlings are
first planted in small polyethylene bags or seedbeds filled with
sandy loam or sandy clay loam soil for about 4 months before
transplanting into large poly bags in the main nursery for
Elaeis
123
another 8 months. Prenursery allows for more efficient
management and use of space and water. A round of culling is
done before transplanting into the main nursery. Shade is not
essential but is often provided in the form of overhead nettings
or cut palm fronds. Shade is reduced progressively and
removed completely before the third month.
competition and pests and disease attacks at this age. Therefore
the best possible care and growing conditions are given to these
palms. Poorly established retarded palms and runts are replaced
within the first 2 years to avoid over-shading by surrounding
palms. Small farmers may plant a cash crop (e.g. pineapple,
banana, papayas, melon) in the inter-rows during this period.
Main nursery The prenursery seedlings are transplanted into
large poly bags spaced at 1 ⫻ 1 m in a triangular pattern in the
main nursery. Germinated seeds may also be planted directly
into the large poly bags as a single-stage nursery. Single-stage
nurseries require more labour, supervision, space and water
but seedlings have been reported to grow faster by about 2
months. Routine nursery practices of watering, fertilizer
application and pest and disease control are carried out. Only
the best uniformly grown seedlings are transplanted into the
field.
MATURE PHASE Under favourable growing conditions, the oil
palm begins to produce flowers and fruit by 15 months after
field planting and regular harvesting commences at 24–30
months. Fresh fruit bunch (FFB) yield increases rapidly over
the first 4 years of production reaching a plateau by 8–10 years
after planting. During peak production, yields of > 30
t/ha/year are common for well-managed plantings in good
growing areas. The economic lifespan of an oil palm planting
ranges from 20 to 30 years depending on growing conditions.
Fields are replanted when the palms become too tall for
economic harvesting. Social and other economic considerations also affect replanting decisions.
Planting density All commercial plantings
adopt an equilateral triangular spacing ranging from 9 to 9.8 m
giving a planting density of 149–120 palms/ha for maximum
light capture by the single terminal palm canopies. The
optimum density depends on the terrain, soils, growing
conditions and genotype. On favourable soils and flat to gently
rolling terrain, a density of 138 palms/ha is optimum. On
more marginal conditions (e.g. peat soils), where palms are
smaller and inter-palm competition is reduced, higher density
planting (up to 160 palms/ha) is preferred to maximize yield
per hectare.
FIELD PLANTING
Planting Planting is usually timed to coincide with the
beginning of the wet season. Planting holes are usually dug
with a tractor-mounted auger and 0.25–0.5 kg of phosphate
rock is applied into the hole before the seedling is planted
manually.
Establishment of leguminous cover crop Due to the long period
(> 3 years) between land clearing and full palm-canopy
covering, a creeping leguminous cover crop is planted
immediately after land preparation to reduce soil erosion,
suppress weed competition and improve soil fertility through
nitrogen fixation, organic matter enrichment and nutrient
transfer and recycling. This improves soil aeration, rooting
and water conservation. A mixture of Pureria phaseoloides,
Calapogonium muconoides and Calapogonium caeruleum
leguminous cover-crop seeds is commonly sown at a rate of
5–10 kg/ha. A small starter dose of compound fertilizer is
applied at 1–3 weeks after germination and three split dustings
of phosphate rock totalling 600–1000 kg/ha given at 1, 3 and 5
months after germination. Mucuna cochinchinesis is gaining
popularity due to its fast-growing habit enabling rapid ground
cover and weed suppression and it is also shade tolerant.
The immature phase from planting to the
start of regular harvesting varies between 24 and 36 months or
more depending on growing conditions and management
considerations. This is a crucial period in that large, fast-growing
and vigorous palms are not only more precocious but are also
higher yielding. The palms are also most susceptible to weed
IMMATURE PHASE
AGRONOMIC PRACTICES Weeding Weeds compete with the
oil palm for water, nutrients and light and can suppress crop
growth and production. Thick weeds also hinder access for
field operations. The preferred method of weed management
in immature plantings is to have a thick leguminous cover to
smother the weeds in the inter-rows. Common weeds in
Malaysia and Indonesia are: Imperata cylindrica (lallang grass),
Mikania sp., Clidemia sp. and Chromolaena odorata (Siam
weed). From planting, a clean-weeded palm circle of
1.5–2.0 m diameter and increasing to 4 m as the palms mature
is maintained primarily to prevent weed competition. Manual
circle weeding is preferred during the first year but the use of
herbicides, at the risk of contaminating the young palm, is now
common practice due to labour constraints. Young palms are
sensitive to systemic and hormonal herbicides (e.g. glyphosate,
diuron, 2,4-D amine) which can cause canopy twisting or
spear snapping or parthenocarpic fruit development. The
recent innovative use of a perforated 2.4 ⫻ 3 m ultravioletresistant plastic mulch over the palm circle after planting
reduces the need for frequent, manual weeding in the first 6
months and minimizes chemical scorching of the lower leaves
in the first 2 years. The whole first year’s requirement of
compound fertilizers is broadcast onto the palm circle before
the plastic mulch is laid down thus saving labour costs for up
to eight fertilizer applications otherwise needed.
For mature palms, clean-weeded palm circles are maintained
to reduce weed competition and to facilitate loose fruit
collection. Inter-row and path weeding is equally important to
suppress the growth of competitive weeds, to provide easy
access for routine upkeep and harvesting operations as well as
to maintain a non-competitive ground cover for soil, moisture
and nutrient conservation. Soft grasses (e.g. Paspulum sp.,
Axonopus compressus) and ferns (e.g. Nephrolepis sp.) are thus
retained. Systemic herbicides applied with medium- and lowvolume sprayers are the preferred method for weed control.
Spraying intervals are aimed to break the seed-production
cycles of the weeds. Biological control of weeds on a research
scale has been attempted, for example the use of the leaf-eating
caterpillar Pareuchaetes pseudoinsulata against Chromolaena in
Indonesia and Malaysia.
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Major pests and diseases The oil palm being a tropical
perennial crop provides a continuous supply of food, host
medium and suitable growth environment for a range of pests
and diseases. Routine pest management is thus needed
throughout the crop’s life. Integrated pest management is
generally advocated. This involves crop monitoring for
infestations/infections and their prevention from exceeding
economic thresholds by employing various cultural, physical,
chemical and biological control measures.
Insect pests commonly reported to cause economic damage
in Malaysia and Indonesia are the leaf-eating caterpillars, for
example bagworms, Metisa plana, Pteroma pendula and
Mahasena corbetti; and nettle caterpillars, Setora nitens, Darna
sp. and Setothosea sp. They can cause severe defoliation
resulting in significant yield losses. The planting of nectarproducing plants such as Cassia cobanensis and Euphorbia
heterophylla to enhance the population of natural enemies of
leaf-eating caterpillars as a preventive measure has been
attempted in Malaysia. The potential of mass production of
microbial pathogens of these leaf-eating caterpillars for
commercial applications is also being explored. The current
method of control used in outbreaks is trunk injection of
systemic insecticides (e.g. methamidophos and monocrotophos) and spraying with chemicals with short residual
effects (e.g. cypermethrin). In recent years, rhinoceros beetles
(Oryctes rhinoceros) have become an important pest of
replanted palms in Malaysia, Indonesia and India. Adult
beetles burrow into the soft meristem tissues of the palms to
feed, disfiguring the developing foliage and this may lead to
the consequent death of the palms. Fortnightly spraying with
cypermethrin is the current control method. The beetles
breed in the decaying palm biomass so experiments are being
conducted using cultural control measures to speed up
decomposition of the old palm biomass to deprive the beetles
of suitable breeding sites. Biological control programmes
involving trapping adult beetles with pheromone baits
followed by infection with a pathogenic virus before being
released, and augmentative release of the pathogenic fungus
Metarhizium sp. into the old palm residues have been
reported.
Coptotermes curvignathus termites cause severe losses of
young palms planted on peat in Malaysia and Indonesia. They
bore into the meristem tissue in the crown to feed, causing
death within 6–12 months after infestation. A current
common control measure is to drench the crown and soil
around the base of the infested palms with the insecticide
Fipronil. The use of baits containing sublethal doses of
insecticides that could be carried to the nests by the workers to
destroy the reproductive queens has been reported.
Rats (Rattus tiomanicus, Rattus diardii and Rattus argentiventer) are a chronic pest of oil palm that require recurrent
control measures. In young palms, rats feed on the frond
bases, male flowers and fruit. In mature palms they feed on the
mesocarp and kernel of the developing fruit causing 5–10%
losses in oil yield. The common method for control is the use
of anticoagulant-based baits. This is sometimes augmented
with biological control using barn owls (Tyto alba).
Jungle fringe plantings are prone to damaging attacks by
larger mammalian pests such as elephants, wild boar,
porcupines and monkeys.
Serious arthropod pests in South America include the
Tetranchus sp. spider mite, and the lepidopterans Leptopharsa
gibbicarina and Sagalassa valida, the former two attacking the
leaves while the latter attacks the roots.
Most reported diseases of oil palms are caused by fungi,
however, some diseases of economic importance have implicated viruses (e.g. ring spot disease in India), bacteria (e.g.
Erwinia sp. in little leaf disease in Zaire) and nematodes
(Bursaphelenchus cocophilus in red ring disease in South
America). In most cases, management of these diseases is
through cultural control by excluding the disease organisms
from areas where it is not known to occur, destroying diseased
palms and control of the associated vectors of these diseases.
In addition, providing the palms with optimum growth
conditions and balanced nutrition would maximize vigour and
minimize infection incidences.
Basal stem rot (Ganoderma sp.) is an important disease of oil
palm in most countries where it is grown. In young palms, the
external symptoms normally comprise one-sided yellowing or
mottling of lower leaves followed by necrosis and retarded
growth. In mature palms, common symptoms are the presence
of multiple unopened leaf spears and pale canopy. Tissues of
the infested stem give a characteristic dry rot and the palm
collapses with severe rotting. Basidiomata may be produced on
the dead palm tissues. Disease incidence increases with age of
the palms with reported incidence as high as 67% at 15 years
resulting in a 46% reduction in yield. Infected young oil
palms normally die within 6–24 months after the first
appearance of symptoms, but mature palms can take up to 2–3
years to die. Cultural measures to exclude and destroy the
inoculum of the disease is the current method of control.
Infected palms are felled and chipped into small slices and
spread out thinly to hasten decomposition, and the soil around
the bole is dug up to expose the infected roots. The
inoculation of oil palm seedlings with vesicular-arbuscular
mycorrhizal fungi to prevent infection of the palms in the field
is being explored. Screening for disease-tolerant genotypes is
also being pursued. Ganoderma disease of palms is the subject
of an international cooperative research effort coordinated by
CABI.
Vascular wilt caused by Fusarium oxysporum elaedis is a
serious disease of oil palm in West Africa and has been
introduced into South America. It is a soil-borne disease
attacking young and older palms. In infected older palms the
older leaves desiccate and the rachis break at some distance
from the base, the disease moves up the spiral leaving
reduced-sized chlorotic green leaves at the crown and the
palm may remain in this state for a while before collapsing. In
young palms, the symptom of ‘lemon frond’ appears at the
upper mid-crown before drying and this then progresses to
the younger leaves and results in the subsequent death of the
palm. Tolerant genotypes are available; a nursery screening
technique has been developed and is routinely used in
breeding programmes in West Africa.
Lethal bud rot hampers oil palm expansion in South
America. Its pathogenic basis is in contention although some
resistant cultivars have been claimed. Sudden wither or
marchitez sorpressiva used to be a serious disease of oil palm
in South America but has been brought under control with
eradication of infected palms. It is caused by the flagellate
125
Elaeis
280
K
240
Nutrient (kg/ha)
Phytomonas sp. which is transmitted by Lincus sp. of the
Pentamidae. Blast is an important nursery-seedling disease in
West Africa. The causal agent is unknown but a vector (Recilia
mica, Jessidae) has been implicated.
Crown disease occurs sporadically on 2–4-year-old palms
with characteristic symptoms of bending young leaves. Palms
generally recover readily from the disease with no apparent
defect but severe and extended symptoms retard early
development and reduce yield. Disease susceptibility is known
to be under the control of a recessive gene and a modifier gene
and can be bred out. The ‘Deli’ material is particularly
susceptible.
200
160
120
40
0
Pruning Inflorescences and fruit bunches occur at the frond
axils. The subtending leaf of a ripe bunch is usually removed
at harvesting. However, not every leaf will carry a fruit bunch
and regular pruning is necessary to remove excess leaves. For
tall palms, excess leaves trap loose fruit and obstruct visibility
of ripe bunches for harvesting. In early palm maturity pruning
twice a year is needed but when fully mature, an annual
pruning round suffices. Over-pruning, especially during early
maturity, is detrimental due to reduction of leaf area. A
general guide is to maintain a full canopy with at least two
leaves below the oldest bunch for young palms (< 8 years) and
one leaf for older palms. Harvesters usually carry out pruning
during low-cropping periods, using the same tools as they use
for harvesting. Pruned leaves are spread over the inter-rows as
organic mulch and are very effective for soil conservation.
Soil and water conservation Soil erosion can be very serious
especially when hilly terrains are cleared for planting. Despite
the high annual rainfall in the tropics, periodic water stress is
not unusual due to uneven distribution of rainfall and high
evapotranspiration demand. The need for effective soil and
water conservation is well recognized and emphasized in wellmanaged commercial oil palm plantations where terrace
plantings, cover-crop planting and mulching are used as
discussed earlier.
Nutrition and fertilization Nutrient demand is very high for a
high-yielding stand of oil palm. This is due to its potential for
very rapid vegetative growth and the precocious and very
rapidly ascending yield pattern from the onset of maturity to
peak yields in years 8–10 after planting. Thus, large quantities
of nutrients particularly nitrogen (N) and potassium (K) are
immobilized each year in the vegetative tissue and exported in
the harvested crop (Fig. A.16). The annual uptake of N and K
for a stand of mature palms yielding 25 t/ha/year FFB is
estimated at 193 kg/ha/year N and 251 kg/ha/year K,
respectively. As the oil palm is mainly grown on highly
weathered acid soils in the tropics, adequate and balanced
fertilization is essential to realize and sustain the palms’ high
genetic growth and yield potential. Fertilizer rates may vary
from 110 to 185 kg/ha/year N and 185–300 kg/ha/year K
depending on soil fertility and actual yield levels. Other
important nutrients are phosphorus (P) and magnesium (Mg)
and these are applied at rates of 50–60 kg/ha/year P and
30–40 kg/ha/year Mg. Among the micronutrients, boron (B)
is most important especially on sandy soils. Applications of
copper (Cu) and zinc (Zn) are often required with peat
N
80
0
1
2
3
4
5
6
7
8
9
10
Palm age (years)
Fig. A.16. Estimated annual nitrogen (N) and potassium (K) uptake
(kg/ha) in oil palm (Source: Ng, 1977).
plantings. Application rates are usually 100–200 g/palm of
borate and copper sulphate or zinc sulphate, respectively.
Consequently, the costs of fertilizers and application
constitute the single largest component of production cost (>
60% of field cost, 20% of total cost) in oil palm plantations.
Understandably, fertilizer usage is a major focus of most R&D
and advisory programmes.
The system of fertilizer recommendations and management
for oil palm is well developed based on a vast database of
experimental results and practical experience. Palm nutrient
requirements are estimated based on the nutrient balance
approach where nutrient inputs from the soil, recycled in palm
residues and mill by-products and from the rain are balanced
against nutrients exported through the crop, immobilized in
the tissues and lost via run-off, leaching or soil immobilization
(Fig. A.17). Any shortfall is augmented as applied fertilizers.
Leaf analysis is usually carried out annually and soil analysis
every 3–5 years to assess and monitor the palm and soil
nutrient status, respectively.
Based on the above nutrient balance approach, an
integrated site-specific fertilizer recommendation system
(INFERS) has been developed for the diagnosis of fertilizer
requirement in Malaysia and has been subsequently expanded
into an oil palm agronomic recommendation system called
ADEPTTM. This system includes an empirical site yield
potential model (ASYP), to estimate the yield potential for
each unique site; INFERS, to provide a balanced nutrition for
DEMAND
Nutrients immobilized
in palm tissues
Nutrient losses,
e.g. via leaching
SUPPLY
Atmospheric return
Nutrient recycled
?
Soil nutrients
Nutrient for growth
and production
Fig. A.17. Components of nutrient balance in oil palm.
Fertilizers
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Arecaceae
optimum growth and yield; and a ‘Best month’ expert system
to optimize timing of fertilizer application. A Global
Positioning System (GPS) and Geographical Information
System (GIS) have also been incorporated in the ADEPTTM
to facilitate implementation, display and communication.
These developments are geared towards precision agriculture
for more accurate fertilizer recommendations to sustain high
growth and production.
Ablation and sanitation Only in less favourable
growing conditions is ablation (removal of first inflorescences)
carried out 20–24 months after planting, to promote vegetative growth and root vigour. Otherwise, ablation is unnecessary. Usually a round of sanitation pruning (removal of dried,
dead leaves, old flowers and rotten early bunches) is carried
out about 6 months prior to regular harvesting.
HARVESTING
Ripeness standard During ripening (15–20 days) carbohydrates accumulated in the fruit are converted into oil. When
the fruit has fully ripened and the oil content is at its
maximum, it loosens and detaches from the bunch. Once
detached, the free fatty acid (FFA) content increases rapidly
due to breakdown of glycerides by the microbial lipase
enzyme.
Harvesting of under-ripe bunches would result in lower oil
content, while harvesting over-ripe bunches would result in
more loose fruit lost and high FFA levels in the oil. The
optimum ripeness standard is in between these two extremes.
A practical guide is five to ten loose fruit per bunch for
<8–10-year-old palms and one to five loose fruit for older
palms when harvesting is done at 10–14 day intervals.
Harvesting interval Fruit bunches are produced throughout
the year although there is a distinct peak and trough in the
annual cycle. Harvesting of ripe bunches is therefore a yearround activity done at intervals of 8–14 days. Harvesting at
intervals of 7 days or less is usually not economic especially
during the trough period. Conversely, harvesting intervals
exceeding 15 days will result in an excessive number of loose
fruit, incurring higher labour costs in loose-fruit collection.
Any ripe bunches inadvertently missed will become seriously
over ripe or rotten by the next harvest. Harvesting intervals of
10 days are recommended but may be extended to 15 days
during peak yield cycles.
Harvesting At the start of production, the young palms
produce many, relatively small (2–5 kg) bunches. As the palms
mature, fewer but bigger bunches (up to 50 kg or more) are
produced. Harvesting is done manually by skilled harvesters.
Young palms are harvested using a long chisel with a broad,
sharp cutting edge. As palms get taller with age, this is
replaced by a sickle attached to a light but strong bamboo or
extensible aluminum pole (up to > 12 m). Prototypes of
mechanical harvesting cutters and machines are being tested.
The harvested bunches and loose fruit are collected and
deposited at specific points along collection roads.
Traditionally hand-baskets, wheelbarrows and buffalo-drawn
sledges are used. Terrain permitting, mechanical in-field
collection is done using a mini-tractor fitted with a mechanical
grabber and high-lift trailer which then dumps the crop onto
bins or trucks for transport to the palm oil mill for processing
within 24 hours.
PROCESSING Upon arrival at the palm oil mill the harvested
bunches are sterilized by pressurized steam (about 40 psig) in
an enclosed chamber to facilitate the stripping of fruit from
the bunch and to stop the enzymatic conversion of oil into
FFA. The sterilized FFB are then mechanically threshed to
strip off the fruit. The empty fruit stalks/bunches (EFB;
about 22% of FFB, containing 0.72% N, 0.08% P, 2.64% K,
0.27% Ca and 0.12% Mg on a dry-matter basis) are returned
to the field as fertilizer/soil ameliorant. The stripped fruit are
then sent to a mechanical digester, which pulps and heats the
mesocarp to rupture the oil-bearing cells. The resultant mash
is transferred to a screw press to extract the mesocarp oil
(crude palm oil or CPO; about 20% of FFB). The CPO is
then screened, purified and vacuum dried and stored in a
specially constructed storage tank for onward shipment to the
buyers (refiners). CPO buyers would usually specify that it
should contain < 5% FFA, and moisture and dirt content
< 0.25%. The nuts are separated from the screw press cake,
cracked and the kernel (about 6% of FFB weight) extracted.
The oil and kernel extraction rates (OER and KER,
respectively) are based on the amounts of oil and kernels
processed from the FFB. The kernel contains about 50% oil,
mainly lauric oil similar to coconut oil. Extraction of kernel oil
is carried out in a separate mill usually outside the plantation.
PRECISION AGRICULTURE AND TECHNOLOGICAL INNOVATIONS
Precision or site-specific agriculture implies the concept of
using information about variability in site, climate and palms to
manage distinct units within a field with best practices for
optimum profitability, sustainability and protection of the
environment. Traditionally, oil palm plantations are divided
into c.40 ha management units. Although soil and terrain are
usually taken into consideration in delineating blocks, it is often
not possible to achieve a satisfactory degree of uniformity
within the blocks for uniform application of inputs.
With recent advances and affordability in GPS, GIS and
Variable Rate Application (VRA) technologies, the reduction in
size of the management units to that of the size of a harvesting
task of about 1 ha for yield monitoring and application of
fertilizer and other inputs is now feasible. GPS/GIS/Remote
Sensing technologies have enabled more accurate mapping of
field boundaries (roads, waterways, terrain) and coupled with
recent innovative technological tools (e.g. information
technology, computers, hand-held organizers, sensors), which
can be linked to mechanized field operations, would allow the
plantation management to be more precise and efficient. Roads,
drains and terraces would be planned more efficiently. It may
be possible in the near future that yield records can be captured
digitally and accurately, yield maps drawn and poor yielding
fields delineated, to be followed by remedial agronomic inputs
(fertilizers, pesticides) perhaps using VRA technology. These
concepts are currently being tested. It is envisioned that the
future plantation manager will eventually become a knowledge
worker with ready access to the case history and potential
of each of their management units and to the latest
knowledge and tools to achieve the best productivity from their
plantation.
Elaeis
Concluding remarks
As per capita intake of edible oil is still low in many countries
and palm oil is the cheapest, most profitable and versatile oil, oil
palm plantations are likely to expand further in developing
countries with suitable climates, particularly in South-east Asia
and South America. In countries with a mature oil palm industry
such as Malaysia with increasing costs of production, the
challenge is to maintain sustainability, profit-wise and
environmentally. This will have to be met through increased
investments in research for new palm oil uses, improved
cultivars and precision farming.
Acknowledgements
We thank our company, Applied Agricultural Research Sdn.
Bhd. and its principals, Boustead Holdings Berhad and Kuala
Lumpur Kepong Berhad for permission to publish this entry.
A.C. Soh, K.K. Kee, K.J. Goh, B.N. Ang and L.H. Ooi
Literature and further reading
Arasu, N.T. (1985) Genetic variation for fatty acid composition in the
oil palm (Elaeis guineensis Jacq). Thesis, University of
Birmingham, UK.
Armstong, D.L. (1999) Oil Palm Nutrition Management. Better Crops
International, Volume 13 No.1, Special Edition. Potash and
Phosphate Institute, Saskatoon, 54 pp.
Chew, P.S. (1998) Prospects for precision plantation practices in oil
palm. The Planter, The Incorporated Society of Planters, Kuala
Lumpur 74, 661–684.
Corley, R.H.V. and Tinker, P.B. (2003) The Oil Palm. Blackwell
Science, Oxford.
Flood, J., Bridge, P.D. and Holderness, M. (2000) Ganoderma Diseases
of Perennial Crops. CAB International, Wallingford, UK.
Hartley, C.W.S. (1988) The Oil Palm, Elaeis guineensis Jacq., 3rd edn.
Tropical Agriculture Series, Longman Scientific and Technical,
UK, 761 pp.
Ho, C.T. (1998) Safe and efficient management systems for plantation
pests and diseases. The Planter 74, 369–385.
Kee, K.K., Goh, K.J., Chew, P.S. and Tey, S.H. (1994) An Integrated
Site Specific Fertilizer Recommendation System (INFERS) for
high productivity in mature oil palms. In: Chee, K.H. (ed.)
Management for Enhanced Profitability in Plantations. The
Incorporated Society of Planters, Kuala Lumpur, pp. 83–100.
Khoo, K.C., Ooi, P.A.C. and Ho, C.T. (1991) Crop Pest and their
Management in Malaysia. Tropical Press, Kuala Lumpur,
Malaysia, 242 pp.
Kok, T.F., Goh, K.J., Chew, P.S., Gan, H.H., Heng, Y.C., Tey, S.H.
and Kee, K.K. (2000) Advances in oil palm agronomic
recommendations. In: Pushparajah, E. (ed.) Plantation Tree Crops
in the New Millennium: the Way Ahead (Vol. 1). The Incorporated
Society of Planters, Kuala Lumpur, pp. 215–232.
Mariau, D., Desmier de Chenon, R. and Sudharto, P.S. (1991) Oil
palm insect pest and their enemies in Southeast Asia. Oleagineux
46 (11) (Special Issue). Centre de Cooperation Internationale en
Recherche Agronomique pour le Developpement (CIRAD),
Montpellier, 476 pp.
Ng, S.K. (1977) Review of oil palm nutrition and manuring: scope for
greater economy in fertilizer usage. In: Earp, D.A. and Newall, W.
127
(eds) International Developments in Oil Palm. The Incorporated
Society of Planters, Kuala Lumpur, Malaysia, pp. 209–233.
Oil World (2002) Statistics Update. Available at: http://www.
oilworld.biz (accessed 13 December 2002).
Ooi, L.H. and Tey, S.H. (1998) Applications of global positioning
system and geographical information system in oil palm estates.
In: Towards Improving Productivity Through Mechanization.
Preprint of National Seminar on Mechanization in Oil Palm
Plantations. Palm Oil Research Institute of Malaysia, Kuala
Lumpur.
Pantzaris, T.P. (1997) Pocketbook of Palm Oil Uses. Palm Oil Research
Institute of Malaysia, Kuala Lumpur, 163 pp.
Rajanaidu, N. (1990) Major developments in oil palm (Elaeis
guineensis) breeding. In: Proceedings of 12th Plenary Meeting of Oil
and Fat. Hamburg, Germany, pp. 37–57.
Rajanaidu, N., Kushari, A., Rafii, M., Mohd Din, A., Maizura, T. and
Jalani, B.S. (2000) Oil palm breeding and genetic resources. In:
Yusof, B., Jalani, B.S. and Chan, K.W. (eds) Advances in Oil Palm
Research. Malaysian Palm Oil Board, Kuala Lumpur, Malaysia,
pp. 171–224.
Singh, G., Lim, K.H., Teo, L. and Lee, D.K. (1999) Oil Palm and
The Environment. A Malaysian Perspective. Malaysian Oil Palm
Growers’ Council, Kuala Lumpur, 277 pp.
Soh, A.C., Wong, G., Hor, T.Y., Tan, C.C. and Chew, P.S. (2003) Oil
palm genetic improvement. In: Janick, J. (ed.) Plant Breeding
Reviews. John Wiley and Sons, New Jersey, pp. 165–219.
Tan, K.S. (1983) The Botany of Oil Palm. Casual Papers on Oil Palm.
Incorporated Society of Planters, Kuala Lumpur, Malaysia, 32 pp.
Turner, P.D. (1981) Oil Palm Diseases and Disorders. Oxford
University Press, Kuala Lumpur, Malaysia, 280 pp.
Wong, G., Tan, C.C. and Soh, A.C. (1997) Large scale propagation of
oil palm clones: experience to date. Acta Horticulturae 447,
649–658.
Yusof, B., Jalani, B.S. and Chan, K.W. (2000) Advances in Oil Palm
Research. Malaysian Palm Oil Board, Kuala Lumpur, 1526 pp.
Elaeis oleifera
corozo
Corozo or American oil palm, Elaeis oleifera (H.B.K.) Cortes.
(Arecaceae), is the only member of this genus of two species
found natively in the Western hemisphere. It is used as an oil
source to some extent throughout its range.
World production and yield
Rarely cultivated, the American oil palm is primarily an
extractive resource from wild stands, and there is very little
production data available. Fruit range from 8.5 to 12.8 g in
weight and clusters rarely more than 22 kg. Annual yields
average about 25 kg of fresh fruit, which equals nearly 13,000
individual drupes (Balick, 1979a, b). About 75–85% of the oil
contained in the fruit mesocarp is recovered. Though the oil
extracted from the fruit is higher quality than that of Elaeis
guineensis, the American oil palm is much less productive.
Uses and nutritional composition
The fruit contains 30–50% oil, the seed (kernel) only slightly
less. The fruit oil is nearly 50% saturated fat (33% palmitic),
but it also contains a sizable fraction of unsaturated fatty acids
of which 50% is oleic and 12% linoleic (Duke, 2001).
128
Arecaceae
Botany
The American oil palm is
the only other species in the genus Elaeis other than the
African oil palm, E. guineensis. Synonyms include Alfonsia
oleifera H.B.K., Corozo oleifera (H.B.K.) Bailey and Elaeis
melanococca Gaertn. emend. Bailey.
TAXONOMY AND NOMENCLATURE
The American oil palm is a fairly small, often
procumbent single-stemmed palm. The stem is 1–6 m long and
up to 0.5 m in diameter. Typically only the upper 1–3 m of the
trunk is erect; the proximal portions lie on the soil surface,
rooting from their lower surface. The arching, feather-like leaves
are several metres long and number 30–40 with 35–90 tworanked leaflets per side, without the basal swelling found in E.
guineensis. The petiole is spiny. Male and female inflorescences
are produced on the same plant, often sequentially. The flower
stem is a short-stalked, congested cluster of 100–200, sharppointed spike-like branches up to 16 cm long. The flowers are
sunken into their surface. The (usually) one-seeded, ellipsoid
fruit are crowded into rounded, conical clusters and ripen from
yellow to red. Each fruit is 2–3 cm long. Many of the fruit are
reportedly parthenocarpic.
DESCRIPTION
ECOLOGY AND CLIMATIC REQUIREMENTS Elaeis oleifera
ranges through Central America and northern Colombia at
low elevation, with scattered occurrences in the Amazon basin,
perhaps the result of human introduction. It is usually found
in low-lying, moist soils along watercourses. While sometimes
observed in drier forests, between 1700 and 2200 mm of
annual rainfall is considered optimal (Duke, 2001). Soils with
a pH of 4–6 are suitable for cultivation, and the palm has no
tolerance of freezing temperatures.
Bees are thought to be the primary
pollinators of American oil palm, with some wind pollination
as well. While the palms primarily produce separate staminate
and pistillate inflorescences, bisexual inflorescences are
sometimes observed.
REPRODUCTIVE BIOLOGY
FRUIT DEVELOPMENT Fruit ripen from January to June, the
clusters held fairly close to the ground. Typically, five clusters
of ripe fruit are produced each year. Palms are generally in
excess of 10 years of age before production begins. Fruit ripen
in 6 months after pollination.
Horticulture
PROPAGATION On the rare occasions when the American oil
palm is deliberately cultivated, seeds are sown directly into
germination beds and seedlings transplanted into a field site in
1–1.5 years. Seed germination begins after several months,
and can continue erratically for over a year. Seedlings can also
be container grown.
Applications of ammonium
sulphate and potassium sulphate at a rate of 225 g/palm are
recommended shortly after planting in the field (Duke, 2001).
On magnesium-deficient soils, 227 g magnesium sulphate or
slowly soluble kieserite should be added. These amounts can
be increased to 450 g/palm over the next 4 years.
NUTRITION AND FERTILIZATION
Young seedlings need to be
kept well irrigated and shaded from too much direct sun. The
seedlings are slow growing, and 7 years may transpire before a
normal-sized leaf is formed. Cover crops and inter-cropping
with other food plants is often practised.
TRAINING AND MAINTENANCE
DISEASES, PESTS AND WEEDS
Many of the same pest and
disease organisms that attack African oil palm in the Americas
can also affect the American oil palm (see entry on E.
guineensis for specific information).
MAIN CULTIVARS AND BREEDING The American oil palm has
been bred with the far more economically important African
species in order to increase the oil quality of the more
productive E. guineenesis. Some of these hybrids have then
been reproduced clonally via tissue culture. Alan W. Meerow
Literature cited and further reading
Balick, M.J. (1979a) Amazonian oil palms of promise: a survey.
Economic Botany 33, 11–28.
Balick, M.J. (1979b) Economic botany of the Guajibo, I. Palmae.
Economic Botany 33, 361–376.
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Euterpe oleracea
assaí
Assaí, Euterpe oleracea Mart. (Arecaceae), is a slow-growing,
clustering palm of the Amazonian rainforest. Assaí grows on
indundated soils of the Amazon basin and similar areas of
northern South America, often close to the coast. The fruit,
and especially the liquid extracted from the pulpy mesocarp, is
an important food resource in tropical America.
Related species with similar uses include Euterpe precatoria,
known as assaí de terra firma, a single-stemmed (solitary) palm,
growing to a maximum of 20–22 m in height. The crown shaft
is composed of 14–19 leaves, each reaching 3.5–4.5 m in
length. A large number of pendent leaflets confer a unique,
ornamental appearance to this palm. The stem and root system
are similar to those of E. oleracea. The inflorescences are larger
than those of E. oleracea, bearing a larger number of rachillae
(70–76) and flowers. The flowers are lighter in colour, usually a
pale yellowish pink (male) and light brown (female). The fruit
are globose, 1.0–1.8 cm in diameter, dark purple when mature,
with a thin (0.5–1.5 mm thick) and juicy mesocarp from which
a beverage very similar to açaí or or vinho de açaí is prepared.
Euterpe precatoria occurs almost exclusively in non-inundated
forests. Its floral biology and reproductive system are similar to
those of E. oleracea.
World production and yield
Few data are available on production of assaí, which occurs
only within the area of the palm’s natural range, since it is
chiefly harvested from wild populations. About 4000 ha of
exploited assaí forest on Marajó Island in Brazil produced 7 t
of pure and sweetened assaí pulp for export to the USA in
2000. One stem on average produces four to eight fruiting
stems annually, each one yielding approximately 4 kg of fruit.
Euterpe
Thus, one stem can provide 16–32 kg of fruit, with a mean of
24 kg/year (Bovi and de Castro, 1993).
Uses and nutritional composition
The assaí palm is an extremely important indigenous resource
for people in areas bordering the Amazon estuary (Strudwick
and Sobel, 1988). It has been the focus of considerable
research directed towards its commercial exploitation
(Calzavara, 1972; Anderson, 1988; Brondízio, 2002). People
harvest the fruit by climbing the palms, cutting the
inflorescence and extracting the fruit pulp mechanically or by
hand. The nutrient content is reportedly as follows:
1.25–4.34% (dry weight) protein; 7.6–11.0% fats; 1–25%
sugar; 0.050% calcium; 0.033% phosphorous; and 0.0009%
iron (Mota, 1946; Campos, 1951; Altman, 1956). It also has
some sulphur, traces of vitamin B1 and some vitamin A.
Calorific content ranges from 88 to 265 kcal/100 g, depending
upon the concentration. It is processed into beverages, ice
cream and pastries and is sold at local or regional markets, and
also by the local businesses that process the fruit (known in
Brazil as as açaílandias). Mixed with cassava flour or rice, it is
consumed in great quantity (2 l/day/capita) by the population
of the lower Amazon River. The flavour has been described as
nutty with a metallic aftertaste. The texture is creamy to
slightly oily. Details of assaí processing, consumption, and
marketing can be found in Strudwick and Sobel (1988). Assaí
liquid, locally called açaí or vinho de açaí (though not
alcoholic), is highly perishable and this factor has restricted its
export. However, the dehydrated liquid can be kept for 115
days after preparation (Melo et al., 1988), and in this form, as
well as frozen pulp, it has been imported into the USA and
Europe in recent years, where it is sold in health food stores
and used an ingredient in so-called ‘energy’ beverages. The
inner stems (palm heart) are also extracted and eaten locally, as
well as canned for export.
Botany
TAXONOMY AND NOMENCLATURE Euterpe consists of seven
species (Henderson and Galeano, 1996) of monoecious
pinnate-leafed, solitary or clustering palms ranging from
Central America to northern South America and Trindad in
the West Indies. Assaí is also known by the common names
murrap, naidi (Colombia), pinot (French Guiana), manaka
(Surinam) and manac (Trinidad).
DESCRIPTION Euterpe oleracea produces several stems
reaching up to 20 m in height. The slender trunks are grey at
maturity and 7–20 cm in diameter. A skirt of roots is usually
visible at the base of the stems, and in swampy conditions,
upward-growing pneumatophores may be formed. The crown
consists of eight to 14 leaves, each 2–4 m long. The leaf bases
are tightly sheathing and form an attractive, smooth crown
shaft that can be green or variously yellow, red or purple. The
50–62 widely spaced leaflets are pendulous in orientation on
the rachis, and can reach nearly 1 m in length. The manybranched flower stems reach about 1 m in length, emerging
from below the crown shaft. The inflorescence is composed of
a central stiff rachis with an average of 54 lateral branches
(rachillae), each of which bears clusters of two lateral
129
staminate (male) flowers and one central pistillate (female)
flower, except at the terminus where only staminate flowers
occur. The flowers are unstalked (sessile). The purplish male
flowers are 4.5 ⫻ 2.7 mm; the purple to light brown female
flowers are 3.2 ⫻ 2.6 mm. The ultimate branches are densely
covered with light brownish-white hairs. The fruit is a 2 cm
diameter drupe that is purple at maturity (some assaí
populations that have mature green fruit are known locally as
white assaí). The mesocarp is sweet and pulpy.
ECOLOGY AND CLIMATIC REQUIREMENTS Assaí can form
extensive stands in swampy forest and along river courses in
rainforest. In the Amazon estuary, it may occur in huge,
monospecific populations. It is intolerant of dry situations and
prefers an organic acidic soil, high humidity and warmth. It is
most adaptable to humid, tropical climates where
temperatures rarely drop below 10°C.
Euterpe oleracea is monoecious;
each inflorescence produces numerous, small, sessile staminate
and pistillate flowers. Staminate flowers mature before the
pistillate flowers, thus promoting outbreeding. However, a
variable amount of self-pollination can occur depending on
the synchronization between inflorescences in the same or
different stems. The pollinators are predominantly small bees
and flies, as well as beetles. The palms can achieve reproductive maturity in as few as 4 years under excellent
conditions. Individuals growing under the forest canopy take
longer to start flowering. Assaí palms flower year-round, but
drought may induce inflorescence abortion (Jardim and
Anderson, 1987). Seed dispersal over short distances is by
rodents. Long-distance seed dispersal is accomplished by
birds (Zimmermamm, 1991) and passively by water.
REPRODUCTIVE BIOLOGY
FRUIT DEVELOPMENT The maximum number of fruit
clusters per plant is about eight, although three or four is
typical. On any stem, there will usually be infloresecences and
fruit at all different stages of development, from flower stems
enclosed in the bracts to clusters of ripe fruit. Fruiting occurs
throughout the year, but the peak period of production is the
dry season, July–December.
Horticulture
Only recently has there been interest in
developing production plantations of E. oleracea (Nogueira et
al., 1995). Slow initial growth and considerable mortality of
seedlings have been the main problems to its successful
establishment in the field (Bovi et al., 1987).
Assaí seeds are recalcitrant and sensitive to both
dehydration and low temperature during storage (Araujo and
de Silva, 1994). Refrigeration will kill the seeds. Seeds are best
air-dried for several days to 1 week after removal from the
fruit, then stored in plastic bags at room temperature, but for
as short a time period as possible. Seed stored for 15 days had
33.3% moisture content and 79% germination (Araujo and de
Silva, 1994). After 2 months of storage germination was
reduced to 28%, and to 8% after 7 months. Seed usually
germinates in 4–8 weeks, but germination may continue for
nearly a year (Bovi and de Castro, 1993).
PROPAGATION
130
Arecaceae
Chup (1999) found beneficial effects on growth of assaí by
inoculating seedlings in containers with arbuscular mycorrhizal
fungi. Ledo et al. (2002) reported successful somatic
embryogenesis from E. oleracea, which may have utility in the
rapid propagation of selected clones in the future.
The most productive assaí
stands are located on moist, organic, acid soils. No data on
mineral nutrition and fertilization are available.
NUTRITION AND FERTILIZATION
DISEASES, PESTS AND WEEDS
No major pests or diseases of
assaí have been reported. Zorzenon and Bergmann (1995)
described fruit and seed predation on E. oleracea by Xyleborus
ferrugineus (Coleoptera: Scolytidae). Seed germination was
reduced by 80%.
No cultivars of assaí palm
have been developed. Though no formal breeding effort has
been carried out on the species, interspecific hybrids between
E. oleracea and Euterpe edulis have been produced in Brazil
(Bovi et al., 1987).
Alan W. Meerow
MAIN CULTIVARS AND BREEDING
Literature cited and further reading
Altman, R.F.A. (1956) O caroco de açaí (Euterpe oleracea Mart.).
Boletim Técnico do Instituto Agronômico do Norte (IAN) 31,
109–111.
Anderson, A.B. (1988) Use and management of native forests
dominated by acai palm (Euterpe oleracea) in the Amazon estuary.
Advances in Economic Botany 6, 144–154.
Araujo, E.F. and Silva, R.F. de (1994) Evaluation of quality in acai
seeds stored in different packages and environmental conditions.
Revista Brasileira de Sementes 16, 76–79.
Bovi, M.L.A. and Castro, A. de (1993) Assaí. In: Clay, J.W. and
Clement, C.R. (eds) Selected Species and Strategies to Enhance
Income Generation from Amazonian Forests. Food and Agriculture
Organization (FAO) Miscellaneous Working Paper 93/6. FAO,
Rome. Available at: http://www.fao.org/docrep/v0784e/v0784e0b.
htm#assaí (accessed 4 December 2006).
Bovi, M.L.A., Godoy, G. Jr and Saes, L.A. (1987) Híbridos
interespecíficos de palmiteiro (E. oleracea ⫻ E. edulis). Bragantia
46, 343–363.
Bovi, M.L.A., Spiering, S.H. and Melo, T.M. (1989) Temperaturas e
substratos para germinação de sementes de palmiteiro e açaízeiro.
In: Anais do Segundo Congresso sobre Tecnologia de Sementes
Florestais. Atibaia, São Paulo, Brazil, p. 43.
Brondízio, E.S. (2002) The urban market of açaí fruit (Euterpe
oleracea Mart.) and rural land use change: ethnographic insights
into the role of price and land tenure constraining agricultural
choices in the Amazon estuary. Urban Ecosystems 6, 67–97.
Calzavara, B.B.G. (1972) As Possibilidades do Acaizeiro no Estuario
Amazonico. Boletim da Fundacao de Ciencias Agrarias do Para,
São Paulo, Brazil.
Campos, F.A.M. (1951) Valor nutritivo de frutos brasileiros. Instituto
de Nutrição, Trabalhos e Pesquisas 6, 72–75.
Chup, E.Y. (1999) The effects of arbuscular mycorrhizal fungi
inoculation on Euterpe oleracea Mart. (açaí) seedlings. Pesquisa
Agropectura Brasiliera, Brasilia 34, 1019–1024.
Henderson, A. and Galeano, G. (1996) A revision of Euterpe,
Prestoea, and Neonicholsonia (Palmae). Flora Neotropica
Monograph 72, 1–90.
Jardim, M.A.G. and Anderson, A.B. (1987) Manejo de populaçães
nativas do açaízeiro (Euterpe oleracea Mart.) no estuário
Amazônico: resultados preliminares. Boletim Pesquisas as Florestais
15, 1–19.
Ledo, A. da Silva, Lameira, O.A., Benbadis, A.K., Menezes, I.C. de,
Oliveira, M.S.P. de and Filho, S.M. (2002) Somatic
embryogenesis from zygotic embryos of Euterpe oleracea Mart.
Revista Brasileira de Fruticultura 24, 601–603.
Melo, C.F.M., Barbosa, W.C. and Alves, S.M. (1988) Obtenção de
açaí desidratado. Boletim Pesquisa 92, 1–13.
Mota, S. (1946) Pesquisas sobre o valor alimentar do açaí. Anais
Associação Química Brasileira 5, 35–38.
Nogueira, O.L., Carvalho, C.J.R. de, Ller, C.H.M., Galvão, E.U.P.,
Silva, H.M. de, Rodrigues, J.E.L.F, Oliveira, M.S.P. de, Carvalho,
J.E.U. de, Rocha Neto, O.G. da, Nascimento, W.M.O. do and
Calzavara, B.B.G. (1995) A Cultura do Açaí. Coleção Plantar 26,
EMBRAPA-SPI, Brasilia, Brazil.
Strudwick, J. and Sobel, G.L. (1988) Uses of Euterpe oleracea Mart.
in the Amazon estuary, Brazil. Advances in Economic Botany 6,
225–253.
Zimmermamm, C.E. (1991) A dispersão do palmiteiro por
paseriformes. Rev. Ciencia Hoje 12 (72), 18–19.
Zorzenon, F.J. and Bergmann, E.C. (1995) Occurrence of Xyleborus
ferrugineus (Fabricius, 1801) (Coleoptera: Scolytidae) in fruits and
seeds of two species of the genus Euterpe. Revista de Agricultura
Piracicaba 70, 17–20.
Hyphaene thebaica
doum palm
Doum palm, Hyphaene thebaica (L.) Mart. (Arecaceae), is an
African palm distributed from western India through the
Middle East and south to tropical Africa. It is most common
in coastal East Africa and in Eritrea. Though never cultivated
as far as is known, people have used the fruit and seeds for
centuries throughout its range. Other species in the genus are
used similarly. In Egypt, the doum palm has been revered as a
sacred tree, symbolizing masculine strength.
World production and yield
The doum palm is merely exploited from wild populations
throughout its habitat, thus no data are available. A mature
tree (6–8 years old) produces about 50 kg of fruit annually.
Uses and nutritional composition
The unripe seeds are edible, but the bony endocarp renders
the ripe ones of sufficient durability to be used as a weapon.
The hardened endosperm of the ripe seed has been used as
vegetable ivory. In Africa they have been crushed and utilized
as a millet substitute. The mesocarp of the fruit, though
fibrous and tough, can be variably sweet and pleasant tasting,
suggestive of carob or even ginger. They are often processed
into a beverage, but in the deserts of northern Africa, the
mesocarp is a significant source of nutrition. Seeds contain
4–5% protein. The plumule of the young seeding just below
ground is eaten as well. Like so many palms, many other parts
of the plant are used for fibre, forage and construction. A
variety of medicinal uses have been reported as well.
Jubaea
Botany
TAXONOMY AND NOMENCLATURE Hyphaene consists of six to
ten species of fan-leafed palms of arid and semi-arid areas of
the Old World. They are most diverse in Africa and there is
little doubt that man has influenced their distribution, either
by deliberate or accidental introduction. Hyphaene thebaica is
the most widely distributed species in the genus. It is a
member of the tribe Borasseae, a group of dioecious palms
distributed about the perimeter of the Indian Ocean and its
islands. Various other vernacular names include zembaba
(Amharic), dom (Arabic), gingerbread palm (English) and
mkoma (Swahili).
The doum is one of the few palms that branch
dichotomously on the aerial stem, one or more times, and
eventually can reach 10 m in height. The stiff leaves are large
and costapalmate (fan-shaped with an extension of the petiole
into the blade). The leaves are ribbed and the petiole is fiercely
armed with sharp teeth. The palms are dioecious. Staminate
and pistillate inflorescences, produced from between the
leaves, are similar catkin-like (male) or club-shaped (female)
branches that arise in clusters from a central rachis. The entire
inflorescence can exceed 1 m in length. The small individual
flowers are in sunken pits densely arranged on these branches.
The fruit is a drupe about 5 cm in diameter that turns
yellowish brown at maturity.
DESCRIPTION
Doum palm populations achieve their greatest size and productivity in moist,
tropical climates and can form thick stands along watercourses
in hot, dry areas. They are found in more arid situations as
well, however, fruit size and production is diminished where
the palms are subjected to great water stress. Doum palm is
also able to withstand a few degrees of frost, though cold
tolerance probably varies with geography.
ECOLOGY AND CLIMATIC REQUIREMENTS
Staminate and pistillate inflorescences
are found on separate plants of the doum palm. Pollination is by
wind. Elephants and baboons are significant fruit dispersal
agents, but human agency has probably had an impact on the
present distribution of the palm.
REPRODUCTIVE BIOLOGY
Doum palms first fruit at 6–8 years of
age. The fruit ripens in 6–8 months, often remaining on the
plant until the next flowering season.
FRUIT DEVELOPMENT
131
of drought. Young plants should be transplanted with care, as
the seedling stem is easily damaged. Field-grown plants
transplant with difficulty until an aerial trunk is formed; even
then root pruning, not necessary for most palms, is
recommended when moving larger plants.
Little is known of the doum
palm’s mineral nutritional needs, but growth is best in a rich,
sandy loam, with a pH of 6.5–7.5. It tolerates moderate salinity.
In Florida, it has proven tolerant of nutrient poor soils.
NUTRITION AND FERTILIZATION
DISEASES, PESTS AND WEEDS The nuts of the palm can be
parasitized by the scolytid beetle Coccotrypes dactyliperda.
Alan W. Meerow
Literature cited and further reading
Moussa, H., Margolis, H.A., Dubé, P.-A. and Odongo, J. (1998)
Factors affecting the germination of doum palm (Hyphaene
thebaica Mart.) seeds from the semi-arid zone of Niger, West
Africa. Forest Ecology and Management 104, 27–41.
Vogt, K. (1995) A Field Guide to the Identification, Propagation and
Uses of Common Trees and Shrubs of Dryland Sudan. SOS Sahel
International, Oxford, UK.
Jubaea chilensis
Chilean wine palm
Chilean wine palm, coquita, Jubaea chilensis (Molina) Baillon
(Arecaceae), is mainland Chile’s only native palm, and was
almost extirpated in the wild for sap collections. Now it is
protected by law. The fruit is sweet and edible, as is the
endosperm of the seed, which has a taste similar to coconut.
World production and yield
The harvest of the fruit of the Chilean wine palm is an
incidental use for the palm, for which no data are available.
Uses and nutritional composition
The sweet, fleshy mesocarp of the fruit is eaten fresh, and the
nuts (endocarp and endosperm) are either eaten fresh or used
to make various confections. The main traditional use for the
palm was the tapping of the stem for sap collection as a sugar
source (and fermented as a source of palm wine). The palm is
decapitated for this purpose.
Botany
Horticulture
Propagation of doum palm is by seed and
secondarily by separating and establishing basal offsets that
sometimes form at the base of the trunk. Seeds germinate
slowly (up to a year) and remotely (the cotyledonary petiole or
‘sinker’ emerges and grows deeply into the soil before the
formation of the seedling shoot axis) and a great deal of
underground development takes place before the first seedling
leaf emerges. It is best if the fruit wall is removed and then the
seed planted. The seeds are best planted singly in deep
containers, or directly in the ground (about 20 cm deep). Soil
needs to be kept moist for 2–3 months to ensure germination;
after that, the seedlings can withstand as much as 10 months
PROPAGATION
TAXONOMY AND NOMENCLATURE Only one species is recognized in the genus. It is closely related to the monotypic
genera Juania from the Juan Fernandez Islands off the coast of
Chile, and is known to hybridize with species of Butia and
Syagrus. Some synonyms include Cocos chilensis (Molina)
Kunth, Jubaea spectabilis Kunth and Micrococos chilensis
(Molina) Philippi.
Chilean wine palm produces a massive solitary
dark grey trunk 1 m or more in diameter and 10–15 m tall that
is usually swollen at the base or sometimes towards its middle.
A dense canopy of 40–50 green to grey-green pinnate leaves is
formed that may be 6–8 m wide and up to 6 m tall. Each leaf is
DESCRIPTION
132
Arecaceae
2–4 m long on short petioles with many 60 cm long, stiff, tworanked leaflets. The inflorescences, which are produced from
among the leaves, are once-branched, 1–1.5 m long, and bear
numerous purple flowers in clusters of a central female flanked
by two males. The round fruit are orange, one-seeded and
about 4 cm in diameter.
fresh fruit, with a local market value of US$1525. It is likely
that large stands of M. flexuosa occupy at least 1000 km2 in
Amazonas. Thus, an estimate of the quantity of buriti fruit
produced in the region would be about 600,000 t, the great
majority of which is consumed by wildlife.
Uses and nutritional composition
The habitat of the
Chilean wine palm is dry river courses or sparsely vegetated
Andean foothills of western Chile between 32° and 35°S
latitude at low elevation, but the range is now much more
restricted. Due to cutting of the palms for sap harvest and
land clearing, the number of Chilean wine palms has
decreased from an estimated 5 million to about 12,000 over the
past five centuries. Jubaea chilensis is adapted to a
Mediterranean climate with wet winters and dry summers,
and performs best in similar climates elsewhere in the world.
As a rule, it does grow very well in humid subtropical and
tropical climates. It is believed to be hardy to ⫺9°C.
ECOLOGY AND CLIMATIC REQUIREMENTS
REPRODUCTIVE BIOLOGY AND FRUIT DEVELOPMENT Flowering is from November to December in habitat (May to June in
the northern hemisphere), with fruit that ripen starting in
January (July in the northern hemisphere). Individual palms
may require as much as 60 years before they flower and fruit.
Horticulture
Seed is sown as soon as it is ripe and can take 6 months or
longer to germinate.
Alan W. Meerow
Mauritia flexuosa
buriti
The buriti palm, Mauritia flexuosa L.f., (Arecaceae) is one of
the most extensively utilized palms of the Amazon region. It is
a dioecious, single-stemmed, fan-leafed palm, broadly distributed throughout northern South America east of the
Andes, most typically in inundated soils. The fruit pulp is
eaten directly, dried into flour or fermented. Most, if not all,
of the fruit consumed is harvested from wild populations.
Common names throughout its range (Dugand, 1972;
Glassman, 1972; Bohorquez, 1977; Borgtoft-Pedersen and
Balslev, 1990) include buriti (from the Tupi Indian language:
mburi’ti), buriti-do-brejo, coqueiro-buriti, buritizeiro, miriti,
muriti, muritim, muritizeiro, muruti, palmeira-dos-brejos,
carandaguaçu, carandaiguaçu (Brazil); moriche (Venezuela,
Trinidad); ita (Guyana); palmier bâche (French Guiana);
achual, aguaje, auashi, bimón, buritisol, mariti, muriti, moriche
(Peru); caranday-guazu, ideuí (Bolivia); and cananguche,
chomiya, ideuí, mariti, miriti, muriti, moriche (Colombia).
World production and yield
It is estimated that an individual buriti palm could yield 200
kg of fruit/year (Lleras and Coradin, 1988), from which
approximately 24 kg of oil could be extracted. A density of 150
plants/ha could result in a harvest of 3600 kg of mesocarp
oil/year. Bohorquez (1976) reports harvests of 19 t of
fruit/ha/year from plantations of 100 palms/ha in Peru, a net
of 190 kg fruit/plant. From the Peruvian aguajale described
by Kahn (1988), Peters et al. (1989) estimated 6.1 t/ha/year of
From the fresh, pulpy mesocarp of the buriti’s fruit, a
beverage known as ‘buriti wine’ or ‘vinho de buriti’ is
prepared. Fruit are first softened in warm water for a few
hours, left under leaves for several days (they are collected and
often fall naturally before fully ripe). The slow treatment is
said to enhance flavour. The pulp is extracted by hand, diluted
with water and strained. Buriti wine is consumed fresh,
sweetened, or mixed with manioc flour, as is done with assaí
(Euterpe oleracea). Fresh pulp is made into a confection, doce
de buriti, and used in ice cream. The liquid is also diluted
further, sweetened and frozen like popsicles (dim-dim in
Brazil, curichi in Peru). The pulp is also dried and ground
into flour. The complete range of products is widely used and
figures importantly in the local commerce of the Amazon
basin (Cavalcante, 1988; Padoch, 1988). Lima (1987) reported
that a 20-day treatment with doce de buriti eliminated all
symptoms of hypovitaminosis A in children, a syndrome that,
ironically, is quasi-epidemic in the region. In the Orinoco
River delta of Venezuela, Indians make a type of bread from
the pulp (Braun, 1968). An alcoholic beverage is also
sometimes made by fermenting a mash of the fresh pulp. Oil is
also extracted from the fruit.
Though only 12–13% of the fruit dry weight, the buriti
pulp is an important source of calories, proteins and vitamins
for the people of the Amazon region (Table A.56). The pulp
contains about 3 g of protein, 10 g fat, and 120 to over 200
calories/100 g of fresh pulp (Chaves and Pechnik, 1946, 1949;
Leung and Flores, 1961; Bohorquez, 1977). The mesocarp oil
is also very high in vitamin A, and may represent one of the
highest concentrations of carotene in the plant kingdom.
Rizzini and Mors (1976) reported 300 mg of -carotene/100 g
of dry mesocarp, a level three times that of African oil palm
(Elaeis guineensis).
Botany
Until recently, buriti was
known as Mauritia vinifera and was considered distinct from
miriti or buriti-do-brejo (M. flexuosa) on the basis of male
inflorescence, fruit size and habitat differences. Mauritia
vinifera was considered endemic to the central planalto of
Brazil. It is now treated merely as an ecological variant of M.
flexuosa. One other species is recognized in the genus.
Mauritia and the related Mauritiella are the only fan palms
classified outside of the palm subfamily Coryphoideae, and are
members of the rattan group, subfamily Calamoideae (Uhl and
Dransfield, 1985).
TAXONOMY AND NOMENCLATURE
The buriti is a robust, solitary-stemmed palm
reaching 30 m in height. The roots often form pneumatophores
in flooded soils (Granville, 1974). The eight to 20 fan-shaped
(costapalmate) leaves form a large crown. Each leaf consists if a
2.5 m long blade divided into several hundred segments, and
DESCRIPTION
Mauritia
133
Table A.56. Nutritional value of 100 g of buriti palm mesocarp (Source: de Castro (1993).
Chaves and Pechnik
(1946, 1949)
Bohorquez (1977)a
FAO (1986)
Altman and Cordeiro
(1964)
Leung and
Flores (1961)
Constituent
Fresh
Fresh
Dry
Dry weight
Dry weight
Calories
Water (%)
Proteins (g)
Fats (g)
Fibre (g)
Ash (g)
Calcium (mg)
Phosphorus (mg)
Iron (mg)
Vitamin A (mg)
Thiamine (mg)
Riboflavin (mg)
Niacin (mg)
Vitamin C (mg)
120.0
71.8
2.9
10.5
11.4
1.2
158
44
5
30.0
–
–
–
50.5
143.0
72.8
3.0
10.5
11.4
1.2
113.0
19.0
3.5
12.0
0.3
0.23
0.7
26.0
–
–
5.5
31.0
23.0
2.4
–
–
–
30.0
0.1
–
–
52.5
–
68
5.2
26.2
27.5
2.9
–
–
–
–
–
–
–
–
265
72.8
3
10.5
11.4
1.2
–
–
–
–
–
–
–
–
a
Exocarp included.
an equally long petiole. A skirt of dead leaves often hangs down
below the crown. Plants are either male or female. The hanging
inflorescences are similar in both sexes and arise from the axils
of the leaves. Each flower stem is about 2 m long and bears
many conspicuous bracts. The flower-bearing branches or
rachillae number several dozen, and are catkin-like on the
males; the females are shorter and thicker. Male flowers are in
pairs on the branches; the female flowers are solitary. The fruit
are spherical or ellipsoidal, 4–5 cm in diameter and 5–7 cm
long. They are covered with brownish-red scales. The pulpy
mesocarp varies from yellow through orange to reddish orange
in colour. The endocarp (inner fuit wall) is soft. Each fruit
contains up to a dozen round seeds with brownish coats and
solid, white endosperm (Wessels-Boer, 1965).
ECOLOGY AND CLIMATIC REQUIREMENTS Mauritia flexuosa is
restricted to permanently or seasonally flooded soils. It is
found throughout Amazonia, from the Orinoco valley of
Venezuela in the north, through French Guiana and the
northern coast of the Brazilian state of Amapá, and west up to
the Andean foothills in Colombia, Ecuador, Peru and Bolivia.
It is common throughout north-eastern Brazil, from
Maranhão to Bahia states. It extends into the cerrado
vegetation of the central Brazilian states of Minas Gerais and
Mato Grosso do Sul (formerly M. vinifera), but is limited to
river margins and swamps. Also it occurs in Trinidad and
Tobago. The palms are found in small groves along
watercourses that wind through the non-flooded upland
rainforests, and eventually form large stands, often to the
exclusion of other tree species, in the major Amazonian
estuaries (Peters et al., 1989). These are known as aguajales in
Peru. In western Amazonas, buriti zones cover thousands of
hectares along floodplains. The cerrado populations of Central
Brazil form groves called veredas along riverbanks and other
hydric sites (Bondar, 1964). The buriti thrives in full sun and
humid, tropical conditions, and has little or no tolerance of
freezing temperatures. Growth on dry soils is slow and the
plants appear stunted.
REPRODUCTIVE BIOLOGY Flowering and fruiting in buriti
occur annually, but irregularly distributed throughout the year.
In central Amazonas, flowering occurs at the end of the rainy
season to the beginning of the dry season, from May to August
(de Castro, 1993). The fruit ripen during the following rainy
season and are offered in local markets from December to June.
Urrego (1987) reported similar phenology in Colombia. In
Iquitos, Peru, mature fruit are abundant in the markets from
February to August, with a distinct shortage from September to
November, due to seasonal fluctuations (Padoch, 1988). In
eastern Amazonas, in Belém, Cavalcante (1988) reports that
mature fruit appear in the markets from January to July. Heinen
and Ruddle (1974) reported two flowering seasons in the
Orinoco delta of Venezuela: one, primarily male plants, appears
to be initiated by the commencement of the rainy season; a
second takes place in December, amidst the rainy season, and is
dominated by female palms. However, mature fruit are found
throughout the year, but with two peaks, one from August to
October, and a less abundant period from February to April.
Gender ratios in natural populations are not well understood.
Urrego (1987) reported that 15–20% of male plants in a
population are sufficient to provide optimum fruit production.
FRUIT DEVELOPMENT The palms begin to bear fruit when
they attain about 6 m of height, usually in 7 or 8 years from
seed germination. They yield for several decades, the amount
of fruit declining after 40–50 years (Bohorquez, 1976; FAO,
1986). Unfortunately, tall palms with large fruit crops are
frequently cut down to make harvest easier, and exploited
populations dominated by male plants are a noticeable
consequence in Amazonas.
The mature fruit are dark red in colour and fall from the
trees. They quickly deteriorate, and must be collected before
they ripen fully if shipped.
Horticulture
Buriti is propagated exclusively by seed.
Bohorquez (1976) observed 100% germination after 75 days for
PROPAGATION
134
Arecaceae
seed collected no more than 10 days before sowing. When seed
was collected 3–4 weeks before sowing, germination was reduced
to 55% at 120 days. Seed from single-seeded fruit that was 8
days old began to germinate after 92 days, with 48% of the seeds
germinating between 120 and 150 days, and a final germination
of 52% (Storti et al., 1989). Seeds from double-seeded fruit
started to germinate after 55 days, with 41% germinating
between 120 and 150 days and a final germination of 64%.
Pretreatment of seed in running water at 29°C for 5 days and
immersion in a 100 ppm solution of gibberellic acid for 72 h,
increased final germination to 58% and 68%, respectively, but
did not alter germination frequencies (Storti et al., 1989). Little
is understood about seedling development and optimum size
and age for transplanting (Bohorquez, 1977). De Castro (1993)
observed seedling development in different levels of shade over
15 months, and concluded that shade is beneficial during the
early stages of growth. Once stem development, referred to as
‘establishment growth’ by Tomlinson (1990), is completed, the
buriti requires full or near full sun for best growth.
NUTRITION AND FERTILIZATION Nothing is known about
mineral nutrition and fertilization for the buriti palm.
DISEASES, PESTS AND WEEDS Little has been reported concerning pests and diseases of buriti palms. The large weevil
Rhynchophorus palmarum (Curculioniodae), the larvae of which
bore through the trunks of palms, is known to infest damaged or
otherwise stressed buriti palms. In fact, palms are sometimes
deliberately damaged to attract the weevils, as the larvae are
eaten by many indigenous people of the Amazon region (Suarez,
1966; Padoch, 1988; Borgtoft-Pedersen and Balslev, 1990).
MAIN CULTIVARS AND BREEDING No cultivar selection or
breeding efforts have been initiated on the buriti palm. De
Castro (1993) suggests that seed could be collected from the best
palms in each population and sown in situ or nursery grown then
transplanted back into the buriti zone to improve the quality of
the population. As the palms grow in sites that are marginal at
best for agricultural development, sustainable exploitation and
wise management of natural populations could provide a role in
Amazonian development without resulting in environmental
degradation (Peters et al., 1989).
Alan W. Meerow
Literature cited and further reading
Altman, R.R.A. and Cordeiro, M.M.C. de M. (1964) A
Industrialização do Fruto do Buriti (Mauritia vinifera Mart. ou M.
flexuosa). Química, Publicacao No. 5, INPA, Manaus, Brazil.
Bohorquez, J.A. (1976) Monografia sobre Mauritia flexuosa L.f. In:
Simpósio Internacional sobre Plantas de Interés Economico de la
Flora Amazonica. Informes de Conf., Cursos y Reuniones No. 93,
IICA, Turrialba, pp. 233–244.
Bohorquez, J.A. (1977) Monografia sobre Mauritia flexuosa L.f. In:
Simpósio Internacional sobre Plantas de Interés Economico de la
Flora Amazonica. Informes de Conferencias, Cursos y Reuniones
No. 93, IICA, Turrialba, Costa Rica, pp. 233–244.
Bondar, G. (1964) Palmeiras do Brasil. Boletim no. 2, Instituto
Botânica, Secao Agricultura São Paulo, São Paulo, Brazil, 159 pp.
Borgtoft-Pedersen, H. and Balslev, H. (1990) Ecuadorean palms for
agroforestry. AAU Reports (Aarhus University Press, Aarhus,
Denmark) 23, 1–122.
Braun, A. (1968) Cultivated palms of Venezuela. Principes 12, 36–64.
Castro, A. de (1993) Buriti. In: Clay, J.W. and Clement, C.R. (eds)
Selected species and strategies to enhance income generation
from Amazonian forests. Food and Agriculture Organization
(FAO) Miscellaneous Working Paper 93/6, Rome. Available at:
http://www.fao.org/docrep/v0784e/v0784e00.htm#Contents
(accessed 4 December 2006).
Cavalcante, P.B. (1988) Frutos Comestíveis da Amazônia, 4th edn.
Coleçao Adolpho Ducke, Museu Emílio Goeldi/Souza Cruz,
Belém, PA, Brazil.
Chaves, J.M. and Pechnik, E. (1946) Estudo da composição química e
do valor alimentício do buriti (Mauritia sp. Mart.). Revista de
Química Industrial 15, 140–141.
Chaves, J.M. and Pechnik, E. (1949) Em dois frutos brasileiros, o
maior potencial de pro-vitamina A que se conhece – buriti e
tucum. Revista de Química Industrial 18, 176–177.
Dugand, A. (1972) Las palmeras y el hombre. Cespedesia 1, 31–101.
Food and Agriculture Organization (FAO) (1986) Food and fruitbearing forest species, 3. Examples from Latin America. FAO
Forestry Paper 44/3, Rome.
Glassman, S.F. (1972) A Revision of B.E. Dahlgren’s Index of American
Palms. Phanerogamarum Monographie Tomus VI. Verlag Von J.
Cramer, Lehre, Germany, 294 pp.
Granville, J.J. de (1974) Aperçu sur la structure des pneumatophores de
deux espèces des sois hydromorphes en Guyane: Mauritia flexuosa
L. et Euterpe oleracea Mart. (Palmae). Cahiers Office de la Recherche
Scientifique et Technique de Outre-Mer sér Biologie 12, 347–353.
Heinen, H.D. and Ruddle, R. (1974) Ecology, ritual, and economic
organization in the distribution of palm starch among the Warao
Indians of the Orinoco Delta. Journal of Anthropological Research
30, 116–138.
Kahn, F. (1988) Ecology of economically important palms in
Peruvian Amazonia. Advances in Economic Botany 6, 42–49.
Leung, W.-T.W. and Flores, M. (1961) Food Composition Table for Use
in Latin America. US National Institute for Health, Bethesda,
Maryland.
Lima, M.C.C. (1987) Atividade de vitamina A do doce de buriti
(Mauritia vinifera Mart.) e seu efeito no tratamento e prevenção
da hipovitaminose A em crianças. Dissertação de Mestrado,
Depto. de Nutrição, Univ. Federal de Paraíba, Joao Pessôa, Brazil.
Lleras, E. and Coradin, L. (1988) Native Neotropical oil palms: state
of the art and perspectives for Latin America. Advances in
Economic Botany 6, 201–213.
Padoch, C. (1988) Aguaje (Mauritia flexuosa L.f.) in the economy of
Iquitos, Peru. Advances in Economic Botany 6, 214–224.
Peters, C.M., Balick, M.J., Kahn, F. and Anderson, A.B. (1989)
Oligarchic forests of economic plants in Amazonia: utilization and
conservation of an important tropical resource. Conservation
Biology 3, 341–349.
Rizzini, C.T. and Mors, W.B. (1976) Botânica Econômica Brasileira.
EPUSP, São Paulo, Brazil.
Storti, E.F., Ferraz, I.R. and Castro, A. de (1989) Tratamentos para
germinação de sementes de Mauritia flexuosa L.fil., Arecaceae.
XL Congresso Nacional Botanica, Cuiabá, Brazil.
Suarez, M.M. (1966) Les utilisation du palmier ‘moriche’ (Mauritia
flexuosa L.f.) chez les Warao du delta de l’Orénoque, territoire
Amacuro, Venezuela. Journal d’Agriculture Tropica et Botanique
Appliquée 13, 33–38.
Tomlinson, P.B. (1990) The Structural Biology of Palms. Oxford
University Press, New York, 492 pp.
Oenocarpus
135
Uhl, N.W. and Dransfield, J. (1985) Genera Palmarum. Allan Press,
Lawrence, Kansas, 610 pp.
Urrego, G.L.E. (1987) Estudio preliminar de la fenologia de la
Cananguchi (Mauritia flexuosa L.f.). Tesis, Universidad Nacional
Colombia, Facultad de Agronomia, Medellin.
Wessels-Boer, J.G. (1965) The Indigenous Palms of Suriname. E.J.
Brill, Leiden, the Netherlands, 172 pp.
Nypa fruticans
nipa palm
Nipa palm, Nypa fruticans Wurb. (Arecaceae), is a mangrove
palm that can form large populations along tidal estuaries in
south Asia and Australasia. The endosperm of the seed is
edible and consumed at various stages of development. Many
other parts of the palm are used as well, with the tapping of
the flower stems for the sugary sap being the most important.
Uses and nutritional composition
The gelatinous stage of the endosperm is consumed directly
or else preserved in syrup. Once solidified, it may be ground
up into a meal. The seeds contain 71–78% carbohydrate,
much of which is starch.
Botany
Nypa fruticans is the only
species classified in the subfamily Nypoideae of the palm
family. It is thus considered quite isolated from the other
palms, and is in fact one of the earliest palms identifiable in
the fossil record (Uhl and Dransfield, 1987).
TAXONOMY AND NOMENCLATURE
Nipa palm grows from a dichotomously
branching, prostrate stem that often remains underground
(Fig A.18). In time, a single plant can form a network of
creeping stems. Each stem branch holds fewer than a dozen,
5–9 m long, sub-erect pinnate leaves, each with numerous
1.2–1.5 m long lanceolate leaflets. The inflorescence is 1–2 m
long and spadix-like, with the male flowers born on catkin-like
lateral branches and the female flowers in a congested head.
The female flowers are much larger than the male. The fruit
are aggregated in a globular head that is dispersed as a unit
capable of floating.
DESCRIPTION
Fig. A.18. Nypa fructicans palm and fruit (with permission from Sitijati
Sastrapradja from Palem Indonesia, Lembaga Biologi Nasional, 1978).
production, rather than fruit production. Natural stands are
sometimes managed by thinning to between 2500 and 3500
palms/ha, at a spacing of 1.5–2 m. Wider spacing (380–750
trees/ha) has been advocated by some (Duke, 2001). Four
hours of inundation appears to be beneficial for the growth of
nipa palms, which may achieve 2 m of height in their first year
from seed. Nuts are harvested throughout the year.
DISEASES, PESTS AND WEEDS Grapsid crabs damage young
nipa palms (Duke, 2001).
Alan W. Meerow
Literature cited and further reading
Nipa palm forms
large populations in heavy muds of tidal estuaries from India
and Sri Lanka to Australia. It also occurs on the Solomon and
Ryukyu Islands, and is naturalized in parts of tropical Africa.
It is extremely salt and flood tolerant. Tropical conditions are
required for growth.
ECOLOGY AND CLIMATIC REQUIREMENTS
Duke, J.A. (2001) Handbook of Nuts. CRC Press, Boca Raton,
Florida, 343 pp.
Uhl, N.W. and Dransfield, J. (1987) Genera Palmarum. Allen Press,
Lawrence, Kansas, 610 pp.
Oenocarpus bataua
REPRODUCTIVE BIOLOGY
Pollination of nipa palm is by
drosophilid flies.
Horticulture
PROPAGATION
Nipa palm is propagated by seed or detached
stem branches.
TRAINING AND MANAGEMENT Management strategies of
natural nipa palm stands are oriented for maximum sap
patauá
Patauá, Oenocarpus bataua Mart. (Arecaceae), the batauá or
patauá palm, is a robust, feather-leafed (pinnate) canopy palm
of northern South America, and one of most common palms
in the region. A beverage is prepared from the fruit, but of
greater importance is the oil obtained from boiling them,
which has been favourably compared to olive oil. The quantity
and quality of this oil has repeatedly resulted in the citation of
this palm as an underutilized plant with great economic
potential.
136
Arecaceae
Related species include Oenocarpus bacaba Martius (bacaba:
Brazil) which is widely used in northern South America as a
source of vinho de bacaba, a thick, somewhat oily juice prepared
from the slurry of mesocarp and water. The bacaba is a large,
single-stemmed palm growing both in flooded and in nonflooded areas in rainforest ecosystems. In the latter ecosystem,
bacaba may form high-density stands. Oenocarpus balickii F.
Kahn (sinamillo: Peru) occurs in Amazonian Peru, Brazil and
Colombia and is similar to Oenocarpus mapora but differs by its
single stem, larger number of leaflets and smaller drupes. It is
frequent on well-drained rainforest soils. A drink is made from
the fruit. Oenocarpus distichus Martius (bacaba-do-leque: Brazil)
is found on sandy soils of seasonally dry forest in south-eastern
Amazonas. The palm differs from bacaba by its opposite leaves;
the latter species has spirally arranged leaves. The fruit of this
species is used to prepare vinho de bacaba by people of the
south-eastern Amazonas. Oenocarpus mapora Karsten
(bacabinha: Brazil) is found from southern Central America
through north-western South America and the fruit of this
species is made into vinho de bacaba. It is a clustering palm of
smaller stature than most Oenocarpus spp. It typically occurs as
dispersed individuals within terra firma or seasonally inundated
forest. Two apparent hybrids also exist in the wild, O. bacaba ⫻
O. bataua and O. bacaba ⫻ Oenocarpus minor. The presence of
natural hybridization bodes well for future breeding
programmes designed to improve yields, tolerance to stress,
reduced time to maturity, etc.
Uses and nutritional composition
The oil extracted from the mesocarp of patauá is virtually
identical to olive oil in appearance and fatty acid composition
(Balick, 1986, 1988). Patauá oil is highly unsaturated, with 78
± 3% monounsaturated fatty acids and 3 ± 1% polyunsaturated fatty acids (Table A.57). Indigenous people in
Amazonas consider it second to no other wild plant as an oil
resource.
In the Amazon basin and northern South America,
Amerindians produce a thick non-alcoholic juice with a nutlike flavour from the fruit mesocarp. Its high protein content
and unsaturated oils make it an excellent nutritional addition
to local diets (Balick and Gershoff, 1981; Table A.58). Patauá
oil is considered a cure for minor bronchial and pulmonary
infections (Balick, 1986).
Botany
Long treated as a species of
the segregate genus Jessenia, O. bataua is one of nine species
distributed throughout from Central America through northern
South America south to Brazil and Bolivia (Henderson, 1994).
Synoynms include Jessenia bataua (Martius) Burret, Jessenia
polycarpa Karsten and Jessenia repanda Engel. Other common
names include batauá, milpesos, seje, trupa, chapel, patawa,
turu, unguraui, komboe, yagua and aricauguá.
TAXONOMY AND NOMENCLATURE
The patauá is a large (15–25 m), singlestemmed palm with a trunk of 15–25 cm diameter at breast
height. The crown consists of eight to 16 spirally arranged,
large, pinnately compound leaves that may reach 10 m in
length (the petiole is about 1 m long, the blade 3–8 m). The
100–200 segments (pinnae) are arranged in a single plane
along the rachis. The paniculate inflorescence has several
hundred branches (rachillae), each about 1 cm long, bearing
numerous cream-coloured flowers. Each panicle may bear over
1000 round, dark purple drupes, each 2.5–4 cm in diameter
and containing a single seed.
DESCRIPTION
World production and yield
Despite acclaim as an overlooked agronomic resource, few data
on yield of the patauá palm have been accumulated. On
average, a patauá palm yields two fruiting stems annually, with
a mean weight of about 15 kg, although individuals producing
as many as five fruiting stems have been reported under
favourable conditions (Balick, 1988). Each fruit contains
6.5–8% oil by fresh weight (Blaak, 1988). Sirotti and Malagutti
(1950) estimated that a natural population in Venezuela
consisted of 400–500 reproductively mature palms/ha, of
which just over 70% bore fruit. Balick (1988) suggests that the
most productive spacing for an agroforestry plantation would
likely be more like 204–216 plants. Balick (1988) estimated that
100 fruiting plants would produce 1.6 t of fruit. Peters et al.
(1989) estimated 3.5 t based on 104 plants/ha. From a hectare
of wild palm stands, yields will vary from 1.6 to 1.5 t of fresh
fruit, from which 112–260 kg of patauá oil can be expected to
be extracted (Balick, 1993). If Balick’s predicted plantation
density is accurate, per hectare yields of 3.27 t of fruit and
240–525 kg of patauá oil could be expected.
Traditionally, the oil was extracted by mashing the
mesocarp, heating and pressing it in a tipitipi, a long woven
tube (Balick, 1986, 1988), which successfully extracted only
about one-third of the oil (Blaak, 1988). Technology similar to
that utilized for African oil palm (Elaeis guineensis) would
increase extractive yield to at least 85–89%. Ironically, South
America is currently a net importer of olive oil. During World
War II, when a world shortage of olive oil occurred, Brazil
exported over 200 t/year of patauá oil (Pinto, 1951).
Unfortunately, most of the harvest was accomplished by
felling mature trees in natural populations.
On upland soils of
terra firma rainforests, patauá has a scattered distribution of
mostly juvenile palms (Kahn, 1988). The species has not been
reported above 950 m elevation (Balick, 1986). In flooded
swamps, O. bataua can form huge, virtually unbroken stands
(Peters et al., 1989). It is to be expected that patauá has little or
no resistance to frost.
ECOLOGY AND CLIMATIC REQUIREMENTS
Table A.57. Fatty acid composition of patauá mesocarp oil (in % total oil).
Fatty acid
Myristic
Palmitic
Palmitoleic
Stearic
Oleic
Linoleic
Linolenic
Unsaturated (%)
a
Mean + SD of 12 samples.
Jamieson
(1943)
Balick and Gershoff
(1981)a
–
8.8
–
5.6
76.5
3.4
–
79.9
–
13.2 ± 2.1
0.6 ± 0.2
3.6 ± 1.1
77.7 ± 3.1
2.7 ± 1.0
0.6 ± 0.4
81.6 ± 4.7
Oenocarpus
137
Table A.58. Amino acid composition of patauá mesocarp protein (mean and + SD of seven samples) (Source:
Balick and Gershoff, 1981).
Non-essential amino acids
Aspartic acid
Serine
Glutamic acid
Proline
Glycine
Alanine
Histidine
Arginine
mg/g protein
122 ± 8
54 ± 3
96 ± 5
75 ± 8
69 ± 4
58 ± 4
29 ± 4
56 ± 2
Little is recorded on flowering
phenology of this palm. Fruit ripen from April to November.
REPRODUCTIVE BIOLOGY
Some authors have reported that
patauá takes 10–15 years to fruit (Balick, 1988). As with most
palms, however, this time may be reduced by modifications in
the agroecosystem, especially by reducing competition and
enriching the nutrient content of the soil. Balick (1986)
observed a plant in Ecuador that fruited precociously, at less
than 2 m from the ground. The patauá germplasm bank being
organized at the Centro de Pesquisas Agropecuárias do
Trópico Umido, in Belém, Pará, will provide information on
the precocity of different genotypes in plantations. This trait
generally has a moderate to high heritability and can be
selected for in an improvement programme.
FRUIT DEVELOPMENT
Horticulture
Fresh seed should be de-pulped, and placed in
warm water (50°C) for 30–60 min. Treated in this manner,
90–98% germination can be expected within 2 months
(Balick, 1988). Viability diminishes quickly, with major losses
occurring in as little as a month. Germination is best
conducted in part shade. As soon as the seedling has at least
one leaf, it should be transplanted to a container with welldrained organic media and grown under 50% shade for the
first year. Inoculation with mycorrhiza (St John, 1988)
increases growth.
Seedlings intended for transplanting into natural stands or
rainforests can be directly planted from the nursery, while
plants that will be established plantation-style must be adapted
to full sun first. Young transplants can be shaded for the first
few weeks using a folded leaf from an adult palm. Blaak (1988)
recommends 7 m square spacing for field plantings.
PROPAGATION
NUTRITION AND FERTILIZATION Blaak (1988) suggested that
1.5 kg of fertilizer/plant should increase yields twofold. No
other information is available on mineral nutrition and
fertilization of patauá palms.
DISEASES, PESTS AND WEEDS
No information on diseases, pests
and weeds is available. Balick (1988) noted that patauá palms in
Colombian agroforestry settings produced more fruit than those
in primary forest, suggesting that lessening competition from
other plants has beneficial effects on production.
Essential amino acids
Isoleucine
Leucine
Lysine
Methionine
Cystine
Phenylaline
Tyrosine
Threonine
Valine
Tryptophan
mg/g protein
47 ± 4
78 ± 4
53 ± 3
18 ± 6
26 ± 6
62 ± 3
43 ± 5
69 ± 6
68 ± 4
9±1
MAIN CULTIVARS AND BREEDING No sustained improvement
efforts have yet been undertaken for the patauá palm. The
presence of at least two putative hybrids in nature, O. bacaba
⫻ O. bataua and O. bacaba ⫻ O. minor, suggest that future
breeding programmes would be a worthy avenue of pursuit.
Alan W. Meerow
Literature cited and further reading
Balick, M.J. (1986) Systematics and economic botany of the
Oenocarpus-Jessenia (Palmae) complex. Advances in Economic
Botany 3, 1–140.
Balick, M.J. (1988) Jessenia and Oenocarpus: neotropical oil palms
worthy of domestication. Food and Agriculture Organization (FAO)
Plant Production and Protection Paper No. 88, Rome.
Balick, M.J. (1993) Patauá. In: Clay, J.W. and Clement, C.R. (eds).
Selected species and strategies to enhance income generation
from Amazonian forests. Food and Agriculture Organization
(FAO) Miscellaneous Working Paper 93/6. FAO, Rome. Available
at:
http://www.fao.org/docrep/v0784e/v0784e0f.htm#patauá
(accessed 4 January 2006).
Balick, M.J. and Gershoff, S.N. (1981) Nutritional evaluation of the
Jessenia bataua palm: source of high-quality protein and oil from
tropical America. Economic Botany 35, 261–271.
Blaak, G. (1988) Mechanical extraction and prospects for
development of a rural industry. In: Balick, M.J. (ed.) Jessenia and
Oenocarpus: neotropical oil palms worthy of domestication. Food
and Agriculture Organization (FAO) Plant Production and
Protection Paper No. 88. FAO, Rome, pp. 65–83.
Henderson, A. (1994) The Palms of the Amazon. Oxford University
Press, New York, 388 pp.
Jamieson, G.S. (1943) Vegetable Fats and Oils. Reinhold Publishing
Co., New York, 437 pp.
Kahn, F. (1988) Ecology of economically important palms in
Peruvian Amazonia. Advances in Economic Botany 6, 42–49.
Peters, C.M., Balick, M.J., Kahn, F. and Anderson, A.B. (1989)
Oligarchic forests of economic plants in Amazonia: utilization and
conservation of an important tropical resource. Conservation
Biology 3, 341–349.
Pinto, P.G. (1951) Contribuição ao estudo químico do sebo da
ucuúba. Boletim Techico do Instituto Agronomico do Norte 18.
Belém, Pará, Brazil.
Sirotti, L. and Malagutti, G. (1950) La agricultura en le territorio
Amazonas, exploitación del seje (Jessenia bataua) palmera
oleaginosa. 1 de enero (manuscript), Caracas, Venezuela.
138
Arecaceae
St John, T.V. (1988) Mycorrhizal enhancement in growth rate of
Jessenia bataua seedlings. In: Balick, M.J. (ed.) Jessenia and
Oenocarpus: neotropical oil palms worthy of domestication. Food
and Agriculture Organization (FAO) Plant Production and
Protection Paper No. 88. FAO, Rome, pp. 140–148.
Phoenix dactylifera
date palm
Date palm, Phoenix dactylifera L. (Arecaceae), is a major fruit
crop in arid regions such as North Africa and the Middle East.
The fruit are the main income source and staple food source
for local populations and have enormous significance in the
economy, society and environment in arid regions. During the
last century, dates were introduced to Australia, Mexico,
Pakistan, South Africa and the USA.
History and origins
Dates are one of the oldest known fruit crops and have been
cultivated in the Middle East and North Africa for thousands
of years (Zohary and Hopf, 2000). The earliest known record
in Iraq (Mesopotamia) shows that its culture was probably well
established as early as 5000 years ago. This long history of date
palm cultivation in the regions and the wide distribution of
date palm beyond its original ranges have made it difficult to
pinpoint its centre of origin. Although the origin of dates is
unknown, date palms most likely originated from the ancient
Mesopotamia area (southern Iraq) or western India. There are
two hypotheses about the origin of dates, one suggesting that
the date palm descended from one or several species of the
genus Phoenix cultivated in their natural habitats, and the
other suggesting it is a hybrid of several unknown Phoenix
species. From its centre of origin, date cultivation spread
throughout North Africa and the Middle East wherever
conditions were favourable.
The earliest dissemination of date palms must have been by
seeds carried by human beings from oasis to oasis. Date
culture had apparently spread into Egypt 3500 years ago,
although it did not become important until later, when it
spread westward across North Africa from Egypt. The spread
of date culture was facilitated by the introduction of the camel
into this area in the 6th century, which made the transport of
off-shoots possible. The spread of date cultivation
accompanied the expansion of Islam and reached the northern
outpost in southern Spain and an eastern extension into
Pakistan. Until recently, expansion of date culture south of the
Sahara Desert was limited because of lack of water. The
Spanish were the first to introduce date palms to America. In
the past century, dates were introduced to the desert areas of
the Colorado River in North America, the Atacama Desert in
South America, the Kalahari of South Africa and the great
central desert of Australia.
Throughout the history of the Middle East, the date has
had a very important influence. Without the date, no large
human population could have been supported in the desert
regions. The caravan routes existed for centuries mainly for
the transport of dates. Early on, date cultivation achieved a
high level of status in the Middle East, and became a sacred
symbol of fecundity and fertility. Many ancient beliefs and
other representations indicate the importance of the date palm
for millennia. Dates had great spiritual and cultural
significance to peoples in the region. The date palm and date
culture are depicted on ancient Assyrian and Babylonian
tablets, including the famous Code of Hammurabi, which
contained laws pertaining to date culture and sales. The date
palm is also found in ancient Egyptian, Syrian, Libyan and
Palestinian writings. It is in Egyptian culture that the date
palm achieves its greatest esteem.
World production and yield
The worldwide production of dates reached 6,259,688 t in
2002. The top ten date producing countries in 2001 were
Egypt, Iran, Saudi Arabia, United Arab Emirates (UAE), Iraq,
Pakistan, Algeria, Oman, Sudan, and Libyan Arab Jamahiriya.
The top five date exporting countries in 2001 were UAE, Iran,
Pakistan, Tunisia and Iraq. The top five date importing
countries in 2001 were India, UAE, Pakistan, France and
Syria. The European and US markets are also important
export markets of dates. Egypt is the largest date producing
country, where production has increased from 439,539 t in
1982 to 1,113,270 t in 2002. Egyptian production accounted
for 17.8% of worldwide date production in 2002 on 29,461 ha
of dates harvested, and it had one of the highest yields/ha in
the world. In UAE, there were about 1.5 million date palms in
1971 when the country was founded, and an estimated 18
million date palms in the mid-1990s. UAE also has one of the
largest date tissue culture (TC) establishments and produces
millions of TC date palms annually.
Uses and nutritional composition
Date fruit are usually eaten fresh or dried. Dates can be eaten
from the middle of the Khalal stage to the Tamar stage
depending on the cultivar (the fruit ripening stages are
explained under ‘Fruit growth and development’). Dates in the
Khalal stage are at their largest size and highest sugar content.
In areas with marginal heat accumulation, there is often not
much ripening beyond the Khalal stage. Consumption of
Khalal dates is generally confined to areas near production in
North Africa and the Middle East. Where there is slightly
more heat, dates must be eaten at the time they begin to soften
as there is not enough heat to dry them out before they
ferment. However, in warmer regions, dates dry down to a soft,
dry fruit stage and then may be classified as ‘soft’, ‘semi-dry’ or
‘dry’. Soft dates pass through the Rutab stage and remain soft
at Tamar, with high moisture content. Semi-dry dates also pass
through Rutab, but dry out further at Tamar. Semi-dry dates
account for the majority of varieties. Dry dates (‘bread’ dates)
do not pass through Rutab but dry out quickly and have the
lowest moisture content at harvest. In humid climates, dry
dates may be relatively soft but in most instances of production
are quite hard and brittle. These categories are somewhat
arbitrary and are influenced by climate and production
practices, some varieties may classify differently depending
upon these factors (Dowson and Aten, 1962).
Dates are a high-energy food source with high sugar
content. About 72–88% of the dry matter in dates is sugar and
few ripe dates have any starch. As ripening progresses, the
sucrose is hydrolysed into ‘invert’ or ‘reducing’ sugars, a
mixture of glucose and fructose, depending upon variety,
Phoenix
cultural practices and other factors. Soft and semi-dry dates
are primarily of the reducing sugar type (having higher levels
of sucrose hydrolysed), while dry dates are mostly of the
sucrose type (having a lower proportion of the sucrose
hydrolysed) (Rygg, 1975; Vandercook et al., 1980; Ahmed et
al., 1995). Dates are a good source of iron and potassium, a
fair source of calcium, chlorine, copper, magnesium and
sulphur, and contain a small amount of phosphorus and 16
kinds of amino acids (Vandercook et al., 1980) (Table A.59).
These minerals accumulate during maturation (Ahmed et al.,
1995). Dates contain small amounts of vitamins A, B1 and B2,
and substantial amounts of nicotinic acid. Aqueous extracts of
date fruit also have potent antioxidant and anti-mutagenic
properties.
In addition to being consumed as fresh or as dried fruit,
dates are pressed into a large cake. Other products include:
date honey made from the juice of fresh fruit; date sugar; date
sap for making beer or wine; date palm flour made from the
pith of the trees; oil from the seeds; and the palm heart eaten
in salads. There are also non-food uses. For instance, seeds can
be used as animal feed or strung as beads. The wood of date
palms can be used for doors, beams, furniture, rafters and
firewood; leaves can be used for matting, baskets, roofing,
fencing and shelter; and fibres from dates can provide thread
and rigging for boats. Also dates are used as folk remedies for
many medicinal purposes. Recently the date palm has been
used as an ornamental and in landscapes in southern Europe
and the USA.
Botany
Date is a diploid (2n = 2x
= 36), perennial, monocotyledonous plant belonging to the
TAXONOMY AND NOMENCLATURE
Table A.59. Proximate fruit composition of dried medjool date edible flesh
(Source: USDA, 2004).
Proximate
Water
Energy (kcal)
(kJ)
Protein
Lipid (fat)
Carbohydrate
Fibre
Ash
Minerals
Calcium
Iron
Magnesium
Phosphorus
Potassium
Sodium
Vitamins
Ascorbic acid
Thiamine
Riboflavin
Niacin
Vitamin A
%
21.3
277
1160
1.81
0.15
74.97
6.7
1.74
mg
64
0.9
54
62
696
1
mg
0
0.05
0.06
1.61
149 IU
139
family of Arecaceae/Palmaceae. The name of date palm
originates from its fruit; ‘phoenix’ from the Greek means
purple or red (fruit), and ‘dactylifera’ means the finger-like
appearance of the fruit bunch. Some authorities claim that the
word phoenix itself has referred to the date palm since ancient
times, and others believe that the date palm has characteristics
of the legendary phoenix bird. The genus Phoenix is
distinguished from other genera of pinnate-leaved palms by
the upward and lengthwise folding of the pinnae and the
furrowed seeds. The taxonomy of Phoenix has been somewhat
confused in the past, with most authors recognizing
approximately 17 genera (but not necessarily the same genera).
The recent revision of Phoenix recognizes 13 species including
P. dactylifera (Barrow, 1998). All species are native to the
tropical or subtropical parts of Africa or southern Asia. This is
in contrast to other species of Phoenix, which have often been
reported under different species names, albeit mostly in the
genus Phoenix (Barrow, 1998). Most Phoenix species are cross
compatible.
Dates are grown on a wide variety of soils throughout the
world, from sand to clay. However, production is improved if
soils have the maximum water-holding capacity consistent
with good drainage. Some sandy soils require high levels of
irrigation and fertilization. Hard pans or perched water tables
can result in shallow root systems or poor growth due to
anaerobic conditions or soil-borne pathogens. Soils with poor
drainage can result in saline or sodic soils, which are found in
many areas of date production.
Date palms are considered having the highest tolerance to
saline conditions among all tree crops, tolerating up to 4.0
dS/m before yield reductions are noted. This assessment was
based upon work done in Indio, California by Furr and
associates. Furr and Armstrong (1962) compared mature date
palms irrigated with water salinized to 4–8, 8–12 and 16–24
dS/m with non-salinized controls. They reported no
reductions in leaf growth rate, yield, fruit size and quality, or
chloride ion (Cl⫺1) content of the leaf pinnae. Growth of date
palm seedlings linearly decreased as electrical conductivity
rose from 6.5 to 39.0 dS/m; however, accumulation of sodium
(Na+) and Cl⫺ ions in leaf tissue was not observed. This
suggests that reductions in seedling growth were due to
osmotic rather than specific ion effects. Young date trees were
more sensitive to these effects than mature trees. None the
less, very saline conditions can cause reductions in yield and
quality. Date palms experience yield reductions if the EC of
the soil exceeds about 4 dS/m or if the EC of applied
irrigation water exceeds about 3 dS/m. If total soluble solids
in irrigation water exceed several hundred ppm, an extra
quantity of water should be applied for leaching. The leaching
ratio for date palms is estimated to be from 5 to 20, depending
upon the salinity.
The date palm has an erect columnar trunk,
40–50 cm in diameter (without leaf sheaths) that may reach a
height of 20–28 m or more (Fig. A.19). The trunk may have
one to several suckering off-shoots at or near the base. In
cultivation, the off-shoots are removed for propagation
purposes (discussed under ‘Propagation’). If the off-shoots are
left attached, it can result in a clumping or thicket-like growth
habit. The root system of the date palm does not have a
DESCRIPTION
140
Arecaceae
Fig. A.19. Leaf, flower and fruit of date (Source: Le Maout, 1877).
taproot. The root is fibrous and pneumatic. It may extend
several metres laterally and 6 m or more in depth, although
the bulk of the roots are distributed within 2 m of the trunk
both laterally and in depth. The crown of the date palm
consists of 60–150 leaves depending upon variety, growing
condition and cultural practices. Morphological differences in
leaf, spine, pinnae and fruit are commonly used to characterize
the varieties. The leaves of the date palm have a lifespan of 3–7
years. The colour ranges from light to dark shades of green.
The leaves are glaucous in different degrees depending on the
variety and age of the leaves. The leaves are 3–7 m in length,
based on the measurement of the blade from the lowest spine
to the tip of the terminal leaflet. Leaves with a blade length
less than 335 cm are considered short, from 335–427 cm as
medium, and long if more than 427 cm. The midrib or petiole
is triangular in cross-section, with its greatest width at the
base, tapering rapidly towards the apex. The midribs are
straight or with different curvatures depending upon the
variety, are vertical in orientation, and are armed with rows of
pinnae (also referred to as leaflets) on each side. The pinnae
are long, narrow and folded upward and lengthwise. Varieties
differ in the number, thickness, length, breadth and relative
stiffness of the pinnae and how they droop or curve down. A
characteristic of the genus Phoenix is that the leaves have
terminal leaflets, and the basal leaflets are modified into spines.
The length and breadth of the pinnae are important
morphological characters for varietal description. Pinnae with
length less than 61 cm are considered short, 61–75 cm as
medium, and long if more than 75 cm. The breadth of pinnae
is considered short if less than 3.8 cm, medium from 3.8 to
4.4 cm, and broad if more than 4.4 cm. The pinnae usually
grow in groups of two or more, relatively close together at the
point of attachment to the midrib and separated from other
groups by a space greater than that between the pinnae within
the group. Groups of two or three pinnae are common, but
some varieties have four or more. The lower leaflets near the
base of the trunk are modified into stout spines. The number
and length of the spines differ with variety and age. Spines
with a length of less than 10 cm are considered short,
10–15 cm medium, and long if more than 15 cm. The shortest
spines are at the base of the spine area and the longest near the
pinnae. Date palms have a phyllotaxic arrangement of leaves in
which leaves are arranged in spirals of five and 13 in one
direction around the trunk, and in spirals of eight in the
opposite direction.
Date palm is dioecious, meaning that it has separate female
and male trees. Occasionally hermaphroditic trees or flowers
are observed. The inflorescences of male and female trees
differ in morphology. Inflorescences of both sexes are enclosed
in a hard, fibrous cover (the spathe) during the early stages of
annual development. The spathe protects the delicate flowers
from heat and sunlight until they are ready to perform their
reproductive functions. The flowers (and later the fruit, on
female trees) are borne on a flat, tapering peduncle or rachis,
commonly known as the ‘fruit stalk’ in female varieties. This is
relatively short (50 cm) and upright in male trees, whereas in
female trees it is longer (100 cm) and upright at the beginning
of the flowering season, later elongating and drooping. The
inflorescence consists of many unbranched rachillae,
commonly known as ‘strands’, arranged in spirals on the
rachis. The rachillae of male spathes are crowded towards the
end of the rachis, and are short and robust. Male flowers are
waxy white, and crowded along the length of the rachilla.
There are usually three sepals and three petals. The yellowishgreen female flowers (also usually with three sepals and three
petals) are borne in clusters of three along the entire length of
the long, slender rachillae, which are more evenly distributed
along the rachis than in the males. Just before flowering, the
inflorescence that arises in the axis of the leaves pushes up
through the fibrous sheaths that form the leaf bases. The
spathe cracks longitudinally at anthesis. Only the portion of
the rachilla that bears flowers is exposed. Only one ovule per
flower is fertilized, producing a single date fruit. After
anthesis, no noticeable change is observed in the size of the
flower-bearing region. The fruit stalk elongates 50–60 days
after flowering and pushes out the portion of the inflorescence
that does not bear flowers to a length of 60–120 cm.
Date fruit are variable in size and shape, depending upon
variety, climate and cultural practices. The shape of the date
fruit is characteristic of the variety, and is most distinct during
the Khalal stage. Fruit shapes are oblong, elliptical, oval, ovate
or obovate. Date fruit are generally 4–7 ⫻ 2–3 cm, and 60 g or
more in weight. The calyx is persistent and useful in varietal
identification. Skin colour is green in the early stages (through
Kimri). At Khalal, fruit become distinctive colours ranging
from yellow to red and brown. As the fruit matures through the
Rutab and Tamar stages, it generally becomes darker and is
usually some shade of brown at harvest. The mesocarp is sweet,
thick and fleshy, or dry and thin depending upon the time of
harvest. Commercial harvest of the earliest varieties begins in
August in the northern hemisphere, with the last of the late
varieties being harvested in November. Date seeds are variable
in size and shape, but are generally elongate with pointed
apices. Seed size ranges from 20 to 30 mm by 5 to 8 mm.
Phoenix
ECOLOGY AND CLIMATIC REQUIREMENTS The date can grow
in very hot and dry climates, and is relatively tolerant to salty
and alkaline soils (Zaid and De Wet, 2002). It needs a long,
intensely hot summer with little rain and very low humidity
from pollination to harvest, but with abundant underground
water near the surface, or irrigation as in desert oases or river
valleys. It can grow in temperatures ranging from 12.7 to
27.5°C, withstanding up to 60°C and sustaining short periods
of frost down to ⫺5°C. The ideal temperature from
pollination to the fruit ripening stages ranges from 21 to 27°C.
Date roots are adventitious and grow horizontally for a long
distance. The fruit of dates vary in size, ranging from 2 to 60 g
in weight, from 2 to 11 cm in length and from 1 to 3 cm in
width depending on the cultivar. For proper maturing of fruit,
dates require prolonged summer heat without rain or high
humidity during the ripening period. Dates are grown in a
wide variety of soil, but soil with maximum water-holding
capacity consistent with good drainage is desirable. Dates are
widely grown in the arid regions between 15° and 35° N, from
the Canary Islands and Morocco in the west to India in the
east. Date flowers when the shade temperature rises above
18°C and forms fruit when the temperature is over 25°C.
Early work using 0°C as a base indicated that a heat
accumulation of over 5100° was necessary for production of
acceptable date fruit (Dowson, 1982). Swingle (1904) used the
figure of 18°C as the lower limit (based upon maximum
temperature), since this is the temperature necessary for
flowering. Using this figure, Swingle found that good quality
dates needed at least 3000° over the period of May–October
(184 days).
Although date palms have a high water requirement, they
are adapted to arid, low rainfall conditions. Rain during the
flowering and pollination season can affect fruit set. High
humidity can increase the incidence of diseases or disorders.
Extremely high and low humidity can cause soft, sticky or
extremely dry fruit, respectively. Ideally, rainfall is nonexistent and humidity is low from bloom to harvest. Winter
rain not damaging the fruit can alleviate saline soil conditions.
REPRODUCTIVE BIOLOGY The date is wind pollinated in
nature but insect pollination is possible. One male tree
produces sufficient pollen for pollination of approximately 50
female trees. Commercially, few male trees are grown in the
gardens and pollen is collected for artificial pollination that is
critical for successful production. Artificial pollination is an
ancient practice in North Africa and the Middle East, and is
mentioned in the cuneiform texts of Ur, Egypt, 4300 years
ago. Fresh pollen is better than stored pollen for pollination.
Male spathes are traditionally cut a few days before or after
they split and then placed in a dry, shaded area for drying.
High temperatures, sunlight and moisture can cause
deterioration of date palm pollen. Strands are cut from the
spathes and stored in bags or containers so pollen that falls
from them will not be lost. If larger quantities of pollen are to
be extracted, the flowers are generally placed over a wire sieve,
which is then placed over a container for pollen collection and
storage. Mechanical pollen collectors are used sometimes.
Date palm pollen can be stored for several months at moderate
temperatures under dry conditions. Longer-term storage
requires stricter conditions. The pollen needs to be well dried
141
and placed in an airtight container, if possible with some sort
of desiccant. Storage at ‘refrigerator temperatures’ (4–5°C) is
usually done, but lower temperatures (–4 to –20°C) are also
appropriate (Nixon and Carpenter, 1978). Cryogenic storage
of date palm pollen for research is also possible. Growers
usually have preferred or special male trees that are selected
based on experience and some general properties. Early
blooming is valuable, since it assures a pollen source when the
females begin to flower. Size and number of male inflorescence
affects the amount of pollen produced. Individual flowers that
adhere to the strands are preferable, and the pollen content of
the individual flowers should be abundant. In areas where
inflorescence rot occurs (discussed under ‘Diseases, pests and
weeds’) pollen should be taken only from healthy males. Pollen
of the earliest and latest inflorescences may have lower quality.
The most common method of pollination used today is to
cut individual strands from male flowers and place two or
three of them within the strands of the female flower within a
few days after flowering. Twine is then tied around the female
strands to make sure that the male strands stay in place and
that the female strands do not become entangled. Pollination
of female flowers can be done before or after the spathes crack
naturally. The spathe can be split or removed artificially and
pollen can be applied to the inflorescence. If male flowers
cannot be used within a few days of collection, it is better to
use dried pollen. Dried pollen may be used undiluted or
mixed with a carrier such as flour or talc. The simplest method
is to coat a cotton swab with pollen, sprinkle this over the
female inflorescence, and then place the cotton ball within the
female strands. A modified technique utilizes hand-made
puffing devices with tubes for delivering the pollen from the
ground. Large-scale mechanical pollinators capable of
pollinating over 30 ha/season are in existence, but are used
mainly in highly developed production areas where labour is
scarce and expensive (Nixon and Carpenter, 1978).
Mechanical pollination is less efficient in terms of both pollen
used and fruit set. However, yield and quality can be
indistinguishable from hand-pollinated dates if thinning is
adjusted accordingly.
Pollination of 60–80% of the female flowers should result in
adequate fruit set. Covering the inflorescence with paper bags
after pollination can increase fruit set. Cultivars differ in the
length of time during which maximum fruit set can be
achieved, from up to 7 days before spathe cracking to 10 days
afterwards. Usually 2–4 days after the spathe opens is best.
Cultivars require different amounts of pollen for maximum
fruit set; usually two to three strands of male flowers are
adequate. Environmental conditions also influence fruit set.
Pollen grains have the highest germination rate from 25 to
28°C, and low temperature (< 21°C) can result in low fruit set.
Pollination is best achieved from mid-morning to midafternoon (10:00 a.m.–3:00 p.m.). Rain or dust storms before
or afterwards can reduce fruit set.
Cultivars differ in their fruit set percentages. For instance,
cultivar ‘Zahidi’ has a high fruit set percentage and ‘Hayany’
has a naturally low fruit set percentage. Incompatibilities or
partial-incompatibilities between different female and male
cultivars result in poor fruit set. The range of compatibility
has not been fully investigated. Pollen of other Phoenix species
are compatible with dates. Different pollen sources can
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Arecaceae
influence the size and shape of seeds (‘xenia’ effect). Different
pollens have direct effects on fruit size, fruit shape, yield and
ripening time. This pollen effect on tissue outside the embryo
and endosperm is called ‘metaxenia’. The ‘metaxenia’ can
reduce fruit ripening time from 10 to 60 days depending on
the growing season, resulting in more return because higher
prices are gained for earlier fruit. Also it may be beneficial if
total heat units are low to enable proper ripening and if rain is
avoided near the end of the season in some areas. Metaxenic
effects in date palms have been reported in many countries.
The fruit develops after
fertilization from one of the three carpels within each of the
pistillate flowers. If pollination or fertilization failed, all or one
of the three carpels can develop into parthenocarpic fruit. A
parthenocarpic fruit with three carpels is called a
‘parthenocarpic triplet’ (PT) and fruit with one developed
carpel is called a ‘parthenocarpic single’ (PS). The PT fruit
are hollow inside and PS fruit contain degenerated seeds. PS
fruit are larger than PT fruit, but smaller than seeded fruit.
All three types of fruit can be found on the same tree with
various proportions. Between 15 and 20 days after spathe
cracking, one of the three carpels begins to develop much
more rapidly than the other two carpels in pollinated flowers
or sometimes in unpollinated flowers. After 60–90 days,
depending on the cultivar, significant differences in fruit size
development can occur between seeded, PS and PT fruit.
Growth rate and development of seeded fruit show a sigmoid
growth curve. Most growth in size and weight occurs near the
end of the Kimri stage and at the beginning of Khalal. Most
fruit growth occurs 40 days after spathe crack and the fruit
reach maximum size about 120 days after spathe crack
(Reuveni, 1986). Natural fruit drop occurs 25–35 days after
spathe crack, when PT and PS fruitlets can drop. For cultivar
‘Deglet Noor’, about 50% of the flowers develop into mature
fruit. There are two distinct waves of fruit drop for ‘Deglet
Noor’. The first wave occurs 30–35 days after spathe crack
and lasts for about 1 month. The second wave starts about 100
days after spathe crack and also lasts for about 1 month. This
second wave of fruit abscission is sometimes referred as ‘June
drop’. The timing and duration of fruit drop vary by cultivars
and locations.
The date fruit goes through four distinct ripening stages.
Arabic terms, Kimri, Khalal, Rutab and Tamar, are used to
represent the immature green, the mature full coloured, the
soft brown and the hard raisin-like stages, respectively
(Reuveni, 1986). In the Kimri stage, fruit increases size and
weight rapidly until the Khalal stage. The colour changes from
green at Kimri to the characteristic mature colour at Khalal.
The rate of gain in size and weight during the Khalal stage
decreases slightly until the fruit reach full size and weight.
The fruit remain turgid, astringent and contain substantial
amounts of water-soluble tannins at Khalal. The fruit in the
Rutab stage are characterized by darkening of the skin to an
amber, brown or nearly black colour, accompanied by
softening, decreasing astringency and increasing insoluble
tannins. In the final ripening Tamar stage, the fruit lose much
of their water to a point where the sugar to water ratio is high
enough to prevent fermentation, similar to raisins or dried
prunes. The physical and chemical composition of the fruit
FRUIT GROWTH AND DEVELOPMENT
change during development and ripening. Water content is
75–80% in young fruit, dropping to 40–60% at the beginning
of ripening and decreasing rapidly afterwards. In general, the
sugar content is about 20% at early Kimri, increasing steadily
to about 50% at the beginning of Khalal, and finally
accumulating at a fast rate and reaching 68–88% of dry matter
at maturation.
Horticulture
The average date palm produces about 40 kg of fruit annually.
Cultivation with high input regularly produces 100 kg of fruit
annually but in underdeveloped regions date palm may
produce 20 kg fruit or less. The female plants start producing
dates at 4–6 years old and reach full production within 15–20
years. The average economic life of a date garden is 40–50
years; some can grow for 150 years.
Traditional date palm culture in North Africa and the
Middle East developed around oases or in riverbeds. Date
palms in oases are often of secondary importance to annual
crops or other tree crops grown beneath them. Date palms in
oases are usually derived from seedlings and off-shoots. The
oasis may be isolated and the seedlings may be quite inbred
and more uniform than trees that hybridized readily. Tree
spacing in these areas is often irregular. Trees are closer
together than in plantation-type settings. Off-shoots often are
not removed and trees retain multiple trunks. The date
growers in the traditional manner are often poor and lack
resources and manual labour is performed for all tasks.
Carpenter (1981) and Ferry (1996) cite the following problems
in traditional date culture: crowding of trees; retention of old
or unproductive trees; planting mixed varieties and/or
seedlings; salt accumulation; poor drainage; insufficient
irrigation, fertilization or tillage; lack of insect and disease
control; competition with other crops; soil degradation; and
water scarcity. Jaradat (2001) considered drought, high
salinity, aged trees, bayoud disease and genetic erosion as the
major constraints for date palm production worldwide.
If irrigation water is available, more industrial-style date
palm plantations are possible if climate and soils are
appropriate. Large-scale plantings of dates in the 20th century
are planted utilizing off-shoots or tissue culture (TC) derived
plants that are uniform. The common spacing is 9–10 m in
both directions, with planting density of 120 trees/ha. In
marginal climate areas, date palms are planted more closely
together. Date palms are generally a cash crop and receive
better care and greater inputs than in traditional oases. Most
literature involving cultural practices of dates were performed
in and aimed at this type of production, and it may not be
applicable to more traditional cultivation.
PROPAGATION Three methods are used for date palm
propagation. The most common method is the vegetative
propagation of off-shoots, which ensures the genetic identity
of maternal varieties. Off-shoots are developed from axillary
buds on the trunk near the soil surface. Dates produce offshoots when they are young and occasionally when they are
mature. Some varieties sucker more than others, but most
trees produce 10–30 off-shoots. After 3–5 years of attachment
to the parental palm off-shoots produce roots and are ready to
Phoenix
be removed. To promote rooting of off-shoots, the base of the
off-shoots should be in contact with moist soil for at least 1
year before cutting. The size and weight of the off-shoots
ready for cutting vary by variety, ranging from 10 to 30 kg in
weight and from 20 to 35 cm in diameter. The best time for
cutting off-shoots is after the soil begins to warm up in late
spring and early summer. Soil temperatures should be at least
20°C. In general, no leaves should be removed from an offshoot until it is cut from the parent palm. If a palm is crowded
with off-shoots, the leaves of smaller ones are sometimes cut
back close to the bud to slow the growth and the larger offshoots are removed first. The smaller off-shoots can be used
for subsequent cuttings. The soil is first dug away from the
off-shoots with a sharp, straight-bladed shovel. A ball of soil,
5–8 cm thick is left attached to the roots. For dry or sandy soil,
irrigation several days before cutting makes it easier to dig and
ball the off-shoots. A sharp chisel with the flat side facing the
off-shoots allows the connection between the off-shoots and
parental palm to be cut. Some pruning is always done after
removal of the off-shoots. The old leaf stubs and lower leaves
are cut off close to the fibre, the basal 0.6–1.5 m of the offshoot being left bare. Between ten and 12 leaves around the
bud are kept and tied close together. The terminal portion of
the leaves beyond the tie is cut off. The root ball should be
kept moist between cutting and planting. Balling with wet
burlap is often used with off-shoots that are to be shipped for
long distances. Most varieties can be planted at a 10 ⫻ 10 m
spacing. After planting, the off-shoots should be kept moist for
a few weeks by light, frequent irrigation. Best results in
planting off-shoots occur when medium or large, rather than
small, off-shoots are used.
The second propagation method is using chance seedlings
from sexual crosses. Seedlings are not identical to the maternal
trees and are not uniform genetically, varying greatly in their
production and fruit quality. Seedlings are also called ‘Khalts’,
‘Balady’, ‘Sairs’, ‘Deguouls’ or ‘Mantours’. About 50% of the
seedlings are male although they cannot be identified until
trees began to flower in 4–6 years. Date groves consisting of
‘Khalts’ are considered marginal. Production and fruit quality
from these marginal groves are greatly reduced, compared
with groves developed from off-shoots. Date seeds usually
germinate readily when planted in well-aerated soil at a depth
of 3–5 cm after the weather warms up in the spring. Seeds
may be planted either in nursery rows or directly in
permanent or semi-permanent locations. Two or three seeds
may be placed in each permanent location to ensure
germination and later all but one of the seedlings can be
removed. Date seeds stored in moderate temperature can
retain viability for at least 5–6 years.
The third date propagation method is through TC from
shoot tips, through either embryogenesis or organogenesis. TC
was first developed in the 1970s to 1980s. The massive
expansion of date palm plantations in Egypt, Saudi Arabia,
UAE and Jordan has led to the extensive use of TC-derived
date palms since there are insufficient off-shoots for
expansion. A TC programme was also initiated under the
threat of bayoud disease in Morocco. Organogenesis can be
achieved using auxiliary buds and apical meristems, while
embryogenesis can be done through a callus stage from various
meristematic tissues like shoots, young leaves, stem, rachilla,
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etc. Cultivars respond to TC differently, and different optimal
conditions are needed for each cultivar. It takes about 6 years
to reach production through the TC process and 8 years to
reach commercial quantities. TC has been used for in vitro
selection against the fungus responsible for bayoud disease. It
offers opportunities for mutation selection and genetic
transformation. In general, TC progenies have similar
characters as those derived from off-shoot propagation. These
include leaf morphology, vegetative characteristics, flowering
and fruit set, fruit physical properties (length, diameter,
dimension ratio, circumference, volume, weight, flesh weight,
flesh weight ratio, hardness and length), seed physical
properties (length, circumference, diameter and weight) and
fruit chemical composition (% moisture, % dry matter, pH,
% protein, % crude fibre, % ash, % pectin, % soluble solid
and % insoluble solid). The TC-derived progenies do tend to
revert into a more juvenile phase.
One of the main problems with TC propagation is
somaclonal variation (off-types). These somaclonal variants
exhibit several typical phenotypes including variegation in
leaves, variation in leaf structure and overall plant growth
patterns, trees that do not form inflorescences or produce
abnormal floral development, and trees that produce seedless
parthenocarpic fruit. Other somaclonal variants occurred
through the organogenesis TC method and have been reported
from countries such as Israel, Jordan, Morocco and Namibia.
These variants grow very slowly after planting with a stunted
appearance (dwarfism), lower number of leaves and a low rate
of total growth. Most somaclonal variants can be detected in
the early stages, however, some can only be detected in the
field, several years after planting or after flowering, fruit set
and maturation of the trees. In Saudi Arabia, dwarfism was
reported in 13.3% (of 1260 trees) of the TC-derived ‘Barhee’
cultivar and 20% (of 403 trees) of the TC-derived ‘Khalas’
cultivar, and 75.6% (of 2000 trees) of the mature ‘Barhee’
trees from TC failed to set fruit. Supernumery carpels (four,
five and six carpels compared with the normal three carpels)
occurred at frequencies of 7.8–16.9%, 2.4–7.7% and
0.7–3.5%, respectively. The frequency of somaclonal variation
in TC-derived date palms can occasionally be very high and
the mechanisms causing this variation are unclear and are
under investigation. In order to reduce the percentage of
somaclonal variation, organogenesis instead of embryogenesis
is used to generate date seedlings in UAE. Various methods
have been utilized to assess the genetic basis of this variation,
including representational difference analysis.
When applied to date palms,
the word ‘pruning’ has a somewhat different meaning than
when applied to deciduous fruit trees. The commonly practised
pruning of date palms is the removal of dead and dying leaves.
This is often done after fruit harvest or in synchronization with
other cultural operations such as tie-down or bagging. Dry,
dead leaf bases are difficult to cut and it is easiest to remove the
senescing leaves before they dry out completely. Sometimes
dead leaves are retained to provide some cold protection. In
addition to the removal of senescent leaves, the basal spines are
removed from the previous year’s leaf growth during the
winter months to facilitate pollination and bunch management
operations and prevent injury to workers.
PRUNING AND FRUIT THINNING
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Arecaceae
In general, living green leaves are kept as the productivity of
date palms is related to the number of leaves retained. An
insufficient number of leaves can reduce fruit quality in the
current season and fruit set and yield the following season.
Date palm leaves remain alive for 7 or more years, and do not
normally fall off. If trees are left un-pruned, an excessive
number of leaves may exist and this increases the relative
humidity (RH) in the fruit zone of the tree’s crown and the
incidence of disorders. An excessive number of leaves may also
increase water stress. In varieties with long fruit stalks,
removing leaves up to the point where the lower portions of
the bunches are exposed is the preferred practice; for varieties
with shorter fruit stalks this results in the removal of too many
leaves for optimal production. With these shorter-stalked
varieties, relatively few leaves should be removed or bunch
removal should be performed in order to retain an adequate
leaf:bunch ratio. This ratio is about eight to ten leaves per
bunch when normal thinning practices are followed. Older
date palms (20 years old and above) usually do not retain green
leaves below the fruiting zone.
Fruit thinning is often practised in date culture and has a
number of beneficial effects. Fruit thinning ensures adequate
flowering in the following season, lessens the chance of a small
crop and reduces alternate bearing tendency. Fruit thinning
also increases the size of the fruit (especially important in
‘Medjool’ and other large-fruited varieties), improves fruit
quality and advances ripening. Bunch compactness and weight
are reduced by thinning, thus facilitating bunch management
operations later in the season. Fruit thinning in date palms is
carried out in three ways: removal of entire bunches;
reduction in the number of strands per bunch; and reduction
in the number of fruit per strand.
Reducing the number of strands per bunch or the number
of fruit per strand is referred to as ‘bunch thinning’. It is
recommended that bunches be uniformly thinned by 50–75%
of the normal load of flowers (Nixon and Carpenter, 1978).
The procedure is slightly different for long- and shortstranded varieties. In long-stranded varieties, such as ‘Deglet
Noor’ the lower third of all the strands in the bunch is
removed. Ideally, the total number of flowers in an average
strand should be counted in order to make this reduction more
precise. Strand length reduction has its maximum effect when
done at the time of pollination. Removal of entire strands from
the centre of the bunch to reduce the total number of strands
by 33–50% is also practised and can be done at the time of
pollination, but is normally delayed until the cluster has
emerged further. The beginning of thinning is delayed until
6–8 weeks after pollination so that set can be observed and the
thinning ratio adjusted appropriately. Short-stranded varieties,
like ‘Khadrawy’, need to be thinned slightly differently. Only
10–15% of the strand length should be removed, while
removal of the centre strands increases to 50% or more. In
some cases, only centre strand removal is used on shortstranded varieties. Timing for these operations is the same as
for long-stranded varieties. Carpenter (1981) states that
thinning should result in 1000–1500 fruit/tree. Over thinning
however, can result in puffing and blistering of the skin, and
cutting back strands can increase the incidence of checking,
blacknose and shrivel. Variety, climatic conditions and cultural
practices all influence the appropriate level of thinning. The
individual grower should observe and record this information
to assist in long-term management of the date garden.
Removal of individual flowers or fruit from the strands is
slightly more effective in increasing fruit size than reducing
the length or number of strands. However, fruit removal is less
practised due to the cost. Fruit removal is time consuming and
so labour costs are high. It is practised mainly with ‘Medjool’
where the premium price obtained for large, high quality fruit
justifies the added expense.
Removal of entire fruit bunches is sometimes practised to
reduce the number of bunches to an appropriate level. This
number is dependent upon the age, size, vigour, variety and
number of leaves on the date palm. Some of these factors may
be influenced by cultural practices such as irrigation and
fertilization. In order to establish date palms, all bunches
should be removed from off-shoots for the first 3 years after
planting. Thereafter, the number of bunches allowed may be
increased by one or two per year. Date palms reach full
production at 10–15 years of age, at which time they may
support 10–15 bunches of dates. Mature date trees may carry
up to 20 bunches if bunch removal is not practised. This can
result in poor crops or alternate bearing. Certain bunch classes
are higher priority for removal: early and late bunches, which
are usually small and poorly pollinated; bunches with poor
fruit set or with broken stalks or other structural damage; and
excess bunches on one side or quadrant of the tree. In the
USA, mature ‘Deglet Noor’ trees with 100–120 leaves are able
to produce 12–15 moderately thinned bunches without
alternate bearing; this results in eight or nine leaves per bunch
(Nixon and Carpenter, 1978).
After pollination, bunches are usually tied to the leaf stalks.
This is done by carefully pulling the bunch through the leaves
below the level of the bunch and tying the fruit stalk to the
midrib near the base of the leaf. This should be done when the
fruit stalk is near the end of elongation but is still pliable. Tie
down prevents breakage of the leaf stalk from the weight of
the fruit, reduces damage from wind or high-pressure spray,
and facilitates other bunch operations. It is not usually
necessary until the fruit is about 75% of its final weight. This
practice is easier with long-stalked varieties. Varieties with
short fruit stalks are sometimes tied to leaves adjacent or even
slightly above the bunch (Nixon and Carpenter, 1978).
Bunches of dates are commonly covered (bagged) in the
USA. This practice has several advantages. Primary among
these is protection from high humidity and rain, which can
cause the physiological defects of checking, blacknose and
splitting (discussed under ‘Diseases, pests and weeds’) and can
also result in rot and souring from secondary pathogens.
Bagging can also reduce losses from birds and minimize
sunburn. Brown craft paper is generally used, although in
some instances white paper has been found to reduce sunburn.
More recently, cotton or nylon mesh bags have been used;
these have the advantage of superior ventilation. It is
important that the bags be open enough that humidity does
not build up within the bag, therefore plastic bags should not
be used. Bunches are usually bagged at the Khalal or late
Kimri stage. In some countries, mesh bags are used to exclude
at least some insects (Carpenter, 1981). Sometimes, bunches
are spread open with wire rings prior to Khalal to reduce
humidity-induced problems (Nixon and Carpenter, 1978).
Phoenix
NUTRITION, FERTILIZATION AND IRRIGATION Date palms
require a large amount of water for vigorous growth and high
yield of good quality fruit. However, they are able to withstand
long periods of drought under high temperatures. Date palms
under drought conditions are stunted and unproductive.
Productivity is determined by the quality and reliability of the
water. Date palms in oases or riverbeds may have some
irrigation management by the construction of basins, borders
or canals. Flood irrigation is the oldest form of irrigation and
is still utilized in many areas. Furrow and basin irrigation are
also quite old in application. These methods are inexpensive
and easy to apply, but they are not efficient in water usage and
are labour intensive. Sprinkler irrigation is the oldest ‘modern’
method of irrigation, and results in a more efficient but
expensive use of water. For young date palms, use of
traditional sprinklers could place water onto the growing point
and cause damage. Recently, micro-sprinklers and drip
irrigation have been utilized in date palm plantings. These
modern irrigation practices allow more precise management of
the amount and placement of the water.
When flood, furrow or basin irrigation is used, bearing date
gardens are usually irrigated every 1–2 weeks during the
summer and every 3–4 weeks during the winter. Frequency
depends upon soil texture and weather conditions. In the main
US production area of the Coachella Valley, 220–300
m3/tree/year are required for mature, producing date palms,
with 24–36 m3/tree/month being needed during the summer
(Nixon and Carpenter, 1978). Consumptive water use by
mature date palms was approximately 104–168 m3/tree/year,
with the highest monthly use being about 15–25 m3/tree in
July. Short stature cultivars such as ‘Khadrawy’ use less water
than other cultivars. Mature trees of various varieties were
reported to use approximately 144 m3/tree/year in central
Iraq. Dowson (1982) notes work by various investigators that
list annual water use per date palm in the range of 138–364 m3
in numerous growing regions. Crop coefficients for estimating
evapotranspiration are not well established for date palms. The
Food and Agriculture Organization (FAO) estimate was 0.95
(Allen et al., 1998). A study done in UAE developed crop
coefficients ranging from 0.66 to 0.90, depending upon the
time of year, while in Iraq crop coefficients were reported to
be 0.75–1.00, with a seasonal average of 0.85.
Responses of date palms to irrigation were based upon the
observation that growth (as reflected by the rate of elongation
of the central unexpanded leaf) reflected soil water depletion.
Using this observation as an index, soil water limits date palm
growth about 4 weeks after irrigation during the summer
months, with soil water potential being about ⫺0.08 MPa at a
75 cm soil depth. When irrigation was withheld to the point
where there was a 15–20% reduction in the rate of leaf
elongation during the summer, fruit size was reduced, fruit
moisture content was decreased, and fruit ripening was earlier.
With ‘Maktoom’, a soft cultivar, under moderate rainfall,
limiting irrigation reduced shrivel and blacknose with no
reduction in size, grade or yield when water stress occurred
during the harvest period. A slight reduction in growth of the
tree occured under water stress, but the number of leaves and
inflorescences was not affected. Excess amounts of water did
not increase tree growth or fruit yield and quality.
Fertilization is needed to sustain production and fruit
145
quality of date palms. However, little information is available.
Many date producers follow the traditional fertilization
practices, which vary by region within countries. In traditional
date gardens, fertilization is often done by application of
animal manure, with chicken manure being preferred due to
its higher nitrogen (N) content. Although use of inorganic
fertilizers is common, manure is sometimes used in
plantation-type production of dates at a rate of 11–34 t/ha.
Manure is generally applied in the autumn or winter, or in the
spring after removal of cover crops (Nixon and Carpenter,
1978; Dowson, 1982).
Most date palm fertilization research has involved N or
other macro-elements. Furr et al. (1951) reported that ‘Deglet
Noor’ date palms receiving N fertilization over a 7-year period
showed no response to fertilization during the first 3 years, but
showed increased growth and yield over the last 4 years. The
N content of pinnae and other tissues was significantly higher
in the fertilized trees, and generally correlated with yields.
However, in a different experiment, ‘Khadrawy’ palms did not
respond to fertilization and there were no differences in
mineral content of tissue. Young, non-bearing ‘Medjool’ date
palms did not respond to N fertilization. In Algeria, no
response to fertilization was found because all mineral needs
of the date palms were supplied in the irrigation water.
Conversely, in Qatar, date palms do respond to fertilization
and yield was positively correlated with leaf N content and
inversely with leaf potassium (K) content. In Saudi Arabia,
heavy N fertilization increased date yields but decreased
quality as compared to a more moderate rate of fertilization.
Responses of date palms to fertilization are dependent upon
growing conditions, including soil texture and pH and other
cultural practices. Date palms remove a certain amount of
nutrients from the soil each season, mostly from fruit and leaf
pruning. In Algeria 72 kg of N, 5 kg of phosphorus (P) and 27
kg of K were removed each year from 120 date palms (1 ha).
In Saudi Arabia, 1 ha of date palms was estimated to lose 56 kg
N, 6 kg P and 50 kg K annually. Lower losses of N and P were
reported in the USA: 25 kg N and 2 kg P; however, loss of K
was greater at 74 kg. Up to 78 kg N was reported to be lost
annually in California. These values are taken by multiplying
the mineral concentrations in leaves and fruit by yield and
pruning mass and do not take into account nutrients that
would be necessary to support annual growth of leaves and the
permanent structure, nor soil-associated losses. A
comprehensive study establishing appropriate nutrient levels
in date palms has not been conducted.
DISEASES, PESTS AND WEEDS The date palm is affected by
many diseases, pests and weeds but the incidence of a
particular problem varies with the area and cultural practices.
Reports of disease and pest outbreaks in specific countries or
locales may be found in technical journals and on the worldwide web (e.g. the Plant Protection and Pest Information
Service of FAO). There have been no reports of viral or viroid
diseases of date palms so far; only fungus- and phytoplasmacaused diseases as reviewed by Elmer et al. (1968), Carpenter
and Elmer (1978), Djerbi (1983) and Zaid et al. (2002). Much
of the following is summarized from those references. A
concise list of reported diseases (as of the date of publication)
is given in Carpenter (1991). A list of diseases and disorders of
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Arecaceae
date palms is in Table A.60. Most diseases of date palms are
fungal pathogens, some disease are caused by phytoplasmas.
The common control of most diseases of date palms include
sanitation practices, use of clean off-shoots or materials from
tissue culture, quarantine, reducing moisture by fruit
thinning, bagging fruit and changing irrigation practices.
A list of reported insect pests of date palms is in Table A.61
and reviewed by Howard et al. (2001). Most insect pests of dates
are controlled by chemicals, biological control, pheromone
trapping, quarantine and sanitation practices. Insect pests
attacking stored fruit are controlled by chemical fumigation.
Rodents can be pests of dates. Various types and species of
rodents attack date palms. Rodents may eat date fruit either on
the tree or in storage. Some also eat the inflorescence.
Underground-dwelling rodents damage roots and also interfere
with irrigation. Control of rodents is by trapping or poisoning.
Many weed species are associated with date palms around
the world. The composition of the weed flora varies with
Table A.60. Common names, causal organisms or factors, symptoms and countries/region where date palm diseases and disorders were reported.
Common names
Causal organisms or factors
Symptoms
Reported countries/region
Bayoud
Fusarium oxysporum f. sp. albedinis
Attacks vascular system, causes browning and
necrosis of the fronds, eventually kills the plant
Algeria, Morocco
Inflorescence rot
Mauginiella scattae
Infection of spathes, destruction of inflorescences
North Africa and Persian Gulf
region
Black scorch (fool’s disease) Thielaviopsis paradoxa
Necrosis of leaves, inflorescences, trunk, vascular
system, and terminal buds
Graphiola leaf spot
Graphiola phoenicis (Moug) Poit.
Infection occurs in sub-epidermis of leaves and
later emerges as small, black spots
Diplodia disease
Diplodia phoenicum (Sacc) Fawc. &
Klotz
Attacks leaves of off-shoots and terminal buds, and
results in death of off-shoots
North and Central Africa, Persian
Gulf region, USA
Brown leaf spot
Mycosphaerella tassiana (De Not) Johns Necrosis of leaves
North Africa
Omphalia root rot
Omphalia spp.
Premature leaf death, reduced growth and decline,
necrotic roots
USA
Belâat disease
Phytophthora spp.
Decline and death of young leaves and death of
terminal bud
North Africa
Pre-harvest fruit rot
Alternaria spp., Sepergillis spp.,
Helminthosporium spp.,
Macrosporium spp.
Fruit rot begins at Khalal through Rutab and Tamar
stages
Al wijam
Mycoplasm-like organism
Decline and death starting from terminal bud
Saudi Arabia
Brittle leaf disease
Mycoplasm-like organism
Chlorotic and dry leaves, and stunting
North Africa
Lethal yellow
Mycoplasm-like organism
Desiccated, greyish-brown leaves, rotting of
terminal bud, loss of tree crown
White-tip dieback
Phytoplasm organism
Dry bone
Unknown
Irregular white blotches and streaks on leaves,
later with reddish-brown margins
North Africa
USA, North Africa
Forun disease
Unknown
Rapid and fatal decline in 3–5 years, starting with
auxiliary bud abortion, stunting of young leaves,
flattering stature of older leaves
Mauritania
Rhizosis/rapid decline
Unknown
Fruit drop or shrivel on the bunch, pinnae of older
leaves become discoloured, young leaves wilted,
death of the fronds
USA
Bending head
Unknown
Central cluster of fronds forms a fascicle with a
bent tip, death of older leaves, necrosis of terminal
bud and a heart rot, trunk bends or breaks
North Africa and Mauritania
Barhee disorder/bending
barhee syndrome
Unknown
10-year-old trees lean to the south at 5–90° angles, Iraq, Israel, USA
reduction in fruiting bunches
Blacknose
Physiological disorder?
Shrivelled and darkened tip of the fruit
Egypt, USA
Iraq, Israel, Pakistan, USA
Crosscuts
Anatomical defect
Breaks at the base of fruit stalks and leaf petioles
Whitenose
Caused by dry wind
Dry wind at early Rutab cause rapid maturation and Iraq, North Africa
drying of the fruit
Black scald
Unknown
Well-delineated blackened areas on the sides and
tips of the fruit
Root-knot nematode
Meloidogyne spp.
USA
Algeria, USA
Phoenix
147
Table A.61. Common names, causal organisms, symptoms and countries/region where date palm pests were reported.
Common names
Causal organism
Description
Reported countries/region
Parlatoria or white scale
Parlatoria blachardii Targ.
Presence in foliage and fruit, fed on leaf stalk basal
tissue, death is rare
All date producing areas except
USA
Red date scale
Phoenicococcus marletti (Ckll)
Symptoms similar to parlatoria scale, un-thriftiness All date producing areas
look in severe infection, small losses
Green scale
Asterolecanium phoenicis
(Ramachandra Rao)
Heavy infestation results in fruit scarring and
economic losses
Egypt, Israel and Persian Gulf
areas
Red palm weevil/Indian
palm weevil
Rhynochophorus ferrugineus Oliv.
Infestation initiates through crown of the palm and
ends in apical bud, palms collapse and die from
heavy infestation
India, Pakistan, Egypt and Persian
Gulf areas
Palm stem borer
Pseudophilus testaceus Gah.
Infestation through trunks and leaves, extensive
damage could happen
Egypt, United Arab Emirates and
Persian Gulf areas
Boring beetles
Oryctes spp.
Feed on tender leaves, spathes and apical buds;
adult insects may kill the tree
Fruit stalk borer
Oryctes elegans Prell
Mining by the borer weakens the tree and results
in frond breakage
Saudi Arabia, Middle East
Nitidulid beetles
Carpophilus dimidiatus F.,
C. hemiperus L., Urophorus humeralis
F., Haptoncus luteolus Erich.
Fruit damaged during ripening and curing stages
on the trees, on the ground or in storage
USA
Carob moth
Ectomyelois ceratoniae Zeller
Larvae attack dates in the field, packing houses
and in the market
Asia, North Africa and
Mediterranean areas of Europe
Indian meal moth
Plodia interpunctella Hbn.
Larvae feed on ripe dates on the tree or postharvest Algeria, Israel, Libya and USA
Greater date moth
Arenispes sabella Hmpsn.
Attack spathes and fruit stalks that results in
dehiscence of the bunch
Raisin moth, almond or
fig moth
Cadra spp.
Attack stored and ripening dates
Desert locust
Schistocera americana gregaria Forsk.
Feed on leaves and fruit; may destroy entire crops
and consume entire canopy
Termite
Microcerotermes diversus Silv. and
other species
Attack roots, trunks and leaves; weakened trees
may collapse; off-shoots may be killed by termites
Old world date mite (Bou
faroua or Goubar)
Oligonychus afrasiaticus McGr.
Abraded and discoloured leaves and fruit after
infestation, webbing can cover fruit bunches in
heavy infestation causing premature fruit drop and
a decrease in fruit quality
North Africa, Middle East, Iran and
Saudi Arabia
Banks grass mite
Oligonychus pratensis Banks.
Damage is similar to old world date mite
North Africa, Middle East and USA
locale, cultural practices and season. Some common weeds
found in date palm plantations are halfa (Imperata
cylindricum), Bermuda grass (Cynodon dacytlon), nutsedges
(Cyperus spp.), Chenopodium spp., Juncus spp. and Johnson
grass (Sorghum halapense). Although weeds have often been
reported to cause more economic losses than other pests, their
deleterious effects – and therefore their control – are often
overlooked. In some cases, the difference between a weed and
a cover crop is not obvious. In addition to their competitive
effects, weeds can also serve as alternate hosts for insects and
pathogens. Weed control, when implemented, may be
mechanical, cultural or chemical.
HANDLING AND POSTHARVEST STORAGE In North Africa and
the Middle East, some dates are harvested at the Khalal stage
when they are yellow or red, depending on the cultivar. Most
American and European consumers find them astringent (high
tannin content). Most dates are harvested at the fully ripened
Rutab and Tamar stages, when they are high in sugar content
India, North Africa, Middle East
North Africa and Middle East
and low in moisture and tannin. The quality of the fruit is
determined by fruit size, colour, texture, cleanliness and
freedom from defects (sunburn, insect damages, sugar
migration to fruit surface and fermentation) and decaycausing pathogens. Dry dates and some semi-dry types are
usually harvested once, at full maturity. Soft dates and some
semi-dry types are harvested on several occasions during the
season due to uneven ripening. Ripening is usually uniform
through a bunch, so it is more economical to harvest entire
bunches rather than individual fruit. Labour costs sometimes
dictate frequency of harvest, even though this lowers the
overall quality of the product. Most date harvesting is done
manually. Pickers climb the trees assisted by climbing gear or
ladders; usually, there is a belt that can be clipped to several
fronds to support the picker. More recently, hydraulic lift
platforms have been used to gain access to the crowns of date
palm trees. In some cases, hydraulic lifts are used for bins but
pickers scale the trees in the traditional manner. The fruit are
then separated from the bunches by a shaker. This type of
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Arecaceae
mechanically assisted harvest is best suited to drier types of
dates. A machine that shakes an entire tree in order to harvest
dates has not yet been successfully demonstrated.
In many traditional date-producing areas, dates are still
hand sorted in the field and then sold at local markets or by
other means. More industrialized date production relies upon
packing and storage facilities and to hold them until shipped.
In many cases the equipment used in date packing houses is
modified from that designed for other large-volume crops.
Date packing operations may be private or cooperative, may
range in size from small to large, and may be modern or less
so. Within the packing house, there are a number of
procedures used to ensure the quality. Fumigation, although
sometimes done in the field, is generally the first treatment
that dates are subjected to upon arrival at the packing house.
By the 1960s and 1970s, methyl bromide had emerged as the
fumigant of choice (Rygg, 1975; Nixon and Carpenter, 1978).
However, environmental concerns have led to a proposed
phase-out of methyl bromide, and use of other treatments.
Alternative fumigation treatments used on dates in the past
have included phosphine, carbon disulphide, hydrocyanic acid
and ethylene oxide. Some of these compounds are quite toxic
and may have their own detrimental environmental effects.
Other alternatives include physical filtering, irradiation and
controlled atmospheres (Glasner et al., 2002).
After fumigation, the dates may be cleaned. In small
operations, this can be done with damp towels attached to
shakers; larger operations use washing devices with revolving
brushes or jets. They are then dried with hot air or placed in a
dehydration room. After cleaning, the dates are sorted to
remove culls and to separate the fruit into uniform lots.
Sorting into size categories may be done mechanically but for
quality standards it requires human sorters. Industrialized
date production generally has to adhere to governmentally
mandated grade standards based on factors such as colour,
uniformity of size and physical appearance. Dates are packed
into various types of containers for storage, shipping and
marketing. Perhaps most common are cardboard boxes or flats.
In poor or marginal date-producing areas, dates may be packed
into whatever container is available or marketed in bulk. In
some countries, whole bunches are packed into containers.
As with many commodities, economics dictate that some
date fruit be stored. The higher the moisture content of the
dates, the more perishable they are. Very dry dates can be
stored without refrigeration. Refrigeration is necessary for
long-term storage of most dates. The optimum storage
temperature is at 0°C for 6–12 months, depending on the
cultivar. Semi-soft dates like ‘Deglet Noor’ and ‘Halawy’ have
a longer storage life than soft dates like ‘Barhee’ and
‘Medjool’. For long-term storage 18°C can be used. These
lower temperatures are also better for soft types of dates.
Generally, stored dates should be maintained at 20–25%
moisture content for optimum quality. Maintaining this
moisture content requires tight control over temperature and
storage humidity. The optimum RH for storage of dates is
about 70–75% when stored at room temperature (24°C),
increasing to 80–85% for storage at 0°C. Soft dates require
slightly lower storage humidity than semi-soft and dry types to
retain their moisture content. Dates are not a chilling sensitive
commodity (Kader, 1992). A few artificial processes can be
utilized to increase the quality of dates, including artificial
ripening, dehydration and hydration. Artificial ripening (also
called ‘curing’) is not practised to a large degree, but is
sometimes used when dates are picked at a less than fully
mature state. This may occur in regions with marginal heat
accumulation; when conditions are such that the Tamar stage
occurs very quickly after Khalal, so that some fruit are at
Khalal while others are shrivelling; and when fruit are picked
at Khalal for other reasons. Artificial ripening is done in
specialized rooms where temperature and humidity are tightly
controlled. The dates are held at high temperature (35–46°C)
and humidity (70–80% RH) for a period of several hours to
several days. The actual conditions depend upon variety (soft
varieties requiring higher temperatures), state of maturity and
other factors, and generally a great degree of experience is
required for successful artificial maturation (Rygg, 1975).
Semi-dry and soft date types must often be dehydrated unless
they are consumed immediately or stored at very low
temperatures. Temperatures are lower than for artificial
ripening (c.30°C), as are levels of RH. Air movement is
important, and RH should not fall below 50%. Dehydration
times range from several days to a week or more. The actual
amount of dehydration depends upon the objective, long-term
storage needing a greater degree of dehydration (moisture
content of 23–30%, or lower in some cases) than for
consumption at a fairly close date (moisture content of
30–35%). An alternative for soft varieties is to cool them to
⫺18°C for storage immediately after cleaning (Rygg, 1975).
Hydration is necessary when dates become overly dry due to
unusually hot and dry climates, inadequate irrigation or
delayed picking. The moisture content of such dates needs to
be increased for consumer acceptance. Hydration is
accomplished by saturating the fruit with water or steam.
Commonly in California, hydration consists of holding
‘Deglet Noor’ dates at 60°C in live steam (0.36 kg/cm2) for
4–8 h. Conditions vary somewhat in other countries (Rygg,
1975).
Dates are a non-climacteric fruit with a very low respiration
rate (Kader, 1992) thus their deterioration rate is lower than
many other commodities. However, there are some conditions
that contribute to deterioration. Physiological deterioration in
dates includes both oxidative and non-oxidative darkening and
sugar spotting. Other physiological conditions that are not
deterioration per se are considered defects or blemishes and
are discussed in the previous section, ‘Diseases, pests and
weeds’. Spoilage caused by yeasts is usually due to
Zygosaccharomyces spp. or Hansenula spp. as these genera are
most tolerant of high sugar contents. Moulds and bacteria are
considered less important in date deterioration (Rygg, 1975).
As mentioned in the section ‘Uses and nutritional
composition’, dates may be processed into various products,
including pitted pressed dates, date paste, pickles, syrups,
sugar, alcohol, candies, etc. For more extensive information on
methods of processing, the reader is referred to Barreveld
(1993). Further information on other aspects of date packing
can be found in Dowsen and Aten (1962), Rygg (1975) and
Glasner et al. (2002).
MAIN CULTIVARS AND BREEDING Nomenclature of date palm
cultivars is very confusing because of the long history of
Phoenix
cultivation, wide exchanges of date palm germplasm, dioecism
and seedling propagation. Large numbers of synonyms and
homonyms exist from country to country for many cultivars.
The same cultivar sometimes may have different names from
oasis to oasis. Furthermore, transliteration of Arabic names
into other languages can further confuse the issue. Different
genetic marker systems have been used to study the genetic
diversity and relationships among date palms, including
morphology, isozymes, restriction fragment length
polymorphisms (RFLP), random amplified polymorphic
DNA (RAPD), amplified fragment length polymorphism
(AFLP) and Representational Difference Analysis.
Worldwide, there are over 3000 cultivars of which some 60
are widely grown throughout the major date-growing
countries. Hundreds of cultivars have been reported in
different countries, particularly in North Africa and the
Middle East. For example, over 450 date cultivars have been
reported in Iraq, over 800 in Algeria and over 220 in Morocco.
Most cultivars currently grown have resulted from thousands
of years of selection from chance seedlings. Descriptions of
additional cultivars are available in Nixon (1950); however,
these are somewhat biased towards those varieties that have
proven useful in the USA. Nixon (1950) lists most of the early
accounts by US Department of Agriculture (USDA) plant
explorers for further reference. Other literature regarding date
cultivars is somewhat limited. Some of the leading date
cultivars are briefly described below.
‘Deglet Noor’, meaning ‘date of the light’ in Arabic, also
known as ‘Deglet Nour’, ‘Deglet Nur’ or ‘Deglet Nuur’, is
believed to have originated near Touggourt in the Algerian
Sahara in the 17th century. It is a dry or semi-dry date and
very popular in European and US markets. It is commonly
grown in Algeria and Tunisia. Fruit have a very attractive
appearance, turning light red in Khalal, amber brown in Rutab
and light brown in Tamar, and a distinctive and delicate
flavour. Fruit size is 40–50 ⫻ 20–25 mm, with medium thick
skin adhering to the 4–5 mm thick flesh, and with a medium
brown seed. ‘Deglet Noor’ performs best in relatively light soil
underlain by loam or silt soil, and in a relatively hot and dry
environment. Weather requirements limit the production of
‘Deglet Noor’ to northern Africa and California; it is not
possible to grow it in the hot and humid Persian Gulf region.
‘Medjool’, meaning ‘unknown’ in Arabic, also known as
‘Medjhool’, ‘Medjehuel’, ‘Mejhul’, ‘Mejhoul’, ‘Tafilalet’,
‘Tafilelt’ or ‘Tafilat’, is believed to originate from the Tafilalt
district of Morocco. It is a large soft date widely accepted by
the market. The fruit has an orange-yellow colour with a fine
reddish-brown stippling at Khalal, ripening to an amber
colour, turning into reddish brown, and more or less
translucent in appearance at Tamar. Fruit size is 38–60 ⫻
26–32 mm, with medium thick skin adhering to 5–7 mm thick
flesh and a dark brown seed. The fruit is moderately soft and
mildly rich in flavour. The ‘Medjool’ in Morocco was nearly
wiped out by bayoud disease in the early 1900s. The USA and
Israel are the major producers of ‘Medjool’. Due to the
superior quality of this cultivar, it is being produced in large
numbers by tissue culture and its importance as a cultivar is
therefore increasing worldwide.
‘Barhee’, also transliterated as ‘Barhi’, ‘Berhi’ and ‘Birhi’, is
a name of uncertain meaning, but possibly is associated with
149
the hot ‘barh’ winds that occur near Basra, Iraq, its place of
origin. ‘Barhee’ is a soft date of high quality. Due to its low
astringency during the Khalal stage, it is one of the major
cultivars marketed at that stage. The fruit is yellow at Khalal,
turning to amber at Rutab and amber to reddish brown at
Tamar. ‘Barhee’ dates are 32–37 ⫻ 23–30 mm, with medium
thick skin. ‘Barhee’ palms are robust with stout trunks but few
off-shoots. Like ‘Medjool’, the high quality of ‘Barhee’ fruit
has made it one of the most widely utilized cultivars for
production by tissue culture.
‘Deglet Beida’ (‘Degla beida’, ‘Daqlah Baydahi’) originated
in Algeria. The fruit ripens early in the season and is a dry
type, medium sized, 37–45 ⫻ 20–23 mm, oblong or narrowly
oblong with skin moderately thick and relatively smooth,
yellow when immature, very light pale brown or buff at
maturity and when cured. The flesh is firm and the flavour is
good. The tree is of medium vigour with a medium-heavy
trunk and is said to be quite salt tolerant.
‘Halawy’, also transliterated as ‘Halawi’, ‘Hallawi’ and
‘Hellawi’, is an important Iraqi cultivar. The fruit is semi-dry,
35–45 ⫻ 17–20 mm, oblong, thin skinned, with soft flesh and
a rich flavour. Colour is yellow at Khalal, darkening to a deep
translucent brown at Tamar. Yields are good at 70–90 kg/tree.
‘Hayany’ (‘Hayani’) is from Egypt and shows prolific offshoot production. The early ripening fruit is large, 45–55 ⫻
22–28 mm, oblong-elliptical, with medium thick skin that is
deep red when immature, but almost black at maturity. The
flesh is soft and watery with a mild flavour, lacking distinct
quality. Trees are of medium vigour with a slender trunk and
moderately arched leaves. It produces good yields (110–140
kg/tree).
‘Khadrawy’, also transliterated as ‘Khadrawi’, ‘Khadhrawi’
and ‘Khudrawee’, is a distinctive cultivar from near Basra,
Iraq. It grows slowly and mature palms are considerably
shorter than other cultivars. The fruit is soft, oblong-ovate,
33–40 ⫻ 20–24 mm, with skin medium thick, yellow at
Khalal, turning amber at Rutab and reddish brown at Tamar.
The ‘Khadrawy’ has good quality fruit with a rich but not
cloying flavour, but it is only moderately productive.
‘Khalasa’ (‘Khalaseh’, ‘Khalasi’, ‘Khalas’, ‘Khulas’, ‘Khlas’)
is from Arabia. The fruit is a semi-dry type, 30–40 ⫻
19–23 mm, oblong-oval, with a thin skin that is yellow when
immature, and amber to reddish brown at maturity. The flesh
is very tender and melting and the flavour is rich and delicate.
The fruit ripens mid-season. The tree is moderately low in
vigour and the trunk is medium heavy.
‘Rhars’ (also called ‘Ghars’) originated in Algeria. The very
early ripening fruit is a soft type; large, 45–55 ⫻ 20–24 mm,
and oblong-ovate. The skin is medium thick and medium
tough, yellow when immature and amber to reddish brown at
maturity. The flesh is soft and melting; the flavour is rich and
sweet but rather cloying, of good quality but very susceptible
to checking from rain or high humidity. It yields 90–115
kg/tree when losses from checking do not occur.
‘Samany’, also transliterated as ‘Samani’ and ‘Samiani’, and
having the synonym ‘Rashedi’, originated in Egypt. The fruit
ripens mid-season and is a soft type, very large, 50–60 ⫻
25–35 mm, and oblong-ovate. The skin is thin and tender,
yellow when immature, dull amber to brown at maturity. The
flesh is soft but rather coarse with a mildly sweet flavour early
150
Arecaceae
in the season, becoming rather insipid later. Fruit quality of
‘Samany’ is good early in the season (Khalal, early Rutab) but
disappointing later.
‘Zahidi’, also known as ‘Zahdi’, ‘Zadie’, ‘Zaydi’, ‘Zehedi’ or
‘Zaheedy’ originated from northern Iraq and is widely grown
in Iraq. ‘Zahidi’ has a compact crown. The fruit ripens in midseason and has a distinctive obovate shape. It is yellow in
Khalal, with the softer portions turning light brown and the
drier basal portions fading to yellow or straw colour in Rutab,
and in Tamar the softer portions turn reddish brown and the
drier portions light brown. Fruit size ranges from 34 to 40 ⫻
23 to 25 mm, with thick and tough skin adhering to the
4–5 mm thick flesh, with a large greyish-brown seed. The
fruit is semi-dry, with no special flavour.
The long life cycle, juvenility period and dioecism of date
palms make breeding a challenge. The determination of the
sex ratio in sexual progenies is believed to be controlled by a
single gene. In general, progenies are segregated for a 50:50
female to male ratio. A sexual chromosome with nucleolar
heterochromatin has been identified that might be used for sex
determination. So far no molecular markers linked to sex
expression of date palm have been identified. Empirical
selections at the local level are common for choice clones from
chance seedlings, and are subsequently clonally propagated
from off-shoots. A backcross-based breeding programme
spanning over 30 years was initiated at the USDA Date and
Citrus Station at Indio, California, USA in 1948. Due to
problems of sterility and low vigour from inbreeding
depression, no useful backcross progenies were produced.
Moreover, the female cultivars produced, while having some
interesting or desirable characteristics, have not proven more
useful than the established commercial cultivars. A similar
programme was carried out starting in 1943 at the Institut de
Technologie et de Developpement de l’Agriculture
Saharienne at El-Oued, Algeria. This programme has also not
shown impressive results. These two programmes illustrate
the long time frames needed for conventional breeding and
the disappointing results of considerable effort.
Because of these obstacles, date palm improvement programmes have turned towards biotechnological approaches,
including the use of TC for mass production of clones and
identification of molecular markers associated with desirable
traits. Most current biotechnologically based date palm
improvement programmes have focused on pest and disease
resistance, particularly related to the bayoud disease.
Research into selecting clones resistant to bayoud has been
carried out in Morocco and Algeria for several decades. Some
resistant clones have been identified or produced and
subsequently propagated by tissue culture for further
evaluation. The future of date palm improvement will be
based even more on biotechnological techniques, such as
marker-assisted selection, somatic hybridization and possibly
the production of transgenic date palms.
Regarding the status of genetic diversity in P. dactylifera,
the long history of exploitation and selection means that
possibly there are no ‘wild’ examples left of this species. There
may be a few wild groves still growing around oases, springs or
seepage areas, but most of the trees that currently exist are the
end results of an unknown number of acts of selection. This
includes trees which are not currently cultivated and may
appear to be growing wild in oases, abandoned gardens, etc.
Probably evolutionary change due to human selection has been
relatively low, so there is a certain amount of genetic diversity
present in date palms. This is reflected in the many local
cultivars, which have been selected for their adaptations to
local conditions. Characteristics such as off-shoot production,
tolerance to humidity and fruit characteristics have been
documented (Krueger, 2001).
Due to the market acceptance of dates like ‘Deglet Noor’
and ‘Medjool’ and successful TC propagation of popular
cultivars, this handful of cultivars is replacing traditional
cultivars in many date-growing countries. Due to their high
quality, ‘Medjool’ and ‘Barhee’ have become the predominant
cultivars propagated by TC, and these cultivars often replace
traditional cultivars when older blocks are replanted.
Additionally, population growth and developments, such as
hydraulic projects and desertification, threaten areas of
traditional date production, where there is the greatest chance
of genetic diversity existing. Due to these factors, genetic
resources of dates are decreasing at a rapid pace and urgent
conservation of diverse date germplasm is needed. There has
been some effort towards establishing ex situ collections of date
palm germplasm. However, there are only about a dozen in the
world, and the majority of these appear to consist mostly of
elite cultivars (Jaradat, 2001; Krueger, 2001). It is hoped that
in the future, more attention will be paid to the conservation
of native or potentially wild date palms.
Chih-Cheng T. Chao and Robert R. Krueger
Literature cited and further reading
Ahmed, I.A., Ahmed, A.W.K. and Robinson, R.K. (1995) Chemical
composition of date varieties as influenced by the stage of
ripening. Food Chemistry 54, 305–309.
Allen, R.G., Pereira, L.S., Raes, D. and Smith, M. (1998) Crop
evapotranspiration: guidelines for computing crop water requirements.
Food and Agriculture Organization (FAO) Irrigation and Drainage
Paper No. 56. FAO, Rome. 300 pp.
Barreveld, W.H. (1993) Date palm products. Food and Agriculture
Organization (FAO) Agricultural Services Bulletin No. 101. FAO,
Rome, 216 pp.
Barrow, S. (1998) A monograph of Phoenix L. (Palmae:
Coryphoideae). Kew Bulletin 53, 513–575.
Carpenter, J.B. (1981) Improvement of traditional date culture. Date
Palm Journal 1, 1–16.
Carpenter, J.B. (1991) Date palm (Phoenix dactylifera L.). Plant
Disease 75, 227–228.
Carpenter, J.B. and Elmer, H.S. (1978) Pests and Diseases of the Date
Palm. US Department of Agriculture (USDA) Agriculture
Handbook No. 527. USDA, Washington, DC, 42 pp.
Djerbi, M. (1983) Diseases of the Date Palm. Food and Agriculture
Organization (FAO) Regional Project for the Control of Bayoud in
North Africa, Baghdad, 112 pp.
Dowson, V.H.W. (1982) Date production and protection with special
reference to North Africa and the Middle East. Food and Agriculture
Organization (FAO) Technical Bulletin No. 35. FAO, Rome, 294 pp.
Dowson, V.H.W. and Aten, A. (1962) Dates: handling, processing, and
packing. Food and Agriculture Organization (FAO) Plant
Production and Protection Series No. 13/FAO Agricultural
Development Paper No. 72. FAO, Rome, 392 pp.
Phytelephas
Elmer, H.S., Carpenter, J.B. and Klotz, L.J. (1968) Pests and diseases
of the date palm. Food and Agriculture Organization (FAO),
reprinted from FAO Plant Protection Bulletin 16 (5) and 16 (6).
FAO, Rome, 32 pp.
Ferry, M. (1996) La crise du secteur phoenicicole dans les pay
méditerranéens. Quelles recherches pour y répondre? Le Palmier
Dattier dans L’Agriculture d’Oasis des Pays Méditerranéens, Options
méditerranéennes, Series A, No. 28. Centre International Hautes
Etudes Agronomiques Mediterraneennes, Paris, pp. 129–156.
Furr, J.R. and Armstrong Jr, W.W. (1962) A test of mature Halawy
and Medjool date palms for salt tolerance. Date Growers Institute
Report 39, 11–13.
Furr, J.R., Currlin, E.C., Hilgeman, R.H. and Reuther, W. (1951) An
irrigation and fertilization experiment with Deglet Noor dates.
Date Growers Institute Report 28, 17–20.
Glasner, B., Botes, A., Zaid, A. and Emmens, J. (2002) Date
harvesting, packing house management and marketing aspects. In:
Zaid, A. (ed.) Date palm cultivation. Food and Agriculture
Organization (FAO) Plant Production and Protection paper No.
156. FAO, Rome, pp. 177–208.
Howard, F.W., Moore, D., Giblin-Davis, R.M. and Abad, R.G. (2001)
Insects on Palms. CAB International, Wallingford, UK, 400 pp.
Jaradat, A.A. (2001) Date palm – a tree with a taste for salt.
Biosalinity News 2 (2), 5–7.
Kader, A.A. (1992) Postharvest biology and technology: an overview.
In: Kader, A.A. (ed.) Postharvest Technology of Horticultural Crops,
2nd edn. Publication 3311, University of California Division of
Agriculture and Natural Resources, Berkeley, California,
pp. 15–20.
Krueger, R.R. (2001) Date palm germplasm: overview and utilization
in USA. In: Proceedings of the 1st International Conference on Date
Palms. Al-Ain, United Arab Emirates, March 1998, pp. 2–37.
Le Maout, E. (1877) A General System of Botany Descriptive and
Analytical, Part II, Descriptions and Illustrations of the Orders.
Longmans & Green, London.
Nixon, R.W. (1950) Imported Varieties of Dates in the USA. US
Department of Agriculture (USDA) Circular No. 834. USDA,
Washington, DC, 144 pp.
Nixon, R.W. and Carpenter, J.B. (1978) Growing Dates in the US. US
Department of Agriculture (USDA) Bulletin No. 207. USDA,
Washington, DC, USA.
Reuveni, O. (1986) Date. In: Monselise, S.P. (ed.) CRC Handbook of
Fruit Set and Development. CRC Press, Boca Raton, Florida,
pp. 119–144.
Rygg, G.L. (1975) Date Development, Handling, and Packing in the
USA. Agriculture Handbook No. 482. US Department of
Agriculture (USDA), Washington, DC, 56 pp.
Swingle, W.T. (1904) The Date Palm and its Cultivation in the Southwestern States. US Department of Agriculture (USDA) Bureau of
Plant Industry Bulletin No. 53. USDA, Washington, DC.
US Department of Agriculture (USDA) (2004) National Nutrient
Database for Standard Reference, Release #17. Available at:
h t t p : / / w w w. n a l . u s d a . g o v / f n i c / f o o d c o m p / c g i bin/list_nut_edit.pl (accessed 17 July 2005).
Vandercook, C.E., Hasegawa, S. and Maier, V.P. (1980) Dates. In:
Nagy, S. and Shaw, P.E. (eds) Tropical and Subtropical Fruits:
Composition, Properties, and Uses. AVI Publishing Company,
Westport, Connecticut, pp. 506–541.
Zaid, A. and de Wet, P.F. (2002) Climatic requirements of date palm.
In: Zaid, A. (ed.) Date palm cultivation. Food and Agriculture
151
Organization (FAO) Plant Production and Protection Paper No.
156. FAO, Rome, pp. 57–72.
Zaid, A., de Wet, P.F., Djerbi, M. and Oihabi, A. (2002) Diseases and
pests of date palm. In: Zaid, A. (ed.) Date palm cultivation. Food
and Agriculture Organization (FAO) Plant Production and
Protection Paper No. 156. FAO, Rome, pp. 227–281.
Zohary, D. and Hopf, M. (2000) Domestication of Plants in the Old
World: the Origin and Spread of Cultivated Plants in West Asia,
Europe, and the Nile Valley. Oxford University Press, Oxford,
UK.
Phytelephas spp.
tagua
Tagua, ivory nut palm, Phytelephas spp. Ruiz & Pav.
(Arecaceae), consists of six species of single-stemmed or
clustering, dioecious, feather-leafed palms found in wet,
tropical forests from Panama to Bolivia. They are the source of
tagua or vegetable ivory (the hardened endosperm of the seed)
which was once the raw material of a significant industry for
the manufacture of buttons. Today, the ivory-like properties of
tagua have garnered renewed interest in the light of the world
ivory ban and current attention to products of sustainable
tropical agroforestry systems. As the six species are similar
enough to each other morphologically and have all been
exploited regionally, it is best to treat them together.
World production and yield
From 1840 to 1841, tagua was a small component of
Colombia’s total exports (Ocampo, 1984). The export volume
mushroomed 20 years later when tagua numbered in the top
five Colombian exports (Tovar Zambrano, 1989), as well as one
of the five most important Ecuadorean forest products (Acosta
Solís, 1944). From 1875 to 1878, tagua constituted just over
3% of all Colombian exports (Tovar Zambrano, 1989). In
1929, a peak year, over 25,000 t of Ecuadorean tagua was
exported, with a value US$1.2 million (Acosta Solís, 1944).
This would be about US$15 million in today’s dollars.
Exports of tagua from Colombia declined in the 1920s and
disappeared about 1935. Ecuador continued to produce tagua,
but exports declined after 1941, and tagua disappeared from
trade almost completely by about 1945 (Barfod, 1989).
In the latter part of the 19th century and into the first half
of the 20th, the manufacturing of buttons from tagua was a
major industry. A fifth of all buttons made in the USA in the
1920s were fabricated from tagua (Acosta Solís, 1944). Plastic
eroded the demand for tagua in the 1930s and the industry
shrunk to almost nothing (Barfod, 1989). Button production
never died off completely, however, and a small button
industry survived in Ecuador. This continued to produce the
tagua disks from which tagua buttons were manufactured in
Japan, West Germany and Italy (Barfod et al., 1990).
Few statistics are available on tagua yields from the boom
years, but Acosta Solís (1944, 1948) estimates that one plant of
Phytelephas aequatorialis produces approximately 30 kg of dry,
husked vegetable ivory annually. Bernal and Galeano (1993)
suggest that a more realistic average is one-half to one-third of
that figure. A stand of wild tagua palms (250–500 palms/ha,
assuming half are female) can yield 2.25–4.0 t/ha/year of
usable tagua (Bernal and Galeano, 1993).
152
Arecaceae
Uses and nutritional composition
The hardened endosperm of the seeds of the tagua palm is its
principal product. Dense and incredibly tough, the off-white
coloured endosperm resembles ivory when polished, and is in
fact often referred to as ‘vegetable ivory’. Unlike true ivory,
tagua is water soluble and will completely dissolve if immersed
for long periods of time. It can be softened by hydration and
then re-hardens when allowed to dry (Acosta Solís, 1944). Tagua
nuts polish very nicely and are easy to colour with various dyes.
The nuts are carved into a diverse assortment of items,
including chess pieces, figurines and tool and utensil handles.
Tagua handicrafts are particularly well developed in Ecuador.
Buttons were the most important end product during the
heyday of the tagua industry, and several initiatives oriented
towards rainforest conservation have resurrected the production
of tagua buttons. The world ban on elephant ivory has furthered
increased interest in tagua jewellery and other fashion items.
The endosperm of the tagua nut is primarily composed of
mannan A (45–48%) and mannan B (24–25%), two longchain polysaccharides (Aspinall et al., 1953, 1958; Timell,
1957). The liquid endosperm of young fruit is made into a
drink, and the gelatinous transitional stage is also sometimes
consumed. Both the outer and the inner mesocarp layers of
the fruit are edible, but the inner is preferred for its flavour
and modest oil content. This, however, is a minor use. The
seeds contain (per 100 g) 5.3 g protein, 1.6 g fat, 91.6 g total
carbohydrate and 9.3 g fibre (Gohl, 1981). No information on
the nutritive properties of the mesocarp is available.
Botany
TAXONOMY AND NOMENCLATURE Phytelephas is one of three
recognized genera in the oldest subfamily of the Arecaceae, the
Phytelphantoideae, with many unique characteristics of flower
and fruit morphology. The treatment herein follows
Henderson et al. (1995), who recognize two of Barfod’s
(1991a) subspecies of Phytelephas macrocarpa as distinct
species. Phytelephas aequatorialis was once treated as the
monotypic genus Palandra O.F. Cook. In addition to tagua, the
species are known vernacularly as yarina in many areas.
Phytelephas aequatorialis Spruce is a singlestemmed palm that can reach 12 m in height. The stem is
25–30 cm in girth, with conspicuous spirally arranged leaf
scars. The crown consists of several dozen large, pinnate leaves,
as much as 8 m in length. A skirt of dead foliage often persists
below the live leaves. Each leaf has over 100 narrow leaflets on
each side of the rachis, up to 90 cm long and 6–7 cm wide.
These are two-ranked, though sometimes the middle pinnae
are clustered and then arranged in several planes.
Phytelephas tumacana O.F. Cook is very similar to
P. aequatorialis, but has fewer leaves.
Phytelephas macrocarpa Ruiz & Pav. is solitary or clustered,
but has a very short stem that may be entirely underground or
else functioning as a pseudo-rhizome. It has fewer leaves with
fewer leaflets, all two-ranked.
Phytelephas schottii H. Wendl. is similar, but always solitary.
Phytelephas seemannii O.F. Cook is also single-stemmed, but
the stem becomes decumbent and roots develop on the lower
side. The older portions of the stem eventually die, thus the
DESCRIPTION
palm creeps along the ground. It bears 25–35 leaves reaching
7 m in length, with about 90 leaflets two-ranked on each side
of the rachis.
Phytelephas tenuicaulis (Batrford) Henderson forms clusters
of two to eight stems, 1.5–7 m tall and up to 10 cm in
diameter. Each stem holds between eight and 20 leaves with
35–73 pinnae on each side.
All species are dioecious, with separate staminate and
pistillate plants. Inflorescences are produced among the leaves
and the two sexes are very different in appearance. Male
inflorescences are long, cylindrical, fleshy spikes, up to 1.5 m
long, densely packed with 300–500 spirally arranged flower
clusters, each with two pairs of cream-coloured male flowers.
The flowers are about 1 cm long and have 150–700 stamens.
Pistillate inflorescences are much shorter than the staminate
ones and may be hidden by the crown or (on short-stemmed
species) buried in leaf litter. Each bears several to as many as
30 stalkless pistillate flowers towards their tips. The pistillate
flowers are very large (up to 15 cm long). The staminate
flowers of P. aequatorialis are borne on long stalks; those of P.
macrocarpa, P. schottii, P. seemannii and P. tenuicaulis are sessile
(unstalked); on P. tumacana, they are short-stalked.
The fruiting stems of the tagua species are round heads up
to 30 cm diameter of 15–20 closely appressed, pressureangled, dark-brown fruit, each up to 15 cm in diameter. The
thick outside layer is thick, woody and spiny; the underlying
yellow or orange mesocarp is thin, fleshy and oily. Each
contains five or six seeds of variable size and shape, but
averaging 3 ⫻ 5 cm. The endosperm is fluid, then gelatinous,
and finally hardens.
AND
CLIMATIC
REQUIREMENTS Species
of
Phytelephas are found from the Pacific lowlands of Panama,
Colombia and Ecuador, the Magdalena River valley in
Colombia, and north-western Amazonas in Colombia,
Ecuador, Peru and Brazil. Phytelephas aequatorialis is endemic
to western Ecuador, from the northern border with Colombia
Azuay province in the south. Phytelephas macrocarpa ranges
through north-western Amazonian Colombia, Ecuador, Peru
and Brazil. The related P. schottii is restricted to the valleys of
Magdalena and Catatumbo rivers in Colombia (Barfod, 1991b).
Phytelephas seemannii is the northernmost distributed species,
from the Chocó region of Colombia to eastern and central
Panama. Phytelephas tumacana is an endangered species in a
small area of south-western Colombia (Bernal, 1989).
Phytelephas species are most common on alluvial soils below
500 m elevation, where soil temperatures remain over 18°C.
Phytelephas aequatorialis and P. schottii can climb to
1000–1200 m. The species grow best in moist, shady areas
with rainfall in excess of over 2500 mm annually. However, P.
schottii can be encountered on steep slopes in fairly dry
locations in north-eastern Colombia.
Phytelephas may form large stands called taguales in
Colombia and Ecuador, that range between 1 and 25 ha or
more, inhabited by 240–500 palms/ha. Stands of the palms
are often left in pastures after deforestation. The palms set
fruit, but establishment of new seedlings is low to nonexistent. With few exceptions (Acosta Solís, 1944; Mora Mora,
1990), tagua palms have not been cultivated. Rice fields have
replaced taguales in many areas of western Colombia.
ECOLOGY
Salacca
Various bees, beetles and flies visit
Phytelephas flowers, and beetles are considered the most
frequent pollinators (Henderson et al., 1995). Acosta Solís
(1948) reported year-round flowering and fruiting of P.
aequatorialis, while Barfod (1991b) claims that flowering of
Phytelephas species is concurrent with the dry season in areas
with one. Tagua nuts are gathered continuously, at least from
perennially moist regions, but peak production does coincide
with the drier months. The seeds are dispersed by rodents,
such as pacas (Agouti paca) and agoutis (Dasyprocta spp.), who
transport the seeds from the taguale, feed on the fleshy
mesocarp or bury the seeds. Palms will continue to produce
fruit for at least 100 years.
REPRODUCTIVE BIOLOGY
FRUIT DEVELOPMENT Tagua nuts are produced only by
pistillate palms. When ripe the fruit disintegrates and the
individual seeds, enclosed by the fleshy mesocarp, fall to the
ground. The tagua nuts are gathered from the ground, after
rodents have cleaned the orange fruit flesh from around them.
The nuts are dried for several weeks to 1 year. The endocarp
surrounding the seeds is brittle after curing, and is removed.
When this is accomplished, the endosperm is still covered in a
thin, brown seedcoat.
Horticulture
Except for small areas in Ecuador (Acosta
Solís, 1944), tagua has never been cultivated. In the heyday of
the tagua industry, campesinos tossed seeds in abandoned
garden spots, and merely suppressed weeds while the palms
established, typically beginning to bear fruit in 15 years.
Germination requires 4–9 months or longer. Shade appears
essential during the juvenile period.
PROPAGATION
MANAGEMENT Male palms are frequently thinned from
taguales in Ecuador to make room for females. Tagua seems to
produce best under partial illumination (as under the thin
canopy found in riverside forest). Palms growing in full light
usually have smaller leaves than palms growing in shade.
However, much basic research on the development and
production of tagua under different conditions remains to be
done and this research is currently being conducted or
planned in Colombia and Ecuador.
In Ecuador, tagua is often found in association with fruitbearing tree species such as breadfruit (Artocarpus altilis),
cacao (Theobroma cacao) or timber species such as Cedrela
odorata and Cordia alliodora. It is believed that these are
remnants of abandoned agroforestry developments in former
taguales, and could serve as models for modern, sustainable
management of remaining ones.
NUTRITION AND FERTILIZATION
DISEASES, PESTS AND WEEDS
153
palms in South America. Inhabitants of the Chocó region of
western Colombia report that P. seemannii is susceptible to the
disease.
Alan W. Meerow
Literature cited and further reading
Acosta Solís, M. (1944) La Tagua. Edicionnes, Quito, Ecuador.
Acosta Solís, M. (1948) Tagua or vegetable ivory – a forest product of
Ecuador. Economic Botany 2, 46–57.
Aspinall, G.O., Hirst, E.L., Percival, E.G.V. and Williamson, I.R.
(1953) The mannans of ivory nut (Phytelephas macrocarpa). Part I.
The methylation of mannan A and mannan B. Journal of the
Chemistry Society 1953, 3184–3188.
Aspinall, G.O., Rashbrook, R.B. and Kessler, G. (1958) The mannans
of ivory nut (Phythelephas macrocarpa). Part II. The partial acid
hydrolysis of mannans A and B. Journal of the Chemistry Society
1958, 215–221.
Barfod, A. (1989) The rise and fall of vegetable ivory. Principes 33,
181–190.
Barfod, A. (1991a) Usos pasados, presentes y futuros de las palmas
Phytelephantoideas (Arecaceae). In: Ríos, M. and Bergmann, B.
(eds) Las Plantas y el Hombre. Edicionnes Abya Yala, Quito,
Ecuador, pp. 23–46.
Barfod, A. (1991b) A monographic study of the subfamily
Phytelephantoideae (Arecaceae). Opera Botanica 105, 1–73.
Barfod, A., Bergmann, B. and Pedersen, H.B. (1990) The vegetable
ivory industry: surviving and doing well in Ecuador. Economic
Botany 44, 293–300.
Bernal, R.G. (1989) Endangerment of Colombian palms. Principes 33,
113–128.
Bernal, R.G. and Galeano, G. (1993) Tagua. In: Clay, J.W. and
Clement, C.R. (eds) Selected species and strategies to enhance
income generation from Amazonian forests. Food and Agriculture
Organization (FAO) Miscellaneous Working Paper 93/6. FAO,
Rome. Available at: http://www.fao.org/docrep/v0784e/v0784e10.
htm#tagua (accessed 4 December 2006).
Gohl, B. (1981) Tropical Feeds: Feed Information Summaries and
Nutritive Values. Food and Agriculture Organization (FAO)
Animal Production and Health Series 12. FAO, Rome.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.
Mora Mora, J.A. (1990) Impacto ambiental por el establecimiento de
palma africana y camarón en selva, Costa Pacífica, Tumaco.
Unpublished Report. Fondo FEN-Colombia, Bogotá, Colombia.
Ocampo, J.A. (1984) Colombia y la Economía Mundial 1830–1910. Ed.
Siglo XXI, Bogotá, Colombia.
Timell, T.E. (1957) Vegetable ivory as a source of a mannan
polysaccharide. Canadian Journal of Chemistry 35, 333–338.
Tovar Zambrano, B. (1989) La economía colombiana (1886–1922).
In: Tirado Mejía, A. (ed.) Nueva Historia de Colombia, Vol. 5.
Edicíon Planeta, Bogotá, Colombia, pp. 950.
No information is available.
The larvae of a large weevil
similar (perhaps identical) to the giant palm weevil
(Rhynchophorus palmarum) infests the stems of tagua, killing
the palm (Acosta Solís, 1948). Rhynchophorus palmarum is the
vector of the nematode that causes red ring disease, which has
caused great losses among cultivated coconuts and African oil
Salacca zalacca
salak
Salak, Salacca zalacca (Gaertn.) Voss. (Arecaceae), is the correct
name, though Salacca edulis is still found in the literature
(Mogea, 1982). It is most commonly referred to as salak (in
Indonesia, Malaysia and the Philippines). Other names are sala
and rakam (ragum, ragahm, rakum) (Thailand); snake fruit,
154
Arecaceae
snake palm and salak palm (English); yingan (Myamar); keshi
sa laka and she pi guo zong (Chinese); fruit à peau de serpent,
fruit de palmier à peau de serpent and salacca aux fruit à peau
de serpent (French); salakpalme, salak, schlagenfrucht and
zalak (German); sarakka yashi (Japanese); and salaca (Spanish).
World production
Salak is widely cultivated in the wetter parts of the IndoMalay region and found as an understorey palm in Java and
southern Sumatra.
rakam in Thailand, salak kumbar and salak renkam in
Malaysia) provide edible fruit. Salacca wallichiana is shorter
and more compact than S. zalacca. The synonyms are Calamus
zalacca Gaertn., Salacca edulis Reinw., Salacca zalacca (J.
Gaertn.) Voss ex Vilmorin, Salakka edulis Reinw. ex Blume.
Subspecies have been described for Indonesia S. zalacca
(Gaertn.) Voss var. amboinensis (Becc.) J.P. Mogea, known in
English as Bali salak palm and in Malay as salak bali, and S.
zalacca (Gaertn.) Voss var. zalacca known in Malay as salak
jawa.
This creeping and tillering palm has a short
stem up to 1.5 m, with very short internodes and shallow
roots. It does form basal suckers but does not form large
clumps. When the trunk comes in contact with the soil it
sends out roots. This palm grows rapidly, reaching 1.5 m
within 4 years. The feather-like pinnate leaves (7 m long) have
grey to blackish, long, thin, sharp spines on the petiole (Fig.
A.20), midrib margins and leaflets (Maggs, 1984). The dark
green leaflets are 20–70 cm long by 2–7 cm wide. To assure
fruiting, ten leaves per plant are necessary for ‘Pondoh’
cultivar and 16 for ‘Bali’.
This usually dioecious palm produces an axillary stalked
spadix that is initially enclosed by the spathe. The male
spadices (7–15 ⫻ 0.7–2 cm) occur in bunches of four to 12
(Schuiling and Mogea, 1992). The larger female flowers are
borne on shorter spadices (7–10 cm long), with 15–40 per
inflorescence and nine to 12 inflorescences/year. The flowers
are borne in pairs in the axils of the scales. The male flower
has six stamens borne on a reddish, tubular corolla with
DESCRIPTION
Uses and nutritional composition
The ripe fruit is normally consumed fresh and there is an
export market for high quality graded fruit in cartons. The
edible part is non-fibrous, has a sweet taste and crisp texture
and is yellowish white to brown in colour. Peeling can be
tedious as the small spines on the skin can cause itchiness. It is
also candied, pickled, dried and fresh unripe fruit are made
into a salad. The fruit is also canned in syrup. The flesh can be
minimally processed with a plastic cover wrap or an edible
coating, and held at 5–10°C for about 1 week.
Harvest indices have not been set up though harvesting is
delayed until the astringency and acidity have reached a
minimum. Other factors that are used when considering when to
harvest are when 160 days have passed since flowering, the fruit
colour changes from dark brown to reddish brown, and firmness
and ease of fruit detachment. Spadices should be individually
harvested at the optimum stage. Overripe fruit is tasteless and
has off-odour. The fruit is low in vitamins and oil (Table A.62).
Botany
TAXONOMY AND NOMENCLATURE Salacca zalacca (salak) and
a related species Salacca wallichiana C. Martins. (known as
Table A.62. Proximate analysis of salak fruit per 100 g (Source: Leung et al.,
1972; Siong et al., 1988).
Proximate
Water
Calories (kcal)
Protein
Fat
Carbohydrate
Fibre
Ash
Minerals
Calcium
Phosphorus
Iron
Sodium
Potassium
g
80
77
0.7
0.1
18.4
0.4
0.6
mg
8
9
0.3
6
168
Vitamins
mg
Ascorbic acid
Carotene
Thiamine
Niacin
Riboflavin
4
0.05
0.2
2.4
0.008
Fig. A.20. Salacca edulis palm and insert showing fruit (with
permission from Sitijati Sastrapradja from Palem Indonesia, Lembaga
Biologi Nasional, 1978).
Salacca
minute pistil lobes that shed pollen in the early morning. The
tubular corolla of the female flower is yellow-green outside,
dark red inside and has a triocular ovary with a short trifid red
style and six staminodes. Inflorescence development from
emergence takes 80–90 days.
The palm thrives
under humid tropical lowland conditions with 1700–3100 mm
rain/year (Kusumo, 1995). The rainfall should be uniformly
distributed, with only a short dry season, otherwise irrigation
or a high water table is needed. Availability of water may
determine seasonality of fruiting patterns; uniform water
availability provides regular flowering and fruiting. A
temperature range of 22–32°C is reported to be required.
Salak grows from sea level to 300–500 m depending upon the
distance from the equator. Fruit yield and quality decline in
cooler areas or at higher elevations. This palm is normally
grown under shade and is sometimes intercropped with other
tree crops such as mango, jack fruit, durian, rambutan,
mangoesteen, banana or rubber. Shaded (25%) young salak
plants grow faster and have higher production. Mature plants
do not normally require shading as they begin to self shade
each other. A free draining soil (pH 6–7) with high organic
matter is preferred. The shallow rooting system does not stand
flooding. It can be grown on sandy soils with irrigation.
ECOLOGY AND CLIMATIC REQUIREMENTS
The palm starts flowering 3–4 years
after sowing from seed and after 2–3 years for suckers. It may be
productive for up to 50 years. Like other palms, a dry period is
not required to induce flowering. This plant is normally crosspollinated, however, some cultivars (monoecious ‘Bali’) are selfpollinated. Insects (weevils and other beetles) are thought to be
the natural pollinators (Mogea, 1978), but hand pollination is
practised when natural pollination is deficient. The palm
flowers and sets fruit continuously but there are harvest peaks
on the island of Bali in June–July and December–February in
other parts of Indonesia. The December–February harvest peak
in Indonesia coincides with flowering in the first half of the dry
season after a smaller June–July harvest.
Pollen is collected by covering the inflorescences with a
paper bag or the cut inflorescences or florets are dried in an
oven at 35°C. The collected pollen can be stored for 6–12
months at 4°C and used for pollination. Stored pollen is
diluted with talcum powder before pollination of
inflorescences that can be done at any time of the day if there
is no moisture (dew) on the stigma. Wet pollen and stigma are
susceptible to fungal attacks.
REPRODUCTIVE BIOLOGY
FRUIT DEVELOPMENT Fruit mature 5–7 months after
pollination. The 15–40 tightly packed globose to ellipsoid
drupes per spadix (about 50 g) are 5–7 ⫻ 5 cm tapering to a
point. The most noticeable feature is the numerous yellow to
brown united scales that end in a small spine and cover the skin.
The scales develop from the exocarp. There are usually three,
2–8 mm thick edible fleshy sarcotesta seeds per fruit (blackish
nuts) and a white somewhat translucent homogenous
endosperm. The taste has been compared to a combination of
apple, banana and pineapple. The aroma is due to esters and
lactones, with carboxylic acids giving the slightly sour, pungent
odour; no terpenoids have been detected (Wong and Tie, 1993).
155
Horticulture
The palm can be propagated from seed,
suckers, layering or stem cuttings. Micro-propagation
procedures are being developed. After removal of seeds from
the fruit, viability is quickly lost, with germination falling
from 55% after 1 week to no germination after 2 weeks. Seeds
are planted directly into the field or in a nursery then
replanted 4 months later. Seeds should be planted into moist
conditions with an organic mulch covering. Young palms
require shade (25%) during the first year of establishment.
Plants from seeds flower in 3–4 years with > 50% of the
plants being male. Layered trees fruit in 2–3 years. Layers and
suckers should come from a mature clump, in which three to
four shoots develop each year, though the thorns make sucker
removal difficult. A split bamboo tube is used to encase the
sucker and this is pushed into the soil until rooting occurs.
After rooting takes place, the bamboo tube and sucker can be
removed with less danger. The stem from mature plants (7–10
years old) with all leaf sheaths removed can also be cut into
sections, each with a lateral bud, dipped in fungicide and
rooting hormone and planted in a nursery.
Drainage is essential to avoid waterlogging and organic
matter needs to be worked into the soil before planting. Since
the crop is frequently interplanted with other tree crops that
provide shade, spacing information is limited, 2–6 m on a
square giving 2000–3000 plants/ha has been recommended.
Male plants, if necessary, are planted at a rate of 2–20%,
dispersed among the female trees.
PROPAGATION
PRUNING AND TRAINING Basal suckers are removed so as not
to reduce yield of the mother palm. Lateral shoots may be
spared to grow into fruiting stems or for vegetative propagation. If the stem becomes tall, it looses vitality; to rejuvenate
it, earth is pushed up around the stem. Alternatively, tall
plants are cut off or bent over to touch the ground and the
stem covered with earth, compost and manure to stimulate
rooting.
Fruit thinning to six to eight per inflorescence is practised
about 3 months after flowering to provide space for the
remaining fruit. The supporting leaf is sometimes also pruned
to allow fruit bunch development and the plant is fertilized
(1–2 kg/plant).
NUTRITION AND FERTILIZATION Manure and compost,
ammonium sulphate, urea, superphosphate and potassium
chloride have been tried. Excess nitrogen is reported to lead to
strong vegetative growth that increases the risk of plants
falling over, and having large fruit with poor postharvest life.
Adequate use of fertilizer may make shading less necessary
than previously assumed (Schuiling and Mogea, 1992).
Besides potassium, magnesium, sulphur, boron and zinc are
reported to limit growth of ‘Bali’ and ‘Pondoh’ cultivars. A
fertilizer rate per plant of 300 g ammonium sulphate, 38 g
urea, 175 g potassium chloride, 200 g dolomite, 3.8 g borax
and 3.4 g zinc sulphate for ‘Pondoh’ and the same, except only
150 g per plant of dolomite for ‘Bali’ has been recommended.
Half is applied after harvest and the other half 30 days later,
broadcasted around the plant at the outer canopy line
(Kusumo, 1995).
156
Arecaceae
Irrigation is necessary if the superficial root system does not
reach the water table. Dry spells in excess of 10 days indicate
the need for irrigation. Irrigation during dry spells can lead to
more even fruiting throughout the year. Plants need 0.7 times
the evaporation rate (4–4.5 mm/day) on a 6 ⫻ 6 m spacing,
about 100–118 l/plant/day.
DISEASES, PESTS AND WEEDS A number of diseases have been
reported (Table A.63), though the importance of each is
unreported. Sanitation is practised to reduce infection pressure.
A layer of granular-looking flesh adheres to the kernel in ripe
fruit and is referred to as ‘masir’. The cause of ‘masir’ is
unknown. Fruit splitting can also occur in fruit approaching
maturity that receives excess rain after a short drought. Postharvest fruit rot caused by Thielaviopsis spp. can be controlled by
dipping fruit in 50°C water for 3 min (Kusumo, 1995).
Larvae of weevils (Omotemnus miniatocrinitus, Omotemnus
serrirostuis) tunnel into the top of the palm and can cause
severe damage. The weevil, Nodocnemis sp., though a
pollinator, can damage young fruit bunches by boring into the
fruit. Other pests include leaf-eating caterpillars, leaf rollers
and scabs. Rodents such as rats and squirrels can cause losses.
Until leaf canopy closure occurs, weed control is essential.
Mechanical weed control is normally practised.
Fruit are harvested
by cutting the bunches, with a mature palm bearing 20
kg/year. In Indonesia, fruit are handled in bamboo baskets
and considerable losses occur due to mechanical injury that
leads to spoilage. Fruit can be washed in water or brushed
with a dry brush. Good, undamaged fruit should be selected
and can be stored at 12–15°C for up to 2–3 weeks. Lower
storage temperatures lead to chilling injury, the symptoms
being skin pitting and discoloration and the flesh can turn
brown and become soft (Mahendra and Janes, 1994). At
ambient temperatures, fruit last about 7–10 days.
POSTHARVEST HANDLING AND STORAGE
Indonesia has numerous
cultivars (Kusumo, 1995) that are distinguished by place of
origin and cultivation. ‘Bali’ is monoecious bearing both
hermaphroditic and male flowers, while dioecious cultivars
include ‘Condet’, ‘Gading’, ‘Pondoh’ and ‘Suwaru’. Cultivars
vary in fruit taste, mesocarp texture, colour of flesh and rind,
and place of production (Yaacob and Sabhadrabandhu, 1995).
‘Pondoh’ is sweeter but has only 52% edible flesh, while the
others are 70–80% edible. ‘Bali’ with about 80% edible flesh
MAIN CULTIVARS AND BREEDING
Table A.63. Some diseases and disorders of salak.
Common name
Organism
Parts affected
Fruit rot
Mycena sp.
Mycelium growth on the
fruit branches
Flower wilt
Fusarium sp.
Marasmius palmivorus
Pestalotia sp.
Corticium salmonicolor
Thielaviopsis spp.
Ceratocystis paradoxa
Fusarium sp.
Aspergillus sp.
Leaf spot
Pink disease
Fruit rot
Flower
Black spots on leaves
Plants and fruit
Fruit
has the same sugar level (20%) as ‘Pondoh’, but has twice the
acid content (0.44 versus 0.23%). The larger ‘Swaru’ fruit,
though sweet and moist, has poor keeping quality. Frequently,
a cultivar will not perform well outside the region in which it
was selected.
The dioecious nature of many cultivars leads to wide
variation in their progeny; hence upon selection of a suitable
clone, vegetation propagation is used. Seedless forms do occur
and are preferred. Soft fruit spines, thick flesh, sweet and
aromatic taste, and high yields are frequently the desired
characteristics. Thornless salak cultivars would be very useful.
A thornless Salacca spp. has been found in Thailand
(Polprasid and Salakpetch, 1989), but unfortunately, it has
poor fruit quality and low productivity. A number of related
species are being investigated for potential use (Hambali et al.,
1989).
Robert E. Paull
Literature cited and further reading
Azwar, R., Sumarmadji, U., Haris and Basuki (1993) Intercrops in
small holder rubber based farming system. Indonesian Agriculture
Research and Development Journal 15 (3), 45–51.
Chandraparik, S., Poonnachit, U., Worakuldamrongchai, S. and
Salakpetch, S. (1996) Sara Khong Sala. Mitkaset Karntalard. Lea
Kotsana, Jatujuck, Bangkok, Thailand, 65 pp. (In Thai)
Hambali, G.G., Mogea, J.P. and Yatazawa, M. (1989) Salacca
germplasm for potential economic use. In: Siemonoma, J.S. and
Wulijami-Soetjipto, N. (eds) Plant Resources of South-East Asia
Fruit. PROSEA International Symposium. Pudoc, Wageningen,
the Netherlands, p. 260.
Kusumo, S. (1995) Salak, a prideful fruit of Indonesia. Indonesian
Agriculture Research and Development Journal 17 (2), 19–23.
Leung, W.-T.W. and Flores, M. (1961) Food Composition Tables for
Use in Latin America. US National Institute for Health, Bethesda,
Maryland.
Leung, W.T.W., Bitrum, R.R. and Chang, F.H. (1972) Part 1.
Proximate composition of mineral and vitamin content of East
Asian foods. In: Food Composition Table for Use in East Asia.
United Nations Food and Agriculture Organization and US
Department of Health and Welfare, Bethesda, Maryland.
Maggs, D.H. (1984) Palmae, Subfamily Calameae, Salak. In: Page,
P.E. (compiler) Tropical Tree Fruits of Australia. Queensland Dept.
Primary Industry Information Series QI 83018, Brisbane,
Australia, pp. 129–130.
Mahendra, M.S. and Janes, J. (1994) Incidence of chilling injury in
Salacca zalacca. In: Champ, B.R., Highley, E. and Johnson, G.I.
(eds) Postharvest Handling of Tropical Fruits. Proceedings of
International Conference, Chiang Mai, Thailand, 19–23 July
1993, pp. 402–402.
Mogea, J.P. (1978) Pollination in Salacca edulis. Principes 22, 56–63.
Mogea, J.P. (1982) Salacca zalacca, the correct name for the salak
palm. Principes 26, 70–72.
Polprasid, P. and Salakpetch, S. (1989) Improvement of Salacca spp.
in Thailand. In: Siemonoma, J.S. and Wulijami-Soetjipto, N. (eds)
Plant Resources of South-East Asia Fruit. PROSEA International
Symposium. Pudoc, Wageningen, the Netherlands, pp. 296–297.
Porcher, M.H. (2005) Multilingual Multiscript Plant Name
Database. The University of Melbourne, Australia. Available at:
http://www.plantnames.unimelb.edu.au/Sorting/Frontpage.html
(accessed 31 August 2005).
Serenoa
Schuiling, D.L. and Mogea, J.P. (1992) Salacca zalacca (Gaentner)
Voss. In: Verheij, E.W.M. and Coronel, R.E. (eds) Plant Resources
of South East Asia. PROSEA Foundation, Bogor, Indonesia,
pp. 281–284.
Siong, T.E., Noor, M.I., Azudin, M.N. and Idris, K. (1988) Nutrient
composition of Malaysian foods. ASEAN Food Handling Project,
Kuala Lumpur, Malaysia.
Sosrodihardjo, S. (1986) Physical and chemical development of salak
fruit (Salacca edulis Reinw.) Pondoh variety. Bulletin Penelitian
Horticulture 13 (2), 54–63.
Tomlinson, P.B. (1990) The Structural Biology of Palms. Oxford
University Press, Oxford, UK.
Uhl, N.W. and Dransfield, J. (1987) Genera Palmarum. Allen Press,
Lawrence, Kansas.
Uhl, N.W. and Dransfield, J. (1988) Genera Palmarum, a new
classification of palms and its implication. Advances Economic
Botany 6, 1–19.
Verheij, E.W.M. and Coronel, R.E. (eds) (1992) Edible Fruits and
Nuts. Plant Resources of South East Asia No. 2. PROSEA
Foundation, Bogor, Indonesia.
Wong, K.C. and Tie, D.Y. (1993) Volatile constituents of salak
(Salacca edulis Reinw.) fruit. Flavour and Fragrance Journal 8,
321–324.
Wuryani, S., Budiastra, I.W., Syarief, A.M. and Purwadaria, H.K.
(2000) Model predicting the shelf life of edible-coated minimally
processed salak. Australian Centre for International Agricultural
Research (ACIAR) Proceedings 100, 559–566.
Yaacob, O. and Sabhadrabandhu, S. (1995) The Production of
Economic Fruits in South-East Asia. Oxford University Press,
Kuala Lumpur.
Serenoa repens
saw palmetto
Saw palmetto, Serenoa repens (Bartr.) Small (Arecaceae), is a
shrubby palm native to the south-eastern USA that forms
large clusters from a prostrate, branching stem. The fruit,
while not particularly palatable, yields an effective treatment
for benign prostate enlargement. Though not approved by the
Food and Drug Administration (FDA) for this use in the
USA, large quantities of fruit are shipped to Mexico and
Europe annually for processing.
157
Botany
TAXONOMY AND NOMENCLATURE Only a single species is
recognized in the genus Serenoa. It is classified in the tribe
Corypheae of subfamily Coryphoideae. Synonyms include
Sabal serrulata (F. Michx). Nutt. ex Schult. & Schult f. and
Serenoa serrulata (F. Michx). Nutt.
Saw palmetto stems typically lie prostrate at
the soil surface but can grow upright and reach heights of
5–7 m. Multiple, persistent, palmate leaves up to 1 m wide
emerge from the stem’s terminal buds. They vary from olive
green to silvery blue. Short recurved spines line the petioles,
giving rise to the common name. Flowers are perfect and
borne on paniculate inflorescences that emerge from among
the leaves. The fruit is a one-seeded ellipsoid drupe with a
fleshy mesocarp, 1.6–2.5 mm long and 1.2–1.9 cm wide. The
fruit mesocarp has a strong odour of butyric acid. Fruit colour
turns from green to yellow to orange, and then to bluish black
when fully ripe.
DESCRIPTION
ECOLOGY AND CLIMATIC REQUIREMENTS Saw palmetto is
endemic to the coastal plain of the south-eastern USA. The
northern limits of its range extend from south-eastern
Louisiana through Tifton Georgia to Charleston County,
South Carolina (Hilmon, 1968; McNab and Edwards, 1980).
Saw palmetto occurs as a major understorey plant in seasonally
wet pine flatwoods, well-drained scrubby flatwoods, and on
sandy berms and dunes along rivers and the coast (Tanner et
al., 1996). It grows in a variety of conditions from shade to full
sun. Saw palmetto occurs on a wide range of soil types but
most commonly is found on seasonally flooded, sandy, acidic
podosols typical of flatwoods ecosystems in the lower coastal
plain, but it readily colonizes calcareous sandy soils near coasts,
and on limestone in southern Florida. Annual rainfall varies
from 114 cm in the northern part of its range to over 150 cm
along the south-eastern coast of Florida. Rainfall also becomes
more seasonal going from north to south, with increasingly
more rainfall occurring during summer. In southern Florida,
the southernmost part of its range, 64% of average annual
rainfall occurs from June to September (Hilmon, 1968).
Fruit production of saw palmetto is primarily from management of wild stands. Typically 0.4–0.5 kg of fruit are
produced on each inflorescence. In some cases, individual
inflorescences can produce up to 12 kg of fruit. Average fruit
yield for a site is approximately 200 kg/ha; however, yields
can vary from less than 100 kg/ha to more than 1500 kg/ha
(Carrington et al., 1997). Fruit collected for pharmaceuticals
sold for over US$6/kg in 1995. Total estimated value of fruit
sold in 1996 was approximately US$5 million (Carrington et
al., 2000).
REPRODUCTIVE BIOLOGY Saw palmettos primarily reproduce
vegetatively through suckers from the main stem. In time,
extensive clumps of genetically identical clones can be formed.
Saw palmettos must be at least 0.6 m in height to flower
(Carrington et al., 2000). Inflorescences emerge from buds at
the bases of previous season’s leaves in February–April, and
flowering occurs from April to June (Hilmon, 1968). Flowers
are insect pollinated (Tanner et al., 1996). The European
honeybee (Apis mellifera L.) is the primary pollinator but over
30 other insect poliinators have been reported. Saw palmettos
flower heavily about every 2–4 years. Fruit are bird and
mammal dispersed (Tanner et al., 1996) and ripen in
August–November.
Uses and nutritional composition
Horticulture
Free fatty acids and phytosterols within the fruit are effective
in treating benign prostatic hyperplasia (Tasca, 1985;
Braeckman, 1994; Wilt et al., 1998).
PROPAGATION
World production
Seed germination ranges from 20% after 15
months in field conditions to 55% after 6 months under lab
conditions (Hilmon, 1968). Seeds can remain viable for up to
158
Arecaceae
1 year, but germination rate is reduced (Carrington et al.,
2000). Seed germination may be enhanced if the seeds pass
through an animal digestive system (Tanner et al., 1996).
MANAGEMENT Growth of saw palmetto is slow, between 0.6
and 2.2 cm/year of stem elongation (Hilmon, 1968;
Abrahamson, 1995). It has been estimated that some saw
palmettos may be 500–700 years old (Abrahamson, 1995;
Tanner et al., 1996). The most cost-efficient practice to
increase fruit production is prescribed burning (Carrington et
al., 2000). Optimal burning frequency is every 5–8 years. Soil
fertilization has been used to increase coverage of saw
palmetto (Carrington et al., 2000).
PESTS AND WEEDS Emerging saw palmetto
inflorescences are subject to attack by cabbage palm caterpillars (Litoprosopus futilis G. & R.) and green fruit are subject
to anthracnose (Colletotrichum gloeosporioides (Penz.) Penz. &
Sacc. in Penz.) (Carrington et al., 2000). Giant palm weevil
(Rhynchorphorus cruentatus) is believed to infest saw palmetto
stems. Removal of competing vegetation in wild stands may
increase fruit production.
Alan W. Meerow
Syagrus spp.
syagrus
Syagrus, Syagrus spp. Mart. (Arecaceae), is distributed broadly
in South America, with a particular diversity of species in
seasonally dry to xeric regions of central and eastern Brazil.
The seeds of virtually all of the species are edible, a number of
which yield oil, and many of the species also have edible fruit.
None of the species is cultivated commercially for its fruit and
seeds.
Uses and nutritional composition
Nutritional composition has been determined for the fruit of
one species, Syagrus coronata (Table A.64). It might be
assumed that profiles for other species would be similar.
DISEASES,
Literature cited and further reading
Abrahamson, W.G. (1995) Habitat distribution and competitive
neighbourhoods of two Florida palmettos. Bulletin of the Torrey
Botanical Club 122, 1–14.
Braeckman, J. (1994) The extract of Serenoa repens in the treatment of
benign prostatic hyperplasia: a multicenter open study. Current
Therapy Research 55, 776–785.
Carrington, M.E., Mullahey, J.J. and Roka, F. (1997) Saw palmetto: a
fountain of youth. Proceedings of the American Forage Grassland
Council 6, 233–237.
Carrington, M.E., Mullahey, J.J., Krewer, G., Boland, B. and
Affolter, J. (2000) Saw palmetto (Serenoa repens): an emerging
forest resource in the southeastern USA. Southern Journal of
Applied Forestry 24, 129–134.
Hilmon, J.B. (1968) Autecology of saw palmetto [Serenoa repens
(Bartr.) Small]. PhD thesis, Duke University, Durham, North
Carolina.
Hilmon, J.B., Lewis, C.E. and Bethune, J.E. (1963) Highlights of
recent results of range research in Southern Florida. Society of
American Foresters Proceedings 1962, 73–76.
McNab, W.H. and Edwards, M.B. (1980) Climatic factors related to
the range of saw-palmetto (Serenoa repens (Bartr.) Small).
American Midland Naturalist 103, 204–298.
Tanner, G.W., Mullahey, J.J. and Maehr, D. (1996) Saw-palmetto: an
ecologically and economically important native palm. Circular WEC109, Florida Cooperative Extension Service, Institute of Food and
Agricultural Sciences, University of Florida, Gainesville, Florida.
Tasca, A. (1985) Treatment of obstructive symptomatology in
prostatic adenoma with an extract of Serenoa repens. Minerva
Urologica e Nefrologica 37, 87–91.
Wilt, T., Ishani, A. and Mulrow, C. (1998) Saw palmetto extracts for
treatment of benign prostatic hyperplasia: a systematic review.
Journal of the American Medical Association 280, 1604–1609.
Botany
TAXONOMY AND NOMENCLATURE Thirty species of Syagrus
are recognized, many of which were once treated as species of
Cocos (Glassman, 1987; Henderson et al., 1995), to which the
genus of Syagrus is thought to be fairly closely related.
Generic synonyms that were still recognized until recent
taxonomic work include Arecastrum (Drude) Becc., Arikury
Becc., Arikuryoba Barb. Rodr. and Rhyticocos Becc. The
species readily hybridize where they occur together.
DESCRIPTION
Syagrus spp. are small to large, mostly solitary-stemmed
palms, with pinnately compound leaves. The leaf sheaths are
frequently fibrous and the petiole is sometimes armed with
fibre spines. The leaflets are most often arranged in clusters
along the rachis, green or grey-green, and with brown scales
on the underside. Specific details about various species are
listed in Table A.65. The flower stems emerge from among the
leaves and are always subtended by a persistent peduncular
bract (spathe). They may be branched to one order or spicate.
The flowers are unisexual and borne on the rachillae in triad
clusters of a central female flanked by two males. The fruit are
ellipsoid, spherical or ovoid, often have a prominent beak, and
can be coloured green, yellow, orange, brown or red. The
Table A.64. Chemical composition of the pulp and nut from the fruit of the
licury palm (Syagrus coronata) (Source: Crepaldi et al., 2001).
Mean and standard deviation
Constituent
Moisture (%)
Calorific value (kcal/100 g)
Ash (%)
Lipids (%)
Nitrogen (%)
Protein (%)
Total carbohydrates (%)
Xanthophyll
␣-Carotene
-Carotene (mg/g)
Vitamin A (ER)
␣-Tocoferol (mg/g)
Ascorbic acid
a
ND, none detected.
Pulp
Nut
77.4 + 0.16
108.6 +
1.4 + 0.06
4.5 + 0.3
0.5 +
3.2 +
13.2 +
Trace
Trace
26.1 + 0.7
4.4 + 0.1
3.8 + 0.4
Trace
28.6 + 0.8
527.3 +
1.2 + 0.01
49.2 + 0.08
2.2 + 0.01
11.5 + 0.03
9.7 +
NDa
ND
ND
ND
ND
ND
Table A.65. Some Syagrus spp. with edible fruits and/or seeds.
Name
Common name(s)
Distribution
Syagrus botryophora
(Mart.) Mart.
Pati, patioba
Syagrus cardenasii
Glassman
Ecology
Habit and size
Leaves
Fruit
Seed
Atlantic coast of
Rainforest on lateritic 10–20 m tall, solitary
Brazil from south
clay below 400 m
Sergipe to north
Espirito Santo states
10–15, 3 m long, arching,
with 100–150 rigid,
ascending leaflets per side
of rachis
Ellipsoid, 3.5–4.5 cm long, 2.2–2.5
cm wide, white to yellow-green,
human consumption not reported
Rich in edible oil
Corocito, saro
Bolivia
Seasonally dry forest 2–3 m tall, clustered or
on dry hills at
solitary, stem mostly
400–1800 m
underground
8–12, 1–2 m long, with
32–74 clustered leaflets per
side, grey
Ovoid, 2–3 cm long, 1.5–2 cm wide, Human consumption
brown, mesocarp with pineapple
not reported
flavour
Syagrus comosa
(Mart.) Mart.
Babão, catolé
Central and eastern
Brazil
Open cerrado
vegetation, often on
rocky slopes to
1200 m
1–7 m tall, aerial or
subterranean stem
6–12, 1.5 m long, with
Ellipsoid, 2.5–3 cm long, 1.5–1.8
38–82 leaflets per side in
cm wide, green, edible
dense clusters of two to four
Syagrus coronata
(Mart.) Becc.
Licuri, ouricuri
North-eastern Brazil
Caatinga, semideciduous forest,
transitional
vegetation
4–15 m tall, solitary
15–30, arranged in five
twisted vertical rows,
greyish green, with 80–130
rigid leaflets per side in
clusters of two to five
Ellipsoid, 2.5–3 cm long, 1.7–2 cm Edible
wide, yellow-green to orange, with
brown hairs, sweet, edible mesocarp
Syagrus flexuosa
(Mart.) Becc.
Acumã, côco de
campo
Eastern and central
Brazil
Cerrado, woodlands, 3–7 m tall, usually
sandy to rocky soils clustering, leaf bases
to 1200 m
persisent
7–15, 1 m long, dark green,
38–80 leaflets per side, in
clusters of 2–5, waxy white
below
Ellipsoid, beaked, 3–3.5 cm long,
1.5 cm wide, yellow, edible
Human consumption
not reported
Syagrus inajai
(Spruce) Becc.
Curua rana,
inaya-y, peh-peh
Guianas and
northern Brazil
Rainforest to 500 m
Solitary, 5–28 m
15–18, 3.5 m long, 51–110
flaccid leaflets per side in
clusters of two to seven
Ellipsoid, 3–4.5 cm long, 2–3 cm
wide, yellow, human consumption
not reported
Edible
Syagrus oleracea
(Mart.) Becc.
Catolé, guariroba
Eastern Brazil
Semi-deciduous
forest to 800 m
Solitary, 7–22 m
15–20, spirally arranged,
2–4 m long, 100–150
leaflets per side, rigid, waxy
green, in clusters of two to
five
Ovoid, 4–5 cm long and 2.5–3 cm
wide, beaked, greenish to yellowgreen, edible and sold locally
Seed oil
Syagrus
romanzoffianum
(Cham.) Glassman
Chirivá, pindó,
jeribá, guariroba,
queen palm
Central and southeastern Brazil,
northern Argentina,
eastern Paraguay
and Uruguay
Various forest types, Solitary, 12–20 m tall
from dry to moist
Syagrus schizophylla
(Mart.) Glassman
Aricuriroba,
licurioba
Atlantic coast of
north-east Brazil
Restinga forest,
sandy soils, low
elevation
Syagrus smithii
H.E. Moore
Catolé
North-west Amazon
region
Lowland rainforest
on non-inundated
soils to 400 m
7–15, 3–5 m long, arching, Ovoid, 2–3 cm long, 1–2 cm wide,
plumose, 150–250 leaflets
yellow to orange, edible
per side in clusters of two to
five, the tips pendulous
Human consumption
not reported
Broadly ellipsoid, 2–3 cm long,
1.5–2.5 cm wide, bright orange,
edible and sweet
Human consumption
not reported
Ellipsoid, 6–8 cm long, 3–4 cm
wide, yellow, human consumption
not reported
Edible and reportedly
delicious
Syragrus
159
Usually solitary, 3–6 m 8–25, 1–2 m long, leaf
tall, leaf bases persistent sheaths with fibre spines,
18–48 leaflets per side,
two-ranked, rigid, regularly
arranged
Solitary, 6–12 m tall
5–18, 2.5–3 m long, 83–94
leaflets per side, irregularly
arranged; sometimes
produces undivided elliptical
leaves
Human consumption
not reported
160
Arecaceae
mesocarp is fibrous and often fleshy. The fruit contain one to
two seeds surrounded by a bony endocarp with three pores at
one end.
ECOLOGY AND CLIMATIC REQUIREMENTS Syagrus spp. occur
from Colombia east to French Guiana and south to Uruguay
and northern Argentina, with a single species in the West
Indies. The greatest diversity of species is found on the central
planalto of Brazil. Most species are found in seasonally dry
vegetation on sandy or rocky substrates. A few species inhabit
wet forests of the Amazon region or Brazil’s coastal Atlantic
rainforest; only two occur in the Andes. The species are
strictly tropical and subtropical in their climatic requirements.
Syagrus romanzoffianum is able to withstand temperatures to
⫺4°C without injury.
REPRODUCTIVE BIOLOGY Syagrus spp. are predominantly
insect-pollinated and the fruit of most species are dispersed by
animals. The seeds of several species are an important, in
some cases the exclusive, food for certain parrots during their
breeding season.
Horticulture
PROPAGATION Solitary-stemmed Syagrus spp. are propagated
exclusively by seed; the few clustering species can be carefully
divided. Fresh seed germinates readily, with germination times
varying from 6 weeks to over a year. The dryland-dwelling
dwarf species with subterranean stems are particularly slow
growing; S. romanzoffianum is the fastest (in fact, one of the
faster growing of all palm species), and is a very common
ornamental in subtropical regions.
DISEASES,
PESTS AND WEEDS Some Syagrus spp. are
susceptible to the lethal yellowing phytoplasma. A number of
fungal pathogens can cause foliar blights or bud rots (Chase
and Broschat, 1991).
Alan W Meerow
Literature cited and further reading
Chase, A.R. and Broschat, T.B. (eds) (1991) Diseases and Disorders of
Ornamental Palms. American Phytopathological Society, St Paul,
Minnestota.
Crepaldi, I.C., Almeida-Muradian, L.B. de, Rios, M.D.G., Penteado,
M. de V.C. and Salatino, E.A. (2001) Nutritional composition of
licuri fruit (Syagrus coronata (Martius) Beccari). Revista Brasileira
de Botanica 24, 155–159.
Glassman, S. (1987) Revision of the palm genus Syagrus Mart. and
other selected genera in the Cocos alliance. Illinois Biological
Monographs 56, 1–230.
Henderson, A., Galeano, G. and Bernal, R. (1995) Field Guide to the
Palms of the Americas. Princeton University Press, Princeton,
New Jersey, 363 pp.