Tropical Fruit

CROP PRODUCTION SCIENCE IN HORTICULTURE SERIES
Series Editor: Jeff Atherton, Professor of Tropical Horticulture,

University of the West Indies, Barbados
This series examines economically important horticultural crops selected from the
major production systems in temperate, subtropical and tropical climatic areas.
Systems represented range from open eld and plantation sites to protected plastic
and glass houses, growing rooms and laboratories. Emphasis is placed on the scientic
principles underlying crop production practices rather than on providing empirical
recipes for uncritical acceptance. Scientic understanding provides the key to both
reasoned choice of practice and the solution of future problems.
Students and staff at universities and colleges throughout the world involved in
courses in horticulture, as well as in agriculture, plant science, food science and
applied biology at degree, diploma or certicate level will welcome this series as a
succinct and readable source of information. The books will also be invaluable to
progressive growers, advisers and end-product users requiring an authoritative, but
brief, scientic introduction to particular crops or systems. Keen gardeners wishing
to understand the scientic basis of recommended practices will also nd the series
very useful.
The authors are all internationally renowned experts with extensive experience
of their subjects. Each volume follows a common format covering all aspects of
production, from background physiology and breeding, to propagation and planting,
through husbandry and crop protection, to harvesting, handling and storage. Selective
references are included to direct the reader to further information on specic topics.
Titles Available:
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Ornamental Bulbs, Corms and Tubers A.R. Rees

Citrus F.S. Davies and L.G. Albrigo
Onions and Other Vegetable Alliums J.L. Brewster
Ornamental Bedding Plants A.M. Armitage
Bananas and Plantains J.C. Robinson
Cucurbits R.W. Robinson and D.S. Decker-Walters
Tropical Fruits H.Y. Nakasone and R.E. Paull
Coffee, Cocoa and Tea K.C. Willson
Lettuce, Endive and Chicory E.J. Ryder
Carrots and Related Vegetable Umbelliferae V.E. Rubatzky, C.F. Quiros and
P.W. Simon
Strawberries J.F. Hancock
Peppers: Vegetable and Spice Capsicums P.W. Bosland and E.J. Votava
Tomatoes E. Heuvelink
Vegetable Brassicas and Related Crucifers G. Dixon
Onions and Other Vegetable Alliums, 2nd Edition J.L. Brewster
Grapes G.L. Creasy and L.L. Creasy
Tropical Root and Tuber Crops: Cassava, Sweet Potato, Yams and Aroids V.
Lebot
Olives I. Therios
Bananas and Plantains, 2nd Edition J.C. Robinson and V. Galn Saco
Tropical Fruits, 2nd Edition Volume 1 R.E.Paull and O. Duarte
Blueberries J. Retamales and J.F. Hancock
Peppers: Vegetable and Spice Capsicums, 2nd Edition P.W. Bosland and E.J.
Votava
Raspberries R.C. Funt
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TROPICAL FRUITS,
2ND EDITION, VOLUME II
Robert E. Paull
Tropical Plant and Soil Sciences
University of Hawaii at Manoa
Honolulu, HI, USA
and
Odilo Duarte
Retired Professor - Escuela Agrcola Panamericana
El Zamorano, Honduras
Now: Professor and Lead Scientist in Agribusiness
CENTRUM Catlica Business School
Ponticia Universidad Catlica del Per, Lima, Per
CABI is a trading name of CAB International

CABI
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R.E. Paull and O. Duarte 2012. All rights reserved. No part of this publication may
be reproduced in any form or by any means, electronically, mechanically, by
photocopying, recording or otherwise, without the prior permission of the
copyright owners.
A catalogue record for this book is available from the British Library,
London, UK.
Library of Congress Cataloging-in-Publication Data
Paull, Robert E.
Tropical fruits / Robert E. Paull and Odilo Duarte. -- 2nd ed.
p. cm. -- (Crop production science in horticulture series ; no. 20)
Includes bibliographical references and index.
ISBN 978-1-84593-672-3 (alk. paper)
1. Tropical fruit. I. Duarte, Odilo. II. C.A.B. International. III. Title. IV.
Series: Crop production science in horticulture ; 20.
SB359.P38 2011
634.6--dc22
2010016776
ISBN-13: 978 1 84593 789 8
Commissioning editor: Sarah Hulbert

Editorial assistant: Chris Shire
Production editor: Simon Hill
Typeset by Columns Design XML, Reading.
Printed and bound in the UK by MPG Books Ltd.
CONTENTS
PREFACE
vii
ACKNOWLEDGEMENTS
ix
1
ANNONAS: SOURSOP AND ROLLINIA
2
BREADFRUIT, JACKFRUIT, CHEMPEDAK AND MARANG
25
3
CARAMBOLA AND BILIMBI
53
4
DURIAN
75
5
GUAVA
91
6
MANGOSTEEN
123
7
RAMBUTAN AND PULASAN
139
8
PASSION FRUIT AND GIANT PASSION FRUIT
161
9
PALMS
191
v
vi
Contents
10
OTHER AFRICAN FRUIT: TAMARIND, MARULA AND ACKEE
223
11
OTHER TROPICAL ASIAN AND PACIFIC FRUIT
255
12
AMERICAN FRUIT
303
INDEX
363
PREFACE
Volume I presented the general aspects of tropical fruit production and

covered the major tropical fruit in international trade such as banana,
pineapple, papaya, mango and avocado. Many other tropical fruit, already
well-known in the tropics, are now appearing in larger temperate city
markets. In this volume, we have selected those that are being increasingly
seen in overseas markets outside of the tropics.
The choice of crops to present in Volume II was the greatest challenge,
especially in the last three chapters dealing with other Asian and Pacic,
African and American Fruits. The fruit crops covered in these last three
chapters is the tip of what is available and that have considerable potential as
fruit crops. A number of the chapter fruit sections show the signicant gaps
in our knowledge of managing these fruit crops in large orchards and not
backyard production. A major gap is lack of breeding effort to develop varieties
suitable for intensive production that have disease and insect resistance, high
yield and desired fruit quality that suit different growing environments and
consumer markets.
We have followed the same chapter layout used in the rst edition and
Volume I of this edition. The information in each fruit chapter deals with
taxonomy, varieties, propagation and orchard management, biotic and abiotic
problems, variety development and postharvest handling. The information
contained should be of use to all readers and students interested in an
introductory text on tropical fruit production.
Many have contributed to the endeavour. Encouragement and help of
Henry in this passion came from many and acknowledged in the First and
Second Editions. Numerous comments and suggestions from colleagues
have been incorporated. All errors and omissions are our responsibility. The
illustrations of many of the crops covered in this Volume and for Volume
I were done by Susan Monden in Honolulu. The family of Dr. Jorge Leon
granted us permission to use the gures from Botnica de los Cultivos
Tropicales. Dr. Lionel Robineau, coordinator TRAMIL gave us permission
vii
viii
Preface
to use the Hylocereus drawing and Dr. Mike Nagao the use of the Pulasan
picture. Mrs. Meg Coates Palgrave kindly agreed to let us use the Marula
drawing from Trees of Central Africa though we are unable to render it in
colour. The editor of Flore Analytique du Bnin let us use the ackee drawing.
Thanks are also due to the Commissioning Editor Sarah Hulbert and
Christopher Shire at CABI for their assistance and patience during the books
development.
We would greatly appreciate receiving all comments and suggestions on
this text. We can be reached at the address in the front of the text or via E-mail
at paull@hawaii.edu or odiloduarte@yahoo.com.
In closing, we both acknowledge the continued support, assistance,
and love of our wives Nancy and Carla, and our children that enabled us to
complete this undertaking.
Robert E. Paull
Honolulu, USA. 2012
Odilo Duarte
Lima, Peru. 2012
ACKNOWLEDGEMENTS
The authors gratefully acknowledge the support, information, and ideas

supplied by Jeff Anderson, Alton Bailey, Jit Baral, Lou Biad, Chris Biad, Anna
Biad, Judy Bosland, Emily Bosland, Will Bosland, Emma Jean Cervantes,
Danise Coon, Deyuan Wang, Natalie Goldberg, Max Gonzalez, Wendy
Hamilton, Steve Hanson, John Hard, Sue Hard, Jaime Iglesias, Sanjeet Kumar,
Jimmy Lytle, Jo Lytle, Ariadna Monroy, Mary OConnell, Jaebok Park, Jennifer
Randall, Adrian Rodriquez, Robert Steiner, Ousmane Sy, Betty Terrien,
Nankui Tong, Manju Vishwakarma, Stephanie Walker, April Ulery, Everardo
Zamora, and the Chile team at New Mexico State University.
ix
1
ANNONAS: SOURSOP AND ROLLINIA
BOTANY
Family
Both soursop and Rollinia belong to the Annonaceae, commonly referred to
as the custard-apple family. The family consists of about 75 genera that are
now widely distributed. Some Annona species are grown as ornamentals, while
others are known for their edible fruit and perfume.
Important genera and species

The soursop belongs to the most important genus Annona, which among its
more than 100 species has seven species and one hybrid that are grown
commercially worldwide. Soursop is the most tropical of these species and has
the largest fruit. Rollinia (or biriba) belongs to the closely related genus Rollinia
and it is not as well known as soursop. The two most important commercial
species are cherimoya and sweetsop. Along with the hybrid, atemoya, all three
are discussed in Chapter 6 of Volume 1.
Annona muricata L. is known as soursop in English and guanbana in most
Spanish-speaking countries. It is also known as catoche (Venezuela), zapote
agrio, zapote de viejas or cabeza de negro (Mexico), guayabano (Philippines),
nangka belanda zuurzak or sisrsak (Indonesia), thurian-khaak (Thailand), sitaphal
(India), fruta de conde, graviola, jaca do Par (Brazil), nona sri kaya or durian
belanda (Malaysia) and corossol epineux (France).
The synonyms for Rollinia mucosa (Jacq.) Baill. are R. orthopetala A. DC.
or R. deliciosa Saff., R. pulcherrima A. DC. and Annona mucosa Jacq. (Sousa,
2008). Other names are birib, fruta da condessa or berib (Brazil) (Manica,
2000), anona babosa or zambo (Mexico), ann (Peru), chirimoya (Ecuador),
Paull and Duarte 2012. Tropical Fruits, 2nd Edition, Volume II

(R.E. Paull and O. Duarte)
Chapter 1
mulato (Colombia), cachiman cochon or cachiman montagne (Guadalupe),

ann cimarrn (Puerto Rico), anonillo (Panama) and candongo (Dominican
Republic).
Area of origin and distribution

Soursop is the most tropical of and produces the largest fruit among the
Annona species. The Caribbean is the area of origin, although soursop was
distributed very early to the warm lowlands of eastern and western Africa and
to south-east China. It is commonly found on subsistence farms in south-east
Asia, and was established very early in the Pacic islands.
Soursop is considered well suited to processing and for use in local markets
for fresh consumption. It is grown extensively in Mexico, from Culiacan to
Chiapas, and from Veracruz to the Yucatan Peninsula in the Gulf region. Large
orchards are seen in this region, with a total of more than 6000 ha being grown.
Venezuela has around 4000 ha, Brazil more than 2000 ha and Peru almost
500 ha. Colombia and Ecuador have also developed some modern orchards in
the last few years for the local fruit markets and for industrial use. Exports of
fresh fruit are very low.
Rollinia originated in the forests of the Brazilian Amazon in the states of
Acre, Rondnia and the Antilles. It has been spread throughout Brazil and
other South American countries, as well as Florida (Manica, 2000; Donadio
et al., 2002). This species, apparently cultivated since pre-Columbian times, is
not widely cultivated for commercial purposes; rather, it occurs in backyard
orchards or small plantings.
ECOLOGY
Soil
Soursop, as with most Annona species, is capable of growing in a wide range
of soil types, from sandy soil to clay loams (Pinto et al., 2005). Nevertheless,
higher yields occur on more well-drained sandy to sandy loam soils. Drainage
is essential to avoid root rot. A. glabra is of interest as a rootstock because of
its tolerance to ooded soils. Soursop can withstand some drought, but will
experience ower abscission. The ideal soil pH is 56.5.
Rollinia also prefers well-drained deep soils with a good content of organic
matter but it can grow in poor acid soils high in exchangeable aluminum. The
tree can withstand periodic ooding (Sousa, 2008).
Annonas: Soursop and Rollinia
Climate
Rainfall
Rainfall and high humidity during the peak owering season greatly enhance
fruit production by preventing desiccation of stigmas, prolonging their
receptive period, and increasing fruit set and early fruit growth. Under very
rainy conditions, as occurs in parts of Costa Rica, soursop has many leaf and
fruit disease problems and is not normally grown commercially. In Colombia,
soursop will grow successfully under rainfall conditions that can consist of
two rainy seasons a year or just one. The alternating rainy and dry seasons
have a positive effect on ower initiation. In addition, dry periods favor
some leaf fall that results in new vegetative growth. Well-distributed yearly
precipitation of 5001500 mm results in adequate production, depending on
its distribution. Yields are low when the rainfall is less than 500 mm (Duarte
1997; SCUC, 2006).
Rollinia grows in hot, humid climates where a short dry period can occur,
but in many cases monthly rainfall can be as high as 300 mm during the rainy
season. It probably tolerates heavy rainfall areas better than the other fruitproducing species of this family. It needs at least 1500 mm of rainfall, and in
many areas in the Amazon valley it will grow with more than 27003000 mm
(Donadio et al., 2002).
Temperature
Temperature is a major limiting factor to production, with frost killing young
trees while older trees show some tolerance. Soursop is the least tolerant of
the Annona species (1525C mean minimum). Donadio et al., (2002) has
reported that Rollinia can withstand mild frosts in the Jaboticabal area of
Brazil, where the plant will defoliate in the winter. Soursop grows best under
average temperatures of 2528C. It can be grown at elevations of up to
1000 m in the tropics and subtropics, as long as winter temperatures do not
drop below 15C. The temperature range for Rollinia is 12C higher than for
soursop, and it prefers the hot, humid tropics. Poor pollination, a frequent
problem with all Annona species, occurs with high temperatures (30C) and
low humidity (30% relative humidity [RH]), even with hand pollination.
Lower temperatures (25C) and high humidity (80% RH) greatly improve
pollination.
Light and photoperiod
Light penetration to the base of vigorous trees with a dense canopy in close
spacing can be 2% of full sunlight, and there is very little fruit set. The soursop
plant also tends to have a conic upright form. Pruning practices and spacing
need to be adjusted to ensure light penetration. No photoperiod responses have
been reported for any Annona species.
Chapter 1
Wind
The soft wood of the trees makes them susceptible to wind damage and limb
breakage. Wind may also be partially responsible for the penetration of collarrot organisms. Productivity can be improved by windbreaks and under-tree
sprinkling to raise the RH above 60%.
GENERAL CHARACTERISTICS
Tree
The soursop is a small, evergreen tree that is slender and upright or lowbranching and bushy. It grows to heights of 4.59 m. Glossy, dark-green
leaves are alternate, simple and entire, with an obovate to elliptic shape, and
are 12.720 cm long (Fig. 1.1). The leaves emit a strong odour when crushed.
Fig. 1.1. The leaves and ower of soursop.
Rollinia generally reaches 610 m in height, but can grow as high as 20

m with a trunk diameter of 60 cm. The wood is soft, and the round or conic
canopy is formed by branches that tend to grow vertically with laterals forming
at their bases. The plant is semi-deciduous, with alternate oblong or elliptical
oblong leaves, 1220 cm long and 810 cm wide with an acuminated apices
base. The leaves are leathery and the petioles are 612 mm long. Flowers occur
once a year after leaf fall.
Flowers
The owers of Annona species are hermaphroditic and are produced singly
or in small clusters on the current seasons growth, although owers arising
in cushions from old wood are not uncommon. All lateral buds can have up
to two vegetative buds and three ower buds. Soursop lateral buds are exposed
in the leaf axil (Fig. 1.1), while the lateral buds of atemoya, cherimoya and
sweetsop are normally buried (subpetiolar). Adventitious buds can arise at
any point on the trunk. New owers continue to appear toward the apex of the
shoot as owers produced earlier at the basal portions mature.
Soursop owers are pale yellow and 2.54 cm long, with three thick, eshy
petals and three smaller inner petals alternating with the outer petals (Fig.
1.1). They have a peculiar odor. Defoliation of A. muricata manually or with
ethephon spray promotes lateral branch growth and induces additional ower
formation near the apex of the branches. Rollinia owers are solitary or form
small clusters on the current seasons growth (Moncur, 1988). They have three
sepals and six petals. The external petals have the form of wings and give the
ower the appearance of a propeller (Fig. 1.2A). They form a tubular structure
at their junction in the center of the ower (Villachica et al., 1996).
Annona species generally require 2735 days for ower-bud initiation to
anthesis. In A. squamosa, owering can extend from 36 months or even longer,
with heavy peaks. Two major owering periods occur after periods of vegetative
ushes, with the second peak coinciding with the onset of the monsoon season
in India (Kumar et al., 1977). Flowering can occur year round with a continuous
warm climate and water availability, while harvest becomes more seasonal in
the subtropics.
Pollination and fruit set

Natural pollination
The owers exhibit both dichogamy and a protogynous nature (Pinto et al.,
2005). This poses a serious problem in obtaining high yields. A. muricata oral
anthesis takes place mostly between noon and 8 pm and from 4 am to 8 am,
with pollen release occurring between 4 am and 8 am (Moncur, 1988).
Chapter 1
Fig. 1.2. Rollinia owers (A) and fruit (B).
Escobar and Snchez (1992) have given a detailed description of the timing
of the pollination process, dividing it into four phases that can take between
96 and 132 h. In phase I, the ower button opens slightly at the basal point of
contact of the outer petals. Sexual structures are whitish and stigmatic liquid
starts to become apparent and viscous, indicating the ower is receptive. In
phase II, after 4860 h the tips and bases of the outer petals have separated,
the ower is more receptive since more stigmatic liquid is present, and the
stamens become yellow, normally in the morning as the ower reaches its nal
size. In phase III, after another 2448 h, the outer petals are semi-open and
have a yellow-greenish color and more stigmatic liquid. The stamens have a
dark-yellow color and the pollen is viable. Phase IV is reached 24 h later, when
the outer petals are completely open and have acquired a sulfur color. Stigmatic
liquid becomes less viscous and the ower is still receptive. The anthers are now
cream in color and release viable pollen. The inner petals do not open, but are
slightly separated in phases III and IV. After this all the petals, stamens and
stigmata fall in 1224 h, with the calyx, receptacle and peduncle remaining
attached.
Natural pollination in soursop is complex and in most cases results in very
low fruit set and yields, with wind- and self-pollination being low (1.5%). The
nitidulid beetles (Carpophilus and Uroporus spp.) are considered important
pollinators of Annona owers, although no signicant effect has been observed
from their presence in some cases. These beetles breed very fast in the remains
of fruit, so it is recommended to maintain the rotting fruit attractant. Some
reports have indicated that the presence of three nitidulid beetles per ower can
increase fruit set by 25% (SCUC, 2006).
In the case of Rollinia, the owers are also protogynous and the two female
and male phases do not overlap to allow for self-pollination (Moncur, 1988).
The petals open only slightly during the female stage. Insects are attracted by
the scent, but nectar is not produced. Later on, the male-stage owers open
widely and insects forage for the pollen. All of this leads to poor fruit set. In
Brazil, four species of leaf beetles (Chrysomelidae) pollinate the owers with only
32% fruit set (Morton, 1987).
Hand pollination
Hand pollination is used to overcome poor pollination. Hand pollination is
often very efficient, resulting in signicant economic returns from the higher
fruit set and larger and more symmetrical fruit. Hand pollination has also
proven to be effective in Rollinia (Moncur, 1988). The pollen grains of owers
appearing early in a owering season have thick walls and are high in starch,
germinate poorly and give poor fruit set. The pollen of later owers shows a
high proportion of individual pollen grains without starch grains, and these
germinate well.
Pollen is obtained from opened owers collected between 4 and 5 pm when
the sacs have turned from white to cream. Flowers on thin branches or at the
end of such branches should be harvested for pollen collection. Pollen can be
obtained directly from picked owers held in a paper bag or cardboard box, not
a sealed container, at phase IV. Flowers picked at phase III will release pollen
the next morning. Pollen from both stages can then be mixed for use. When
the owers are shaken over a shallow tray or in a plastic jar, the stamens and
pollen separate and the pollen will stick to the jar walls. The pollen is then
transferred to a small container. Pollen obtained in the afternoon can be held in
a refrigerator for use the next morning. The moist pollen is applied to owers in
phase III or IV using a hair brush or even by rubbing the pollen on the stigma
with an index nger. Some people remove one of the inner petals to make it
easier to apply the pollen. Flowers can be tagged to keep control of the process.
Pollination is performed between 6 and 10 am, and earlier if the days are hot
and dry.
Success in hand pollination is sometimes variable, being less successful on
very humid overcast days and with young, vigorous trees. About 150 owers
can be pollinated by a skilled laborer in 1 h with a success rate of 80100%.
Flowers for hand pollination should preferably be taken from strong branches
at the center of the canopy of trees older than 4 years.
Growth regulators for fruit set
Hand pollination in commercial orchards is tedious, time-consuming and
costly. Attempts have been made to use growth regulators to regulate fruit set,
with considerable variations in response. Auxin-induced fruit grow very slowly
with less fruit drop, while gibberellic acid promotes fruit set and growth rate;
however, it does not assist in post-set retention (Yang, 1988).
Chapter 1
Fruit
The soursop fruit is a syncarp that varies from less than 0.4 kg to more than
4.5 kg, with some fruits reaching 12 kg. The size depends on genotype, extent
of pollination and fertilization. A normal fruit is generally heart-shaped to
oval (Fig. 1.3), but poor pollination results in unfertilized ovules that lead to
small, distorted, irregular shapes. The skin is dark green with many recurved,
soft spines. The esh is juicy and white with a cottony texture, and contains
many dark brown seeds that are about 2 cm long. The pulp has an agreeable
sub-acid avor with a distinct aroma. Soursop produces fruit throughout the
year, but in most areas peak production is during summer and early autumn,
sometimes with a secondary peak in the early spring.
The Rollinia fruit is also a syncarp that generally weighs from 200 to 1000 g
(Fig. 1.2B) and sometimes up to 4 kg. The avor is like that of the soursop, but
it is sweeter and less acidic; the aroma is also appreciated by consumers. The
weight of 1000 seeds is about 315 g. The ripe fruit normally has a yellow skin
and the pulp is cream or white, mucilaginous, soft and juicy.
Fruit growth shows the typical sigmoidal curve, with maturation occurring
in 1624 weeks, depending on the species and growing conditions (Fig. 1.4).
Low humidity (<60% RH) and temperature (<13C) near fruit maturity can
increase the severity of fruit-skin russeting, as well as delaying fruit maturation.
High temperatures can cause premature fruit ripening and fermentation of the
fruit.
CULTIVAR DEVELOPMENT
Genetics, cytogenetics and breeding
The chromosome numbers of most Annona species are 2n = 14 or 16. A
desirable hybrid would be between cherimoya and soursop. This would
combine the larger fruit size and acidity of soursop with the sweetness, avor
and texture of cherimoya. Attempts to cross the soursop with cherimoya,
ilama, bullocks heart or sweetsop have not been successful, and may reect a
considerable genetic distance of soursop from the other species (Samuel et al.,
1991).
Problems with breeding

Existing commercial cultivars show considerable variation in growth, fruit
set, fruit size and quality. No single variety has all the desirable characteristics.
The length of the juvenile period varies, with earliest production occurring
Fig. 1.3. Fruit of soursop, sweetsop and atemoya.
Time from Anthesis (weeks)
Fig. 1.4. Increase in fruit diameter from anthesis for soursop, sweetsop and
cherimoyas. (After Thakur and Singh, 1964; Worrel et al., 1994.)
10
Chapter 1
in 2 years and full production in 56 years. This juvenile period is extremely

variable with scions on seedling rootstocks. The seedling rootstocks are
derived from extremely heterogeneous open-pollinated seeds. Breeding
programs have focused on selections from seedling populations. Early
maturity, better fruit appearance and, in the subtropics, greater cold tolerance
are the most frequent objectives.
Cultivar development
Except for cherimoya and atemoya, very few named clonal cultivars have
been developed among the Annona species, since most plantings have been of
seedlings. In many Latin American countries, eld selections have been made
separating the sweet (actually less acid) from the acid fruited types. There are
also differences in fruit form, color and consistency (juicy and hard). These
two groups are recognized in Colombia, with the sweet type having fruits of
about 1 kg, while the acid can have fruits of up to 5 kg. Producers in Costa
Rica selected some superior types that were given names and are now being
propagated clonally. Selections have also been made in Brazil, where cultivars
(Morada, Lisa and Blanca) have also been introduced from Colombia. Of
these, Morada has the highest yields per tree (up to 40 kg) and the largest size
of fruit (310 kg). It also has rm pulp, a sub-acid avor and is more tolerant
to fruit and stem borers, which makes it the most desirable cultivar (Pinto
and da Silva, 1996). Brazilian selections include Cerradinha, Ibrimirina,
Gigante de Alagoas and FAO II.
Rollinia has apparently seen no breeding programs, although some selections
have been made in Brazil by the native tribes. Some selections produce fruit that
weigh 4 kg (Clement et al., 1982).
CULTURAL PRACTICES
Propagation
The Annona species are usually propagated by seed. The recalcitrant seeds
rapidly lose viability (6 months) and should be sown as soon as possible
after removal from the fruit. Seeds can take up to 30 days to germinate and
gibberellic acid signicantly increase germination and enhance seedling
growth. Annona seedlings require at least 34 years to bear fruit (Sanewski,
1991).
Clonal propagation by cuttings, layering, inarching, grafting and budding
have been tried for many of the Annona species. In some reports, grafting is
superior to budding in percentage takes and subsequent growth, with side-whip
and cleft graft techniques giving the best results. Escobar and Snchez (1992)
11
found patch budding to be the best, with an 82.5% success rate for soursop,
while the side graft was second with lower success rates with A. muricata, A.
reticulata and A. montana. Cleft graft and inverted T budding did very poorly
for all four rootstocks. The success of cleft and side grafting or inverted T and
patch budding was practically zero with A. squamosa. The branches should be
defoliated 12 weeks before scion wood is cut to induce bud swelling, and petioles
should be left on the branch. There are considerable graft incompatibilities
among Annona species.
Rollinia is normally propagated by seeds that germinate in about 1 month.
Seedling growth is very fast during the initial years. In some cases, grafting has
been used successfully (Sousa, 2008).
Field preparation
A soil sample should be taken 46 months before planting to determine lime
requirements and soil nutrient levels. Soil phosphorus can also be adjusted
at this time or in the planting hole. Minimal tillage can be achieved with a
2 m-wide band cultivated where the trees are to be planted. Drainage should
be installed at this time to avoid ooding, with either contour or subsurface
drains. Windbreaks should be established prior to transplanting.
Transplanting and spacing

Transplanting should be done at the beginning of the wet season if there are
seasonal dry periods and no irrigation facilities. In the subtropics, planting
should not be performed if there is a risk of frost. Plants should have attained
a height of 3050 cm at transplanting, with the union of grafted or budded
plants placed approximately 15 cm above the ground. Trees should be
irrigated as soon as possible after transplanting, and wind and sun guards are
sometimes required.
Soursop trials suggest spacings of 44.6 u 67 m do not affect growth or
interfere with cultural practices. Spacings of 67 u 68 m are also used. A
triangular layout is recommended whatever planting distance is selected, with
the rows running north to south to avoid shading. For Rollinia, a 6 u 6 m or
7 u 7 m spacing is normally used (Vargas et al., 1999).
Irrigation practices
Annona species are grown without irrigation in many areas where rainfall is
well distributed. Except for pond apple (A. glabra), most Annona species can
stand periods of drought and prefer rather dry conditions. Adequate soil
12
Chapter 1
moisture is required to encourage vegetative growth, however, since owering

occurs on new growth. The amount and frequency of irrigation is determined
by experience for a particular location and soil type. Water stress should be
prevented during owering, fruit set and fruit development, as fruit are more
sensitive than leaves.
High soil moisture to increase humidity during the owering season may
prolong stigma receptivity and fruit set and growth. Low-rise sprinklers beneath
the tree canopy during owering can increase humidity. The stomata of Annona
species respond to RH not water stress. They will continue to lose water if the
humidity is greater than 80%, making maintenance of soil moisture crucial
(Marler et al., 1994).
Annona species, especially soursop and sweetsop, can show hardening of the
fruit pulp with brownish lumps apparently caused by the sudden movement
of water into the fruit (George et al., 1987). The effect is enhanced by boron
deciency. This disorder is reported as being common in north-eastern Brazil
under conditions with limited or no irrigation. As in most crops, any water stress
slows plant growth and decreases fruit size. Water quality is also important.
Sodium chloride can negatively affect growth, especially at a leaf concentration
of 0.3% sodium and can cause leaf burn and defoliation. Elevated levels of
chlorine and boron in the water are phytotoxic to fruit and leaves, and are
difficult to control (Pinto and da Silva, 1996).
Pruning
Training of trees begins 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, after which it should be headed back to produce lateral
branches. The lateral branches should be spaced 1525 cm above each other
and allowed to grow in different directions to develop a good scaffold. After
about 2 m, trees are left to grow naturally. Pruning is carried out when the
trees are dormant. For heavy fruit-bearing trees, pruning involves the removal
of lower limbs that touch the ground and branches in the center that may be
rubbing against each other. The aim is to allow sunlight access to the center of
the tree.
The soursop will, by nature, usually produce a symmetrically conical tree
and is well adapted to the central-leader system. It has an erect habit of growth,
and thus height has to be controlled if trees are to remain short to make harvest
and other tree-management practices easier. An alternative is to develop a
mushroom-shaped tree that is topped at 22.6 m. The fruit in this system are
borne on the lateral branches and hang down for ease of harvest. This process
starts with formation pruning, where plants are topped at 6080 cm once they
start growing following transplanting. After topping, three or four primary
13
branches are selected so that they are evenly spaced around the trunk and care
is taken that they do not arise too close together so that no weak zone occurs.
These primary branches can be topped at about 50 cm from their origin to
induce secondary branching. 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. Plants should not be allowed to grow above 2.54 m, depending
on the formation method used.
Diseased or insect-infested branches should be removed periodically, as
should branches growing in the wrong direction or with undesirable sprouting.
Any excess vegetation inside the canopy should be removed to allow greater light
penetration and ventilation. Rejuvenation by heavy pruning is occasionally
needed, but very severe pruning reduces subsequent fruiting.
Fertilization
Annona species have an indeterminate growth habit (axillary owering) and
applying nitrogen in a somewhat excessive amount does not greatly interfere
with oral initiation, as is the case with plants with a determinate growth
habit. However, excessive tree vigour is usually associated with reduced owering and yields in many trees.
Fertilization starts in the nursery, with in particular nitrogen applied in
small amounts. During transplanting some fertilizer, especially phosphorus, is
added to the bottom of the hole. In the vegetative growth period prior to fruiting,
phosphorus, potassium and sometimes calcium from dolomitic rock are
usually applied. Aviln (1975) found that soursop has a high requirement for
phosphorus and potassium. According to Pinto and da Silva (1996) a 10:15:15
or 10:13:15 ratio is adequate for growth and production. The fertilizer (250
g) of any of these formulas is applied in four applications (approximately 63
g each) per tree per year, during the rst 4 years. In the fth and following
years, this amount is increased to 1 kg/tree applied over four applications.
Observations in Hawaii and Mexico have indicated the desirability of providing
1.3 kg of a triple-15 fertilizer formulation during the rst year of production,
split into two applications. Each year thereafter, up to approximately the sixth
bearing year, the total amount can be increased by approximately 0.45 kg/tree/
year.
In Brazil, a recommendation of 40 g nitrogen per tree in the rst year to 180
g in year 5 and thereafter has been made. For phosphorus, the recommendation
was none for the rst year to 40180 g/tree/year in year 5 and thereafter; and
for potassium, from 3060 g/tree in the rst year to 60180 g/tree/year after
year 5, with the amounts being related to soil analysis results (Table 1.1).
14
Chapter 1
Table 1.1. Nitrogen (N), phosphorus (P) and potassium (K) requirements for soursop
according to the age of the plant and the availability of soil phosphorus and
potassium (Silva and Silva, 1997).
P-resin (g/dm3)
Age (years) N (g/plant)
K-exchangeable (g/dm3)
010
1120
>20
P2O5 (g/plant)
045
>90
4690
K2O (g/plant)
01
40
60
40
30
12
80
80
60
40
80
60
40
34
120
120
80
60
120
80
60
>4
180
120
80
40
180
120
60
Another recommendation suggests that non-irrigated bearing plants should be

fertilized with 3 kg ammonium sulfate, 660 g triple superphosphate and 500 g
potassium chloride (SCUC, 2006). This is applied in three equal portions during
the year, preferably at the start, the middle and toward the end of the rainy
season. The fertilizer should be lightly incorporated into the soils around the
tree. Soursop also responds well to manure applications: either 15 kg per plant
of decomposed cow manure or 34 kg of decomposed poultry manure.
In cool subtropical areas, most vegetative growth takes place during the
warmer months from spring to autumn. A reduction in nitrogen during the
winter minimizes new vegetative growth in young trees that are vulnerable to
cold temperatures.
As with other perennial fruit trees, soil and plant tissue analyses are the
techniques most used to evaluate the nutritional state of the plants. Soil
sampling in adult soursop orchards should be similar to that recommended
for other crops. For leaf sampling, the recommended method depends on the
age of the plant, the position of the leaf in the canopy and on the branch and,
as with many fruit crops, whether the branches are fruiting or not. Pinto and
da Silva (1996) recommended collecting 8- to 9-month-old leaves that are free
from fertilizer or agrichemical residues. The sample should consist of about
100 leaves for every 57 ha. Four leaves should be obtained from each of 25
randomly selected plants. The orchard should be divided into smaller plots
according to soil characteristics. Samples should not be taken from sick or
abnormal plants. Flowering time and periods of heavy rain should be avoided.
Select plants of similar size and age, and avoid recently fertilized plants.
Normal leaf concentrations for nitrogen and potassium in Brazil are 1.6- to
2.0-times greater than those from decient leaves. Comparing data from Aviln
(1975) and Silva and Silva (1997), there is a greater difference in Venezuela
than in Brazil between normal and decient leaves with regard to nitrogen,
while the variation is less for potassium (Table 1.2).
15
Table 1.2. Leaf nutrient concentration for soursop in Venezuela and Brazil.
Venezuela
(Aviln, 1975)
Brazil
(Silva et al., 1984)
Normal
17.6
2528
Decient
11.0
1316
2.9
1.4
1.1
0.60.7
Normal
26.0
26.1
Decient
12.6
26.4
Normal
17.6
10.8
Decient
10.8
4.5
0.2
1.51.7
Element
Concentration
Nitrogen (g/kg)
Phosphorus (g/kg)
Normal
Decient
Potassium (g/kg)
Calcium (g/kg)
Magnesium (g/kg)
Normal
Decient
Boron (mg/kg)
0.08
1.11.3
Normal
3547
Decient
614
Pest management
Diseases
A number of diseases of soursop have been reported (Table 1.3).
Anthracnose, caused by Colletotrichum gloeosporioides, is the most serious
disease on soursop, particularly in areas of high rainfall and humidity
and during the wet season in dry areas. This disease causes twig dieback,
defoliation and dropping of owers and fruit. On mature fruit, the infection
causes black lesions. Black canker (Phomopsis annonacearum) and diplodia
rot (Botryodiplodia theobromae) occur mostly on neglected trees and cause
similar symptoms of purplish to black lesions, resulting in mummied fruit.
Marginal leaf scorch is also caused by P. annonacearum, while B. theobromae
causes twig dieback. Diplodia rot has darker internal discoloration and
causes deeper, more extensive corky rot in fruit. Cylindrocladium fruit and leaf
spot is caused by a soil-borne fungus, C. 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, which spread along
the sides, enlarge, become dry and crack. Infection is skin-deep, but the fruit
become unmarketable. Control measures recommended are good orchard
sanitation with heavy mulching and lower-branch pruning to prevent
splashing of soil during heavy rainfall (Sanewski, 1991).
16
Chapter 1
Table 1.3. Major diseases of soursop.

Common name
Organism
Parts affected,
symptoms
Region or
country
Anthracnose
Colletotrichum
gloeosporioides
(Glomerella)
Flowers, fruit, leaves,

dieback, seedling
damping off
Universal
Armillaria root rot Armillaria luteobubalina
Roots, base of trees,

decline
Australia
Bacterial wilt
Pseudomonas
solanacearum
Tree wilt
Australia
Black canker
(diplodia rot)
Botryodiplodia
theobromae
Universal
Black canker
Purple blotch
Phomopsis
annonacearum
Phytophthora palmivora
Leaf scorch, twig

dieback, peel
blackening, graft
union rotting
As for diplodia rot
Rust fungus
Australia
Phakopsora cherimoliae
Spots on immature
fruit, fruit drop, twig
dieback
Leaves
Australia
Florida
Fruit rot
Gliocladium roseum
Fruit
India
Rhizopus rot
Rhizopus stolonifer
Fruit
Brazil
Seedling rot
Rhizoctonia solani
Cylindrocladium spp.
Seedlings
Universal
Insects
Insect pests of soursop occur in numerous growing areas (Table 1.4). One
of the most serious insects in Mexico, Central America, Trinidad, Surinam,
Colombia, Venezuela and Brazil is the Cerconota moth, which lays its eggs on
young fruit. The emerging larvae tunnel into the pulp, causing blackened,
necrotic areas. It is not uncommon to nd every fruit larger than 7.5 cm
infested. For this moth, the use of light traps is recommended, as well as
picking and burying fallen fruit. The use of specic approved insecticides and
the release of parasitoid have also proven effective (Escobar and Snchez,
1992). The Bephrata or Bephratelloides wasp is also widely distributed throughout the Caribbean, Mexico, Central America and central and northern South
17
Table 1.4. Major insect pests of soursop.

Common name
Organism
Parts affected
Country/region
Bephrata wasp
(soursop wasp)
Bephrata meculicollis
Fruit
Wasp
Bephratelloides
paraguayensis
Cerconota anonella
Fruit
Thecla ortygnus
Flowers,
young fruit
Young fruit
Stem, leaves
Fruit
Fruit
Fruit
Fruit
Leaves
Leaves,
owers
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
Cerconota moth
(soursop moth)
Thecla moth
Banana spotting
Mealy bug
Citrus mealy bug
Southern stink bug
Caribbean fruit y
Queensland fruit y
Potato leaf hopper
Red spider mite
Scale insects
Coconut scale
Amblypelta lutescens
Dysmicoccus spp.
Planococcus citri
Nezara viridula
Anastrepha suspensa
Bactrocera tryoni
Empoasca fabae
Several genera,
species
Saissetia coffeae
Aspidiotus destructor,
other genera and
species
Fruit
Universal
Caribbean
America. This wasp is considered to be the most important pest in Florida.

Considerable damage to the soursop fruit has been observed in Mexico and
Central America by the authors. The larvae infest the seeds and damage the
pulp as they bore through the esh to emerge when the fruit matures. Control
measures include preventive spays with approved products during initial
fruit growth. For both of these insects, very good control can be obtained by
bagging the fruit using plastic bags either with holes or with the basal end
open (Escobar and Snchez, 1992; Broglio-Micheletti et al., 2001).
The Thecla moth is widespread throughout parts of the Caribbean and in the
American tropics, but it is not considered to be as serious a pest as the Cerconota
moth and Bephrata wasp. Damage is primarily to the owers. The larvae feed on
ower parts, such as the perianth, stamen and stigmas, and the owers fail to
set fruit.
Mature-green annonaceous fruit have been shown to be rarely infested by
the Mediterranean fruit y (Ceratitis capitata) and the Oriental fruit y (Dacus
dorsalis), but they are found occasionally in tree-ripened fruit. Bait sprays and
eld sanitation are recommended measures to minimize fruit-y infestation.
Fruit bagging also provides protection.
18
Chapter 1
Mealy bugs and various species of scale insects are found universally and
usually become serious pests 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 owers and leaves
in the Tecomn area of Mexico during the prevailing dry period, with trees
showing heavy ower drop.
Weed management
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.
Shallow root systems limit the use of cultivation under the tree.
HARVESTING AND POSTHARVEST HANDLING

Harvesting season, yield and harvesting
The harvesting season is quite similar in most areas, especially for soursop
and sweetsop, differing only in range (Table 1.5). A major problem in
soursop cultivation is obtaining commercial yields and large fruit with a
symmetrical shape. To increase fruit set and size and achieve a better shape,
hand pollination has become an important aspect of cultivation practices in
Table 1.5. Peak harvesting seasons for soursop.

Country/region
Month(s)
Caribbean
Year round
Brazil, center
MaySeptember
Brazil, north-east
Year round
Florida
JuneNovember
Hawaii
JanuaryOctober
Indonesia
Year round
Mexico
JuneSeptember
The Philippines
JuneAugust
Colombia
Year round
Puerto Rico
MarchSeptember
19
some areas. Rootstocks have been shown to greatly inuence yield (Sanewski,
1991).
In Hawaii, soursop yields from trees grown in a marginal eld have shown
approximately 43 kg/tree on 4-year-old trees, increasing to 83 kg/tree on
6-year-old trees. In Paramaribo, Surinam, soursop yields of 54 kg/tree at 278
trees/ha have been reported.
Fruit is harvested when fully mature and rm. The skin-color changes as the
fruit approaches maturity. The immature soursop fruit is dark green and shiny,
losing its sheen and becoming slightly yellowish-green on reaching maturity.
Determining harvest time by dating oral anthesis is impractical as owering
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 is hand harvested and put into lug boxes or baskets. Harvesting is
more difficult and time-consuming for soursop, because the trees are generally
taller than those of other Annona species and the fruit are much larger. In
large soursop orchards, mechanical harvesting aids are feasible and accelerate
handling.
Rollinia fruit turn yellow at maturity and should be harvested before they
start to change color or as the process starts, but before they are completely
ripe. Ripe fruit are soft and difficult to handle. The fruit should be harvested very
carefully by cutting the peduncle with a sharp knife or pruning shears. Yields
can vary from 25 to 60 fruit/tree/year for 5-year-old trees, while 15-yearold trees can produce 100150 fruits/year (Vargas et al., 1999). Harvesting
in Brazil is normally done between January and June, 4 months after ower
anthesis.
Postharvest handling
Harvested fruit should be handled with care to prevent bruising of the
skin. This is especially important for fruit that are marketed for fresh
consumption. Firm soursop fruit need to be held after harvest for 47 days
at room temperature, with optimum quality processing occurring 56 days
after softening begins (Paull, 1983). The skin of the ripening soursop fruit
will gradually turn dark brown to black, but the esh is unspoiled. Storage
temperatures below 15C cause chilling injuries and a failure to develop
full avor. Pre-cooling of fruit is essential to help extend the shelf life.
Protuberances on the skin of fully ripe Rollinia are easily injured and they turn
brown to almost black, making the fruit unattractive (Morton, 1987).
When processed, soursop fruit are stored on racks in the shade and
inspected daily. All fruit that yield to nger pressure are removed for processing.
Slightly immature fruit will ripen but they lack the full avor and aroma, and
nectars prepared from the puree of such fruit have a at taste. Pulp-recovery
percentages have been reported to range from 62% to 85.5% (Paull, 1982).
20
Chapter 1
Differences in recovery percentages are caused by differences in equipment,

extraction methods, cultivar and cultural practices, including environmental
inuences. The number of seeds per fruit also inuences pulp recovery.
Compositional changes during fruit ripening

All Annona species bear climacteric fruit. Soursop respiration begins to
increase within a day of harvest and reaches its peak at days 68. Ethylene
production is initiated approximately 48 h after initiation of the respiration
rise, and reaches its peak at about the same time as the respiration peak
reaches a plateau (Paull, 1983). Total soluble solids increase from around
10% to 16% during the 3 days of ripening. The major titratable acids are
malic and citric acids. Days 6 and 7 are considered to be the optimum edible
stage and coincide with the peak of ethylene production.
UTILIZATION
Soursop fruit is marketed fresh to local markets. This fruit, of all the Annona
species, has the best processing potential because of the excellent avor
characteristic of the pulp and high recovery from large fruit. Unfortunately,
soursop has to be hand peeled and cored, an expensive and time-consuming
operation. The fragility of the skin and the fruits irregular shape and softness
limit machine processing.
Soursop pulp is viscous and requires dilution to produce a desirable nectar
viscosity; however, this diluted product is at and weak. To overcome this
dilution effect, the pH needs to be adjusted to 3.7 by adding citric acid and
sugar to 15% total soluble solids to create a desirable balance between acidity,
sweetness and avor. Unsweetened and sweetened soursop pulp processed
below 93C show no changes in organoleptic properties. Freeze preservation
produces a higher-quality product. Enriched pulp, sweetened or unsweetened,
can be processed and stored frozen for re-manufacture as various products or
reconstituted directly by the consumer. Puree can be used to prepare an iced
soursop drink or mixed with other juices, or it can be made into sherbets and
gelatin dishes. Soursop is a good source of potassium, riboavin and niacin
(Table 1.6).
Rollinia pulp is normally eaten fresh, though in some parts of Brazil it is
used to make a fermented wine. Sugar can be added to the pulp to make some
desserts. The seeds have insecticidal properties (Vargas et al., 1999).
21
Table 1.6. Composition of 100 g edible portion of soursop

(Wenkam, 1990) and Rollinia (Collazos et al., 1975, cited by
Villachica et al., 1996).
Constituent
Proximate
Water (g)
Energy
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Magnesium (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A
Seed/skin (%)
Soursop
Rollinia
80.10
247 kJ
0.69
0.39
18.23
0.95
0.58
85.0
53.0 cal
1.1
0.4
12.9
1.2
0.6
9.00
0.82
22.00
29.00
320.00
22.00
16.40
0.07
0.12
1.52
0
34
3.40
0.07
0.23
0.79
0
FURTHER READING
Bayogan, E.R. and Paull, R.E. (2008) Soursop Annona muricata. In: Janick, J. and Paull,
R.E. (eds) The Encyclopedia of Fruit and Nuts. CAB International, Wallingford, UK,
pp. 4246.
Campbell, C.W. (1985) Cultivation of fruits of the Annonaceae in Florida. Proceedings of
the Tropical Region of the American Society for Horticultural Science 29, 6870.
Coelho de Lima, M.A. and Alves, R.E. (2011) Soursop (Annona muricata L). In: Yahia,
E.M. (ed) Postharvest Biology and Technology of Tropical and Subtropical Fruits. Volume
4: Mangosteen to White Sapote. Woodhead Publishing Ltd., Cambridge, pp. 363
391.
Love, L., Paull, R.E. (2011) Rollina. University of Hawaii at Manoa, College of Tropical
Agriculture and Human Resources. Fruit and Nuts Publication F_N-21. Available
from: http://www.ctahr.hawaii.edu/oc/freepubs/pdf/F_N-21.pdf. Accessed August
20, 2011.
22
Chapter 1
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. Volume II: Subtropical and Tropical Crops.
CRC Press, Boca Raton, Florida, pp. 200206.
Pinto, A.C. (2002) Soursop. In: Crisstomo, L.A. and Nuamov, A. (managing eds) and
Johnston, A.E. (ed) Fertilizing for High Yield and Quality Tropical Fruits of Brazil.
International Potash Institute Bulletin, IPI, Horgen, Switzerland, pp. 202217.
REFERENCES
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composicin qumica de la guanbana (Annona muricata L.) cultivada en soluciones
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tropical fruits annonas. In: Scheaffer, B.C. and Andersen, P.C. (eds) Handbook of
Environmental Physiology of Fruit Crops, Vol II. Subtropical and Tropical Crops. CRC
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23
Nakasone, H.Y. (1972) Production feasibility for soursop. Hawaii Farm Science 21, 10
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Paull, R.E. (1982) Postharvest variation in composition of soursop (Annona muricata
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Produo. Embrapa-SPI, Braslia.
Pinto, A.C. de Q., Cordeiro, M.C.R., de Andrade, S.R.M., Ferreira, F.R., Filgueiras, H.A.
de C., Alves. R.E. and Kinpara, D.I. (2005) Annona species. International Centre for
Underutilized Crops, University of Southampton, UK. Available from: http://www.
icuc-iwmi.org/les/R7187_-_Annona%20monograph%202005.pdf.
Accessed
August 20, 2011.
Samuel, R., Pineker, W., Balasubramaman, S. and Morawetz, W. (1991) Allozyme
diversity and systematics in Annonaceae a pilot project. Plant System Evolution
178, 125134.
Sanewski, G.M. (ed.) (1991) Custard Apples Cultivation and Crop Protection. Information
Series Q190031. Queensland Department of Primary Industries, Brisbane,
Australia.
SCUC (Southern Centre for Underutilized Crops) (2006) Annona: Annona cherimola, A.
muricata, A. reticulata, A. senegalensis and A. squamosa. Field Manual for Extension
Workers and Farmers. University of Southampton, Southampton, UK.
Silva, A.Q. and Silva, H. (1997) Nutrio e Adubao de Anonceas. In: So Jos,
A.R., Souza, I.V.B., Morais, O.M. and Rebouas, T.N.H. (eds.) Anonceas, Produo e
Mercado. Universidade Estadual do Sudoeste da Bahia, Vitria da Conquista, Bahia,
pp. 118137.
Silva, H.A., da Silva, A.Q., Cavalcante, A.T. and Malavolta, E. (1984) Composio
mineral das folhas de algunas fruteiras do Nordeste. Anais do 7mo. Congresso
Brasileiro de Fruticultura (Florianpolis), pp. 320325.
Sousa, N.R. (2008) Rollinia mucosa Birib. In: Janick, J. and Paull, R.E. (eds) Encyclopedia
of Fruit and Nuts. CAB International, Wallingford, UK, pp. 6870.
Thakur, D.R. and Singh, R.N. (1964) Studies on pollen morphology, pollination and fruit
set in some annonas. Indian Journal of Horticulture 22, 1017.
Vargas, O., Alix, C., Lobo, A.D. (Authors), Duarte, O. and Sanchez, J. (Technical
Reviewers). (1999) Frutales y Condimentarias del Trpico Hmedo. CURLA; PDBL;
AFE/COHDEFOR; DICTA; SETCO; PROFORFITH, La Ceiba, Honduras.
Villachica, H., de Carvalho, J.E.U., Muller, C.H., Diaz, C. and Almanza, M. (1996) Anona
(Rollinia mucosa (Jacq.) Bailln), In: Frutales y Hortalizas Promisorias de la Amazona.
Tratado de Cooperacin Amaznica, Secretara Pro-Tempore, Lima, Peru, pp. 20
24.
Wenkam, N.S. (1990) Foods of Hawaii and the Pacic Basin. Fruits and Fruit Products, Raw,
Processed, and Prepared. Vol. 4, Composition. Research Extension series 110. HITAHR,
College of Tropical Agriculture and Human Resources, Honolulu, 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, 715.
24
Chapter 1
Yang, C.S. (1988) Application of plant growth regulators on Annona culture. In: Lin,
H.S., Chang, L.R. and Lin, J.H. (eds) The Application of Plant Growth Regulators on
Horticultural Crops. Symposium Proceedings. Special Publication No. 12, Taichung
District Agricultural Improvement Station, Changhua, Taiwan (Chinese, English
summary), pp. 305320.
2
BREADFRUIT, JACKFRUIT, CHEMPEDAK
AND MARANG
The family Moraceae includes the g and mulberry. The genus Artocarpus,
which includes breadfruit, jackfruit, chempedak and marang, contains about
50 species with milky latex. Most species are native to Asia, and 15 produce
edible starchy fruit that are frequently staples. The genus name comes from
the Greek words artos (bread) and karpos (fruit). The three most important
species are the more tropical breadfruit A. altilis (Parkinson) Fosberg (syn A.
communis, Foster; A. incisus L.; Communis incisa), the jackfruit A. heterophyllus
Lam. (syn A. integer [Thumb.] Merrill; A. integrifolius) and its close relative
chempedak A. integrifolia L., (syn. A. polyphema Persoon; A. champeden [Lour.]
Stokes).
Other Artocarpus species are also grown: A. odoratissimus (marang), A.
camansi (breadnut) Blanco, A. lakoocha Roxb. (monkey jack) and A. mariannensis
Trcul. (dugdug). Species with edible fruit that are not commercially grown,
but are collected and consumed in their native range, include A. anisophyllus
Miq., A. chama Buch-Ham., A. fulvicortex Jarrett, A. hypargyreus Hance, A.
kemando Miq., A. lanceifolius Roxb. subsp. lanceifolius and clementis Merr., A.
nitidus Trcul., A. rigidus Blume, A. sarawakensis Jarrett., A. sericicarpus Jarrett,
A. styracifolius Pierre, A. tonkinensis, A. Chevalier and A. vrieseanus Miq. (Love,
2008).
BREADFRUIT
Introduction
Breadfruit originates from New Guinea and possibly the Moluccas, with
numerous varieties spread throughout the islands of the Pacic. It has
been distributed throughout the humid tropics since the late 1700s. The
tree was the reason behind Captain Blighs voyage to Tahiti and the mutiny
on The Bounty (Spary and White, 2004). In many regions, the seeded and
25
26
Chapter 2
seedless cultivars have different common names. Seeded breadfruits are

called breadnut (English), kelur or kelor (Indo-Malaya) and kamansi or pakok
(Philippines), while seedless are sukun (Indo-Malaya) and rimas (Philippines).
Other names include arbre pain (French), sake (Thai and Vietnamese), rbol
del pan or panapen (Spanish), fruta pao (Portuguese) and ulu, uru, kuru, uto, mei,
lemae and mos (Pacic islands).
Two closely related species that possibly contributed to breadfruit are
breadnut A. camansi Blanco from New Guinea, the Indo-Malay region and
possibly the Philippines; and dugdug A. mariannensis Trcul from western
Micronesia. Breadnut is a wild ancestor of the breadfruit indigenous to the
lowlands of New Guinea, where it grows in ooded riverbanks, secondary and
primary growth forest, and freshwater swamps, and in cultivation. It may
also be indigenous to the Moluccas and possibly the Philippines. Dugdug is
morphologically very distinct from A. altilis and grows wild in Palau, Guam
and the Northern Mariana Islands. Introgression between the two species has
occurred in Micronesia and there are a number of hybrid varieties (Ragone,
1997).
Breadfruit is principally grown as a subsistence crop in most areas of the
world, with Pacic and Caribbean islands being the major production areas.
Fruit range from 0.2 to 4.5 kg, depending on the cultivar. Yields vary from as
low as 50150 to as many as 700 fruit/tree, with an estimated yield of 1650
t/ha based on a density of 100 trees/ha. Canopy volume is a good measure of
the potential yield.
Ecology
Soil
A variety of soils with sufficient depth and good drainage are suitable. Soils
with high levels of organic matter and fertility are recommended. On Pacic
islands, breadfruit does grow on shallow coralline soils, demonstrating its
considerable varietal adaptability.
Climate
Regular rainfalls of 15003000 mm/year and humidity of 7090% are
preferred. Rainfall is necessary for vegetative growth, owering and fruit
growth, with a bimodal pattern preferred with a 36 month dry season. The
tree is sensitive to chilling, with no growth at temperatures of 5C or lower.
The tree is well suited to hot, humid, tropical lowlands, with temperatures of
up to 38C and altitude below 1500 m. Best production takes place below 650
m in the tropics. Full sun is required, and no photoperiodic events have been
noted.
Breadfruit, Jackfruit, Chempedak and Marang
27
General characteristics
Tree
This fast-growing evergreen tree can grow up to 30 m in humid and wet
areas, and can live for as long as 90 years. The tree is partially deciduous
under drought or during the dry part of a monsoon climate. The alternate
and ovate leaves, which are 2075 cm in length, are dark green with none
to as many as 13 lobes (Fig. 2.1A). The trunk is straight, with thick branches
terminating in branches of 1020 cm in length with two large deciduous
stipules enclosing the terminal bud. Root suckers begin bearing in 35 years
and seedling plants in 810 years.
Fig. 2.1. Breadfruit leaf (A), male (B) and female (C) owers and immature fruit (D).
Jackfruit and chempedak inorescences are similar in shape. (From Nakasone and
Paull, 1998, with permission from CAB International.)
28
Chapter 2
Flowers
This monoecious species has the staminate (Fig. 2.1B) and pistillate
inorescence on a 415 cm peduncle in separate leaf axils. The drooping,
spongy, club-shaped male (1520 u 34 cm) inorescence has minute owers,
each with a single stamen. The globose pistillate inorescence (610 cm) is
covered with numerous tiny owers on a spongy axis. Each pistillate ower is
reduced to a tubular calyx with a two-celled ovary, and a two-lobed stigma on
a short style.
Rain encourages vegetative growth and owering. Some cultivars can ower
throughout the year under the right environmental conditions. Crosspollination is assured by the staminate inorescences maturing before the
pistillate. Following wind or insect pollination, fertilization occurs over
36 days in seeded cultivars. A high percentage (75%) of the orets are set,
with the percentage being reduced in rainy weather. This reduction suggests
that pollination is necessary to stimulate parthenocarpic growth. However,
pollination is difficult as the rudimentary perianth acts as a physical barrier to
pollination and argues against the fruit being parthenocarpic. Pollen sterility
is also a factor contributing to reduced fertility and seed production.
Paclobutrazol, naphthalene acetic acid and ethephon inhibit vegetative
growth but fail to stimulate owering. In the West Indies, methanol spray in
the dry season has been found to enhance vegetative growth and bring about
earlier and more profuse owering. Solar radiation signicantly inuences
the onset of owering and female inorescence production. Fruit set is also
related to tree width, with fewer fruit setting on trees with a greater width;
this is possibly associated with uneven light interception, and suggests that
tree management including pruning and plant spacing can impact fruit
yield.
Fruit
The fruit develops from the entire inorescence as the perianths of the
individual owers attached to the central axis or core, fuse together and
become eshy (Fig. 2.1D). The fruit is normally round to oblong, sometimes
cylindrical and 1030 cm in length. The thin, reticulated skin is pale green or
yellowgreen when the fruit is mature, turning yellow-brown when ripe. The
core is surrounded by a pale-yellow or creamy white edible pulp (Fig. 2.2).
Most cultivars are seedless, but the seeded wild types have 10150 brown
seeds of 2.5 cm in length.
Depending on the stage of maturity and cultivar, the core and the esh
will exude a white viscous latex that discolors greenish or reddish-brown on
exposure to air. The skin also exudes latex and dried, hardened drops are an
indication of fruit maturity in some varieties. The fruit matures in 1321 weeks
from the time the pistillate inorescence is rst detectable in the terminal leaf
29
Fig. 2.2. Mature breadfruit (front right), jackfruit leaf and bud (top right) and
jackfruit (back).
sheath (Fig. 2.3). Optimum maturity occurs at 1519 weeks and fruit at this
stage are preferred, as it provides a 5-week period during which the fruit can be
harvested and still be acceptable to the consumer.
Seedless cultivars are typically triploid (2n 84), with the reduced seed
number in diploid (2n 56) cultivars probably due to accumulated mutations
in clonally propagating plants using root suckers. Numerous varieties have
been described (Table 2.1), although few have been compared at the same
location.
Triploidy is common in seedless types. Many Micronesian cultivars are
hybrids or triploids of A. altilis, A. camansi and A. mariannensis, with some of
the diploids being fertile. Molecular data support the conclusion that breadfruit
is a complex of these three species. Seedy types are more common in the
Chapter 2
Total Sugars (g)
30
Fig. 2.3. Growth of breadfruit, showing the pattern of fruit diameter, fruit fresh
weight, total sugars and fruit starch. (Redrawn from Worrell et al., 1998.) AIS,
alcohol-insoluble solids.
Table 2.1. Fruit characteristics of selected widely distributed breadfruit cultivars.
(From Ragone, 2011 and others.)
Variety
Origin
Shape
Flesh, seeds
Maafala
Polynesia
Small oval
Maopo
Polynesia
Puou
Polynesia
Oval to broad
ovoid
Round, oval or
heart-shaped
White esh, seedless,

occasionally with
one or two seeds
Pale white or creamy,
seedless
Creamy paleyellow, seedless,
occasionally with
one or two seeds
Light yellowgreen,
seedless
Creamy-yellow,
seedless
Meinpadahk Micronesia Oval to

asymmetrical
Yellow-heart Caribbean Oval
Fruit weight
(average kg)
0.8
2.5
1.9
1.1
2.0
31
western South Pacic. Evaluation and selection trials are currently underway
in an extensive germplasm collection of Pacic Island breadfruit cultivars at
the National Tropical Botanical Garden in Hawaii. This organization maintains
an extensive website describing many varieties, although no breeding work has
been reported. Variability has been observed in growth form, leaf shape, fruit
quality, time to bearing, seasonality, keeping quality of fruit and salt tolerance.
Cultural practices
Propagation and nursery management
Seeds have 9095% viability as soon as they are removed from the fruit, but
this is lost in 2 weeks. Breadfruit is typically clonally propagated traditionally
from root shoots or cuttings, although losses can be high with root suckers.
Roots of 1.56 cm in diameter are cut into sections from 12 to 30 cm long.
These are placed in clean, washed sand or potting media and kept moist.
The roots can be placed horizontally below the surface of the medium or
diagonally with the upper few centimeters exposed. The percentage of rooting
ranges from 80% to 85% and takes 25 months, if kept well watered. Some
success has been achieved with air layering, budding and grafting.
There is interest in grafting seedless cultivars to rootstock of atoll-adapted
seeded and seedless types. The cultivar Maafala has been used as a rootstock
in Samoa.
Field preparation
No special orchard preparation is reported.
Plants are set out at 715 m, depending on the variety and growing
conditions. Shade is provided for the rst year as the young trees growth
resumes.
Continued vegetative growth requires irrigation, especially during periods of
drought. Irrigation reduces fruit drop during the dry season.
Pruning
The tree may be pruned to improve its shape, although regular pruning is not
normally carried out. Trees that have grown too tall to readily harvest are
often topped or trimmed back to keep the tree at a more convenient height.
Fruit-bearing branches may break during heavy fruiting periods and these
need to be removed. Some growers suggest that pruning branches that have
borne fruit stimulates new shoots and limits tree height.
32
Chapter 2
Fertilization
General requirements have not been determined. The application of 100200
g ammonium sulfate per tree 1 month after planting and again at 6 months
is recommended (Coronel, 1983). The amount should be gradually increased
until the trees start to produce fruit; thereafter, 5001000 g complete fertilizer
may be applied to each tree twice a year. A full bearing tree may require at
least 2 kg complete fertilizer per application. Mulching and the application of
organic manure two to three times a year, sometimes mixed with fertilizer, are
used to increase and maintain growth rate.
Pest management
Pingelap causes dieback from the top branches, and tree death is caused by
an unknown organism with no method of control. This was a major disease
in Micronesia in the 1960s. Other diseases include dieback (Fusarium,
Pythium and Rosellinia), leaf spot (Cercospora), leaf rust (Uredo artocarpi),
root rot (Phellinus noxius), fruit rot (Phytophthora, Phyllosticta and Rhizopus)
and stem end rot (Phomopsis, Dothiorella). Fruit rot tends to be more of a
problem on rough- than smooth-skinned varieties. Control measures involve
removing affected fruit from the tree and not allowing fruit to ripen on the
tree or rot on the ground. Mealy bugs, scales and twig borers are the major
pests, with no control usually practiced. The fruit is a fruit-y host. Recently,
mealybugs have become a major problem affecting breadfruit in Kiribati,
while Phellinus is causing crown rot and dieback of trees in Samoa (Brooks,
2002).
Weed management
Mulching around the base of the trunk is practiced to control weeds, conserve
moisture and provide nutrients.
Orchard protection
A windbreak is not usually necessary. However, branches may break in high
winds during periods of heavy fruiting. Breadfruit trees are occasionally used
as windbreaks and shade for other crops.
Harvesting and postharvest handling

Mature-green fruit are harvested as a starch vegetable, while some people
prefer to eat the ripe sweet fruit. Harvested green fruit produce copious latex,
especially from the cut peduncle and injuries on the fruit. Maturity is indicated
by larger size, a slight change in the skin color to yellowish-green, small drops
of latex on the rind and rm esh texture. In addition, the segments are more
rounded and smoother than in less mature fruit. As the fruit starts to ripen,
the skin changes to a yellowish-green and begins to soften. Latex needs to be
33
allowed to drain from the fruit after harvest and before washing in water to
avoid latex stain.
Fruit that are physiologically mature, with green skin, rm esh, uniform
shape and free from decay, sun-scald, cracks, bruises and mechanical damage,
are marketed. Fruit at different growth stages are harvested to meet different
market needs. There are no US or international grade standards. The fruit is
graded according to appearance, blemishes, maturity and size. Various fruit
counts are used depending on berboard carton size (918 kg). Fruit are sold
on a weight basis. Telescope two-piece berboard cartons or one-piece cartons
with dividers to minimize fruit movement and rubbing are used.
The fruit is cooled as soon as possible after harvest and stored at 1214C
and 9095% relative humidity for a maximum of about 20 days. Hydrocooling
is not recommended as it leads to skin browning. Film wrapping and coatings
delay the softening and skin discoloration of fruit stored at 13C. Controlled
atmosphere studies have indicated that at 12C, the best storage atmosphere
is 25% O2 and 5% CO2 for up to 3 weeks. Chilling injury symptoms begin to
develop within 7 days at 10C. Symptoms are a brown scald-like discoloration
of the skin, failure to fully soften, poor avor development, and an increase in
decay.
Utilization
Breadfruit is typically eaten while still mature, rm and starchy (Fig. 2.3). In
some areas, round immature fruit are also eaten cooked. The sweet ripe fruit
is eaten as a dessert and can be used to make pies, cakes and other sweets. The
fruit can be roasted, boiled, dried, pickled, used in bread making or fermented,
while slices can be fried or stored in brine. The edible esh comprises 70% of
the fruit, and is 6085% water, 1.22.4% protein, 2237% carbohydrate and
0.20.5% fat (Table 2.2). The carbohydrate in mature fruit is mainly starch.
Alcohols are the major aroma compounds, with cis-3-hexenol3-hydroxy2-butanone, cyclohexanediol and 2-pentanone making up 62% of the
detected volatiles. The cooked seeds are also eaten, and contain 47.766.2%
water, with 13.319.9% protein, 26.676.2% carbohydrate and 2.529%
fat. The leaves and fallen fruit are fed to animals. The collected latex is used
medicinally and as a caulk, glue and chewing gum.
JACKFRUIT AND CHEMPEDAK

Introduction
The jackfruit A. heterophyllus Lam. (Moraceae) and its very close relative
chempedak A. integer (Thunb.) Merr. originated in India and Malaysia.
34
Chapter 2
Table 2.2. Composition of 100 g edible portion of breadfruit, jackfruit, chempedak

and marang (Dignan et al., 1994; Wenkam, 1990).
Constituent
Edible portion (%)
Proximate
Water (g)
Energy (kcal)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
Vitamin C (mg)
Breadfruit
Jackfruit
Chempedak
Marang
70
28
22
2433
62
561
1.3
0.18
37
1.45
1.2
83
301
1.6
0.2
25.4
5.6
2.2
67
490
2.5
0.4
25.8
3.4
1.2
21
0.26
48
551
13
37
1.7
26
292
48
40
1.1
5
246
25
0.12
0.06
1.54
41
20.5
0.06
0.4
66
7.9
0.15
0.5
48
17.7
65.784.2
265510 kJ
0.81.5
0.20.3
32.4
0.60.77
0.50.8
17
2.1
35
Jackfruit is also known as jacquier (French), nangka (Javanese and Malay),

langka (Philippines), khnaor (Cambodia), makmi, khanum, banum (Thailand)
and mit (Vietnamese). The English name is most likely derived from the
Portuguese jaca, taken from the Malaya tsjaka.
Jackfruit has been spread to Sri Lanka, southern China and south-east Asia,
and further to tropical Africa. It was probably introduced into the Philippines
in the 12th century and domesticated soon thereafter. The writings of Pliny the
Elder as early as 100 AD mention jackfruit as being essential to the traditions
in its place of origin (Campbell and Ledesma, 2003). The tree is still highly
regarded by subsistence farmers from India and through south-east Asia for its
fruit, timber and medicinal uses.
Chempedak, also known as bankong, baroh (Malaysian/Indonesia),
sonekadat (Burmese), champada (Thai), cempedak, jack tree (English), kathal,
kathar (Hindi), campedak, cempedak, comedak (Javanese), chakka, pilual (Tamil)
and mit to nu (Vietnamese) is distributed in Burma, peninsular Thailand and
Malaysia, the Indonesian islands and western New Guinea. Chempedak is
separated from jackfruit by having a smaller size with a slender peduncle, a
male inorescence that is pale green to yellow and not dark green, smaller and
roundish fruit with a thinner rind, and more juicy esh that is a darker yellow
35
when ripe. In addition, the embryo radicle is immersed while in jackfruit it is

supercial (Jarrett, 1959). Chempedak is restricted to south-east Asia, with
some trees in Australia and Hawaii, while jackfruit is spread throughout the
tropics.
Ecology
Soil
A variety of well-drained soils with a pH 57.5 can be used for jackfruit. Deep
alluvial sandy and clay loams are preferred. The soils used for chempedak are
normally uneroded and well-drained, although the tree tolerates temporary
water-logging.
Climate
Cold, drought and ood tolerance limits the distribution of jackfruit to areas
with more than 1500 mm rainfall evenly distributed throughout the year,
without a prominent dry season. A warm and humid frost-free climate with
minimum temperatures of 1622C and mean temperatures of 2530C,
an altitude below 1000 m, and regions 25 north and south are desirable for
good jackfruit bearing. It is grown in protected subtropical regions 30 north
and south. Temperatures below 5C severely damage trees and frost will kill
developing shoots and fruit, and sometimes main branches. The trees do not
do well in exposed locations with drying winds. They have some salt tolerance,
but poor drought and ood tolerance.
Chempedak is found at 01200 m in areas with a mean annual temperature
of 1347C and mean annual rainfall of 12502500 mm. It is frequently an
understory tree.
Tree
These monoecious, evergreen, latex-producing trees reach up to 25 m in
height with a straight stem that branches near the base at an angle of 3288.
All parts of the plant produce a milky-white gummy latex. The diameter of
the normally dome-shaped dense canopy is 3.57 m in 5-year-old trees. The
trunk is rarely buttressed with a girth of 3080 cm and a grayish-brown,
rough, uneven, somewhat scaly bark. Minute white hairs up to 0.5 mm long
are found on the surface. The tree produces a long taproot.
The glossy leaves are 425 u 212 cm (Fig. 2.2) and are usually hairy, with
a dark-green top and pale-green underside. The leaves are arranged alternately
on horizontal branches and spirally on ascending branches. Midrib and main
veins are greenish-white to pale greenish-yellow. At the nodes, the stipules are
36
Chapter 2
fused around the stem that leaves an encircling scar after the leaf abscises.
Chempedak has long wiry brown hairs (<3 mm long) on the leaves, stipules
and twigs.
Flowers
The owers in mature trees are found on short shoots from the trunk and
older branches. In young trees, the fruit are borne on branches. The elongated
hanging or drooping male inorescence is 515 cm long and 24.5 cm wide,
and produced singly. The whitish- or dark-green spikes have a smooth skin
that becomes yellowish and rough when mature. The female spike is either
solitary or paired and elliptical or round, with rough, light to dark-green skin,
515 cm on a 89 mm thick peduncle. When young, the male and female
inorescences are enclosed by a pair of stipules that abscise. Twice as many
male than female inorescences occur on one tree.
The chempedak male inorescence is cylindrical, 35.5 cm long and 1 cm in
diameter. The male and female inorescences are similar to those of breadfruit.
Seedling trees start to bear in 414 years, with no photoperiodic response
reported. In suitable hot, humid conditions with evenly distributed rainfall,
jackfruit bear owers and fruit throughout the year. In areas with distinct
wet and dry seasons, owering occurs in the rst 2 months of the dry season
and the wet season. A load of fruit, however, may suppress further owering.
The male inorescence matures 35 days before the female. The sticky yellow
pollen has peaks of release between 2.004.00 am and 4.006.00 pm, with a
sweet scent that attracts small insects; however, the owers may be also wind
pollinated. Anthesis commences 23 weeks after emergence and lasts about
2 weeks. The ower normally rots before abscission, attracting numerous
insects by the smell. The stigmatic surface is composed of papillae that
becomes sticky 12 weeks after exertion and remains so for a further 2 weeks.
Jackfruit may be an outcrossing species with some self-incompatibility.
The juvenile period for chempedak from seed is 36 years, and 24 years
for clonal trees. Chempedak is more seasonal than jackfruit, with blooms
being more common in February to April and August to October in peninsular
Malaysia. In western Java, the fruit owers in July and August with fruit
ripening between September and December. Female ower heads are found
only on cauliorous shoots, while most male heads are on peripheral shoots of
the canopy, possibly to facilitate pollination. Chempedak provides sticky pollen
to attract diverse nocturnal insects and is pollinated by them. Female heads
offer a protein-rich liquid. Stigmas remain receptive for 12 weeks.
The use of potassium nitrate and plant growth regulator sprays to induce
owering has been trialed without success. Potassium nitrate sprays do induce
vegetative growth.
37
Fruit
The multiple fruit are pear- or barrel-shaped syncarps, borne on a 510
cm stalk (Fig. 2.2), with jackfruit having a larger fruit (4.550 kg) than
chempedak. Each achene that makes up the syncarp is indehiscent, oneseeded, and 410 cm long and 24 cm wide when mature. The fruit is pale or
dark green when young, turning to greenish-yellow, yellow or brownish when
mature. The thick rubbery rind (1 cm) in jackfruit has short blunt spines. The
jackfruit receptacle is not separable from the waxy, rm-to-soft, golden yellow,
eshy, edible perianth (2540% of the total fruit) that surrounds the seed (5%
of the total weight). Unfertilized owers develop as strap-like tissue between
fertilized developing fruitlets. The fruit can have up to 500 seeds, 24 cm
long by approximately 2 cm, with each surrounded by a horny endocarp and
subgelatinous exocarp. The seeds are rm and waxy, and weigh up to 15 g.
Initial fruit growth is rapid after stigma emergence and for the rst couple
of weeks after anthesis. Fruit drop is then signicant (approximately 35%) with
the peak occurring 6080 days after anthesis. Fruit growth follows a sigmoid
growth pattern (Fig. 2.4). The fruit matures in 34 months for different varieties
and may take up to 6 months or longer when grown at higher altitudes and in
cooler areas.
Length and Girth (cm)
High variability occurs in both jackfruit (2n = 56 [tetraploid]) and chempedak

(2n = 56) characteristics: length of juvenile stage, seed germination, tree
vigor, fruit shape and size, esh appearance, amount of latex, edible esh,
Time After Fruit Set (Days)
Fig. 2.4. Change in jackfruit cv. NS1 girth and length after fruit set. (Redrawn from
Muda et al., 1996.)
38
Chapter 2
avor, aroma and fruit maturation time. No longstanding breeding programs

have been undertaken. Jackfruit and chempedak occasionally hybridize and a
clone has been selected in Malaysia called Nangka-chempedak CH/NA. This
selection Cheena has smaller fruit (2.5 kg) and 33% edible esh. This ability
to hybridize between the two species attests to their close relationship.
Ripe jackfruit are divided into two types based on edible pulp. The rst type
has thin, brous, soft, edible esh, acid to very sweet with a strong aroma.
The other type has thick, rm to crisp esh with less aroma. There are many
varieties of both types. Some are more suited to canning than other fresh fruit
varieties. A number of chempedak clones have been selected in Malaysia. Some
of the preferred clones have an attractive orange esh and higher yields.
Australian selections are Black Gold, Cochin and Golden Nugget. All
three have deep-orange esh. Cochin has little latex, while Black Gold has a
strong, sweet avor. Dang Rasimi and Mong Tong (Golden Pillow) are two Thai
cultivars. These tend to have a mild sweet avor. Dang Rasimi has larger fruit
(8 kg) with 32% edible esh. Tabouey is a popular Indonesian selection with
an average weight of 12 kg and crunchy, pale-yellow esh. Twenty types
of jackfruit have been identied throughout India and are currently being
perpetuated through vegetative propagation. Torres, Jo and Jo2 are major
cultivars in the Philippines, while 13 cultivars have been characterized in Sri
Lanka.
An extensive selection program in Malaysia has lead to the selections
J-30 and J-31. These produce fruit of 7 and 12 kg, respectively, with
3638% edible esh. The deep-orange rm esh of J-30 has a sweet taste
and only a slight aroma, while the deep-yellow esh of J-31 has a strong
aroma. Another Malaysian program evaluated 400 samples, which led to
the hybridization of CJ1 and CJ6 from which the Mastura cultivar was
developed. Commercial plantings of Mastura began in 2000 in Malaysia.
It has a golden color with a sweet aroma, and is rapidly developing into a
popular cultivar in the region.
Cultural practices
Seed from selected trees is the major means of propagation. The seed loses
viability within 3 months of removal from the fruit, so is planted immediately.
Seedlings are best grown under shade. Germination can be improved by
soaking in naphthalene acetic acid or gibberellic acid solutions.
Root cuttings are used to propagate a desirable tree, with stem cuttings and
air layers also being successful with some varieties. Grafting and budding are
now widely used in India and south-east Asia. Jackfruit can also be propagated
in vitro. Budding, grafting and inarching are made onto 12-month-old root
stocks of A. integer, A. heterophyllus, other Artocarpus species and the same
39
species. However, the suitability of these rootstocks has not been evaluated in a
range of environments.
Chempedak is generally grown from seeds taken from ripe fruit with desirable
qualities. The seeds are recalcitrant and do not remain viable for long after
removal from the fruit, and are sown immediately after cleaning with water.
Chempedak can be grafted to like rootstock, and some success with grafting
chempedak to jackfruit and other Artocarpus species has been reported.
Field preparation
Orchards are prepared as for other tree crops.
Jackfruit transplanting needs to be carried out with care to avoid damage
to the tap root and is best done before the trees are around 1 year old. The
traditional spacing of 612 m on a square or triangular pattern has been
recommended for these slow-growing trees. Narrower spacings of 3 m
between trees in a row are now common in trees pruned to 35 m.
Chempedak seedlings are ready to transplant in 1 year and are placed 1214
m apart, generally at the onset of the rainy season. Again, care is taken not to
damage the long taproot.
Due to poor drought tolerance, irrigation is required especially during
establishment. In the west Bengal dry season, watering with 30 l per 8-yearold plant at 30-day intervals has been found to signicantly increase fruit
retention, fruit weight and date to rst harvest (Table 2.3). Drainage is
essential if the land is subject to ooding.
Table 2.3. Effect of watering and grass mulching during the dry season on jackfruit
production. Eight-year-old trees received 30 l water per plant every 30 days (Ghosh
and Bera, 2006).
Treatment
Control
Mulching, no irrigation
Irrigated from November to
April, with mulching
Irrigated from December to
April, with mulching
CD at 5%
Fruit
retention
(%)
Fruit
weight
(kg)
Fruits per
plant
Date of rst
harvest
50
60
83
3.7
3.5
5.5
9
10
17
4 June
15 June
20 June
77
3.9
21
28 June
4.8
0.2
1.2
40
Chapter 2
Pruning
Shoots are sometimes thinned and branches cleared to allow harvesting
access, although often it is only the dead wood that is removed. Newer pruning
strategies aim for a tree that is 35 m high.
Fertilization
The Malaysian recommendation is for nitrogen, phosphorus, potassium
and magnesium (ratio 8:4:2:1) at 30 g/tree for those 6 months old, doubled
every 6 months to 2 years. Older trees receive 1 kg/tree at a ratio of 4:2:4:1,
every 6 months. Higher rates of 23 kg are recommended in the Philippines.
Application is before and at the end of the wet season around the outer
canopy drip line.
Pest management
Seed and blossom rots, leaf spots, pink disease and fruit rot occur on jackfruit.
The blossom and fruit rot are caused by Rhizopus artocarpi to both developing
and mature fruit. Bacterial dieback (caused by Erwinia canetorora) can be a
problem with most Artocarpus species. Corticium salmonicolor causes pink
disease of jackfruit. Root rots caused by Fusarium and Phytophora are major
problems, especially if the root system is ooded for a few days. Leafspot
caused by Phomopsis artocarpina, Colletotrichum lagenarium and Septoria
artocarpi is a problem in many areas.
Jackfruit is reported to be attacked by shoot borers, bark borers, bud weevils,
spittle bugs, mealy bugs, scale insects and aphids. Larvae from oriental jackfruit
y (Dacus umbrosus Fabricius and D. dorsalis) have been found in jackfruit and
marang, but are controlled with modern baits and protective bags covering the
fruit as it develops. In Asia, monkeys, bats and elephants are common pests.
Weed management
Once the tree is established, weeds are not a problem because of the dense
shade under the canopy. Weeds should be controlled between trees by
cultivation and the use of mulch.
Orchard protection
The tree can withstand moderate wind and is occasional planted as a
windbreak with closer spacing. Depending on wind strength and duration, a
windbreak may be required in commercial orchards.

Fruit quality after harvest is very dependent on maturity at harvest (Fig.
2.5). Fruit maturity can be judged by a dull hollow sound when tapped, skin
color changing from green to greenish-yellow or yellowish-brown, and a
41
Acidity (% Citric)
Solids and Starch (%)
Time After Fruit Set (Days)

Fig. 2.5. Changes in titratable acidity, total soluble solids and starch during jackfruit
growth and development. (Redrawn from Muda et al., 1996.)
characteristic odor and attening of the surface spines. The tapping method
is regarded as being the most reliable. On peninsular Malaysia, the nearly
mature fruit are often wrapped in palm leaves or bagged ostensibly to protect
against bats, rats and fruit ies, and to attract ants that keep other insects
away. Fruit are harvested by cutting the peduncle with a knife and handled
carefully to avoid mechanical injury. The latex is allowed to drain in the
eld, and fruit are then moved from the orchard. Latex ow is greater early
in the morning and least in late morning and early afternoon. The fruit are
sometimes allowed to fall and must be collected daily because they have a shelf
life of only 23 days. Letting the fruit fall can cause damage, loss of shelf life
and premature ripening.
Grading is not normally practiced and fruit are rapidly transported to the
markets. Fruit from larger orchards and at packing sheds, and accumulation
sites will be graded on size and appearance. A large fruit would be more than 15
kg and small fruit less than 78 kg.
This climacteric fruit shows the typical increase in respiration production
of a strong sweet aroma, softening, increase in aril yellow carotenoids and
conversion of starch to sugars over the 79 days of ripening (Fig. 2.5). During
ripening, tannins also decline while acidity shows little change. Fruit harvested
after 12 weeks in tropical areas should be organoleptically acceptable after
ripening. Mature undamaged fruit can be stored at 1012C for 23 weeks.
Fruit ripen in 37 days at 2227C, depending on the stage of maturity at
harvest.
42
Chapter 2
World production and utilization

World production
Jackfruit is spread throughout the tropics, while chempedak is mainly
restricted to south-east Asia with some trees in Australia, Hawaii and
elsewhere. Jackfruit can produce 20250 fruit/tree/year and sometimes up to
500, with each fruit weighing from 1030 kg. Both species have a tendency to
be somewhat biennial in bearing. Commercial yields average 70100 kg/tree/
year. Orchards in Malaysia report a yearly average of 17,000 kg/ha. For the
year 2000, the Philippines reported 11,735 ha of jackfruit in production with
4.08 t/ha. India is probably the largest producer.
Chempedak is widely grown in southern Thailand and peninsular Malaysia.
It is also cultivated in Indonesia, especially in Sumatra, Borneo, Sulawesi, the
Moluccas and Irian Jaya, as well as in west Java. Chempedak yields are similar
to those of jackfruit.
Utilization, including nutritional principles
Fruit are eaten unripe at 2550% full size as a vegetable or ripe as a fruit
(Fig. 2.6). Jackfruit is a good source of carbohydrates and vitamin A and
a fair protein source (Table 2.2). Chempedak is very similar to jackfruit in
composition, although many Westerners nd the musky smell and taste of
chempedak only slightly less objectionable than that of durian. The major
volatiles of jackfruit are esters (31.9%), which are important contributors
to the avor. Esters vary with variety and include isopentyl isovalerate,
butyl isovalerate, butyl acetate, ethyl isovalerate and 2-methylbutyl acetate.
Chempedak fruit volatiles are 37.4% alcohols and 32.2% carboxylic acids. The
main constituents are 3-methylbutanoic acid (28.2%) and 3-methylbutan-1ol (24.3%). The 3-methylbutanoic acid has a rancid cheese, sweaty and putrid
smell, and most likely contributes to the musky odor of chempedak.
Because of the large fruit size, ripe jackfruit are frequently cut open and
sliced into pieces for sale (Fig. 2.6). The copious quantities of sticky latex, with
the amount varying by cultivar, makes preparation and clean-up difficult. The
application of vegetable oil to knives and surfaces, along with the use of rubber
gloves, reduces the clean-up problem. The edible pulp is removed, with the fruit
placed in trays, given a plastic overwrap, and sold (Fig. 2.6). When used as a
vegetable, the fruit is peeled, sliced and boiled, then seasoned or mixed with
other food. Jackfruit curry is fairly common in south India and Malaysia. The
seeds of both species are boiled or roasted, then eaten. Jackfruit pulp is canned
in syrup or used to make jams, jellies and ice cream. Dried jackfruit and jackfruit
chips are growing in popularity.
43
Fig. 2.6. Young (A) and mature (B) jackfruit after harvest are allowed to fully ripen
(C). The edible esh is removed (D) and placed in trays with a plastic wrap for
sale (E).
44
Chapter 2
MARANG
Introduction
Marang, A. odoratissimus Blanco (Moraceae), is also called terap (Malaysia),
loloi (the Philippines), khanun sampalor (Thailand) and keiran (Indonesia). The
synonyms are A. tarap Becc. and A. mutabilis Becc. Marang is generally smaller
than jackfruit. It is round to oval, with short brittle spines protruding from a
thick and eshy rind. It originated in Borneo and, similar to jackfruit, has been
introduced into neighboring countries.
Anthesis occurs about 3 weeks after the emergence of the inorescence. The
stigma remains receptive for 12 weeks after emergence (Dela Cruz, 1992).
Marang secretes a nectar-containing fructose to attract diverse nocturnal
insects, by which it is pollinated. Female heads offer a protein-rich liquid
(Momose et al., 1998).
Ecology
Marang has similar temperature requirements to jackfruit. In the Philippines,
it is generally grown in shaded areas up to 800 m altitude where there is
abundant and equally distributed rainfall, in rich, loamy, well-drained soils. In
Sarawak, marang is found up to 1300 m altitude in sandy clay soils.
Marang is an evergreen tree that often achieves a height in excess of 25
m. Twigs are 410 mm thick with long, yellow to red hairs. The stipules
are ovate and 18 cm long with yellow to red hairs. The leaves are elliptic
to obovate, 1650 u 1130 cm (Fig. 2.7A). The inorescences occur in leaf
axils and are solitary. The male heads are ellipsoid to clavate, 411 u 26
cm, while the female heads are pubescent peltate bracts that are mostly shed
and have simple styles that are exserted to 1.5 mm. The fruit are borne at
the end of long branches and average 16 cm in length by 13 cm diameter
(Fig. 2.7B) in the Philippines. The number of edible seeds varies greatly,
averaging close to 100. Pericarps including seeds are ellipsoid, sized about
12 u 8 mm.
In northern Queensland, Australia, marang owers in October and
November with harvesting the following February. In Hawaii, mature marang
trees will ower and fruit year-round, given sufficient rainfall or irrigation.
45
Fig. 2.7. Marang male ower and leaf (A), and fruit (B). (From Ken Love, used with
permission.)
Cultivated marang is much larger than wild fruit and is believed to be a
tetraploid derivative of a diploid ancestor. A variety Maraguinto has been
described in the Philippines. This variety has a large thick aril, superior avor
and small seeds, and can be used as a fresh or processed fruit. This selection
was made from a germplasm collection at the University of Southern
Mindanao. Other varieties in the Philippines are Evergreen and Brown.
Cultural practices
Marang is generally grown from seeds taken from ripe fruit with desirable
qualities. Seeds are recalcitrant and do not remain viable for long, and are
generally sown immediately after cleaning with water. Seedlings are ready for
planting in 1 year and placed 1214 m apart, often at the onset of the rainy
season. Care is taken not to damage the long taproot. The species can be grafted
to like rootstock and some success with grafting to other Artocarpus species has
been reported. Marang has been inarched successfully with breadfruit (A. altilis).
46
Chapter 2
Pruning
Shoots are sometimes thinned and branches cleared to allow harvesting
access, although often it is only the dead wood that is removed. Newer pruning
strategies aim for a tree that is 35 m high.
Fertilization
Marang plants are given 100200 g ammonium sulfate after planting toward
the end of the rainy season. Bearing trees are given 0.51 kg complete
fertilizer per tree twice a year.
Pest management
Seed and blossom rots, leaf spots, pink disease and fruit rot occur on jackfruit
and are likely problems with marang. Bacterial dieback (Erwinia caratovora)
can be a problem with most Artocarpus species. Root rots caused by Fusarium
and Phytophthora are major problems, especially if the root system is ooded
for a few days.
Marang, like jackfruit, is probably attacked by shoot borers, bark borers, bud
weevils, spittle bugs, mealy bugs, scale insects and aphids. Larvae from oriental
jackfruit y (D. umbrosus Fabricius and D. dorsalis) have been found in marang,
but are controlled with modern baits and protective bags covering the fruit as it
develops. In Asia, monkeys, bats and elephants are common pests. Borers can
also be a problem.
Weed management
Once the tree is established, weeds are not a problem because of the dense
shade.

The fruit are sometimes allowed to fall and must be collected daily because
they have a postharvest life of only 23 days. Fruit in the southern Philippines
are harvested 8090 days post anthesis. Letting the fruit fall can cause
damage, loss of shelf life and premature ripening. Grading is not normally
practiced and fruit are rapidly transported to market. Mature undamaged fruit
can be stored at 1012C for 23 weeks. Fruit are dried or preserved in simple
sugar syrup for future use.

Marang is spread throughout the tropics and is cultivated in the Philippines
and Malaysia. A cultivated mature marang tree produces an average of 180
fruit/year. Fruit of up to 2.5 kg have been reported in Sarawak and up to 1.8
kg in the Philippines.
47
Fruit are eaten unripe at 2550% full size as a vegetable or ripe as a fruit. Ripe
fruit are frequently cut open and sliced into pieces for sale. The esh is snowy
white, very sweet when ripe, juicy, very aromatic and of excellent avor. The
esh is separated into segments clinging to the central core and each segment
contains a seed. The edible portion of eshy perianth is 2433% of fresh fruit
weight (Table 2.2). The seeds can be eaten after boiling or roasting. Marang is
considered to be one of the best-avored dessert fruit in the Philippines and is
sometimes used to avor ice cream.
FURTHER READING
Campbell, R.J. and Ledesma, N. (2003) The Exotic Jackfruit: Growing the Worlds Largest
Fruit. Fairchild Tropical Botanic Garden, Miami, Florida.
Carrington, C.M.S. and Sankat, C.K. (2011) Breadfruit (Artocarpus altilis (Parkinson)
Frosberg). In: Yahia, E. (ed.) Postharvest Biology and Technology of Tropical and
Subtropical Fruits: Volume 2. Woodhead Publishing Limited, Cambridge, pp. 251
271.
Kanzaki, S., Yonemori, K., Sugiura, A. and Subhadrabandhu, S. (1997) Phylogenetic
relationships between the jackfruit, the breadfruit and nine other Artocarpus spp.
from RFLP analysis of an amplied region of cpDNA. Scientia Horticulturae 70,
5766.
Love, K. (2008) Minor Artocarpus spp. In: Janick, J. and Paull, R.E. (eds) Encyclopedia of
Fruit and Nuts 2008. CAB International, Wallingford, UK.
Roberts-Nkrumah, L.B. and Badrie, N. (2005) Breadfruit consumption, cooking
methods and cultivar preference among consumers in Trinidad, West Indies. Food
Quality and Preference 16, 267274
Sangchote, S., Wright, J.G. and Johnson, G.I. (2003) Diseases of breadfruit, jackfruit and
related crops. In: Ploetz, R.C. (ed.) Diseases of Tropical Fruit Crops. CAB International,
Wallingford, UK, pp. 135144.
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Primack, R.B. (1985) Composite studies of fruits in wild and cultivated trees of chempedak
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3
CARAMBOLA AND BILIMBI
BOTANY
Family
The family Oxalidaceae is primarily herbaceous, often with tubers and bulbs.
There are some shrubs and two woody genera. Only two of the woody species
in one genera are of interest because of their fruit: Averrhoa carambola L. and
A. bilimbi L. The species is named for Averroes, a 12th century Muslim doctor
and philosopher in Christian Europe who lived in Cordoba, Spain.
Common names for A. carambola are carambola, starfruit and ve ngers
(English), belimbing (Indo-Malay), babingbing (Philippines), caramba, yangtao (Chinese), carambolier (French), ma fueng (Thai), fuang (Laos) and khe
(Vietnamese).
A. bilimbi is closely related to carambola though with marked differences in
fruit appearance, fruiting and avor. Bilimbi is the common Indian name and
the name is widely used throughout south-east Asia. The other English name
is the cucumber tree.

The center of origin of carambola is not clear; Indochina, Malaysia and
Indonesia are considered the most likely places of origin. Other suggestions
include the Moluccan archipelago, India and Sri Lanka. It probably does
not exist in the wild state. Knight (1982) suggested a secondary center of
diversication in northern South America around Guyana, where carambola
has been established for more than 150 years. Carambola was taken to the
Philippines and India in prehistoric times. A Sanskrit name Karmana, in
India, attests to its long presence there. It was reported in Rio de Janeiro
in 1856, and in Florida and Trinidad and Tobago in 1887. The crop was
53
54
Chapter 3
probably introduced to Hawaii sometime after 1789 and to Australia at

the end of the 19th century. The rst introduction into the Canary Islands
occurred in 1881 (Galn Saco, 1993).
Carambola is now cultivated in many places in the tropics and subtropics.
The main producing countries include Taiwan, Malaysia, Brazil, Guyana, India,
the Philippines, Australia, Israel and the USA (Florida and Hawaii). Statistics
are not currently available about world production and export, but worldwide
commercial carambola planting is probably less than 10,000 ha.
Bilimbi is cultivated throughout Indonesia, Malaysia, the Philippines, India
and Sri Lanka on a small scale and is frequently found as a backyard tree. It is
also common in other south-east and south Asian countries, and is now found
worldwide. Because it is grown as a backyard crop, yield data are not available.
ECOLOGY
Soil
Carambola thrives in almost any soil type, from sand to heavy clay loam and
rocky calcareous soil that are reasonably well drained. Low-lying areas and
those where water ponds for more than 12 h are unsuitable. A pH range of
5.56.5 is preferred, but carambola can be grown in alkaline soil to pH 7.7. It
may not produce well under saline conditions. Bilimbi also grows in a range of
soil types.
Climate
Both species are adapted to climates from sea level to about 500 m in the
tropics and warm subtropics.
Rainfall
Well-distributed rainfall of between 1500 and 3000 mm is reported to be
suitable for carambola. The tree grows well in dry areas and is tolerant of
seasonal drought once established. Good-quality fruit have been obtained in the
Canary Islands with rainfall and irrigation of 800 mm. Water stress limits root
and leaf growth, and initiates ower development (Fig. 3.1). Drought during
ower and fruit development can signicantly reduce the yield by reducing
ower development and inducing abscission (Fig. 3.2). Well-watered plants
show delayed owering of up to 3 months and extensive vegetative growth (Fig.
3.1).
Temperature
The ideal temperature for carambola is considered to be between 21C and
32C. Below 15C, growth ceases and ower opening is restricted. Young
Leaves
(Control)
55
Flowers
(Severe)
Leaves
(Moderate)
Leaves
(Severe)
Flowers
(Moderate)
Flowers
(Control)
Flower Number per Plant
Leaf Number per Plant
Carambola and Bilimbi
Time from Start of Treatment (Weeks)
Yield (tonne/ha)
Fig. 3.1. Inuence of soil moisture availability on leaf and ower number per plant
of carambola B-17. Control = 88100%; moderate = 6570%; severe = 4248%.
(After Ismail et al., 1996.)
Irrigation (l/day)
Fig. 3.2. Fruit yield of B-17 carambola trees that were either not irrigated or
irrigated with 4, 12, 30, l/tree/day, only during the drought months, when the
rainfall was less than evaporation or rain did not fall for 2 weeks. (After Bookeri,
1996.)
shoots are damaged by temperatures approaching 0C and die below 0C,

with complete defoliation following exposure to 1C. Adult carambola
trees are more tolerant but are killed if exposed to 4C for more than 24 h.
Bilimbi is less cold tolerant than carambola. Carambola fruit produced during
the cool season can have 6 lower total soluble solids than in the summer.
56
Chapter 3
Temperatures greater than 3035C during carambola owering reduce

pollen germination and pollen tube growth.
Light and photoperiod
Maximum carambola fruit production is obtained in direct sunlight, with
more than 2000 h of sun per year being recommended. Flowering occurs
with photoperiods between 8 and 16 h, though 1416 h of daylight reduces
the owering rate and 812 h increases the number of carambola owers per
inorescence.
Wind
Hot and dry wind can harm or kill carambola trees, while cold winds of 10C
and greater than 80 km/h can defoliate trees; the tree will require a few
months to recover. Fruit injury caused by rubbing on branches and other fruit
seriously reduces fruit quality. Windbreaks, either temporary or permanent,
are essential to produce high-quality unblemished fruit. The bilimbi tree is less
wind and cold tolerant than carambola.
Tree
Carambola is a small, slow-growing, evergreen tropical tree. It rarely reaches
more than 89 m in height and has a useful economic life of about 25 years.
The sparsely branched bilimbi tree is up to 15 m tall. Young carambola trees
generally have a pyramid shape, changing to a symmetrical rounded top. The
trunk is smooth grayish to dark, with a tendency to form low branches. The
roots branch very close to the base of the trunk with very thick lateral and
deeply penetrating anchoring roots.
Carambola leaves are arranged alternately (Fig. 3.3). They are petiolate and
pinnate, and may reach 20 cm. The young leaves are bronze red, changing
to pale to dark green when mature. The pinnate leaets vary in number from
two to 11, but usually number three to six. The leaet shape is ovoid, ovate
lanceolate or oblong elliptic (Fig. 3.3). The laminae are 2.57.5 cm long and
14 cm wide, pubescent on the upper surface and almost smooth on the
underside. The compound leaves of bilimbi have 2040 leaets, each of which
is 510 cm long.
Flowers
Grafted carambola plants produce owers in 9 months, while seedlings may
not ower until 46 years old. The owers occur on a loose panicle produced
57
Fig. 3.3. Carambola leaves, ower panicle, ower and fruit. A transverse section
through a fruit shows the star shape and seeds.
on basally branched slender twigs (18 cm long) toward the periphery of the
tree. The panicles occur in leaf axils and occasionally on leaess branches and
terminally of new shoot growth or from latent buds on older shoots. Bilimbi
ower panicles have 1864 owers that form on the trunk and older branches
(Fig. 3.4).
Perfect carambola owers have a calyx with ve pink sepals (Fig. 3.3)
surrounding the purple corolla, and are 512 cm long attached to a short (1
mm) round, dark-red pedicel. Heterostyly (distyly) is a characteristic. Some
cultivars bear long-style (2 mm), short-stamen (34 mm) owers, while others
have short styles (0.51 mm) and long stamens (56 mm) (Table 3.1). There
are ve sepals with 10 stamens, of which the ve adjacent to the petals are
reduced to staminoids with no or aborted anthers. Four or ve styles develop
from the ovary and fuse with the stigmatic surface, having numerous papillae.
The ovary is 1525 mm long with four or ve loculi each containing two to
four superimposed ovules (Galn Saco, 1993). Bilimbi owers are also
58
Chapter 3
Fig. 3.4. Bilimbi owers and fruit on the tree trunk.
Table 3.1. Flower style length varies between carambola

cultivars and are described as short or long. (After
Watson et al., 1988) Cultivars starting with the letter
B are from the Malaysian Agricultural Research and
Development Institute.
Short style
Long style
B-2
B-8
B-10
Fwang Tung
Lu Tho
Wheeler
Sri Kembangsaan
Kara
B-1
B-6
B-11
B-16
Star King
Maha
Arkin
Kary
59
heterotristylous, with free petals 1030 mm long and yellowish-green to

reddish-purple in color; the short stamens are fertile.
Carambola owering can be induced by a period of water stress and/or root
restriction and is dependent on cultivar. Continuous owering occurs in the
tropics, with different sections of the tree apparently owering at random. The
asynchrony of owering can be reduced by management practices such as
shoot pruning.

From a visible ower bud (1 mm) to open ower takes about 2 weeks in
carambola. The terminal owers on a peduncle open rst. Flowers open
between 8 am and 10 am and close between 2 pm and 6 pm on the same day
(Salakpetch et al., 1990). Petals fall during the morning following anthesis.
After opening, though no longer receptive, owers may remain on the plant
for up to 16 days. Insects are apparently needed for good pollination, though
some wind pollination occurs. The bright-colored owers have nectar and are
attractive to bees; one hive is recommended for each 0.4 ha of trees.
Heterostyly is a cause of pollination failure and self-incompatibility. Failure
is almost complete in short-by-short crosses (approximately 0.5% on the same
tree and between owers on different trees). Fruit set is low with long-by-long
style crosses (approximately 0.5%) and high with long-by-short style crosses
and vice versa (320%). Microscopy indicates that germination occurs in noncompatible crosses, but pollen tubes are usually inhibited in the style.
Bilimbi owering occurs continuously, with marked peaks following a period
of drought. Flowers take about a week from bud burst to open ower, with
owers opening in the morning. It takes about 15 days for the whole panicle to
ower. Bilimbi fruit set occurs 1520 days after inorescence emergence and
is indicated by petal fall with the stigma persisting on the end of the fruit. The
owers are self-fertile.
Fruit
The carambola fruit is a large, indehiscent, eshy berry of 512 cm long and
36 cm across. In cross-section, the fruit has a ve- (occasionally six-) pointed
acute star appearance (Fig. 3.3). It is yellow to orange when ripe, with 1012
ovoid seeds approximately 1 cm long. Within 710 days after pollination, fruit
set is indicated by swelling of the ovary or shedding. Fruit set is regarded as
having occurred when the petals have fallen, ovary expansion begun and the
color changes from white to green. Initial fruit growth is via division until
1215 mm long, then via expansion and elongation. In Florida and Hawaii,
fruit maturity occurs 6065 days after fruit set (Fig. 3.5). Oxalic and malic
Acids Sugars (mg/g FW)
Chapter 3
Fruit Masses (g)
60
Time after Fruit Set (Days)
Fig. 3.5. Increase in Arkin fruit mass and total sugars and decrease in oxalic acid
content after fruit set. (After Campbell and Koch, 1989.)
acid levels decline during development and especially during fruit ripening,
while total sugars increase. Fruit sugar level only increases when the fruit is
attached to the tree, becoming progressively sweet up to the full yellow/orange
stage. The tart or acid varieties do not show a similar marked decline in oxalic
acids.
The waxy pale green bilimbi fruit is slightly lobed, and 510 cm long and 5
cm across (Fig. 3.4). Unlike carambola there is no aril on the seeds, which are
8 mm long. Fruit color changes from green to light yellow when ripe. The fruit
show a typical sigmoid growth pattern and reach maturity 5060 days from
anthesis.
Carambola is reported to have a 2n = 22 or 24. Limited data are available
on gene frequencies, linkages or inheritance of desirable horticultural traits.
Polymorphism has been found in isozyme alleles of seedling populations from
controlled pollination. Seeds from specic crosses are easily obtained with the
anthers from short-styled owers removed with forceps, without damaging
the ower. Long-styled owers are more difficult to use as a female parent. Selfsterility further limits breeding.
61
Selection and evaluation

Sweet and acid types of carambola fruit were frequently used to separate
seedlings before named varieties appeared. The most intensive selection
programs have been carried out in Malaysia, Taiwan, Thailand, Florida and
Hawaii. The Malaysian Agricultural Research and Development Institute
maintains a numbered series with different desirable characteristics.
An extensive list of desirable characteristics for carambola varieties includes
early bearing habit, abundant regular production and ease of training.
Plasticity of growth, owering and fruit set to withstand water stress, salinity
and adverse soil and weather conditions would also be helpful. A major focus
of all selection programs has been for desirable fruit characteristics, including
weight, wing-to-body ratio, color, sugar-to-acid ratio, low seed number,
esh texture, fruit esh browning and resistance to mechanical injury. Wide
variation occurs in fruit quality, with color varying from pale yellow to deep
orange. The star shape and rib depth can vary signicantly, with shallow ribs
being preferred for packing and avoidance of mechanical injury. The cultivars
Fwang Tung and B-6 have bigger ribs than B-10 and Arkin. Sugar content
can also vary from 5% to 13% total soluble solids. A sugar level greater than
10% total soluble solids is preferred, as consumers will otherwise regard the
fruit as bland.
The use of fruit is sometimes a factor in selection. Dessert fruit and minimally
processed slices should preferably have a high sugar-to-acid ratio. The acid
level, mainly oxalic acid, shows the greatest variation between sweet and tart
cultivars, ranging from 1 to 6 mg/g fresh weight. Oxalic levels are higher in
the ribs than the body of the fruit and decline during ripening. The malic acid
content is higher in sweet varieties (1.2 mg/g fresh weight) than in tart varieties
(0.4 mg/g fresh weight). The juice pH can vary from 2.4 to 5.0.
Major cultivars
Seedling populations are very heterozygous and cannot be relied on for
commercial production, with those from short style types regarded as yielding
greater diversity than long-style types. Many named varieties are available
with acceptable yield, fruit and disease tolerance (Table 3.2), although the
climactic suitability of individual cultivars varies widely. Taste similarly varies
considerably with stage of maturity at harvest (Table 3.3) and this needs to
be standardized before making comparisons. Some cultivars show different
amounts of change in total soluble solids and titratable acidity during
ripening (Table 3.3); for example, Arkin has higher titratable acidity and
lower total soluble solids when fully ripe than Kyra.
Color
Name
Arkin
Golden Star
Sri
Kembangsaan
Kary
B-2
B-10
B-17
Fwang Tung
Star King,
Sweetie,
Similar B-2
Collected in
Malaysia
selection
Kary
Chum Choi
Maha 66;
slowgrowing
tree,
medium to
large fruit
Ching Sing
Keow;
short styles,
vigorous
tree, large
fruit,
moderate
resistance to
fruit y
Cristal Honey
or Honey
Carambola
*TSS, total soluble solids..
Ribs
Country
Weight (g)
Mature
Full ripe
Size
Angle
Flesh
texture
Flavor
TSS* (%) Use
Florida,
USA
90200
Golden
yellow
Yellow
orange
Thick
Large
Crisp
4.9
6.011.5 Fresh
processing
Florida,
USA
Hawaii,
USA
100200
Thick
Large
150200
Golden
yellow
Lemon
yellow
Crisp, very
juicy
Slightly
6.5
crisp
Indonesia
Singapore
Taiwan
Up to 315
Light green Orange
Thin
Sharp
Malaysia
100200
Greenishyellow
Yellow
Deep
furrow
Sharp
Malaysia
100200
315
Greenishyellow
Yellow to
golden
reddish
to
orange
Malaysia
Thailand
100300
7.013.0
Slightly
crisp
Crisp
78
More
Slightly
compact
rounded
B-2
Crisp
912
Fresh
processing
Golden
yellow
Large
5.3
1518
Crisp
5.5
9.212.6
Chapter 3
Kaput
Lang Bak
Cheng Tsey
Other names;
notes
62
Table 3.2. Fruit characteristics of some carambola varieties grown in different countries. (From OHare, 1993; Galn Saco, 1993;
Campbell, 1971; Watson, et al., 1988; Sedgley, 1983.) Flavor is ranked 1 = strongly dislike to 9 = strongly like.
63
Table 3.3. Changes in total soluble solids and titratable acidity of three Hawaii
cultivars and the Florida cultivar Arkin of carambola during fruit ripening on the
tree.
Total soluble solids (%)
Stage of
ripeness
Green
Green
yellow
Yellow
orange
Orange
Titratable acidity (mEq/100 ml)
Kajang
Arkin
Sri
Kembangsaan
5.5
5.9
5.7
5.5
5.7
6.2
6.3
5.9
5.8
5.3
6.9
6.5
5.8
6.3
5.6
6.0
6.4
6.4
7.1
6.3
5.2
5.4
5.2
5.5
7.6
7.3
9.6
8.1
4.1
5.0
4.3
3.1
Kyra
Kajang
Arkin
Sri
Kembangsaan
Kyra
Sweet and acid forms of bilimbi fruit exist, although no named cultivars have
apparently been developed. The sweet forms are too acid for the fresh market.
Variable progeny is derived from seeds because of out-crossing.
CULTURAL PRACTICES
Mature carambola seeds germinate readily within 7 days if sown immediately
after removal from the fruit. If kept moist, seeds can be stored for about 2
weeks in the refrigerator. Well-drained medium should be used in trays, with
the seedlings later transplanted to pots or bags when the rst true leaf is
mature. Bilimbi is usually seed propagated, but can be vegetatively propagated
by air layering and grafting. There are no reports of rootstock suitability for
bilimbi.
For carambola, side-veneer grafting, chip budding, wedge grafting, modied
Forkert bud grafting and splicing are used. Approach grafting is rarely used
commercially, being slow and requiring the vegetative potted trees to be
adjacent to each other. A range of seedling types are used as rootstock with
incompatibility not being reported, although seedlings vary in vigor. Defoliated
hardened (brown) scion wood is used, with leaves removed from the scion 3 or
4 days before collection. After 4 weeks, if the scion is still alive, the upper part
of the tie is removed and the stock decapitated just above the graft. Watering is
reduced to above one-third normal until the scion makes signicant growth.
Air layering is difficult with slow root growth, and cuttings are not reliably
rooted under mist.
64
Chapter 3
Top-working can be done with side-veneer or wedge grafting on new growth.

Conventional propagation on regrowth from stumped trees can be used, which
allows existing trees to be top-worked when a better selection becomes available.
Trees can be expected to start yielding within 918 months.
Field preparation
Deep ripping along the tree row is recommended, and should follow the
contour if on hillsides. Drainage is essential and areas in which ponding
occurs for more than 12 h should be avoided. Lime may be applied as
required in acid soils, with sulfur in alkaline soils, to achieve pH 5.56.5.
Animal manure or organic matter should be dug into at each plant hole,
46 months before planting. Fertilizer (NPK) should be mixed with top soil at
transplanting.

Holes should be dug just before planting, which can be done at any time of the
year. Spring and early summer are recommended in the subtropics. Eighteenmonth-old seedlings with the roots carefully pruned, if necessary, are used.
After planting, seedlings should be watered regularly during the rst 24
weeks. Seedlings may need to be staked during the rst year.
Spacing of carambola trees varies widely, depends on the cultivar, soil type
and pruning practices. Mature trees can reach 7.5 m in diameter. Minimum
distances of 57 m between trees and 69 m between rows have been
recommended. Spacings, however, can vary from 4.2 u 3.6 m to 10 u 10 m,
with the smaller spacing giving 900 trees/ha. Higher densities are used for
younger trees, with surplus trees being later removed. Trellises allowing
denser tree spacing are used in Taiwan (Fig. 3.6A and B), and have been used
in Florida and tested in Hawaii. Higher-density planting also shows promise.
Intercropping with short-season crops is frequently practiced.
Irrigation is recommended, particularly following planting and during dry
periods. Soil moisture stress appears to be a key factor in inducing owering
in areas with low rainfall. Drought during and after owering leads to poor
owering, early fruit abscission, reduced yield and small fruit. Applications of
up to 2000 l of water per mature tree per week has been recommended during
high-demand periods, applied in two to three applications per week. Irrigation
should be increased as owering occurs and as the fruit develop, then reduced
65
Fig. 3.6. Production practices for carambola. (A) Trellises are used in Taiwan, with
(B) the bagged fruit hanging under the trellis. Wind can cause severe damage to
fruit that rub against the stem and other fruit, necessitating protection with either
a tree or screen wind-break (C). Birds can be a problem in some areas, requiring
protective structures (D).
as the fruit begin to ripen. Water stress is avoided when trees are being cycled
for owering (Fig. 3.7), where pruning is the main management strategy.
Mulching around the base of the tree is also very benecial.
Flood irrigation, sprinkler and microspot sprinklers are used, with the last
of these being now more widely accepted. Real-time evapotranspiration-based
irrigation controllers result in signicant water conservation. Flooding is to be
avoided, as it reduces the productivity of mature trees by inducing severe leaf
abscission.
Pruning and training

Three to ve well-developed branches 5080 cm from the soil, with wide
angles, are allowed to develop on carambola. The annual pruning removes
narrow-angled, low-hanging and overlapping branches. All side shoots up to
1 m are also removed. Pruning aims to retain recently maturing shoots and
generate new shoots that bear the highest fruit yield; new shoots have smaller
fruit. Flowers on main branches are also less productive. Bilimbi trees can be
66
Chapter 3
Fig. 3.7. Management practices utilized to induce three harvests per year in
southern Taiwan. Double fertilization is carried out 12 months before the rst
pruning in every year during the cool dry season and then every 2 months. The
time of pruning are varied to achieve year-round production. Other essential
management practices include maintaining the water supply and fruit thinning (T)
to improve fruit size and reduce the length of the harvest period.
pruned and trained to various shapes. Heavy pruning of bilimbi suppresses

owering since owering occurs on thicker branches.
In more intensive production systems, the tree height of carambola
is maintained at less than 2.5 m to allow fruit bagging, if practiced, and
harvesting from the ground. Trellis pruning of trees at 1.8 m is used to assist
in maintaining tree height in an inverse conical shape. Alternatively, trees are
tied down to achieve a weeping willow structure for ease of harvesting from the
ground (Fig. 3.6A and B).
In Taiwan, heavy pruning at the end of the cool season is used to induce
ower-bud initiation 715 days later. This heavy pruning reshapes the tree,
reducing its height as well as removing slender and senescent branches. Lighter
pruning at about 3-month intervals can then be used to develop further
owering and fruiting (Fig. 3.7), giving three harvests a year. Light pruning
removes dry, slender and senescent branches to open up the canopy, produce
new shoots and ower buds and prevent fruit drop. Any water shoots that
develop between regular prunings are removed.
Flower and fruit thinning are frequently used to increase fruit size, reduce
the occurrence of misshaped fruit and prevent interference of ower cycling.
Thinning to three fruit per cluster 34 weeks after owering has been
recommended. If out-of-season owering is required, all or most owers and
fruit from the previous cycle are removed. Where fruit thinning is practiced, the
individual fruit are often bagged when they reach about 6 cm in diameter. This
bagging avoids mechanical damage and insect stings. The bags can be made of
white paper, newspaper or polyethylene. Commercial bags of 20 u 10 cm with
a wire tie at the open end are available, and can be tied on the fruit with one
hand.
67
Fertilization
Animal manure or organic matter should be dug into each planting-hole
site for carambola, 46 months before planting. Fertilizer (NPK) is mixed
with top soil at transplanting. Organic matter or manure, 10 kg/tree/year
for young trees and 1025 kg/tree/year for older trees, is applied in a single
application in some production systems. The fertilizer is applied in a 1 m ring
around the tree base or along the drip-line. There are no reports of fertilizer
recommendations in the literature for bilimbi.
In more intensive production systems, young bearing carambola trees
receive 0.40.8 kg/tree/year of NPK (ratio 11:12:1715:15:15), depending on
soil analysis results. Older trees (8 years) may require 625 kg/tree/year. The
fertilizer can be applied at intervals of about 3 months during fruit production,
though limited during owering. An application is made after the last fruit
harvest of the year, followed by irrigation to stimulate new growth. The trees
are pruned heavily when growth slows 12 months later.
Each 1 t of carambola fruit can remove 1.02 kg nitrogen, 0.12 kg phosphorous, 1.58 kg potassium, 0.1 kg magnesium, 0.1 kg sulfur and 0.05 kg
calcium from the soil. Leaf analysis from non-bearing trees has found leaves
contain 1.4% nitrogen, 0.12% phosphorous, 0.12% potassium, 0.98% calcium,
0.64% magnesium and 0.24% sulfur. These gures are given as guides and
may not directly relate to yield.
Pest management
Diseases
Leaf spot (Cercospora averrhoa Petch.) can cause serious loss of carambola
leaves and can also affect the fruit. The small chlorotic spots (up to 5 mm)
are at rst brown, then turn grayish-brown and lead to premature leaf
loss. Cupric fungicides and Difolatan provide control. Leaf spots caused by
Phomopsis species, Phyllosticta species and Corynespora cassiicola have been
reported in different areas.
Fruit rot or blemishes caused by Botrytis, Ceratocystis, Colletotrichum,
Aspergillus, Dothiorella, Alternaria, Phoma and Phomopsis species also occur. Fruit
blemish can lead to rejection on the packing line. Symptoms of anthracnose
(Colletotrichum spp.) include thin, light-brown patches that enlarge and
coalesce into salmon-colored patches, which then blacken. There are no
recommendations for postharvest disease control. Precooling and refrigeration
do reduce the disease development rate.
Fusarium decemcellulare can cause serious inorescence growth disorder of
bilimbi, with Penicillium reported as pathogenic to the fruit. Leaf spots have
been seen, but the causal organism is unknown.
68
Chapter 3
Insect pests
Fruit ies such as Dacus dorsalis (the Oriental fruit y) in Asia are major
commercial pests of carambola and a reason for fruit bagging. This and other
fruit ies are also a quarantine restriction for export from tropical countries to
some markets. Several moths, including Othreis species, pierce nearly mature
to mature fruit and suck the juice, with the area subsequently rotting and
fruit falling prematurely. This is particularly a problem in south-east Asia and
Australia. Control measures are limited in their effectiveness.
Various other beetles, fruit borers, thrips, mealy bugs and scales can attack
the branches, fruit and owers of carambola. Citrus red mite (Panonychus citri)
is a severe problem, especially in the dry season in southern Taiwan. Ants
are sometimes found around the peduncle depression at the end of the fruit,
collecting honey dew produced by tree hoppers (Membracidae spp.) that feed on
the peduncle.
No serious insect pests have been reported for bilimbi.
Non-pathogenic problems
Browning and rotting of the area between the ribs has been reported from
Florida. This should not be confused with the surface browning frequently
found in cooler months or following moisture loss. Chilling injury leading to
brown patches has also been reported for carambola. This, however, may be
related to dehydration and not low temperatures. Bird and bat damage can be
a major problem (Fig. 3.6C and D). Cultivars with sharp rib edges are more
susceptible to bruising and discoloration than those with rounded edges. This
bruising and rubbing is frequently caused by wind damage.
Weed management
Trees are not good competitors when small, and weed control is essential.
Organic or plastic mulches are used to maintain a 1 m diameter weed-free
area around the base of the tree.
Orchard Protection
Carambola fruit are very susceptible to wind damage, characterized by
rubbing and marking of fruit. The trees can also become defoliated, with
dieback of twigs and stunted growth. Wind barriers and/or a protected site
are needed (Fig. 3.6D).
69

Harvesting
Grafted trees can start bearing fruit within 9 months of planting, with
satisfactory yields in another 23 years. Fruit thinning and bagging are used
to achieve good yields of adequately sized fruit, with a good shape and free
from wind or insect damage. Although trees can produce total yields of 20
kg, increasing up to 500 kg/tree when the tree matures in 67 years, 90% of
the fruit can be rejected. Yields of 100250 kg/tree/year are more common,
in two to three crops per year. Fruit are harvested by hand and should be
carefully accumulated in the eld and transported to the packing shed.
Accumulating fruit in deep containers or buckets should be avoided to prevent
mechanical injury.
Harvesting is based on physiological and horticultural maturity, as indicated
by changes in skin color from green to yellowish-green, then to full yellow or
yellowish-orange, and the increase in fruit sugar levels that only takes place
when the fruit is attached to the tree. Optimum sugars are achieved at the full
yellow color; however, ripe fruit are more fragile and easily damaged, and so fruit
are often harvested at the color break to 50% yellow stage, depending on market
requirements. Fruit that are half to three-quarters yellow are rmer than fullcolor fruit and hence regarded as commercially mature. Sugar increases and
acidity decreases as skin color develops on the tree (Fig. 3.5). There is no further
increase in sugars after harvest, although color development continues.
Fruit should be graded and sorted to remove small, misshaped, insect and
wind damaged, and diseased fruit, then sorted by size and degree of color
development. There are no US or international grades. Fruit is sold in 3.5 kg
ats, 10 kg single layers, 9 kg suitcases and clam shells (16 fruit in one layer
or 32 fruit in two layers). Careful packing is required to reduce damage,
coupled with the use of either a plastic or foam sleeve or waxed paper wrap.
Careful handling can ensure fruit reach the consumer with a minimum
of injury, rubbing or browning. Cartons holding 3.5 kg of fruit are used for
export, while larger cartons holding 14 kg of fruit are used within south-east
Asia. The carton should allow for display and still have sufficient padding of
either paper or polyethylene to provide protection against crushing, bruising
and desiccation.
Cooling to 510C by forced air or room cooling as soon as possible is
desirable, with storage at 510C and 9095% relative humidity for 2135
days. These storage temperatures lead to minimum changes in sugars and
a slight decline in organic acids. The length of storage varies with stage of
70
Chapter 3
Table 3.4. Composition of 100 g edible portion of carambola

(Wenkam, 1990) and bilimbi (Leung et al., 1972).
Constituent
Edible pulp (%)
Proximate
Water (g)
Energy (kJ)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Magnesium (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
Carambola
90.23
155
0.85
0.9
7.52
1.47
0.5
1
0.06
9
11
145
2
35
0.04
0.044
0.71
21
Bilimbi
86
92.5
0.61
0.3
6.3
0.6
0.3
5
0.6
13
130
4
35
0.02
0.04
0.02
105
ripeness when placed into storage. Lower humidity result in more severe ribedge browning, and fruit have a storage life of 34 days if held at 20C and 60%
relative humidity. Moisture loss is a major problem during storage and can lead
to skin browning, which has been mistakenly described as chilling injury. Fruit
held at 7C in 2.24.2% O2 with 88.2% CO2 retain better color and rmness
than fruit held in air. Sealed polyethylene lm bags delay degreening and with
no effect on avor after 1 week at 20C of either green or full-colored fruit.
Waxing also delays water loss and degreening. Ethylene can be used to enhance
color, although it can cause blemishes.
Irradiation (250 Gy), cold treatment (12 days, 1C) and heat treatments
(fruit center to approximately 49C) have the potential for insect disinfestation
to address quarantine concerns in importing countries. These treatments
occasionally lead to some skin discoloration, dehydration and pitting. Hotwater treatments can reduce shelf life. Fruit at the greenyellow stage are more
easily damaged by irradiation and hot-water treatments than riper fruit.
71
UTILIZATION
Carambola are normally consumed fresh as a dessert fruit. Dessert fruit and
minimally processed slices should preferably have a high sugar-to-acid ratio.
Carambola can also be consumed as jams, preserves, pickles, candy, juice and
liquor. Tart fruit are preferred in Taiwan for processing into juice. Green fruit
are sometimes consumed as a vegetable and sliced ripe fruit added to salads.
The fruit is a good source of potassium and vitamin A and a moderate source
of vitamin C (Table 3.4). The fruit has a sweet, oral avor due to esters, with
methyl benzoate and ethyl benzoate being major active aroma components.
Beside esters, aldehydes, alcohols, ketones and norisoprenoids are also
present, with the stage of fruit ripeness signicantly changing the aroma
prole.
The bilimbi is too acid to be used as a fresh fruit, and its major use is in
pickles, curries, chutney and preserves. The fruit is widely used in traditional
medicine. The sour taste is caused by a high oxalic acid content, which ranges
from 10.514.7 mg/g in green fruit to 8.4510.8 mg/g in ripe fruit.
FURTHER READING
Galn Saco, V. (1993) Carambola cultivation. FAO Plant Production and Protection Paper
No. 108.
Nunez-Elisea, R. and Crane, J.H. (2000) Selective pruning and crop removal increase
early-season fruit production of carambola (Averrhoa carambola L.). Scientia
Warren, O. and Sargent, S.A. (2011) Carambola (Averrhoa carambola L.). In: Yahia, E.
(ed.). Postharvest Biology and Technology of Tropical and Subtropical Fruits: Volume 2.
Woodhead Publishing Limited, Cambridge, pp. 397413.
Wilson, C.W. (1990) Carambola and bilimbi. In: Nagy, S., Shaw, P.E. and Wardowski,
W.F. (eds.). Fruits of Tropical and Subtropical Origin Composition, Properties and Uses.
FSS Inc., Lake Alfred, Florida, pp. 276301.
REFERENCES
Bastos, C.N. and Santos, A.O. (2001) Superbrotamento de inorescencias do lumao-decaiena causado por Fusarium decemcellulare. Fitopatologie Brasileira 26, 222225.
Bookeri, M. (1996) Effect of irrigation on carambola (Averrhoa carambola) production
ill drought prone areas. Proceedings of the International Conference of Tropical Fruit,
Kuala Lumpur, Malaysia, pp. 317320.
Campbell, C.A. and Koch, K.E. (1989) Sugar/acid composition and development of
sweet and tart carambola fruit. Journal of the American Society for Horticultural
Science 114, 455457.
Campbell, C.W. (1989) Carambola production in the United States. Proceedings of the
InterAmerican Society for Tropical Horticulture 33, 4754.
72
Chapter 3
Chung, R.C.K. and Lim, A.L. (1998) The embryology of Averrhoa (Oxalidaceae).
Sandakania 12, 3755.
Crane, H., Knight, R.J. Jr, Rodriguez, O. and Crane, L.C. (1998) Cultivar tree growth
and fruit quality evaluation of young carambola (Averrhoa carambola L.) trees.
Proceedings of the Florida State Horticulture Society 111, 299302.
Crane, J.H., Willis, L., Lara, S.P. and Ellenby, M. (1991) Short-term effects of manual
topping of ve-year old Arkin carambola trees. Proceedings of the Florida State
Horticultural Society 104, 5760.
Dahanayaka, S. and Wijesundera R.L.C. (1994) Penicillium purpurogenum on fruits of
Averrhoa bilimbi in Sri Lanka. Journal of National Science Council of Sri Lanka 22,
2324.
Galn Saco, V. (1993) Carambola cultivation. FAO Plant Production and Protection Paper
No. 108.
George, A.P. and Nissen, R.I. (1994) Carambola. In: Schaffer, B. and Andersen, P.C.
(eds). Handbook of Environmental Physiology of Fruit Crops, Volume 2, Subtropical and
Tropical Crops. CRC Press, Boca Raton, Florida, pp. 206211.
George, H.L., Crane, J.H., Schaffer, B., Li, Y. and Davies, F.S. (2000) Effect of polyethylene
and organic mulch on growth and yields of Arkin carambola (Averrhoa carambola
L.) in south Florida. Proceedings of the Florida State Horticultural Society 113, 511.
George, H.L., Crane, J.H., Schaffer, B. and Li, Y. (2001) Effect of polyethylene and organic
mulch on owering, pollen viability, percent fruit set, and fruit production of Arkin
carambola (Averrhoa carambola L.) in South Florida. Proceedings of the InterAmerican
Society for Tropical Horticulture 44, 7277.
Hernndez Delgado, P.M. and Galn Saco, V. (2003) Evaluation of carambola cultivars
in the Canary Islands. Fruits 58, 1926.
Ismail, M.R. and Awang, M. (1992) Growth and physiological changes of Averrhoa
carambola as inuenced by water availability. Pertanika 15, 17.
Ismail, M.R. and Noor, K.M. (1996) Growth and physiological processes of young
starfruit (Averrhoa carambola L.) plants under soil ooding. Scientia Horticulturae 65,
229238.
Ismail, M.R., Yusaf, M.K. and Mastnri, A. (1996) Growth and owering of water
stressed star fruit plants and response to ameliorated water stress. Proceedings of the
International Conference of Tropical Fruit, Kuala Lumpur, Malaysia 2, 97106.
Joseph, J. and Mendonca, G. (1989) Oxalic acid content of carambola (Averrhoa
carambola L.) and bilimbi (Averrhoa bilimbi L.). Proceedings of the InterAmerican
Joyner, M.E. and Schaffer, B. (1989) Flooding tolerance of Golden Star carambola trees.
Proceedings of the Florida State Horticultural Society 102, 236239.
Kisekka, I., Migliaccio, K.W., Dukes, M.D., Schaffer, B. and Crane, J.H. (2010)
Evapotranspiration-based irrigation scheduling and physiological response in a
carambola (Averrhoa carambola L.) orchard. Applied Engineering in Agriculture 26,
373380.
Knight, R.J. Jr (1965) Heterostyly and pollination in carambola. Proceedings of the Florida
State Horticultural Society 78, 375378.
Knight, R.J. (1982) Response of carambola seedlings population to Dade countys oolitic
limestone soil. Proceedings of the Florida State Horticultural Society 95, 121122.
Knight, R.J. (1989) Carambola cultivars and improvement programmes. Proceedings of
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Lennose, A. and Ragoonath, J. (1990) Carambola and bilimbi. Fruits 45, 497501.
Leung, W.T.W., Butrum, R.R. and Chung, F.H. (1972) Food Composition Tables for Use in
East Asia Part 1. US Dept of Health, Education and Welfare, Washington, DC and
FAO, Rome.
Marler, T E. and Zozor, Y. (1992) Carambola growth and leaf gas exchange responses to
seismic or wind stress. HortScience 27, 913915.
Mathew, L., George, S.T., Babylatha, A.K. and Geetha, C.K. (1991) Flowering and fruit
development in bilimbi (Averrhoa bilimbi L.). South Indian Horticulture 41, 4142.
Matthews, R.F., Sims, C.A. and West, P.F. (1988) Utilization of carambola in a tropical
beverage. Proceedings of the InterAmerican Society for Tropical Horticulture 32, 25
33.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resources System, Inc. Winterville,
North Carolina. Available from: http://www.hort.purdue.edu/newcrop/morton/
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Nunez-Elisea, R. and Crane, J.H. (2000) Selective pruning and crop removal increase
early-season fruit production of carambola (Averrhoa carambola L.). Scientia
Nunez-Elisea, R., Schaffer, B., Zekri, M., OHair, S.K. and Crane, J.H. (2000) Monitoring
soil water content in tropical fruit orchards in south Florida with multisensor
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OHare, T.J. (1993) Postharvest physiology and storage of carambola (Starfruit): a
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Razi, M.I., Awang, M. and Razlan, S. (1992) Effect of water stress on growth and
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Salakpetch, S. (1987) Flower ontogeny and pollen germination in carambola (Averrhoa
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Journal of Agricultural Research of China 42, 387395.
4
DURIAN
BOTANY
The tropical family Bombacaceae is known for its showy owers and pods,
with seeds covered with cotton-like bers. The family is found in both South
America and Asia. The fruit of the genus Durio differ in having fruit with
large seeds enclosed in a eshy aril. The genus Durio has 29 species, six of
which produce edible fruit, with D. zibethinus Murray (syn. D. acuminatissima
Merr.) durian being the most widely cultivated. The genus-specic zibethinus
is derived from the Italian zibetto for civet, a cat-like animal with a musky
smell. The ornamental D. kutejensis has highly regarded fruit, with no offensive
smell and soft pliable spines, but has proved less adaptable outside of its native
Borneo.
The genus is native to South East Asia, with wild durian found in Borneo
and Sumatra. The tree is mainly cultivated in Sri Lanka, South India,
Southern Burma, Thailand, Cambodia, Vietnam, Malaysia, Indonesia, Borneo,
Mindanao (the Philippines) and New Guinea. It has been spread throughout
the tropical world, with the general name of durian (Indo-Malaysia) or variants
duren (Indonesia), duyin (Burma), thureen (Cambodia), thurian (Thailand) and
saurieng (Vietnam).
ECOLOGY
Soil
Deep, well-drained sandy clay or clay loam are best. Poor drainage in heavy
soils is conducive to Phytophthora root rot.

75
76
Chapter 4
Climate
The tree requires abundant rainfall. Most production occurs in areas with a
mean yearly rainfall of 15002500 mm, well distributed throughout the year,
although higher rainfall is recommended. Drought for more than 3 months
leads to irreversible tree damage; irrigation overcomes this limitation.
Growth is limited below a mean monthly temperature of 22C, although
owering and fruiting appear to be more prolic. Temperatures below 10C
cause premature leaf abscission, while temperatures of up to 46C can be
tolerated. Durian is grown successfully at up to 800 m near the equator to
18 north and south. Shade (3050%) is necessary for young trees before they
reach 0.8 m in height following eld planting. The shade is reduced slowly over
12 months to full sun. Durian is not photoperiodic for ower initiation.
Tree
Durian trees of 2040 m tall are not uncommon, although grafted trees
are normally 812 m. The tree has a straight, low-branched trunk with an
irregular and dense crown tending to conical. The leaves are simple, alternate,
leathery, drooping, oblong or elliptic, and 820 cm long and 46 cm wide,
with a light- to dark-green upper surface and a silvery or brown rusty color
underneath. The wood is brittle, grainy and dark brown to reddish in color,
and shrinks on drying. The effective root depth for water and nutrient uptake
is shallow (2030 cm).
Flowers
Flower clusters arise on laterals, main branches and occasionally on the tree
trunk (Fig. 4.1A). Clusters have up to 825 long-stalked (58 cm) pendulous
white petal owers, which open over a period of 23 weeks. The eshy outer
covering, the epicalyx, splits into two to four parts on the day when full
anthesis occurs between 3 pm and late evening, when owers emit nectar
and a sour-milk odor. Five distinct bundles of shaving brush clusters of
stamens emerge during the late afternoon to midnight (Fig. 4.2). The epicalyx,
sepals, petals and stamens then all drop. The pistils fall 37 days later, if not
pollinated.
Flowering normally occurs during or near the end of a dry period that
reduces shoot growth (Fig. 4.3). In monsoon climates, owering takes place in
the dry season. A 1014 day continuous dry period is necessary for cv. Chanee
to ower. Some areas near the equators have two crops a year (e.g. early March
Durian
77
Fig. 4.1. Durian owers occur on main branches and sometimes the trunk itself (A).
When split open, the fruit show the locules with aril-covered seeds (B). To prevent
fruit abscission and damage from the fruit falling to the ground as they approach
maturity, the fruit are tied to the tree (C). After harvest, the arils are removed from
the fruit and sold in trays covered with plastic wrap (D).
Fig. 4.2. Durian ower, leaf and fruit.

The diagram shows part of the fruit rind
removed to exhibit seeds enclosed by
the aril.
78
Chapter 4
Fertilize and
Irrigate
Harvest
Vegetative
Growth
Growth
Suppression
Flowering
Fruit
Development
Continuous
Dry Period
(714 days)
Fig. 4.3. Harvesting to the owering cycle in durian requires a continuous dry
period (1014 days for cv. Chanee). The length of the required dry period can
be reduced with the use of paclobutrazol. (After Chandraparnik et al., 1992a;
Salakpetch, 2005.)
and April, and September or October in Malaysia). In Thailand, owering

occurs from March, after the cool dry season. Low doses of growth retardants
stimulate owering, especially as a growth ush nishes, supporting the idea
that a period of slow vegetative growth is needed for owering. Ethephon,
diaminozide and naphthalene acetic acid do not induce early owering, and
gibberellic acid (GA3) delays owering. Flower forcing can be achieved by
applying growth retardant (paclobutrazol), removing ower clusters from the
rst crop, and applying fertilizer and irrigation following a dry period.
Flower buds normally emerge in 68 weeks, while other buds can remain
quiescent as small clusters for up to 32 weeks before full development. Dormancy
at this stage is observed when young ower buds at the dot stage are exposed to
a small amount of rainfall and continuous growth is interrupted. Some owers
do develop, leading to more than one cycle of fruit set on a tree per year.

Self-incompatibility is common and cross-pollination between different
cultivars is often necessary. Poor pollination gives poor seed and aril
development and irregular-shaped fruit. However, single-variety plantings in
Thailand show no apparent reduced fruit set. Pollination is by nectarivorous
bats and moths at night, when bees are not active. Because of the size of
the sticky pollen, which tends to clump, wind pollination is probably not
important. Application of GA4 paste to durian pedicel at 6 weeks after anthesis
reduces fruit drop, and increases fruit size and weight by up to 30%.
If pollination is successful, the color of the young fruit changes from light
brown to light green, and the ovary starts to enlarge within a week. The ovary
has ve locules, with each locule containing seven to nine ovules. However, not
all the ovules develop and a mature fruit contains an average of only 13 mature
seeds.
Durian
79
Fruit
Dry Mass (g)
The fruit is a round, ovoid, cylindrical or ellipsoid capsule (Fig. 4.1B and 4.2),
derived from a single ovary, that will dehisces into three to ve segments when
ripe. The green to brownish fruit weighs up to 8 kg, and can be up to 30 cm
long and 20 cm in diameter. It is densely covered with stout, pyramidal, long
sharp spines (1 cm) on a thick brous rind that is white on the inside. The
spines have trichomes and lenticels on the surface. The fruit is divided into
ve smooth-walled compartments, each containing one to six glossy, creamy
to red-brown seeds up to 26 cm long, enclosed by a white to yellowish soft
sweet aril (Fig. 4.1B). The aril may be odorless, but can also have a powerful
odor suggesting garlic, onion and strong cheese with a background of a fruity
smell. The smell is caused by thiols, esters, hydrogen sulde and diethyl sulde.
The edible aril (2035% total weight) has a smooth, custard-like to rmer
texture. In selected cultivars, the seeds are rudimentary or small compared
with the wild types.
Excessive owering and fruit set, such as in Thailand, may require thinning
of owers or young fruit, 46 weeks after anthesis. The degree of thinning
depends on the variety. One to two fruit are left per panicle, with 50150 fruit
per tree. There is a positive relationship between aril weight and seed number.
The fruit matures in 90150 days with a sigmoid growth pattern, depending
on variety and temperature (Fig. 4.4). At maturity, the fruit abscises at the
Time from Anthesis (Weeks)

Fig. 4.4. Growth of durian fruit, showing total fruit weight and the increase in fruit
rind, aril and seed weight. (After Valmayor et al., 1965.)
Chapter 4
Starch and Soluble Solids (%)
80
Time at 30C (Days)
Fig. 4.5. Changes in starch, soluble solids and rmness of the aril of harvested
Chanee durian during ripening and fruit dehiscence. (After Ketsa and Pangkool,
1994; Sriyook et al., 1994).
articulation of the pedicel with the inorescence, ripening in 24 days, with

the fruit normally splitting into three to ve segments of irregular width at
the stylar end. Ripening-related increases in soluble sugars and a decrease in
starch and aril rmness are still ongoing when dehiscence occurs (Fig. 4.5)
Leaf ushing during early fruit development, 2156 days from anthesis,
may reduce the number of fruitlets and increase the percentage of deformed
fruit. Later ushes can also reduce esh quality, leading to hard esh, brown
esh discoloration and uneven esh color development. Use of high potassium
and phosphorus and no foliar fertilizer and growth retardant may therefore
have a role in modifying young leaf ushing and fruit development needs as
sinks for available carbohydrate from mature leaves.
Breeding
Durian (2n = 56) shows considerable diversity, with only a few of the 300
named varieties in Thailand being commercially grown. Hybridization within
the genus is reported to be relatively easy, and wild species may be able to
contribute to disease resistance.
Pollen is released over 34 h, and collected pollen remains viable for 24 h
in a refrigerator for hand pollination the next day. Pollen viability can decline
Durian
81
from 96% when released to 75% in 48 h. Hand pollination is carried out on

emasculated owers a few hours before dusk when the stigma becomes sticky;
receptivity is considerably reduced the following day. The long period before
bearing (710 years) limits crop-improvement research.
Selection criteria include an aril with sweet avor and good texture, few or
small seeds, large aril percentage, tree retention before fruit abscission, high
yield of fruit with marketable weight (1.5 kg), elongated to round shape, good
shelf life, good rind color and thickness, reduced rind dehiscence and increased
length of harvest season. Superior varieties have a thick, yellow, berless, crisp
pulp and small aborted or rudimentary seeds (Table 4.1).
Major cultivars
Selection programs in Thailand and Malaysia have involved hundreds of
cultivars, with only a few being recommended. Little work has been reported
from Indonesia. The variety Chanee is in high demand in Thailand as it has
less of a garlic avor than other varieties (Table 4.1). Others in Thailand
include Monthong (Golden Pillow), Kan Yao (Long Stalk), Luang and
Kradum Thong. Malaysian varieties have a D and a number, indicating
characterization by the government department and later a research institute.
Table 4.1. Comparison of three commercial Thai durian varieties (Hiranpradit et al.,
1992).
Characteristic
Mon Thong
Chanee
Kan Yao
Fruit set
Very good
Effect of high fruit retention
Poor
Phytophthora resistance
Fruit
Seed number
Seed size
Aborted seed
Edible esh
Flesh texture
Flesh ber
Flesh color
Odor
Physiological disorders
Ripening
Full ripe
Processing
Preserved
Frozen
Poor
Good
Good
Poor fruit quality,
branch die-back
Poor
High
High
Coarse, sticky
High
Mild
Mild
Few
Not uniform
Firm esh
Delicate
High
Attractive
Many
Uniform
Wet, pungent
High
Large
Delicate, sticky
Low
Uniform
Few
Firm eshy
Very good
Very good
Medium
Acceptable
Poor
82
Chapter 4
Examples include D-2 Dato Nina, D-7 Repok B-2, D-10 Durian Hyan,
D-24, D-98 Katoi, D-99, D-114 and D-117 Gombak, Sitokong, Sukun,
Mas, Parang and Bakul. Old Singapore varieties include Tan Chye Siam
and Jurong.
In Malaysia, a variety mix of 60% D-24, 30% D-99 and 10% D-98 or D-10
is recommended to ensure pollination. Similar mixed stands extend the harvest
period.
CULTURAL PRACTICES
The seed is sexually produced and monoembryonic, and gives viable progeny.
It is short-lived (1 week), especially if exposed to sunlight, and can germinate
in 38 days after the aril is removed. Uniform trees are always obtained by
grafting and rooting of cuttings; air layering is not successful. Inarching
has a 50% success rate, but is not popular because of the many months
necessary for success. Patch or cleft budding from ushing shoots onto pencilthin 2-month-old rootstock is used. Chanee rootstock is frequently used in
Thailand, with related species used to achieve some root-rot resistance.
Field preparation
Deep ripping and disking is recommended, followed by incorporation of
manure at the tree planting site 612 months before planting. The site is often
prepared in the wet season.

Planting distances are commonly 816 m (40156 trees/ha) in either
a square or triangle. The wider spacings are required to prevent canopy
competition between mature trees. Narrower spacings can be used initially for
some production before the canopy closes, with trees thinned in years 810.
In Thailand, interplanting is sometimes practiced with rambutan and langsat
before the durian comes into full production. Transplanting is performed just
before the wet season using 1-year-old trees.
Total rainfall and its distribution are crucial for regular production and tree
growth. Irrigation is essential if the dry period is longer than 3 months and
Durian
83
during owering and fruit development. Microsprinklers and drip tubes

are used, with the shallow root depth (2030 cm) suggesting application in
frequent small amounts. The crop coefficient for different stages of bearing
tree development has been determined (Table 4.2). Mild water stress is needed
to initiate owering, and this requirement is indicated in the crop coefficient
of 0.00. During ower development, late fruit growth and ripening, the crop
coefficient is 0.75 and higher. Chanee is reported to require less irrigation
than other varieties.
Pruning
Trees are trimmed to remove laterals and watershoots, leaving only a central
leader. In Thailand and Malaysia, trees may be left to grow unrestricted for
the rst 23 years before the interior of the tree is thinned. Flower and fruit
thinning are carried out to limit fruiting and to prevent the upper limbs from
bearing fruit, because the fruit load can cause limb breakage. Fruit are also
removed from the main trunk to prevent excessive competition. Heavy crops
may require propping or limb tying (Fig. 4.1C). Trees are sometimes topped at
10 m to aid in crop management.
Fertilization
Animal manures and mulches are still used in Malaysia and Indonesia,
but are not favored in Thailand. Up to 15 g of phosphorus is added to the
planting hole. Fertilizer can be applied in the irrigation water. The basal
Table 4.2. Crop coefcient (Kc) for durian

to estimate the daily water requirement
(WR) for different stages of development
based on evapotranspiration (ET). WR = Kc
u ET (Hiranpradit et al., 1998).
Stage of development
Vegetative growth
Floral initiation
Floral development
Fruit setting
Early fruit growth
Late fruit growth
Fruit maturity
Kc
0.60
0.00
0.75
0.50
0.60
0.85
0.75
84
Chapter 4
dressing of fertilizer should be applied only after budded or grafted plants

are well established in the orchard. Periods of active growth, particularly
the 2 months before owering, should be supported by adequate fertilization
(Fig. 4.3). During the rst 5 years, a 14:4:3.5 mix is recommended for nonbearing trees, then 12:4:7 twice yearly, increasing from 100 g to 4 kg/tree/
year to 12 years of age. Compound fertilizer is recommended at ower-bud
emergence, supplemented by a side dressing if there is a good fruit set, and
a third application after harvest. The nutrients removed by 1 ton of fruit
2.4 kg nitrogen, 0.4 kg phosphorus, 4.2 kg potassium, 0.3 kg calcium and
0.4 kg magnesium indicate a need for adequate nitrogen and potassium
fertilization. Calcium fertilizer may also be needed.
Pest management
The major problems of durian are patch canker and root rot. Patch canker,
caused by Phytophthora palmivora, occurs at or just above ground level,
eventually girdling and weakening the tree. Good drainage, removal of
vegetation and use of resistant rootstocks are the most common control
measures. The disease cannot be controlled by cultural practices once trees
have been infected, although newer fungicides show promise. Inarching of
seedlings to mature trees is practiced to control the diseases. Root rot can also
be caused by the same organism, with Mon Thong being very susceptible.
The same organism, P. palmivora, causes rotting of immature and mature
fruit, leading to high losses during rainy weather. Fruit on the ground can be
attacked by Sclerotium rolfsii.
Various stem borers and leaf-eating larvae have been reported on durian,
although damage is minimal. The fruit is not regarded as a fruit-y host if the
rind is unbroken. Rats, monkeys, orangutans, pigs and elephants are attracted
to ripening fruit, sometimes leading to signicant losses. Four-meter-high
electric fences are used to deter elephants.
Failure of the aril to soften and uneven softening are frequently observed
disorders. Another disorder leads to a watery aril with a at and dull taste,
and occurs especially during the rainy season. The cause of both disorders is
unknown.
Weed management
The soil around the tree should be kept free of weeds, especially in the rst
year. Control is by hand or with an approved herbicide. Mulches that do not
touch the trunk are also used.
Durian
85
Orchard protection
Shade is needed after eld planting, and is gradually reduced over 12 months.
Windbreaks are benecial to prevent ower damage, dropping of young fruit,
and the breakage of branches laden with fruit. Branches may have to be
propped up if heavily laden.

Harvesting
To prevent natural fruit abscising, fruit may be tied to the limb or harvested
at maturity (Fig. 4.1C). Maturity can be judged by the appearance, a hollow
sound when tapped with the nger, aril dry matter content and days from
anthesis. Fruit at 85% maturity, based on days from anthesis and rind
characteristics, ripen to excellent quality in less than 1 week at 22C. Fruit
that is 95% mature has already commenced ripening, while 75% mature
fruit may ripen but with inferior quality. Fruit collected from the ground after
falling are more subject to disease and fracture, and have a short postharvest
life (23 days, instead of the 78 days when picked).
Postharvest treatments
In Thailand, the fruit is harvested with the peduncle attached and the
peduncle is wrapped in a leaf or paper, reportedly to extend shelf life. Fruit
are cleaned and, if to be exported, brushed to remove insects, especially scale.
Fruit are graded on weight, shape, size and defects. Defects include disease,
insects, mechanical injury and esh disorders. The standards vary with
variety. The fruit is packed into a truck bed and bamboo baskets for the local
markets and cardboard cartons (46 fruit/carton) for export. In general,
fruit should not be stored at temperatures below 15C to avoid chilling injury
unless fully ripe. This depends on the variety, however; Mon Thong has
a slower ripening rate and can be held for 23 weeks at 15C. Symptoms of
chilling injury include blackening between the spines and failure of the aril
to soften. The postharvest life can be extended by holding the fruit at 57%
oxygen at 22C, with high CO2.
For Chanee and Kan Yao, the respiratory and ethylene peaks plateau or
decline when the fruit is overripe, while in Mon Thong the peak occurs when
overripe. Chanee is at its optimum eating stage for only a few days, while Mon
Thong remains at this stage for a longer period.
86
Chapter 4
Marketing
The fruit is marketed fresh, with a consumer preference for fruit that weigh
24 kg. The relatively short season can cause a glut at the markets and low
prices. The main season in Thailand lasts for 23 months. Choice varieties
demand and receive higher prices than other varieties. Soft arils are sold
fresh in shrink- or stretch-wrapped trays (Fig. 4.1D), and ripe fruit and arils
are frozen especially for export. Fruit that matures out of the peak season
command high prices.
Singapore and Hong Kong are major markets. Consumers in Singapore
prefer fully ripe fruit with no splitting, while those in Thailand prefer the rmer
pulp of less ripe fruit with fewer volatiles. Partially ripe fruit are difficult to open
without damaging the aril.
WORLD PRODUCTION AND UTILIZATION

World production
Peak tree production occurs at 1215 years and annual production averages
50100 fruit/tree (418 t/ha). Individual fruit generally weigh from 1.5 to 4
kg, occasionally reaching 8 kg. Commercial cultivation occurs in Australia,
Borneo, Indonesia, Malaysia, the Philippines, Thailand and Vietnam.
Thailand is the largest producer, with 711,371 t being produced in 1992 from
97,238 ha, while peninsular Malaysia produced 384,500 t from 61,294 ha.
Indonesia records 228,668 t from 44,016 ha.
Total world production is unknown. The fruit is highly prized in the markets
of south-east Asia and probably has limited appeal elsewhere. There is a
demand among ethnic groups familiar with the fruit in large temperate cities.
Utilization
The fruit is most frequently eaten fresh. Minimally processed, ready-toeat durian is becoming popular and easier for people who have difficulty in
dehusking or opening the fresh fruit (Fig. 4.1D). This product is suitable for
those who want to eat only one or two locules of the pulp. The durian aril
with the seeds intact is removed from the fruit locules, placed in a foam tray
and overwrapped with polyvinyl chloride lm. Durian pulp can be frozen,
dehydrated and processed into durian powder, durian paste (durian cake)
and durian chips. Its products can be used in salads, tempura, chowder, cakes,
ice cream, cookies, candy and biscuits. The dried product is used as a avoring
in ice cream, confectionery, pastry and soft drinks. The mature seeds can be
roasted and eaten as a snack (Subhadrabandha and Ketsa, 2001).
Durian
87
Table 4.3. Composition of 100 g edible

portion of durian (Ketsa and Paull,
2008).
Constituent
Edible portion (%)
Proximate
Water (g)
Energy (kcal)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
Vitamin C (mg)
Durian
22
64
640
2.7
3.4
27.9
0.9
1.0
40
1.9
44
70
40
0.35
0.2
0.7
90
23
The aril contains 67% water, 2.5% protein and 2.5% fat. It is high in
carbohydrate at 28.3%, and also contains 0.02% calcium and 25 mg/1000 g
vitamin C (Table 4.3).
FURTHER READING
Anon. (1987) Fruits in Thailand. Department of Agricultural Extension, Ministry of
Agriculture and Co-operatives, Bangkok.
Ketsa, S. (1997) Durian. In: Mitra, S.K. (ed.) Postharvest Physiology and Storage of Tropical
and Subtropical Fruits. CAB International, Wallingford, UK, pp. 323334.
Ketsa, S. and Paull, R.E. (2008) Durian Durio zibethinus, Bombacaceae In: Janick, J. and
Paull, R.E. (eds) Encyclopedia of Fruit and Nuts. CAB International, Wallingford, UK,
pp. 176182.
Morton, J.F. (1987) Bombacacea. In: Fruits of Warm Climates. Creative Resources Systems,
Inc. Winterville, North Carolina, pp. 287291. Available from: http://www.hort.
purdue.edu/newcrop/morton/durian_ars.html. Accessed 21 February 2011.
Salakpetch, S. (2000) Durian production in Thailand. In: Proceedings of the Tenth Annual
International Tropical Fruit Conference, 2022 October 2000. Hilo Hawaiian Hotel,
88
Chapter 4
Hilo, Hawaii, pp. 2529. Available from: http://www.hawaiitropicalfruitgrowers.

org/10th%20Inter national%20Tropical%20Fruit%20Conference%20
Proceedings%202000.pdf. Accessed 25 February 2011.
Siriphanich, J. (2011) Durian (Durio zibethenus Merr.). In: Yahia, E. (ed.) Postharvest
Biology and Technology of Tropical and Subtropical Fruits: Volume 3. Woodhead
Publishing Limited, Cambridge, pp. 80114.
REFERENCES
Baldry, J., Dougan, J. and Howard, G.E. (1972) Volatile avoring constituents of durian.
Phytochemistry 11, 20812084.
Brooncherm, P. and Siriphanich, J. (1991) Postharvest physiology of durian pulp and
husk. Kasetsart Journal (Natural Science) 25, 119125.
Chandraparnik, S., Hiranpradit, H., Punnachit, U. and Salakpetch, S. (1992a)
Paclobutrazol inuences ower induction in durian Durio zibethinus Murr. Acta
Chandraparnik, S., Hiranpradit, H., Salakpetch, S. and Punnachit, U. (1992b) Inuence
of Thiourea on ower bud burst in durian, Durio zibethnius. Murr. Acta Horticulturae
321, 636640.
Chua, S.E. and Teoh, T.S. (1973) Propagation by approach grafting and woody cutting
of some tropical fruit and ornamental trees. Singapore Journal of Primary Industries
1, 8795.
Chua, S.E. and Young, S.K. (1978) The use of approach, bud, and wedge grafting
technique to propagation durian (Durio zibethinus Murr.), rambutan (Nephelium
lappaceum, L.) and mango (Mangifera indica, L.). Singapore Journal of Primary
Industries 6, 94101.
Germplasm Resources Information Network. (2011) Durio Ans. United States
Department of Agriculture. Available from: http://www.ars-grin.gov/cgi-bin/
npgs/html/genus.pl?4046. Accessed 11 March 2011.
Hasan, B.M. and Yaacob, O. (1986) The growth and productivity of selected durian
clones under the plantation system at Serdang, Malaysia. Acta Horticulturae 175,
5558.
Hiranpradit, H., Lee-Ungulasatian, N., Chandraparnik, S. and Jantigoo, S. (1992a)
Quality standardization of Thai durian, Durio zibethinus Murr. Acta Horticulturae
321, 695704.
Hiranpradit, H., Someri, S., Chandraparnik, S. and Detpittayanan, V. (1992b) Clonal
selection of Durio zibethinus Murr. Acta Horticulturae 321, 164172.
Hiranpradit, H., Chandraparnik, S. and Salakpeteh, S. (1999) Integrated Technology to
Improve Durian Production, 2nd edn. Kasetsart University Press, Bangkok, Thailand.
Ketsa, S. and Pangkool, S. (1994) The effect of humidity on ripening of durians.
Postharvest Biology and Technology 4, 159165.
Ketsa, S. and Paull, R.E. (2008) Durian Durio zibethinus, Bombacaceae In: Janick, J. and
pp. 176182.
Lim, T.K. (1990) Durian Diseases and Disorders. Tropical Press SDN, BHD, Kuala
Lumpur, Malaysia, pp. 6072.
Malo, S.E. and Martin, F.W. (1979) Cultivation of Neglected Tropical Fruits with Promise.
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Part 7. The Durian. US Department of Agriculture, Science and Education

Administration, New Orleans, Louisiana.
Mamat, A.S. and Wahab, A.A. (1992) Gibberillius in the developing ower and fruit of
Durio zibethinus Murr. Acta Horticulturae 292, 101106.
Morton, J.F. (1987) Bombacacea. In: Fruits of Warm Climates. Creative Resources
Systems, Inc., Winterville, North Carolina, pp. 287291.
Moser, R., Ruvel, D. and Greve, R. (1980) Volatile constituents and fatty acid composition
of lipids in Durio zibethinus. Phytochemistry 19, 7981.
Nanthachai, S. (1994) Durian: Fruit Development, Postharvest Physiology, Handling and
Marketing in ASEAN. ASEAN Food Handling Bureau, Kuala Lumpur, Malaysia.
Ng, S.K. and Thamboo, S. (1967) Nutrient removal studies on Malaysian Fruits: durian
and rambutan. Malaysian Agricultural Journal 46, 164182.
Niranpradit, N., Chandraparnik, S. and Salakpetch, S. (2003) Technology of Durian
Production. Kasetsart University, Bangkok, Thailand.
OGara, E., Guest, D.I. and Hassan, N.M. (2004a) Occurrence, distribution and utilisation
of durian germplasm. In: Drenth, A. and Guest, D.I. (eds) Diversity and Management
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OGara, E.O., Sangchote, L., Fitzgerald, S., Wood, D., Ching Seng, A. and Guest, D.I.
(2004b) Infection biology of Phytophthora palmivora Butl. in Durio zibethinus L.
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Salakpetch S. (2005) Durian (Durio zibethinus L.) owering, fruit set and pruning. In:
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Salakpetch S., Chandraparnik, S. and Hiranpradit, H. (1992a) Pollen grains and
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Salakpetch, S., Chandraparnik, S., Hiranpradit, H. and Punnachit, U. (1992b) Sourcesink relationship affecting fruit development and fruit quality in Durian, Durio
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Soegeng-Reksodihardjo, W. (1962) The species of Durio with edible fruits. Economic
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Sriyook, S., Siriatiwat, S. and Siriphanich, J. (1994) Durian fruit dehiscence water
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Stanton, W.R. (1966) The chemical composition of some tropical food plants. VI durian.
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Tai, L.H. (1973) Susceptibility of durian clones to patch canker disease. MARDI Research
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Takakazu, T. (2000) Bird-pollination of three Durio species (Bombacaceae) in a tropical
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Tongdee, S.C., Chayasombat, A. and Neamprem, S. (1987b) Effects of harvest maturity
on respiration, ethylene production, and composition of internal atmospheres of
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surface coatings on weight loss and internal atmosphere of durian (Durio zibethinus
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Yaacob, O. and Subhadrabandhu, S. (1995) The Production of Economic Fruits in SouthEast Asia. Oxford University Press, Kuala Lumpur, Malaysia.
5
GUAVA
BOTANY
Introduction
Guava belongs to the family Myrtaceae, which has more than 80 genera
and approximately 3000 species distributed throughout the tropics and
subtropics, mostly in the Americas, Asia and Australia. Species range from
tall trees to shrubs and woody creepers. Many are cultivated as ornamentals
and for products such as timber, oil, gum, tannin, resin, spices and fruit. Spices
found in this family include cloves, nutmeg, cinnamon and allspice.

Five genera of interest because of the fruit: Psidium, Eugenia, Syzygium,
Myrciaria and Feijoa. All usually have simple opposite leaves and bisexual
owers, with four or ve calyx lobes, that are separated or united at the base.
Flowers have four or ve petals and numerous stamens. The ovary is usually
inferior and has one to three or more cells. The fruit is a berry or capsule, or
rarely a drupe or nut-like. It has few to many seeds.
Psidium
This genus is composed of approximately 150 species of evergreen trees
and shrubs in the American tropics. A good taxonomic classication of this
genus is lacking. P. guajava L. is by far the most widely known and distributed
species. Common names include guava, guajaba, gujaba, goyavier and jambu
bat (Malaysian). P. friedrichsthalianum (O. Berg) Medenza is known as the
Costa Rican jelly guava. It is a shrub or small tree with square branchlets
and glossy leaves above and pubescent below. The fruit is small (2.53.0 cm),

91
92
Chapter 5
sulfur-yellow and very acid. The plant has some difficulty fruiting at sea level,
but does well at higher elevations. It is cultivated as a backyard plant in Costa
Rica.
P. guineense Swartz. (P. araa Raddi; P. molle Bertol), the Brazilian jelly guava,
has highly acid yellow fruit of 2.53 cm diameter when ripe. It bears heavily
at sea level and is quite tolerant to lower temperatures. P. littorale Raddi var.
longipes (O. Berg) Fosb. and P. littorale Raddi var. littorale were formerly known
as P. cattleianum Sab. and P. cattleianum var. lucidum (Degner) Fosb., respectively.
The variety longipes is popularly known as the purple strawberry guava or
Cattley guava, while var. littorale is known as the yellow strawberry guava or
yellow Cattley, and they are vegetatively indistinguishable. The fruit of both
varieties are about 1.52.5 cm in diameter and seedy. They are somewhat subacid, but with a pleasant aroma.
Eugenia
This genus comprises a large and heterogeneous group of evergreen trees
and shrubs of the American tropics and the Old World. Most of the Asian
species were transferred to the genus Syzygium when Eugenia was revised, thus
making Eugenia a smaller group. The Eugenia differ from Syzygium in having
cotyledons that are usually united; the seed-coat is smooth and free from the
pericarp, and the inorescence is generally a raceme of pedicelled owers.
E. brasiliensis Lam (E. dombeyi [K. Spreng.] Skeels) is commonly called the
Brazilian cherry, and produces cherry-size, dark-red to black fruit on trees up
to 15 m tall. E. uniora L. (E. michelii Lam), commonly called Surinam cherry
or pitanga, is a native of Brazil. It has an eight-carpellate (furrowed) fruit,
which is red to purple when fully ripe and spicy and acid. It is frequently used
as an ornamental.
Syzygium (Jambosa DC)
This genus contains 400500 species, mostly from south-east Asia. It is
described in full in Chapter 11.
Myrciaria
This genus contains some very interesting species such as M. cauliora and
M. jaboticaba, which are closely related species called jaboticabas. The trees
and shrubs have good-tasting fruit that are borne on old stems and branches
(cauliory). Another important species is M. dubia (HBK) Mc Vaugh or M.
paraensis O. Berg, the camu camu. This fruit has a high vitamin C content
(2780 mg/100 g) and is exported from Peru, Bolivia and Brazil as a direct
source of vitamin C, normally as fruit pulp. For several months of the year the
plant grows wild on lands that are ooded by rivers in the Amazon valley, but
it can also grow under normal soil conditions with irrigation.
Guava
93
Feijoa
This genus is represented by two evergreen species in South America. F.
sellowiana O. Berg is well known as feijoa or pineapple guava and is found wild
in southern Brazil, Paraguay, Uruguay and northern Australia. It grows well
in parts of California under dry subtropical climates.

The most well-known species in this family, guava (P. guajava L.), is native to
the American tropics. The extent of its dissemination in the pre-Columbian
period is obscure. The English name guava probably came from the Haitian
name guajaba. The Spanish explorers took guava to the Philippines, while the
Portuguese disseminated it from the Philippines to India. It spreads easily
and rapidly throughout the tropics because of an abundance of seeds with
long viability. The species has become naturalized to the extent that people
in different countries consider guava to be indigenous to their own region. It
is now also grown in the subtropics. The hardiness of guava has made it an
invasive pest, especially in pasture lands.
ECOLOGY
Soil
Guava is adapted to a wide variety of soil types. Trees will thrive on shallow,
infertile soils, although growth and production are poor (Duarte, 1997). It
responds well to soils with good drainage and high organic matter, with a pH
range of 57. Cultivation in soils with a pH of less than 5 or higher than 7 will
usually lead to symptoms of zinc and iron (Fe) deciency, respectively. Guava
is fairly tolerant to salt. The threshold for seedling growth is 1.2 dS/m, and at
14 dS/m seedling survival is 25% after 50 days (Tavora et al., 2001). Fruit size
and ascorbic acid decrease with increasing salinity to 5.0 dS/m.
Climate
Rainfall
Guava does best with abundant moisture; 10002000 mm is optimal. Rainfall
should be well distributed throughout the year, and should not be less than 600
mm/year. Adequate moisture is required during vegetative growth, and for
optimum owering and fruit development. The tree does tolerate drought for
up to 6 months, but prolonged drought induces defoliation. New growth and
94
Chapter 5
owers occur soon after the beginning of the rainy season. In the dry tropics,
owering is greatly inuenced by water availability. The ideal rainfall pattern
for guava fruit production is alternating dry and wet conditions. Articial
cycling to induce owering attempts to mimic this wet and dry cycle. In regions
with prolonged dry seasons, ower induction can be managed by irrigation.
Drought and very low humidity during owering drastically reduce fruit set,
with almost complete post-set drop being observed. Low moisture conditions
during fruit enlargement reduce fruit size and puree recovery, caused by
shrinkage of the inner pulp and its separation from the inner rind. Waterstressed trees will drop most fruit of 12 cm in diameter following heavy
irrigation or rain as the tree resumes vegetative growth.
Temperature
Guava does best in warm areas with abundant moisture. It will grow from
sea level to elevations exceeding 1500 m, if frost-free (Maggs, 1984). The
optimum temperature is reported to be 2328C, with fruit set signicantly
reduced by temperatures outside of this range during owering (Fig. 5.1).
However, guava production in Hawaii thrives very well between 15.5C
and 32C. In spite of guava being native to the tropics, it can produce in
subtropical areas or at altitudes of up to 1700 m. This adaptability has led to it
being found in many places of the world and sometimes viewed as an invasive
weed. Young plants are reported to be killed at 2C, if this temperature is
prolonged. In areas where winter night temperatures are 57C for a few
hours a night, such as in Okinawa, growth ceases and leaves become purple.
Commercial production is difficult in subtropical regions with insufficient heat
calories during the winter months, and the time from anthesis to fruit harvest
can increase to 220 days.
Fig. 5.1. The effect of temperature during owering on guava fruit set. (After Huang,
1961.)
Guava
95
Low winter temperatures during the dry season lead to natural defoliation,
and owering will commence as soon as warm weather and rainfall induce
new growth ushes and fruit set. In some places, defoliation is induced by
withholding irrigation during the winter (16 months) to protect the plant
from cold damage (Gonzlez et al., 2000).
Light
Light saturation for this typical C3 plant is high: greater than 925 mol/m2/s
photosynthetic photon ux. Guava does not show any visible response to
photoperiod. Seedlings normally require 27 years before owering. Longer
sunlight duration leads to greater shoot growth.
Wind
Guava, though hardy, does benet from windbreaks. Trees grafted on seedling
rootstocks have tap roots that provide substantial anchorage. However, trees
produced from rooted cuttings are subject to uprooting by winds of 6580
km/h for the rst 3 years, probably because of faster top growth than root
growth, and the shallower root system of these plants in relation to seedling
roots. Trees exposed to prevailing winds of 1632 km/h gradually develop
branches away from the winds, with little branch growth against the wind.
Windbreaks are crucial if high-quality dessert-type cultivars are being grown
for the fresh market.
Tree
Guava is a shrub but, under high-moisture conditions, will grow to 69 m
in height and spread, with trunk diameters of 30 cm or more. The trunk is
short, freely branching from the base. Under cultivation, a single trunk tree
is developed by proper pruning and training. The bark is smooth but peeling,
and greenish-brown to brown in color. Branches are pliable and hence rarely
broken by winds. The leaves, arranged in pairs, are oblong or oval, 1018 cm
in length, smooth on the upper surface, nely pubescent on the undersurface
and prominently veined (Fig. 5.2). Young green twigs are square, becoming
rounded as they age.
Flowers
Flowers occur singly or in clusters of two to three at the leaf axils of current
and preceding growth (Fig. 5.2). The perfect bisexual owers are white and
2.53.5 cm in diameter, with four or ve petals, numerous stamens and one
96
Chapter 5
Fig. 5.2. The leaf, ower and fruit of guava.
style. The tubular calyx encases the bud and splits into four or ve segments
at anthesis. The ovary is inferior with four or ve carpels, each containing
numerous ovules in axile placentation. Floral morphology favors selfpollination, but considerable cross-pollination occurs. Flowers open between
5 and 7 am, depending on the cultivar and morning temperatures; the calyx
splits on the previous day. The anthers usually dehisce at anthesis or shortly
before. Bees are the principal pollinators. It takes approximately 30 days from
oral initiation to anthesis, and approximately 3.5 months from anthesis
to fruit maturation. Two owering peaks occur in India and Hawaii. These
natural peaks can be altered by changing weather conditions or cultural
manipulation (Figs 5.3 and 5.4).

Natural pollination
Most guava clones have few problems with fruit set. Guava pollen generally
has high rates of germination, except for triploid clones such as Indonesian
Guava
97
Fruit Thinning
and Bagging
Harvest Duration (Days)
Cycling to Harvest (Days)
Defoliate, Prune
and Fertilize
Cycle to Harvest (Days)
Fig. 5.3. The steps involved in cycling guava trees for fruit production. The
relationship between month of cycling, induced by pruning and fertilizing, and
days to fruit harvest (after Bittenbender and Kobayashi, 1990) and the relationship
between days to harvest from cycling and the duration fruit are available for harvest
(H.C. Bittenbender, unpublished data) on the island Kauai, Hawaii. Fruit thinning
and bagging are only used for the production of dessert fruit, not processing fruit.
Seedless. Fruit set in the triploid cultivars is good when grown together
with diploid clones as a pollen source. Fruit set as high as 90% is obtained
when owers 48 h old are pollinated. The period of stigma receptivity in
Beaumont, a Hawaii cultivar, is about 48 h. Post-fruit-set drop does occur as
a result of factors other than pollination.
Some degree of self- and cross-incompatibility among guava clones has been
shown; some combinations are totally incompatible, while reciprocal crosses
produce some fruit (Table 5.1). Beaumont produces 100% fruit set when selfpollinated and 6080% fruit set by cross-pollination with other guava cultivars
(Ito and Nakasone, 1968). These incompatibilities are the result of inhibition
of pollen-tube growth in the style.
Fruit set by chemicals
Use of growth regulators has been tried, primarily to produce seedless fruit.
Auxin-like compounds do not reduce fruit drop. Fruit set is increased when
50 g/ml gibberellic acid is applied, and the fruit contain fewer seeds and
98
Chapter 5
Fig. 5.4. Change in the monthly production of guava in 1975, when cycling was
not used, and in 1990, when a major portion of Hawaiian production was cycled to
give a single owering peak.
Table 5.1. Cross-compatibility of some guava clones as percentage mature fruit

from self- and cross-pollination (Ito and Nakasone, 1968).
% Fruit set
Male parent
7197
7199
Beaumont
Lucknow
Indonesian
Seedless
7197
67
33
33
29
7199
33
64
50
50
Beaumont
62
67
100
100
Lucknow
57
33
33
27
Indonesian Seedless
57
14
33
Female parent
improved quality characteristics (e.g. total soluble solids, ascorbic acid,

reduced fruit drop). When unopened ower buds are emasculated and styles
cut off, and the gibberellic acidlanolin mixture is applied to the cut surface,
seedless fruit are obtained. Fruit obtained by this treatment are larger than
those from untreated owers (Shanmugvelu, 1962). The fruit, however, show
six to eight prominent ridges and swelling at the calyx end.
Guava
99
Fruit
The fruit is botanically a many-seeded berry, varying in size from 2.5 to 10
cm in diameter (Fig. 5.2). The shape can be globose, ovoid, elongated or pearshaped. The skin is yellow when ripe, while the esh can be pink, salmon,
white or yellow. The skin texture can be smooth or rough. The inner wall of
the carpels is eshy and of varying thickness, and the seeds are embedded in
the pulp (Fig. 5.2). Flavor and aroma vary widely among seedling populations.
There are low-acid, sweet types, bland types that are low in both sugars
and acidity, and high-acid types. The undesirable musky aroma is more
pronounced in the fully ripened low-acid, sweet types.
Fruit growth follows a simple sigmoid curve (Fig. 5.5) and pulp growth
parallels total fruit growth. The time from anthesis to harvest can vary from
about 120 days to more than 220 days (Fig. 5.3), depending on the temperature
during fruit development. Cultivars also vary by up to 60 days from anthesis to
fruit maturity.
Cytogenetics and genetics
The genus Psidium, except for P. guajava, is represented by di-, tetra-, hexaand octoploid species (2n = 22) (Hirano and Nakasone, 1969a). Plants of P.
Mass (g)
Total Fruit
Time from Anthesis (Days)
Fig. 5.5. The growth of total fruit and pulp mass from anthesis to maturity in
Hawaii. (After Paull and Goo, 1983.)
100
Chapter 5
guajava are preponderantly diploid, but triploidy, tetraploidy and aneuploidy

have been reported. Seedless cultivars have highly abnormal meiotic division,
with univalents, bivalents and laggards, as well as bridging of chromosomes,
indicating a hybrid origin. Pollen is functional, although pollen-tube growth
is inhibited in the style, indicating a gametophytic type of self-incompatibility
(Table 5.1). Pollination stimulates fruit development.
Breeding
The ower morphology of guava favors self-pollination, with 35% outcrossing
reported. This provides a heterozygous, open-pollinated seedling population,
with adequate genetic variation for selection of desirable commercial
types. Because of the many-seeded fruit, guava lends itself to controlled
hybridization if there are specic objectives such as disease or insect
resistance. Otherwise, open-pollinated progenies are adequate for cultivar
development. A collection of germplasm is a necessary part of a breeding
program. Seth (1960) reported cross-incompatibility when crossing Behat
Coconut with Lucknow-49, SI with Behat Coconut and Behat Coconut
with Apple Colour. No fruit set was obtained, and the cause was attributed to
triploidy or other genetic factors.

Selection criteria or objectives in a breeding program would include all or
some of the following. Fruit selection criteria include: (i) large (200340 g),
with few seeds and thick pulp; (ii) white for dessert fruit or a dark-pink pulp
color, particularly for processing; (iii) avor and aroma characteristic of
fresh guava, with no woody off-avor or muskiness; (iv) more than 10% total
soluble solids; (v) acidity of 1.251.5% for processing fruit and 0.20.6% for
dessert guava; (vi) vitamin C content of 3000 mg/kg or higher; (vii) minimum
number of stone cells; (viii) good postharvest shelf-life; and (ix) resistance
to fruit diseases and insects. Tree selection criteria include: (i) vigorous,
spreading, low-growth type; (ii) resistance to tree diseases and pests; (iii) high
yield; and (iv) dwarng rootstock.
Cultivars
There are many cultivars of guava and many orchards are planted using
seedlings. In general, the white- and cream-eshed cultivars are sweeter and
more suited for consumption as fresh fruit, while the pink- and red-eshed
cultivars are more acid and suited for industrial purposes. Some cultivars
Guava
101
developed in Thailand have very large fruit and cream to white esh. These
fruit are consumed in the green-ripe stage while still rm and have almost no
odor. These types have been expanded to Taiwan and are now being grown
in Central America. Low-acid cultivars with white esh such as Allahabad
Safeda and Apple Colour have been developed in India (Table 5.2), though
Sardar (formerly Lucknow 49) remains highly recommended. South
Africas Malherbe and Fan Retief are mild, sweet, dessert types, with lightpink esh, for out-of-hand consumption. They are also suitable for canning
as halved fruit. The principal cultivar in Bangladesh is Kazi Piara (Paull and
Bittenbender, 2008). China (Guangdong) recommends the cultivar Zhenzhu
as a dessert type (Chen et al., 2002).
Beaumont, Kahua Kula and Waiakea are recommended for processing.
Beaumont was the rst processing cultivar introduced to the Hawaii industry,
and was the only recommended processing guava until the introduction of
a selection Kahua Kula (Nakasone et al., 1976). Beaumont produces large
fruit, ranging from 145 to 235 g, averaging about 170 g (Table 5.2) and a 78%
puree yield. The fruit are mildly acid, with total titratable acidity ranging from
less than 1.0% during the summer to about 1.25% in winter. Soluble solids
range from about 7% in winter to 10% during summer. The fruit characteristics
of Kahua Kula are similar. Both varieties are good yielders, exceeding 227 kg/
tree/year after the fth year, with some pruning.
Taiwan has developed and introduced some guava varieties to Central
America through its cooperative work schemes. The fruit are mostly harvested
at the mature-green stage for eating at that stage. These varieties include
Tai-kuo-bai, Pai-bai, Lay-a-bai, An-a-bai and Taiwan-yeh-bai. The fruit
of Tai-kuo-bai generally weigh 400800 g, and sometimes more than 1 kg.
They have a crisp and juicy white pulp, are pale green when ripe and are for
fresh use only. The fruit of Pai-bai is the most popular in Taiwan. The 180
g fruit are pale yellow and pear-shaped, with white esh. The branches of the
tree tend to grow horizontally. Lay-a-bai produces a 300 g elongated fruit with
greenyellow peel and white succulent pulp, and 7.2Bx. An-a-bai produces
an elongated fruit with pale-yellow peel, white pulp and very good aroma. The
mean weight is 320 g. The fruit has a short postharvest life when ripe and is
mostly used for juice. Taiwan-yeh-bai has year-round production, producing
300 g pale-green fruit. This is one of the most-used varieties for processing
(Romn Zeledn and Wann Fuh, 1994).
In recent times, producers in South-east Asia and Australia have concentrated
on growing seedless cultivars such as Thai seedless and Indonesian Seedless.
Both dessert and processing cultivars of excellent quality have been developed
in Florida. Ruby, Supreme and the hybrid, Ruby u Supreme are excellent
dessert types (Campbell, 1963). Cuba has an interesting dwarf guava cultivar
called Roja Enana Cubana or EEA 1840 that is used by the industry or for
fresh consumption. The fruit are pear-shaped and have a reddish-pink pulp.
White Selection of Florida, Pentecosts, Pedra Branca, Branca de
Varieties
Origin
Fruit
weight (g) Flesh color
102
Table 5.2. Fruit analyses of selected guava varieties (Nakasone et al., 1967).
Cavity
diameter
(cm)
Total
diameter
(cm)
Seeds
(%)
Soluble
solids (%)
Total
acidity*
(%)
173
136
235
White
White
Pink
5.0
4.6
5.2
7.5
6.5
7.3
3.2
6.3
3.0
12.5
8.8
12.0
0.6
0.4
1.3
Burma
Hong Kong Pink
Burma
Hong Kong
210
218
White
Pink
4.9
4.0
6.4
7.6
4.5
1.8
10.7
10.0
0.2
Hong Kong
White
Indonesian White
Indonesian
Seedless
Patillo (7197)
Pink Acid (7198)
Hong Kong
181
White
4.7
6.7
4.2
12.5
0.4
Indonesia
Indonesia
105
176
White
White
4.1
4.3
6.3
7.3
1.4
0.7
10.5
12.5
0.3
0.4
Florida
Florida
113
167
Pink
Dark pink
4.0
4.7
5.8
6.9
3.3
2.7
11.9
11.7
1.7
1.7
Sardar (Lucknow
49)
6362
India
227
White
4.4
7.3
3.3
11.0
0.4
Florida
176
Pink
4.5
6.7
2.0
10.5
0.4
6363
Florida
250
White
4.6
7.1
2.1
11.4
0.3
7199
Florida
153
Pink
5.1
6.9
1.9
11.3
0.4
*As percentage citric acid.
Sweet, lumpy surface

Sweet, seedy, musky
Processing, mild
acid, large-fruited
Sweet, seedy
Sweet, thick-eshed,
few seeds
Sweet, seedy
Sweet, few seeds
Sweet, thick-eshed,
no to few seeds
Processing, high acid
Processing, high
acid, good color
Sweet, thick-eshed,
medium seedy
Sweet, few seeds,
good color
Sweet, large-fruited,
few seeds
Sweet, thin-eshed,
few seeds, slight
muskiness
Chapter 5
Allahabad Safeda India

Apple
India
Beaumont
Hawaii
Comments
Guava
103
Valinhos, Branca de Kumagai or Kumagai, Ogawa Branca, Red selection

of Florida, Pirassununga Vermelha, Pirassununga Blanca, IPA B 22 and
Paluma are cultivated in Brazil (Gonzaga Neto and Monteiro Soares, 1994).
Venezuela produces many varieties, including those in the Mara 1 to Mara 8
series. Local selections exist in most other producing countries.
CULTURAL PRACTICES
Propagation
Sexual
Seed germination is used to produce seedlings in breeding and selection
programs or to produce rootstocks for grafting of desirable cultivars. Seeds are
sown in a well-draining media and more than 90% of fresh seeds germinate
in 1520 days. Guava as a short-cycle fruit tree starts producing about a year
after planting, reaching maximum production when trees are 4 years old
(Avilan, 1988).
Asexual
Container-grown seedlings may be budded or grafted when stem diameters
are 1220 mm, with greater diameters being especially suitable for budding.
Budding is preferred over other grafting techniques because as bud growth is
faster and each bud on a scion or bud wood is a potential plant. The patch-bud
technique or the modied Forkert method give good results. Success requires
a vigorously growing seedling, where the bark peels readily and well-prepared
bud wood with swollen axillary buds. A skilled propagator can achieve 90% or
better success (Hamilton and Nakasone, 1967).
The side-wedge method is most frequently used in grafting. The scion wood
or bud wood should be prepared approximately 1014 days before cutting by
removing the leaves from the branch. Normally the branches that will serve to
obtain budwood should be girdled or defoliated to encourage axillary buds to
enlarge and greatly accelerate growth when budded or grafted. Wood that is
shedding or has already shed its bark and is smooth, grayish-green and without
leaves gives good results in budding or grafting.
Rootstocks have been created for specic purposes, in Cuba a nematodetolerant and vigorous guava stock called Cotorrera is used for grafting or
budding. In other cases, rootstocks are used to control guava wilt and rootknot nematode (Marin et al., 2000). Other species such as P. molle Bertol. or
P. guineense Sw. have also been tried as rootstocks (Alix and Duarte, 2000).
Rooting of greenwood cuttings has been attempted in the past. In 1962,
Jelicoeur reported 44% rooting using leafy cuttings treated with indole-3butyric acid (Jelicoeur, 1962). Cuttings with two to four leaves can be treated
with naphthalene acetic acid and indole-3-butyric acid, with the latter
104
Chapter 5
being more effective and practical for producing large quantities of plants
in a relatively short time (Abdul et al., 2002). Rooting is achieved within 6
weeks to 2 months, although there are clonal differences in rooting ability. A
subterminal leafy cutting is used in Cuba, with one or two pairs of leaves taken
when the branch bark is yellow, not green or brown. Cuttings are kept moist
with mist or microsprinklers for 68 weeks, then repotted into plastic bags
and acclimatized before being taken to the growing-out area of the nursery
until they attain transplant size. Root cuttings and suckers can also be used
by stimulating supercial root development via wounding with a heavy spade.
For root cuttings, a medium diameter root piece of 1225 cm long and about
0.51.0 cm diameter should be used. In vitro methods have also been developed
for rapid propagation of guava for disease resistance (Vos et al., 2000).
Field preparation
Field preparation follows conventional procedures. Soil pH is best maintained
between 5 and 7, and lime should be incorporated during land preparation.
Transplanting and plant spacing

Well-grown rooted cuttings of approximately 68 months old can be eldtransplanted after hardening in the direct sun for several weeks. Transplanting
can be done at any time of the year if irrigation facilities are installed.
Planting holes should contain approximately 225 g of a complete fertilizer,
mixed into the soil at the bottom of the hole. Manure may also be used.
Postplant application of 115 g of fertilizer is spread around the plant, mixed
with the surface soil and irrigated.
Guava trees tend to have a spreading growth habit and are relatively fast
growers. Trees are spaced from 47 u 67.6 m (about 250420 trees/ha).
Closer spacing requires severe pruning and thinning, but trees planted at 600
1000 trees/ha in a northsouth orientation in Taiwan have been reported to
give higher yields in the rst 5 years. The dwarf Cuban varieties are planted at
1.52.5 u 45 m, resulting in 1000 to almost 1500 plants/ha. Other varieties
under tropical temperatures with good soil moisture are planted at densities
of almost 1000 plants/ha, with pruning to maintain the height below 3 m. In
Central America, the Taiwanese varieties are planted at 2 u 3 m spacing when
interplanted with mango, while 4 u 6 m is normally recommended for a single
crop. At even higher densities (73,000 plants/ha) trees grow in height with
few laterals, as compared with controls, although the lowest density (27,000
plants/ha) has been found to produce the highest yield (Mohammed et al.,
1984). The spreading growth habit of guava does not appear to support such a
high-density system as the tree ages or the cultivar tested was unsuitable.
Guava
105
Irrigation
Guavas thrive in areas with long dry periods and over a wide range of rainfall.
Adequate moisture is required during vegetative growth, and for optimum
owering and fruit development. Almost complete post-fruit-set drop is
observed during drought. In the dry tropics, owering is greatly inuenced by
water availability. Heavy owering follows the onset of the rainy season.
Drip and under-tree irrigation (microsprinklers) are being used increasingly
in production to replenish daily water loss (2550 mm/week). In large orchards,
where irrigation is done by sections, the microjet or a low-sprinkler system is
more desirable. These irrigation systems also make it easier to apply fertilizer
for immediate effect. During fruit growth, irrigating at 7080% of panevapotranspiration gives best yields. Where there is a danger of frost damage,
such as in certain areas of Mexico, irrigation is withheld during the coldest
months to induce leaf fall and hardening of the plants (Gonzlez et al., 2000).
Pruning
Branches can be pruned immediately or soon after harvest. Trees are
pruned and fertilized to induce new axillary growth on which owers will be
produced. Heading-back branches induces new, long, whip-like shoot growth,
with sparse owering compared with cutting at a fork. Branches grown
horizontally are far more productive than vertical ones. For dessert cultivars,
an ideal tree shape is one with no branches 0.51.5 m from the ground and
three to six horizontal branches. This at shape allows for ease of orchard
management, fruit thinning and bagging, and reduced labor for harvesting.
However, this shape requires the ends of the branches to be supported.
Pruning is more effective than chemical defoliation to concentrate the new
growth and owering. In some trials, branches with a diameter of more than 3
cm and all terminals of more than 0.8 cm in diameter are removed once a year,
compared with the regular system that eliminates all terminals of 0.81.5 cm
diameter and cutting back to 1.01.5 m all branches to the height of 1.51.8 m.
In the subtropics, low winter temperatures reduce vegetative growth, similar
to in the dry season in the tropics. Growth in the tropics resumes with the
onset of rain. In these cases, the plant is pruned when growth is minimal. It is
difficult to change the harvest season under these conditions, but pruning does
concentrate the harvest peak (Lopes et al., 1984).
Cycling
In the tropics, guavas produce varying amount of fruit throughout the year.
In Hawaii, a small harvest peak occurs between April and May and a larger
peak between September and November (Fig. 5.3). Flowering is prolic under
106
Chapter 5
natural conditions of drought and/or low temperatures followed by irrigation

or warm temperatures, especially if the trees have shed their leaves. This leads
to a short harvest season and results in over-supply and lower prices. The
fruiting peaks can deviate with prevailing weather conditions and cultural
practices. Since owers are produced on new growth, factors that stimulate
new growth such as irrigation, fertilization, pruning and defoliation also
stimulate owering (Fig. 5.6). Varying these production practices can shift
and concentrate the harvest period (Fig. 5.3), avoid fruit gluts, increase yields
and even out year-round fruit availability and labor needs for harvest. A 25%
urea spray plus wetting agent applied to 15-month-old guava seedlings of the
variety GA9-EX39 produced a three-fold increase in yield over untreated
trees at 22 months of age. The harvesting period was shortened from 15
weeks in the control to 4 weeks for sprayed trees. Pruning is repeated after
harvest every 79 months, the period between one pruning and the end of
harvest. Initial owering after pruning occurs in about 56 weeks (Fig. 5.6),
and the harvest period is concentrated to 46 weeks at the end of the cycle.
Starting the cycle at 4- to 6-week intervals in different sections of the orchard
allows continuous year-round production.
Defoliation is often used with fertilization to induce more intense owering
and a shorter harvest period (Fig. 5.3). Urea is a powerful defoliant and can be
Time from Defoliation (Weeks)
Fig. 5.6. The pattern of leaf production on new shoots after defoliation, ower
production, fruit set and fruit abscission: 67% of owers abscised, followed by
abscission of 69% of fruit, before harvest in Nambour, Queensland, Australia. Fruit
matured 18 weeks after anthesis. (Menzel and Paxton, 1986)
Guava
107
combined with ethephon and a detergent. A 1525% urea solution causes 90%
of leaves to abscise, and a 5% urea solution with ethephon and a wetting agent
is equally effective; the latter is easier to prepare (Shigeura et al., 1975; Shigeura
and Bullock, 1983). Gibberellic acid acts synergistically with ethephon to
induce abscission. Following defoliation, new leaves appear on new shoots in
34 weeks, with a peak of owering occurring 912 weeks after defoliation
(Fig. 5.6). This is followed by fruit set and a peak of abscission of young fruit.
This fruit abscission may be related to the ability of the new leaves to develop
all the fruit set, as the percentage fruit abscission correlates with the number
of fruitlets set per tree. A single fertilizer application after defoliation, rather
than continuous or periodic applications, can be more effective in sustaining
high yields.
Continuous pruning
Continuous light pruning is more labor-intensive and is used especially under
tropical conditions where temperatures are warm and water is available
year round. Under this system, plants are continuously pruned by heading
or pinching back the current seasons growth so they keep producing new
shoots, owering and fruit. The procedure is to pinch the current seasons
branches above their fourth to sixth pair of leaves. This is practiced in Cuba
with the dwarf cultivars Roja Enana Cubana and 123, which are heavy
producers (IICF, 1998b; 2000; RELAFRUT, 1999). Fruit can be harvested
practically every week of the year, except when cold spells reduce vegetative
growth and owering. Continuous production is more suited to small
orchards, as pickers need to go through the whole orchard looking for mature
fruit to harvest. In larger orchards that are cycled into owering, fruit pickers
concentrate only on the blocks that are maturing.
Fertilization
It is a general practice in most areas to apply 110225 g of a complete
fertilizer three to four times a year for the rst 2 years. When trees begin to
produce commercial quantities of fruit, fertilizer is usually given after harvest,
with pruning and irrigation to encourage new axillary growth. A foliar spray
of urea and phosphate solution at the preowering stage increases mean
yields by as much as 45% and enhances fruit total soluble solids, sugars and
ascorbic acid. In Central America, the recommendation for the Taiwanese
varieties is to fertilize four times a year (Romn Zeledn and Wann Fuh, 1994)
since production is almost continuous (Table 5.3). In subtropical climates
where winter temperatures are relatively low, only one major crop season
occurs; fertilization is adjusted accordingly, with an application in the spring
as new growth starts.
108
Chapter 5
Table 5.3. Fertilization recommendations for Taiwanese guava cultivars in Costa

Rica (Romn Zeledn and Wann Fuh, 1994).
Element (g/plant)
Age of
plant Nitrogen, Phosphorus, Potassium
(years)
N
P2O5
K2O
1
2
34
56
78
910
11
40
60
120
200
250
300
400
40
60
120
120
140
180
200
Product (g/plant)
Ammonium Ammonium Potassium
sulfate
phosphate
sulfate
40
60
120
200
250
300
400
200
300
600
1000
1250
1500
2000
220
330
660
660
770
990
1100
80
120
240
400
500
600
800
Table 5.4. Fertilizer recommendations for guava plants according to age, cultivar
and soil analysis (Natale et al., 2008).
P-resin (mg/dm3)
<6
Age
(years)
N (g/plant)
cv. Rica
01
120
12
240
23
480
612
1330
K-exchangeable (mmolc/dm3)
>30
<0.8
P2O5 (g/plant)
0.81.5
1.63.0
3.0
K2O (g/plant)
120
90
60
30
120
80
40
240
180
120
60
240
160
80
480
360
180
90
cv. Paluma
01
100
100
80
50
30
12
200
100
50
30
200
150
100
50
23
400
200
100
60
400
300
150
80
Fertilizer recommendations have been prepared in Brazil according to

plant age, plant cultivar and soil analysis (Table 5.4) (Natale et al., 2008). A
tentative foliar-analysis guide to guava fertilization on the basis of dry matter
is: 1.7% nitrogen, 0.25% phosphorus, 1.5% potassium, 1.25% calcium and
0.25% magnesium. The microelement recommendation is 20 ppm zinc, 60
ppm manganese, 8 ppm copper and 20 ppm boron (Shigeura and Bullock,
1983). Several studies have shown response to micronutrients such as zinc
and manganese. Different varieties of guava show different uptake patterns of
macro- and micronutrients (Table 5.5) (Natale et al., 2008).
Guava
109
Table 5.5. Uptake of macro- and micronutrients by the guava cultivars Rica and
Paluma in experiments in the municipalities of Jaboticabal and Sao Carlos, Sao
Paulo state, Brazil (Natale et al., 2008).
cv. Rica
Macronutrient
g/kg
g/m
Nitrogen
Phosphorus
9.80
1.20
Potassium
cv. Paluma
kg/ha
g/kg
g/m
kg/ha
1353
166
66.8
8.3
8.6
0.9
1146
121
84.3
8.9
1.57
2167
107.1
12.4
1662
122.8
Calcium
0.80
110
5.4
0.7
94
6.9
Magnesium
0.80
110
5.4
0.9
114
8.4
Sulfur
1.10
152
7.5
0.9
114
8.4
mg/kg
g/m
g/ha
mg/kg
g/m
g/ha
0.83
41
0.67
50
Micronutrient
Boron
Copper
1.11
54
11
1.48
109
Iron
15
2.07
98
14
1.88
139
Manganese
28
3.87
188
14
1.88
139
Zinc
13
1.73
84
15
1.95
144
Fruit thinning and bagging

Fruit thinning and bagging are frequently performed with fruit that are to
be consumed fresh. Taiwanese varieties that are harvested year round and
eaten in the green-mature stage are normally thinned, especially varieties
with large fruit, leaving one to two fruit per node and a maximum of four
fruit per branch. In Brazil, the recommendation is to leave one fruit out of
four in order to obtain larger size fruit, with the central fruit at the node being
preferred. In other cases, fruit are thinned and bagged to one or two fruit per
new shoot, and one or two fruit per node, 3040 days after ower anthesis
(Fig. 5.3).
Where labor is available, young fruit destined for the fresh-fruit market are
thinned and covered with a plastic or paper bag or wrapped with newspaper
to protect them from insects. This does not alter the growth rate or nal fruit
quality (Lam, 1987), although the fruit will be cleaner. The Taiwanese varieties
that are eaten in the green-mature stage are normally thinned and bagged when
the fruit are 1 cm in diameter. Paper bags are preferred in hot climates; if plastic
bags are used, the fruit must be protected with a net sleeve of polyurethane or a
polyethylene foam sock to avoid direct contact with the plastic bag, which can
110
Chapter 5
cause burns (Romn Zeledn and Wann Fuh, 1994), or to provide additional
protection from bruising. The fruit are sprayed with an approved preharvest
fungicide before the bags are placed.
Yields
The average production in Hawaii is 26.9 t/ha. In experimental planting of
5-year-old Beaumont, annual production was estimated at 50.7 t/ha at
a plant density of 198 trees/ha. To emphasize the importance of selecting
high-yielding cultivars, in Australian trials, at a plant density of 805 trees/
ha, with ve seedling lines of Hawaiian origin, 24.7, 37.2 and 61.5 t/ha were
obtained the rst three harvests at the same plant density, with no signicant
differences among seedling lines (Chapman et al., 1981).
A problem with young guava trees is that they carry light and variable
crops, and both young and old trees ower and fruit over protracted periods.
Management practices for fruit cycling greatly increase production. In some
trials, the mean yield has been 28 t/ha for cycled trees versus 9 t/ha for controls
at a density of 805 trees/ha. Fruit size was not affected by cycling. There was
a second crop 12 months later, also induced with urea sprays, with 40% more
fruit than in the rst year. There were also reductions in harvest period and
labor costs, and better control of fruit ies (Chapman et al., 1979). A yield of
up to 120 kg/tree can be obtained using the open-center at-pruning method
in Taiwan. The average is 8090 kg/tree, giving an average yield of 59.5 t/
ha at 600800 trees/ha. Reports from Cuba indicate that the dwarf variety
Enana Roja Cubana can produce 80105 t/ha with 1000 or 1481 trees/ha,
respectively, using the year-round pruning and harvesting method, in the fth
year after planting (IICF, 2000).
Pest management
Diseases
Some diseases are region-specic, while others are widespread wherever
guavas are grown (Table 5.6). Anthracnose is widespread and is considered
an important disease in most countries. Algal spot is also very common
but is not a serious problem, except if dessert-type fruit are sold on the
fresh market, because of the unsightly appearance of the fruit. Other types
of fruit rots are attributed to a number of organisms. Guignardia fruit rot
becomes serious in Hawaii when fruit are left to over-ripen on the tree or the
ground. Wilting of guava trees has been reported in South Africa and can
be controlled by using rootstock resistant to Penicillium vermoensenii (Vos
et al., 2000). In India and Pakistan, however, wilting seems to be related to
attacks by Fusarium oxysporum and Colletotrichum gloeosporioides. This can
Guava
111
Table 5.6. Important diseases of guava.

Common name
Organism
Anthracnose
Colletotrichum
gloeosporioides
Glomerella cingulata
(perfect stage)
Botrytis cinerea
Phomopsis psidii
Physiological
Pestalotia psidii
Macrophomina spp.
Rhizopus stolonifer
Gliocladium spp.
Fusarium solani
Guignardia spp.
Mucor hiemalis
Cephaleuros virescens
Glomerella fruit rot

Blossom end rot
Fruit canker
Fruit rot
Guava fruit rot
Guava wilt
Fusarium wilt
Guignardia rot
Mucor rot
Algal spots
Part(s)
affected
Region
Fruit
Worldwide
Fruit
Puerto Rico
Fruit
Fruit
Fruit
Fruit
Fruit
Fruit
Root
Root
Fruit
Fruit
Leaf, fruit
South Africa
India, South Africa
Hawaii, Australia
Australia
Caribbean
Hawaii
South Africa, Australia
India
Hawaii
Hawaii
Florida, Hawaii,
Caribbean
be controlled with the use of fungicides and antagonistic fungi (Ansar et al.,
1994).
Mucor fruit rot rst appears as a water-soaked area that later becomes covered
with yellowish, fuzzy mycelia and fruiting bodies of the fungus (Kunimoto et al.,
1977). The infection rate can be as high as 8090% and, as a wound parasite,
it is commonly associated with fruit-y oviposition wounds (Ito et al., 1979).
Culture control is possible by removing fallen fruit from the eld at intervals of
24 days, crushing underfoot during harvesting or lightly rolling the orchard
oor. Low-acid, sweet cultivars are more tolerant to this disease than acid types,
although some acid selections have shown high tolerance (Ito et al., 1976).
Blossom end rot of fruit appears to be widespread. In Hawaii, no organism has
been isolated and fungicidal sprays have been ineffective. Calcium application
to guavas largely alleviates this disease.
Insects
Many insects are common to all guava-growing areas (Table 5.7), with most
being present in small numbers and causing little damage. In Hawaii, Mitchell
(1973) compiled a list of approximately 45 species of insects and six species of
mites attacking guavas. These included species of aphids, thrips, scales, mealy
bugs, beetles, moth larvae, false spider, eriophyid and spider mites. There are
many species of parasitic wasps and predators that keep scale insects and
112
Chapter 5
Table 5.7. Important insect and nematode pests of guava.

Common name
Organism
Part(s) affected
Region
Mediterranean fruit
y
Natal fruit y
Oriental fruit y
Melon y
Caribbean fruit y
Ceratitis capitata
Fruit
Hawaii, South Africa
Ceratitis rosa
Dacus dorsalis
Dacus cucurbitae
Anastrepha striata
Fruit
Fruit
Fruit
Fruit
Stem, branches
Leaf
Leaf, fruit
South Africa
Hawaii
Hawaii
Caribbean,
American tropics,
Amazon area
South American
tropics
Amazon area
Australia
Worldwide
Bark
India
Root
Australia
Root
Root
Caribbean
Caribbean
Guava weevil
Conotrachelus
psidii
Guava stem moth
Timocratica albella
No common name Monolepta australis
Red-banded thrips
Selenothrips
rubrocinctus
Bark-eating
Indarbela
caterpillar
quadrinotata
Root-knot nematode Meloidogyne
incognita
M. arenaria
M. acrita
Fruit
mealy bugs under reasonable control. Fruit ies cause serious fruit damage
and fruit rot within 12 days on ripening. Fruit bagging, along with thinning
3040 days after anthesis, can signicantly reduce the problem and produce
high-quality, blemish-free dessert fruit. A review of guava pests in South
Africa included bird and bat control (Villiers and Grove, 2000). Bats frequently
attack ripening fruit in Cuba.
Thrips can be troublesome, causing leaf silvering and fruit scarication. The
skin of scarred fruit becomes russeted by disruptions of the epidermal layer and
corky-tissue development in the subepidermal area. Young fruit that are severely
damaged will often fail to develop and become mummied. Natural enemies
can keep thrips under fair control, although outbreaks do occur, especially
during the fruiting season. Several sweet cultivars, such as Allahabad Safeda,
Ruby u Supreme and Lucknow-49, have a higher degree of resistance to
thrips than others. This resistance increases somewhat with increasing foliar
levels of nitrogen and potassium. In the Amazon region, Anastrepha striata
(fruit y), the lepidopteran Timocratica albella (a stem and branch borer) and
Conotrachelus psidii (guava weevil) are potentially serious enemies (Vsquez et
al., 2002). The guava weevil spends its larval time in the fruit and feeds on the
seeds, producing petrication and premature fruit ripening. It is important to
stress that insecticides should be applied in the afternoon to protect honey-bees;
thus, better pollination and harvests will be obtained.
Guava
113
Weed control
Weed control is crucial during the rst 23 years of orchard establishment.
After that, the tree canopy provides adequate shade to minimize interference
by weeds. Mulching with black polyethylene sheets or heavy mulching with
organic materials, such as straw, dried grass, wood shavings or whatever
material is available, immediately around the plant reduces weed growth. A
herbicide such as glyphosate may be applied by rope wicks or rollers. Grass
alleys for harvest vehicles are maintained by mowing.
Orchard protection
Windbreaks are particularly recommended for plants coming from rooted
cuttings or air layers, since they will have a shallow root system and can be
uprooted with wind velocities of 6580 km/h, especially during the rst 3
years. Trees are not lost to winds after the fourth year, and this problem does
not occur at all if the root system is from a seedling plant. Windbreaks are
essential if dessert-type fruit are being produced, as the fruit skin is easily
damaged by rubbing.

Harvesting
Skin color is used to measure maturity and ripeness. The stage of fruit harvest
depends on the variety and the stage at which the fruit is to be eaten. If eaten
green, the fruit is harvested at the mature rm stage without any signs of
ripening. Fruit to be consumed when soft and ripe are harvested when they
show some sign of color change from green to yellow and initial softening.
Later harvesting, when the fruit is riper, can lead to a high number of fruit-y
stings and later larvae in the esh. Soluble solids can vary from 3% in green
fruit to more than 10% when ripe, and the acid from 0.2% to 1.5%; cultivars
vary greatly in sweetness and acidity. The fruit is commonly shipped in 4.5 kg
single-layer cartons, with foam sleeves or wrapping to prevent injury.
Harvesting is done manually, although tests with machine harvesting have
been conducted in some countries for fruit that is destined for processing.
Dessert fruit are always hand-harvested at the mature-green stage and
carefully handled to avoid injury, graded to size and packed carefully in cartons
for shipment. Processing fruit should be picked at the rm yellow to half-ripe
stage. Overripe fruit and those severely infected with fruit ies and diseases
should be destroyed rather than left to fall and rot in the eld, as these fruit
will otherwise become a source of continuous eld infection. Losses of 3540%
114
Chapter 5
can occur when fruit are harvested on a 3-day cycle, as the fruit rapidly ripen
and abscise. Harvesting fruit showing some yellow skin to the half-ripe stage
allows the interval between harvest cycles to be lengthened to about 3 days
with minimal losses.
Fruit are harvested into plastic buckets or picking bags worn by the harvester
with bottom delivery. They are then transferred into larger bins with capacity
for approximately 227 kg of fruit or into small wooden boxes. Harvested fruit
should be held in the shade until delivered to the processing plant. Sorting
of fruit according to maturity in the eld allows those that are less than fully
ripe to be held at ambient temperatures for ripening. When the supply of fruit
exceeds the processing capacity, ripe full-yellow fruit may be stored at 7.2C for
1 week and at 2.2C for as long as 2 weeks without injury.
Fruit-detachment force is related to the stage of fruit development and fruit
quality (Fig. 5.7), and is an important consideration if mechanical harvesting
is to be used. Fruit requiring a detachment force of 75 N or more are maturegreen fruit; the force declines to 10 N in overripe fruit, with a concomitant
decline in fruit softness and titratable acidity and an increase in total soluble
solids. Any mechanical harvester should be adjusted so that the force detaches
only yellowing fruit and minimizes loss of green fruit.
Guava is generally grown for processing. Color-break fruit held at 20C
develop full-yellow skin in 68 days. Fruit for processing can be held at 15C
to allow gradual ripening and delay the deterioration of quarter- and halfripe fruit. Ethylene can be used on mature-green fruit to accelerate ripening.
Immature fruit do not ripen properly and develop a gummy texture (Reyes
and Paull, 1995).
In areas free from fruit ies or where fruit bagging is practiced, low-acid
cultivars can be grown for fresh consumption. The marketability of these
mature-green fruit decreases as the storage period increases beyond 10 days,
although fruit packed in polyethylene bags can be stored at temperatures
of 810C for 14 days and be 100% marketable. Forced air, hydrocooling or
holding in a cold room can be used to cool the fruit to about 10C. Shelf life is
about 7 days when stored at 20C, while soft ripe fruit can be kept for about 7
days at 58C and a relative humidity of 9095%. Chilling injury can occur,
and the symptoms include skin scald, pitting and a failure to ripen if maturegreen or partially ripe when chilled. Browning of the esh can also occur.
Chilling injury will increase the incidence and severity of decay.
Packaging in polyethylene bags and the use of wax coatings create a modied
atmosphere that delays the ripening and softening of mature-green fruit.
Short-term treatment (24 h) with 10% oxygen and 5% CO2 before storage in air
at 4C for 2 weeks delays color development and reduces the development of
115
Acidity (meq/100 g FW)
Guava
Fig. 5.7. Relationship between fruit-detachment force, fruit softness, total soluble
solids (TSS) and titratable acidity of Kahua Kula guava and days from anthesis. FW,
fresh weight. (After Paull and Goo, 1983.)
chilling-injury symptoms, compared with fruit held in air (Bautista and Silva,
1997). The modied atmosphere generated in polyethylene and other plastic
packages reduces weight and ascorbic acid losses and extends the shelf life of
the fruit. Skin blackening is a problem with some wax coatings.
UTILIZATION
Guava is grown widely throughout the tropics and subtropics and is either
consumed fresh or processed. India, Mexico and Brazil are the largest
producers. Mexico has an estimated 21,000 ha (Gonzlez et al., 2000) and
produces both high-acid (processing) and low-acid (fresh, dessert) cultivars.
The leading export countries are Brazil, Mexico, the Dominican Republic,
India, Pakistan, Ecuador, Colombia, the Philippines, South Africa, Thailand
and Taiwan. Most of Indias production is consumed within the country.
Mexico is increasing its volume of exported dessert fresh fruit. Thai and
116
Chapter 5
Taiwanese cultivars have been developed for fresh consumption at the

mature-green stage.
When guava is consumed fresh, it is harvested at the mature-green stage
and eaten either as a rm, crisp, apple-like product or as a ripe fruit, depending
on the habits of consumers. In Latin America, people prefer to eat raw guava
fruit when it is fully ripe and has a very strong aroma. The cultivars eaten
fresh are sweet, larger and normally have white esh, compared with the more
acid, red-eshed processing cultivars. The undesirable musky aroma is more
pronounced in fully ripe, low-acid, sweet types, and is regarded as undesirable
by some consumers.
Guava is an excellent source of ascorbic acid, dietary ber, vitamin A and
Ca (Table 5.8) (Wenkam, 1990). The ascorbic acid is mainly located in the
skin, with a slightly lower concentration in the esh. The esh is high in pectin,
making it useful for jam and jelly, although pectin necessitates dilution when
Table 5.8. Composition of 100 g edible portion of
common guava, with seeds removed (Wenkam,
1990).
Constituent
Proximate
Water (%)
Energy (kJ)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
Common guava
8.5
230
0.3
0.1
15
2.4
0.5
15
0.3
16
292
6
190
0.06
0.06
1.3
109
Guava
117
guava puree is used in juice. The puree is used in juices, cakes, puddings, sauces,
ice creams, jams and jellies. Guava can also be dehydrated and powdered. In
many countries of Latin America, a relatively hard guava paste is made from
the puree by adding more sugar and evaporating water through boiling. This is
called ate or carne and is eaten instead of jelly or combined with cheese and
crackers.
Conventional processing requires hot lling of containers or freezing as a
means of preserving the product without adding preservatives. To eliminate
preservatives, aseptic methods with reduced heating that affect quality or
the more costly freezing are used. The aseptic-processed product requires no
refrigeration and the container is disposable. The puree shows little loss of
ascorbic acid and avor, but there is a signicant, though not objectionable,
change in color. Loss of the pink puree color in storage is more rapid at 38C
than at ambient temperatures (Chan and Cavaletto, 1982). Lowering the
storage temperature of aseptically processed guava juice to 10C retards, but
does not prevent, color change and the slight ascorbic acid loss.
Guava seed is generally considered a waste product. The seed contains 16%
crude protein, 18% crude oil, 20% crude ber and 44% carbohydrate on a dry
weight basis. The oil is 77% linoleic acid and can be used as an edible oil or for
paints.
FURTHER READING
Ali, Z.M. and Lazan, H. (1997) Guava. In: Mitra, S.K. (ed.) Postharvest Physiology and
Storage of Tropical and Subtropical Fruits. CAB International, Wallingford, UK, pp.
145165.
Bttenbender, H.C. and Kobayashi, K.D. (1990) Predicting the harvest of cycled
Beaumont guava. Acta Horticulturae 269, 197204.
Menzel, C.M. and Paxton, B.E (1986) The pattern of growth, owering, and fruiting of
guava varieties in subtropical Queensland. Australian Journal of Experimental Agriculture
26, 123128.
Reyes, M.U. and Paull, R.E. (1995) Effect of storage temperature and ethylene treatment
on guava (Psidium guajava L.) fruit ripening. Postharvest Biology and Technology 6,
357365.
Shigeura, G.T. and Bullock, R.M. (1976) Management of guava: cycling fruit set for
continuous production. Proceedings of the Tropical Region of the American Society for
Singh, S.P. (2011) Guava (Psidium guajava L.) In: Yahia, E.M. (ed) Postharvest Biology and
Technology of Tropical and Subtropical Fruits. Volume 3. Cocona to Mango. Woodhead
Publishing Ltd, Cambridge, pp. 213245.
Wilson, C.W. III (1980) Guava. In: Nagy, S. and Shaw, P.E. (eds) Tropical and Subtropical
Fruits. AVI Publications Inc., Westport, Connecticut, pp. 279289.
118
Chapter 5
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6
MANGOSTEEN
The family Clusiaceae (alternate name Guttiferae) is now regarded as

pantropical. It has approximately 50 genera and 1200 species, with eight
genera producing edible fruit. Linnaeus described the genus Garcinia based
on G. mangostana (mangosteen) and the genus Rheedia based on R. laterifolia.
The genus Garcinia was initially regarded as an Old World tropical genus
of dioecious evergreen trees or shrubs. However, taxonomists are now
including species from the New World that were formerly in the genus Rheedia
in Garcinia. Two thirds of the more than 200 species in the genus Garcinia
produce edible fruit. These range from 1 to 12 cm in diameter and have a
woody skin that varies from thin to thick.
The most praised of tropical fruit is G. mangostana L. (syn. Mangostana garcinia
Gaertner [1790]), otherwise known as mangosteen (English), mangostanier
(French), mangostan (Spanish), manggis (Malay), manggustan (Philippine),
mongkhut (Cambodia), mangkhut (Thai) and cay mang cut (Vietnamese).
Commercial production occurs in South-east Asia (Thailand, Malaysia and
Indonesia), where it is thought to have originated as an understory tree. In
the last two centuries, mangosteen has been spread to Sri Lanka, southern
India, Central America, Brazil, northern Australia, Hawaii and other tropical
countries.
Other Garcinia with edible fruit include the following. G. atroviridis a native
of peninsular Malaysia, Thailand, Burma and Assam (India); G. dulcis (Roxb.)
Kurz., a native of the Philippines and Indonesia; G. pseudoguttifera Seemann
from the Solomon Islands, Vanuatu, Fiji and Tonga; and G. xanthochymus,
from India, are cultivated for their fruit. G. humilis (Vahl) C.D. Adams (formerly
Rheedia laterifolia [L.] Herzog) grows wild in the oriental parts of Bolivia. In
Spanish, it is called achachair, shashair, ibaguaz, cachicheruqui and tapacuari.
Wild trees and fruit from recent plantings are harvested to sell in local markets
and for export. G. indica (Thouin) Choisy is known as kokum, the kokum butter
tree or Indian butter tree. It is native to the evergreen forests of the Western

123
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Chapter 6
Ghats mountain range of India at elevations of up to 1800 m, and is also found

in Assam and West Bengal.
These species are not considered further in this chapter. The focus here is on
mangosteen.
ECOLOGY
Soil
The mangosteen tree grows on a wide range of slightly acidic soils (pH 56),
with a high amount of organic matter being preferred. Heavy soils are
reported to be tolerated even with a weak root system. The tree is not adapted
to limestone soils, sandy alluvial soils or sandy soil that are low in humus. The
tree can withstand some waterlogging, and has a preference for a high water
table of 1.21.5 m below the surface.
Climate
The tree is regarded as a shade-tolerant, lower-canopy tree that is adapted
to humid tropical lowlands. The tree is found in the humid tropics. A short
dry season (1530 days) is required to stimulate owering, and then an
uninterrupted water supply. Stress should be avoided and irrigation may be
needed during the dry season, if the annual rainfall is less than 1270 mm.
Growth is slow below 20C and the trees are killed at 35C. The
photosynthetic rate is steady over a temperature range of 2736C, in 2050%
shade. The upper temperature limit is 3840C, with both leaves and fruit being
susceptible to scorching and sunburn. In the tropics, mangosteen is grown at
elevations of up to 1000 m, although at a slower growth rate than at lower
sites. Shade is essential during the rst 24 years in the nursery and eld.
Failure to provide shade leads to stunted growth and reduced leaf size, burnt
leaves, reduced frequency of leaf ushing and sometimes death. There are no
reports of photoperiod responses.
GENERAL CHARACTERICS
Tree
The 625 m tall, dioecious tree has a straight trunk, symmetrical branching
and pyramidal crown. Leaves are opposite on short petioles (Fig. 6.1), with
oblong or elliptical blades that are 1525 cm long by 713 cm wide. The
leaves are thick, leathery and glabrous, and olive green above and yellow
Mangosteen
125
Fig. 6.1. The leaf, ower and fruit of Garcinia mangostana.
green below. The small number of large stomata are found only on the bottom
of the leaf. Early seedling growth is dominated by shoot development and slow
root development. Secondary and tertiary root development does not occur
until 34 months after germination. Root hairs are very sparse on all roots.
The tree can reach 60 cm in 2 years, with one or two pairs of laterals. There
is a very large shoot-to-root ratio (6.24) at the seedling stage (Table 6.1), which
declines with age to 4.94 at 24 months. The juvenile phase lasts for about 16
pairs of laterals and the rst crop appears after 57 years. If growing conditions
limit growth, this phase may last for 1015 years. Young trees can make up
to six shoot ushes in 1 year. The effective root depth for water and nutrient
uptake is 90120 cm.
Flowers
The solitary or paired owers are found on the terminal branchlets inside the
outer canopy. Each ower has four sepals in two pairs, four thick and eshy
yellowgreen with reddish edges to the petals. The ovary is globose, with a
Chapter 6
126
Table 6.1. The shoot/root ratio of mangosteen

seedlings.
Seedling age (months)
Shoot-to-root ratio
6
12
18
24
6.24
5.75
5.5
4.94
four to eight lobed stigma and staminodes (Fig. 6.1). Flowers are borne at the
tips of older mature shoots and on more hidden branches. Initiation is noticed
as tip swelling, and the bud stage to anthesis takes 25 days. Male owers
are thought to be non-existent; although stamens and pistils are present at
the early stage of ower development, the staminate growth is stunted and
aborted.
Trees tend to bear in alternate years and bearing varies from tree to tree.
Trees generally ower after vegetative growth ushes and frequently twice a
year, depending on growing conditions and the number of growth ushes,
and especially after dry weather (Fig. 6.2). In Sri Lanka, low-elevation trees
fruit in May and July and those at higher elevations in September and October.
In Puerto Rico, unshaded trees fruit in July and August, and shaded trees in
November and December. India has two owering periods: one during the
monsoon season (July to October) and another from April to June.
Flower induction, pollination and fruit set

Flowering can be induced by a continuous dry period of 1530 days, which
limits apical bud growth. Three factors play a role in ower induction: the
threshold leaf water stress, the age of apical buds (at least 10 weeks) and
vigorous vegetative growth. The threshold leaf water stress is about 1.0 MPa
(Fig. 6.3). This water stress or dry period is followed by two heavy waterings
spaced 7 days apart. This irrigation regime can lead 3035% of the apical
buds to produce owers. Delaying the irrigation regime can lead to fruit
shedding of the early owers, while too early induction of irrigation can lead
to the remaining buds being vegetative. Fruit set is not a problem, although set
of 1015% of the owers can inhibit the remaining buds, leading those buds
to vegetative development.
Paclobutrazol (a gibberellin biosynthesis inhibitor) applied as a soil drench
or a spray to the leaves and buds also alters the owering cycle of mangosteen.
Treated trees ower about 40 days sooner. Application as a soil drench has a
residual effect of 2 years or longer. Coiling a wide band (3 cm) of wire tightly
around the trunk, 3040 cm above the soil, is also reported to induce earlier
owering.
Mangosteen
127
Apical Buds
>8 Weeks Old
Days
Heavy Irrigation
+ Second Irrigation
7 Days Later
Within
2 Weeks
Time to First Flowers (Days)
Fig. 6.2. Phenology of steps required to induce owering in mangosteen. The

tree must be vigorous with bright green leaves and wide and thick leaf blades at
owering. Flowering intensity depends on plant vigor, apical bud, age and length of
the drought period. (After Salakpetch, 1996.)
0.50
0.70
0.90
1.10
1.30
Fig. 6.3. The inuence of leaf water stress level induced by withholding irrigation
on days from onset of irrigation to appearance of owers and number of fruit per
tree. At a leaf water potential of 0.86 MPa, the leaves show slight wilting while at
0.08 MPa the leaves are noticeably wilted and at 1.12 MPa show severe wilting.
(After Salakpetch et al., 2006.)
Fruit
The globose and smooth berry develops parthenocarpically. The fruit is 48
cm in diameter and weighs about 80150 g, with a persistent calyx at the
stem end (Fig. 6.1). The pericarp is 510 mm thick and turns purple when
ripe. The skin contains a bitter yellowish latex and purple-staining juice. The
edible white aril (approximately 30% of total fruit weight) occurs in four to
128
Chapter 6
eight segments. The one or two segments containing brown, almond-shaped

apomictic seeds are larger.
There is no true seed, as the seed develops from the inner carpel wall, is
sometimes polyembryonic and has an underdeveloped embryo. At anthesis, it is
obvious which fruit segment will be seeded. Fruit development takes 100120
days from anthesis and up to 180 days in cooler areas or at higher elevations
(Fig. 6.4). Growth is initially dominated by the pericarp with aril dry matter,
not increasing until 20 days from anthesis and continuing throughout fruit
growth.
The 2n number has been variously reported as 5676, 8890, 96 and 120
130; the most commonly accepted number is 8890. The difficulty is in
counting the numerous small chromosomes in shoot tips. Mangosteen is only
known as a cultivated plant. Based on morphological characteristics it may be
an allopolyploid hybrid, as a female between G. hombroniana Pierre (2n = 48)
and G. malaccensis (2n = 42).
Apomictic seed and an absence of male trees or owers suggest all trees
belong to a single clone. However, some differences in tree and fruit shape,
rind thickness, leaf shape and area, and petal color do occur. This variation
has been conrmed by molecular DNA markers and may be a result of the
accumulation of natural mutations. Another hypothesis is that mangosteen
arose from more than one hybridization. The nucellar nature of the seeds,

Fig. 6.4. Mangosteen fruit growth as dry mass, showing total mass and separate
components of the rind, esh and seed. (After Poonnachit et al., 1992.)
Mangosteen
129
absence of diversity and long juvenile period (615 years) limit selection work.
Collections are maintained in South-east Asia, principally in Thailand and
Malaysia. No described cultivars have been reported. The exposure of seed
callus and cell cultures to irradiation has produced seedlings with increased
genetic variability.
CULTURAL PRACTICES
The apomictic seeds are viable for a short period (3 days, if dried) and are
best kept in moist peat moss or left in the fruit. Heavier seeds (>1.3 g) give
the best seedling survival, as initial growth is slow. Seeds should be planted
in freely draining growing medium under high humidity and shade. On
germination (23 weeks), a radicle and plumule emerge from opposite ends;
as soon as an adventitious root develops at the base of the young shoot, the
radicle dies. Very slow seedling growth is a major problem and is attributed
to poor seedling root development, having no root hairs and few laterals. A
porous medium is best for seedling growth, with peat moss, bark and coarse
sand being ideal. Growth increases signicantly when side shoots emerge. The
shoots generally emerge from every node.
Grafting onto mangosteen seedlings is not difficult, although such plants are
slower-growing and small-fruited with a shorter juvenile phase. Other rootstock
have been tested with variable results. Positive results have been obtained with
top-wedge grafting. Rooting of cuttings and air layers from mature trees have
failed, although cuttings from seedling can be rooted under mist.
Field preparation
No specic information is available regarding land preparation, although
high rates of organic matter are recommended. Practices normally follow
recommendations for other tree crops in the area. Deep-ripping is recommended
for compacted soils.

The long, delicate taproot and poor lateral root development mean that
transplanting should be performed with care. Planting holes are prepared (1.2
u 1.3 m) in advance and organic matter added a month before transplant.
Plants should have reached 60 cm before transplanting and a deep ball of
earth set out, then watered heavily.
Chapter 6
130
Because of the need for shade and humidity, trees are often planted in mixed
stands with durian, rambutan and coconut used as the dominant trees. An area
of 4080 m2 is allowed per mangosteen tree and trees 0.6 m high are planted
at 810 m (110140 trees/ha) or 1112 m if a mechanical harvesting aid is
to be used. Shade is maintained for 24 years, then gradually reduced to full
sunlight.
Mangosteen trees can withstand some water-logging but not drought, so
a constant supply of water is required. A continuous dry period of 1530
days should be imposed to limit apical bud growth (Fig. 6.2) and encourage
owering. The recommended crop coefficient at this stage is 0.00 (Table
6.2). This dry period is followed by two heavy waterings, spaced about 7 days
apart. The trees should then be regularly watered during fruit growth and
development at 8085% of evapotranspiration (Table 6.2). Trickle irrigation
or microsprinklers may be ideal for this crop.
Pruning
The regular pyramidal crown and slow overall growth limits pruning.
However, the tall nature of the tree (25 m) and fruit being borne singly make
harvesting difficult; dwarf rootstocks and pruning may therefore be useful.
Inside shoots and dead branches are removed along with suckers at the base
of the main trunk. Water sprouts should also be removed. Severe pruning is
never desirable. The limited pruning is carried out when there are no owers,
fruit or leaf ushes.
Table 6.2. The crop coefcient (Kc) for
mangosteen to estimate the daily water
requirement (WR) for different stages of
development, based on evapotranspiration
(ET). WR = Kc u ET (Hiranpradit et al., 1998).
Vegetative growth
Floral initiation
Floral development
Fruit setting
Early fruit growth
Late fruit growth
Fruit maturity
Kc
0.60
0.00
0.75
0.75
0.80
0.85
0.85
Mangosteen
131
Fertilization
Manure is recommended for young trees, along with mulching around
the tree base. Non-bearing trees require a low, steady supply of nutrients,
applied every 3 or 4 months. One recommendation is for 70 g nitrogen, 6 g
phosphorus and 50 g potassium per tree per year until maturity. About 27 kg
of complete fertilizer (10:10:19) per year is required for mature bearing trees.
Half is applied when vegetative growth is being stimulated after fruit harvest
(Fig. 6.2) and the remainder 25 weeks after anthesis. Dolomite can be applied
at 0.2 kg per tree per year of age to 15 years, with a constant application rate
for older trees. Manure is also used.
Pest management
On the Malay Peninsula, Cankers on stems and young and older branches
are caused by Zignoella garcineae P. Henn. The foliage on infested branches
withers and eventually the whole tree dies. Trees should be cut and destroyed
to arrest the spread. Thread blight, caused by Pellicularia koleroga Cooke, has
been reported in Puerto Rico under conditions of excess shade and humidity.
The smaller stems are rst attacked, with the blight becoming severe when it
attacks the leaves forming a whitish lm over the blade. The leaves turn a clear
brown, then darken before abscising. Removal of some shade and application
of a Bordeaux mixture or other copper fungicides give control. Postharvest
decay can be caused by Botryodiplodia theobromae.
Only a few insect pests have been reported, possibly due to the bitter sap.
Ants nesting in the tree can damage the growing tips. Mites can attack the
fruit surface and make it unattractive for market. Caterpillar larvae and grass
hoppers can cause some leaf damage. Fully ripe fruit are attacked by monkeys,
rats and bats.
Weed management
The slow growth of young trees means they can be quickly overtaken by
weeds. Organic mulch is often used to assist with weed control and reduces
evaporation from the soil around the trees. Mulch should not be placed
against the tree trunk. Plastic mulch can also be used.
Orchard protection
Shade is essential during the rst 24 years. Shading (3050%) can be
achieved with mixed stands or crops such as pigeon peas, bananas, plantains,
Chapter 6
132
rambutan, durian and coconut, placed at least 1.5 m from the mangosteen
tree. Cloth is also widely used to provide shade. Excessive shade (>50%)
produces tall and skinny trees. Cover crops (e.g. Crotalaria, cowpeas, tropical
kudzu) also help and have been recommended as long as the area around the
tree is clear. Mangosteen trees must be protected from strong winds and salt
spray.

Harvesting
The fruit are picked when soft and dark purple with the peduncle attached.
Harvesting is by hand or with a pole and basket every 23 days. Mangosteen
harvest indices have been based on the extent and intensity of purple pericarp
development (Table 6.3). The stage at harvest depends on whether the fruit
is intended for the local market or export; export fruit are picked at an earlier
Table 6.3. Mangosteen harvest index stages. Fruit are normally harvested between
stages 14 and eaten from stages 47. The nal total soluble solids gure was
obtained after the fruit was allowed to ripen to stage 5 at 24C (Tongdee, 1985) and
fruit attachment force from the tree (Tongdee and Suwanagul, 1989).
Stage Pericarp color
Detachment
force (kg) Latex
Seed/aril
Yellowish-white
2.2
Severe
Light greenishyellow with

scattered
pink spots
Irregular pink
red
Uniform pink
background
Red to reddishbrown
Reddish-purple
2.09
Severe
1.19
Moderate Difcult
1.24
Slight
1.32
2
3
4
5
6
Dark purple,
slight red
coloration
Dark purple
black
Not
separable
Not
separable
Final
eating
avor
Final total
soluble
solids
Inferior
15.2
Inferior
16
Moderate
Minimum
stage
Export
None
Readily
Export
17.7
1.32
None
Easy
18.3
1.32
None
Easy
Eating
stage
Eating
1.32
None
Easy
Eating
Mangosteen
133
stage. Fruit should not be harvested before the pericarp is a light greenishyellow with distinct irregular pink red spots over the entire fruit (stage 1).
Fruit with less color development have excessive latex exudation at the
peduncle and an inferior avor when they do darken to the full purple stage
in about 5 days. Care is essential to avoid mechanical injury, as a 20 cm fall
causes signicant damage to the aril. Fruit ripening on a tree takes place over
612 weeks. The amount of latex declines with maturity, while total soluble
solids increase and acidity remains relatively constant after stage 1 (Fig. 6.5).
Burst latex vessels on the fruit skin leave dried yellow latex (gamboge) that
may be scraped off.
Acidity (% Citric)
Total Soluble Solids (%)
Fruit are graded to remove damaged fruit and for size and color. There are no
US or international standards. Fruit are usually sold in single-layer berboard
cartons of 2.25 kg with padding, sometimes in trays and individually wrapped
to prevent injury (2024 fruit/carton). In South-east Asia, the fruit are sold
either in baskets or strung in long bundles of 1025 fruit. The thick fruit wall
hardens as the fruit ripens and during storage at low temperatures (<10C).
Current practice is to store at 1214C for a storage life of about 20 days
without chilling injury. Precooling and tray packs are useful. The application of
Maturity Stage
Fig. 6.5. Mangosteen aril acidity and total soluble solids at different maturity stages,
as described in Table 6.3. (After Tongdee, 1985.)
134
Chapter 6
surface coating and storage in polyethylene bags reduces fruit weight loss and
prevents calyx wilting during storage. It is unclear if the effects are due to the
prevention of water loss or to the modied atmospheres.
Marketing
The fruit is regarded as a fruit-y host, which limits its movement in
international trade. Shipments are made by air to Europe, which has no fruity disinfestation requirements. Harvested fruit can be carefully cut open
and the aril inspected before the fruit is frozen whole and shipped to Japan.
Although it has been shown that mangosteen fruit is not a host for oriental
fruit y (Bactrocera dorsalis [Hendel]), vapor heat treatment is needed before
fresh fruit can be exported to Japan.
Fruit damage during harvesting and marketing affects 20% or more fruit.
The cause of gamboge disorder, where latex seeps into the esh giving a bitter
taste, is unknown. The disorder makes separation of the aril and surrounding
tissue difficult even in ripe fruit and leads to pericarp hardening. This should not
be confused with impact injury. Pericarp hardening at the point of impact and
aril collapse, dehydration and pinking or browning are common signs of injury.
A drop of just 10 cm can cause slight pericarp damage, indicated by hardening
at the point of impact. Higher drops cause signicantly greater damage, leading
to downgrading of the fruit.
WORLD PRODUCTION AND UTILIZATION

Most commercial production occurs in South-east Asia (Thailand, Malaysia,
Indonesia), where mangosteen is thought to have originated. Full-grown trees
produce 2002000 fruit/tree. The average yield in Thailand is reported to
be 4.5 t/ha. Older trees (45 years) can yield up to 3000 fruit/tree, but yields
decline in very old trees.
No gures are available for worldwide production. There have been reports
of declining production associated with the costs of the long juvenile stage,
labor for harvesting and the irregular bearing habit. Production is highest in
Thailand, followed by peninsular Malaysia and the Philippines (Mindanao and
the Sulu Archipelago); no data are available from Indonesia. There is some
production in Central America.
The aril makes up to 2030% of the fruit, having 20% dry matter, about
20% total soluble solids and 0.49% acid. The fruit is devoid of vitamin A and
has a low mineral content (Table 6.4). The delicate aril avor is due to hexyl
acetate and cis hex-3-enyl acetate. Hex-3-en-1-ol and six sesquiterpenes have
also been identied. The fruit is normally consumed fresh or in a partially
frozen state. The delicate avor has made attempts to use mangosteen as a juice
Mangosteen
135

portion of ripe mangosteen (Leung and
Flores, 1961; Siong et al., 1988).
Constituent
Edible portion (%)
Proximate
Water (g)
Energy (kcal)
Protein (g)
Fat (g)
Carbohydrate (g)
Fibre (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Thiamine (mg)
Niacin (mg)
Vitamin A (mg)
Vitamin C (mg)
Mangosteen
24
88
34
0.6
1.0
5.6
5.1
0.1
7
4
13
45
7
0.06
0.3
0
4.2
unsuccessful. Preserves are made, although the product darkens and does not
possess a unique avor. Immature fruit can be canned, but mature fruit have
little avor after canning. The pericarp is used to tan leather and dye fabric
black. The rind and bract are used in traditional medicine. The wood is dark
red, coarse and strong, and can be used in carpentry.
FURTHER READING
Achmad, S., Mohamed, Z.A., Teck, C.S., Hamidah, W. and Hussein, W. (1983) Past,
present and suggested future research on mangosteen with example of research
and production in Malaysia. International Workshop for Promoting Research on
Tropical Fruits, Jakarta, 30 May6 June.
Bin Osman, M. and Milan, A.R. (2006) Mangosteen Garcinia mangostana. Southampton
Centre for Underutilised Crops, University of Southampton, Southampton, UK.
Available
from:
http://www.icuc-iwmi.org/les/Publications/Mangosteen_
Monograph.pdf.pdf. Accessed 9 March 2011.
Diczbalis, Y. (2009) Farm and forestry production and marketing prole for mangosteen
(Garcinia mangostana). In: Elevitch, C.R. (ed) Specialty Crops for Pacic Island
136
Chapter 6
Agroforestry. Permanent Agriculture Resources (PAR), Holualoa, Hawaii. Available

from: http://www.agroforestry.net/scps/Mangosteen_specialty_crop.pdf. Accessed
11 March 2011.
Ketsa, S. and Paull, R.E. (2011) Magosteen Garcinia mangostana (L.) In: Yahia, E. (ed.).
Postharvest Biology and Technology of Tropical and Subtropical Fruits: Volume 4.
Woodhead Publishing Limited, Cambridge, pp. 130.
Paull, R.E. and Ketsa, S. (2002) Mangosteen. In: Gross, K.C., Wang, C.Y. and Saltveit, M.
(eds). Agriculture Handbook No. 66. The Commercial Storage of Fruits, Vegetables and
Florist and Nursery Stocks. Produce Quality and Safety Laboratory, USDA, Beltsville,
Maryland. Available from: http://www.ba.ars.usda.gov/hb66/092mangosteen.
pdf. Accessed 9 March 2011.
Salakpetch, S. and Paull, R.E. (2008) Mangosteen Garcinia mangostana L. clusiaceae/
guttiferae. In: Janick, J. and Paull, R.E. (eds) Encyclopedia of Fruit and Nuts. CAB
International, Wallingford, UK, pp. 263267.
Sobir, Poerwanto R. (2007) Mangosteen genetics and improvement. International Journal
of Plant Breeding 1, 105111.
Yaacob, O. and Tindall, H.D. (1995) Mangosteen Cultivation. FAO Plant Production and
Protection Paper. No. 129. Food and Agriculture Organization (FAO), Rome.
REFERENCES
Alexander, D. McE. (1983) Guttiferae. In: Page, P.E. (compiler) Tropical Tree Fruits for
Australia. Queensland Department of Primary Industry, Brisbane, Australia, pp.
6668.
Almeyda, N. and Martin, F.W. (1976) Cultivation of Neglected Tropical Fruits with Promise,
Part 1. Mangosteen. United States Department of Agriculture ARS-S-155.
Augustin, M.A. and Azudi, M.N. (1986) Storage of mangosteen (Garcinia mangostana
L.). ASEAN Food Journal 2, 7880.
Campbell, C.W. (1967) Growing the mangosteen in Southern Florida. Proceedings of the
Florida State Horticulture Society 79, 399400.
Chay, P. (2010) Crop Management of Mangosteen. Queensland Department of Primary
Industry and Fisheries, Queensland, Australia. Available from: http://www.dpi.qld.
gov.au/26_16134.htm. Accessed 10 March 2011.
Germplasm Resources Information Network (GRIN) [Web Database] USDA, ARS,
National Genetic Resources Program, National Germplasm Resources Laboratory,
Beltsville, Maryland. Available from: http://www.ars-grin.gov/cgi-bin/npgs/html/
genus.pl?4842. Accessed 9 March 2011.
Gonzalez, L.G. and Anoos, Q.A. (1951) The growth behavior of mangosteen and its graft
affinity with relatives. The Philippine Agriculturist 35, 379395.
Hume, E.P. (1947) Difficulties in mangosteen culture. Tropical Agriculture 24, 3236.
Hume, E.P. and Cobin, M. (1948) Relation of seed size to germination and early growth
of mangosteen. Proceedings of the American Society for Horticulture Science 48,
289302.
Krishnamurthi, S., Rao, V.N.M. and Ravoof, N.A. (1964) A note on the owers and oral
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biology in mangosteen (Garcinia mangostana L.). South Indian Horticulture 12, 99

101.
Leung, W.T.W., Butrum, R.R. and Chung, F.H. (1972) Food Composition Tables for Use in
East Asia Part 1. US Department of Health, Education and Welfare, Washington,
DC and FAO, Rome.
MacLeod, A.J. and Pieris, W.M. (1982) Volatile avor components of mangosteen
Garcinia mangostana. Phytochemistry 21, 117119.
Marshall, J.R. and Marshall, J. (1983) Mangosteen (Purple). Rare Fruit Council of
Australia, Fact Sheet No. 3, Cairns, Queensland, Australia.
Poerwanto, R., Efendi, D., Widodo, W.D., Susanto, S. and Purwoko, B.S. (2008) Offseason production of tropical fruits. Acta Horticulturae 772, 127133.
Poonnachit, U., Salakpetch, S., Chandraparnik, S. and Hiranpradit, H. (1992) Integrated
Technology to Improve Mangosteen Production. Chanthaburi Horticultural Research
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Popenoe, W. (1928) The Mangosteen in America. Journal of Heredity 19, 537545.
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Ramage, C., Sando, L., Peace, C., Carroll, B. and Drew, R. (2004) Genetic diversity
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Richards, N.J. (1990) Studies in Garcinia, dioecious tropical fruit trees: the origin of the
mangosteen (Garcinia mangostana L.). Botanical Journal of the Linnean Society 103,
301308.
Rukayah, A. and Zabedah, M. (1992) Studies on early growth of mangosteen (Garcinia
mangostana L.). Acta Horticulturae 292, 93100.
Salakpetch, S. (1996) Technology to Improve Mangosteen Production. Orchard
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Salakpetch, S. (2000) Mangosteen production in Thailand. In: Proceedings of the
Tenth Annual International Tropical Fruit Conference, 2022 October 2000,
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7
RAMBUTAN AND PULASAN
The family Sapindaceae is composed of around 150 genera and 2000 species of
trees, shrubs and a few herbs and vines. The species are usually monoecious
and are distributed widely in the warm tropics. Among the numerous genera,
four related genera and ve species are of interest to fruit horticulturists,
with litchi and longan the most widely known. The genera Nephelium has 22
species of which two have been commercialized: rambutan and pulasan.
N. lappaceum L. is generally known as rambutan (Indonesia, Malaysia,
Philippines, English), litchi chevelu (French), ngoh, phruan and ngo (Thailand)
and hong mao dan (China). The three varieties of N. lappaceum L. (vars. lappaceum,
pallens and xanthioides) exist on the basis of variation in leaet characteristics.
N. ramboutan-ake (Labill.) Leenh. (Litchi ramboutan-ake Labill., N. mutabile
Blume) is known as pulasan (Malay, English), ngo-khonsan (Thailand), kapalasan
(Indonesia) and bulala (Tagalog).
RAMBUTAN
Rambutan (N. lappaceum L.) is thought to be indigenous to the Malay
Peninsula, though its long history of domestication makes its origins difficult
to ascertain. Rambutan is now well distributed in South-east Asia, and is
mostly grown in humid, high-rainfall areas. The characteristic environment
of rambutan is one of high rainfall (evenly distributed), high humidity, low
evaporation rates and average minimum temperatures above 20C. The
crop is now grown in number of locations outside of its natural distribution,
including Central America, Sri Lanka, India, New Guinea, tropical Africa,
Hawaii and northern Australia.

139
Chapter 7
140
Ecology
Soil
Rambutan thrives on a wide range of soil types, as long as drainage is good
enough to prevent water-logging and supplementary irrigation is available
during prolonged periods of dry weather. Trees thrive best on deep clay loam
soils and prefer a slightly acid soil (pH 5.06.5). Soil types that support lush
growth may be counterproductive to regular fruit production, particularly in
environments where a check in growth caused by either dry or cool conditions
does not occur.
Climate
High rainfall and humidity induce good growth in
rambutan. Ample annual rainfall is considered to be around 15003000 mm.
The tree is relatively tolerant of drier environments, as long as supplementary
irrigation is available.
Rambutan owering correlates with the end of the dry season, leading to a
second smaller crop each year on branches that did not bear fruit in the previous
fruiting. A dry period of at least 1 month is essential to initiate rambutan
owering (Fig. 7.1). The intensity of owering appears to be closely associated
Flushing or Flowering (% of Canopy)
RAINFALL AND MOISTURE
Fig. 7.1. Mean rainfall, vegetative ushing and owering pattern for eight cultivars
of rambutan over 4 years at Hilo, Hawaii, which has no marked dry season.
Vegetative ushing parallels rainfall, with owering occurring following a period of
lower rainfall. (Redrawn from Kawabata et al., 2007.)
Rambutan and Pulasan
141
to the duration of water stress. Rambutan requires adequate moisture, from

either rainfall or irrigation, during fruit set and growth (Fig. 7.2). There are
numerous stomata on the fruit spinterns and low humidity during fruit
development can lead to fruit desiccation.
Rambutan has no cold requirement for owering and is suited
to tropical areas with a temperature range of 2230C. Temperatures less than
22C reduce leaf ushing and can extend fruit development to nearly 6 months.
TEMPERATURE
There is little information on the

response of rambutan to varying light levels. It is thought to be day-neutral.
LIGHT INTENSITY AND PHOTOPERIOD
WIND Windbreaks are necessary as young and mature trees buffeted by the
wind establish, grow and yield poorly. Common windbreak species in Australia
are found in the genera Acacia, Artocarpus, Casuarina, Melaleuca, Pennisetum
and Syzygium.
Tree
The tree grows to 25 m in height and has a straight trunk of up to 60 cm wide,
with a dense, usually spreading crown. The evergreen alternate leaves are 40
cm long, pinnately compound and divided into two to four pairs of leaets,
usually alternate elliptic to oblongelliptic or rather obovate (Fig. 7.3). Leaets
are 7.623 cm u 3.89.0 cm. They are pale when young, becoming medium
to dark green in color.
Flowers
Rambutan owers are small, less than 6 mm wide, greenish and apetalous.
They are produced in panicles (Fig. 7.3) that are axillary near the branch
apex (Allen, 1967). Trees are normally classied into three types based on
ower type: male trees that produce only staminate owers (4060% of a
seedling population); trees with functionally female hermaphrodite owers;
and trees with functionally female and functionally male hermaphrodite
owers (Fig. 7.3). Male trees can have 3000 male owers on a panicle, with
approximately 500 open on one day. Hermaphrodite trees are classied as
female and produce two types of owers, with about 500 owers per panicle
and 100 opening per day. Male functional owers have well-developed
stamens and pistils that fail to function normally; the bid stigmas split
but remain erect, preventing the exposure of the sticky stigmatic section
required for pollination. Female functional owers have well-developed pistils
with non-functional stamens, reduced to ve to seven staminodes. The bid
stigmas split open and curve downward to expose the sticky stigmatic surface
Temperature >22C
Low Water, High N
Temperature >22C
Low Water, High N
Temperature >22C
High Water
Flowering
Floral Induction
and Development
Temperature >22C
Low Water, High N
Temperature >22C
High Water
Temperature >22C
Water Stress
Floral Induction
and Development
Prune
Fertilize
Irrigate
No Flowering
Temperature <22C
Fertilize
Irrigate
Temperature >22C
High Water, High N
142
Temperature >22C
High Water, High N
No Irrigation
Induce Stress
First Flowers
Remove Plastic
and Irrigate
Fertilize
Fruit Set
and Growth
Irrigate
Chapter 7
Fig. 7.2. The fruiting cycle of rambutan, as affected by fertilization and water availability, under normal conditions (A) and when water
stress is used to induce owering (B). Soil surface covers and ditches are used to take away rainfall (C). The plastic cover is removed
when tree owering occurs (D).
143
Fig. 7.3. Rambutan leaves (A), panicle (B), hermaphrodite ower (C), male ower
(D) and fruit bunch (E) showing the spintern on the pericarp, and one fruit with half
the pericarp removed to expose the aril that surrounds the seed.
and remain receptive for 1 day. Nectar production for both ower types begins
at anthesis.
Hermaphrodite trees with both functional male and female owers are more
desirable and are most commonly found in some rambutan cultivar selections,
with male owers in the range 0.50.9% (Lam and Kosiyachinda, 1987).
Flower induction, pollination and fruit set
Rambutan is not believed to have a cold requirement for owering and is
suited to tropical areas with a temperature range of 2230C. In Australia,
owering in the dry tropics (Darwin, 12.5S) usually follows the onset of cool
nights (1812C) in July to August (Diczbalis et al., 1996). In the wet tropics,
however, owering is reported to occur throughout the year regardless of
the climate (Watson, 1988) but usually occurs from September to October
following a short dry season. In Thailand, a drop in night temperatures of
23C with the onset of the dry season has been suggested as the prompt for
owering. Multiple regression analysis showed that for every 1C decrease in
144
Chapter 7
night temperature, ower induction increased by 6.7% (Manakasem, 1995).

Temperatures lower than 22C reduce leaf ushing and can extend fruit
development to nearly 6 months.
Management practices in Thailand are directed toward producing a large crop
in the May to June period. Cultivars vary in their requirements for induction,
leading to early- and late-season bearers. The cultivars Jitlee and R137 are
more suited to areas with some water stress than R4 or R7. Foliar-applied
paclobutrazol does not produce off-season owering. Ringing and covering a
narrow area of the bark (<2 cm), then spraying with potassium nitrate (3040
g/l) 1 month after ringing brings about earlier owering of mature trees. The
ringing treatment changes the translocation of photosynthetate to the roots and
increases starch in the shoot. Ringing can only be performed in alternate years.
Because rambutan owering is correlated with the end of the dry season,
farmers in Vietnam cover the soil with plastic sheets with ditches at the edge
of the canopy to carry away any rainfall (Fig. 7.2). A dry period of at least
1 month is needed to initiate rambutan owering, with periods of up to 3.5
months being used. The soil covers are removed when the rst owers appear
on the trees. The overall system used involves harvesting the previous crop,
pruning the tree and fertilizing to induce vegetative growth, and covering the
soil and leaving the soil cover on until the rst owers appear. The trees are then
irrigated on removal of the soil covers, with a second application of fertilizer
at fruit set. The intensity of owering is associated with the duration of water
stress, and harvesting can be scheduled for every 6 months. This protocol is
used to induce rambutan owering even during the wet season.
Only 12% of owers form fruit on a rambutan panicle. Since no pollen is
apparent on the hermaphrodite ower anthers, pollen transfer or apomictic
fruit development may occur. In mixed plantings with no male trees, pollination
problems are not seen if owering periods overlap. The cultivar Seechompoo
has low numbers of functional male hermaphrodite owers and is notorious for
poor fruit set in a pure stand.
Fruit
Usually only one of the two locules in the rambutan ovary develops to
form a normal fruit; the other aborts and remains at the base of the fruit.
Occasionally, however, both carpels develop equally to form two normal fruit.
The fruit occur in clusters of 1018 and are large (2545 g in weight), ovoid
or globose, and about 4.5 cm long and 2.53.7 cm broad. The outer skin on
the pericarp is 24 mm thick and covered with long soft spines (spinterns)
(Fig. 7.3) that turn red or yellow when the fruit is ripe.
Early rambutan fruit growth (Fig. 7.4) is dominated by skin growth; the aril
is attached to the single large seed and does not begin to grow until around
60 days after anthesis. In some rambutan cultivars the hard testa comes away
with the aril, making these cultivars less desirable. The major phase of growth
145
Mass (g)
Color
Break

Fig. 7.4. The growth pattern of rambutan cultivar Seechompoo fruit and its parts
from anthesis. (Redrawn from Wanichkul, 1980.)
occurs 5080 days after anthesis. As with litchi, there is a rapid decline in
titratable acidity from more than 1% to less than 0.5% and an increase in total
soluble solids from about 13% to 20% in the last 35 days of fruit growth. Total
starch declines during the same period and fruit begin to change color.
The diploid number for rambutan is 2n = 22.
Breeding and selection
Breeding and crop improvement attempts appear to be limited or poorly
reported. A large-scale evaluation of F1 hybrids based on two maternal parents
(R99 and R134) and 14 popular Malaysian cultivars was undertaken in
Malaysia (Sarip et al., 1996). Six years after its establishment, 50% of the
population had owered and about 40% were males. Seven percent of the
population owered 2 weeks earlier than both maternal parents and less
than 1% produced high-quality fruit with the combined attributes of good
appearance, high recovery and cling-free. This suggests that improved cultivars
will continue to come from grower selection and potentially via biotechnology.
Chapter 7
146
Table 7.1. Fruit characteristics of selected rambutan cultivars.

Edible
content
(%)
Cultivar
Synonyms
Vigor
Fruit
weight
(g)
Jit Lee
Deli
Medium
3055
35
Binjai
Medium/
small
Medium
3241
41
2233
46
4051
36
4045
49
4050
41
2835
40
R3
Peng ting
ching
R9
Tau po cheng Medium/
large
R137
Gajah mati
Medium/
large
Rongrien Roengrean
Medium/
large
Chompoo Seechompoo Dwarf
TSS
(%)
Skin color
2022 Orange/red,
green tips
1821 Orange/red,
green tips
22.5 Red
2023 Pink/crimson
red
2223 Red
1821 Dark red,
green tips
1820 Orange
The selection criteria for rambutan are similar to those of litchi. Adherence
of the aril to the testa is an important criterion. The criteria include:
x Fruit: large, with small seed and a high proportion of edible aril. Bright red or
yellow skin color, long shelf-life and ability to retain skin color under storage
conditions. Firm esh with an acceptable sugar-to-acid ratio and resistance
to disease.
x Tree: vigorous, precocious, regular and high yielding. Resistant to water
stress, wind, soil salinity, diseases and insects.
Numerous rambutan cultivars (Salma, 1986) with a wide range of
characteristics (Table 7.1) are grown in major production areas. Salma (1986)
identied and developed a key to 31 cultivars of rambutan grown in Malaysia
out of an accession list that exceeds 65. A large number of accessions have
been collected in major centers in Australia, Hawaii, Indonesia, Malaysia,
Mexico, the Philippines, the Seychelles and Thailand (IBPGR, 1986).
Cultural practices
Propagation
Valmayor et al. (1971) reported that planting from seed results in a high
proportion of male-owering trees. Seedling trees are less precocious and
extremely variable in character, and hence commercial production is based on
clonally propagated trees. Vegetative propagation is used to propagate selected
147
female-owering trees. The male tree is seldom found as orchards are usually
based on clonal budwood. In rare cases where wood below the bud graft has
been allowed to develop, shoots with male owers are clearly seen.
SEXUAL The seeds are recalcitrated and rapidly lose viability after removal
from the fruit. Seeds germinate in 715 days.
ASEXUAL Little success has been reported with rooting rambutan cuttings.
Grafting is difficult and a number of methods have been tried (e.g. shield,
patch and modied Forkert budding; various approach graft techniques and
cleft grafting). Modied Forkert or patch budding appears to be the preferred
technique of commercial propagators worldwide. Approach grafting, although
much more successful, is cumbersome and time consuming, and reliant on
scaffolding surrounding the parent tree. Research and experience suggest that
budding or grafting operations should occur during the active growing period
(the rainy season). Despite the availability of a range of budding and grafting
techniques, the overall success rate is generally low. Therefore, air layering is
also utilized.
In air layering young, actively growing shoots are girdled and the wound
surrounded with peat or similar moisture-absorbing material until roots
emerge. The rooted shoots are removed from the parent tree and established
in a pot or directly into the eld. This technique is growing in popularity in
Australia because of the shortage and expense of budded trees. The long-term
survival of air-layered trees has been questioned.
The selection of rootstocks is generally based on seed availability. Little
research has been conducted on stock scion relationships. In the Philippines,
N. intermediam (bulaba) has been shown to be a successful stock for grafted
rambutan (Valmayor et al., 1961). A trial was established in Australias
Northern Territory to evaluate the production of two cultivars (Jitlee and
R134) on eight stocks (Jitlee, R156, Rapiah, Rongrien, Gula Batu,
R9, R37 and Bogor). To date, no commercial differences in growth and
yield performance have been detected (McMahon, personal communication,
2004). Field observations have noted that certain stock/scion combinations
are semi-dwarfed. There is a need to explore whether this is due to those specic
combinations or chance.
Field preparation
Deep ripping may be necessary if the soil is compacted. Liming to pH 5.05.5
should be carried out and manure incorporated before planting. Rambutan
require well-drained alluvial soils for good development. In South-east Asia,
rambutan is frequently interplanted with other tree crops such as durian or
duku/langsat. Monoculture orchards are the preferred method of establishment
in Hawaii, Australia, Central America and, increasingly, South-east Asia.
Chapter 7
148

Budded or grafted trees are generally ready for eld planting within 1224
months. Air-layered trees are generally removed from the tree with 34
months of girdling and established in pots (36 months) prior to eld
planting. Growers in Australia are nding success with the direct eld
establishment of air-layered trees. Rambutan are planted with 510 m
between trees and 810 m between rows. One male rambutan tree is often
planted per 810 functional female hermaphrodite trees. Windbreaks are
necessary for young and mature trees.
Irrigation
Irrigation is required for rambutan grown for high-value domestic and export
markets, as water stress during ower and fruit development reduces yields
and fruit size. Irrigation is also essential during the vegetative ushing stage
after harvest, and is withheld during oral initiation (Fig. 7.2). Preowering
water stress, which does not induce leaf wilting, leads to earlier owering
and improved harvest synchrony without affecting yield. High rainfall
during blooming can lead to poor fruit set. In the monsoonal environment of
northern Australia, irrigated rambutan trees have a shallow root system with
80% of the roots in the top 15 cm and not extending beyond the tree canopy.
Such trees can require irrigation up to three times a week (Diczbalis et al.,
1996). In trees grown on sandy soils in hot and dry growing environments,
severe leaf loss can occur within 410 days of withholding irrigation. The
amount of irrigation should, at a minimum, replace that lost by evaporation,
particularly from owering to harvest (Table 7.2).
Table 7.2. Crop coefcient (Kc) for rambutan to estimate

the daily water requirement (WR) for different stages of
development based on evapotranspiration (ET). WR =
Kc u ET (Hiranpradit et al., 1998). Higher Kc values are
reported during fruit set and fruit development in the wet/
dry tropics of Australia (Diczbalis et al., 1996).
Vegetative growth
Floral initiation
Floral development
Fruit setting
Early fruit growth
Late fruit growth
Fruit maturity
Kc
0.6
0.00/0.60
0.75
0.75
0.8
0.85
0.85
149
Pruning
Little research has been published on the need for and effectiveness of
pruning. Early training aims to achieve a strong framework of branches to
encourage secondary lateral branches. Annual pruning is designed to remove
water sprouts, pest- and disease-infected shoots, dead branches and crossing
branches. In some areas the panicle is pruned after harvest to induce vigorous
canopy regrowth. The effect of pruning on rambutan cropping in the wet/
dry tropics of Australia has been evaluated (Menzel et al., 2000). Pruning
delays harvest and yields are 2040% lower than in non-pruned trees, but
on much smaller canopies. The reduction in cropping is greatest when shoottipping in June is followed by structural pruning in February. The trial clearly
demonstrated that in the wet/dry tropical environment of northern Australia,
structural pruning can occur without a complete loss of production the
following season. Structural pruning and structural plus shoot-tip pruning
treatments both delayed owering and harvest dates relative to the control
treatments. Pruning treatments also improved the synchronization of harvest.
The ability to manipulate harvest dates and synchronization are potentially
important management tools.
Commercial growers in Australia are currently using mechanical pruners
to reduce tree size and shape trees to allow machinery access. The effect of
mechanical pruning on tree productivity depends on the time of pruning and
the amount of wood removed. In situations where relatively heavy pruning has
taken place, owering may be delayed until the following season.
Fertilization
The fertilization practices of commercial rambutan orchards differ because
of differences in climate, soil, availability of different kinds of organic and
inorganic fertilizers and other factors.
Rambutan is reported to have similar nutritional requirements to litchi.
A hectare of rambutan (7080 trees/ha) yielding 6750 kg/ha in north
Queensland removed 13.6 kg nitrogen, 2.1 kg phosphorus, 12.1 kg potassium,
3.7 kg calcium, 1.9 kg magnesium and 1.3 kg sulfur (Diczbalis, 2002). Fertilizer
management in rambutan can be enhanced by the use of soil and leaf analysis
(early panicle emergence), with nutrient replacement based on nutrient removal
plus losses due to leaching, runoff or volatilization. Fertilizer application needs
to correspond with differing needs at various stages of the growth cycle. Critical
periods are before owering and fruit set, several weeks after fruit set and after
harvest. Leaf samples for analysis are taken twice a year, once at the end of
harvest and again at owering, with only phosphorous and potassium applied
during fruit set to avoid vegetative growth from nitrogen application. Tentative
leaf nutrient standards (north Queensland) at early panicle emergence are:
2.01% nitrogen, 0.21% phosphorous, 0.66% potassium, 1.2% calcium, 0.32%
150
Chapter 7
magnesium, 0.21% sulfur, 485 mg/kg manganese, 102 mg/kg iron, 54 mg/
kg copper, 26 mg/kg zinc and 51 mg/kg boron (Diczbalis and Alvero, 2005).
Minor elements are applied once a year at the end of the cold period.
Erratic bearing
In shoots that have not fruited, vegetative shoots arise from the terminal buds
and 57% ower and fruit in the following season. Poor synchronization of
ushing and owering within an orchard increases management problems.
Cultivars vary in their requirements for induction, leading to early- and lateseason bearers.
Pest management
DISEASES The major diseases of rambutan are powdery mildew on young
growth, stem canker, pink disease and sooty mold. The importance of these
diseases varies according to location. The ascomycete Dolabra nepheliae is
thought, in association with various insect larvae, to be the cause of stem canker.
Algal leaf spots caused by Cephaleuros virescens can be problematic in mature
orchards. Postharvest diseases are similar to those of litchi and include various
fruit rots (Colletotrichum gloeosporioides, Phomopsis spp., Dothiorella spp.).
A number of insects and arachnids infest rambutan. A total of 48
insect and mite pests have been recorded on rambutan in Australia (Astridge,
2003), with similar numbers reported in other production areas. Approximately
16 species are considered to be major pests of leaves, stems or fruit. None are
regarded as serious, and most are controlled with minimal spraying programs.
The insects reported include loopers (Achaea janata, Oxyodes tricolor), swarming
beetles (Monolepta australis, Rhyparida spp.), fruit-sucking moths (Eudocima spp.),
ower/fruit caterpillars (Conogethes punctiferalis), plant hoppers (Colgaroides
acuminata), stink bugs (Lyramorpha spp., Plautia affinis), fruit-spotting bug
(Amblypelta lutescens), weevils (Myllocerus spp.), mites (Brevipalpus spp.), thrips
(Selenothrips rubrocinctus) and mealy bugs. Scale insects (Ceroplastes rubens,
Coccus viridis, Icerya spp.) and mealy bugs (Planococcus citri) can be a problem,
especially if they infest the fruit and have to be removed for export. Rambutan
are considered to be a poor or non-host for fruit ies. The Mediterranean fruit y
(Ceratitis capitata) does not attack this fruit, except where the fruit skin has been
broken by other means and the pulp is exposed. The Oriental fruit y (Dacus
dorsalis) does infest ripe fruit, causing punctures that are often the focus of
entry for fungal organisms that cause fermentation and decomposition. Other
fruit ies can be serious pests in particular countries.
INSECTS
OTHER PESTS Other pests include birds and fruit bats. In Hawaii, the white
eye (Zosterops palpebrosus) and the bulbul (Pycnonotus cafer) can cause severe
damage as fruit approach maturity, while the rainbow lorikeet (Trichoglossus
151
haematodus) is a problem in Australia. Fruit bats (Pteropus spp.) and ying foxes
have caused severe losses in South Africa and Australia.
Weed management
Weed control is most important from the time of eld transplanting up to
3 or 4 years old. As trees grow and expand horizontally, shading results in
a decreasing amount of weed growth underneath the canopy. The use of
polyethylene mulch about a meter square around the plant at transplant time
reduces weed growth near the tree. Organic mulches are highly recommended
around the base of rambutan trees.

Harvesting
There are numerous stomata on the fruit spinterns, and low humidity during
fruit development can lead to fruit desiccation. Trees require adequate
moisture, from rainfall or irrigation, during fruit set and growth.
Grade standards are generally set between the shipper and buyer. Small, poorly
colored, immature or damaged fruit are culled. Rambutan spinterns (hairs)
are easily damaged and care is essential to avoid mechanical injury (OHare,
1995). Soluble solids should be greater than 15% and acid levels low.
Fruit packaging varies by country, and fruit destined for local markets in
Asian growing regions is generally sold on the stalk. Fruit destined for local
supermarkets or export is generally wrapped in plastic lm to avoid water loss
and packed in cardboard boxes weighing 2.311.0 kg. Packaging sizes reaching
UK markets are generally 2.0 kg in weight, with 4.0 kg packs arriving from
Thailand (Anon, 1999). For longer-term storage (2 weeks) the fruit is best kept
at high humidity (>95% relative humidity) and low temperature (1015C).
Fruit will brown rapidly once exposed to cool dry conditions such as experienced
in supermarket displays, despite careful attention to harvesting and postharvest
storage. Hence, exporters should strongly consider the use of retail-ready
packaging such as bags or punnets (clam shells) in sizes that suit the intended
markets.
Utilization
World production gures are difficult to obtain, given that the major and
smaller centers of production do not regularly update production data.
Estimates based on data available from the mid 1990s suggest that total
world production is in the vicinity of 1.2 million t (Vinning and Moody,
152
Chapter 7
1997). Recent production statistics suggest that the world area under
rambutan exceeds 200,000 ha and production is approximately 1.52.0
million t/year (Campbell and Ledesma, 2003; Crane et al., 2003; Huang et
al., 2003; Poerwanto, 2003; Salakpetch, 2003). Thailand, Indonesia and
Malaysia account for approximately 80% of the total world production. The
amount traded internationally is substantially less. UK fruit markets record
receivables of fresh product from Honduras, Indonesia, Malaysia, Panama,
the Philippines, Sri Lanka and Thailand. The trade of rambutan is somewhat
limited by quarantine regulations, with a number of potential importing
countries imposing strict disinfestation requirements.
Central American producers such as Costa Rica, Guatemala, Honduras
and Mexico supply local and mainland USA markets. Australia is capable of
producing up to 1000 t of rambutan in favorable years, with approximately
100 t exported to Japan. Hawaii produces approximately 120 t, with most
being consumed locally and some exported to the mainland USA and Canada
after irradiation.
Rambutan is primarily consumed as a fresh fruit, but is also frozen, juiced,
canned and dried in limited quantities (Anon, 1979). The proximate fruit
composition shows high levels of potassium and some vitamins (Table 7.3).
PULASAN
Botany
The pulasan or poolasan, N. mutabile Blume (Sapindaceae), is sometimes
confused with its close relative rambutan, which it closely resembles. The
common names include ngo-khonsan (Thailand), rambutan-kafri and kapalasan
(Indonesia), meritam (Sabah, Sarawak), bulala (Tagalog) and rambutan-kafri
and rambutan paroh (Malaysia). The Dutch name in Java is kapoelasan. The
species is native to the region from Burma to the Malay Peninsula, and also
occurs in India (Assam), Burma, Indonesia, Malaysia and the Philippines.
Wild trees are infrequent in lowland forests around Perak, Malaya, but
abundant in the Philippines at low elevations from Luzon to Mindanao. It is
grown to a limited extent in northern Australia and Hawaii. Although the
fruit is considered superior to rambutan, it is not as commonly seen in the
market. The lack of proven varieties, suitable production and postharvest
handling practices, and consumer awareness of pulasans qualities contribute
to its limited production.
Ecology
Pulasan is a lowland primary forest tree. It grows well at below 350 m in the
tropics, with annual rainfall of 15003000 mm. Well-drained rich soil that
153
Table 7.3. Composition of 100 g edible portion of rambutan and

pulasan (Nga, 1980; Ortiz and Cordero, 1984; Wills et al., 1986;
Morton, 1987; Wenkam, 1990).
Constituent
Edible portion (%)
Proximate
Water (g)
Energy (kJ)
Protein (g)
Lipid (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Magnesium (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
Rambutan
43
83
297
0.8
0.45
14.5
4
0.3
25
3
10
13
140
20
48
Trace
0.065
0.8
0
Pulasan
3545
8491
0.82
0.55
12.9
0.14
0.43
0.010.05
0.002
is high in organic matter is preferred, and the tree does not do well on sandy
soil.
Flowering is more common following a longer dry season than required
for rambutan. In many ways pulasan is similar to rambutan in its climatic
requirements, although it is less tolerant of high light during establishment. It
is often found on riverbanks, but rarely in swamps.
Tree
Pulasan trees grow to 1015 m in height. The alternate leaves are pinnate
or odd-pinnate and 1745 cm long. There are two to ve pairs of opposite or
nearly opposite leaets, which are oblong- or elliptic-lanceolate, 6.2517.5
cm long, and up to 5 cm wide (Fig. 7.5). The leaves are slightly wavy, dark
154
Chapter 7
green and barely glossy on the upper surface; they are pale and somewhat
bluish with a few short, silky hairs on the underside.
Flowers
The very small, greenish, petalless owers have four to ve hairy sepals and are
borne singly or in clusters on the branches of the erect, axillary or terminal
panicles. Male owers possess ve to eight well-developed stamens with
dehiscent anthers that shed viable pollen (Fig. 7.5). The hermaphroditic female
owers possess ve to eight stamens attached to the base of the ovary and do
not extend beyond the stigma (Fig. 7.5). The anthers are indehiscent. The pistil
consists of a bi-lobed ovary with a single style, topped with a bifurcated stigma.
Reproduction
Seedling trees can be grouped into male trees that produce only male
owers and do not produce fruit, and female trees that produce functionally
female hermaphroditic owers. Pollen germination tests indicate that the
male owers produce viable pollen. Some hermaphroditic female trees may
also produce functional male owers since lone Seebabat trees have been
reported to set fruit on their own. Trees ower during June and August, and
fruit mature during October to December in Malaysia and April to May in
Fig. 7.5. Pulasan trees, owers and fruit clusters are very similar to those of
rambutan. Male inorescence (A) and hermaphrodite owers (B). The fruit clusters
(C) are tighter than those of rambutan and the fruit have short, thick, stubby spines
or tubercles on the pericarp rather than spinterns. (From Encyclopedia of Fruits and
Nuts 2008, Plate 78, courtesy of Dr. Mike Nagao.)
155
Thailand. Flowering can occur twice a year (February to March and August to
October) in locations such as Hawaii. Mature fruit can be harvested in spring
or fall, roughly coinciding with the rambutan owering seasons.
Fruit
Fruit are borne in clusters (Fig. 7.5). Although two carpels are present on
each ower at anthesis, only one usually develops while the other aborts but
remains attached to the base of the developing fruit. The fruit is 46 cm wide
and 47 cm long, with a dark red or occasionally yellow, thick, leathery skin
when ripe, covered with short, thick, stubby spines or tubercles. Fruit weigh
between 5080 g, and the edible pulp is 2530% of the total fruit weight. The
aril is white, juicy and sweet with 1625% total soluble solids; it is not always
attached to the seed testa. The seed is ovoid, oblong or ellipsoid, light brown
in color and somewhat attened. It is 23.5 cm long and weighs 2.02.5 g.
Underdeveloped seeds are occasionally found in some fruit, which will also
have low esh weights.
The single-seeded fruit exhibit a simple sigmoid growth pattern and mature
at about 1518 weeks after anthesis in the warm tropics. This fruit maturation
can be extended by up to 6 weeks in cooler locations. As fruit mature, heavy
and inconsistent rainfall can cause the pericarps to crack and result in loss of
production. The development of at fruit is often observed. These fruit, which
are normal in length, fail to develop fully and have a attened appearance. These
fruit are seedless and do not possess a well-developed aril. Lack of pollination
may be responsible for this abnormal fruit development.
Two forms of pulasan are reported from Java: Seebabat and Kapoolasan
seebabat. The fruit are mostly dark red and the tubercles are crowded
together. The esh of Seebabat is very sweet and juicy, and separates easily
from the seed. In the other group, the fruit is light red and smaller, and the
tubercles are not so closely set. The esh adheres rmly to the seed.
Cultural practices
Propagation
Propagation by seed is not favored as the seedlings may be male or female.
Seeds are sown fresh and germinate rapidly. Air layers are easy to propagate,
but are thought to be generally very short-lived. Budding grafting is difficult
and has a low success rate. Successful budding has been reported if done in
the rainy season on rootstocks already set out in the eld. Superior rootstocks
have not been identied and bud grafting is done on seedling rootstocks.
156
Chapter 7
Planting distances of 810 m are recommended. Grafted trees begin to fruit in

35 years. Trees often receive little or no fertilizer or other cultural attention.
Flowering occurs from axillary and terminal buds, and regular pruning after
harvesting is necessary to maintain tree size. Trees are highly branched as
for other sapindaceous fruits, and early pruning and training are necessary
to develop a strong scaffold of branches that are not subject to breakage. An
open-center system is often used.
Nutrition and fertilization
Information on the nutritional requirements for pulasan has not been
developed. However, based on the similar phenology and growth pattern to
rambutan, fertilizer practices used in rambutan production will probably
apply to pulasan trees. In the Philippines, a 15:15:6 mix with magnesium is
recommended for non-bearing trees and 12:12:12 for bearing trees.
Diseases, pests and weeds
Diseases and pests have not been widely reported. Powdery mildew can occur
on the leaves and sooty mold on the fruit. Leaf rollers, fruit borers and weevils
also cause damage, and mite infestations have been observed in rootstocks
grown in the greenhouse.

The fruit bunch is harvested when most fruit are dark red or yellow in color.
The fruit bunches are accumulated in baskets before sorting. Postharvest
handling is similar to that for rambutan.

Yields of 50006000 fruit/tree are reported for mature trees. Records in the
Philippines indicate mature trees can produce 180 kg of fruit.
The fruit is consumed fresh and has a similar nutrient composition to that
of rambutan (Table 7.3). The fruit can be frozen and dried or used in juices.
The frozen and dried product can be used in ice cream, desserts and preserves.
Boiled or roasted seeds are used to prepare a cocoa-like beverage.
Hydrocyanic acid has been detected in the bark and leaves. The volatiles
are mainly aliphatic hydrocarbons (70%), of which pentadecane (61%) and
aliphatic alcohols (19%) are the most dominant. The dried seed kernels yield
74.9% of a solid white fat, which melts at 4042C to a faintly perfumed oil that
has a possible use in soap-making. The wood is light red and harder and heavier
157
than that of rambutan. It is of excellent quality but is rarely available. The

leaves and roots are employed in poultices. The root decoction is administered
as a febrifuge and vermifuge. The roots are boiled with Gleichenia linearis Clarke,
and the decoction is used for bathing fever patients.
FURTHER READING
Diczbalis, Y.A. and Paull, R.E. (2008) Rambutan Nephelium lappacium L., Sapindaceae.
In: Janick, J. and Paull, R.E. (eds) Encyclopedia of Fruit and Nuts. CAB International,
Lam, P.F. and Kosiyachinda, S. (eds) (1987) Rambutan: Fruit Development, Postharvest
Physiology and Marketing in ASEAN. ASEAN Food Handling Bureau, Kuala Lumpur,
Malaysia.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resource Systems, Winterville,
rambutan.html. Accessed 2 March 2011.
Nagao, M.A. and Paull, R.E. (2008) Pulasan Nephelium mutabile Bl. Sapindaceae. In:
Janick, J. and Paull, R.E. (eds) Encyclopedia of Fruit and Nuts. CAB International,
Salakpetch, S. (2003) Rambutan production in Thailand. Acta Horticulturae 665, 67
72.
Tindall, H.D. (1994) Rambutan Cultivation. UN-FAO Plant Production and Protection
Paper #121. Rome.
Wall, M., Sivakumar, D. and Korsten, L. (2011) Rambutan (Nephelium lappaceum L.). In:
Yahia, E. (ed.) Postharvest Biology and Technology of Tropical and Subtropical Fruits:
Volume 4. Woodhead Publishing Limited, Cambridge, pp. 312333.
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8
PASSION FRUIT AND
GIANT PASSION FRUIT
INTRODUCTION
The passion ower family (Passioraceae) includes about 18 genera with around
530 species of dicotyledonous, herbaceous or woody vines, usually with
axillary tendrils. Erect shrubs and trees are rare in the family. The species are
native to the tropical and subtropical regions of both hemispheres, growing
at medium to high elevations where temperatures are moderate. Only two
genera, Passiora and Tetrapathaea, are cultivated. The most important genus is
Passiora, the species of which are mostly vines with axillary tendrils; the fruit
is a many-seeded berry. The owers of many species are conspicuous in their
form and color and are grown for their ornamental value.
Only 5060 Passiora species bear edible fruit, and most are unknown
outside their area or origin or cultivation by the native people (Table 8.1). A
number of species are considered commercial, and the fruit of P. quadrangularis,
P. ligularis, P. laurifolia and P. mollissima are often found in village markets in
Latin America. However, only the purple passion ower fruit (P. edulis Sims),
the yellow passion ower fruit (P. edulis f. avicarpa Deg.) and hybrids between
the two are considered to be of value in international commerce (Table 8.2).
This chapter discusses P. edulis, P. edulis f. avicarpa and P. quadrangularis L.
Commercially, the former two are referred to as passion fruit (the name used
in this chapter) and the latter as giant passion fruit.
The common names, besides purple or yellow passion fruit, include granadilla
(English), grenadille and couzou (French), markisa and buah susu (Malaysia),
linmangkon (Bangkok, Thailand), benchawan (Chiang Mai, Thailand), limangkan
(Laos), maracuy morado (purple) and maracuy amarillo (yellow) (Spanish),
gulupa (purple in Colombia), maracuya peroba (Portuguese), mara-cuia and
maracuja (Brazil), fruta de pasion and pasionaria (Tagalog, Philippines), parcha
and marora (Ilokano, Philippines), parcha, parchita and parchita maracuy
(Venezuela), chinola (Puerto Rico), lilikoi and yellow lilikoi (Hawaii) and
mountain sweet cup (Jamaica).
161
Chapter 8
162
Table 8.1. Selected Passiora species, their areas of origin and use (Martin and
Nakasone, 1970).
Use
Common name(s)
Species
Area(s) of origin
Banana passion fruit
P. alata Dryand
P. antioquiensis
Karst
P. banksii Benth
P. caerulea
P. coccinea Aubl.
Blue passion ower

Red granadilla
Purple granadilla
Wild passion ower,
maypop, apricot
vine
Yellow granadilla,
water lemon,
Jamaica
honeysuckle
Sweet granadilla
Sweet calabash,
sweet cup, conch
apple
Banana passion fruit,
curuba
Giant granadilla
Ornamental
Fruit
Brazil, Peru
Colombia
X
X
Australia
Brazil to Argentina
Venezuela to
Bolivia
Brazil
Southern USA
X
X
X
X
X
P. laurifolia L.
West Indies to
Brazil and Peru
P. ligularis Juss.
P. maliformis L.
Mexico to Bolivia
West Indies to
South America
X
X
P. mollissima
(HBK) Bailey
P. quadrangularis
L.
P. vitifolia HBK
Venezuela to
Bolivia
Unknown
P. edulis Sims.
P. incarnata L.
Nicaragua to
Venezuela and
Peru
X
X
The giant passion fruit is also known in Spanish as granadilla gigante and
granada (Central America), badea and corvejo (Colombia and parts of Venezuela),
tumbo costeo (Peru), granadilla de fresco (El Salvador), granadilla real and
sanda de passion (Bolivia), parcha granadina and parcha de Guinea (Venezuela),
maracuja-assu, maracuya-acu, maracuja mamao and maracuja grande (Brazil),
groote markoesa (Surinam), parola and kasaora (Philippines), markiza, markesa
and markesa (Indonesia), timun belanda, marquesa and mentimum (Malay) and
barbadine (France, Vietnam).
Passion Fruit and Giant Passion Fruit
163
Table 8.2. Similarities and differences between Passiora edulis Sims and P. edulis
f. avicarpa Deg. (Knight, 1980).
Characteristic
P. edulis Sims
P. edulis f. avicarpa
Ecology
Cooler elevation
Low elevation
Vine
Less vigorous
Vigorous
Leaves
Similar shape, smaller
Larger
Flowers
Fruit
Smaller, fragrant, less

protandry, anthesis in
the morning
Purple, small
Larger, stronger fragrance,

stronger protandry, anthesis
in the afternoon
Yellow, large
Juice
Mild acid
Acid
Chromosome number
2n = 18
2n = 18
Meiosis
Normal
Normal
Ovules, pollen grains
Fully viable
Fully viable
Compatibility
Self-compatible
Self-incompatible

The well-known purple passion fruit is considered native to southern Brazil,
and was widely distributed to other countries of South America and the
Caribbean and into Asia, Africa, India and Australia during the 19th century.
It is grown commercially mainly in Colombia, Ecuador, Kenya, South Africa,
the Philippines, Australia and New Zealand. The fruit is sold locally and
exported in its fresh form and as processed pulp.
The origin of the yellow form, P. edulis f. avicarpa, is unknown. It may
have originated in Australia as a mutant form of P. edulis or seeds may have
been obtained from tropical America. A hybrid origin of P. edulis f. avicarpa
is unlikely, as normal meiosis with fully viable ovules and pollen grains occurs.
Both P. edulis and P. edulis f. avicarpa have 2n = 18. An interspecic hybrid would
exhibit some meiotic irregularities, leading to irregularities in fertility.
The giant passion fruit is a native of tropical America, but the exact place
of origin is unknown. It can be found from Mexico to Brazil and Peru, in the
Caribbean, Malaysia, Vietnam, India, Sri Lanka, the Philippines and tropical
Africa, and in the warmer parts of Australia, Hawaii and south Florida. There
are no large plantations like those of yellow or purple passion fruit, and it is sold
mainly in local markets.
164
Chapter 8
ECOLOGY
Climatic preferences of the different species vary rather widely. The purple
passion fruit is more adapted to the subtropics and higher elevations of the
tropics with cool periods. The yellow passion fruit is grown commercially in
more tropical areas or lowlands, with the main world producers being Brazil,
Ecuador, Peru and Colombia. These countries are also the largest exporters
of concentrated juice. In Brazil, P. edulis appears to be a variable species, with
forms having different ecological preferences, including those represented by a
yellow fruit type (Martin and Nakasone, 1970).
The giant passion fruit is truly tropical, preferring high humidity and warm
days and nights, although it will produce in the subtropics. It can also grow at
up to 2000 m in the Andes in protected locations, and in other elevated areas
of the world where there is no danger of frost.
Soil
Passion fruit can tolerate a wide range of soil types, although the vines are
highly susceptible to poor drainage and waterlogging. Soil pH may range from
5.5 to 6.8 and even higher. On the Peruvian coast, the yellow passion fruit has
been shown to be fairly salt tolerant.
The giant passion fruit prefers deep and fertile soils, with good drainage and
rich in organic matter. The vine does well in alluvial and sandy soils, even if
slightly alkaline. Too heavy soils are not recommended. The ideal pH is from
5.5 to 6.5.
Climate
Rainfall
A well-distributed annual rainfall is necessary for passion fruit production,
especially if supplemental irrigation is not available (Duarte, 1997). However,
rainfall must be minimal during the owering period, as pollen wetted by
free moisture bursts open and becomes non-functional. Furthermore, rain
minimizes insect activity and hinders pollination. The yellow passion fruit
has been grown quite successfully with rainfall of 8001750 mm uniformly
distributed throughout the year or with supplemental irrigation during dry
periods. Yields of around 40 t/ha have been obtained with a total water supply
of 13001470 mm.
Moisture stress of less than 1.3 MPa leads to a signicant decline in leaf
area, owering and yield (Staveley and Wolstenholme, 1990). Moisture stress
may be one of the major environmental factors responsible for seasonal
uctuations in passion fruit yields (Menzel et al., 1986). When hybrid vines are
165
subjected to water stress, leaf potential recovers within a day of irrigation, leaf
growth returns to prestressed levels in about 4 days and net CO2 assimilation in
6 days (Menzel and Simpson, 1994).
The owers of the giant passion fruit open in the morning, so rains in the
tropics are less likely to bother the pollination processes. It produces satisfactory
yields with a well-distributed rainfall of 15002500 mm. Supplemental
irrigation is needed with rainfall of less than 8001000 mm. During the rainy
season, the withered and dried ower parts get soggy and should be removed
from the peduncle to avoid rotting of that end of the fruit.
Temperature
Different temperature regimes for growing passion fruit in the tropics can be
based on altitude above sea level. The economic life of the purple passion fruit
at about 800 m is 34 years, while plants between 1200 and 1500 m produce
reasonable crops for about 8 years. Low temperatures (15C day/10C
night) reduce vegetative growth and potential yield, while high temperatures
(30/25C) can prevent ower production (Menzel et al., 1987). Some hybrid
cultivars show differences in the optimum temperatures for growth and yield.
The cv. Lacey does not ower at 25/20C, while K-23 and Purple Gold have
reduced owering (Menzel and Simpson, 1994). Purple passion fruit thrives
and yields well at night temperatures of 4.513C and day temperatures of
1830C. Mature vines of the purple passion fruit can withstand light frosts,
but are injured at 1 to 2C (Beal and Farlow, 1984). The yellow passion
fruit is tropical in its requirements, exhibiting more vigor and a wider range of
adaptability. The vine grows and owers well from sea level to 600 m or even
higher, depending on proximity to the equator. In areas with cool winters,
owering stops with the onset of the cold season and the plant will ower for
only 69 months of the year.
The giant passion fruit prefers a tropical climate, like that of the yellow
passion fruit. It will do well on the dry coast of Peru where relative humidity is
normally high. In more humid tropics, it does well as long as excessive rainfall
does not occur during owering. It can also grow in the subtropics, but its
growth is signicantly reduced during the cold months. The ideal average
temperature is 24C, without extremes in temperatures and no cold spells.
Light
Seasonal changes in solar radiation can signicantly inuence productivity.
Lower average irradiance in the cool season, during the wet season with
cloudy weather and with self-shading reduces plant growth, the number of
oral buds and open owers and vine growth (Fig. 8.1). Short periods (1 out
of 4 weeks) of heavy shade signicantly reduce foliar area, dry weight, owerbud numbers, owering and potential yield. Cultivars differ in response; cv.
Purple Gold is more precocious at lower light levels than Lacey (Menzel and
Simpson, 1988). There is an interaction of sunlight with temperature, as no
Length (cm) or Area (cm2 x 10)
Open Flowers or Buds per Vine
Chapter 8
166
Solar
Radiation(MJ
(MJ/m
/Day)
Solar
Radiation
m22
Day1)
Fig. 8.1. The effect of solar irradiation on the hybrid E-23 passion fruit vine growth,
leaf area, ower buds and number of open ower buds. (Redrawn from Menzel and
Simpson, 1988.)
owering occurs at high temperatures with low irradiance. A similar behavior

has been observed for the giant passion fruit.
Photoperiod
Flowering of yellow passion fruit was suggested to be photoperiodic (Watson
and Bowers, 1965). Articially induced short days (8 h) prevent vines from
owering and no owers are produced under natural day lengths of about 11
h or less in Hawaii (21N). In the absence of light-interruption studies, these
results can be interpreted as being due to greater solar radiation or higher
temperatures (Menzel and Simpson, 1994). No data are available for the giant
passion fruit, but it is assumed to be similar to the yellow passion fruit in this
aspect.
Vine
As owers develop on new vine growth, growth of the vine is essential for
continued owering. The vigorous perennial passion fruit vine has medium to
large, toothed leaves (Fig. 8.2). The purple passion fruit has green tendrils,
while the yellow passion fruit possesses reddish or purplish tendrils. The leaves
of the yellow passion fruit are somewhat larger than those of the purple. The
vine can grow to 10 m long (Ruggiero et al., 1996).
167
Fig. 8.2. The stem, leaf, ower and fruit of Passiora edulis f. avicarpa.
The giant passion fruit has thick four-angled stems (hence its name) and
these angles are winged. It also has axillary tendrils that can reach 30 cm,
anked by stipules. The leaves are alternate, oblong-ovate to broad-ovate,
815 cm wide and 1020 cm long, round or cordate at the base, and abruptly
pointed at the apex. The roots are supercial, with 60% in the upper 30 cm of
soil (Borges and Lima, 2008).
Flowers
The solitary large showy passion fruit owers (7.510 cm in diameter) consist
of ve sepals and ve white petals. These form a tubular calyx tube, usually
surrounded by a thread-like crown or corona in the center (Fig. 8.2). The ve
stamens unite into an elongated stalk bearing the ovary, with three horizontal
styles (0.5 cm in diameter). The ovary is superior, one-celled and with three
parietal placentas. Flowers of the yellow passion fruit are larger and more
Chapter 8
168
fragrant than those of the purple passion fruit, with a strong tendency towards
the protandrous habit (Table 8.2). Floral anthesis occurs about 40 days after
the buds become visible in the yellow passion fruit.
The giant passion fruit has even larger solitary fragrant owers of about 12
cm wide, with ve greenish or reddish-green sepals on the outside and pink,
white or purple inside. The petals are white and pink. The corona laments are
56 cm long and purple and white below, blue in the middle and pinkish-blue
above around the pistil, style and stigma. The time from bud formation to oral
anthesis in P. quadrangularis ranges from 17 to 24 days.
The name passion ower was given by early Spanish Jesuit missionaries,
to whom the ower represented the passion of Christ and illustrated the
crucixion. The 10 sepals and petals represent the 10 apostles at the crucixion;
the fringed crown represents the crown of thorns; and the ve stamens and
three styles represent the wounds and nails, respectively. The tendrils are the
cords or scourges, while the lobed leaves represent the hands of the persecutors.
The white symbolizes purity and the blue the heavens.
Flower buds are produced at every node of new growth. After four to 10
owers have set fruit, further setting of the remaining owers ceases, even when
hand-pollinated. Fruit set resumes when the initially set fruit begin to mature.
This leads to peaks of fruit production with continuous vine growth (Fig. 8.3).
350 l/Vine/Week
N and K 35% Total
Jan
200 l/Vine/Week
350 l/Vine/Week
N and K 30% Total
N and K 35% Total
Feb Mar Apr May Jun
Jul
Aug Sep Oct Nov Dec
Month
Fig. 8.3. The growth, owering and yield of hybrid passion fruit vines in relation
to pruning, irrigation and fertilization. (Redrawn from Menzel et al., 1989, 1993.)
Mean of 2 years data from southern Queensland, Australia. K, potassium; Mg,
magnesium; N, nitrogen; P, phosphorus.
169
The alternation of fruit setting and cessation of setting, which leaves fruitless
spaces on long vines, could be a physiological mechanism to balance the number
of fruit to leaves and photosynthate supply.

The ower is large with a strong fragrance, attractive colors, abundance of
nectar and large sticky pollen, all of which is conducive to insect pollination.
The pollination process and the self-incompatibility problem of yellow
passion fruit and sometimes giant passion fruit are normally solved by insects,
especially carpenter bees (Xylocopa). Other frequent visitors are honey-bees
(Apis mellifera) and various ies. Carpenter bees are much more efficient
pollinators because of the size of their bodies; their backs are more likely to
make contact the anthers and stigma (Aubert, 1975).
Fruit set is about 70% by natural pollination, while hand-pollination fruit
set can reach 100%. Hand-pollinated fruit are larger, heavier and higher in
juice content, because of the increase in seed number (Akamine and Girolami,
1959). In Honduras, hand pollination at 1 pm or 4 pm resulted in 53.5%
and 58.6% fruit set compared with 44.6% for natural pollination. The handpollinated fruit were 2025% heavier, and juice weight per fruit was 73.8% and
82.0% higher than for the natural-pollinated control. Seed numbers increased
by 40%. Hand pollination at 4 pm gave better results, although not statistically
greater than at 1 pm (Duarte and Sierra, 1997).
Style curvature is crucial for passion fruit ower pollination and fruit set.
When the owers open, which is normally at noon, the styles are upright. They
gradually recurve during the afternoon to approximately 55 in the rst hour
after anthesis and 85 by 6 pm, before reverting to an average of about 20
curvature at ower-closing time (Hardin, 1986). The most effective time for
pollination is after the styles have completely recurved, with the stigmas being
receptive only on the day of anthesis. Ruggiero et al. (1976, 1996) reported
three kinds of owers with respect to style curvature, sometimes on the same
vine: total curvature (about 71% of owers), partial curvature (2023%)
and upright styles (610%). Flowers with upright styles fail to set fruit with
pollen from compatible vines and wither under natural conditions or when
hand pollinated. Pollen of owers with upright styles effect good fruit set when
deposited on stigmas of total- and partial-curvature owers of different vines,
indicating high pollen viability.
The purple passion fruit is self-compatible, as are its hybrids, with the
yellow passion fruit often requiring cross-pollination. On pollination, stigmas
must be kept dry for at least 2 h the time necessary for pollen germination
(Akamine and Girolami, 1959). Pollen bursts and becomes useless on contact
with free water, although germinated pollen is not affected. Since owers of the
170
Chapter 8
yellow passion fruit open during the afternoon, rain or irrigation by overhead
sprinklers during the afternoon can greatly reduce fruit set. This is a problem in
many tropical areas, where heavy afternoon rains result in low yields.
The giant passion fruit is normally self-compatible but is sometimes
protandrous. Certain lines can be self-sterile, needing the presence of other
plants for cross-pollination. The owers start to open at 56 am and take about
1 h to complete the process. At that time the styles are vertical; by 9 am they
are completely curved and the anthers start releasing pollen. In the afternoon,
petals and sepals close and the ower is accid. When hand pollination is
practiced, the best time is between 9 am and noon when the styles are curved;
pollination in the afternoon is less successful. An average fruit set of 8.8% was
observed in one study (Duarte and Marn, 2002).
Fruit
Fruit range from 6 to 8 cm in diameter by 7 cm in length in the yellow passion
fruit with a weight of 70130 g, although fruit weighing 200 g are not
uncommon. EMBRAPA in Brazil has released selections with fruit weighing
around 200 g. These have been introduced to Ecuador, Peru and Colombia.
Fruit weighing as much as 650 g have been reported in Brazil (CampoVivo,
2010). This is achieved through selection, hybridization and good eld
management, especially nutrition and hand pollination. The purple fruit is
normally 3.57 cm in diameter and 49 cm in height, with a weight of 60
100 g (Ruggiero et al., 1996).
Both passion fruit types are round to oval in shape and have a hard exocarp
(Fig. 8.2). The mesocarp is a thin green layer beneath the exocarp and above
the white endocarp, forming a shell of 36 mm. Each seed has a hard black
testa and is surrounded by a juicy edible aril. The seeds are attached by peg-like
funiculi to the endocarp. The yellow passion fruit has up to 350 seeds. The
yellow to orange juicy aril pulp of the purple passion fruit is aromatic, with a
very desirable avor. Total soluble solids usually exceed 15% and total titratable
acidity ranges from around 2.5% to 4.0% (Table 8.3), depending on the season.
The yellow passion fruit pulp is more acid than the purple passion fruit (Seale
and Sherman, 1960).
The purple and yellow passion fruit take 6090 days to mature (Fig. 8.4).
Growth follows a sigmoid growth curve, reaching maximum size in about 21
days, when sclerication leads to a hardening of the shell. Subsequent fruit
mass increase occurs at a slower rate and reaches a maximum at about 50 days
from anthesis for purple passion fruit and 60 days for yellow passion fruit. The
increase in sugars, with a concomitant decline in starch, occurs during this last
phase. When maximum mass is achieved, titratable acidity begins to decline
as maturity approaches. A positive correlation exists between the number of
171
Table 8.3. Pulp characteristics of Passiora edulis, P. edulis f. avicarpa and some
hybrids (Beal and Farlow, 1984).
Pulp
(%)
Total
soluble
solids (%)
Titratable
acidity
(%)
Total
volatile
esters
(ppm)
122
49
15.3
2.4
159
72
50
37
15.3
2.4
83
46
14.5
3.2
122
3568
13.416.1
2.33.6
49200
No. of
seeds
P. edulis
36
P. edulis
f. avicarpa
3-1
Other hybrids
Fruit Mass (g)
Acidity (% Citric) and Soluble Solids (%)
Fruit
weight
(g)

Fig. 8.4. Change in purple passion fruit mass, skin yellowing, total soluble solids
and titratable acidity during growth and development. (Redrawn from Shiomi et al.,
1996.)
seeds that develop and fresh fruit mass and juice content. Juice content reaches
a maximum on the vine and declines due to dehydration after abscission.
Growth of the giant passion fruit also follows a single sigmoid curve, reaching
its nal size 25 days after anthesis and ripening in 6580 days (Duarte and
Marn, 2002).
172
Chapter 8
Genetics and cytogenetics
On the basis of established chromosome numbers of relatively few species, the
2n = 18 group is the largest, with all of the horticulturally important species
and hybrids included (Storey, 1950). The horticultural species are hexaploids
with fairly high interspecic compatibility and regular meiosis.
Purple tendril color is dominant over green. Fruit-shell color is controlled by a
single pair of genes lacking dominance, with three color types being recovered in
the F2 (Nakasone et al., 1967). Inheritance data for crown rot indicate a simple
dominant gene for resistance to Fusarium oxysporum f. passiorae, with P. edulis f.
avicarpa being the resistant parent. The yellow passion fruit has also been found
to be resistant to a similar disease in South Africa, attributed to Phytophthora
nicotianae var. parasitica. The purple passion fruit is susceptible to passion fruit
woodiness virus (PWV), while some yellow passion fruit lines are tolerant.
Breeding and selection

In establishing a passion fruit breeding program, it is highly desirable to
develop a collection of as many species as possible in at least two ecological
zones (sea level and a higher elevation). The breeding objectives in Hawaii are:
(i) oval-shaped fruit (10% more juice than in round fruit); (ii) avor of purple
passion fruit and acidity, fruit size, juice recovery and vine vigor of yellow
passion fruit; (iii) bright tangerine-colored juice with high total soluble solids;
(iv) high degree of self- and cross-compatibility and good fruit set; (v) resistance
to wilting disease and Alternaria brown-spot disease; and (vi) resistance to
broad mite and other insects. For other regions, breeding for cold tolerance
and resistance to PWV, Fusarium and Phytophthora crown or collar rot and
other problems must be added to the set of objectives (Farlow et al., 1984).
Where the purple passion fruit is the primary crop, rootstock breeding for
resistance to crown and root rots is a major objective.
In Australia, lines produced from P. edulis u P. incarnata have shown
improved cold hardiness and tolerance to the severe strain of PWV. The
original F1 hybrid and selections from the F2 and F3 populations backcrossed
to P. edulis hybrids and to P. edulis f. avicarpa have shown promise as improved
rootstock. In South Africa, P. caerulea is tolerant to both Fusarium oxysporum
var. passiorae and Phytophthora parasitica, and hence is used as a rootstock
(Grech and Frean, 1986). Australia has developed purple hybrids (3-1, E23, Lacy and Purple Gold), which are maintained by grafting on seedlings
of the yellow passion fruit.
Breeding through interspecic hybridization has not been explored
adequately in any sustained program. Numerous hybrids have been made and
173
the most frequent cross is between P. edulis and its form P. edulis f. avicarpa. This
cross can be accomplished only if P. edulis is used as the female parent, as the
reciprocal is strongly incompatible. The latter condition has discouraged the
development of clonal cultivars in the yellow passion fruit. A cross between
yellow passion fruit and P. alata produces fertile hybrids with high-quality fruit.
Yellow passion fruit can be transformed using Agrobacterium tumefaciens,
offering the possibility of incorporating genes for virus resistance and other
desirable traits.
Cultivars
Many growers plant selected seeds or use purple passion fruit vines grafted
onto Fusarium-resistant lines of the yellow passion fruit. The higher acidity
of the yellow passion fruit, along with the need to develop disease-resistant
cultivars, has led to the development of hybrids (Table 8.3). The selection of
hybrids between purple and yellow passion fruit has focused on winter and
summer cropping and tolerance to passion fruit mosaic virus, nematodes,
Alternaria spot and Fusarium wilt. In Australia, two or three hybrids are
grown to spread production peaks, with E-23 and Purple Gold being
most widely used. The hybrids Supersweet, Land 3 and Misty are used
in the subtropical parts of Australia, while the purple-skinned selection
of the yellow passion fruit Panama is grown in tropical areas. The most
recent release in Hawaii was Noels Special, a yellow passion fruit with an
unusually bright orange-colored juice and tolerance to Alternaria brown
spot. The round fruit averages 90 g in weight, yielding 4356% juice by
weight and with total soluble solids of 1519.8%. Hybrids have been
developed and released in Taiwan, with Tainung No. 1 being one of the
most common. The yellow passion fruit Hawaiana is used in Colombia and
Venezuela, along with some of the modern Brazilian selections. There are
a number of selections in Brazil, including Muico, Peroba and Pintado
(all purple) and the yellow Miram and Grande. More recently, Brazil
has released selections (Sol do Cerrado, Gigante Amarelo and Ouro
Vermelho) from crosses of the yellow type with fruit that can weigh as much
as 650 g (CampoVivo, 2010). The released hybrids are larger, with increased
acidity and more orange aril.
Giant passion fruit has no dened varieties, but there are some types
or selections. Colombia has two types. One, from the province of Choc, has
a relatively small fruit of 1015 cm long and 810 cm diameter, with
sweeter arils and a thinner white pulp of about 11.5 cm and sparse foliage.
The other is the giant or normal type, with fruit that are 2530 cm long and
1215 cm wide, and a white pulp of 2.54.0 cm thick that can amount to
60% of total fruit weight. The arils and juice are not as sweet as in the smaller
variety.
174
Chapter 8
CULTURAL PRACTICES
Propagation
Passiora species are readily propagated by seeds, cuttings, air layers or grafting
on a selected seedling rootstock. Leafy yellow passion fruit cuttings can also
be propagated easily with naphthalene acetic acid. High levels of naphthalene
acetic acid cause a reduction in root number and vigor, although the rooting
percentage is similar (Duarte and Franciosi, 1976). Grafting and budding are
fairly easily performed. Cleft grafting is more successful with adult scions, while
whip grafting works better with juvenile material. When the purple passion
fruit or its hybrids are the desired cultivars, plants are propagated by grafting
on seedlings of the yellow passion fruit. Grafted vines are more vigorous than
their seedling counterparts and have longer lifespans. Grafting is done 5055
cm above the ground to prevent soil contact with the scion.
When yellow passion fruit is grown or used as a rootstock, plants are produced
exclusively from seed. The yellow passion fruit has vigor and resistance to root
and stem rots and nematodes. Each seedling has a different genetic make-up
and cross-pollination can take place. Asexually propagated material shows selfincompatibility, and requires an orchard to be planted with vegetative material
from a number of seedling plants that are randomly chosen.
Seeds can be sown immediately or stored at about 1013C for future
use. Seeds stored at room temperature for 3 months give better than 85%
germination. Seeds germinate in about 2 weeks, although germination can
extend over 23 months because of seed-coat dormancy. Cracking the seed
coat increases germination, but scarifying with sandpaper or fermenting seeds
with wall-degrading enzymes has no effect. However, seed cracking is feasible
only for small quantities of seeds. Twelve-hour periods alternating between
20C and 30C also increases germination compared with a constant 30C.
Giant passion fruit seeds from 45-day-old fruit have about 23% germination,
while seeds from 60-day-old fruit have 98100% germination. The best
treatment for optimum germination is to ferment the seeds in their juicy pulp
for 23 days, followed by washing and supercial drying out for 3 days. Seeds
at room temperature lose viability in 1 month (Duarte and Marn, 2002). Seeds
put in seedbeds germinate in 23 weeks, and plants can be taken to the eld
when they are 2030 cm tall. This plant can also be propagated fairly easy
by semi-hardwood or mature wood cuttings of 3040 cm long and 12 cm
diameter, stuck directly in nursery bags (Alix and Duarte, 1999).

Seedlings at the two- to four-leaf stage are transplanted into individual plastic
containers, grown in semi-shade for 12 months and then gradually provided
175
with more sunlight. Seeds can also be sown in 60 cell trays and taken directly
to the eld. Seedlings are considered ready for eld transplanting when they
have attained heights of 2550 cm and have been hardened in full sunlight
for 12 months. For grafted vines, the scion portion should have grown
until about 25 cm and hardened. The practice of applying fertilizer in the
planting holes varies widely, from no fertilizer to 1 kg of superphosphate in
South Africa. In Hawaii and Australia, fertilizer (60114 g of 10:5:20) and/
or manure is incorporated in a circular area (approximately 0.8 m diameter)
around each planting site.
The trellis is the principal initial cost of production. There are a number of
different trellis types, each having variations in height, number of strands and
placement of wires, length of cross-arms (if arms are used), spacing of posts
and method of construction. The two most commonly used types are the I or
vertical or fence trellis, with one to several wires strung parallel, one below
the other, on upright posts and the T- or cross-type trellis, with three strands
of wires, one running on top of the posts and the other two attached at the ends
of each cross-arm. With taller posts, two to three strands of wires are used,
spaced about 0.60.9 m apart. In the cross-type trellis, the cross-arm is 1.21.5
m long and is placed 0.40.6 m from the top of the post. For yellow passion
fruit, some authors recommend the fence-type trellis when manual pollination
is performed since the other trellis types make this activity more difficult. In
Hawaii, the cross-type trellis was found to give higher yields over the fencetype trellis. Trellis height is around 2.0 m.
A higher trellis provides a longer time for vines to trail from the wires to the
ground with less piling of vines at the top, due to better spreading of growth on
the three strands of wires. The high trellis and spreading of vines by the crossarms allows greater exposure of the vine to sunlight. A pergola-type trellis can
also be used but if the foliage gets too dense then some of the ripe fruit will stay
on top and are lost, since harvesters cannot see or reach the fruit. Minimum
row spacing for grape trellises should be about 3 m to permit mechanization.
If the cross-type trellis with a 1.0 m bar is used, there should be a distance of
2 m between the ends of bars on the parallel trellis of the adjacent row. Plant
spacing within the trellis row is dependent on the type of passion fruit being
grown, with too high densities leading to lower yields (Fig. 8.5). In cooler
subtropical areas, the purple passion fruit or its hybrids may be spaced 2.53
m apart because of less vigorous growth. In-row spacing may be up to 5 m if
very vigorous growth occurs. For the yellow passion fruit in warm areas, plants
can be initially spaced at 2.42.5 m; after the rst harvest year, every other
vine is thinned to provide a permanent spacing of 4.85.0 m (694 plants/ha).
Taking advantage of the high yields in the rst crop year may be well worth the
added cost of the additional plants. Under poor soil and watering conditions,
the initial spacing between plants can be much lower with better yields during
not only the rst but also the second year.
The trellis systems used for giant passion fruit are very similar. Plants can
Chapter 8
Yield (t/ha)
176
Fig. 8.5. Vine spacing signicantly affects the yield of yellow passion fruit during
its third year at Viamao, Rio Grande do Sul, Brazil. (Redrawn from Manica et al.,
1985.)
be transplanted once have reached about 2530 cm for seedlings and 4050
cm for plants from cuttings. With this species a pergola-type trellis with wires
crossing at 5060 cm can be used with posts in rows separated 3 m and 36 m
between posts in the row, with more posts for a sturdier trellis. The fence-type of
trellis is also used, with a height of 2.32.5 m and ve lines of wire, separated
by 4550 cm (Martnez, 1981). If the plants have been set too close and the
foliage starts to overcrowd then they can be thinned by eliminating every other
plant in the row. The T or cross-type trellis can also be used.
Hand pollination
Self-incompatibility in yellow passion fruit and some types of giant passion
fruit is often solved by insects, especially the carpenter bee. If the work of the
carpenter bees is not satisfactory or they are absent then hand pollination is
performed. Pollination is conducted when the owers are open, normally in
the afternoon for yellow passion fruit and in the morning for the giant type.
The nger of a gloved hand will trap pollen when touched to an open ower;
this will transfer to the stigmas of the new ower and at the same time the
glove will catch pollen from this new ower. This captured pollen is repeatedly
transferred through the orchard. In the case of a fence trellis, the pollinator
would go from a ower in one fence to a ower in the opposite-side fence, to
make sure the owers are from different plants and avoid incompatibility. An
individual can pollinate 5060 owers per minute, and two or three people
177
can cover 1 ha during an afternoon. Hand pollination should be performed

whenever owers are present, not only at owering peaks, to ensure high
yields from more and larger fruit (Ruggiero et al., 1996).

Planting holes are made 0.61.0 m away from the posts. In Hawaii, seedlings
4560 cm tall are topped at the time of transplanting to force lateral shoots
to develop, of which two to four leaders are allowed to grow. One or two of
the lateral shoots are trained to grow in one direction on the trellis, with the
remaining shoots growing in the other direction. Bamboo stakes with short
branches and about the height of the wires provide excellent support for vines.
Tendrils wrap around the bamboo branches, so tying the vines to the stakes is
unnecessary. All other trailing branches are removed. If tying is used, care has
to be taken to eliminate the ties before they strangle the stems. A single leader
is preferred in some systems and is trained along the wire in one direction. In
the fence system, a leader is allowed to grow until it reaches just above the rst
line of wire. At this point it is pinched and three shoots are allowed to grow:
one will be directed to climb to the next line of wire, while the other two are
directed along the wire in opposite directions. This is done for every line of
wire. At the last wire, after pinching only two shots are left, which should go in
opposite directions.
Passion fruit vines produce owers on the current seasons growth, and
practices that encourage new lateral growth increase owering (Fig. 8.3).
Occasional pruning is needed, although the degree varies. Once a lateral has
completed its fruiting it should be cut back to a developing side shoot as close to
the main leader as possible. If no side shoot has formed then the lateral should
be cut off at the fourth or sixth node from the main stem. Once the laterals
reach the ground after 1215 months, they should be cut back to 30 cm of the
main leader. Pruning should be performed when fruit load is small (Joubert,
1984). If not pruned, laterals begin growing on the ground, the vines tend to
form a tangled mass on the wires, and only the new outside growth bears fruit.
Pruning of old growth can limit the current crop, while removal of new growth
restricts future cropping. The following general conclusions are drawn:
1. Light, selective pruning, particularly at the end of the annual production
cycle, enhances new growth and maintains high yields the following year. This
consists of removing all vines about to reach the ground or growing on the
ground. Vines are cut near the trellis wires, but a few nodes away from the main
stems. Vines hanging only halfway to the ground are left.
2. Long vines should not be thrown over the trellis, as this only increases
entanglement on the trellis and depresses yields.
Chapter 8
178
3. Some vine growth on top of the entangled tops of the trellis may be pruned,
as fruit produced on these vines is apt to be lost in the maze of vines, especially
when the planting is 2 or 3 years old. Pruned vines on the trellis should be left
to dry in place, as attempts to remove the cut vines can damage uncut vines.
4. Cross-arm trellises of 2.43.0 m high, although more costly initially, offer
some advantages, such as better spread of vines and greater height for vines to
grow and reach the ground. It is also easier to remove old vines hanging down
from the wires.
Pruning is also recommended for the giant passion fruit. The formation
pruning consists of eliminating all side shoots to leave a single stem, and is
similar to that used for purple and yellow passion fruit. Production pruning
consists of eliminating shoots that have already produced fruit in order
to stimulate new shoots that will ower. At the same time, all damaged or
diseased shoots should be removed (Martnez, 1981).
Irrigation
Node No., Open Flower No., Root Mass (g)
Adequate soil moisture is required to sustain vegetative growth and production

(Fig. 8.3). No oral buds are initiated under dry conditions (Fig. 8.6) as vine
extension and growth are curtailed. Production is generally associated with
higher rainfall in the 2 months before owering. In Australia, 300400 l/vine/
week is required during the summer. Passion fruit are herbaceous and thus
need a good supply of water to develop and produce properly.
Node No.
Open
Flower No.
103
102
101
100
Soil Water Potential (MPa)
Fig. 8.6. Effect of different moisture stress applied for 10 weeks on the number of
nodes produced, leaf area, open-ower number and root dry mass. (Redrawn from
Menzel et al., 1986.)
179
Fertilization
Flowers per Vine
Vine Extension (mm)
Recommendations for fertilization vary widely, with all directed at

encouraging new growth throughout the season. Since all owers occur on
new growth, there is a high demand for nitrogen: leaf nitrogen levels above
4.5% are recommended (Fig. 8.7). Fruit mineral analysis indicates that to
produce each 1000 kg of fruit/ha requires 33 kg/ha of fertilizer, based on a
10:5:20 formulation, assuming 100% uptake efficiency by the fruit; a more
realistic uptake value is 50%. Based on 667 plants/ha (3 m u 5 m spacing),
each plant should receive 4.4 kg of 10:5:20 fertilizer. In Brazil, Borges et
al. (2002) prepared more specic formulations for amounts of nitrogen,
phosphorus and potassium according to soil analysis and yields for irrigated
yellow passion fruit (Table 8.4). Deciency symptoms have been described for
nitrogen, phosphorus and potassium and the microelements iron, boron, zinc,
magnesium, calcium and sulfur (Abanto and Muller, 1976; 1977a,b).
About 60% of passion fruit roots are in the upper 30 cm of soil, and almost
90% are between 0 and 45 cm from the base of the stem. In young orchards,
fertilizers should be applied in a 20-cm-wide ring around and 10 cm from the
trunk, gradually increasing this distance with the age of plants. In mature
vines, fertilizer should be applied in a band 2 m long and 1 m wide, on both
sides of the plants, and 2030 cm from the trunk.
The time of fertilizer application depends on the climatic conditions, general
appearance and vine performance. In the rst year in Hawaii about 113 g
of 10:5:20 fertilizer is placed in the hole at planting, with the same amount
Leaf Nitrogen (%)
Fig. 8.7. Effect of leaf nitrogen on vine growth and ower production of E-23
passion fruit. (Redrawn from Menzel et al., 1989.)
Chapter 8
180
Table 8.4. Fertilization recommendation in the plantation, formation and

production phases of irrigated yellow passion fruit (Borges et al., 2002).
Phosphorous resin
(mg/dm3)
015
Nitrogen
(kg/ha)
1640
>40
Potassium soil (cmolc/dm3)

00.07
0.08
0.15
P2O5 (kg/ha)
At planting
150a
120
80
During growth (days after planting)
0.16
0.30
0.31
0.50
>0.50
K2O (kg/ha)
0
30
10
20
10
60
20
30
20
10
90
30
40
30
20
10
120180
40
60
40
30
20
0
0
During fruit production (expected yield, t/ha)

<15
50
50
30
20
100
90
70
50
1525
70
90
60
40
160
120
90
70
2535
90
120
80
50
200
160
120
80
120
150
100
60
250
200
150
100
>35
a
As bovine manure.
applied every 68 weeks. In the second year, when vines become productive,
application should be based on vine growth. In Malaysia, where temperatures
allow year-round growth, 5.4 kg of a 10:5:10 fertilizer per plant per year
in four applications is recommended for the yellow passion fruit. Similarly,
four applications of 15:4:11 at 500 g per mature vine per application,
alternating with four applications of urea at 460 g per vine per application, are
recommended in Queensland for purple passion fruit and its hybrids (Menzel et
al., 1993).
No specic recommendations exist for the giant passion fruit, but it does
better if planted in soils high in organic matter. A soil analysis should be
performed, taking into account that this plant requires good amounts of
nitrogen and phosphorus (Martnez, 1981). Yellow passion fruit dosages could
initially serve as a reference.
Pest management
Diseases
Passion fruit has very few serious diseases, with the extent of disease
depending on the species and the growing environment (Table 8.5). Alternaria
brown-spot disease has been reported from various passion fruit regions of the
181
Table 8.5. Diseases of passion fruit.

Common name
Organism
Parts affected
Region
Anthracnose
Colletotrichum
gloeosporioides
Leaves, fruit
Probably
universal
Brown spot
Alternaria
passiorae, A.
tenuis, A. tomato
Alternaria alternata
Fruit, leaves
Universal
Fusarium
oxysporum f.
passiorae
Roots, crown
Alternaria spot
Fusarium wilt
Phytophthora blight Phytophthora

nicotianae var.
parasitica, P.
cinnamomi
Fruit scab
Stem-end rot of
giant passion
fruit
Woodiness virus
(insect vectors)
(a long, exuous
virus, 750 nm
long)
Cladosporium
herbarum
Botryodiplodia
theobromae
Australia
Probably
universal
Leaves, stem, fruit, Probably

roots
universal
Fruit
American tropics
Stem end
East Africa
Stem, leaves, fruit
American tropics,
Australia,
Malaysia,
South Africa,
Caribbean area
tropics and subtropics on both the purple and the yellow passion fruit. Fruit
of the yellow passion fruit in high-rainfall areas can be 6698% infected and
most of the leaves defoliated. The reddish-brown to brown, sunken spots,
1.35 cm in diameter, are easily identied. When grown in low- to moderaterainfall areas, vines with high tolerance, combined with a good fungicidal
spray program, can produce up to 8090% marketable fruit.
Fusarium wilt can cause devastating losses on the purple passion fruit, with
a very rapid onset of 2448 h. Symptoms include browning of the vascular
system of roots, crown and stem. Phytophthora blight is serious where purple
passion fruit is grown. It affects the vines, causing defoliation and rotting fruit.
The disease may completely girdle the stem, and causes root rot in soils with poor
drainage. In the giant passion fruit, rotting sometimes occurs at the proximal
end where the dried ower parts remain attached and cover a concavity at the
peduncle insertion. This holds rainwater, and if it does not quickly evaporate
then rotting can occur. Therefore, elimination of the ower remains after fruit
set is recommended.
182
Chapter 8
PWV is an almost universal, aphid-transmitted disease of both purple and

yellow passion fruit. It is a serious problem in Australia, South Africa, Malaysia,
Taiwan, Panama and the Dominican Republic. Infected plants have leaves
with a light-green mosaic pattern and yellow speckling and crinkling. The
fruit is misshapen, with a thick woody rind and shrunken cavities (McCarthy,
1982). PWV is caused by a long exuous virus, 750 nm in length, belonging
to the potyvirus group. It is transmitted by the aphid Myzus persicae, as well as
mechanically by grafting or contaminated tools. There is no evidence of seed
transmission.
Insects
The most troublesome pests are fruit ies (Table 8.6), as they are difficult to
control even using a well-executed spray program. The ies can be particularly
injurious to passion fruit grown at low elevations. Most fruit-y damage
occurs on young passion fruit, where the fruit will shrivel and drop. The larvae
develop better in soft, immature fruit than in mature ones (Anon, 1972).
Fruit-y ovipositing can also cause crater-like scars on mature fruit.
Aphids can attack young leaves and growing points, as do stinkbugs. Mealy
bugs and several scale insects, including barnacle scale, can become a problem.
Other pests, such as red-banded thrips (Selenothrips rubrocinctus), occasionally
build up to serious proportions and cause defoliation. Mites are particularly
prevalent in areas of low rainfall and during the dry season, damaging and
defoliating vines; they are effectively controlled with sulfur sprays. The plants are
also very susceptible to nematodes such as the root-knot eelworm (Meloidogyne
sp.) and other species. In many cases, grafting on the more resistant yellow
passion fruit is recommended (Villiers, 1982).

Yield
Passion fruit yields vary with climate, species of passion fruit, cultivation
practices, trellis type, presence of disease and the abundance of appropriate
pollinating agents. Among the two most commonly cultivated passion fruit,
the yellow passion fruit has the highest yield potential, with 44.8 t/ha/year
in Hawaii and 41.861.9 t/ha/year at the Malaysian Agricultural Research
and Development Institute research station in Sungai Baging, Malaysia (Chai,
1979). In commercial orchards, a more realistic yield is between 20 and 30 t/
ha/year. The purple passion fruit is less vigorous. Yields of 510 t/ha/year can
be obtained in Australia, with the hybrids having yields of up to 25 t/ha/year
(Beal and Farlow, 1984). Average yields for the giant passion fruit are 1520
t/ha/year in Colombia (Martnez, 1981). This can be increased with improved
irrigation and fertilizer management.
183
Table 8.6. Insect and mite pests of passion fruit.

Common name
Organism
Mediterranean fruit Ceratitis capitata

y
Parts affected
Region
Mostly fruit
Hawaii, Central
America,
Panama, Mexico
Hawaii, Panama,
other areas
Hawaii, Asia,
parts of Africa,
western Pacic,
Malaysia
Australia
Melon y
Dacus cucurbitae
Oriental fruit y
Dacus dorsalis
Mostly fruit, also

vegetative parts
Fruit
Queensland fruit
y
Caribbean fruit y
Dacus tryoni
Mostly fruit
Anastrepha sp.
Fruit
Leaf caterpillar
Leaf caterpillar
Leaf eater
Dione juno
Agraulis vanillae
Diabrotica
speciosa
Cryptorhynobus
sp.
Atta cephalotes
Selenothrips
rubrocinctus
Brevipalpus
papayanis
Brevipalpus
phoenicis
Tetranychus
cinnabarinus
Hemitarsonemus
latus
Meloidogyne spp.
Rotylenchus
reniformis
Leaves
Leaves
Leaves
Borer
Leaf-cutting ant
Red-banded thrips
Red spider mite
Red and black at
mite
Carmine mite
Broadmite
Nematodes
Caribbean,
American tropics
American tropics
American tropics
American tropics
Root, stem,
branches
Leaves
Leaves
Panama
Vegetative parts
Universal
Bark, diebacks,
fruit
Mature leaves
Universal
Young terminal
leaves
Roots
Universal
American tropics
Panama, Hawaii
Universal
Universal
Harvesting
Hand-harvesting is the most costly operation in passion fruit culture and can
account for approximately 4050% of the variable costs. Normally, fruit are
allowed to ripen on the vine and abscise. Fallen fruit are gathered once or
twice per week, depending on the quantity. Fruit are gathered more frequently
during rainy periods, and daily if sunburn is a problem in summer. Vineyard
layout and management, such as locations of access roads, line lengths,
row widths, trellis types, eld grading, trellis orientation and pruning before
184
Chapter 8
vines trail on the ground, inuence harvesting efficiency. Trellis posts aligned
on the raised part of the bed, with ground sloping away on both sides of the
posts, allows falling fruit to roll to the side, concentrating the fruit outside the
canopy and making it easier to gather. Harvesting using hand-raking and
a machine that straddles the fruit and sucks them up has been developed.
Alternatively, fruit are picked up from a mown sward with minimal damage.
Despite the fruits thick skin, it is very susceptible to mechanical damage.
Mechanical harvesting aids cause damage and make the fruit generally
suitable only for processing.
The giant passion fruit should be harvested when violet skin appears at
the distal end of the fruit and the peel becomes yellowish and translucent,
starting at the apex. It will fail to ripen properly if harvested too early. The skin
is susceptible to mechanical injury even a slight pressure with the ngers will
leave a mark. The fruit are cut from the vine using sharp knives or pruning
shears and handled very carefully.
Fruit for the fresh market are carefully handled in small, 510 kg berboard
cartons. Marketing standards for the fresh-fruit market in Australia require
half- to full-ripe fruit, not less than 35% pulp and larger than 4 cm in
diameter. Diseased or badly blemished fruit are culled. Dark-purple fruit with
120140 fruit/carton attract the best prices. Growers wax their fruit to extend
shelf-life and secure higher prices. Purple passion fruit can be kept for up to
45 weeks with little weight loss at 5C and 8090% humidity (Fig. 8.8). The
yellow passion fruit can only be stored for about 1 week at 57.5C. Ethylene
can be used to enhance the skin-color development of mature fruit without
affecting soluble solids or juice pH.
The giant passion fruit is a very susceptible to injury when ripe, so it is
normally harvested during early ripening. The fruit can be transported in
berboard boxes or wood crates in layers with some protective cushioning,
although single-layer boxes are better. Alternatively, partially ripe fruit are
packed in a vertical position with the proximal end at the bottom. It is preferable
to wrap every fruit with paper or to cover it with a plastic foam net sleeve. It
should be stored at 10C and 8590% RH for maximum life.
UTILIZATION
Passion fruit is valued more for its unique avor and aroma than for its
nutritional value. According to Wenkam (1990), yellow and purple passion
fruit are good sources of provitamin A, niacin, riboavin and ascorbic acid
(Table 8.7). Yellow passion fruit juice has 2410 mg/100 ml of vitamin A,
while purple passion fruit has only 717 mg (Ruggiero et al., 1996). The
185
Total Soluble
Solids
Loss
Time in Storage (Days)
Fig. 8.8. Changes in mass and total soluble solids of purple passion fruit stored at
5C, 10C and 15C. (Redrawn from Arjona et al., 1992.)
Table 8.7. Composition of 100 g edible portion of passion fruit (Wenkam, 1990)
and giant passion fruit (Morton, 1987).
Giant passion fruit
Constituent
Proximate
Water (g)
Energy (kJ)
Protein (g)
Fat (g)
Carbohydrate
Fiber (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
P. edulis
P. edulis f.
avicarpa
85.6
213
0.39
0.05
13.6
0.04
84.9
222
0.67
0.18
13.72
0.17
94.4
0.112
0.15
0.7
78.4
0.299
1.29
3.6
3.6
0.24
12.5
3.8
0.36
24.6
0.7
0.7
0.7
3.6
3.6
3.6
29.8
Trace
0.131
1.46
717
20.0
Trace
0.101
2.24
2410
14.3
0.033
0.378
0.004
0.003
0.120
15.3
0.019
Thick white
esh
Aril and seed
186
Chapter 8
passion fruits characteristic juice avor and aroma must be retained during
processing. Pulp from Alternaria brown-spot-infected fruit can seriously affect
juice avor. Yellow passion fruit has an average juice yield of 3033%, while
purple passion fruit has a yield of 4550%. The extracted juice is quickfrozen as a puree and kept frozen for later processing of nished products. The
juice can be pasteurized (85C, 1 min) prior to freezing. However, the avor
constituents of passion fruit are extremely sensitive to heat treatments and the
pasteurization process can cause the loss of 35% of the volatile components.
The strong, pleasing avor of passion fruit permits its use as a pure juice or a
constituent in various frozen and heat-processed punches. It adds an excellent
avor to other products, such as pies, cakes, sauces, salads and sherbets. The
fresh juice and concentrates are refreshing mixers with alcoholic beverages,
such as gin, vodka and rum. Other products include tropical fruit cocktail,
passion fruit sherbet and ice, and jelly and jam combinations. The juice can
be boiled down into a syrup that is used in the ice cream and pastry industries.
Passion fruit processing produces large quantities of waste in the form of
rind and seed, which creates a disposal problem. The waste is composed of 51%
rind and 11% seed by mass for the yellow passion fruit. Dehydrated rind has
been fed experimentally in rations to swine and dairy cattle, with good results.
The seeds contain more than 20% oil; this is similar to that of soybean or
safflower, and can be used for the same purpose. The oil has more than 80%
unsaturated fatty acids.
Giant passion fruit is normally consumed fresh. Its nutritional value and
vitamin content are not as high as those of the yellow or purple passion fruits
(Table 8.7). The white esh of the ripe fruit, with the outer peel and the inner
skin removed, can be cut into pieces and eaten like papaya. However, it does not
have a very distinct avor so is usually added to pieces of other fruits such as
papaya, pineapple, melon, watermelon or banana and seasoned with orange,
lime or lemon juice, or with its own juice and arils. The esh pieces can also be
sprinkled with sugar, orange liquor and orange juice, and eaten as a fruit salad.
The arils can be eaten with the seeds.
Young fruit can be cooked or steamed and eaten as a vegetable. The roots
of old vines are eaten in Jamaica as a substitute for yams. The leaves, skin
and immature seeds contain a cyanogenic glucoside and the pulp contains
passiorine. In excess passiorine can cause somnolence, so in some places the
esh is used as a sedative to relieve asthma, diarrhea, headaches, neurasthenia
and insomnia. The leaf and root decoctions are used as a vermifuge (Morton,
1987).
FURTHER READING
Bora, P.S. and Narain, N. (1997) Passion fruit. In: Mitra, S.K. (ed.) Postharvest Physiology
and Storage of Tropical and Subtropical Fruits. CAB International, Wallingford, UK,
pp. 375386.
187
Martin, F.W. and Nakasone, H.Y. (1970) The edible species of Passiora. Economic Botany
24, 333343.
Menzel, C.M. and Simpson, D.R. (1994) Passion fruit. In: Schaffer, B. and Anderson,
P.C. (eds) Handbook of Environmental Physiology of Fruit Crops. Vol II. Subtropical and
Tropical Crops. CRC Press, Boca Raton, Florida, pp. 225241.
Menzel, C.M., Winks, C.W. and Simpson, D.R. (1989) Passion fruit in Queensland. 3.
Orchard management. Queensland Agricultural Journal 115, 155164.
Sao Jose, A.R., Ferreira, F.R. and Vaz, R.L. (1991) A Cultura do Maracuj no Brasil.
Fundao de Etudos e Pesquisas em Agronomia, Medicina Veterinaria e Zootecnica
(FUNEP). Jaboticabal, Brazil.
Schotsmans, W.C. and Fischer, G. (2011) Passion fruit (Passiora edulis Sim.). In: Yahia,
E.M. (ed) Postharvest Biology and Technology of Tropical and Subtropical Fruits,
Volume 4. Mangosteen to White Sapote. Woodhead Publishing Ltd, Cambridge,
pp. 125142.
Winks, C.W., Menzel, C.M. and Simpson, D.R. (1988) Passion fruit in Queensland. 2.
Botany and cultivars. Queensland Agricultural Journal 114, 217224.
Yockteng, R., dEeckenbrugge, G.C. and Souza-Chies, T.T. (2011) Passiora. In: Kole,
C. (ed.) Wild Crop Relatives: Genomic and Breeding Resources. Springer Berlin,
Heidelberg, pp. 129171.
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Abanto, A.M. and Muller, L.E. (1977a) Alterations produced in passion ower plants
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Abanto, A.M. and Muller, L.E. (1977b) Alterations produced in passion ower plants
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Akamine, E.K. and Girolami, G. (1959) Pollination and Fruit Set in the Yellow Passion
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Chapman, K.R., George, A.P. and Cull, B.W. (1978) A mechanical harvester for passion
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Correa, S.L., Ruggiero, C. and Olivera, J.C. (1979) Propagation of yellow passion fruit
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9
PALMS
The palms (family Palmae) have been classied into 200 genera in six
subfamilies (Uhl and Dranseld, 1988). The subfamilies are separated on
four major characteristics: leaf form; number of empty bracts subtending the
inorescence; ower arrangement; and the number and form of carpels in
the gynoecium. The fossil record attests to the diversity of palm morphology
and ecology, and shows palms to be the one of the oldest recognizable families
of monocotyledons. Most palms grow in non-seasonal or mildly seasonal
climates, with the rate of the phases of development determined by the
environment.
Numerous palms are used by humans, but only a small number have taken
on signicant commercial importance. Palms are regarded as multipurpose
trees, with their many products including include oil (e.g. Elaeis guineensis,
African oil palm; Cocos nucifera, coconut), waxes (e.g. Copernicia prunifera,
carnauba wax), ber (e.g. Cocos nucifera, coconut coir, Calamus spp., rattans),
food (e.g. Cocos nucifera, coconut; Phoenix dactyliferas, date; Bactris gasipaes,
peach palm or pejibaye; Salacca zalacca, salak; Nypa frutican, nipa palm, water
coconut; Arenga pinnata, sugar palm; Borassus abellifer, palmyra or toddy palm),
narcotics and medicines (e.g. Areca catechu, betel nut), thatch and ornamentals
(numerous species). In the food category, besides fruit, products include dried
foods, palm hearts (the edible portion of the growing terminal bud), seed, sap
(also fermented into wine and distilled into spirits), sugar and starch (sago).
This chapter discusses only two palms that provide fresh fruit: coconut and
salak. These are used as examples of the diversity of palms.

191
192
Chapter 9
COCONUT
Introduction
Coconut (Cocos nucifera Linn.) is also known as coco and cocotero (Spanish),
cocotier (French), kerala (India), tennai (Tamil), pol (Sinhala) and niu
(Hawaiian). Its fruit is used fresh at both the immature and mature stages.
The genus Cocos is monotypic, containing only the highly variable C. nucifera
Linn. Previously the genus contained over 30 other species that occurred in
Central and South America, but these have now been reassigned to several
other genera. No truly wild coconuts are known. The genus Cocos occurs
within the Arecoideae subfamily, tribe Coroeae, as do peach palm, oil palm and
betel nut.
The coconut palm is found throughout the tropics at low elevations. Two
centers of origin were originally considered: south-east AsiaMelanesia, and
Central and South America. South-east Asia has the greater diversity of types,
with numerous local names and uses. There are also unique insect and crab
associations in south-east AsiaMelanesia that attest to a long association.
Coconut cultivars on the Pacic coast of tropical America, from Mexico to
Peru, are indeed distinct from those on the coasts and islands of the Caribbean
and the Atlantic coasts of South America and West Africa, suggesting the
ends of eastward and westward movement. From the 16th century onwards,
Portuguese and later Spanish shipments and introductions could account for
coconuts in Africa, the Caribbean and the Atlantic coast of the Americas.
The closest botanical relatives of coconut (once classied as other Cocos
species) are found in South America, southern Africa and Madagascar, raising
the possibility that the true center 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. 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-Pacic. Subsequently, domestication in South-east Asia and the
south-west Pacic, followed by introgression of wild and domestic types, may
have led to human dissemination inland, upland and by boat to coastlines to
which the wild type could not oat. Coconuts possibly moved to New Guinea
and Polynesia with humans or by drifting.
Ecology
The numerous useful products obtained from the palm and its fruit have led
to its wide cultivation in the tropics. It is commercially grown in the warm
Palms
193
tropics along sea coasts with an adequate water supply, as well as inland,
between 23N and 23S latitudes.
Soil
Although the palm is grown on a wide range of soil types, it produces its
best yields on rich river alluvial deposits with good drainage. In most tropical
countries, it grows on beach sands with low nutrient levels. In these sandy
conditions, coconut requires higher land or fresh water swamps to carry
nutrients via percolation toward the beaches. Management of coconuts on
clay soil is difficult, as good drainage is essential. Soil pHs of acid clays (pH
5.0) to coral-derived sands (pH 8.0) are tolerated.
Climate
RAINFALL It takes about 44 months from ower primordium initiation to fruit
maturity, including the 12 months from anthesis to fruit maturity. Drought or

an extended dry season of 3 months in a row leads to inorescence abortion,
button shedding, premature nut fall and low nal nut yield (Fig. 9.1). Hence,
rainfall in the rst 3 months of nut development determines crop size 12
months later. The effects of prolonged drought can persist for up to 30 months.
Total rainfall between 1300 and 2300 mm/year is required for good production,
although the pattern of rainfall is more important than the total amount.
A mean temperature of 27C and diurnal variation of 67C is
considered optimum. These conditions are found on tropical sea coasts, where
the sea acts as a buffer against rapid temperature changes. The palm is killed
by frost and inorescence abortion occurs at low non-freezing temperatures
(<15C). Temperature is the deciding variable in the altitude of cultivation,
with a mean monthly temperature of 20C being the minimum. At the equator,
the limit for commercial production is 1000 m; at 18N (Jamaica), it is 150 m.
High maximum temperatures can also decrease yield.
TEMPERATURE
RADIATION Shade or very cloudy conditions leads to poor palm growth. The
inuence on yield has not been studied because of the long nut development
period. Approximately 2000 h/year of sunshine has been suggested as the
minimum.
WIND Coconuts can tolerate high winds if adequate soil moisture is available.
Windward trees may show some reduction in yield. Strong winds from
hurricane or cyclones can lead to tree death through being blown over or the
crown breaking off. However, windbreaks are not normally provided.
Chapter 9
194
No. Female Flowers/Spadix or %
Leaf No. or Days
Female
Flowers/Spadix
Month
Fig. 9.1. Effect of rainfall distribution on coconut leaf production and days between
leaves (A) and number of female owers per spadix, percentage spadices producing
fruit and percentage abortion (B) (Menon and Pandalai, 1957).
LIGHTNING Lightning can cause signicant damage and leads to disease at the
damage site. Producers in Malaysia and Sri Lanka both report lightning as a
major contributing factor to disease.
Stem and roots
Coconut trees can reach 2030 m in height and live for 80100 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 growing point leads to plant death. Branching of the stem has been
reported but is a rare event. Early and rapid stem growth occurs until fruiting,
after which the rate of stem growth declines. The stem gradually increases in
thickness during the initial years of growth. The stem diameter is maintained
Palms
195
until the tree reaches a height of about 10 m, and then gradually reduces. The
absence of a lateral cambium layer means there is no secondary thickening
and no capacity to repair injury. Stem strength is due to the brous sheath
surrounding the numerous vascular bundles in the stem periphery and a large
number of smaller bundle bers in the stem vascular bundles.
Like all monocotyledonous plants, coconut has an adventitious root system
with uniformly thick primary roots. The secondary and tertiary roots are also
uniform in thickness, with progressively smaller diameters. These roots are
produced from the base throughout the plants life. A 25-year-old tree can have
15004000 roots arising from the base of the stem.
The long-lived primary roots produce branch roots of about the same
thickness, and have short-lived rootlets. About 70% of the roots are found
within a 1 m radius of the stem to a depth of 0.10.5 m (Cintra et al., 1993).
A quarter of the roots grow vertically downward, while the majority spread
horizontally and can reach 20 m from the tree and about 2 m below the
surface. The initial roots of the germinating seedling can grow at a rate of 1
cm/day, although this rate slows after about 3 months of growth. Pruning of
roots induces root branching and continued growth.
Leaves (fronds)
The crown of fronds has about 3040 open leaves of 36 m in length. Around
1014 of these subtend fruit bunches at different stages of development
(Fig. 9.2). There are also 3050 unopened fronds in the crown. Leaves are
produced in succession at a rate of eight to 20 per year, and each takes 45
months to emerge from the sheath. The rate of leaf production is higher
in dwarf trees (17 per year) than tall trees (12 per year). Leaves survive for
33.5 years after fully opening. There is a gradual reduction in the rate of
leaf emergence, life and length after about 30 years, and therefore a lower
nut yield. Each leaf has 200250 parallel veined leaets that are 7080 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.
Inorescence and owers
This monoecious palm carries both male and female owers borne on an
axillary inorescence (spadix) of each leaf (Fig. 9.2). Dwarf forms may
begin owering in 3 years, and tall forms in 57 years. The rst inorescence
may be all male, with later inorescences producing both male and female
owers. The immature spadix is enclosed by a prophyll and peduncular bracts
(spathes). The spadix is 1.21.8 m long when mature and after emergence
from the bract. It is straw to orange colored and has a simply branched rachis.
Each branch (rachillae) bears one or more female owers near the base and
numerous male owers above. There are generally about 50 female owers
per spadix and possibly thousands of small male owers.
196
Chapter 9
Fig. 9.2. Male (A) and female (B) owers, tree (C) and germinating nuts (D) of
coconut. (With permission from Vozzo, J.A. [2002] Tropical Tree Seed Manual.
United States Department of Agriculture, Agriculture Handbook #721.)
The inorescence primordium is formed very soon after the leaf, although
little primordium growth occurs until the subtending leaf has nished its
expansion. The inorescence then begins to rapidly differentiate, followed by a
phase of elongation with the inorescence opening about 1213 months later.
The number of spadices produced depends on the rate of leaf production and
amount of spadices aborted. Spadix abortion is more common in young trees
and during drought. The number of female owers per spadix also varies with
variety, tree age, the season and tree management.
The male owers are small (3 mm) and non-symmetrical, with small sepals,
three longer petals and six stamens (Fig. 9.2A). The globose female owers
are about 2.5 cm long and 3 cm in diameter, with a reduced round perianth
surrounding the base (Fig. 9.2B). There is a short style with three stigmas.
Three ovules are produced, although usually only one is fertile.
Palms
197

The male owers begin opening at the distal end of the spadix during the
morning, falling in the afternoon. The whole process continues for upwards of
1 month on the same spadix. Flowers produce a sweet scent at anthesis and
nectar is produced from the pistillodes at the base of the male owers. The
female owers produce nectar as they open from the distal end, and anthesis
lasts from 615 days. Male ower anthesis has usually nished before the
female owers on the same spadix are receptive, forcing cross-pollination in
most cultivars. During the warmer months, however, inter-spadix pollination
on the same tree is possible in 4050% of trees. Some dwarf cultivars have
coinciding male and female phases, leading to variable outcrossing. The
presence of nectar and the sweet scent leads to considerable bee activity on
both male and female owers. Flies and other insects can also be involved in
pollination, and wind pollination occurs to a limited extent.
After anthesis, there is a period of immature nut (buttons) shedding,
mostly during the rst month. This shedding can vary from 55% to 95% of
female owers and is possibly caused by defective pollination, drought, disease,
insects or poor tree condition. A further shedding of more mature nuts occurs
before they are fully grown and before the endosperm has begun to form. This
shedding is often greater following a drought and is accentuated by heavy
rainfall following drought. A higher number of female ower and nut set is
sometimes observed after tapping for toddy (see later). In addition, dwarf
palms carry a heavy load of nuts when they rst come into production. This
last observation and biannual bearing suggests a role for assimilate supply and
demand in button-nut shed and immature nut fall. Several hormone sprays,
particularly 2, 4-D and coconut water, can double button set and increase nut
yield when applied after fertilization.
Fruit
The fruit is a brous drupe, called a nut (Fig. 9.2D). The mature nut is usually
ovoid and 2030 cm long, weighing 12 kg. The nut has a thin epidermis,
brous mesocarp (coir) and hard lignied endocarp (shell) that is brown when
mature. Wild types of nuts have three or more prominent longitudinal keels
that are absent in the more rounded commercial types. Inside the endocarp is
the white esh. This is 12 cm thick (endosperm) and encloses an opalescent
liquid endosperm (coconut water), which lls three-quarters of the large
central cavity at maturity. The rod-like embryo lies in the esh under one of
the three generative pores at the basal end of the nut. The embryo weighs
about 1/1000 of the fruit weight. Occasionally, there are two or three viable
embryos, one under each generative pore (eye).
The fruit take about 12 months to reach maturity (Fig. 9.3). The proportion
of dry matter partitioned to the fruit can be as high as 62% in this C3 plant.
198
Chapter 9
Time from Anthesis (Months)

Fig. 9.3. Coconut fruit growth and development from anthesis. (Jayaswriya and
Pandalai, 1957.)
About 32% of the endosperm is deposited in the rst 8 months and 94% by
month 11. The mesocarp comprises the major portion of the nut when young,
and increases in thickness and number of bers up to maturity. 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 part to develop. It begins as a liquid containing free
nucleic and some cells. These cells begin to coalesce toward 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
esh 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 (Fig. 9.4). Coconut
water is of cytoplasmic origin, but in mature coconut is free on any free
cells. As maturity approaches, the brous mesocarp begins to dry, becoming
reddish-brown. This dehydration and the shrinkage in the amount of liquid
endosperm mean that the mass of the nut declines from 34 kg at 9 months to
1.52 kg at 12 months.
199
Total Soluble
Solids
Total Soluble Solids (g/100 ml)
Dry Weight (g) or Volume Water (ml)
Palms
Endosperm
Dry Weight
Time from Anthesis (Months)

Fig. 9.4. Changes in coconut water volume and total soluble solids in relation to
husk and endosperm growth from anthesis. (Shivashankar, 1991.)
This very variable species has a 2n = 32. The use of the word variety may not
be appropriate, although a distinction is made between tall (typica) and dwarf
(nana) types. The tall types tend to be slower to begin owering and are mainly
outbreeders, while dwarf types are inbreeding (Table 9.1). The dwarf types are
more variable, showing several color variants in the petioles and fruit that are
not seen in the tall types. However, they are less productive and adaptable, 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, further
characterizing the different types (Batugal et al., 2009; Perera et al., 2010).
Of the population types listed (Table 9.1), the most interesting for fresh or
green coconut use is macapuno. Whereas the mature endosperm is normally
hard and compact around the periphery of the cavity, in the macapuno
type it is soft and curd-like, lling the entire cavity. The endosperm cells of
macapuno are large, multinucleate cells with low intercellular adhesion and
higher cytokinin activity than non-macapuno nuts. In addition, macapuno
endosperm has a lower hemicellulose content in the cell walls and a different
cell-wall sugar composition. A type similar to the Philippine macapuno has
Chapter 9
200
Table 9.1. Characteristics of dwarf and tall types of coconut, and example of
populations.
Height (m)
Leaf production per
year (n)
Juvenile period (years)
Pollination
Fruit
Disease resistance
Undesirable features
Populations
Dwarf type
Tall type
8
1618
25
About 12
1.53
Self-pollination
Smaller
Some resistance to lethal
yellows
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, MalayanMalaysia/Ivory Coast
Red, Malayan-Malaysia/
Ivory Coast
Cameron-Cameron,
West Africa
68
Regular bearing
Outcrossing
Large
Susceptible
Height for harvesting
Laguna, Philippines
Macapuno, Philippines
Tahiti, Tahiti
West African Tall, Ivory
Coast
Rennel, Solomon Islands
been reported in India as Thairu Thengai, in Sri Lanka as Dikiri, in Thailand

as Maphrao Kathi, in Indonesia as Korpyor and in Vietnam as Dua Dac
Ruot. The macapuno character is probably held on a single recessive gene,
as the homozygous trees need to be grown in isolation to ensure 100% yield
of macapuno. Dwarf types have been selected from this normally tall-type
variety. However, macapuno 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 variety (Nam Hom, smells like pandanas leaf)
in the dwarf group, which has been produced in Thailand and is now used for
export. The water in Nam Hom is not as sweet as that of another Thai dwarf
variety Nam Wan, which has 6.57% total soluble solids at the green stage. In
Sri Lanka, a variety Tambili (king coconut) is grown only for its sweet water.
Palms
201
Most breeding programs aim to increase the yield of copra per unit area
(Santos, 1986), with selections for the fresh market being made in Thailand
and Brazil. Yields can be increased by increasing the number of nuts per
tree (rather than amount of copra per nut) and improving earlier bearing
(possibly tied to number of leaves per year), tree vigor 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 poor storability, high outcrossing of tall
types leading to highly heterozygous offspring and the diverse environments
used for coconut production. Successful crossing between dwarf and tall types
has lead to the cultivars MAWA (Malayan yellow dwarf u West African tall)
and MAREN (Malayan red dwarf u Rennel tall). MAWA achieves higher
yields via early bearing and an increased number of nuts, although it is not
particularly adaptable and has a higher susceptibility to bud rot and smaller
nut size.
Cultural practices
Propagation
Ripe, large nuts with adequate liquid from selected mother trees are used
for propagation. The nuts should be handled carefully to avoid damage and
loss of seed viability, and not dropped to the ground. Planting can take place
immediately or be delayed, with the nuts stored for up to 4 months in a well
ventilated, cool, dry place. The nuts are often stored for about a month to
complete maturation before planting. Plantlets can also be obtained from
excised embryo after 56 months of culturing, avoiding the need to handle
large and heavy nuts.
The nuts are normally germinated in nurseries using a sandy soil, with the
use of polybags (450 u 450 mm) becoming more common. Polybags minimize
root disturbance during transplanting and reduce transplanting shock.
The nuts must initially be watered daily to assure uniform germination and
development. The epidermis is sometimes trimmed from the germ end of the
nut to facilitate water penetration and germination.
A shoot appears after several weeks. There is a relationship between the speed
of germination and vigor, and the productivity of the tree grown; therefore,
slow-germinating and less vigorous seedlings are culled. As germination
begins, a shoot grows toward the soft eye while the other end develops a spongy
haustorium into the seed cavity for nutrient absorption (Fig. 9.5). Coconuts
generally have adequate stored nutrients in the nut until transplanting,
although the endosperm is about 40% absorbed after 918 months. Fertilizers
can stimulate growth. The application of 50 g per plant of equal parts calcium
202
Chapter 9
Fig. 9.5. A germinating coconut, showing the endosperm and haustorium.
phosphate, potassium chloride and magnesium sulfate on two occasions is

recommended.
Transplanting can take place soon after the nuts have sprouted (46
months), and sometimes at up to 9 months. This should be scheduled so that
germination occurs in time for transplant at the start of the wet season.
The cleared land, which should not be subject to waterlogging, is commonly
planted in a triangular pattern, with rows running northsouth and 610 m
between plants. This gives the optimum number of trees per hectare. Wider
spacings (810 m) are used for tall types and narrower (68 m) for dwarf
types. At 9 m spacing, there are 160 trees/ha. Organic manure is frequently
added to the planting hole, which should be big enough to accept the plant
without signicant root disturbance. Mulch may then be put around the plant
after the hole has been relled. If irrigation is available, the seedling should be
watered as soon as possible to stimulate root development.
Palms
203
Irrigation
Irrigation is needed until the root system reaches the dry-season water table.
Trickle irrigation is recommended in the absence of a water table. Large
plantations are not normally irrigated, though nut and copra yields are
reduced by drought. The rate of water application varies with soil type, and
4050 l/week/plant on an Oxisol has been recommended in India during the
dry months. The mean transpiration rate of the mature coconut palm is about
7.5 g/cm2/s, and leads to a loss of about 90 l/day/tree.
Pruning
Pruning is not practiced. Since all parts of the tree can be used, however,
leaves are sometimes trimmed. This trimming can inuence nut yield (Fig.
9.6). The leaves intercept about 44% of the incident light, depending on
season and plant density. This light interception makes coconut suitable for
mixed cropping. The functional leaf area in a plantation is dependent more
on the intensity and duration of seasonal drought and less on plant density.
Insect damage, storm damage, pruning or clipping can lead to signicant
reductions in nut yield if more than 40% of the leaf area is lost (Fig. 9.6).
The reduction in nut yield is due to nut shedding and premature nut fall. The
impact of 50% leaf loss on yield can continue for 5 months, and the loss of
70% of leaves for 17 months after defoliation.
Fertilization
The lack of a uniform response of planting material, large areas required
for trials, difficulty in recording data on tall trees, seasonal effects and long
period before fertilizer effects are seen are major difficulties in fertilization
experiments (Murray, 1977). Nitrogen, potassium and magnesium have
been shown to signicantly inuence production. The amount of minerals
Fig. 9.6. Effect of leaf defoliation on the percentage average decline in mature
coconut production after 9 months of defoliation. (After Bailey et al., 1977.)
204
Chapter 9
exported in harvested nuts is greatest from the husk: 67% of the potassium
and 85% Cl. Leaf analysis data have suggested critical levels for the leaets of
the 14th leaf: nitrogen 1.82.0% dry mass, phosphorous 0.12%, potassium
0.81.0%, calcium 0.30.4% and magnesium 0.3% is recommended for
tall varieties; and 2.2% nitrogen, 0.12% phosphorous, 1.4% potassium and
0.2% magnesium for hybrids. Chloride has been shown to signicantly affect
growth, with a critical concentration of 0.50.6%.
Pest management
DISEASES Bud rot, basal stem rot and gray leaf blight can be important
diseases (Table 9.2). Other fungal diseases of coconut can cause losses, such as
leaf rot (Helminthosporium halodes) and lethal bole rot (Marasmiellus cocophilus).
The viroid diseases caused by single-stranded RNA such as cadang-cadang (the
Philippines) and tinangaya (Guam) can be tested for using molecular probes,
thus avoiding transmission of infected material. Lethal yellowing disease is
epidemic in parts of the Caribbean and West Africa, and related diseases have
been identied in Indonesia. 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 South-east Asia show high to medium levels of resistance to lethal
yellowing disease, as does the F1 hybrid Maypan. The website of the Centre for
Information on Coconut Lethal Yellowing makes current information on this
disease available online.
More than 700 species of insects associate with
coconut, of which 165 are peculiar to this tree. Only a few are serious pests.
Oryctes species, especially the rhinoceros beetle (O. 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
9.2), while other Rhynchophorus species cause damage by laying eggs in the
petiole and foliage. Scales insects (e.g. Aspidiotus destructor) attack the leaets
and cause loss of tree vigor. Leaf-eating and leaf-mining beetles (Brontispa and
Promecotheca) and caterpillars (Lepidoptera) 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 in
the Solomon Islands. The coconut eriophyid mite, Aceria guerreronis, which
seriously reduces fruit size, was rst 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.
In localized areas, monkeys, squirrels and rats will chew on nursery
seedlings, young palms and immature nuts; 5-month-old nuts are particularly
INSECTS AND OTHER PESTS
Table 9.2. Important diseases and nematodes of coconut. (After Ploetz, 1994; Nambiar and Rawther, 1993.)
Common name
Organism
Bud rot and nut fall Phytophthora palmivora,

other Phytophthora spp.
Basal stem rot
Gray leaf blight
Stem bleeding
Lethal yellowing
Red-ring disease
Heart leaves become chlorotic, wilted and easily

removed, extending to stem apex; affected nuts are
shed
Ganoderma spp.
Similar to drought, with gradual wilting leaf collapse
and slowed drooping growth
Pestalotiopsis palmarum
Small yellow to brown spots on the leaets and
rachises, coalescing into irregular necrotic areas
Chalara paradoxa
Infected roots; trunk pith decays when infected, leading
(Thielaviopsis paradoxa)
to slow tree decline, reddish-brown exudate from
trunk infection site
Viroid (RNA), lethal disease Slow infection; nuts become rounded and leaets ne;
(CCCVd)
non-necrotic translucent yellow spots, inorescence
becomes necrotic
A related viroid is found outside the Philippines, but
cadang-cadang is only recognized in the Philippines
Mycoplasma-like organism, Widespread; rst recognized in Jamaica in the 1800s;
plant hopper as vector
rapid spread and rapid palm death; the rst symptom
is nut drop after a latent phase and new inorescence
is black; yellowing of foliage from the oldest leaves
on tall types, brown in dwarf types; bud death, no
control, resistant varieties
Bursaphelenchus
Severe insect damage to crown and infecting the palm;
cocophilus (obligate
leaf yellowing, internal red-ring symptom about 70
parasite of many palms);
cm around the inoculation point of the stem
weevil as vector
Sanitation reduces the vector population
Distribution
India, Jamaica, the
Philippines, Indonesia,
Ivory Coast, Sri Lanka
India, Sri Lanka
All coconut-growing
areas
Indonesia, India
The Philippines
Palms
Cadang-cadang
Parts affected
Caribbean, West Africa,

Mexico, Texas
205
206
Chapter 9
preferred for their water content. Rats can cause heavy losses. 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 does so to
escape danger, rather than to feed off the nuts. The real relationship between
this land-living crab and coconut is to enable the crabs short-lived aquatic
larval stage to disperse long distances to other islands, by oating.
Weed management
Weed control is essential during plantation establishment. Failure to control
weeds in the 24 m area around each palm can lead to shorter leaf length and
fewer new leaves produced per year, fewer open bunches and fewer nuts set
per tree. The larger diameter (4 m) circle is more benecial for palms of 3950
months old.
The open canopy of coconut intercepts only about 4050% of the incident
light. The little change occurs in canopy spread with age and limited root
spread makes this an ideal tree for intercropping. Numerous crops are grown
under coconut: fodder grass and other pastures for grazing, coffee, cacao,
long-gong, duku, pineapple, banana, maize, mango, citrus, ginger, yams,
sweet potatoes, beans, peanuts and medicinal, aromatic and spice crops (black
pepper). Intercropping can increases the income of small growers, but this is
very dependent on the prot margin for each crop and selection of the most
protable combination. Cash crops are used among young coconut plantings,
while more long-term crops such as bananas can be planted in alternate rows.

Harvesting
For fresh consumption, the fruit is harvested at the immature stage (78
months) when the endosperm has begun to form and is jelly-like, before it
gradually thickens and hardens. The juicy, jelly-like endosperm of young nuts
is highly prized for eating out of the shell, for the water and for use in cooking.
The epidermis is uniform green in color and smooth, while the coir is white.
Green nuts are harvested approximately every 20 days. The nuts are cut using
a blade on a pole, or a climber goes to the top of the tree, cuts a bunch and
either carries or lowers the nuts to the ground with a rope to avoid damage.
Mechanical damage will cause the white coir to turn brown and can cause
nut cracking. Younger nuts have a lower rupture force than mature nuts
(108 versus 537 kg) (Tongdee, 1991). The nuts are held in a cool place until
processed or sold at market.
Mature coconuts (1011 months) are also used after hardening of the
endosperm, when they are green with brown patches to fully brown, and the
coir is also brown. The nuts can be collected from the ground if desiccation is
not a problem, but more often all nuts on one or more bunches, 8 months or
Palms
207
more old, are cut and collected at regular intervals throughout the year. Trained
monkeys are also used to collect the nuts. Mature nuts are light and bulky
compared to green nuts, which are up to three times denser. The endosperm
is scraped and squeezed to produce coconut milk; the scrapped endosperm is
preferred over the dehydrated product.
Unopened coconut spadices are tapped and the exudate (toddy) is collected
and fermented for an alcoholic brew (palm wine) of up to 1213% alcohol. It is
also distilled to produce a whiskey. Alternatively, the toddy can be 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 and 5.7% reducing
sugars. It can contain about 5% alcohol, depending on collection frequency
and hence the time allowed for fermentation.
The dehusked green fruit is about 10 cm in diameter and weighs around
500 g (100 g endosperm, 120 g shell and 250 g water) (Fig. 9.5). The liquid
endosperm in fresh green coconuts can contain 130620 ml water and 48
g/kg sugar, depending on the stage of harvest, and is at a maximum at 78
months after anthesis. In Thailand, the green nut is trimmed and shaped,
removing most of the husk. The nal product is at bottomed with a round
body and pyramid top, and the eyes showing (Fig. 9.7A). To prevent browning
of the remaining husk, the nuts are dipped in 13% sodium metabisulte for
Fig. 9.7. Processed coconut. A shaped and wrapped immature nut for eating the
jelly-like endosperm and coconut water (A), and processed and bottled macapuno
endosperm (B).
208
Chapter 9
25 min and then wrapped in plastic lm. Fungicide is sometimes included

in the sulte solution. Alternatively, the husk is completely removed before
dipping in the sulte solution. Coconut water has a maximum of 6% soluble
solids, 89 months after fertilization. Immature nuts can be stored at 12C for
4 weeks; the skin turns brown at lower temperatures.
Mature nuts enter the fresh market chain dehusked. The dry nuts should
be brown, free from ber, damage and cracks, and of the required weight or
size (3545 cm circumference). The nuts are shipped in sacks or cardboard
cartons. Postharvest stress cracks are directly related to coconut weight loss.
Waxing and wrapping the nuts minimizes water loss and dramatically reduces
fruit cracking. The shelf life is 23 months at 12C, before the liquid endosperm
evaporates or the shell cracks because of desiccation. Low humidity and high
temperatures are to be avoided.
Marketing
Humans have used the coconut palm for thousands of years for food, thatch,
ber and wood, and it has entered commercial trade as oil for lamps, candles
and soap. Most worldwide trade is based on dried copra (endosperm): the oil is
extracted from copra for soap, detergents and margarine; dried grated copra
is used in confectionaries; and ber from the husk is used for ropes, mats
and potting mixture ller. More recently, the residual coir dust (cocopeat)
is replacing peat moss for environmentally sustainable horticultural seedsowing and potting mixtures.
Coconut fruit is used fresh at both the immature and mature stages. The
fresh kernel (28% whole nut) contains 3544% oil, 3562% water, 911%
carbohydrate and about 0.01 g/kg vitamin C, as well as vitamins B1, B2 and B3
(Table 9.3). The juicy jelly endosperm in the green nuts, especially macapuno,
is highly prized for eating out of the shell, for the water and for use in cooking
(Fig. 9.7). The spongy haustorium from germinating nuts (Fig. 9.5) is eaten as
a vegetable delicacy in Sri Lanka. The kernel (solid endosperm) from mature
coconuts can be shredded and dehydrated to desiccated coconut. Coconut water
is now available internationally in sterilized long-life packs. Fresh coconut
water is an important adjunct in media for tissue culture.
The nut rst entered commercial trade for its oil, which was extracted from
the copra and used for lamps and for soap. The discovery in 1890 of vegetable
butter or margarine expanded the demand for copra (Woodroff, 1970; Ohler,
1999). Coconut oil becomes solid at temperatures below about 25C and was
used as vegetable butter in competition with other hydrogenation vegetable
oil, a process developed in the 1890s. The use of glycerin, a byproduct of soap
manufacture, in making nitroglycerine expanded demand for copra during the
rst and second World Wars, when Japan took control of all major coconutproducing countries.
Numerous coconut recipes exist for its use in breads, waffles, cakes,
candies, cookies, pies, ice cream, soup and other main-course cooked dishes,
Palms
209
Table 9.3. Composition of 100 g edible portion of palm fruits (Leung et al., 1972;
Siong et al., 1988).
Coconut
Constituent
Proximate
Water (g)
Energy (kCal)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Carotene (mg)
Immature
Mature
Water
Toddy
Salak
fruit
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
80
77
0.7
0.1
18.4
0.4
0.6
11
1.1
42
257
51
13
1.8
83
340
16
24
0.3
18
130
5
Trace
Trace
109
34
8
0.3
9
168
6
3
Trace
Trace
0.1
0
29
0.02
0.01
Trace
Trace
7
0.05
0.03
0.8
Trace
4
0.04
0.03
0.6
0
4
0.2
0.008
2.4
0.05
and the health benets of dietary coconut oil are becoming better known. In
the Philippines, acetic acid bacteria produce a cartilaginous material from
coconut water called nata de coco, which is used in a number of desserts.
SALAK
Introduction
Salak is a small, very spiny tillering palm. It is widely cultivated in the wetter
parts of the Indo-Malay region and is found as an understory palm in Java
and southern Sumatra. Salacca zalacca (Gaertn.) Voss. is the correct name,
although S. edulis is still found in the literature (Mogea, 1982). It is most
commonly referred to as salak (Indonesia, Malaysia, Philippines), snake fruit
(English) and Yingan (Myanmar). The genus contains 21 species with four
varieties. Two other species, S. wallichiana Mar. (known as rakam) and S.
210
Chapter 9
rumphii Wall. ex Blume (called sala), are grown in Thailand. S. wallichiana is

shorter and more compact than S. zalacca. In Malaysia, S. zalacca, S. glabrescens
Griff. (called local salak) and another Indonesian-introduced species, S.
sumatrana Becc., are grown. Salak is normally grown for its fruit, but is also
planted as a hedge fence because of its dense tillering growth and spines.
Ecology
Soil
Free-draining soil (pH 67) with high organic matter is preferred. The shallow
rooting system does not stand ooding. It can be grown on sandy soils with
irrigation.
Climate
RAINFALL The palm thrives under humid tropical lowland conditions with
rainfall of 17003100 mm/year. The rainfall should be uniformly distributed

(200400 mm/month) with only a short dry season, otherwise irrigation
or a high water table is needed. The availability of water can determine the
seasonality of the fruiting pattern; uniform water availability provides regular
owering and fruiting.
A temperature range of 2232C is reported to be required.
Salak grows from sea level to 300500 m, depending on the distance from the
equator. Fruit yield and quality decline in cooler areas and higher elevations.
TEMPERATURE
RADIATION Salak is normally grown under shade (5070%), sometimes
intercropped with other tree crops such as mango, jackfruit, durian, rambutan,
mangosteen, 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.
Stem and leaves
This creeping and tillering palm has a short stem of up to 1.5 m (Fig. 9.8),
with very short internodes and shallow roots. It does form basal suckers,
but does not form large clumps. The trunk sends out roots when it comes
into contact with the soil. This palm grows rapidly, reaching 1.5 m within 4
years.
The feather-like pinnate leaves (7 m long) have gray to blackish, long, thin,
sharp spines on the petiole, mid-rib margins and leaets. The dark green leaets
Palms
211
Fig. 9.8. Salak palm and leaves (A), and male (B) and female inorescences and
fruit (C). (Plant drawing used with permission from Lemaga Biologi Nasional and Dr
Setijati Sastrapradja from Palem Indonesia 1978; inorescence drawing used with
permission from PROSEA Bogor, Indonesia.)
are 2070 cm long by 27 cm wide (Fig. 9.8). To ensure fruiting, 10 leaves per
plant are necessary for the Pondoh cultivar and 16 for Bali.
Inorescence and owers
The palm starts owering 34 years after sowing from seed and 23 years for
suckers. It may be productive for up to 50 years. Like other palms, a dry period
is not required to induce owering. This usually dioecious palm produces
an axillary stalked spadix that is initially enclosed by the spathe. The male
spadices (715 u 0.72 cm) occur in bunches of four to 12. The larger female
inorescences are borne on shorter spadices (710 cm long), with 1540
212
Chapter 9
spadices per inorescence and nine to 12 inorescences per year. The owers
are borne in pairs in the axils of the scales. The male ower has six stamens
borne on a reddish, tubular corolla with minute pistil lobes that shed pollen
in the early morning. Pollen is collected by covering the inorescences with
a paper bag, or cut inorescences or orets are dried in an oven at 35C. The
collected pollen can be stored for 612 months at 4C and used for pollination.
The tubular corolla of the female ower is yellowgreen on the outside and
dark-red on the inside. It has a triocular ovary with a short trid red style
and six staminodes. Inorescences development from emergence takes
8090 days. Stored pollen is diluted with talcum powder before pollination
of inorescences. Pollination can occur at any time of the day if there is no
moisture (dew) on the stigma. Wet pollen and stigma are susceptible to fungal
attacks.
This plant is normally cross-pollinated, although some cultivars (e.g.
monoecious Bali) are self-pollinated. Insects (weevils, other beetles) are
thought to be the natural pollinators, but hand pollination is practiced when
natural pollination is decient. The palm owers and sets fruit continuously,
although there are harvest peaks from June to July on the island of Bali and
during December to February in other parts of Indonesia. The December to
February harvest peak in Indonesia coincides with owering in the rst half of
the dry season after a smaller June to July harvest.
Fruit
Fruit reach maturity 57 months after pollination (Fig. 9.9). The 1540
tightly packed globose to ellipsoid drupes per spadix (about 50 g) are 57 u 5
cm in size, tapering to a point. The most noticeable feature is the numerous
yellow to brown and dark-brown united scales that end in a small spine
and cover the skin. The scales develop from the exocarp. The Thai Rakam
and Sala varieties are reddish-brown. There are usually three seeds with a
28 mm thick edible eshy sarcotesta per fruit (blackish nuts), containing a
white, somewhat translucent, homogenous endosperm.
Salak has a chromosome number of 2n = 28, with 11 metacentric
chromosomes and three sub-metacentrics. Indonesia has numerous cultivars,
which are distinguished by place of origin and cultivation, and fruit color and
taste (e.g. salak Bali, Suwaru, Condet, Padang Sidempuan, gula pasir,
pondoh, putih and gading). The variety Bali (S. zalacca [Gaertn.] Voss var.
amboinensis [Becc.] J.P. Mogea) is monoecious, bearing both hermaphroditic
Time from Pollination (Months)
213
Vitamin C (mg/100 g) and Sugars (%)
Acidity and Tannins (%)
Fruit and Flesh Weight (g)
Palms
Fig. 9.9. The growth and development of Pondoh salak from 3.5 months after
pollination. (After Sosrodihardjo, 1986.)
and male owers. Dioecious cultivars include Condet, Gading, Pondoh

and Suwaru. Cultivars vary in fruit taste, mesocarp texture, color of the esh
and rind, and place of production. Pondoh which is divided into Pondoh
super, Pondoh hitam and Pondoh manggala is a sweeter variety but
has only 52% edible esh, while others are 7080% edible. Bali, which has
about 80% edible esh, has the same sugar level (20%) as Pondoh but twice
the acid content (0.44% versus 0.23%). The larger Swaru fruit is sweet and
moist, but has poor keeping quality. Salak varieties have been released by the
governments in Indonesia and Malaysia. A cultivar often fails to perform well
outside of the region in which it was selected.
The dioecious nature of many cultivars leads to wide phenotypic variation,
hence vegetative propagation is used following selection of a suitable clone.
Seedless forms do occur and are preferred. Soft fruit spines, thick esh, sweet
and aromatic taste, and high yield are the commonly desired characteristics.
Thornless salak cultivars would be very useful. A thornless Salacca species
has been found in Thailand, but unfortunately has poor fruit quality and low
productivity. A number of related species are being investigated for potential
use.
214
Chapter 9
Cultural practices
Propagation
The palm can be propagated from seed, layering of suckers or stem cuttings.
Micropropagation procedures are being developed. Viability is quickly lost
after seeds are removed from the fruit, with germination falling from 55%
after 1 week to zero after 2 weeks. Seeds are planted directly into the eld or in
a nursery, then replanted 4 months later. Seeds should be planted into moist
conditions with an organic mulch covering. Plants from seeds can be selected
for males and females when owers are produced in 34 years. The percentage
of male plants is higher than 50%.
Vegetative propagation allows the preferential selection of female plants
with high yields and desirable fruit qualities. Layered trees and suckers fruit in
23 years. Layers and suckers should come from mature clumps that produce
three to four tillers each year, although mortality can be high. The thorns make
sucker removal difficult, so a split bamboo tube is used to encase the sucker and
pushed into the soil until rooting. After rooting has taken place, the bamboo
tube and sucker can be removed with less danger. The stems from mature plants
(710 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. Suckering of selected plants can be achieved by removing or killing the
apical meristem of the parent tree, leading to ve to 10 suckers in 23 months
that are separated 46 months later. Diesel fuel is recommended in Malaysia to
kill the parent tree meristem and induce suckering.
Field preparation and spacing
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:
26 m on a square, giving 20003000 plants/ha. Male plants, if necessary,
are planted at a rate of 220%, dispersed among the female plants. Young
palms require shade (approximately 25%) during the rst year. In South-east
Asia, salak is frequently intercropped with banana, durian, rubber, oil palm,
coconut and cocoa.
Fertilization and irrigation
Manure and compost, ammonium sulfate, urea, superphosphate and
potassium chloride have been trialed as fertilizers. Excess nitrogen is reported
to lead to strong vegetative growth that increases the risk of plants falling
over and developing large fruit with a poor postharvest life. The adequate
use of fertilizer may make shading less necessary than previously assumed.
Besides potassium, magnesium, sulfur, boron and zinc are reported to limit
the growth of Bali and Pondoh cultivars. A fertilizer rate per plant of 300
g ammonium sulfate, 38 g urea, 175 g KCl, 3.8 g borax and 3.4 g ZnSO4 has
Palms
215
been recommended, plus 200 g dolomite for Pondoh and 150 g dolomite for
Bali. Half is applied after harvest and the other half 30 days later, broadcast
around the plant at the outer canopy line.
Irrigation is necessary if the supercial root system does not reach the water
table. Dry spells of more than 10 days indicate a need for irrigation. Irrigation
during dry spells can lead to more even fruiting throughout the year. Plants
need 0.7 times the evaporation rate (44.5 mm/day) on a 6 u 6 m spacing:
about 100118 l/plant/day.
Pruning and fruit thinning
Basal suckers are removed so as not to reduce the yield of the mother palm.
Lateral shoots may be spared to grow into fruiting stems or for vegetative
propagation. Tall stems lose vitality; earth is pushed up around the stems to
rejuvenate them. 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.
Thinning to six to eight fruit per inorescence is practiced about 3 months
after owering to provide space for the remaining fruit. The supporting leaf is
sometimes pruned to allow fruit bunch development and the plant to be fertilized.
Pest management
A number of diseases have been reported (Table 9.4) in salak, but the
importance of each is unknown. Sanitation is practiced to reduce infection
pressure. The larvae of weevils (Omotemnus miniatocrinitus, O. serrirostris)
tunnel into the top of the palm and can cause severe damage. The weevil of
the Nodocnemis species, 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.
A layer of granular-looking esh adheres to the kernel in ripe fruit and is
referred to as masir. The cause is unknown. Fruit splitting can also occur in
fruit approaching maturity that receive excess rain after a short drought.
Table 9.4. Important diseases and disorders of salak.
Common name Organism
Fruit rot
Mycena spp.
Flower wilt
Fusarium spp., Marasmius

palmivora
Pestalotia spp.
Corticium salmonicolor
Thielaviopsis spp., Ceratocystis
paradoxa, Fusarium spp.,
Aspergillus spp.
Leaf spot
Pink disease
Fruit rot
Parts affected
Mycelium growth on fruit
branches
Flowers
Leaves
Plants and fruit
Fruit
216
Chapter 9
Weed management
Weed control is essential until the leaf canopy closes. Mechanical weed
control is normally practiced.

Harvest indices have not been created, although harvesting is delayed until
the astringency and acidity have reached a minimum (Fig. 9.9), about 160
days from owering. Fruit skin color starts to change from dark brown to
reddish-brown, and the esh from white to yellowish-white. The esh color of
the Thai varieties is yellowish-white when immature, changing to creamy or
yellowish-orange. Firmness and ease of fruit detachment have been used as
harvest indices. Seed color also changes to black to blackish-brown. Spadices
should be individually harvested at the optimum stage. Overripe fruit are
tasteless and have an off odor.
Fruit are harvested by cutting the fruit bunches. A mature palm can bear up
to 20 kg/year. In Indonesia, fruit are handled in bamboo baskets from the eld
and considerable losses occur from mechanical injury leading to spoilage. Fruit
can be washed in water or brushed with a dry brush. Good undamaged fruit
should be selected and graded. In Indonesia, fruit are graded on size: grade 1
has 1215 fruits/kg, grade 2 has 1517 fruits/kg, and smaller fruit are rejected.
ASEAN grades have been developed based on size, appearance and defects.
Fruit are shipped to market in 20-kg boxes and can be stored at 15C for up to 4
weeks. Lower storage temperatures (12.5C) lead to chilling injury in 14 days;
the symptoms are skin pitting and discoloration, and the esh can turn brown
and become soft. Fruit last for about 710 days at ambient temperatures.
The ripe fruit are normally consumed fresh, and there is an export market
for high-quality graded fruit in cartons. The edible part is non-brous and has
a sweet taste and crisp texture. It is yellowish-white to brown in color. The taste
has been compared to a combination of apple, banana and pineapple. The
aroma is caused by methyl esters of carboxylic acids, which give a sweet, slightly
sour, pungent odor. Methyl pentanoate is regarded as the major impact volatile,
with 2-methyl pentanoic, 3-methyl butanoic and an unknown responsible for
the sweaty odor. No terpenoids have been detected. Peeling can be tedious as
the small spines on the skin can cause itchiness. Salak is also made into candy,
pickled and dried. Fresh unripe fruit can be made into a salad. The fruit is also
canned in syrup and made into a drink. The esh can be minimally processed
with a plastic cover wrap or an edible coating and held at 510C for about 1
week. The seeds of Pondoh can be eaten. The fruit is low in vitamins and oil,
and is a good source of antioxidants (Table 9.3).
Palms
217
FURTHER READING
Centre for Information on Coconut Lethal Yellowing (CICLY). Available from http://
groups.yahoo.com/group/CICLY. Accessed 28 March 2011.
Foale, M. and Harries, H. (2011) (revised) Farm and forestry production and marketing
prole for coconut (Cocos nucifera). In: Elevitch, C.R. (ed.) Specialty Crops for Pacic
Island Agroforestry. Permanent Agriculture Resources (PAR), Holualoa, Hawaii.
http://agroforestry.net/scps
Gunn, B.F., Baudouin, L. and Olsen, K.M. (2011) Independent Origins of Cultivated
Coconut (Cocos nucifera L.) in the Old World Tropics. PLoS ONE 6(6): e21143.
Harries, H.C. (2001) The coconut palm (Cocos nucifera) pp. 321338. In: Last, F.T. (ed)
Tree Crop Ecosystems. Elsevier, The Netherlands.
Kusumo, S. (1995) Salak, a prideful fruit of Indonesia. Indonesian Agriculture Research
and Development Journal 17, 1923.
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
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559566.
10
OTHER AFRICAN FRUIT: TAMARIND,
MARULA AND ACKEE
TAMARIND
This well-known, adaptable, drought-tolerant fruit tree has many different
uses in food, medicine and industry. It is also used as a shade tree in many city
streets and parks. The tamarind tree is distributed throughout the tropics and
warm subtropics. It is frequently regarded as an ornamental with useful fruit,
and few commercial orchards exist.
Botany
Tamarind, Tamarindus indica L., is a member of the Fabaceae family. It has
many common names, including tamarinier, tamarin, tamarinier des Indes and
tamarindier (French), tamarinde (German), Indian date (English), mak kharm
(Thai), imli and ambli (Hindi), asam jawa (Malaysia), me (Vietnam), mkwaju
(Swahili) and tamarindo (Spanish and Portuguese). Despite being well known,
this tree has received little attention from researchers.
The genus Tamarindus contains only one species, which has two synonyms:
T. occidentalis Gaertn. and T. officinalis Hook. The common name is used for a
number of other unrelated plants, such as sweet tamarind (Inga spp.), Manila
tamarind (Pithecolobium dulce) from South America, velvet tamarind (Dialium
guineense) and Spanish or African tamarind (Vangueria madagascariensis), a
relative of coffee from Africa that produces round fruit with sweet, tamarindavored pulp.
The species origin is still in doubt. The most accepted theory is that it is from
Central and East Africa, including Madagascar. Central Africa is the most
223
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Chapter 10
likely origin, as it has the largest genetic diversity. From East Africa tamarind
was apparently taken to India, where traders took it to south-east Asia. The
alternative hypothesis is that it originated in south-east Asia and was then
taken to Africa by Portuguese traders; signicant genetic diversity is also
found in Asia. A third proposal is that it originated in India, where it derived
its name, tamere or tamar hindi (the date of India) and from which Linnaeus
took the botanic name T. indica. The fruit was known to the Greeks and the
Egyptians before the Common Era.
Two types of tamarind are found sweet and sour with the latter being
the most prized. It is estimated that the sour type comprises 95% of world
production. India is the main producer, at about 300,000 t/year. This
production is obtained by harvesting trees along roadsides and in windbreaks,
backyards and trees used for shade. Thailand is the other large Asian producer,
growing 30% of the sweet type in addition to the sour type. The sweet type is
gaining in popularity on world markets and commercial orchards are being
established in Thailand. Another important product in big demand is tamarind
kernel powder, with around 20,000 t/year produced in India. Mexico is the
largest producer in the Western hemisphere, where tamarind is extensively
grown in established orchards. Many other countries also produce tamarind,
but not on a large commercial scale.
Ecology
The tamarind is best adapted to semi-arid tropical and subtropical conditions.
It also performs well in many humid tropics, providing there is a dry season
that allows fruit ripening.
Soil
Tamarind can grow in a wide diversity of soil types, such as deep alluvial,
sandy, rocky and oolitic limestone. It will produce in poor and rocky soils with
little or no cultivation and also tolerates saline and sodic soils, although yields
are not as high as on deep, well-drained alluvial soil. Ideally soils should be
slightly acid.
Climate
RAINFALL Tamarind prefers rainfall in the range of 6001500 mm. Growth
does occur at 250 mm, but yields are low. In high rainfall areas (35004000
mm) the tree remains vegetative and does not ower. A dry period is very
important to stimulate heavy owering and high yields. Dry weather is also
needed during fruit ripening to avoid fruit spoilage.
Other African Fruit: Tamarind, Marula and Ackee
225
Young trees are very susceptible to frost, but mature trees can
withstand brief periods of light frost (2 to 3C) without serious injury. The
canopy will occasionally show some injury, but the tree will sprout from the
base. The tree grows well and owers in cooler areas, but fruit will not set or
develop. At the other extreme, adult trees can stand temperatures as high as
47C. The maximum annual temperature range is 3036C, with a minimum
of 918C. The tree is very sensitive to re.
TEMPERATURE
LIGHT This species should be planted in full sun.

WIND Strong, pliant branches and a deep, extensive root system anchor the
tree during strong winds and violent typhoons, and tamarind is regarded as
a hurricane-resistant tree (National Academy of Sciences, 1979). The plant
also tolerates salt spray and can be planted near seashores.
Tree
Tamarind is a long-lived, slow-growing and highly wind-resistant tree that
can reach 2530 m height with a canopy spread of 814 m in diameter
(Duarte, 2001). The tree is normally multi-stemmed, which sometimes results
in a poor tree form. The bark is scaly, dark gray to brown, strongly ssured and
rough. Blood-red gum exudes from the bole and branches when damaged.
The leaves are pinnate and 815 cm long, with 1020 pairs of 1.252.5 cm
long and 56 mm wide oblong leaets that close at night. The leaves may
abscise during long dry periods. The foliage is bright green and has a feathery
appearance, which in combination with the open branch structure makes this
a very good shade tree. The tree can produce fruit for more than 50 years and
live for 80100 years.
Flowers
Flowers occur on small drooping racemes (513 cm in length) that appear
normally along new branches. The owers are inconspicuous at just 22.5
cm in diameter (Fig. 10.1). The ower buds are often pink because of the outer
color of the four sepals that are shed as the ower opens. The four sepals, up
to 1.5 cm long, are unequal, ovate and pink, cream or pale yellow. Of the
ve petals, the posterior and lateral petals are large and showy and slightly
exceed the calyx (Fig. 10.1). The anterior petals are reduced to bristles. The
individual petals are oblong in shape and white, cream, pale yellow or pinkish
in color, streaked with red or orange. The owers can be bisexual, as well as
dichogamous and protogynous. Flower-bud development takes about 20 days
from the rst visible initiation.
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Fig. 10.1. Botanical drawing of Tamarindus indica (tamarind) showing a new

branch, leaves, ower raceme, owers, fruit and young seedlings. (From Vozzo et
al., 2002.)

Stigma receptivity is at its maximum on the day of ower opening. The peak
of anthesis takes place at 6 am the next day, with peak anther dehiscence
at 10.30 am. Flowers usually develop in the spring or summer, with
synchronous leaf growth. In some countries, such as Sri Lanka and India, two
owering periods have been reported.
As with other legume trees, bees seem to the main pollinating agents.
Honeybees are frequent visitors, attracted to collect nectar and pollen.
The tamarind appears to be preferentially cross-pollinated. Flowers are
227
hermaphroditic, but the stamens are shorter than the style and the stigma is
receptive 1 day before anthesis. Self-pollination results in poor fruit set. Fruitset rates range from 1% to 15%. Approximately 75% fruit set can be obtained
in controlled cross-pollination studies, with only 26% of fruit set from selfpollination.
Fruit
The fruit is a pendulous indehiscent pod that can be 720 cm long and 1.5
3.0 cm wide (Fig. 10.1). It can be straight, irregularly curved or sometimes
curved in a horseshoe shape. The skin is cinnamon to grayish-brown and
scaly, and normally has constrictions between the seeds. As the pod ripens it
becomes brittle and can be easily broken. The pod thickens as it matures and
the green pulp surrounding the seeds turns reddish-brown or brown. This
pulp dehydrates to a sticky paste with a few coarse stands of the vascular ber
and separates from the brittle pod wall. The pods may contain one to 12 large
reddish-brown or brown at seeds, with an irregular obovate or rhomboid
shape, that are embedded in the brown edible pulp. Indian types often have
longer pods with six to 12 seeds, while shorter west Indian types contain only
three to six seeds. The pods reach maturity about 7 months after fruit set, and
are fully ripe and dry around 1 month later. The pod loses half of its water
content during drying.
The genus Tamarindus contains only one species with 2n = 24 and a narrow
germplasm base (El-Siddig et al., 1999). Large phenotypic variation exists in
seed pods in different growing areas, with less variation in tree form. The pod
variation occurs in pod size (long or short), pulp taste (sour, sweet or bitter)
and ber content. Variation has also been reported in tolerance to drought,
wind, poor soil, waterlogging, high and low pH, and grazing. Recent work with
molecular markers has dened three major populations from West Africa, East
Africa and Cameroon (Diallo et al., 2008).
Breeding
No program of systematic breeding has been reported. Selections have been
made and provide a foundation for a breeding program. However, because of
the out-crossing nature they must be vegetatively propagated. The owers lend
themselves to controlled pollination. The main aims of breeding programs
should be to improve yield and quality, and to obtain varieties with fruit that
separate more readily from the tree to make harvesting easier.
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Chapter 10

Two major types of tamarind are recognized: sweet and sour. The cultivars
can be further differentiated by pulp color, which may be reddish or brown.
The red-colored pulp is considered superior because of its sweet taste. Indian
cultivars have more pulp content than those from Africa (Lefvre, 1971).
Cultivars
Most tamarind cultivars that are grown are unselected. They vary in fruit
size and degree of tartness. Producers in India mainly grow the sour types in
commercial orchards. The seedlings are not identical in their genotypes. They
grow very slowly, attaining full production after 1213 years, and become
very large.
Some of the major tamarind-growing countries have selected improved
cultivars from among the natural populations. Thailand has more than 50
sweet cultivars, which include Muen Chong, Sri Tong, Nam Pleung, Jae
Horn, Kun Sun, Kru Sen, Nazi Zad and Sri Chompoo. In the Philippines,
Cavite, Batangas, Bulacan, Laguna and Manila Sweet are selected sweet
varieties. In India, selections such as Prathisthan from Maharashtra and
Urigam from Tamil Nadu are well established. The most preferred cultivar is
Periyakulam 1, a clonal selection from a local sour variety that starts bearing
in the third year after grafting.
Cutural practices
Propagation
SEXUAL Tamarind is traditionally grown from seed. Seeds retain their
viability for at least 6 months when kept at ambient temperature in dry
conditions (Alix and Duarte, 1999). Viability can be extended for over a year
if the seeds are well dried, mixed with sand and kept in airtight containers,
and for several years if stored in airtight packs at 10C and 715% moisture
content. Seed germination usually begins within 7 days and can take up to 25
days to complete. Pretreatment of the seeds by soaking in cold water for 24
h can increase germination from 3070% to 80% (Coronel, 1991; Prins and
Magehembe, 1994) and scarifying or cutting the seed coat can increase this
further to 92% (Masano, 1994). Soaking the seeds in gibberellic acid for 24 h
results in taller seedlings with thicker stems at 5 months from sowing. Spraying
4-month-old seedlings with gibberellic acid results in signicant gains in height
and stem diameter after 4 months, but the stem diameter differences are small
at 15 months (Duarte et al., 2002).
Seedling growth is rapid, with the taproot growing 30 cm or more in the
rst 2 months (Fig. 10.1). Transplanting should occur when the seedlings are
at least 80 cm tall. Seed propagation results in signicant variability due to out-
229
crossing and can delay the time before bearing. Seedlings may take 78 years
to bear fruit and 1012 years to produce a good crop. Other selections start
bearing in 4 years.
Vegetative propagation must be used to reduce tree-to-tree
variability and ensure earlier bearing and uniformly high-quality fruit.
Cuttings, air layers, budding and grafting have been successfully used.
Propagation by softwood terminal cuttings has been developed and the
protocol standardized (Srivasuki et al., 1990). The cuttings showed a 94%
success rate with the use of indolebutyric acid. Both air- and ground-layering
methods are widely practiced, but are not as successful as grafting (Ramos,
1976). Air layering has been successful using peat moss as a substrate (Duarte
et al., 2002), with no signicant response to the use of indolebutyric acid. Cleft
grafting is one of the most successful methods; veneer grafting is also popular.
Patch and shield (T or inverted T) budding are also often used, although T
budding is more complicated because of the characteristics of tamarind bark.
Rootstocks for budding or grafting are propagated from seeds. After sowing,
preferably directly into nursery bags, the seedlings should reach 0.81.0 cm
in diameter at the base of the stem to allow grafting or budding with scions or
buds from selected plants.
ASEXUAL
Field preparation
For commercial plantings, the eld should be prepared as for any other fruit
tree. This operation should be done before the onset of the rainy season to be
able to transplant and take advantage of the rain.
Planting distances vary from 710 m between rows and plants in the
row for seedlings, to 45 m for smaller, asexually propagated material. At
transplanting, care has to be taken not to disturb the root ball and to eliminate
the bent portion of the main root. A mixture of old manure and topsoil should
be added to the bottom of the planting hole, sometimes together with 30 g
superphosphate, and covered with 35 cm of soil. Mulching the soil around
the planting hole will help conserve soil moisture and control weeds.
Irrigation
The plant is well adapted to semi-arid regions of the tropics and can withstand
drought conditions, though yields will be low. Young trees require adequate
soil moisture until they become established, although mature trees do well
without supplemental irrigation. Overwatering that results in waterlogged
soils should be avoided. In many arid regions, the plant will grow and produce
very well with articial irrigation. If no irrigation is available, the ideal is to
plant at the beginning of the rainy season and water the plants with buckets
in the dry season during the rst years until they are well established.
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Chapter 10
Pruning
Young trees should be formed so that about four well-spaced main scaffold
branches are allowed to grow. During the rst years very low, hanging or long
upright branches should be removed. Maintenance pruning consists mainly of
eliminating weak, damaged or dead material as well as shoots growing in the
wrong direction, and trying to create an open center to allow light penetration
and make harvesting easier. Failure to prune can result in overcrowding of the
canopy. The plants require very little attention after the overall tree scaffold
has been established.
Fertilization
According to Hughes (2008), tamarind trees fruit well with or without the
application of fertilizer because of their deep and extensive root system.
However, fertilization does improve yields and keeps the trees in a healthy
condition. It is recommended to apply 5060 g/tree of 16:20:0 and 100
120 g/tree of 12:12:12 at 12 months after transplanting, normally at the
beginning of the rainy season; a similar amount should be added toward
the end of the rainy season. The amount of fertilizer should be gradually
increased as trees grow. Plants that are starting to produce should be given
500600 g/tree of 12:12:12 twice a year while full-bearing trees should be
given 34 kg/tree/year of 12:12:12, split into two or three applications. In
alkaline soils, microelements such as zinc and iron may be required and can
be applied as foliar sprays. As with many leguminous species, tamarind roots
nodules are believed to x atmospheric nitrogen under appropriate conditions
(Quiniones, 1983; Ding et al., 1986).
Pest management
DISEASES Several diseases have been reported to infect tamarind. In the
nursery, mildew caused by Oidium species is a very common disease. Other

diseases include leaf spot, stem disease, root and wood rot, stem rot, trunk and
collar rot, stem canker and a bark parasite (Parrotta, 1990). Cercospora leaf
disease results in severe defoliation, while stony fruit disease, caused by the
fungal pathogen Pestalotia macrotricha, makes the fruit hard and stony with
brous structures (Hughes, 2008).
INSECTS A host of pests can attack tamarind in different parts of the world,
including mealybugs, beetles, bagworms, leaf-feeding caterpillars, aphids,

white ies, thrips, green locust, shot-hole borers and a variety of scales (often
spread by ants). Various weevils and borers can also infest the ripening pods or
stored fruit. The most serious insect pests are thrips, aphids, coccids and white
ies that attack new growth. Flowers, young fruit and seeds can be damaged by
caterpillars and some beetles. Fruit are also attacked during storage (Hughes,
231
2008). Nematodes such as Radopholus similis, Longidorus elongatus, Xiphinema

citri and Meloidogyne incognita can attack the roots.
In India, the larvae of the tamarind weevil (Caryedon serratus) attack the
fruit and seeds in the eld as well as the fruit and roasted seeds during storage
(Butani, 1978). This insect has also been reported in Puerto Rico (Timyan,
1996) and other places in Latin America (Vlez, 1972). In Florida and Hawaii,
Calandra linearis attacks the fruit and seeds and is considered a common pest
(National Academy of Sciences, 1979).
Weed management
With small trees, care has to be taken to leave a clean circle around the plant,
preferably covered with a mulch to keep the soil cool and reduce soil moisture
loss. Weeding of adult trees is carried out with brush cutters, mechanical
weeding machines, or machetes or by spraying herbicides.

Harvesting
The fruit develop during the rainy season and take 610 months to mature
(normally in late spring to early summer). The fruit start to dehydrate about
4550 days before they reach the ripe stage. Mature fruit can be left on the tree
for as long as 6 months to allow the moisture content to decline to 20% or less.
Fruit can be harvested half-ripe (malasebo stage) or when fully ripe. The
pulp of half-ripe fruit is yellowish-green and is normally used for avoring,
while ripe fruit are used for processing and fresh consumption. Half-ripe fruit
are harvested by being pulled individually. Ripe fruit have a brown shell and
the pulp is brown or reddish-brown and sticky. The fruit can be tapped with
the nger and a hollow sound will be heard if ripe. The shell is very brittle, and
care has to be taken not to damage it during harvesting. Most of the fruit is
harvested when fully ripe and the moisture content of the pulp is around 20%,
as fruit at this stage command better prices.
Individual fruit on the same tree mature at different times. This makes
selective harvesting necessary, and ripe fruit are often gathered from the ground
after shaking the branches. In humid climates ripe fruit are readily attacked by
beetles and fungi, so they should be harvested and stored under refrigeration.
Non-harvested fruit can remain on the tree until the next owering season if
there is no spoilage or other factors. Poles are not normally used for harvest as
they can damage developing new leaves and owers.
Postharvest treatment
A tamarind pod is composed of 3055% pulp, 1130% shell and bers, and
3340% seeds. Much of the tamarind harvest goes to processing factories
232
Chapter 10
where the pulp is separated from the bers and seed. The pulp is sometimes
made into balls and sun-dried for a week. In rural households, the pods are
dried in the sun or occasionally using small-scale dehydrators. After this, the
shells, bers and seeds are removed and the pulp is pressed and preserved
in large masses and sold by weight, although there are variations between
countries. It is possible to extract the pulp mechanically; a tamarind dehuller
with a hulling capacity of 500 kg/h has been developed in India. The huller
has an efficiency of 80% for large fruit and 58% for small fruit (Hughes,
2008). The most effective method of storing tamarind pulp is by mixing with
salt and storing in transparent containers. The pulp can be stored in a cool,
dry place for 36 months. Freshly harvested fruit can be stored for a few days
in a refrigerator or freezer, or for 46 months by packing in high-density
polyethylene bags and storing dry at less than 10C.

Isolated adult seedling trees can yield 150225 kg/tree/year. A more realistic
yield is 3050 kg/tree/year (Vargas et al., 1999), although the yield can be
higher if superior types are grown. The yield in a well-managed orchard with
good clones could be 1018 t/ha.
The tender pods can be eaten as a vegetable, cooked or pickled. Unripe
fruit are roasted and later eaten. Ripe tamarind is most often consumed as a
fresh or semi-processed fruit. It is marketed worldwide in sauces, syrups and
processed foods. The sticky pulp, which tastes both sweet and sour, contains
the richest natural source of tartaric acid (810%). The pulp has a pleasing
sweet or sour avor and is high in acid and sugar. It is also a rich source of
vitamin B and calcium (Table 10.1). This soft, succulent pulp is used as an item
of confectionery and an ingredient in curries, chutneys, preserves, pickles,
sherbets and beverages.
Ripe tamarind fruit have proven medicinal value. The American
pharmaceutical industry processes 100 t of tamarind pulp annually. The pulp
is used for many medicinal purposes including as a laxative, for fevers and
inammation, in a gargle for sore throats and mixed with salt as a liniment
for rheumatism. The pulp is also used as an astringent on skin infections. Its
effectiveness against scurvy is well documented, and it is a common ingredient
in blood sugar-reducing and cardiac medicines (von Maydell, 1986).
Tamarind seeds are gaining importance as a rich source of protein with
a favorable amino acid composition. The seeds also contain pectin, which
has been found to be superior to fruit pectin for making jellies, jams and
marmalades. Seed powder is used in the textile industry for sizing and nishing
cotton and jute. The leaves are used as fodder for cattle and goats. They are also
fed to certain species of silkworm in India and West Africa. Leaves and owers
are good mordants in dyeing.
233
Table 10.1. Composition of 100 g edible portion of tamarind ripe pulp

and young leaves. (After Morton, 1987.)
Constituent
Proximate
Water (g)
Energy (kJ)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Invert sugarsa (g)
Tartaric acid (mg)
Minerals
Calcium (mg)
Iron (mg)
Magnesium (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Copper (mg)
Sulfur (mg)
Chlorine (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A
Ripe pulp
Young leaves
28.252.0
115
3.1
0.1
67.4
5.6
2.9
3041
8.023.8
70.5
5.8
2.1
18.2
1.9
1.5
35170
1.310.9
54110
375
24
101
5.2
71
140
2.09
63
94
0.73.0
0.16
0.07
0.60.7
15.0 IU
3.0
0.24
0.17
4.1
250.0 Pg
Seeds contain approximately 63% starch, 1418% albuminoids and 4.56.5% of

a semi-drying oil.
The tamarind tree is used in agroforestry systems in many parts of the

tropics. The increasing integration of tamarind with other trees and crops
on farmland offers a strategy to minimize the risk of crop failure. Tamarind
is intercropped with annual crops and other tree species. This provides a
constant income from the farm. In addition to intercropping, tamarind trees
are particularly well adapted as windbreaks. The deep root system anchors the
trees during storms and the thick foliage prevents the growth of vegetation
below the tree, providing a natural break.
234
Chapter 10
MARULA
Botany
Marula, Sclerocarya birrea (A. Rich.) Hochs. (Anacardiaceae), is a very
important and well-known fruit tree of dryland Africa. The name comes
from the Greek sklers meaning hard and kryon meaning nut, in reference
to the stone inside the eshy fruit. The marula tree is one of the great trees
indigenous to southern Africa. It is venerated and preserved by Africans
for its reliable supply of edible fruit under drought conditions. The tree is
considered sacred among the people of the region where it grows and it plays
an important role in tribal life. Nicknamed the marriage tree, it is considered
a provider of vigor and fertility for those who marry under its branches. The
stones inside the fruit are sometimes dried and used to make necklaces that
symbolize love (Hall et al., 2002; National Research Council, 2008).
The best-known names are marula in southern Africa, sakoa in Madagascar,
mngongo in East Africa, nobiga in Burkina Faso and neighboring countries, birr
(hence birrea) in Senegal and homeid in Sudan. In English, it is called jelly plum,
cat thorn, morula, cider tree, marula and maroola nut or plum. It is also known
as maroela (Afrikaans), canhoeiro (Portuguese Mozambique), dania (Hausa),
mngongo (Swahili) mutsomo, mukwakwa, mushomo, muganu and mupfura
(Shona), morula (Tswana) and amaganu (Zulu) (Hall et al., 2002).
The genus Sclerocarya belongs to the Anacardiaceae, along with mango and
cashew. The genus has two species: S. birrea is the best known and most
important species of this genus; S. gillettii is from Kenya. Three subspecies of
S. birrea are recognized: subsp. birrea, which is found from Senegal to Ethiopia
and south to Tanzania; subsp. caffra, which is found from Tanzania to South
Africa and Madagascar; and subsp. multifoliolata, which is found in Tanzania
(Hall, 2008). The subspecies differ in their inorescences and compound
leaves. The best known is S. birrea, subsp. caffra.
Marula is considered a species of African origin, especially in the regions
south of the equator. It has been distributed to as many as 30 African
countries including some north of the equator, from Senegal on the Atlantic
seaboard (17W) and inland to Eritrea and Ethiopia (Hall, 2008). In eastern
Africa, the species reaches the Indian Ocean coast in Kenya but is absent from
the equatorial humid forest region. The latitudinal range is from about 17N
in Niger to 31S in South Africa. It was introduced into Madagascar in the
distant past and more recently into Israel to be tried as an alternative crop for
arid areas (Mizrachi and Nerd, 1996).
235
The distribution of marula throughout Africa has followed the Bantu

migrations, as it has been an important item in their diet for at least 10,000
years. Nuts have been found in Zimbabwean caves, with carved tools and piles
of shells in close proximity (Hutchings et al., 1996; Wynberg et al., 2002), and
it has been present in West Africa for more than 1000 years (Neumann et al.,
1998). Similar tools are still used to crack the stones and extract the kernels.
Marula is not commercially cultivated, even in Africa; exact yield gures are
therefore not available. There are no plantings outside Africa, although small
evaluation orchards have been established.
Ecology
The drought resistance of marula makes it ideally suited to Namibia,
Botswana, Zambia and Zimbabwe, from bushveld to woodlands, with heat,
harsh sunlight and difficult soil conditions. This woody tree is found in open
wooded grassland, and individual trees in these ecosystems tend to be well
separated and with full sun exposure. Fewer than ve trees with boles of 5 cm
or more in diameter are found per hectare (Nghitoolwa et al., 2003).
Soil
Marula is found on a wide range of deep, well-drained sites from sand to loamy
sand to sandy loams. It occurs most frequently on deep, well-drained sandy or
loamy soil and on heavier soils not subject to waterlogging. It grows on acid
soil with a pH of 4.75.5 and on alkaline basalt or dolerite soils. It has shown
a high tolerance to irrigation with brackish water (EC 32 dS/m).
Climate
RAINFALL Marula is found in arid to semi-arid areas with highly seasonal
rainfall. The mean annual total rainfall is 5001250 mm. The dry season
generally lasts for 47 months, with less than 50 mm mean precipitation. In
southern Africa, the plant is said to be best suited to 250800 mm of rainfall.
The roots store water, and the rainfall of the previous season is possibly more
important to the harvest than that of the current fruiting season (National
Research Council, 2008).
Marula grows and sets fruit in warm to hot subtropical to
tropical climates. Hall (2008) indicated that marula is mostly recorded at
elevations below 1700 m and mean annual temperatures higher than 19C.
The tree is very tolerant of high temperatures. No visible damage has been
observed with summer temperatures of 45C. It tolerates light frosts when
TEMPERATURE
236
Chapter 10
leaess in winter, although young trees are more susceptible to frost damage.
Mature trees are damaged by air temperatures of 4C (National Research
Council, 2008).
Tree
This single-stemmed, medium-sized, deciduous tree generally reaches 912 m
and sometimes up to 18 m. It has an extensive taproot and lateral root system.
The bark of adult trees is rough, with a gray or silvery mottled color and
roundish, reddish-brown scales that are initially lighter colored after leaf shed
(Hall, 2008). In comparison, the bark of young trees and shoots is smoother
and pale.
The crown is rounded, with older trees having a wide-spreading canopy. The
foliage is dense and concentrated at the ends of the branches. The leaves are
alternate and imparipinnate, mostly with six to 18 opposite, glabrous, oblong
to ovate leaets that are usually entire (Fig. 10.2). The leaets are 29 cm long
and gray-green in color, turning pale yellow prior to being shed. After leaet
shedding the top branches appear abnormally thick and erect, like upturned
ngers. The tree remains bare for several months of the year during the seasonal
dry period.
Flowers
The juvenile period is from 7 to 10 years. After 1520 years trees will produce
signicant quantities of fruit, with quantities increasing with age (Shackleton,
2002). The small, dark-red owers are unisexual (dioecious) and found in
fragrant clusters at the ends of branches (Hall, 2008). Reddish male owers
are seen in racemes that arise in the axils of new or recently shed leaves (Fig.
10.2). Each male ower has 1525 stamens 34 mm long, that surround a
eshy disc. The reddish female owers arise in spikes of one to four owers.
The owers have ve short sepals and ve petals that are 56 mm long. Each
female ower contains an ovary encircled by a disc, with 1525 staminodes.
The ovary is of two or three cells, each with a single ovule. Three short lateral
styles terminate in globular stigmas.
Although not very fragrant, the owers do produce nectar that attracts
insects, with bees being the main pollinators. The owers start opening early
in the morning and remain open until midday. The anthers release pollen over
2436 h. These pollen grains remain viable for at least 12 h and often for more
than 48 h. In the female owers, the stigmatic surfaces are receptive when
the owers open and remain so for up to 72 h. Secretion of nectar ceases on
fertilization (Hall, 2008).
237
Fig. 10.2. Botanical drawing of Sclerocarya birrea (marula), showing growth at

the stem apex with owers and fruit. (Used with kind permission of Meg Coates
Palgrave. From Trees of Central Africa, 1957.)
Fruit
The fruit is a drupe that is 34 cm in diameter when ripe, covered by a
thick, tough exocarp (Fig. 10.2). Under the exocarp is the brous, eshy
mesocarp, which adheres to a very hard, obovoid endocarp (stone). The
endocarp can have two or three compartments, each of which contains
a seed that is 1520 mm long and attened. The seeds make up a small
portion of endocarp stone (10%) and less than 2% of the whole fruit
weight. When ripe, the fruit has a light-yellow skin with white esh. The
endocarp, when dry, exposes the seeds by shedding two (sometimes three)
small circular plugs at one end.
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Chapter 10
Flowering occurs during the dry season (September to November) and the
fruit ripen 25 months later, depending on temperature and plant moisture
status. The fruit are dispersed by the start of the next rainy season (January to
March). Elephants are the main seed dispersers after eating the fruit.
The chromosome number is 2n = 26 (Paiva and Leitao, 1989). National
multilocation provenance trials have been conducted in eastern and southern
Africa to assess variations in marula germplasm of southern and East
African origin (Hall, 2008). Selection criteria include fruit larger than 100
g, sweetness, tartness, juice and vitamin C content. Nine lines have been
identied in South Africa based on esh and stone quality. These improved
clones produce fruit weighing up to 100 g, with a variety of skin colors. They
are already distributed in South Africa and neighboring nations.
Cultural practices
Propagation
SEXUAL Seed propagation is possible but not recommended, as only half
the plants will be female and it will take 710 years until rst owering to
determine this. The endocarp is generally soaked for 2448 h before sowing to
loosen the plugs that cover the germination openings. The rst seedlings should
appear within a week and the others may take up to a month. The seeds are
sown in a mixture of coarse sand and decomposed manure, and this substrate
is kept moist during germination. The seedlings should be 2540 cm tall after 3
months and ready to be taken to the eld.
ASEXUAL Vegetative propagation by cuttings is possible and ensures the same
sex as the mother plant. The cuttings should be large and thick, as thinner
cuttings do not root as well (Wynberg et al., 2002). Grafting has been reported
by Soloviev et al. (2004) using seedling rootstock, but this is difficult and is not
used on a large scale. Tissue culture is another possibility (Mollel and Goyvaerts,
2004).
Field preparation, transplanting and plant spacing
In the absence of commercial orchards, normal planting practices for an
agroforestry plant should be tested. There are no studies on plant spacing and
the plants are managed as forest trees.
239
Irrigation
Trees are usually grown with the seasonal rainfall pattern with no
supplemental irrigation, unless they are being grown in a garden or backyard
planting. Since owering follows a period of drought and leaf abscission,
supplemental irrigation might be useful to produce out-of-season fruit.
Pests
According to Hall (2008), fruit pests include a beetle (Carpophilus hemipterus),
a moth (Cryptophlebia leucotreta) and a y (Ceratitis cosyra). The main pests
include several loranthaceous mistletoes, among which Erianthemum dregei
and Pedistylis galpinii are strongly associated with the tree in South Africa
(Dzerefos et al., 2003). The impact of these has not been quantied.

Harvesting
Marula fruit naturally abscise spontaneously from the tree at the maturegreen stage. Ripening and skin color changes are completed over the next few
days. Fruit are harvested by picking the fallen fruit before they are eaten by
animals or the fruit rots. Rodents gnaw at the plugs to gain entry to the seeds.
Postharvest treatment
Marula fruit are climacteric. Fungal rots are a postharvest problem, possibly
caused by mechanical injury after abscission and falling to the ground.
Preliminary trials showed that storage at 4C can extend the life of the fruit
for 16 days and reduce disease incidence and severity (Weinert et al., 1990).
Fruit remain green and rm after 14 days of storage at 4C, while those kept
at 12C and 20C develop a yellow color and can be squeezed for juice (Nerd
and Mizrahi, 1993). However, fruit stored at 4C for 14 days can develop
chilling-injury symptoms when transferred to 20C, including brown spots
on the green skin and an off avor. Fruit can be stored at 12C to slow the
ripening rate.
Utilization
Every part of the tree is utilized for an incredible variety of domestic needs.
As a traditional food plant in Africa, marula has the potential to improve
nutrition, boost food security, foster rural development and support sustainable land care. A tree can produce around 6090 kg of fruit (20003000
fruit). Most of the production comes from trees in the wild. A few trees have
been identied that produce as many as 10,000 fruit.
Ripe fruit are succulent and tart. They are high in protein and oil, and rich in
Chapter 10
240
vitamin C (Table 10.2). The vitamin C content is about eight times that found in
an orange. The fruit has a strong and distinctive nutty avor (Wickens, 1995).
The seed kernel, with its subtle nutty avor, is rich in energy, phosphorus
and potassium (Table 10.2) and is an important emergency food. Crude fat
accounts for 5565% of the dry weight and protein accounts for 2035%. The
oil is poor as a source of vitamin E, but is noteworthy for its favorable saturated
to unsaturated fatty acid ratio and oxidative stability (Hall, 2008).
Marula fruit are commonly eaten fresh. On a small scale, the fruit is
processed into juices, jellies, chutneys and alcoholic drinks, especially in South
Africa, Namibia and Botswana (Hall, 2008). A liquor with a distinctive avor
has been exported from South Africa for around 20 years. Slightly green fruit
are collected and allowed to ripen to a creamy yellow color and processed for a
beer called Mukumbi. Whole endocarps are stored in bulk off the ground until
processed for oil and seed cake. The pale, yellowish-brown oil from the seed is
a delicious additive to meals and can be used as a skin-care product. The oil
contains antioxidants and oleic acid.
Table 10.2. Composition of 100 g edible portion of marula
pulp (Wehmeyer, 1966) and nut (Arnold et al., 1984).
Constituent
Proximate
Water (g)
Energy (kJ)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Magnesium (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Zinc (mg)
Copper (mg)
Vitamins
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin C (mg)
Pulp
Nut
91.7
247
0.5
0.1
7.01
0.5
0.2
4.0
2703
28.3
57.3
3.7
2.9
3.8
6.2
0.1
10.5
8.7
4.8
Trace
0.08
118
4.87
462
808
601
3.81
5.19
2.81
0.03
0.05
0.25
67.9
0.42
0.12
0.72
241
The fruit skin is used to make a drink that replaces coffee. The bark has
medicinal properties and is used in decoctions as malarial prophylaxis, for pain
relief and for scorpion or snake bites. The inner bark can be used to make rope and
the soft wood is good for carving. The leaves are chewed to help indigestion and to
treat heartburn. Gum exudates from the stem are mixed with water and soot to
make ink. The bark also yields a redbrown dye that is used in coloring traditional
craftware. The fruit infusion is used to bathe tick-infested livestock, since the fruit
is regarded as a potent insecticide. Other signicant uses are as utility timber and
livestock feed (fruit and foliage) (von Teichman, 1983; Hall, 2002).
ACKEE
Introduction
The ackee, Blighia sapida Koenig (syn. Cupania sapida Voigt.), belongs to the
Sapindaceae family. The fruit contains a potent poison that has to be removed
before eating. In many places it is more known for its poisonous properties than
for its useful aspects (Vargas et al., 1999). The shiny dark-green leaves and
beautiful, red-colored mature fruit make the ackee primarily an ornamental
tree. The fruit is consumed mainly in Jamaica, where commercial orchards of
more than 1000 ha exist. The eshy cream aril of the ripe fruit is eaten. It is
often boiled or fried, avoiding the pink tissue between the seeds and the arils.
The name ackee or akee apparently derives from the West African akyc. Other
common names include akee apple and vegetable brain. In Latin America, it is
called palo de seso or arbol de seso (Cuba), huevo vegetal (Panama, Colombia), fruto
de huevo (Guatemala), pera roja (Mexico), merey del diablo (Venezuela), bien me
sabe and quesito (Colombia), aki (Costa Rica) and seso vegetal (Central America).
It is known as castanha, castanheiro de Africa, huevo vegetal and castanha in
Portuguese, and elsewhere as arbre a fricasser or arbre fricass (Haiti), yeux de
crabe or ris de veau (Martinique), kaka or nzan (Ivory Coast), baha or nza (parts
of Nigeria) and akye, ishin and akyen (other parts of Africa).

The genus Blighia in the Sapindaceae family has four species, which are found
in tropical Africa from Guinea to Kenya. The genus is named after Captain
William Bligh of the HMS Bounty who brought samples of ackee back to
England in 1793. The fruit had apparently been taken to Jamaica from West
Africa, probably on slave ships sometime earlier. The ackee B. sapida Koenig is
a highly variable species found in tropical West Africa and Jamaica (Stair and
Sidrak, 1992).
242
Chapter 10

The ackee tree is native to the forests of the Ivory and Golden Coasts of
tropical West Africa. In Ghana, it is much used as an ornamental street tree.
In Jamaica, trees are frequently found in backyards, along fence-rows or in
small groves, especially on marginal land. The tree was also introduced into
Trinidad and Haiti, many of the West Indies islands and to the Atlantic coast
of Central America by Jamaican slaves eeing the island. However, it was
outlawed in Trinidad after causing some fatalities. It has further spread to
other parts of America such as Brazil and Florida. It has also been introduced
into India and the Philippines.
Ecology
Soil
The tree does well on soils ranging from fertile to stony, but seems to grow best
on fertile, slightly alkaline, well-drained soils at elevations of up to 900 m in
the tropics. It also does well in sands and oolitic limestone (Crane and Balerdi,
2011). Well-established ackee trees tolerate dry soil conditions, although fruit
production may be reduced. The tree is not ood tolerant and may decline
under ooded soil conditions.
Climate
The tree thrives in places with 15002000 mm annual rainfall. It
can withstand prolonged dry periods once established, but production will be
low.
RAINFALL
This species is adapted to tropical and warm subtropical

areas and can be found from sea level to 900 m in Jamaica and Honduras. It
is regarded as a forest climax tree in West Africa. Trees are moderately cold
tolerant and, if mature, may withstand brief periods at 3.3C.
TEMPERATURE
WIND Ackee trees can tolerate moderately windy areas. If pruned regularly to
limit tree size and open the canopy to wind movement then they can withstand
hurricane-force winds without toppling (Crane and Balerdi, 2011). High winds
will cause defoliation and desiccation.
243
Tree
The tree has a moderately dense round to oval crown and stiff branches. It is
a medium to large tree that can reach 824 m in height. The trees are very
variable in plant form and height, color, size and shape of capsules, quality
of arils, yield, and owering time and intensity. The leaves are alternate and
pinnate with short petioles and three to ve pairs of glossy green leaets,
620 cm long and up to 7 cm wide, with the upper leaves being the longest.
Leaets are opposite or sub-opposite, elliptical to oblongobovate and shortly
acuminate (Fig. 10.3). Although the foliage and bright-red fruit of ackee are
beautiful when planted in home gardens, it is not recommended unless the
owners know this tree and the risks involved with the improper handling of its
fruit (Cohen and Paull, 2008).
Fig. 10.3. Botanical drawing of ackee, Blighia sapida, showing leaves, male and
female owers, and fruit before and after opening. (Composite from USDA Forest
Service Handbook 449, Trees of Puerto Rico and the Virgin Islands, Volume 2. and
owers from Backhuys Biological Books [2006] Flore Analytique du Bnin.)
244
Chapter 10
Flowers
The owers of ackee are borne together in pendulous racemes (415 cm long)
from pseudo-terminal shoots with both male and hermaphrodite owers (Fig.
10.3). The proportion of male and hermaphrodite owers varies with each
owering, tree and climatic conditions (Stair and Sidrak, 1992). The fragrant
owers are small (about 5 mm long). The calyx is ve- (or six-)parted and light
green; the ve petals are cream and a nectary disc is present at the base.
Flowering, pollination and fruit set
Ackee can ower throughout the year and may produce three or more crops
annually, with the heaviest bloom during spring. In subtropical areas, trees
probably respond to wetdry periods and cool temperatures, which also inhibit
vegetative growth. In Florida, owering occurs in the spring and fruiting in
midsummer, although there may be a light owering during the fall and fruit
during the winter. The major ower-inducing trigger in the tropics is probably
wet weather after a pronounced dry period. During the dry period the trees
vegetative growth slows or stops, allowing the buds in the leaf axils to mature
and form owers. Subsequently, under natural conditions, the tree owers
when the rainy season begins. With two main rainy seasons in the Caribbean,
there are two distinct fruiting peaks: January to March and July to September.
Considerable variability occurs in the time and extent of owering between
trees, even at the same site. A period of regeneration seems necessary before
owering recurs on a particular branch. Ackee owers are easily crosspollinated. The main pollinators are insects, especially bees (Suah, 1975) and
the wind.
Fruit
The pear-shaped fruit is a eshy, dehiscent, usually three-lobed (sometimes
one-, two- or four-lobed) capsule (70150 g) with a leathery skin. It
measures 610 cm in length (Fig. 10.3). Initially green, it turns to a red or
to a yelloworange color at maturity. The time from anthesis to maturity
is about 5060 days. The fruit doubles in size during the rst 23 weeks;
fruit drop can be very high during this period. Approximately 5.6% of
hermaphrodite owers set fruit and 44% of fruit mature. At maturity
the three lobes split longitudinally apart, with each showing a partially
embedded seed in a kidney-shaped, cream-colored, eshy, glossy aril (usually
3.04.5 cm long and 2.53.5 cm wide). Each aril is attached to a portion of
a black-colored seed. The seeds are about 1 cm in diameter, almost round,
hard, shiny and very poisonous. There are usually three seeds, although one
or two may abort. The aril is attached to the stem end of the capsule by a
pink or orange- to red-colored brous membrane (sometimes called a raphe)
that extends into the groove between the two lobes of the aril attaching it to
the placenta. The red tissue and veins that attach the aril to the seed and the
capsule must be removed before eating.
245
Cytogenetics, genetics and breeding
Little is known about the cytogenetics and genetics of this tree. Selection or
breeding programs for this semi-domesticated species should focus on traits
important for consumer needs (Eku et al., 2010). Preferred fruit traits
include size and shape, and aril color, size and taste. Some selections have
been made in Jamaica based on productivity, fruit quality and time of harvest.
A collection of ackee exists near St. Catherine, Jamaica, with two main
accessions types identied by the color of the aril. The type with a soft yellow
aril is known as Butter; and fruit with more esh around the seed, hard yet
with a smooth texture and cream-colored aril, is known as Cheese. Ease of
cross-pollination has produced types with intermediate characteristics (Royes
and Baccus, 1988). Vegetative propagation of selected, desirable types has
been attempted with limited success.
Cultural practices
Propagation
SEXUAL Traditionally ackee has been propagated by seeds that germinate
readily if sown fresh without any special pre-germination treatment (Vargas
et al., 1999). The seeds are short-lived, and should be sown a few days after
removal from the fruit. Seeds can take 23 months to germinate and seedlings
will bear fruit after 34 years. In Jamaica, the main producer of ackee, most
of the trees originate from seeds. This explains the large variability in tree
characteristics, fruit quality, production and seasonality found on that island.
ASEXUAL Cutting propagation has more recently been attempted. This has
produced very promising results and vegetative propagation is now encouraged

(Lindsay, 2000). The treatment of stem cuttings with rooting formulations
containing auxin may not be necessary as rooting occurs readily under good
conditions. Success has also been reported with grafting and air layering
(Wright, 1999). Superior types may be propagated by grafting (side-veneer or
cleft) or budding (patch) onto seedling rootstock. The tree begins to bear after
12 years with vegetative propagation.
The normal eld-preparation procedures as for any fruit tree are followed,
sub-soiling when necessary. Ackee trees should be planted on a square, 610
m apart. Trees planted too close to other trees or structures may not grow
normally or produce as much fruit because of shading or competition.
246
Chapter 10
Irrigation
Irrigation is not usually practiced. The ideal is to plant at the beginning of the
rainy season and to improve tree-establishment water using a bucket if no
irrigation system is available. In ackee trees more than 4 years old, irrigation
is benecial to plant growth and crop yields during prolonged dry periods.
The specic water requirements for mature trees have not been determined.
However, as with other tree crops, the period from bloom and through fruit
development is important. Drought stress should be avoided at this time, with
periodic watering to ensure proper yields.
Pruning
The tree exhibits vigorous growth and pruning is recommended. In Florida or
subtropical places with frost, pruning should be done soon after this danger
has passed. Pruning may reduce fruit production for one to several seasons.
Formation pruning should include topping the main stem at 8090 cm
when it reaches 11.2 m. The ideal is to keep three or a maximum of four
main branches as evenly distributed as possible around the main trunk and
at different heights so that the weight of the canopy is evenly distributed.
Selectively removing a few upper limbs back to their origins (crotches) each
year will help prevent the loss of the lower tree canopy due to shading by the
upper canopy. Topping the tree at 45 m is suggested to facilitate harvesting
and minimize wind damage. In addition, maintaining a small tree will
facilitate tree care and make spraying easier.
Fertilization
A generalized recommendation is for 450 g of 10:10:10, 4 weeks following
transplanting, followed by 400500 g of ammonium sulfate after 6 months
and 500 g 10:10:10 in year 2; the application rate should be increased by
450 g/year until the tree is 5 years old (Royes and Baccus, 1988). Specic
recommendations are not available and Jamaican orchards use a variety of
regimes (Williams, 1993). Ackee trees are susceptible to iron deciency under
alkaline soil conditions; this can be prevented or corrected by soil applications
of iron chelates. Periodic applications of ferrous (iron) sulfate may be made to
trees growing in low-pH soils (Crane and Balerdi, 2011).
Pest management
DISEASES Some diseases of ackee include leaf spots caused by Colletotrichum
and Cercospora, and sooty mold caused by Phaeosacchardinula anomala (Williams,

2000). Some trees are susceptible to fruit rot, which is associated with
Botryodiplodia species, Glomerella cingulata, Phomopsis species and Pyrenochaeta
species, and to Verticillium dieback (McMillan et al., 2002).
247
Although no major pests or diseases have been reported from Jamaica

(Lindsay, 2000), stem galls caused by an unknown agent are common and can
cause serious damage. Infestations of pustule scales, caused by Asterolecanium
pustulans, produce smaller galls on stems and petioles (Naylor, 1974). Mealy
bugs and black beetles (Macraspis tetradactyla) are reported and the fruit host
fruit ies (Williams, 2000). Plant parasitic nematodes have been recorded
(Dixon and Latta, 1965), but not as serious pests (Hutton et al., 1982).
INSECTS
Weed management
Manual weeding is normally practiced for young plants, while bush cutters
and herbicides are more commonly used in older plantations.

Harvesting
Ackee fruit production data are not available; however, observations suggest
that well-cared for mature trees can produce 4568 kg/year. Some reports
indicate that production can vary from 300 to 2000 fruit/tree/year (Vargas et
al., 1999).
Ackee fruit should be harvested after the lobules have split longitudinally.
It is important not to wait too long after this opening as the fruit will become
overripe and poisonous. Immature fruit and those that have not split open
naturally, or where the aril is soft or discolored, are extremely poisonous. Ripe
fruit can be separated by cutting with knives or pruning shears, but are usually
just pulled with the hand. In Jamaica, the main bearing seasons are between
January to March and June to August.
Fruit for home consumption are harvested after they have split open. Fruit for
canning are harvested while still closed and allowed to split in the holding area
of the canning plant. Fruit that do not split open within 3 days are discarded.
The arils are usually extracted, cleaned and consumed within 13 days,
keeping them under refrigeration. Canning is practically the only method of
prolonging the life of arils.
Utilization
The only edible part of the fruit, the aril, is eaten fried with seasoning or in
pastry. This delicacy is enjoyed by many at breakfast or as an entree. Ackee and
salt sh is regarded as a traditional dish in Jamaica. In West Africa, the aril is
eaten raw, fried or roasted. The fruit arils are a moderate source of calcium,
Chapter 10
248
iron, potassium and ascorbic acid (Table 10.3). They also possesses a small
amount of antioxidants (IC50 value = 6.6 g/ml). The major fatty acids observed
are linoleic (55% of the total), palmitic and stearic acids; ackee contributes to
the fatty acid intake of many Jamaicans (Crane and Balerdi, 2011).
Arils from unripe fruit have a high concentration of a heat-stable, watersoluble amino acid called hypoglycin A (D-E-amino methylenecyclopropylpropionic acid) in addition to hypoglycin B, a J-glutamyl derivative that is half
as toxic (Kean and Lewis, 1981). In children, consumption of unripe fruit can
induce vomiting, convulsions and coma within 648 h (Henry et al., 1998),
a condition referred to as Jamaican vomiting sickness syndrome (JVS). In
Cheese ackees, hypoglycin A decreases from about 8000 mg/kg in green arils
and seeds to 271 and 1451 mg/kg, respectively, in ripe fruit (Fig. 10.4), while
hypoglycin B levels in seeds increase from 1629 to 11,774 mg/kg. This strong
inverse relationship demonstrates that hypoglycin B in the seeds serves as a sink
for hypoglycin A from the ripening aril, and is involved in the detoxication
mechanism of the fruit during ripening (Bowen-Forbes and Minott, 2011).
Fruit that split naturally have safe levels of hypoglycin. It is recognized that
the nutritional status of the consumer is important to his or her propensity to
poisoning, since patients diagnosed with JVS generally show manifestations
of chronic malnutrition and vitamin deciency. Although JVS has resulted in
some fatalities in the past, the increased awareness of the importance of eating
only ripe, opened ackees means that such cases are now rare.
Table 10.3. Composition of 100 g
edible portion of raw ackee arils. (After
Morton, 1987.)
Constituent
Proximate
Water (g)
Protein (g)
Fat (g)
Fiber (g)
Carbohydrate (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Aril
57.6
8.75
18.78
3.45
9.55
1.87
83
5.52
98
65
0.1
0.18
3.74
249
Half
Ope
n, A
ril
and
Seed
Visib
le
Although Jamaica is the traditional ackee producer, orchards have been

established in Costa Rica and Mexico and export has been reported from West
Africa (Wright, 2000). Fresh ackee is not generally exported but the arils are
canned or frozen. In preparation, the aril is usually rst cooked by boiling in
salt water and this water discarded. The canned product is exported to ethnic
markets worldwide and continues to be enjoyed by both visitors to the island and
Jamaicans residing overseas. In 1972, the US Food and Drug Administration
banned the import of the canned product unless safety concerns regarding
hypoglycin could be met. The ban was lifted in 2000, but importation of canned
ackee into the USA is subject to controls to ensure a safe product is imported.
The Jamaican External Trade Statistics for 2001 show that 1.74 million kg of
canned ackee was exported, mainly to Canada, the UK and the USA, with small
amounts going to other Caribbean territories, Europe and Japan. Frozen ackee
is also exported, but in much smaller quantities.
The hard and durable wood of the ackee is called achin in Ghana, and is used for
building and making oars and casks. The owers can be used to extract perfume.
The plant parts have medicinal uses in many places in Africa and America. These
include ingesting aril mixtures for dysentery and using seed extracts against
parasites. Topical treatments use ripe arils, crushed leaves and leaf, seed or bark
extracts for ulcers, conjunctivitis, headaches and severe pain (Lindsay, 2000).
Fig. 10.4. Changes in ackee seed and aril toxic compound, hypoglycin A, during
fruit ripening. (After Brown et al., 1992.)
250
Chapter 10
FURTHER READING
Tamarind
Diallo, B.O., Joly, I.H., Hossaert-McKey, M., McKey, D. and Chevallier, M.H. (2007)
Genetic diversity of Tamarindus indica populations: any clues on the origin from its
current distribution? African Journal of Biotechnology 6, 853860.
Gunasena, H.P.M. and Pushpakumara, D.K.N.G. (2007) Tamarind Tamarindus indica L.
In: Pushpakumara, D.K.N.G., Gunasena, H.P.M. and Singh, V.P. (eds) Underutilized
Fruit Trees in Sri Lanka. World Agroforestry Centre, South Asia Office, New
Delhi, India, pp. 352388. Available from: http://www.worldagroforestry.org/
downloads/publications/PDFs/BC07320.PDF. Accessed 20 September 2011.
Hughes, A. (2008) Tamarindus indica, Tamarind. In: Janick, J. and Paull, R.E. (eds)
Encyclopedia of Fruit and Nuts. CAB International, Wallingford, UK, pp. 400405.
Yahia, E.M. and Salih, N.K.-E. (2011) Tamarind (Tamarindus indica L.). In: Yahia, E.M.
(ed.) Postharvest Biology and Technology of Tropical and Subtropical Fruits, Volume 4.
Mangosteen to White Sapote. Woodhead Publishing Ltd, Cambridge, pp. 442457.
Marula
Hall, J.B. (2008) Sclerocarya birrea, marula. In: Janick, J. and Paull, R.E. (eds) Encyclopedia
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, 2741.
Ackee
Cohen, J.E. and Paull, R.E. (2008) Blighia sapida, ackee. In: Janick, J. and Paull, R.E. (eds)
Emanuel, M.A. and Benkeblia, N. (2011) Ackee fruit (Blighia sapida Konig). In: Yahia,
E.M. (ed.) Postharvest Biology and Technology of Tropical and Subtropical Fruits, Volume
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Vargas, O., Alix, C., Lobo, A.D. (authors), Duarte, O. and Sanchez, J. (technical reviewers)
(1999) Frutales y Condimentarias del Trpico Hmedo. CURLA; PDBL; AFE/
COHDEFOR; DICTA; SETCO; PROFORFITH, La Ceiba, Honduras, pp. 282283.
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Wehmeyer, A.S. (1966) The nutrient composition of some edible wild fruits found in
the Transvaal. South Africa Medical Journal, Supplement South African Journal of
Nutrition 104, 11021104.
Weinert, I.A.G., van Wyk, P.J. and Holtzhausen, L.C. (1990) Marula. In: Nagy, S., Shaw,
E. and Wardowski, W.F. (eds) Fruits of Tropical and Subtropical Origin; Composition,
Properties and Uses. Florida Science Source, Lake Alfred, Florida, pp. 88115.
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V8929E/v8929e06.htm. Accessed 11 October 2008.
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Wright, A. (2000) Marketing ackee challenge for the industry. JAGRIST (The Jamaican
Agriculturist) 12, 3842.
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property, rights and benet-sharing. Southern African Forestry Journal 196, 6777.
11
OTHER TROPICAL ASIAN AND
PACIFIC FRUIT
LANGSAT AND DUKU

Botany
The Meliaceae, or the mahogany family, has about 50 genera and 560 species
of mostly trees and shrubs, including many important timber and ornamental
trees. About six or seven species of this pantropical family native to India,
Malaysia, Indonesia, Borneo and the Philippines have eshy fruits. The best
known edible fruit species are langsat and duku (Lansium domesticum Jack)
and santol (Sandoricum koetjape [Burm.] Merrill). The genus Lansium has only
three members.
Areas of origin and distribution

Members of the langsat group of fruit (Lansium domesticum Jack) originated
in the area from peninsular Thailand to Borneo, where wild species are still
found. Various common names are used for the different members of the
group, including langsat (Malaysia), longkong (Thailand), duku and lanzones
(Philippines), duku trengganu (formerly duku langsat) and duku johore. Only
langsat and longkong are commercially grown in the main production areas
of South-east Asia. It has been distributed to most tropical areas, though
commercial production is limited to tropical Asia.
Ecology
Soil
Sandy loam that is slightly acid and has a high content of organic matter is
preferred. Good drainage is essential to avoid root rot. The tree does poorly on
clay soils that crack on drying.
255
256
Chapter 11
Climate
RAINFALL Longkong grows well under hot humid conditions with uniform
distribution of rain throughout the year. Optimum rainfall is 25003000 mm/

year with 150200 rainy days/year, except during the 12 months of owering.
Optimum temperature is in the range 2030C;
leaf burn is experienced at higher temperatures. Shading is necessary, and the
tree is therefore frequently intercropped. It can be found at elevations of up to
700 m in the tropics.
TEMPERATURE AND LIGHT
Tree
Both langsat, which is a rather slender, open tree, and duku, which has a
broad canopy with dense foliage, can grow to 30 m, although in cultivation
trees are managed to 510 m tall (Table 11.1). The bark is irregularly uted
and mottled gray and orange, and has a sticky milky sap. The alternate darkgreen leaves are glossy on the upper surface and dull underneath. The leaves
are 3050 cm long on petioles up to 7 cm long (Fig. 11.1). Langsat leaves are
faintly hairy underneath, while duku leaves are hairless.
Flowers
Langsat and duku bear many-owered racemes (100300 mm long),
sometimes in groups of two to ve on the trunk, and large branches (Fig.
11.1). The individual perfect white to pale yellow sessile owers (2030 per
raceme) are 12 mm wide and 5 mm long, with ve sepals and ve petals. The
stamens are united to form a eshy tube of 10 anthers.
NATURAL POLLINATION Flowers are perfect and cross-pollination is rare.
Langsat and duku ower-bud differentiation occurs early in the dry season
and owering occurs after the start of the monsoon rains in about 7 weeks.
In Thailand, longkong plants begin to form ower buds during April and June
and fruit can be harvested during September to November. In Malaysia, langsat
generally owers twice a year.
FLORAL INDUCTION AND FRUIT SET BY CHEMICALS Flowers can be induced
by the amount and timing of controlled irrigation. Watering is stopped for

915 days before the desired longkong owering period, and no irrigation is
alternated with periods of less watering to support continuous growth and
induce leaf wilting and yellowing, and nally abscission. Fallen leaves should
be removed to allow the soil underneath the tree canopy to dry. After leaf
Other Tropical Asian and Pacic Fruit
257
Table 11.1. Comparison of three types of Lansium domesticum.

Character
Longkong
Trunk
White-striped trunk,
Round, upright, little
roughness, no white
upright branches
stripe
Long, tapered, thin,
Large, thick, dark
shallow veins,
green, glossy, tip
hairy underneath,
oval and wavy,
sparse foliage
deep veins, smooth
underneath
Smaller than
Large, oval and
longkong and
compact raceme,
duku, ovoid, 20
1040 fruit,
fruit per raceme,
translucent and
white-translucent
hard esh, aroma,
esh, one or two
few or no seeds,
seeds per fruit
seeds bitter
Leaf
Fruit
Peel
Seed
Langsat
Thin, whiteyellow,
Thick, a little rough,
very short hairs
hairless, light(fuzzy), very
yellow when ripe,
smooth, soft skin
no latex, hard skin
Small, green, esh
Few large, oval,
sticks to seed,
yellow-greenish,
germinates with
fertile, germinate
only one seedling
with many seedlings
Duku
Knobs on trunk,
spreading domeshaped canopy
Long, tapered, thin
oval tip, shallow
veins, smooth
underneath, very
similar to coffee leaf
Round, larger than
longkong and
langsat, eight to 12
fruit per raceme,
less esh with
strong aroma, all
carpels have nonbitter seeds
Thick, light-yellow,
short hairs, rough
skin, easily peeled
Long, germinates with
only one seedling
abscission, the trees should be heavily watered to initiate a new leaf ush along
with ower buds.
Fruit
Duku fruit are round (4050 mm), while langsat are slightly ovoid (3050
mm). There are 1525 fruit per langsat raceme and four to 12 in duku (Table
11.1). The pale-green immature fruit with white latex ripen to a pale yellow,
often with brown blemishes. Langsat pericarp is thin with a sticky sap, while
duku has a thicker pericarp and no sap. The pericarp peels easily to reveal a
clear, white, translucent and juicy adhering aril. Both fruit types have ve
separate segments, with one to ve seeds in langsat and one or two in duku
(Fig. 11.1). The esh adheres to bitter seeds that are green and about 2 cm
long by 1.5 cm wide.
The fruit can develop parthenocarpically and the seed apomictically (Salma
and Razali, 1987). Langsat and duku parthenocarpic fruit take 180220 days
to develop from anthesis. Fruit development is slow for the rst 90120 days
from anthesis, after which the growth rate increases to maturity. During this
258
Chapter 11
Fig. 11.1. Leaf, ower and fruit of Lansium domesticum. (From Nakasone, H.Y. and
Paull, R.E. [1998] Tropical Fruits. CAB International, Wallingford, UK.)
nal stage, aril sugar increases while acids and phenols decrease. Longkong
fruit can be harvested when the skin has changed from light to dark yellow and
the peduncles and calyxes have dried. Longkong fruit must be harvested as soon
as they are ripe, otherwise they abscise and have a short shelf life.
The species (2n = 144) has been subject to little genetic or taxonomic study.
Appreciable diversity occurs in the wild and domesticated forms. Molecular
analysis of Malaysian accession gave three main clusters: one (mostly dokong
259
and langsat) possesses thin-skinned fruit, the second (mostly duku-langsat)

has thick fruit skin and the third has only one member.
The cultivated forms range from langsat to duku, with many forms in
between. Since the cultivated forms do not apparently form pollen because
of androecium degeneration, the wild type must seemingly produce viable
pollen to explain the heterozygosity. The intermediate types duku-langsat
(Malaysia), longkong (Thailand) and kokosan (Indonesia) are also variable,
with longkong being nearly seedless and having a brittle skin and soft aril.
Kokosan has hairy leaves and very compact fruit branches, and is dark yellow
with a sour aril. This heterozygosity has led to considerable problems in
describing cultivars. Langsat and duku have general characteristics that allow
classication (Table 11.1); intermediate forms are not as readily described.
Major cultivars
Longkong grown commercially in Thailand is divided into three types (Yaacob
and Bamroongrugsa, 1991): dried longkong, wet longkong and longkongkalamae. Dried longkong is preferred because the fruit is very sweet (17
19% total soluble solids) and has a pleasant aroma. The fruit has ve carpels
and the largest carpel has one seed. Ripe fruit have a dark-yellow skin and dry
and translucent esh. The ripe fruit of wet longkong have a light yellow skin
and are softer than dried longkong fruit. The peel is thin and tough and is not
easily peeled. The esh is dark and juicy, though not as sweet as that of dried
longkong (1618% total soluble solids). The ripe fruit of longkong-kalamae
has a whiteyellow, smooth, thin, soft skin with no latex. The esh is dry,
translucent and soft, with 1618% total soluble solids. The taste is a blend of
sweet and sour, and the fruit has fewer, smaller seeds.
Cultural practices
Longkong can be propagated sexually and asexually. However, only about 10
seeds from 100 fruit will germinate. Care is necessary during seed collection
to avoid mixing seeds from orchards that grow mixed longkong, langsat and
duku plants. The seeds are recalcitrant and quickly lose viability if dried. A
mixture of soil, manure and sand (2:1:1 vol./vol./vol.) is recommended for
germination.
Asexual propagation can be done by approach, cleft, whip or side-veneer
grafting with high success rates. Grafting is preferred during the rainy season
because plants have better growth under high relative humidity. Seedlings of
langsat and duku plants are preferred as rootstock because of their root systems
resistance to disease, resulting in faster growth of the scions. Longkong plants
260
Chapter 11
propagated from seed ower and fruit in 710 years, while asexual propagated
plants ower and fruit in 35 years.
Field preparation
The tree is often grown as a backyard plant or in mixed stands. No general
eld-preparation procedures are followed.
Spacing varies widely from 8 u 8 m to 12 u 12 m (70156 trees/ha), often
intercropped with companion trees such as coconut. Wider spacing is used for
the longkong types in Thailand.
Water stress should be avoided during owering and fruit development
to ensure good fruit size and yield. During these stages, trees should be
frequently watered to prevent fruit drop and reduced vegetative growth. A
drought during fruit maturation followed by rain causes fruit cracking and
abrasion. Maintaining irrigation allows uniform vegetative growth and ower
induction, and prevents alternate bearing, fruit cracking and abscission.
Pruning and thinning
Pruning can be divided into two periods. Longkong plants of 17 years old
that are not yet yielding are trained to an open shape. The lowest branches
of plants derived from asexual propagation should be removed 1 m above the
ground, after which the tree is allowed to grow freely.
Mature plants are pruned after harvest, which is normally at the start the
rainy season as the tree is entering a dormant period. If pruning is not performed
before the dormant period then the plants will be unable to accumulate sufficient
food reserves for the next owering and fruit growth cycle.
All owers in a raceme do not bloom at same time; therefore, fruit do not ripen
uniformly. Flower and fruit thinning must be performed when fruits are about 0.5
cm in diameter. Flower thinning is necessary to prevent small fruit and lower fruit
set in the following owering cycle. Flowers are thinned by hand or knife when
the inorescence is 510 cm long: imperfect owers are removed and one or two
perfect owers that are hanging down or horizontal are left per raceme. Racemes
in the axils of branches or on distal branches should be removed, while those on
the main trunk or proximal large branches should be kept as they frequently have
perfect owers. Branches of about 2.5 cm in diameter can support three to ve
racemes, while those of 3.5 cm in diameter can support 1015 racemes.
Fertilization
Young non-yielding trees respond to fertilization by producing new shoots and
branches and expanding the canopy size. Inorganic fertilizer is applied with
manure or compost three to six times per year. For bearing plants, fertilizers
261
are applied after harvest; general recommendations for NPK application are
unavailable.
Pest management
Diseases (Table 11.2) and insects (Table 11.3) can severely restrict growth
and development, resulting in lower yield and quality of longkong. Collar rot
(Stibella cinnabarina) of langsat and duku is controlled by copper treatments.
Table 11.2. Major diseases of longkong (Suanphiroch et al., 2003).

Common name
Organism
Pink disease (pink Corticium

salmonicolor
mold)
White disease
(white mold)
Unknown
Root and stem rot Phytophthora

palmivora
Algal leaf spot,

red rust
Cephaleuros
virescens
Sooty mold
Meliola sp.
Die back
Unknown
Fruit rot
Cylindrocladium
sp.
Parts affected and symptoms

Petioles, branches and trunks; infected
area is water-soaked and covered with
a white mycelia lesion that becomes
pinkish then turns pink
Branches; knobs are induced by insects
that attack under the bark; the mycelia
grows and spread to petioles and
leaves
Roots and lower part of the trunks;
infected young plants have watersoaked leaves; leaves abscise and plant
death occurs; brown xylem tissue
Leaf lesions (35 mm diameter) occur
on the upper surface; the lesions are a
mixed green, yellow, orange or brown
Infected fruit have dark lesions covered
with mycelia and spores
Leaves turn yellow and abscise in 23
weeks
Infected fruit have light-brown lesions
that become dark and pitted
Table 11.3. Major insect pests of longkong (Suanphiroch et al., 2003).

Common name
Organism
Parts affected
Bark-eating caterpillar
Decadarchis sp.
Hypatima sp.
Prasinoxena sp.
Cossus chloratus
Rastrococcus sp.
Aulacaspis sp.
Araecerus fasciculatus
Nasutitermes sp.
Bark of branches and trunks

Leaves
Leaves
Leaves
Roots and trunks
Mealy bug
Scale bug
Coffee beetle
Termite
262
Chapter 11
Root rot and pre- and postharvest fruit anthracnose (Colletotrichum gloeosporioides) can cause serious losses. The fruit are fruit-y hosts and are subject
to quarantine control in some countries.
Weed management
Control around the trunk is desirable. Any mulch used should not touch the
trunk.
Orchard protection
Windbreaks are essential in windy areas. Strong winds can break branches,
leading to ower and fruit abscission and wilting.

Harvesting
Maturity is judged by skin color (green to yellow) and the lack of latex. All
fruit on one raceme ripen within a short time, but not all racemes on one
tree will be ready to harvest at the same time. Fruit are harvested at the full
ripe stage, indicated by a change in the skin color from light to dark yellow,
dryness of the sepals and the peduncle losing most of its green color. The esh
is transparent when ripe. Fruit on bunches that have been subject to ower
and/or fruit thinning generally ripen together, and over a very short period.
Four to ve harvests are necessary to harvest a tree. It is essential to harvest as
soon as possible because overripe fruit will abscise from the peduncle. Fruit to
be shipped long distances are harvested when 7080% ripe to avoid excessive
fruit drop. Fruit should be picked when dry, as wet fruit can quickly become
moldy when packed wet.
The fruit raceme is normally sold intact at local markets, with abscised
individual fruit sold at a lower price. There are no international standards; fruit
are generally graded by size and color. They are normally sold in single-layer
berboard cartons (2.25 kg) with padding, and sometimes in trays with liners.
Langsat and duku require careful handling to avoid bruising and skin
discoloration. Abrasion and impact injury, water loss and chilling injury are
the three major postharvest disorders. Mechanical injury (abrasion, impact
and compression) leads to skin darkening and browning. Symptoms of chilling
injury include pitting and brown scalding of the skin. Skin browning occurs
rapidly if stored at temperatures of less than 12C. The recommended storage
temperature is 18C and 90% relative humidity, giving 21 days of postharvest
life (Suanphiroch et al., 2003). Holding in plastic bags (0.08 mm thick) reduces
weight loss but increases surface browning. Preliminary recommendations for
modied-atmosphere storage are 5% oxygen and 0% CO2.
263

World production
Cultivation occurs in the Philippines, Vietnam, Burma, India, Sri Lanka,
Australia, Surinam and Puerto Rico. Southern Thailand and Malaysia are
major production areas; langsat and duku are grown in Malaysia, while
longkong and langsat are grown in Thailand. Longkong fruit tastes sweeter,
has less latex and is regarded as more pleasant than langsat fruit. Therefore,
longkong fruit are two to three times more expensive than langsat fruit in
Thailand.
The yields of langsat and duku are highly variable. Longkong trees (10 years
old) can produce 4050 kg/tree, increasing to 80150 kg after 30 years. Other
data suggests 25 t/ha/year in the Philippines to 5.6 t/ha/year for the duku type
in Thailand.
Utilization
The fruit is easily peeled from the stem end and the esh is commonly eaten
out of hand or served as a dessert. Ripe longkong fruit have a unique aroma
and pleasant taste. Less mature fruit are acid. However, longkong fruit have
low nutritional value, especially in terms of vitamins A and C (Table 11.4).
The fresh peel contains 0.2% of a light-yellow volatile oil, a brown resin
and reducing acids. A dark, semi-liquid oleoresin composed of 0.17% volatile
oil and 22% resin is extracted from the dried peel. The resin is non-toxic and
administered to halt diarrhea and intestinal spasms. The dried peel can be
burned, with the aromatic smoke serving as a mosquito repellant; the smoke
is also inhaled by those with tuberculosis (Phantumas, 1998). The pulverized
seed is used as a poultice for scorpion stings. An astringent bark decoction,
sometimes combined with leaves, is taken as a treatment for dysentery and
malaria. The leaf juice is used as an eye drop to dispel inammation.
SANTOL
Botany
Santol, Sandoricum koetjape (Burm. f) Merrill (Meliaceae), is a well-known
tropical Asian fruit. It is probably native to South-east Asia and was
introduced into south Asia long ago. It has become naturalized in the
Philippines. The synonyms are Sandoricum indicum Cav. and S. nervosum
Blume. Other names for santol include kechapi and sentol (English), faux
mangostan (French), sentul, kecapi and ketuat (Indonesian), thitto (Burmese),
toongz (Laotian), sau (Vietnamese), sathon, matong, krathon and kathon (Thai),
katul and kantol (Filipino), kechapi (Singapore) and kechapi, setul, sentoi and
setia (Malayan).
Chapter 11
264
Table 11.4. Composition of 100 g edible portion of longkong, langsat and lanson
(Anon, 1987; Phantumas, 1998).
Constituent
Edible portion (%)
Proximate
Water (g)
Energy (kcal)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
Vitamin C (mg)
Langsat
Longkong
Lanson
41
82
166
0.9
0.1
15.3
0.3
157
0.9
0.2
12.2
86
230
0.9
0.3
12.1
0.4
0.5
5
0.7
35
19
1.1
25
28
12
0.3
30
142
2
0.04
0.08
15
24
0.04
0.07
0.1
1.4
0
1.7
Ecology
Soil
Santol can be successfully grown on a number of soil types. Well-drained soils
with a pH of 6.5 are preferred.
Climate
RAINFALL The drought-tolerant santol can survive and bear fruit in areas
with rainfall of just 800 mm/year. It does best in a wet monsoonal climate, as is
characteristic of South-east Asian countries.
The trees can withstand 40C but grow best at a mean
temperature of 22C. Santol is cold tolerant and is grown at altitudes of up to
1200 m in Java. Trees are reported to recover from severe damage caused by
frosts of 3C.
TEMPERATURE
265
LIGHT Santol is grown in full sun. It is sometimes used as a shade tree for other
crops.
Tree
Santol is a semi-deciduous tree with milky sap, a straight trunk and hairy
young branches and leaves. It reaches 1530 m in height. Asexually
propagated trees tend to be smaller with a more bushy habit of growth and
bear fruit earlier (35 years after planting). The spirally arranged leaves are
compound, with three leaets that are elliptic to oblong, 2025 cm long,
blunt at the base and pointed at the apex, with entire margins on long petioles
(Fig. 11.2). The leaves are glossy and green above and light-green below, and
change to red when about to fall.
Fig. 11.2. Leaf, ower and fruit of Sandoricum koetjape. (From Nakasone, H.Y. and
Paull, R.E. [1998] Tropical Fruits. CAB International, Wallingford, UK.)
266
Chapter 11
Flowers
The numerous owers are produced in panicles of 1020 cm in length, mostly
in the axils of young shoots (Fig. 11.2). The individual owers are 12 mm
wide and 5 mm long, with ve sepals and ve reexed petals. The petals are
larger than the sepals and cover the internal ower parts. The 10 stamens fuse
as a tubular structure. The style becomes elongated as the owers mature and
reach anthesis. The owers are slightly scented and pale green to greenishwhite in color.
Insect-pollinated ower sepals open simultaneously as the rst whorl of ve
segments at the periphery of the oral primordium, followed by the petals.
Flowers open between 6 and 8 pm, and the peak of anther dehiscence does
not overlap stigma receptivity. Self-incompatibility is the rule, with few owers
setting fruit. Under natural pollination, fruit sets of 0.38% and 1.0% occur in
Bangkok and Native varieties, respectively. Flowering lasts about 3 months
following a brief deciduous or partial leaf shed.
Fruit
The santol fruit is a golden-skinned berry (50100 mm across) that is rm,
rounded, somewhat attened and rather pubescent at maturity (Fig. 11.2).
The stout fruit peduncle attaches at a shallow cavity. The skin exudes a thin,
slightly milky juice when damaged. The skin can be smooth or wrinkled
and is green when immature. The outer eshy pericarp is thick, tough and
leathery, and sub-acid to acid. The aril is thin, white, juicy and translucent. It
is somewhat brous and is edible after removal of the skin. The aril surrounds
three to ve seeds, which are large (1520 mm long) and more or less
triangular in transverse section. The seeds have a thin brown covering.
The fruit take 56 months to mature. At maturity, individual fruit weigh
from 50 g (cv. Native) to more than 300 g (cv. Bangkok). In Thailand, quality
fruit are produced by regulating fruit load and wrapping young fruit (Yaacob
and Subhadrabandhu, 1995).
The two principal cultivars in the Philippines are the Native and Bangkok.
The Native santol is a diploid (2n = 22), while the Bangkok santol is a
tetraploid (2n = 44) (Ramirez, 1961). A cross between the two cultivars using
the Bangkok as the female parent may produce triploid (2n = 33) hybrids
that may in turn produce seedless fruit. It has been observed, however, that
fruit produced from this cross drop 2125 days after pollination (Pimentel,
1980).
267

Most Native santol trees bear small and sour fruit with relatively large
seeds in proportion to the edible portion. The leaves of Bangkok are larger
and darker than those of Native. The owers and fruit are also larger in
Bangkok. Bangkok has a superior eating quality compared with Native.
Major cultivars
Santol has previously been divided into yellow and red species (S. indicum,
yellow; S. nervosum, red), but the characteristics used to separate the types
are inconsistent. The yellow santols leaves turn yellow before abscission,
while the leaves of the red variety turn red. A number of santol cultivars
have been described. In Thailand, santol cultivars include Tep Ros, Peai
Fai, E-la, Tong Yib, Ta Chaum, Nim Nuan, Ma Tuuam, Tong Ta Wee,
Took Thong and Tab Tim (Wanichkul et al., 1996; Wattanayothin and
Tunterawattana, 1996). Bangkok santol in the Philippines is referred to as
Manila in Florida, which may be a Thai cultivar (Teparod or Kan Tong).
Kan Tong has also been referred to by Bautista (1994) as Bangkok.
Cultural practices
Santol seeds are cleaned by rubbing them with ne sand and washing with
tap water to remove the brous covering. Seeds are recalcitrant and rapidly
lose their viability. Seeds can be kept for 90 days if packed in moist sphagnum
moss, with fair percentage of germination. Germination takes 2023
days and the seedling has very rapid early growth. Seed propagation is not
recommended as the resulting plants may produce undesirable fruit.
Santol can be propagated by inarching or approach grafting, cleft grafting
and shield budding. In shield budding, non-petioled and fairly mature budwoods that possess fully developed buds about to begin active growth are used.
Seedling rootstocks should be about 12 years old with stems that are the
diameter of a pencil. Santol seedlings grow fast and may be used as rootstock
in a year or less. Propagation by air layering is slow, though fairly successful.
To separate the air layers from the mother plant takes about 56 months. Air
layering is suitable for small-scale propagation. Propagation from leaess stem
cuttings of 12 cm in diameter is slow (2964 days from planting) with a low
rooting percentage.
Field preparation
A eld that has been well prepared by deep plowing and several harrowings
is desirable. Trees are often planted at the start of the rainy season, if no
supplementary irrigation is available.
268
Chapter 11

Prior to planting, old leaves are removed and the remaining ones pruned to
half their size to reduce transpiration. Spacing varies from 8 u 8 m to 14 u 14
m (51156 trees/ha) with companion trees.
Newly planted and young santol trees require adequate water. Water stress
should be avoided during owering and fruit development to avoid abscission
and small fruit size.
Pruning
The trees are pruned to develop two or three well-spaced main branches,
from which lateral branches develop during the rst 23 years of vegetative
growth. The overall objective in pruning and training is a low and spreading
crown. Trees are rarely pruned once fruiting has started, except to remove
dead, crossed, diseased and infested branches.
Fertilization
Specic recommendations for NPK rates are not available. General
recommendations include the application of 100200 g/tree ammonium
sulfate 1 month after planting and an equal quantity 6 months later or just
before the rainy season ends. For bearing trees, about 500 g of a complete
fertilizer is applied per application. Full-grown trees may require at least 2 kg
of complete fertilizer per application. Mulching is recommended.
Pest management
Collar rot (Stibella cinnabarina) occurs, although pink disease (Corticium
salmonicolor) is more common (Table 11.5). In pink disease, leaves of infected
branches suddenly yellow and wilt, followed by masses of pinkish mycelium
developing over the infected areas. Pink disease can be controlled by sulfur
application at weekly intervals.
Phytophthora phaseoli Thaxter causes blight disease in seedlings. Leaves,
young shoots, stems and cotyledons show blighting that results in seedling
death. This disease is more severe during warm and wet weather. It can be
controlled by soil sterilization, sanitation and spraying with Bordeaux mixture.
Larvae of various species can lead to defoliation and damage to ower buds.
A gall-forming mite (Eriophyes sandorici) is a serious pest of santol. The mites
form galls on leaves, making the tree very unsightly. Severely infested trees
produce small fruit. In the Philippines, Bangkok santol is more susceptible to
the gall-forming mite than Native trees. Santol, with its thick rind, may have
some resistance to the Caribbean fruit y.
Weed management
Weeds need to be controlled after planting out. Mulching, plastic cloths and
hand weeding are used.
269
Table 11.5. Major diseases and insect pests of santol.

Common name
Organism
Parts affected and

symptoms
Country
Pink disease
Blight disease
Gall-forming mites
Scale insects
Corticium salmonicolor
Phytophthora phaseoli
Eriophyes sandorici
Leaves, twigs
Seedling
Leaves, shoots, fruits
Twigs, branches
Philippines
Philippines
Philippines
Philippines
Orchard protection
Windbreaks are needed in exposed situations to avoid broken branches and
bruised fruit.

Fruit are harvested when fully ripe or when fruit skin turns yellow or
yellowish-brown. Immature fruit do not ripen properly and have poor avor.
Fruit are usually harvested with a net attached to a long bamboo pole and
transferred to a bamboo basket. Santol fruit has a storage life of 3 weeks at 8C
and 8590% relative humidity.

World production
Santol is widely distributed as a backyard plant in tropical Asia and in semiwild second-growth forests. It is grown in association with other fruit trees
and very seldom grown in monoculture. Santol yield data are very scarce,
although experimental observations suggest annual yields of up to 14 t/ha.
In the Philippines, the 1991 census showed more than 1.6 million productive
trees (Falla, 1999). A fully grown tree may produce up to 22.5 t/ha (Ramos,
1972).
Utilization
The fruit taste can range from sweet to sour. The rind and pulp that does not
adhere to the seed are edible. Fruit are usually eaten fresh, particularly from
sweet-bearing trees, but can also be processed into candies, chutneys, jellies,
marmalades and preserves. In Bangkok, freshly peeled fruit is eaten with a
chili-based sauce. The pulp is fairly high in carbohydrates and fair in iron, but
decient in calcium (Table 11.6).
Santol has a number of medicinal uses. The pounded bark may be applied to
treat ringworm. Fresh leaf decoctions are used in baths to treat fever. Roots are
Chapter 11
270

portion of santol (Intengan et al., 1968;
Satarath et al., 1988).
Constituent
Santol
Edible esh
6168%
Proximate
Water (g)
Energy (kcal)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Magnesium (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
84.5
57
0.4
0.7
13.9
1.0
0.5
9
1.2
17
328
3
Ascorbic acid (mg)

Thiamine (mg)
14
0.05
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
0.03
0.9
14
used for diarrhea and spasm, either as an infusion or decoction. Santol is also
used as a general tonic and as a preventive after childbirth. In Indonesia, local
people use the stem bark against colic and leucorrhoea.
In the Philippines, the bark is used in tanning shing lines. The wood can
be used for construction purposes as the red timber is fairly hard. Santol is a
fast-growing species and has good potential to be developed as a landscaping or
ornamental shade tree.
WAX APPLE
Botany
Wax apple, Syzygium samarangense (Blume) Merrill and L.M. Perry
(Myrtaceae), is one member of a genus that includes a number of popular
species that are cultivated for their colorful, edible eshy fruit. The genus
271
Syzygium contains about 1000 species of trees and shrubs native to the Old
World tropics (Ellshoff et al., 1995). The name Syzygium is derived via Latin
from the Greek syzygos meaning yoked together, possibly referring to the
paired leaves.
The Myrtaceae is made up of about 80 genera and 3000 or more species,
which are native to tropical and subtropical areas worldwide and to
temperate Australia. The family has traditionally been divided into two
main groups: eshy fruited and dry fruited. Myrtoideae is a subfamily with
eshy fruits (berries or drupes), while members of Leptospermoideae have dry
fruit (Cronquist, 1981). The Myrtoideae are characterized by eshy fruit and
opposite leaves. The largest genera are Eugenia, Myrcia, Psidium and Syzygium.
The Syzygium species are cultivated for their economically important
edible eshy fruit. Species include S. aqueum, water apple, a tree native to India
through Malaysia; S. cumini, Java plum, a tree native to India, Sri Lanka and
Malaysia; S. jumbos, rose apple, a tree that probably originated in Malaysia and
possibly South-east Asia; S. malaccense, mountain apple, and the more popular
S. samarangense, most commonly called wax apple, are trees with a native
range from Malaysia to South-east Asia. In addition, there are numerous other
Syzygium species with edible eshy fruit but of no economic importance.
The synonyms for S. samarangense (Blume) Merr. and L.M. Perry are: S.
javanicum Miq., Eugenia javanica Lam., E. alba Roxb., E. formosa Wall. and
Myrtus samarangense Blume. The common names are Java apple, Java rose
apple, Samarang rose apple, water apple, wax jambu, wax apple (English), jin
shan pu tao, lian wu, yang pu tao (Chinese) and nan yang pu tao (Hong Kong),
Curacaose appel (Dutch), pomme deau de formose and pomme de Java (French),
Java-apfel (German), amrool, jamrool, jamrul and jumrool (Hindi), renbu
(Japanese), jambu air mawar, jambu ayer rhio and jambu klampok (Malay), jambu
semarang (Indonesia), cajuil de solim (Dominican Republic), cashu di surinam and
makopa (Costa Rica), manzana de Java and mara n de Curacao (Panama), makopa
(Tagalog), chom phuu, chom phuu kaem maem, chomphuu kao, chomphuu khieo
and chomphuu nak (Thai) and man roi (Vietnamese).
Ecology
Soil
Good soils, rich in organic matter with loam to clay-loam texture, rather than
sandy and nutrient-decient soils, are recommended for better-quality fruit.
Climate
The tropical trees of the wax apple grow in the warm humid lowlands at
elevations of up to 1200 m, with no or light frost. They apparently need a dry
272
Chapter 11
season to ower. Low temperatures slow fruit growth and development of the
red color. In subtropical climates the fruit harvested in winter are generally
better to eat than those harvested in summer. The large leaf size and ease of
fruit bruising necessitate some protection from wind.
Tree
The tree is 515 m tall with a short trunk that is 2530 cm thick. The open,
wide-spreading crown has a pinkish-gray, aking bark. The opposite yellowish
to dark bluish-green leaves are nearly sessile and are ellipticoblong, rounded
or slightly cordate at the base, and 1025 cm long and 512 cm wide (Fig.
11.3). The leaves emit a very aromatic aroma when crushed.
Fig. 11.3. Leaf, ower and fruit of the wax apple, Syzygium samarangense.
(From Nakasone, H.Y. and Paull, R.E. [1998] Tropical Fruits. CAB International,
Wallingford, UK.)
273
Flowers
The inorescences are borne in drooping panicles of three to 30 owers at
the branch tips or in smaller clusters in the axils of fallen leaves. The owers
are fragrant and yellowish-white with four petals. They are about 3 cm across
with numerous stamens of about 2 cm in length (Fig. 11.3).
Flowering and fruiting of the Syzygium species is usually seasonal and
12 months in duration. It varies widely from place to place and even from
year to year. The normal production of wax apple is from May to July in
Taiwan and Hawaii, March to May in Sri Lanka and June to August in Java.
However, fruit can be harvested from one to three times a year, depending on
when conditions led to owering. Early owering is preferred for commercial
growers, although the formation of ower buds does not mean early owering.
Flowering can be triggered by a number of protocols (Fig. 11.4). These were
developed in Taiwan and are now being applied elsewhere in South-east Asia.
The protocols take into account that ower panicles are borne at the tips of
shoots and axils of fallen leaves. However, ower forcing is signicantly affected
by tree vigor, and eld and weather conditions.
1st Flush
Bald Cut and

Shading
1st Flush
2nd Flush
2nd Flush
Fig. 11.4. Wax apple production systems in Taiwan.
274
Chapter 11
The owering induction protocols (Fig. 11.4) do not induce more owers
than untreated trees. However, the owers that are induced appear earlier
and more uniformly over a short period of time. The ve protocols used for
fruit production are: (i) natural, (ii) simple ower forcing, (iii) improved ower
forcing, (iv) bald cut and (v) bald cut and shading.
NATURAL The original production system that most resembles natural growth
conditions. Fruit are harvested from May to July.

Simple ower forcing was developed in the late
1970s. It is similar to the natural protocol, but ower forcing is carried out by
pruning and/or with chemicals in October. Fruit are harvested from February
to July.
SIMPLE FLOWER FORCING
IMPROVED FLOWER FORCING Improved ower forcing was developed in the
early 1980s. The improved forcing protocol is used on fertile soils and is most
common in Taiwan. This protocol can increase fruit yield by 153%. Shading
of the tree canopy with nets (6095%) for 40 days produces 30 times more
inorescences and owers per tree than in control trees (Lai, 1997). Cultural
and/or chemical treatments are also used to retard vegetative growth. Cultural
practices, which include trunk girdling and/or root pruning, are performed
in late August. The trees are then treated with owering chemicals in early
September. Sprays include Cycocel (a gibberellin inhibitor), ethephon (ethylene
source) and 1-naphthaleneacetic acid to control tree vigor and induce
preformed ower buds to ower. Fruit are harvested from December to July.
BALD CUT This is a sophisticated and labor-intensive production system
that was developed by a farmer in 1985. The low fertility and high vegetative
growth of trees grown in sandy soils result in fruit after March that are inferior
in quality to those produced on loam or clay soils at the same time. Instead of
producing the low-quality, low-value fruit, farmers delay owering till August
or September and fruit are harvested in December to February, when trees
under the other protocols are not producing fruit.
In the bald cut protocol, trees are subjected to severe pruning. This involves
removing all owers, fruit, leaves and small branches, leaving only large limbs.
After this pruning, the tree is heavily fertilized with nitrogen and irrigated
heavily to induce new growth. Girdling, root pruning and shading are used
to impede vegetative growth in favor of ower formation in late June and July.
Flowering then occurs in late August or September. Failure to trigger owering
leads to at least a 2-month delay in owering. Cultural practices used to retard
vegetative growth, such as trunk girdling and/or root pruning, are performed
in late July or August, with the trees treated with chemicals in early September.
Fruit are harvested from December to February.
275
BALD CUT AND SHADING The cultural practices used are the same as in the
bald-cut system, with the addition of shading. Trees are shaded for 3040 days
to improve ower forcing.
Fruit
The waxy fruit are usually light-red to crimson, and sometimes creamy or
greenish-white in color. The pear-shaped fruit is 4 cm long and 5 cm wide. It is
narrow at the base and has a very broad, attened, indented shape at the apex,
and is adorned with the four eshy calyx lobes. The skin is very thin. The esh
is white, spongy, dry to juicy, subacid and very bland to very sweet in avor.
There may be one or two somewhat rounded 0.5 cm wide seeds, or none.
Fruit growth follows a single sigmoid curve and takes about 80 days from
anthesis. The position on the tree inuences growth and nal quality. Inner fruit
are smaller, redder and have a higher sugar content. Cytokinins and gibberellic
acid used either alone or in combination improve the size and quality of the
fruit. A synthetic cytokinin (N-(2-chloro-4-pyridyl)-N phenylurea) increases
fruit size by about 1.35-fold.
Varying gures are given for the 2n value of wax apple: 33, 42, 44, 66 and
88. S. aqueum has 2n = 44, S. cumini 2n = 66 and S. malaccense 2n = 22.
Major cultivars
Some superior clones have been selected, but breeding has apparently not been
carried out. In Thailand, there is a green-fruited cultivar called Hiew Savoey.
Taiwan has a number of clones based on fruit skin color: pink, deep red, light
red, white and green, with the pink variety occupying 95% of the planted
acreage (Table 11.7).
Cultural practices
Seeds lose their viability soon after removal from the fruit. Germination is
rapid if the seeds are planted immediately. Preferred trees are readily obtained
by air layering or cutting. Trees can also be obtained by budding or tissue
culture. Budding onto S. pycnanthum Merrill and L.M. Perry (wild rose apple)
provides some protection, as it is resistant to termites. The trees require little
attention during the juvenile period of from 37 years, with air-layered trees
setting fruit in 35 years.
276
Table 11.7. Fruit characteristics of wax apple cultivars grown in Taiwan.

Pink mutants
Light red
Dark red
Green
White
Malacca
Edible portion (%)

Skin color
Fruit size
Fresh weight (g)
Streak
Pink to crimson
Large
97
No or occasional
80
Scarlet red
Small
38
No
95
Dark red
Smallest
28
No
95
Green
Medium
59
No
95
White
Small
34
No
95
White
Medium
56
Yes
Variation
SSC (%)
Fruit length (cm)
Fruit width (cm)
Water content
Dry weight
Seed number
Maturity date
Great
10 (13 possible)
6.4
6.2
90%
10%
2
Early
Red to crimson
Largest
200
No or
occasional
Great
10
6.4
7
90%
10%
2
Early
Not available
7
4.3
4.7
12
Medium
Greatest
4.8
4
4.4
Not available
8.8
5.1
5.4
13
Late
Not available
7
5
4.4
2
Late
Not available
56
4.9
5.4
SSC, soluble solid concentration.
Chapter 11
Pink
277
Field preparation
No special preparation is required.
Trees are commonly spaced 510 m apart in rows, with about 10 m between
rows.
The trees require a regular water supply with a short period of drought. They
should be protected and received additional irrigation if subject to drying
winds. An adequate water supply is needed to replace evaporation loss during
owering and fruit growth.
Pruning
There are no easily accessible reports on pruning or thinning to develop tree
shape. In Taiwan, several labor-intensive production systems have been
developed and are described above.
Fertilization
Fertilization is recommended after the inorescence has formed, since new
ushes compete for nutrients with the ower and fruit. Leaf analysis is used to
monitor the nutritional status of the tree and as a guide for adjusting fertilizer
application. Supplementing the manganese supply through foliar application
increases the coloring of wax apple fruit. Both fruit color and rmness have
been improved after calcium amendment to the soil.
Pest management
Commonly found diseases on wax apples are Pestalotiopsis fruit rot
(Pestalotiopsis eugeniae Thuem.), bacterial wilt (Ralstonia solanacearum), algal
leaf spot (Stomatochroon sp. on the leaf lower surface and Cephaleuros virescens
on the leaf upper surface), sooty mold (Aithaloderma clavatisporum), wax
apple shoot dieback (Fusarium sp.), wax apple anthracnose (Colletotrichum
gloeosporioides Penz.), Rhizopus rot (Rhizopus sp.), Phytophthora fruit rot
(Phytophthora palmivora), Botryodiplodia fruit rot (Botryodiplodia theobromae),
Cylindrocladium fruit rot (Cylindrocladium sp.), Dothiorella fruit rot (Dothiorella
sp.), Phyllosticta fruit rot (asexual generation Phyllosticta sp. and sexual
generation Guignardia sp.) and Pseudocercospora fruit rot (Pseudocercospora sp.).
More than 50 species of insect pests have been recorded for the wax apple
since 1913. The more serious insect pests are the Oriental fruit y (Bactrocera
dorsalis Hendel), smaller green leafhopper (Edwardeiana arescens Fabricius),
citrus lamentosus scale (Nipaecoccus lamentosus Cockerell), horned wax
scale (Ceroplastes pseudoceriferus Green), red coffee stem-borer (Zeuzera coffeae
278
Chapter 11
Nietner), wax-apple thrips (Grape thrips, Rhipiphorothrips cruentatus Hood),

ower thrips (Thrips hawaiiensis Morgan), green beetle (Anomala expansa Bates),
Protaetia brevitarsis Lewis, red wax scale (Ceroplastes rubens Maskell), Chlorita
avescens Fabricius and Lasiognatha mormopa Meyrick (Platypeplus mormopa
Meyrick). These insects cause various amount of injury to fruits, owers, leaves
and twigs all year round, especially during the fruiting season. The Oriental
fruit y, thrips and scales are the major pests (Chiu, 1988).
Placing paper bags on the fruit is used to prevent fruit damage, along with
biological and chemical control methods. It is essential to remove and burn out
all the infected fruit and branches to minimize the inoculum source. The fruit
is regarded as a fruit y host, and should therefore not be left on the tree or
ground.
Weed management
Weed control is important during tree establishment, but weed growth is
limited by the canopy of mature trees. Weed growth between trees can be
managed by mechanical means and herbicides. Mulches can also be used to
suppress growth.
Orchard protection
Windbreaks are desirable when there are strong prevailing winds from one
direction. Numerous species are used as windbreaks.

Harvesting
The pink wax apple (Table 11.7) produces fruit that vary from pink to deep
red, depending on environmental and cultural conditions. The best prices are
acquired from fruit that are large, crisp, thick-eshed, juicy, sweet and deep red
to crimson in color. Fruit quality is inuenced and improved by many factors,
including light, temperature, position on the tree, growing stage, leaf-to-fruit
ratio and adequate mineral element concentrations, especially of calcium
and manganese. The lower fruit on the trunk are heavier and have the largest
fruit volume. Fruit on the upper inner portion of the tree are the smallest, but
redder in color. Fruit with the highest soluble solid concentrations are found in
a lower inner tree position.
The thin delicate skin of the wax apple is easily damaged, so extra care is
required in harvesting and handling. Fruit are harvested two to three times
279
a week when the skin is rm and has nearly full color. The fruit are sorted for
size and shape, and blemished fruit are removed before packing a single layer
in tray packs with padding to limit injury, sometimes with a paper wrap on
each fruit. Cracking of mature fruit can be a serious problem that currently
has no solution. The fruit are probably non-climacteric, as are other species
in this genus, and are chilling sensitive. The waxy skin does not readily
lose water, but a loss of 2% makes the fruit look slightly shriveled; at 4% it
is shrunken and soft. The fruit last 46 days at ambient temperatures, and
become pitted and readily decay if stored at 010C. Sugar content declines
about 1% within 4 days after harvest. Water loss and sugar decline are
reduced by holding the fruit in polyethylene-wrapped pack at 12C.

World production
All ve Syzygium species with edible eshy fruit (S. aqueum, S. cumini, S.
jambos, S. malaccense and S. samarangense) are widely grown throughout the
tropics. These trees often occur in home gardens, with limited commercial
production in the Philippines, Taiwan, Malaysia, Australia and Central
and South America. Rose apples are more popular in Central and South
America. Indonesia has the most extensive collection of water apple
and wax apple varieties, but production is scattered. Taiwan has a more
extensive commercial industry of wax apple production, which is centered
around Pingtung at the southern end of the island. Thailand has quite a
few commercial growers of water apples and wax apples in Phetchaburi
and other provinces (Yaacob and Subhadrabandhu, 1995). A fully mature
wax apple tree yields from 700 to more than 1000 fruit, with each weighing
approximately 65100 g.
Utilization
Wax apple is primarily grown for its fruit. The green fruit are eaten raw with
salt or cooked. The ripe fruit are eaten fresh, with the edible portion being
about 90% water (Table 11.8). The pink fruit are juicier, and more avorful
and suitable for eating out of hand or cooking without accompaniments
(except sugar). However, well-cultivated fruit are very sweet, juicy, crispy and
thick-eshed, and therefore are eaten raw. The proximate analysis indicates
wax apple has very low nutritional value, similar to all eshy fruited Syzygium
species (Table 11.8). Salt or sugar is added when the fruit tastes bland. In
Malaysia, fruit may be cooked as a sauce.
Chapter 11
280
Table 11.8. Composition of 100 g edible portion of ve Syzygium species (Leung

and Flores, 1961; Morton, 1987; Siong et al., 1988; Wenkam, 1990; Dignan et al.,
1994).
S. aqueum S. cumini S. jambos S. malaccense
Edible portion (%)
Proximate
Water (g)
Energy (g)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (mg)
Vitamin C (mg)
80
95
91
80
0.6
0.1
8
0.7
0.4
87
43
0.6
0.2
11.8
0.9
0.4
82.7
60
0.7
0.1
5.8
0.3
0.7
87
30
0.6
0.2
14.2
1.5
0.4
91.4
30
0.5
0.1
7.6
1
0.4
8
1.1
9
8
0.2
13
37
0.8
21
116
34.1
18
0.4
12
50
2
0
0.03
0.6
0
5
Trace
0.01
0.2
Trace
23
0.1
0.04
0.65
20
S. samarangense
0.03
0.02
0.2
0
17
6
2
16
114
0.03
0.03
0.3
Trace
13
PUMMELO
Botany
The genus Citrus is found in the Rutaceae family. The taxonomy and
systematics of the genus are complex and the number of species is much
debated. Many of the named species are clonally propagated hybrids.
Examples of hybrids are oranges, grapefruit and lemons.
The mandarin orange, sour orange and pummelo (Citrus maxima [Burm.]
Merr., formerly C. grandis Osbeck.) are considered to be natives of Southeast Asia. Grapefruit originated as a hybrid of the pummelo and sweet
orange in the West Indies, perhaps in Barbados. The lack of adequate early
281
descriptions of collected Citrus, knowledge of their origin and range of

natural occurrence has made it difficult to dene precisely the centers of
origin and ancestors of Citrus. The multitude of natural interspecic hybrids
and cultivated varieties, including spontaneous mutants, has increased the
difficulty. Scora (1975) and Barrett and Rhodes (1976) suggested there are
only three Citrus types: citron (C. medica L.), mandarin (C. reticulata Blanco)
and pummelo (C. maxima), and that these three constituted valid species. The
remaining Citrus species have been created by natural hybridization events
between various forms that were stabilized and perpetuated by adventitious
nucellar embryogenesis. Kumquats (Fortunella spp.) and trifoliates (Poncirus
trifoliata) are classied as separate but related genera. This classication has
been further supported as new molecular phylogenetic data have become
available.
Areas of origin and distribution
Citrus species have been cultivated in South-east Asia (India, China, Bhutan,
Burma, Thailand, Vietnam and Malaysia) for at least 4000 years. This
cultivation occurred long before these species were introduced into North
Africa, spreading to southern Europe in the Middle Ages. In the 1500s, with
European global exploration, Citrus species were brought to the Americas
from Europe and from South-east Asia. The pummelo is native to southern
Thailand and Malaysia, and is thought to have been introduced into southern
China around 100 BC. Production is found especially in China and southern
Thailand, and also in Taiwan and southern-most Japan, southern India,
Malaysia, Indonesia, New Guinea, Fiji and Tahiti.
Common names for pummelo include pomelo and pummelo (USA),
West Indian pomelo, Thai grapefruit and Chinese grapefruit (English), le
pamplemousse, le pamplemousse doux des Antilles, chadec and shadek (French),
Pumelo, Shaddock, Riesenorange and Lederorange (German), jeruk besar, jeruk bali,
limau kibau and limau balak (Indonesian), pampaleone (Italian), kroch thlng
(Khmer), limau betawi and limau tambun (Malaysia), sam-o and ma-o (Thai),
suh and lukban (Philippines) and cimboa and pampelmusa (Spanish).
Ecology
Soil
Pummelo is most often found on coastal silty soil overlying sand and deep clay
loams on ood plains with a pH of 6.58. It is somewhat tolerant of brackish
tidal water. It can be found on raised beds on salty mud ats. In southern
Florida and the Bahamas, it grows and produces some fruit on oolitic
limestone.
282
Chapter 11
Climate
The pummelo is tropical or near tropical and grows best at low altitudes
(<400 m) near the sea. A major growing area is around Bangkok in southern
Thailand. Here, the mean temperature is 28C (range 2530C), with a mean
annual rainfall of 1430 mm, falling mainly between May and October. A dry
season with slightly cooler temperatures lasts for 34 months. As with most
Citrus species, the higher the temperature, the lower the fruit acid content and
the higher the sugar content. Pummelo is an understory tree in the wild and
can therefore tolerate some shade. It is rarely cultivated commercially in the
shade, although it is often interplanted with other tree crops.
Tree
The small evergreen tree ranges from 5 to 15 m tall with low spreading
branches and angular, often densely pubescent twigs. Spines are found on the
branchlets, old limbs and trunk, especially on younger trees, with fewer spines
on mature trees. The alternate, ovate to ovateoblong or elliptic, leathery
leaves are 520 cm long and 212 cm wide, and moderately serrated (Fig.
11.5). Leaves of the pummelo are larger than those of other Citrus species.
The petioles are broadly winged to occasionally nearly wingless. The stem is
green.
Flowers
The large owers occur singly or in small clusters of two to 10 in the leaf axils
or 1015 in terminal racemes. The racemes are 1030 cm long. The ower
rachis and calyx are hairy and the four or ve petals are yellowish-white, 1.5
3.5 cm long, somewhat hairy on the outside and dotted with yellowgreen
glands. The stamens are white and prominent, and occur in bundles of four or
ve with orange anthers.
In the tropics, pummelo trees can ower two to four times a year, often in
conjunction with shoot ushes. Flower-bud induction commences with the
cessation of vegetative growth during dry periods in the tropics and in winter
in the subtropics. Drought periods longer than 30 days or temperatures below
25C are required to induce a signicant number of ower buds. Flower-bud
formation starts when the meristem starts to broaden and atten, and sepal
primordial followed by carpel development begins. If the terminal apex forms
sepals then the lateral buds will also form owers. If the apex forms leaves
then the lateral buds will form thorns; these are modied axillary owers, not
leaves.
283
Fig. 11.5. Leaf and fruit with ower of pummelo. (From Santiago, 1962.)
Most commercially important cultivars produce 100,000200,000 owers

on a mature tree, but only 12% of these owers will result in harvested fruit.
Pummelo, unlike most Citrus species, produces abundant pollen and most
cultivars are self-pollinated. An initial fruit drop occurs in the rst 34 weeks
following anthesis. This involves the abscission of weak owers and fruitlets
with defective styles or ovaries, or owers that receive insufficient pollination.
Preharvest drop occurs when the fruit have reached maturity, just prior to
harvest. This drop is mainly a result of natural aging and abscission. Insects,
pathogens, environmental factors and mechanical causes can also cause fruit
drop.
Fruit
The fruit is a hesperidium berry (a single enlarged ovary surrounded by a
leathery peel). The outer rind, known as the avedo (exocarp and endocarp),
is covered with tiny pockets containing aromatic oils and can be bitter in
taste. The albedo (mesocarp) contains ascorbic acid and pectin. The pulp
(endocarp) contains the juice sacs. The nearly round to oblate or pear-shaped
284
Chapter 11
fruit is widely variable in size, from 10 to 30 cm in diameter (Fig. 11.5). The

peel, 1.252.5 cm thick, ranges from smooth to coarse and greenish-yellow to
pale yellow, and is sometimes pink-blushed and dotted with tiny green glands.
The esh is greenish-yellow or pale-yellow to pink or red. The esh is divided
into 1118 segments and can range from juicy to dry. Each segment is easily
skinned and the enclosed sacs may adhere to each other or be loosely joined.
The fruit can contain a few to numerous large, yellowish-white seeds that are
white inside.
The peel can cling or be easily removed, and the albedo is soft, white or
pink. The avor varies from mildly sweet and bland to subacid or rather acid,
sometimes with a faint touch of bitterness. A pummelo cross-pollinated by
another pummelo is apt to have numerous seeds; if cross-pollinated by sweet
orange or mandarin orange, the progeny will not be seedy.
Fruit development takes 67 months and follows a sigmoid pattern.
Harvesting may not take place until the fruit is 89 months from owering.
Development occurs in four phases. The rst phase is cell division, which
determines the nal fruits potential size. This is followed by cellular
differentiation into various tissue types, followed by the third phase of rapid
expansion growth and an increase in soluble solids. In the nal maturation
phase, the peel changes from green to yellow. This external color change is a
function of climate more than ripeness and is a poor indicator of maturity.
Pummelo, as with all Citrus species, has 2n = 18 and is heterozygous, with
only a few important traits showing single-gene inheritance. Heterozygosity
produces wide variations in sexually derived progeny, but genetically and
phenotypically uniform progeny from nucellar embryos.
Problems with breeding
There is no sterility barrier between Citrus species, though cross-compatibility,
nucellar embryony and different owering times make cross-breeding more
difficult. Most Citrus cultivars have resulted from natural hybridization or
spontaneous mutation. The long juvenile period of seedlings makes pummelo
breeding a difficult and costly process. Chandler is an example of traditional
breeding by crossing Siamese Sweet pummelo (white, acidless) with Siamese
Pink (acid). This was performed in Riverside, California, and the variety was
released in 1961. The tree is vigorous, with an open drooping canopy. The
early maturing, medium-sized, round fruit have a moderately thick rind and
pink to red esh. The esh is more acid than that of pummelo from South-east
Asia and somewhat coarse in texture.
285
Major cultivars
The highly variable pummelo has numerous cultivars divided into the Thai,
Chinese and Indonesian groups. Care is needed in comparing varieties as
they often have different names when grown in different regions. The Thai
group generally has smaller fruit than the Chinese group, and is regarded
as of better quality than the other two groups. Thai cultivars include Khao
Namphung (White Honey) and mid-season Khao Phuang (White Tassel,
Siam), both of which are pear-shaped and have white esh, and Thongdee
(Golden), which is more rounded and has pink esh. The Chinese group
consists of white-esh, pear-shaped fruit types with thick rinds. The main
Chinese varieties include Shatianyou (J8) in Guangdong and Guangxi,
and Guanximiyou and Wendanyou in Zhejiang and Fujian. Cultivars in
the Indonesian group are larger and have white and red esh, with low acid
content. These include Nambang, Sri Nyonya (thinner skin, sour), Jeruk
Bali merah (red esh) and Jeruk Bali putih (white esh).
Cultural practices
Seeds are used for propagation, although the progeny are not uniform. Air
layering is most commonly used in South-east Asia. When virus-free mother
trees are available then budding (e.g. shield budding) is also practiced, with
seedlings being used as rootstock. Calamandarin is used as a rootstock in the
Philippines. Trees come into bearing 68 years from seed and earlier if from
air layers.
Field preparation
In areas subject to high water tables and ooding, pummelo is grown on raised
beds that require continuous maintenance.
The plant-to-plant spacing is 810 u 68 m, depending on the terrain and soil
fertility. This is equivalent to a population density ranging from about 125 to
210 plants/ha. In Thailand, where raised beds are used to prevent ooding,
two rows are planted at 34 u 45 m on the bed, with 6 m between beds. Prior
to planting-out, the seedling trees are cut back to reduce leaf area and balance
the root-to-shoot ratio. Shade and frequent watering are needed during
establishment.
Pummelo plants are sensitive to drought during vegetative ushes, owering,
fruit setting and fruit enlargement. At these stages, a mature tree requires
286
Chapter 11
100200 l/day water. As a practical guide for irrigation, plants should be

watered when 2030 cm of the topsoil is dry. Mulching reduces the need for
irrigation, but is only occasionally used in commercial production. Surface
ood irrigation, sprinklers and trickle or mist irrigation are employed.
Pruning and fruit thinning
The trees are pruned after planting-out to leave three main branches as a
framework, with the lowest branch 3050 cm above the ground. Subsequent
pruning aims to create an open-centered tree. Dead-wood is removed, along
with overcrowded shoots and branches that sag when loaded with fruit and
touch the ground. Alternatively, sagging branches can be propped up with
poles. Pruning should be carried out in the dry season when little vegetative
growth is occurring.
Fruit thinning is practiced when the fruit are small (approximately 4 cm in
diameter) to increase the size of the remaining fruit. The extent of thinning
varies with variety, with 4060 fruit being left on Thongdee and 2040 on the
lower-yielding Kao Namphung.
Fertilization
The fertilizer practices used for other Citrus species are also applicable to
pummelo. Mature trees are fertilized two or three times a year, with an annual
application of manure. The quantities applied vary with the soil, fertility,
growing conditions, intensity of cultivation and tree age or size. The aim is
to replenish nutrients lost during harvest and leached from the root zone. As
with other Citrus, attention needs to be paid to magnesium nutrition and the
other micronutrients (zinc, manganese, copper, boron). In Thailands Nakhon
Pathom province, about 5 kg/tree/year of 16:16:16 NPK is recommended in
bi-monthly applications, along with foliar fertilizer application for every new
vegetative ush. Other recommendations are for two applications per year: the
rst before owering and the second during fruit growth 46 months later.
Pest management
Citrus trees are subject to many diseases, insects and pests. These affect the
roots, trunk and branches, foliage and fruit. All the pests of Citrus also attack
the pummelo. These include the common leaf miners (Phyllocnistis citrella),
leaf-eating caterpillars, fruit-boring caterpillar (Citripestis sp.), scales, red
mites, fruit ies, nematodes and rats.
The major disease of pummelo is bacterial canker caused by Xanthomonas
citri. Symptoms include oily spots on the leaves and fruit that later turn brown
and corky. Control methods include defoliation and, in severely infected plants,
burning to prevent spread. Root rot, trunk gummosis and fruit brown rot are
caused by Phytophthora and can be a signicant problem. Both fruit and leaves
can also be infected by scabs caused by Elsino fawcettii. To control fungal
287
diseases, repeated spraying of chemical fungicides is recommended. The leaves,

fruit and sometimes the branches are likewise prone to sooty mold, which is
caused by Capnodium citri or Meliola citricola. Sooty mold can be prevented
by proper insect pest control. Huanglongbing (also known as greening) is a
problem in pummelo production, causing a blotchy mottled appearance of the
leaves.
Pummelo varieties show differential susceptibility to different Citrus tristeza
virus isolates. Citrus tristeza virus is spread through infected budwood sources
and in the eld by aphids, which transmit the disease with varying degrees of
efficacy.
Weed management
Weed control is crucial during orchard establishment. Control can be either
mechanical or through the use of herbicides. Mulching under the tree and
managed cover crops also help to control weeds.
Orchard protection
Some protection from strong winds is desirable. Bananas are used in some
countries as a windbreak, for shade and as an initial source of income before
the pummelo reach full production. Palms, such as areca, are planted as an
intercrop and around pummelo orchards.

In China, pummelo is available in January to March and August to December.
In the Philippines, it is generally reported to be in season from November
to January. In southern Vietnam, fruit are available in the markets from
September to February, with the peak from November to January. In the cooler
north of Vietnam, pummelo are in season from August to November, with
differences in fruit maturation by cultivar.
As the fruit matures the skin changes from a dull gloss to shiny, starting at
the base of the fruit. The fruit is ready to harvest when full gloss is achieved
(this will be 67 months after owering). Fruit are often left on the tree and not
harvested until 8 months, but before the esh becomes dry and corky at the
stem end. In Thailand, the minimum grade standard for total soluble solids is
8% for Thongdee and 7% for Kao Namphung. The sugar-to-acid ratio is 8 for
both varieties.
Fruit can be cut from the tree with clipper or broken off by pulling. They are
accumulated in buckets and baskets, and either packed into containers in the
eld for the local market or transported to packing facilities. At packing sheds,
the fruit are washed, graded and packed into cartons. Grading removes fruit
with surface blemishes, sunburn, sooty mold, external damage caused by
288
Chapter 11
insects and disease, poor shape, thick peel, puffiness or peel color more yellow
than green.
This non-climacteric fruit must be handled carefully during harvesting and
handling to minimize damage and postharvest decay. The pummelo can be
held at ambient temperature for 514 days. It can be stored at 1215C and
95% relative humidity for up to 12 weeks. Fruit stored at 610C last for 24
weeks.

World production
Data for pummelo are frequently included with those of other Citrus species
such as grapefruit. Grapefruit and pummelo are produced commercially in
74 countries worldwide, on about 264,000 ha of land. Thailand is the major
producing country. Its mature and sustainable pummelo production has
maintained a strong export trade for about 30 years. Thongdee is the major
variety in the export trade to Hong Kong and China in the main, as well as
to smaller markets in South-east Asia. Singapore is served by both Indonesia
and Malaysia. A major constraint to development of the export market is
the availability of a consistent supply of quality fruit. China, with its large
acreage, is providing fruit for its own consumers; with increasing production,
it will have sufficient quantities to export quality fruit.
Utilization
Pummelo is consumed as a fresh fruit or minimally processed and sold in
plastic covered trays. It is commonly eaten after the segments have been
removed from the peel and the segment membrane is peeled away, and
occasionally like grapefruit with a spoon. The fruit segments are used in fruit
salad and sometimes the juice is extracted. The juice is not as bitter as that of
grapefruit, having a sweet, mild avor.
The fruit is high in vitamin C, folate and carotenoids (Table 11.9). Like most
Citrus fruit, pummelo contain various phytochemicals that may play a health
role. Some pummelo varieties contain up to six different avonoids, including
narirutin, naringin, hesperidin, eriocitrin, neohesperidin and neoeriocitrin.
Pigmented red pummelo fruit contain lycopene and the juice is an excellent
source of antioxidant compounds, including vitamin C, total phenolics and
carotenoids, and exhibits an excellent scavenging ability for different forms of
free radicals (Tsai et al., 2007).
The essential oils are used in cosmetics, soaps, perfumes and aromatherapy,
and for pharmaceutical purposes such as sedatives and antispasmodics. The
inner rind of all Citrus species provides a rich source of pectin for jams and
jellies.
289

portion of pummelo fruit.
Constituent
Proximate
Water (g)
Energy (kcal)
Protein (g)
Fat (g)
Carbohydrates (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Potassium (mg)
Sodium (mg)
Vitamins
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Vitamin A (IU)
Vitamin C (mg)
Carotene (mg)
Pummelo
8594
2558
0.50.74
0.20.56
6.312.4
0.30.82
0.50.86
2130
0.30.5
2027
0.040.07
0.02
0.3
20
3043
AMBARELLA
Ambarella (Spondias dulcis Soland Ex. Forst., syn. S. cytherea Sonn.) belongs to
the Anacardiaceae family. In English, it is known as ambarella, otaheite apple,
golden apple, Jew plum, wi- or vi- or evi-apple, great hog plum, Polynesian
plum, Tahitian quince or apple and golden apple. Other names include June
plum (Jamaica), kedondong and kedongdong (Malaysia, Indonesia), ma kok
farang (Thailand), mokak (Cambodia), coc, pomme cythere and pommier de
cythere (Vietnam), heri (Philippines), gway (Burma), kook hvaan (Laos), juplon
(Costa Rica), hobo de racimos (Colombia), mango ciruelo and taperib (Peru), jobo
de la India and mango jobo (Venezuela) and caj-manga (Brazil).
Important Genera and Species
Important fruit-producing genera in this family are Mangifera (the mango
and relatives), Pistacia (pistachio) and Anacardium (cashew). Other species
of the Spondias genus include S. purpurea L. (red mombin), S. mombin (yellow
290
Chapter 11
mombin or hog plum) and S. tuberosa Arruda (the imbu or umbu, from Brazil).
The other three related species from South-east Asia that have edible fruit
are S. acida Bl. from the western Indo-Malayan area, which has acid fruit; S.
novoguineensis Kostermans from New Guinea to the Solomon Islands, which
is semi-cultivated and frequently confused with S. cytherea; and S. pinnata
(Keonig ex. Linn. F) Kurz. (Indian hog plum) from Burma, India and Thailand
(Andall and Paull, 2008).
Origin and Distribution
Ambarella is native to the South Pacic islands from Melanesia through to
Polynesia. It has been introduced to South-east Asia, where it can be seen
on the markets of Vietnam, Laos, Cambodia and India. It was later taken to
Africa and to most tropical areas, including Central America, the warm
areas of Australia and Caribbean islands such as Cuba, Trinidad, Jamaica,
the Dominican Republic and others. Although widely distributed in tropical
areas, its cultivation in the Caribbean and Latin America is limited to scattered
plants or mixed plantings (Marte et al., 1992). Export markets have become
established during the last couple of decades and the large-type fruit is
exported from several Caribbean countries, including Trinidad and Tobago,
Grenada, St Vincent, Guyana, Surinam, Jamaica, the Dominican Republic and
Dominica, and in the mature-green stage to Europe and the USA (Mohammed
et al., 2011), to ll the demand of ethnic markets.
Ecology
Soil
Ambarella is not very demanding on soils. It adapts to a wide range of soils,
including the oolitic limestone of Florida, as long as they are well drained. It is
susceptible to saline conditions.
Climate
RAINFALL This is a plant for the humid and semi-humid tropics and subtropics.
The tree is drought tolerant and may briey shed its leaves under water stress
or a cold climate. During long periods of drought the trees remain small and
produce fewer undersized fruit, while excess rainfall leads to soft fruit that
bruise easily (Mohammed et al., 2011). The fruit are of better quality when they
mature and ripen during periods of low rainfall (Bauer et al., 1993; Vargas et
al., 1999). This variation in fruit quality is due in part to the sugar content,
which depends on exposure to full sunlight.
291
The ambarella tree grows well in the hot tropics at elevations

of up to 1000 m and in the warm subtropics with mild winters. The tree grows
poorly in cold winters and will bear only few, low-quality fruit. It is severely
damaged by temperatures below 0 to 1C, and does not resist prolonged
periods of temperatures below 10C.
TEMPERATURE
LIGHT AND PHOTOPERIOD Shaded trees produce little fruit and full exposure to
sun is necessary, although light shade is benecial during establishment (Ochse

et al., 1961). A photoperiodic response is apparently not a factor, as it owers
from the tropics to the subtropics.
WIND Ambarella is not very resistant to strong winds because the branches
are brittle and easily broken. The tree is also susceptible to salt spray. A sheltered
location is required.
Tree
The rapidly growing, upright tree can reach 1520 m and has a symmetrical
shape with a rounded crown. The bark is a light grayish-brown and nearly
smooth, often with four to ve small buttresses. The leaves are pinnate (2060
cm long) and are composed of nine to 25 pairs of glossy, elliptic or obovate
oblong leaets (6.010 cm long) on a short petiole (Fig. 11.6). The leaves
normally become yellow in color and fall at the start of the cool-dry season,
leaving an attractive bare tree (Morton, 1987).
The tree grows rapidly, and owers and bears fruit in 4 years from seed.
The tree produces more or less continuously in the humid tropics, while in the
subtropics it becomes more seasonal because of the cold period. A dwarf type
developed in the Far East starts owering after 1 year when it is less than 1 m
high.
Flowers
The whitish owers are small and inconspicuous and are borne in large
terminal panicles (50 cm long) appearing before the leaves. In each panicle,
male, female and perfect owers occur on short pedicels (14 mm). The
calyx lobes are triangular (0.5 mm) and the petals (2.5 u 1 cm) are ovate
oblong. Vegetative ushing and owering occur together. In areas with a dry
season, owering occurs on trees that are nearly leaess from lack of soil
moisture. Subtropical conditions lead to owering in the spring. Fruit bats
may assist in seed dispersal. No data have been reported on pollination and
fruit set.
292
Chapter 11
Fig. 11.6. Leaf, ower and fruit of ambarella (Spondias dulcis Soland Ex. Forst.).
(From Little and Wadsworth [1964] Trees of Puerto Rico. USDA Handbook #249.)
Fruit
The fruit weigh up to 450 g in the regular type and around 60 g in the dwarf
type. They are borne on long peduncles in bunches of a dozen or more, and
have tough thin skin that is often russeted. The fruit is an ellipsoid or globose
drupe (410 u 38 cm) and changes from bright green to bright orange or
golden-yellow on ripening. In rm fruit, the esh is crisp, crunchy, juicy and
slightly sour, with a somewhat pineapple-like fragrance and avor. Like the
skin, the esh becomes golden-yellow when it ripens (Morton, 1987). The
endocarp has conspicuous irregular and brous protuberances or spines
extending from the rough ridges of the ve-celled woody core, which contains
one to ve at seeds. In overripe fruit, the mesocarp becomes soft and musky
in avor and aroma and difficult to slice because the tough spines of the
endocarp (Mohammed et al., 2011). The fruit take 68 months to mature
(Bauer et al., 1993).
293
Attempts have been made in Grenada to cross the normal ambarella with
the dwarf type. Little has been reported on the selection and evaluation of
new varieties. Plants are usually clonally propagated so superior types are
maintained.
Cultivars
There are selections, but no recognized cultivars. Large variations in fruit
quality (acidity, sweetness, size and seed spines) have been reported. The
average fruit weight is 140225 g, and in some cases fruit of 450 g have been
observed (Bauer et al., 1993). The dwarf type that grows to a height of 1.53
m and produces fruit of about 6065 g, starting 1 year after sowing the seed.
This very precocious plant produces throughout the year. In addition, picking
is much easier than in regular types and the plants can be established at very
high densities, resulting in high yields per area.
Cultural practices
Propagation
The tree is often propagated from seeds that germinate in about 1 month.
More than one plant can be obtained from one seed since the stone contains
up to ve seeds. The most vigorous seedlings start to bear about 34 years after
planting. The dwarf type starts producing after 1 year.
Vegetative propagation is possible with large hardwood cuttings and
air layering, while grafting or shield budding can be performed on S. dulcis
seedling rootstock. S. pinnata Kurz. is used as rootstock in India. Some authors
have indicated that T budding on S. mombin L. is successful. Normally large
hardwood cuttings are used, thus ensuring a good initial tree size. This is
practical if the material is available otherwise, grafting or budding will be
needed to rapidly increase plant numbers. Asexually propagated plants will
start producing after 23 years.
Fields are prepared as for other fruit tree plantings. For normal-sized trees,
spacing varies from 7 to 12 m (Andall and Baldeo, 2008); Vargas et al. (1999)
have recommended 10 u 10 to 12 u 12 m. The dwarf type should be planted
at 4.5 u 4.5 m or even closer (Ministry of Agriculture of Jamaica, 2007).
Irrigation
Trees are not normally irrigated, but irrigation is strongly advised during
the dry season, especially if it is excessively long. An estimated 4 l/day/tree
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Chapter 11
should be applied during early growth, increasing to 32 and 157 l/day/tree at

10 years for the dwarf and tall types, respectively (Ministry of Agriculture of
Jamaica, 2007).
Pruning
Large-type trees should be formed to obtain a plant with three or four main
branches. These are later topped to obtain secondary branches and prevent
the tree from becoming too tall. After that, any diseased or damaged branch
should be removed, as well as any branch growing in the wrong direction
or too low. In the dwarf type, care must be taken to leave a short main stem
from where branches will arise after certain height, removing excessively lowgrowing branches and vertical branches competing with the main stem. Fruit
pruning within the panicle increases the size of the remaining fruit but total
yields will be lower, while inorescence pruning reduces both fruit size and
yields (Andall and Baldeo, 2000).
Fertilization
No specic data exist with respect to fertilization. In Jamaica, a general
recommendation is to apply 24 kg organic matter during the early years,
increasing to 10 kg in later years. In the rst year, 200 g diammonium
phosphate and 200 g ammonium sulfate are recommended; in the second
year, 900 g of a 14:28:14 fertilizer should be applied. From years 36, 1.35,
1.8, 2.0 and 3.0 kg/tree, respectively, of a 15:5:35 fertilizer should be applied,
reaching 4.0 kg in year 10 (Ministry of Agriculture of Jamaica, 2007).
Pest management
DISEASES In the Caribbean, a disease called gummosis results in black spots
on the fruit, debarking of the stem and subsequent death of the tree. A canker
problem caused by the Lasiodiplodia fungus or the Xanthomonas campestris
bacteria may also occur. The fruit is attacked by Guldnardia, Asteromella and
Colletotrichum species, which cause small (8 mm) black lesions or as well as
large, round, black spots (1.5 mm) in green fruit, with gumming that remains
supercial (3 mm deep) and does not rot the pulp. The incidence is higher
in the rainy season (Bauer et al., 1993). Growers in Jamaica have reported
anthracnose and some fungal leaf spots (Ministry of Agriculture of Jamaica,
2007). Phytophthora and bacterial attacks have also been reported.
INSECTS The fruit can be attacked by Caribbean fruit y (Mohammed et al.,
2011), and in Costa Rica the bark is eaten by a Trigona wasp that causes necrosis
(Vargas et al., 1999). In Jamaica, mite attacks as well as ddler and June beetles
occur (Ministry of Agriculture of Jamaica, 2007). In Indonesia and Malaysia,
the kedongdong spring-beetle Podontia affinis Grond and P. punctata cause
295
serious defoliation problems. Ants, termites, epiphytes, birds and lizards are
occasional pests (Mohammed et al., 2011).
Weed management
Weeds are only problematic in the very young stages of growth. They can be
managed manually or with herbicides.
Orchard protection
Some commercial plantations are sited in areas subjected to strong winds.
The ambarella wood is very brittle, so a sheltered location should be selected
with the protection of neighboring forests and wind barriers established, if
necessary.

It is estimated that a large tree in its adult stage can produce almost 1000 fruit
(Vargas et al., 1999), which can result in yields of 1020 t/ha/year. Bauer et
al. (1993) have reported yields as high as 900 kg of fruit per tree per year. For
the dwarf type in Jamaica, the estimate of yield is 4 t/ha in the second year,
increasing to 40 t/ha after 9 years (Ministry of Agriculture of Jamaica, 2007).
Harvesting
The fruit are usually harvested when they start to yellow (Ministry of
Agriculture of Jamaica, 2007). Fruit are detached by pulling with the hand, or
harvest shears are used to cut the peduncle.
Ambarella is a climacteric fruit. The fruit from both the regular and dwarf
types are very susceptible to mechanical damage. Fruit should be harvested
at a uniform stage of maturity and handled very carefully between harvest
and packing to avoid bruising. Padded containers should be used and the
fruit shaded before and during transport to the packinghouse. Fruit should
be hydrocooled to remove eld heat within 3045 minutes of arrival using
water at 710C. The fruit are kept at 1213C and 8590% relative humidity
during postharvest handling (Mohammed, 2003). Chilling injury will occur in
green mature fruit at temperatures below 8C.
To market the fruit in the mature-green stage, the recommendation is to
keep the fruit at 1012C and 9095% relative humidity at all times and avoid
exposure to ethylene from other ripening fruit. Fruit can be ripened by being held
at room temperature for 24 days or treated with Ethrel at 500 ppm at 22C
and 9095% relative humidity for 2436 h, and holding at 78C and 9095%
relative humidity until marketed and consumed. Fully ripe fruit are less sensitive
to chilling injury than mature-green fruit. Reduction of the ethylene action or
biosynthesis can also help to extend the storage life (Mohammed et al., 2011).
Chapter 11
296

portion of ambarella fruit. (After Morton,
1987; Mohammed et al., 2011.)
Constituent
Proximate
Water (g)
Energy (cal)
Protein (g)
Fat (g)
Fiber (g)
Ash (g)
Sucrose (g)
Acid (g)
Vitamin
Vitamin C (mg)
Ambarella
59.685.4
157.3
0.50.8
0.281.79
0.853.60
0.440.65
8.0510.54
0.47
4.655.86
Utilization
The pulpy esh can be eaten fresh when still rm and crisp or allowed to ripe
to a soft stage. The green fruit is used for salads, curries, pickles and juices. It
is sometimes eaten with salt, vinegar and hot chili pepper. Ripe fruit are also
stewed and used for jams, jellies and juices and canned. In Jamaica, ambarella
juice is sold in supermarkets mixed with ginger, which gives it a special avor.
The fruits vitamin C content is reported to be 36 mg/100 g and it is a good
source of iron (Table 11.10). Impact aroma volatiles include ethyl(S)(+)
2-methyl butyrate, ethyl isovalerate, ethyl propionate, ethyl butyrate, linalol
and trans-pinocarveol (Fraga and Rezende, 2001). The young leaves are eaten
raw or steamed as a vegetable with salted sh.
The fruit, leaves and bark have been reported to have medicinal value in the
treatment of sores, wounds and burns. The bark is mixed with that of other
species to treat diarrhea. The light-brown wood has a low density and little
timber value, although it has been used for canoes in the Society Islands (Morton,
1987). The gum has a high viscosity, with galactose as the main component.
FURTHER READING
Andall, R. and Paull, R.E. (2008) Spondias cytherea Ambarella. In: Janick, J. and Paull,
R.E. (eds) Encyclopedia of Fruit and Nuts. CAB International, Wallingford, UK, pp.
2931.
Bayogan, E.R.V. and Paull, R.E. (2008) Santol Sandoricum koetjape (Burm. f) Merrill,
Meliaceae. In: Janick, J. and Paull, R.E. (eds) Encyclopedia of Fruit and Nuts. CAB
297
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12
AMERICAN FRUIT
Hundreds of edible fruit grow in tropical America. In this chapter, we discuss

ve fruit that are already being exported to a limited extent: (i) mamey sapote,
Pouteria sapota (Jacq.) H.E. Moore and Stearn (Sapotaceae); (ii) chiku or
sapodilla, Manilkara zapota (L.) von Royen (Sapotaceae); (iii) pitaya or dragon
fruit, Hylocereus species (Cactaceae); (iv) acerola, Malpighia emarginata DC.
(Malpighiaceae); and (v) naranjilla, Solanum quitoense Lam. (Solanaceae).
MAMEY SAPOTE
Introduction
The name sapote is believed to be derived from the Aztec tzapotl, which is
used for soft sweet fruits. The black sapote (Diospyros digyna Jacq.) belongs to
the family Ebenaceae and is therefore not a true sapote. The family Sapotaceae
contains a number of species with edible fruit in different genera: Manilkara
zapota (L.) von Royen (chiku, chico zapote or sapodilla), Chrysophyllum cainito
L. (caimito, star apple), Pouteria sapota (Jacq.) H.E. Moore and Stearn (mamey
sapote), P. campechiana (HBK) Beahni (canistel), P. obovata HBK (lucmo or
lcumo), P. caimito (Ruiz and Pav.) Radlk (abiu or caimo) and Calocarpum viride
Pittier (green sapote or injerto).
The mamey sapote (P. sapota) is the largest fruit of this genus. Synonyms
include P. mammosa (L.) Cronquist, Calocarpum sapota Merr., C. mammosum
Pierre, Lucuma mammosa Gaertn., Achras mammosa L., Achradelpha mammosa
Cook, Vitellaria mammosa Radlk. and Sideroxylon sapota Jacq. The taxonomy of
mamey sapote was confused in the past with that of chiku (Manilkara zapota
[L.] P. van Royen; syn. Achras sapota [Mill.] Fosb.). However, the morphology
of these two plants differs signicantly and they were subsequently placed in
separate genera.

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Common names for mamey sapote (English) include sapote, sapota, sapote
mamey, mamey, mamey colorado, zapote, zapote grande, zapote de carne, zapote
mamey, chachaas and tezonzapote (Spanish America), mamey de la tierra
(Panama), sapotier jaune doeuf and grand sapotillier (Haiti), sapote a crme
(Guadeloupe), grosse sapote (Martinique), sapote mamey and sapotier (French),
sapote (Portuguese), ciko mama (Indonesian), chico-mamey (Malay and Filipino),
trung ga (Vietnamese) and marmalade fruit or marmalade plum (Jamaica).
Mamey sapote is native to southern Mexico to northern Nicaragua in
Central America (Morton, 1987). Although wild populations are found from
Mexico to Panama (Leon, 1983), it seems that it is not native to Panama
and Costa Rica, where it is replaced by P. fossicola and P. viridis (Pennington,
1990). The plant is distributed throughout the tropics and some warm
subtropical areas (e.g. south Florida). It is grown commercially in many
tropical countries and fresh fruit is sold in local markets. However, a major
proportion of mamey sapote fruit is produced by scattered trees in backyards
and home gardens, with only a few large orchards. Mexico and Guatemala
have recently increased their cultivated areas. The large Cuban population in
Florida has increased the demand and price for this fruit, and has lead to the
establishment of orchards in southern Florida. There are no reliable statistics
about worldwide production. Frozen pulp is exported from Central America.
Ecology
Soil
Mamey sapote trees prefer deep clay and clay loam soils with medium fertility,
rich in organic matter and slightly acid, but they will also grow in shallow
calcareous soils such as those of the Yucatan Peninsula. Soils should be well
drained, since this species is intolerant of ooded soils, soils with a high water
table or those with impermeable subsoil (Almeyda and Martn, 1976). In
addition, the trees are intolerant of soil salinity and low pH.
Climate
RAINFALL Mamey sapote is adapted to hot, humid, tropical lowland areas with
moderate rainfall (Martin et al., 1987). It is very susceptible to drought, which

causes shedding of leaves. It can be found in areas of Central America with
5002000 mm rain and in areas of Mexico with average rainfall of 16003000
mm (Galvn and Nez, 1994); the ideal is 15002000 mm (Azurdia, 2006).
In areas with low precipitation (400700 mm rain), satisfactory production
can be obtained with supplemental irrigation during the dry season.
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305
This tree is tropical, but is also found in the subtropics and

at higher elevations in the tropics. It may be found from sea level to 600 m,
occasionally up to 1000 m in tropical Central and South America, and
sometimes at 2000 m in Guatemala (Leon, 1983). In Mexico it is normally found
from 100 to 2000 m, and higher where average temperatures range from 19C
to 27C (Galvn and Nez, 1994). The best production is obtained at altitudes
of 0800 m and the ideal temperature range seems to be 2032C (Azurdia,
2006). At low temperatures, leaf color changes to yellowish or reddish and
leaves eventually abscise. Young trees are damaged or killed by temperatures of
0C or lower and mature trees at 2.0C or lower (Campbell et al., 1977). Frosts
cause burning of the foliage and branches, and induce fruit drop.
TEMPERATURE
LIGHT Wild plants are normally found in forests near canopy openings that
allow about 60% full sun. In monoculture, trees grow well in full sun with
proper care. Temporary shade is needed during the rst years in the eld.
Photoperiod responses have not been reported.
WIND Wind barriers are needed where strong winds are common, with
staking necessary for young trees. Mamey sapote trees can stand hurricaneforce winds, although severe limb damage and defoliation does occur (Crane et
al., 1993).
Tree
Mamey sapote is an erect tree that usually grows up to 18 m (Kennard and
Winters, 1960) and sometimes to 3050 m. The trunk can be 1 m thick and
is often narrowly buttressed. The branches tend to be horizontal and the
canopy is pyramidal, with sympodial branching. The bark is scaly and reddishbrown, and young branches are covered with a light-brown pubescence.
The taproot extends to a great depth and the brous lateral root system is
extensive, especially under good soil conditions. The evergreen or deciduous
simple leaves are clustered at the branch tips in a spiral arrangement. Petioles
are about 35 cm long, pubescent or glabrous; the blades are obovate,
prominently veined, pointed at the apex and 410 cm wide by 1030 cm long
(Fig. 12.1). All parts of the plant exude a white gummy latex when wounded.
Flowers
The hermaphroditic owers are white, pale yellow or greenish-cream in
color. They are subsessile and are borne along small-diameter (1.35.1 cm)
branches, in the axils of the fallen leaves and further back to a proximal
distance of 12 m from the branch tips, appearing in clusters of three to
six. Individual owers are small; the calyx has eight to 12 lobes, the tubular
306
Chapter 12
Fig. 12.1. Mamey sapote showing leaves (A), fruit (B) and seed (C). Fruit shape (D)
varies greatly. (Used with permission from Leon, J. [2000] Botnica de los Cultivos
Tropicales. Agroamerica del IICA, Costa Rica.)
corolla has ve lobes and there are ve stamens, ve staminoids, one pistil and
a ve-celled ovary. Trees can ower several times a year in tropical areas, while
in areas with cooler winters owering is usually in the spring.
Floral buds develop in periodic bursts and the length of bud development
varies greatly among cultivars (Davenport and ONeal, 2001). The owers
begin to open in groups in the late afternoon and most open during the night.
Flowers are open for about 1 day during the warm season and for up to 6 days
at cooler temperatures. Initial fruit set is often greater during summer (38%)
than in spring and winter (approximately 5%). Fruit distribution along the
stem is greater on the lower (49%) part than on the sides (2022%) and upper
part (10%) although initial fruit set is the opposite, with more fruit drop from
the upper parts.
Cross-pollination improves fruit set (Balerdi and Crane, 2009) although
87% of isolated trees set fruit, suggesting a degree of self-pollination. Cross-
American Fruit
307
pollination is 100% in wild populations, but more domesticated and cultivated

trees have a lower percentage of cross-pollination. Several buttery species
visit the owers and it has been suggested that honey-bees and other insects
are important agents (Azurdia, 2006).
Fruit
The fruit is a round, ovoid or elliptic berry that is often acutely pointed at the
apex (Fig. 12.1). It is 823 cm long and 1016 cm in diameter, and weighs
2302300 g or more. The rough, leathery peel is brown and scurfy, becoming
less scurfy at maturity. In wild types the skin can be thin or leathery and gray
in color (Leon, 1983). Pulp color varies from brown to orange, salmon-orange
or red, and is sometimes dark red or even dark yellow. The pulp is very sweet
and soft when ripe with an almond- or pumpkin-like avor, and variable
amounts of ber according to cultivar (Campbell and Lara, 1982). The one
to four seeds are large (511 cm long and 32 cm wide), spindle shaped and
pointed, with a hard and brittle coat that is glossy brown or black and has a
fairly wide and slightly rough hilium on the ventral side. The cotyledons smell
like bitter almond.
The owering season lasts 12 months. The time from anthesis to fruit
maturity is 1324 months, depending on cultivar and temperature. Fruit in
all stages of development may be found simultaneously on the same branch
(Balerdi and Crane, 2009). The main season is from March/April to July/
August in subtropical south Florida; from June/July to November/December in
the Dominican Republic; and from December to March and from May to July in
Costa Rica. Some trees can produce almost year-round (Azurdia, 2006).
There is currently no known breeding program. In Guatemala, the northern
Department of Petn has the richest mamey sapote biodiversity; new
phenotypes have been selected there with fruit sizes ranging from 341 to
1096 g and one to four seeds, and tree heights from 12.5 to 72 m. (GranadosFriely, 1995). Some collaborative work is ongoing between CATIE in Costa
Rica and PROFRUTA/San Carlos University in Guatemala, with the Fairchild
Tropical Gardens of Miami looking for phenotypes with superior fruit quality
that can be asexually propagated. Since there is much variability between
seed-propagated plants in fruit quality and yields, superior types that can be
vegetatively propagated are desirable (Azurdia, 2006). The Florida program
uses material from Central America, Mexico and the Caribbean (Campbell et
al., 1997a).
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Selections of mamey sapote can be divided into groups, but cannot be
differentiated as individuals. Molecular markers have to be used to better
differentiate the types (Carrara et al., 2002). The main criteria are fruit size
and pulp quality, color, absence of ber and few seeds, harvest season and
low-branching habit. There is interest in cultivars with fruit that mature in
different seasons to ll market niches (Campbell et al., 1997a).
Cultivars
Some of the most important cultivars are Magaa and Tazumal from El
Salvador; Pantin and Pace from Florida; Cayo Hueso from the Dominican
Republic (which is very popular among Cubans); and Chenox from Belize.
Older cultivars used on a limited scale commercially or as backyard trees are
Copan from Cuba; Cuban No. 1, which is of Cuban origin brought from El
Salvador; Francisco Fernndez from Cuba; Mayapan from the Isle of PinesCuba; Progreso from Belize; Flores from Guatemala (Morton, 1987); and
Valiente and Rivera from El Salvador. There are several local selections in
Cuba, Mexico, Nicaragua, Guatemala, Costa Rica and the Dominican Republic
that need to be evaluated more closely to extend their use to other places.
Balerdi and Crane (2009) have listed some cultivars that are being grown or
tested in Florida (Table 12.1).
Cultural practices
Propagation
SEXUAL Mamey sapote trees may be propagated by seed, but this should
be only used to raise rootstock or search for new cultivars because of the
variability of the species and long juvenile period (712 years). The shortlived, recalcitrant seeds (Duarte and Suchini, 2001) should be planted within
a few days of removal from the fruit as viability is lost in 14 days at ambient
temperatures. Removal of the seed coat hastens the germination speed, but
reduces percentage and seedling size when compared with seeds with intact or
cracked coats. The best germination is obtained by cracking the seed coat and
sowing in a lay. Gibberellic acid treatment reduces the germination time from
34 to 24 days, but does not increase germination percentage (Duarte and Cruz,
2007), and hastens initial growth to the seedling stage from 4 to 2.5 months.
ASEXUAL Propagation using air layers and cuttings has been attempted with
little or no success (Almeyda and Martn, 1976; 1979). Tissue culture has also
given discouraging results (Azurdia, 2006). Grafting is the most feasible asexual
propagation method. A major problem with grafting is that mamey sapote has
a discontinuity of vascular cambium tissue in the terminal parts of the shoots,
Table 12.1. Characteristics of mamey sapote cultivars evaluated in Florida and their adaptability for commercial production.
(Adapted from Balerdi and Crane, 2009.)
Cold
Commercially
tolerant Precocity grown
High
Yes
No
No
High
No
Yes
Yes
Red
Pink
Pink
Pinkred
Salmon
Salmon
Red
Pink
Dark red
Pinkred
Medium,
branched
G
Small, slow
growth
G
Tall, slender
G
Medium
Fair-G Medium
E
Medium
E
Tall
E
Tall
E
?
G
Medium
E
Branched
E
Slender
High
High
Medium
Medium
High
High
?
Medium
Medium
Medium
Yes
Yes
?
No
Yes
Yes
Yes
?
?
No
No
Yes
?
No
Yes
Yes
?
?
?
?
Under evaluation
No
No
Under evaluation
Yes
Yes
Under evaluation
No
No
Under evaluation
Red
Dark red
E
E
?
?
?
Yes
?
Yes
Under evaluation
No
Harvest
Fruit size (g) Pulp color Flavor Tree size
Copan
JulySept
425900
Red
Magaa
JulyAug
7402400
Pink
Maypan
Tazumal
AREC 3
Piloto
Pace
Florida
Lara
Chenox
Abuelo
Francisco
Fernandez
Flores
Viejo
JulyAug
JanFeb
JulySept
AugSept
MarApril
MarApril
AugSept
MayJune
OctNov
AugSept
5101135
400850
400740
400740
425900
425900
4001130
400850
7402400
560700
NovDec
Dec
7402400
400560
Slender
Branched
American Fruit
Yield
Cultivar
?, Unknown.
309
310
Chapter 12
thus making cambial contact difficult. In addition, ber strands in the cortex
might also interfere with the process (Ogden, 1984). Superior cultivars are
normally propagated by veneer, cleft, four-ap or approach grafting; budding
can also be used (Rodrguez and Gurdian, 1986; Marler, 1991; Campbell
and Lara, 1992; Alix and Duarte, 1999). Veneer grafting is one of the most
popular methods (Quilantn-Carren, 1979). Vigorous seedling rootstocks
should be utilized. The juvenile period for grafted trees is 35 years. Terminal
portions of shoots are commonly used for taking scions after preparing them
by girdling the shoots 2530 cm below their tip 23 weeks before grafting and/
or removing the leaves and leaving a small section of the petiole to stimulate
axillary bud break. Shoots from defoliated trees are usually in a better condition
for obtaining adequate scions. Alternatively, scions may be taken from newly
developed lateral shoots initiated by pruning back mature limbs.
Removal of the apical bud of the rootstock 2448 h prior to grafting improves
grafting success, especially in the summer season when graft take is often poor
(Ogden, et al., 1984). The rootstock is cut twice during grafting, the second time
after latex ow has stopped. Grafting is best accomplished when there are warm
days, slightly cool nights and relatively low humidity (Ogden, 1984; Ogden et
al., 1986).
Field preparation
The eld should be prepared as for any other tree crop if a commercial
plantation is planned.
Seedling plants require more space than asexually propagated material.
Generally, plant spacing ranges from 4 to 12 m in-row and 712 m between
rows. The exact spacing depends on the vigor of the variety and planned
pruning practices (Morera, 1992; Granados-Friely, 1994).
Irrigation
Irrigation is recommended during prolonged dry periods to ensure fruit set
and good fruit size (Morera, 1992). Most of the mamey sapote plantations in
Central America and Mexico have no irrigation and rely entirely on rainfall,
and therefore have low productivity. Well-managed orchards in south Florida,
Mexico and Central America irrigate by applying 2.5 cm of water once or
twice per week during dry periods (Balerdi and Crane, 2009).
Pruning
Mamey sapote responds well to most pruning treatments for size management,
although there may be a negative effect on fruiting (Wasielewski and Campbell,
1999b). Formation pruning is advised for young trees and consists of removing
the tip of the main stem to induce low branching. According to Morera (1992),
the tree should be left with a single trunk and ve whorls of four to six limbs
American Fruit
311
each. Maintenance pruning consists of eliminating limbs forming narrow

angles to avoid their breakage under heavy fruit load, as well as those too
closely spaced, weak or damaged; this includes removal of water sprouts and a
reduction in the tree height to 4.05.5 m by removing or topping certain limbs.
Pruning should be performed when the plants are inactive and fruit are absent.
Topping and/or hedging with mechanical machinery is carried out to a limited
extent in Florida (Balerdi and Crane, 2009).
Fertilization
Fertilizer practices are based on observation and experience, and should
include the application of all essential elements (Morera, 1992). In Florida,
increasing amounts of fertilizer are used until the trees are 10 years old,
starting with 0.5 up to 12 kg/year of a complete fertilizer (8:3:9:2 or 8:8:8:2,
N:P2O3:K2O:Mg). Six applications are made during the rst 3 years and four
applications a year thereafter (Balerdi and Crane, 2009). Foliar application
of micronutrients and drenching the soil with chelated iron materials is
recommended two to three times per year. Similar recommendations are made
in Costa Rica, Puerto Rico and the Dominican Republic.
Yields
Depending on tree size and pruning practices, mature trees may produce more
than 200500 fruit/year, which is equivalent to 130240 kg. The plants tend
to show alternate bearing and in some years trees with no care will produce
very few or no fruit (Azurdia, 2006).
Pest management
DISEASES Several fungi attack the leaves of mamey sapote. These include
anthracnose (Colletotrichum gloeosporioides), which also infects owers and fruit;

leaf spot (Phyllosticta sapotae); rust (Uredo sapotae); black leaf spot (Phyllachora
sp.); and, in certain areas of Guatemala, Pestalotia species. Root rots (Pythium
splendens and Rhizoctonia spp.) can be a problem in poorly drained or ooded
soils. Red alga (Cephaleuros virescens) attacks during the moist rainy season.
INSECTS Insects are not much of a problem in small plantations, but probably
become more important in commercial orchards. Various scales are seen

occasionally but the damage is seldom serious; leaf hoppers can also be found.
The roots are attacked by the Cuban May beetle (Phyllophaga bruneri), which is
an important concern in young orchards (Granados-Friely, 1994). The larvae
of the Diaprepes weevil also attack the roots. An arboreal termite (Nasutitermes
costalis) has been reported to attack trees in Costa Rica and more recently in
Florida (Scheffrahn et al., 2002). Fruit ies (Anastrepha sp.) are probably the
most important concern, especially for fruit harvested during the rainy season,
and are likely to become a more serious pest as planting increases. The release
312
Chapter 12
of the parasitoid Diachasmimorpha longicaudata can be of help to control some

pests (Azurdia, 2006). Other insects in Guatemala include Diabrotica species.
Leaf miners (Phyllocnistis sp.) can damage foliage and leaves. Cutting ants
(Atta) can be a problem, especially in young plants, because of defoliation
(Vargas et al., 1999). Finally, mites (Tetranychus bimaculatus, Brevipalpus sp.)
can become serious, especially in neglected plants.
Weed management
Weed control is an important activity, especially during the early years.

Harvesting
Mamey sapote fruit must be harvested when mature or green mature;
immature fruit do not ripen properly. Fruit destined for commercial
purposes should be harvested when not completely mature, while for home
consumption it is better to wait until the fruit are fully mature. The time of
harvest will depend on cultivar and climatic conditions as great variability
occurs. Fruit maturity is established by scratching the scurfy fruit peel: if the
tissue immediately beneath the peel is pinkbrown, clear orange or clear red
orange then the fruit is ready for commercial use; if this tissue is darker in
color then the fruit is ready for home consumption; and if green or greenish
then the fruit is immature. In home gardens, fruit drop can be used as an
indication of maturity. The fruit is picked by twisting or cutting it from the
tree with a knife or pruning shears. The fruit stem is trimmed after picking to
prevent it scarring adjacent fruit.
Mamey sapote is a climacteric fruit. It can be harvested before it is completely
mature and will soften at room temperature (2228C) in a few days.
Fruit ripen in 613 days at 20C (Diaz-Perez et al, 2000) and can be stored
at 1318C and 8590% relative humidity (McGregor, 1978). Chilling
injuries occur at temperatures below 1015C, depending on the duration
of storage. Magaa can be stored for 7 days at 13C and then transferred
to 20C and show no injury on ripening (Ergun et al., 2005). Treating with
1-methylcyclopropene before storage and holding the fruit at 20C can extend
the storage period to 2 weeks when adequate decay control is obtained.
Utilization
Mamey sapote is commonly eaten fresh out of hand or as a part of various
desserts, including ice creams, milkshakes and pastries. It is also used in
American Fruit
313
marmalades and preserves, such as the one Cubans call crema de mamey
colorado. The moisture of the ripe fruit pulp ranges from 55% to 73% and
it contains moderate amounts of phosphorus and vitamin C (Table 12.2).
Mamey sapote puree has been exported to the USA in volumes ranging from
10,000 to 100,000 kg/week.
Constituents of the seed have been reported to be poisonous; however, the
seed oil is used in soaps and cosmetics and as a sedative, skin tonic and hair
revitalizer; some people eat it when freshly extracted. The milky tree sap is
said to be poisonous or irritant, and has been used to induce vomiting, expel
intestinal parasites and remove warts.
CHIKU OR SAPODILLA
Introduction
Chiku or sapodilla, Manilkara zapota (L.) von Royen, has also been named
Achras zapota L., Pouteria mammosa (L.) Cronquist, Manilkara achras (Mill)
Fosberg, Achras zapota L. var. zapotilla Jacq., A. zapotilla Nutt., Manilkara
zapotilla (Jacq.) Gilly., Sapota achras Miller and S. zapotilla (Coville). This tree
has numerous common names, including chico (Philippines), chicle tree and
sapodilla (English), naseberry, neeseberry, dilly (West Indies), lamut (Thai),
chicle and zapotillo (Mexico), chico zapote, nspero, chico and sapotillo (Honduras,
Guatemala, Mexico), zapota (Venezuela), nspero (Puerto Rico), korob (Costa
Rica), mispel and mispu (Netherlands Antilles, Surinam), naseberry (Jamaica),
chiku and chikoo (Hindi), baramasi (Bengal and Bihar, India), ciku, sawo londo
and sawo Manila (Indonesia), buah chiku (Malaya), lamut (Thailand), hong
xiem, tarn luc and xaboche (Vietnam) and sapota and sapoti (Brazil).
A related species, M. elata (Fr Alen ex Mig.) Monachino (syn. M. huberi
[Ducke] Chew. Mimusops huberi Ducke) from the eastern Amazon basin
has edible fruit. This is an upper-canopy tree (3040 m) and it is difficult to
harvest the fruit before they are eaten by birds. The fruit avor is similar to
that of M. zapota and the fruit is 34 cm in diameter. Fruit are available in the
Amazon area from February to April from owering in the previous October to
November.
Chiku is native to the warm humid areas of southern Mexico, the Yucatn
Peninsula, Central America and the north of South America, especially
Venezuela. It has been cultivated by the native people since ancient times for
the fruit and to extract chicle, which is used for making chewing gum. It
spread to the Caribbean islands, parts of South America and Florida, and was
Constituent
Naranjilla
Mamey
sapote
Sapodilla/
chiku
Red pitaya
Yellow pitaya
Acerola
55.377.3
114
0.21.9
0.10.3
1.429.7
1.23.2
0.91.3
75.0
393
0.5
1.1
23.0
1.6
0.4
89.4
36
0.5
0.1
9.2
0.3
0.5
85.4
50
0.4
0.1
13.2
0.5
0.4
81.991.1
59
0.71.8
0.080.1
714
0.61.2
0.770.82
91.6
28
0.7
0.1
6.8
0.4
0.6
87.5
45
1.2
0.2
10.9
4.0
0.7
28.2121.0
0.52.6
22.933.1
24.0
1.0
10
6
0.4
19
10
0.3
16
8.234.6
0.171.11
16.237.5
8
0.4
14
11
0.6
41
8.440.0
0.0020.025
0.0060.046
1.572.58
0.050.67
15
0.01
0.01
0.02
10.0
20004500
0.20.4
0.040.08
0.340.53
4081000
65
0.6
0.4
1.5
50
48
0.7
0.4
1.5
70
8
0.4
0.4
0.2
0.01
0.2
Pulp
Pulp and seed
Chapter 12
Proximate
Water (g)
Energy (cal)
Energy (kJ)
Protein (g)
Fat (g)
Carbohydrate (g)
Fiber (g)
Ash (g)
Minerals
Calcium (mg)
Iron (mg)
Phosphorus (mg)
Vitamins
Ascorbic acid (mg)
Thiamine (mg)
Riboavin (mg)
Niacin (mg)
Carotene (mg)
Carotene (IU)
314
Table 12.2. Composition of 100 g edible portion of different American fruits. (After Leung and Flores, 1961; Morton, 1987; INCAP and
FAO, 1992; Villachica et al., 1996; Gallozzi and Duarte, 2007.)
American Fruit
315
introduced into the Philippines by the Spanish conquerors. It then expanded

to other countries in the region, especially south and south-east Asia, where
it is very popular not only for home consumption but increasingly for export
(Campbell et al., 2005). There is more planted area in coastal India than
in Mexico, and it is also cultivated in Sri Lanka, the Philippines, Thailand
and Malaysia (Paull, 2008). Chiku is an important as subsistence crop in
tropical America, and as a local and export commodity as well as for chicle
production. Mexico a greater area devoted to chicle than to fruit production,
but the latter is becoming more important in Mexico, Guatemala and
Venezuela as the demand for chicle diminishes. Despite this, little information
is available on the specics of production, cultivars and marketing (Campbell
et al., 1997b).
Ecology
Soil
Chiku grows best in well-drained soils, from sand to heavy clays with acid to
neutral pH (Campbell et al., 1967), but also grows well in deep organic soils.
Calcareous marl and disintegrated limestone (pH 68) are the soils of its
homeland, and it therefore does well in the oolitic limestone of Florida. It is
unclear if the tree is tolerant of saline soils. The best production is obtained in
deep, well-drained, loamy soils (Vargas et al., 1999).
Climate
RAINFALL Chiku grows in drier areas with 10001200 mm rainfall and wetter
humid areas with 18003000 mm. Although the tree is very drought resistant,
supplemental irrigation is needed for proper production and fruit size in dry
areas or when the dry season is too long. It grows well on the dry Peruvian coast
with irrigation. A short dry spell apparently produces better owering.
The tree is more adapted to the tropics or subtropics where
winters are not cold. However, mature trees can withstand 2C or short frosts
with little damage. Too-cold nights are considered the most limiting factor.
The best growth occurs within 12 of the equator and at elevations of up to
10001400 m, and in warmer areas to within 25 of the equator. The average
annual temperature is 26C in the chikus area of origin, with maximum and
minimum temperatures of around 37C and 15C, respectively.
TEMPERATURE
Seedlings may require some shade during

establishment, with mature trees growing in full sun (Marshall, 1991). The
species is apparently not photoperiodic.
LIGHT AND PHOTOPERIOD
316
Chapter 12
WIND Strong winds are tolerated, but better yields can be obtained in protected
conditions because there will be less wind-damaged fruit. Young trees are
normally staked in areas where strong winds occur. The plant can withstand
salt spray (Patil and Patil, 1983).
Tree
Chiku is an evergreen, upright to spreading, low-branching tree (520 m) that
is somewhat irregular in shape. The tree is slow growing and has an extensive
root system. Grafted chiku trees are shorter (up to 15 m). The alternate, glossy,
entire leaves (515 cm long and 2.54 cm wide) are pointed at both ends and
tend to cluster at the end of the shoot. They have short internodes and give the
tree a very ornamental aspect (Fig. 12.2). The upper part of the leaf has more
stomata (Meza and Bautista, 2001). All of the plants parts produce latex
when wounded.
Flowers
Chiku owers are hermaphrodite. They have a bell-like form and are small
(812 mm), with three brown and hairy outer and three inner sepals. These
enclose a pale green to white tubular corolla and six stamens, and the stigma
extends beyond the corolla. Flowers are attached to short slender stalks arising
from leaf-base clusters at the ends of small branches (Fig. 12.2). Flowering
peaks occur in the summer in Australia (February to April), in October to
December in Mexico, at the start of the rainy season in the Philippines (April
to June) and from December to March in India. A reduction in vegetative
growth results in ower induction (Gonzalez and Feliciano, 1953).
Full petal opening occurs during the night, 4560 days after ower initiation.
The stigma that extends beyond the corolla is receptive 1 day before and 3
days after ower opening. A strong scent is produced on the day of opening
and the stigma is covered with a sticky uid. Cross-pollination by insects is
apparently necessary for low-yielding chiku cultivars. These produce little
pollen, and hand-pollination can be used to increase yields. A high degree
of self-incompatibility as well as varying degrees of cross-compatibility have
been reported for chiku; thus, cross-compatibility must be determined when
planting a new orchard for maximum production (Piatos and Knight, 1975).
Some varieties appear to be self-fertile.
Fruit
The fruit is a pendulous berry (Fig. 12.2) that can be round, oblate or
ellipsoidal to oval. It is 510 cm wide and up to 12 cm long, and can weigh
American Fruit
317
Fig. 12.2. Chiku leaves (A), owers (B), whole fruit and the transverse section of a
fruit showing the seed (C).
from 100 to 400 g (some varieties weigh up to 1 kg). The skin is thin and
smooth, and covered with light-brown scurf until it ripens. Immature fruit
are hard, gummy (from copious latex) and astringent. At maturity, the esh
color ranges from yellowish to light brown (sometimes reddish-brown) with
no latex and a smooth or sometimes coarse and grainy texture. The esh is
very juicy and sweet with a low-acid taste, and resembles the avor of a pear
(Lakshminarayana, 1980). Shiny black seeds (none to 12) are oblong and up
to 2 cm long (Fig. 12.2).
When the brown scurf is scrubbed off, the skin color is dark green in
immature fruit and changes to a light yellowish-green when it matures. The
seeds also change from soft and white in color when immature to hard and black
when mature. The fruit takes 168240 days to mature (Fig. 12.3), with growth
occurring in three stages: in the rst stage of 112 days the fruit increases in
diameter; the second stage is a 28-day transitory stage; and in the third stage
(the last 63 days) the fruit increases in length to achieve its characteristic shape
Fruit Weight (g)
Eating
Acceptability
Acceptability and Texture

(1 = like to 9 = dislike)
Chapter 12
318
Fig. 12.3. Growth of chiku cv. Subang fruit from anthesis (A) and changes in total
sugars, fruit texture and eating acceptability after the major fruit growth period (B).
Fruit harvested before 195 days (6.5 months) from anthesis did not ripen properly.
(Redrawn from Ali and Lin, 1996.)
(Selvaraj and Pal, 1984), thus following a single sigmoidal pattern (Yahia and
Gutierrez-Orozco, 2011).
There are a few genetic studies of chiku (2n = 26). Hybridization studies
were started in India in the 1950s, but new cultivars have not been produced.
The plants high degree of self-incompatibility and various degrees of crosscompatibility make a breeding program feasible (Piatos and Knight, 1975).
There is considerable variation among the seedling population in plant and
fruit characteristics. Some seedlings never bear fruit because of pollen sterility.
Large fruit size, reliably high yields, good eating quality and seedlessness are
the major objectives of selection programs. There are no well-documented
genetic collections to systematically improve this species. The emphasis so far
has been on better quality fruit and horticultural traits (Campbell et al., 2005).
The commercial development of chiku has been limited by a lack of information on its production, scarce genetic resources, serious pests and diseases,
and little marketing effort. The genetic resources available to producers have
considerably improved during the last few decades because of more efficient
propagation techniques and the identication of improved cultivars, but superior
American Fruit
319
genetic collections are still limited and there remains a need to keep looking for
improved fruit and tree characteristics (Campbell and Ledesma, 2002).
Cultivars
Numerous named chiku varieties exist, although many are probably
synonyms. The most popular varieties include Sawo Manila with oval fruit,
and Sawo Kulan and Sawo Betawi with round fruit in Indonesia; the highyielding Pineras, the large-fruited, poor-pollen Ponderosa and Sao Manila
in the Philippines; and the small-fruited Krasuey, the large-fruited Kai
Hahn and the medium-sized Makok in Thailand. India has many varieties,
including the small-fruited Kali Patti, the very large-fruited and crisp Cricket
Ball and Dwarapudi, with its large sweet fruit.
The improved cultivars Ipacuru and Tropical were released in north-east
Brazil a few years ago (EMBRAPA, 2003). Important varieties in Venezuela
include Hiplito, Martn and Reina (Aviln et al., 1988), as well as
Santiago and Delna (Meza and Bautista, 2001). Other cultivars include
Caluco, Meja and Rodriguez from El Salvador; 02, 04 and OB from
Mexico (Mrida); and Recodo-1and Recodo-2 from the Dominican Republic
(Vargas et al., 1999). In addition Tikal from Mrida is a heavy and regular
bearer with excellent avor and no astringency (Table 12.3). The most highyielding variety in Florida is the consistently early-bearing Prolic. Others
varieties include Russel and Brown Sugar (Campbell and Malo, 1973).
Several superior selections from tropical Asia and America have been
introduced to the Caribbean region, including Alano, Betawi, Hasya,
Morena, Molix, Modelo, Tikal, Oxkutzcab, Seku and Makok(Table
12.3). These cultivars were not well characterized and there was confusion
over their identity, so they were evaluated under local conditions (Campbell et
al., 1997b) and more concrete information is now available. Makok is a good
variety and is used mainly in home gardens in Florida (Campbell, 2000).
Of the numerous local selections, those of tropical America tend to have large
fruit of excellent quality and heavy production, but inferior horticultural traits.
Conversely, those of Thai origin have smaller, more elongated fruit, excellent
quality and superior horticultural traits (Campbell and Ledesma, 2002).
Cultural practices
Propagation
SEXUAL Dry chiku seeds germinate readily (within 34 weeks) after several
years storage. The sowing depth should be between 2 and 4 cm. Germination is
epigeous (Maciel et al., 1996). A trial in Honduras found that fresh seeds sown
under 60% shade started germinating after 40 days. The best germination was
seen in seeds soaked for 24 h in water (65% vs 45% for the control). Seeds
Chapter 12
320
Table 12.3. Chiku cultivars evaluated in the Fairchild Tropical Garden, Miami,
Florida. (Adapted from Campbell and Ledesma, 2002.)
Fruit
weight
(g)
Tree
Name
Origin
Fruit shape Color
Alano
Thailand
Spindle
Brown/green
150
Makok
Thailand
Spindle
Brown/green
70
Seku
Indonesia Spindle
Brown/gold
70
Modelo
Tikal
USA
USA
Flattened
Oblong
White
Brown
280
250
Betawi
Mexico
Oblong
Brown/red
230
Hasy
Mexico
Oblong
Brown/red
300
Molix
Mexico
Oblong
Brown/red
300
Elongated
Oblong
Brown/red
Brown/red
250
310
Morena
Mexico
OxKutzkab Mexico
Semi-dwarf,
multiple crops,
manageable
Dwarf, multiple
crops,
manageable
Dwarf, multiple
crops,
manageable
Large, manageable
Medium,
manageable
Large, splitting,
difcult to manage
Large, splitting,
difcult to manage
Large, splitting,
difcult to
manage
Large, manageable
Large, manageable
with cracked coats plus 24 h in 1250 ppm gibberellic acid were the fastest to
germinate, but the tallest plants were obtained with intact seeds soaked for
24 h in 2500 ppm gibberellic acid. Seed-coat removal led to the seeds rotting
(Duarte and Hurtado, 2001). The shape and weight of the seeds seemed to
have no inuence on germination, but it is better to use the largest seeds from
large fruit to obtain vigorous seedlings. Light negatively affects germination.
Seedling trees need 58 years, and sometimes longer, before they start bearing.
ASEXUAL Vegetative propagation should be used to obtain uniform planting
material and avoid the initially slow growth of seedling trees. Mist propagation
of leafy chiku stem cuttings treated with a root-promoting powder has been
successful (Rowe-Dutton, 1976). Marcotting during the rainy season is
successfully practiced in the Far East.
Cleft and wedge grafting during the wet season are used on seedling trees
of chiku or more vigorous related species such as Manilkara hexandra (Roxb.)
Dubard in India, M. kauki (L.) Dubard in Indonesia (Gonzalez and Fabella,
American Fruit
321
1952) and balata (Pouteria bidentata A. DC.) in Trinidad and Tobago (Alix and
Duarte, 1999). Veneer grafting completely covering the scions with plastic
strips that are removed after 1 month has also been successful (Malo, 1967).
T budding can be used with seedlings of about 1 year old. Scion wood should
be obtained from mature branches where the bark has turned from green to a
brownish color and with well-developed buds.
Cooler and drier periods are more suited to successful grafting. It is important
to wipe off the latex from the wounds of the scion and the rootstock with the
grafting knife just before joining them so it does not interfere with a proper
cambial contact. Vegetatively propagated chiku trees should bear in 23 years.
Field preparation
No special eld preparation is needed. Plowing and discing or harrowing will
normally be conducted until the soil reaches a proper tilth, and the positions
of the holes will then be indicated by putting stakes or marks with lime. If
the soil has been worked with machinery for many years or in other special
situations, ripping or subsoiling might be necessary to break any hard layers
in the subsoil.
Trees are planted out when 12 years old. Spacing of 1014 m is recommended for trees with a low-spreading habit and 79 m for those that are
upright (Vargas et al., 1999). If planted in exposed situations, the trees are
staked and provided with shade for the rst 12 months.
Irrigation
Young trees require irrigation to become established. Higher yields and larger
fruit are obtained if mature trees receive supplemental irrigation during the
dry season. Flowers and small fruit abscise under moisture stress.
Pruning
Chiku shows variable responses to pruning, depending on the origin of the
cultivar. Cultivars from Thailand have shown good size control and fruiting,
while those from tropical America have responded poorly to all treatments
(Wasielewski and Campbell, 1999b). As a tree with a central leader and
whorls of laterals, chiku requires little pruning. Grafted trees may require
some initial trimming to stimulate lateral growth and properly form the
canopy. Pruning of mature trees is limited to removal of dead and thin
branches and branches of the lower whorl that bend toward the ground.
Fertilization
According to Marshall (1991), mature chiku trees require at least 1.5 kg
nitrogen, 0.5 kg P2O5 and 0.5 kg K2O per tree per year. Applications are split
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into two or three per year, with one before the rainy season and one just
before the end of or after each harvest. Manure is also used. Liming and other
nutrients may be needed.
In Brazil, Simo (1971) has recommended 30 g/plant of nitrogen, P2O5 and
K2O in the rst year, 60 g in the second year, 90 g in the third, 120 g in the
fourth and 150 g in the fth year after planting. This should be applied in two
parts, at the start and in the middle of the rainy season. Plant size and yield will
determine adjustments after the fth year.
Studies in Venezuela have shown that low foliar concentrations of iron, zinc,
manganese and copper are related to poor vigor, with manganese apparently
being the most important element (Meza et al., 2001). Cutting off the top of young
plants has no signicant effect on the foliar content of nitrogen, phosphorous,
potassium, calcium and magnesium, but pruned plants show signicantly
higher iron, copper and manganese contents (Meza and Bautista, 2002).
Yields
Annual yields of 2030 t/ha (Florida), 2025 t/ha (the Philippines) and 20
80 t/ha (India) have been reported. Mature trees can yield 10002500 fruit
annually (Paull, 2008) and even 3000 fruit when 30 years or older (Vargas et
al., 1999).
Pest management
DISEASES A canker that kills affected chiku branches (Corticium salmonicolor,
pink disease) and leaf spots caused by Phaeophleospora indica in India and Septoria
species in Florida have been reported. Leaf rust (Uredo sapotae) can occasionally
cause serious damage (Campbell et al., 1967). Fruit rot caused by Diplodia
theobromae (Aviln et al., 1989) and Phytophthora palmivora is common on lower
fruit in the tree.
INSECTS Leaf-eating larvae, twig borers, scales, aphids and mealy bugs attack
various parts of the tree. As they ripen, the chiku fruit become hosts for fruit
ies such as Anastrepha suspensa in Florida. These ies are frequently quarantine
pests and limit the fruits export potential. Fruit bats often mean the tree has to
be netted for protection as the fruit ripen.
Weed management
The most common practice for controlling weeds is to mulch around the
base of the tree or keep that area clean using a hoe. Care is needed to avoid
the mulch touching the trunk. Weeds in the general area are controlled by
mowing or using a cover crop.
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Orchard protection
Although chiku is tolerant of strong winds, higher yields of less winddamaged fruit are obtained in sheltered conditions. Staking may be necessary
for young trees in strong wind areas.

Harvesting
Maturity for a particular chiku variety is judged by size and appearance.
Rubbing the brown powdery scruff bloom off the skin to determine if there has
been a change in the underlying green of immature fruit to light greenishyellow of mature fruit is not always successful as a maturity index. Waiting
for the rst ripe fruit to fall and then harvesting the largest fruit is more
reliable. Little or no latex from the skin when scratched is another indicator;
alternatively, with experience, the changes during abscission can be judged.
Some varieties shed the scruffy coat when mature. Chiku starch content
during the last 4 months declines with maturity, along with the tannin
content (Abdul-Karim et al., 1987). This coincides with an increase in sugar,
fruit softening and eating acceptability (Fig. 12.3). A study in Brazil found it
took 6 months for chiku fruit to reach physiological maturity, with a soluble
sugar content of around 21.3%. Fruit harvested at 180 days ripened 510
days after harvest (Miranda et al., 2003).
Chiku fruit have a climacteric respiration pattern (Miranda et al., 2003) and
should be carefully handled, as they are readily damaged by abrasion and
impact. After harvest, the stalk is pruned and latex is allowed to drain into
water. Harvested immature fruit shrivel as they soften, without the full sweet
taste. Mature fruit ripen in 37 days at 25C and can be stored at 15C for
about 2 weeks (Broughton and Wong, 1979). Storage under marketing
conditions should be at 20C and 85% relative humidity, which gives a
marketable life of 6 days. An ethylene peak precedes the rise in respiration
by 1 day. In a trial by Baez et al. (1997) using mature fruit with 13.2Bx at
harvest kept at 20C, a 5.1% weight loss was recorded after 6 days while
the soluble solid content increased to 25.8%. Acidity did not change during
marketing.
Chiku fruit can be stored at 5C and 12C in PVC packaging for 2 weeks.
When taken to room temperature (28C), however, they will show a rmness
loss that is associated with higher galactosidase activity. This loss is faster in
fruit kept at 12C (Miranda et al., 2002). Lower storage temperatures lead to
chilling injury and a failure to soften. Ripening can be accelerated with ethylene
(Shanmuganelu et al., 1971). According to Yahia and Gutierrez-Orozco (2011),
chilling injuries can occur at 610C depending on stage of ripeness, variety
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and handling such as previous waxing. Waxing can allow fruit to be kept for 40
days at 10C, although others have recommended 14C.
In some South-east Asian markets, the fruit is dyed brownish red to improve
its visual appeal. The fruit shape lends itself to shipping in cartons, with cell
packs of fruit graded to the same size. The limited postharvest life, difficulty of
grading for similar maturity and the problems with using low temperatures for
storage limit the fruits market potential.
Utilization
Chiku fruit has an excellent aroma and avor. It is normally served fresh
as a dessert, and is frequently chilled, cut in half and the esh eaten with a
spoon or out of the hand. Varieties with a gritty taste are avoided. Fruit juice is
sometimes used to make syrups, jellies and beverages. About 84% of the chiku
fruit is edible, containing 6975% water (Table 12.2). The chiku fruit is low in
vitamins A and C, and its acidity is due to malate.
The ripe pulp can be used dried or in sherbets, jams, fruit salads and yoghurt. It
is difficult to retain the characteristic taste and aroma of chiku during processing.
The avor volatiles imparting the distinctive odor are methyl benzoate and
methyl salicylate (MacLeod and de Troconis, 1982). The delicate avor comes
from the small quantity of volatiles produced compared with other fruits. In
Brazil, Alves et al. (1999) found that fruit had a soluble solids (SS) average of
26% and 22% total soluble sugars, of which 68% were reducing. Titratable
acidity (TA) was 0.12%, giving a high TA/SS ratio (216). The high starch and
pectin content in ripe chiku may impair juice stabilization in processing.
Canning is apparently not satisfactory, but freezing pulp puree and keeping
it at 15C gives good results (Rivas and Martos, 1978). Reports suggest that
slices can be canned with sucrose syrup, with treatment at 70C for 10 min and
irradiation with 4 u 105 rads at room temperature.
The latex collected from the tree is used to produce chicle (15% rubber, 38%
resin), a component of chewing gum that was used before synthetics. The timber
is valuable, as it is deep red in color, very hard and strong. The wood is used for
furniture and jewelry. A leaf tea is used to treat fevers, wounds and ulcers. The
plant is a source of saponin. The tree is widely planted as an ornamental and
can be an invasive weed.
PITAYA
Introduction
Pitaya (or pitahaya) is a generic term that includes several species of
columnar and climbing cacti belonging to the Cactaceae family. The climbing
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types have been classied and given different generic and specic names by
several authors. Only the climbing species is discussed in this section, using
the nomenclature of Le Bellec and Vaillant (2011). All of the so-called pitayas
that are climbers are grouped into the genus Hylocereus (Table 12.4), which
comprises 15 species. While many of these species have ornamental value
for their beautiful owers that open at night, only ve are important as fruit
producers. Four of these species have red or scarlet skin with bracts and white,
red or purple esh, containing small seeds. A fth species is the yellow pitaya
(H. megalanthus Bauer), which has yellow peel and spines and white pulp,
and a sweeter avor and larger seeds than the red species. This yellow species
has been previously named Cereus triangularis Haw., Acanthocereus pitajaya
(Jacq.) Dugand, A. colombianus Britt. & Rose, C. pitahaya DC and Selenicereus
megalanthus Haw. It has even had a new genus (Mediocactus) created for it.
The common names for pitaya include pitajaya and pitahaya (Spanish),
cuauhnochtli (Nahuatl), cierge lzard and poire de chardon (French) and nightblooming cereus, strawberry pear and dragon fruit (English). Many columnar
cacti are also called pitaya, pitahaya or pitajaya in Latin America, but these are
different species.
The commercial species of the genus Hylocereus are discussed in this section
(Table 12.4). Of the ve most cultivated pitayas, four are red pitayas. H.
costaricensis (Web.) Britt. & Rose is the most vigorous vine. It has very waxy
stems and produces scarlet fruit of 250600 g, with a redpurple pulp
and pleasant texture and taste. H. ocamponis (Weing) Britt. & Rose (syn. H.
purpusii) has very large owers (25 cm). The outer perianth segments are
reddish and the fruit are scarlet with large scales. They weigh 150400 g and
have a pleasant texture. H. monocanthus Bauer, which includes H. polyrhizus,
has very long owers (2530 cm), reddish outer perianth segments and
scarlet fruit. These weigh 200400 g and have red esh and a pleasant texture
Table 12.4. Fruit characteristics of different Hylocereus species (Le Bellec and
Vaillant, 2011).
Common name Species
Red pitaya
Yellow pitaya
Red pitaya
Red pitaya
Dragon fruit
H. costaricensis
H. megalanthus
H. ocamponis (syn. H.
purpusii)
H. monocanthus
H. undatus
H. undatus subsp.
luteocarpa
Weight (g)
Peel color
Flesh color
250600
120250
150400
Red
Yellow
Red
Redpurple
White
Red
200400
300800
100480
Purple
Rosy-red
Clear yellow
Redpurple
White
White
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and avor. H. undatus (Haw) Britt. & Rose has long green stems and 2830 cm
owers with green outer perianth segments and white inner segments. The
rosy-red fruit measure 1522 cm and weigh 300800 g. They have very long
scales, white esh and a pleasant texture and taste. The fruit of a subspecies,
H. undatus subsp. luteocarpa, have yellow skin and large scales. The fruit are
rather small with white pulp. This species is found in Nicaragua. The fth
species of this genus, according to Bauer (2003), is H. megalanthus Bauer
(syn. Selenicereus megalanthus), the yellow pitaya. This species is from South
America. It has long and slender green stems, white areoles and yellow fruit
covered with clusters of spines. The fruit weigh 120250 g and have a white
colored, pleasant and sweeter pulp than the red types.
Other cactus genera and species bear edible fruit. Lemaireocereus thurberi Britt.
& Rose, also called pitahaya, is a columnar cactus of Mexican origin that has
edible and fairly sweet but small (47 cm) fruit. The prickly pear cactus Opuntia
cus-indica Mill. is another popular species in cool, semi-arid climates, with a very
sweet and pleasant pulp. Pereskia aculeata Plum. ex Mill., the Barbados gooseberry
or lemon vine, from tropical America, is also a cactus that maintains its leaves. It
grows in areas with adequate rainfall so its leaves do not have to be shed.
Red pitayas are originally from the semi-humid areas of Mexico, Central
America and the Antilles, while the yellow pitayas are native to northern
South America, especially Colombia, Venezuela and northern Brazil.
The native people of these regions cultivated them before the arrival of
Columbus. Pitayas were practically unknown outside of their areas of origin
until about 20 years ago. Today they are distributed round the world, with
commercial production in Indonesia, Malaysia, Taiwan, Vietnam, Cambodia,
Thailand, Israel, Australia, New Zealand, the Philippines, Reunion, Spain
and the warmer parts of the USA. Latin American producers include Mexico,
Nicaragua, Colombia, Guatemala, Costa Rica, Ecuador and the Caribbean
islands. The red types are marketed as dragon fruit, with active international
trade from Vietnam, Israel and Nicaragua. It was introduced into Vietnam
more than 100 years ago by the French, and several thousands of hectares are
under cultivation.
Ecology
Soil
Pitayas prefer loamy, sandy or stony soils with good drainage. They do best in
a loose soil, rich in organic matter, with a pH of 5.56.5 and not more than
50% slope.
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Climate
The plant can stand low to high rainfall. It grows naturally in areas
with 6001300 mm annual rainfall, and sometimes even 3500 mm, although
it can grow in arid areas in Israel or Peru with adequate irrigation. It prefers
areas where wet and dry seasons alternate. It can stand some prolonged dry
periods, as in Nicaragua or Honduras where it grows with 6 months of dry
season. Irrigation is not recommended when rainfall is 15002000 mm. Excess
rain or irrigation will result in young fruit and ower abscission and plant rot
(Le Bellec, 2004).
RAINFALL
Red pitayas prefer warm tropical lowlands with average

temperature of 2129C. The red type is found growing in Mexico and Central
America from sea level to almost 2800 m and it can be grown commercially at
altitudes from 0 to 800 m, with the ideal being 300500 m. In Colombia, which
is very close to the equator, the recommendation is to plant the yellow type at
altitudes of 14001700 m, where temperatures uctuate from 1426C.
Pitayas can tolerate temperatures up to 3840C and as low as 12C with little
damage.
TEMPERATURE
LIGHT AND PHOTOPERIOD Being semi-epiphytes, pitayas usually prefer semi-
shaded locations, such as under a tree canopy. Some species, such as H. undatus,
H. costaricensis and H. purpusi, can be grown under full sun, but growth or
owering can be inhibited. In the Negev desert in Israel, 30% shade is best for
H. polyrhizus, while in the French West Indies H. trigonus grows best under 50%
shade. In Colombia, the recommendation is for 4060% shade, instead of the
full sun exposure used in the past. Longer days seem to trigger owering and
articial light is sometimes used to extend day length.
WIND The plant is fairly tolerant of wind when attached to a trellis. Problems
can occur with single-post trellises, where the plant forms a fairly bulky canopy,
depending on the sturdiness of the trellis.
Plant
Pitaya is an epiphytic climbing cactus that becomes semi-terrestrial under
cultivation. The green stem section is triangular and winged (Fig. 12.4), and
becomes grayish with age due to a waxy coating. The 23 mm circular areoles
in the stem are 35 cm apart, with three to six spines that are 14 cm long
and originate at a common swollen base. The stems emit thin aerial roots that
allow the plant to adhere to a support made from either live trees or concrete
or wooden posts, or other type of trellises. Roots that grow in the soil tend to
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Chapter 12
be rather supercial, normally not growing beyond 3050 cm and tending to

be parallel to the ground.
Flowers
The owers appear individually in a lateral position under the areoles on the
stem segments, usually on newly matured growth. They are usually large
(about 2830 cm) and white or ivorywhite and can be slightly fragrant (Fig.
12.4). In general, they open once at the onset of night (88.30 pm) and close
the next morning or mid morning. The owers have perianths with reddish to
green to yellowgreen outer segments, depending on the species. The middle
segments can be golden and the inner segments whitish or pure white. The
numerous stamens have a slender stalk and in most owers the stigma lobes
are yellow to yellowish and attached to a very long style. Depending on the
species, the plants can have from ve to eight owering ushes between May
and October/November in the northern hemisphere and from November to
April in the southern hemisphere. These ushes occur after periods of 34
Fig. 12.4. Hylocereus species showing the green stem (A), open ower (B), areoles
(C) and fruit with eshy bracts (D). (Used with permission from Dr. Lionel Robineau,
coordinator for TRAMIL [Traditional Medicines of the Islands].)
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weeks. In many cases owering will occur without fruit set because of selfincompatibility or no cross-pollination. The period from the appearance of
ower buds to anthesis is 23 weeks. From anthesis to fruit ripening takes 45
weeks for red pitaya and 45 months for yellow pitaya.
Some clones of H. undatus and H. costaricensis do not produce when taken to
new areas because of self-incompatibility or a lack of pollinating agents. The
anthers dehisce before the owers open. In their native habitats, pollination
occurs at night via bat species that feed on the nectar or via a Sphingidae
buttery of the genus Maduca. Honey-bees can play a role in pollination
during the day if owers stay open during the early hours. Manual crosspollination is performed either before or after anthesis. Before anthesis, the
bulging part of the ower is pinched so that the stigma appears and pollen is
applied on it. Usually a brush or an anther is used to put the pollen onto the
stigma of another ower. The pollen of one ower can be used to pollinate 50
owers. Pollen can be stored frozen for 39 months at 18C to 196C, while
fruit from pollen stored at 4C for 4 months loses it viability (Le Bellec and
Vaillant, 2011).
Fruit
The fruit of most Hylocereus species are red, scarlet or rosy-red; only the
yellow pitaya and H. undatus subsp. luteocarpa are yellow (Table 12.4). Red
pitayas produce large fruit that normally weigh from 150300 g to up to 800
g, depending on the species, pollination and management. The esh contains
numerous small black seeds. The yellow pitaya has smaller fruit (120250
g) and its seeds are slightly larger. The pulp is white and sweeter than that of
the red pitayas because of a higher sugar content. Scales can be short or large
and have the same color as the peel, usually combined with a green tint at the
tips. The yellow type has spines instead of scales. The esh is red or purplered
except for in H. undatus and its subsp. luteocarpa and the yellow pitaya, where
the esh is white.
Red pitayas take about 3045 days from anthesis to harvest, compared
with almost 150 days for the yellow type. Fruit growth shows a double sigmoid
curve, with the pulp betacyanin content paralleling fruit weight (Fig. 12.5).
Fruit acidity begins to increase about 10 days after pollination and peaks just
before the fruit begins to ripen, then rapidly declines (Fig. 12.5). Total soluble
solids peak as the fruit begins to ripen.
Pitayas have a chromosome number of 2n = 22 (Barbeau, 1990). Hybrids
have been produced between H. undatus and H. costaricensis to overcome
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330
Total Soluble
Solids
Total Soluble Solids (%)
Titratable Acidity (% Citric Acid)
Betacyanin (mg/kg)
Time from Pollination (Days)
Time from Pollination (Days)
Fig. 12.5. Growth of dragon fruit from pollination and the increase in fruit pulp
betacyanin content (A) and the increase in total soluble solids and acidity during
fruit growth (B). (Redrawn from Jamaludin et al., 2010; total soluble solids data from
Dr. Phebe Ding, UPM, Malaysia.)
self-incompatibility. Molecular markers that have been used to categorize

Mexican and Colombia varieties showed 92.54% polymorphism among,
within and between populations. The Colombian selection is related to most of
the Mexican selections, suggesting a common origin.
Cultivars
There are many types of red pitaya based on selections made by farmers
according to a special characteristic of the plant, not necessarily yield or fruit
quality. In Nicaragua, for example, three varieties are asexually propagated
for the export market: Lisa, Rosa and Cebra (Vargas et al., 1999). These
same varieties are produced in Costa Rica. In many cases different species
are involved, often with natural crosses among or between them. The market
distinguishes pitaya only by the color of the fruit (yellow or red) and the esh
(white or red).
Cultural practices
Propagation
Pitayas can be propagated using the small seeds, but this normally results
in variability in terms of differences in plant characteristics and lack of
uniformity, especially in productivity and fruit quality. The other problem is
that seedlings will start bearing only after 37 years. Seeds are used breeding
purposes to obtain new genotypes.
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Cuttings are normally used for asexual propagation. Superior or selected

mother plants are identied and used to supply cuttings. Cuttings should
preferably be taken at an internode on a more mature part of the branch, as
these will root more readily. After taking cuttings the pieces (2530 cm) are
disinfected and left in a cool, dry and shaded place for 25 days to cure. After
curing, the cuttings are taken to the eld and planted directly or taken into the
nursery to be planted in 25 u 30 cm plastic bags. Plants from cuttings will start
to produce after 1824 months, reaching maximum yield in about 5 years.
Pitaya cuttings put to root in sand have the greatest number of roots. Tissue
culture has also been used successfully, but the time to owering is long.
Field preparation
For an intensive cropping system the eld should be prepared as for any
fruit crop. When grown in a traditional agroforestry situation, trees are
usually already there or living posts are planted with the pitaya. This
system provides semi-shade, organic matter from the shed leaves and milder
temperatures.
Pitayas can be planted directly at the side of a trellis and tied to it. Large
cuttings (1.8 m u 810 cm diameter) are used in the traditional agroforestry
method. Several tree species are recommended for trellises in Central America
and Mexico, including Gliricidia sepium and Leucaena leucocephala, with one or
two pitaya planted beside each tree. Plant spacing depends on the cultivation
method. When a vertical trellis is used, a distance of 23 m between lines
and 20003500 cuttings per hectare will be required. With horizontal or
inclined support, the cuttings or plants can be planted much closer (5075
cm) around the trellis, resulting in 6500 plants or cuttings per hectare. A
simple system used in Central America is to bury posts of about 1.5 m long
50 cm into the ground, with the rooted or unrooted cuttings planted at their
side. These posts are spaced 2.5 m along the rows and rows are separated by
4 m, giving a total of 1000 plants/ha. The stems grow up the post; when they
reach the end of the post they will start to curve over, producing new branches
until a canopy is formed. Many other trellis systems exist, each of which has
advantages and disadvantages. Commercial plantings can be protable for
more than 10 years before requiring renewing.
Irrigation
Pitayas are usually grown with natural rainfall but irrigation is necessary
for optimal results, especially in areas with a long dry season that affects
production. Pitayas do not consume large amounts of water due to their
anatomy, but long dry periods will have a negative effect on ower and fruit
numbers and fruit size. Irrigation during dry periods signicantly improves
production. Drip irrigation is often used for these shallow-rooted plants.
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Chapter 12
Pruning
Pitayas need formation pruning. This includes eliminating all lower sprouts on
the stem while the plant climbs to the top of the trellis or support structure.
Production pruning implies removal of the tips that have been harvested
so that new shoots with owers will appear. Maintenance pruning includes
eliminating all dead, diseased or damaged material.
Fertilization
Pitayas need proper nutrition for optimal production. The plant has a
supercial root system, giving it a very quick response to fertilizer. In a
modern trellis production system, macro- and microelements should
be applied based on the results of soil and tissue analysis. Nitrogen and
phosphorus are added in formulas such as 15:15:15 or 13:20:6, with
microelements including boron (Infante-Garca, 1996). In Colombia for
yellow pitaya at a density of 1100 plants/ha, 4 t/ha of poultry manure and
1100 kg/ha of a 17:6:18:2 formula is recommended, split into three to four
applications per year (Becerra-Ochoa, 1990).
Pest management
DISEASES The most important diseases affecting pitaya are anthracnose
caused by the fungus Colletotrichum gloeosporioides; basal rot caused by

Fusarium oxysporum; sh-eye disease caused by Dothiorella species; and stem
necrotic lesions caused by Curvularia lunata. Bacteria such as Erwinia carotovora
can cause moist stem necrosis with a strong odor. Some viral diseases can also
attack the plants. The use of clean planting material, clean tools, preventive
and timely spays, and elimination and burning of damaged material helps.
INSECTS The main insect pests reported are the stem-borer beetle Cotinus
mutabilis; the sap sucker Leptoglossus zonatus (a chinch bug); the adult and
larval forms of Metamasius fabrei striatoforatus; cutting and black ants (Atta
and Solenopsis spp.) that attack the ower; and Photinus scintillans, a sucking
insect that causes ower drop. Fruit ies (Bactrocera spp.) and others have been
reported to be hosted by pitaya. Nematodes such as Meloidogyne, Helicotylenchus,
Pratylenchus and Radopholus have been reported in Colombia. Mice, rats, birds
and iguanas eat the mature fruit.
Weed management
Weeds can be a problem, especially around the plants and between rows.
The supercial roots of pitaya are very sensitive to competition from weeds
for water and nutritive elements. The semi-shade produced by live trees can
reduce aggressive weed growth.
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Orchard protection
No special requirements are known. Wind damage is not as important as for
trees, because of the size, shape and growth habit (attached by the roots to a
tree or trellises) of pitayas.

The harvest season for yellow pitaya in Colombia is from February to March
and from July to August, depending on the growing zone. In Vietnam, the
main harvest period for the red pitayas goes from July to October. The annual
productivity of an adult planting can reach 30 t/ha using a trellis system.
Yields are lower under low-input management and agroforestry systems, at
19 t/ha/year (Rodriguez-Canto, 1996; Vargas et al., 1999). For the yellow
pitaya under Colombian conditions with 2200 plants/ha, three or four fruit
per plant can be expected in year 2 with a continuous increase until year 5,
when 45 kg/plant and 10 t/ha/year are obtained.
Harvesting
Both red and yellow pitaya fruit are non-climacteric and usually are harvested
when almost fully colored. Harvested fruit should be handled carefully. For
the local market the fruit are left to mature in the shade, while for the export
market they have to be classied by weight and diameter. Harvesting yellow
pitayas is difficult because of its spines, and laborers have to wear protective
gear and use special harvest tools. The spines are removed using a knife blade
under water to avoid eye injuries. A curved harvest shear is used to detach the
mature fruit.
Fruit can be held at 6C, although chilling injury can occur after 3 weeks
and deterioration is rapid on return to room temperature. Fruit can be held for
2 weeks at 14C and for 1 week at 20C. Modied-atmosphere packaging in
polyethylene bags extends the fruit life to up to 35 days at 10C (Le Bellec and
Vaillant, 2011).
Utilization
The fruit has a texture similar to that of the prickly pear and kiwi fruit. It is
normally cut in halves and the pulp scooped out with a spoon to be eaten fresh
(especially for the yellow type, which is sweeter than the red). Chilled fruit
are preferred. The pulp, especially of the red or purple types, can be blended
as a drink or used for sherbets and salads or to make syrup. The fruit can be
sold as ready to eat after peeling and/or slicing and put into microperforated
polyethylene bags at 48C, where it can be kept for about 15 days. Pulp can
334
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be frozen at 20C and exported, although it is difficult to separate the seeds

from the pulp. Dried slices can also be produced, but their water loss and
shrinkage is great and prolonged exposure to heat affects the quality.
The composition of the edible part (Table 12.2) indicates that the pulp of
yellow pitaya is sweeter because of a higher sugar content. The fruit of both
red and yellow pitaya is low in most nutrients. The red color of the pulp is
caused by betalains, natural pigments that replace anthocyanins and have high
antiradical activity. The peel can be used to produce betacyanin and coloring
pigments, and the mucilage in the food or cosmetics industries. The fruit is
also used as an ornament on banquet or buffet tables, either entire or in slices.
The owers can be cooked and eaten, as can the young shoots. The stems and
owers of Hylocereus species have been used for medicinal purposes to treat
diabetes, as a diuretic and to help wound healing. The fruit has been found to
have a preventive effect on cardiovascular disease.
ACEROLA
Introduction
Acerola (Malpighia emarginata DC) belongs to the Malpighiaceae family, which
comprises about 60 genera and 850 species of dicotyledonous trees, shrubs
and vines. The family is mostly conned to the American tropics, with a few
native to the Old World tropics. Some species produce narcotics, while others
are grown for their fruit or as ornamentals such as the Singapore holly (M.
coccigera L.), a common ornamental from the Caribbean with holly-like leaves.
The genus Malpighia has about 30 species of evergreen shrubs and trees
native to the American tropics, through Mexico, Central America and the
Caribbean islands.
Acerola, M. emarginata DC (syn. M. glabra L. and M. punicifolia L.), is also
known as Barbados cherry, chercese, French cherry, garden cherry and West
Indian cherry. Other names include cereza de la sabana and grosella (Spanish),
semeruco and cemeruco (Venezuela), cereza nortea and corazn de paloma (Peru),
cerise de St Domingue and cerisier (French), cerejeira (Portuguese) and malpi
(Philippines). The acerola fruit is well known for its unusually high amounts
of vitamin C (1033 g/kg edible pulp). Other species of this genus tested for
ascorbic acid have generally shown lower amounts except for two species, M.
souzae and M. shaferi, with 20 and 5 g/kg of vitamin C, respectively (Asenjo,
1980).
Acerola is presumed to be a native of the Caribbean islands, Central America
or northern South America (Moscoso, 1956). Outside of the Caribbean and
Latin America, the primary source of acerola plants has been Puerto Rico,
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335
Florida and Hawaii, where cultivar development has been a primary objective.
The acerola has not attained the level of commercial production anticipated
considering its high vitamin C content. Plantings are found in Brazil (the
largest world producer with more than 10,000 ha) (Alves et al., 2007), Puerto
Rico, Barbados and the east Caribbean (the Windward Islands). The fruit is
delicate and highly perishable, and so appears unlikely to become important for
fresh consumption except in village markets. There is considerable potential for
processed products and also for the food and pharmaceutical industries, if the
costs of production, harvesting and processing can be minimized.
Ecology
Soil
A wide variety of soil types are tolerated, provided they have good drainage.
Growth is generally poor in heavy soils with poor drainage. A suitable soil pH
appears to be in the range of 56.5. Application of 4.5 t/ha of lime to soil with
a pH of 5.4 has increased yields by 400% in Puerto Rico, indicating that a soil
pH closer to 6.0 is more desirable.
Climate
Adequate moisture throughout most of the year is required for
good production because of the repeated owering cycles. The number of
cycles depends on temperature and the rainfall pattern. Although adapted for
cultivation in semi-arid regions, the greatest production occurs in regions with
an evenly distributed annual rainfall between 1200 and 1750 mm/year. Under
high-rainfall conditions (25003125 mm) Cercospora leaf spot (C. bunchosiae)
is a potential problem and can defoliate trees. Leaf abscission occurs in drier
areas, with growth resuming during the rainy season (Gonzaga-Neto and
Soares, 2004). Irrigation is required for successful commercial production in
areas with seasonal rainfall.
RAINFALL
Acerola thrives in the warm lowland tropics. It is best adapted

to sea level, but does fairly well at all elevations in Puerto Rico and to 150 m in
Hawaii. The ideal temperature is around 26C. Growth continues throughout
the year at 1532C, while night temperatures of 1015C completely prevent
growth. The plant does not tolerate frost or temperatures below 7C for extended
periods.
TEMPERATURE
The quality of acerola fruit is largely inuenced by

solar radiation, with a positive correlation between ascorbic acid content and
light intensity. Shading increases the leaf size and reduces the level of ascorbic
acid in the fruit.
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Chapter 12
WIND Plant height can be readily controlled by pruning, but acerola still
benets from the use of windbreaks. Trees established from rooted cuttings can
be blown down by gusts of 64 km/h.
Tree
Acerola is a shrub that can be pruned and trained to a single trunk to form
a tree of about 4.68 m tall. The root system is shallow (mostly 610 cm
below the surface) and equal in diameter to the canopy. Growth habit among
seedling populations varies from semi-prostrate to compact or open types
with upright, spreading or reclining branches. It has been observed among
the experimental population grown in Hawaii that trees producing acid fruit
are generally more upright and compact, while those producing sweet fruit
generally exhibit rank growth and multiple leaders that are widely spread
and rather easily torn by winds. In both types the branches are woody, with
numerous lenticels, and are characterized by short lateral spurs.
The leaves are simple, entire and oppositely arranged, and the shape ranges
from elliptical to oval and ovate with an apiculate apex (Fig. 12.6). Leaf size
varies from 2.53.8 cm wide and 5.06.4 cm long, with shaded leaves being
much larger. Leaves of the sweet type are somewhat undulated. Young leaves
and stems are slightly pubescent.
Flowers
The owers are 2.02.5 cm wide, with ve petals ranging in color from white
to various shades of pink, depending on the clone (Fig. 12.6). The owers have
10 stamens, three styles and three carpels fused into a superior ovary. Flowers
are produced in clusters from leaf axils on new terminals and lateral spurs.
Floral initiation occurs 810 days before bud emergence, with anthesis 7
days later. In Hawaii, owering occurs in cycles approximately a month apart,
commencing in March or April and extending into November or later.
Mean fruit set from open pollination ranges from 1.3% to 11.5%; controlled
self-pollination increases fruit set from 16% to 55%. Cross-pollination
signicantly increases fruit set farther, from 32% to 74%. Reciprocal crosses
sometimes show different fruit-set percentages, suggesting a degree of
incompatibility. Pollen-dissemination studies indicate poor wind distribution;
owers picked during the mid afternoon showed only 1.24 pollen grains per
stigma. Insects are considered to be the major pollinating agents. Honey-bees
and syrphid ies are the only insects readily seen in small numbers. However,
honey-bee hives placed in the orchard do not increase insect activity and fruit
set is not improved. Three days after pollination there is a period of abscission
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337
Fig. 12.6. Acerola owering shoot with leaves (A), fruit and longitudinal section
showing seeds (B), and ower showing 10 stamens and stigmas (C).
at the peduncle, with a second period 1618 days after pollination. Studies
on the reproductive morphology of the owers found that seedlessness
may be the result of: (i) failure of fertilization, followed by parthenocarpic
development; (ii) successful fertilization with developing-embryo abortion,
caused by failure of endosperm development; and (iii) ovule sterility, caused
by absence of division or delayed divisions of the megaspore mother cells,
disintegration of nucellar cells and abortion of the embryo sac before
anthesis.
Fruit
The fruit resembles the true cherry but is a three-carpellate drupe. On the
surface, the carpels are faintly discernible as three shallow lobes (Fig. 12.6).
The eshy mesocarp contains three triangular ridged stones or pyrenes, each
capable of holding a seed. The seeds have extremely low viability. The embryo
is morphologically mature about 15 days after anthesis. Fruit size ranges from
approximately 34 g to as large as 10 g. The color of mature fruit ranges from
orangered or red to deep purplish-red. The skin is delicate and easily bruised.
In Hawaii, the fruit matures 2122 days after oral anthesis (Fig. 12.7).
Green fruit 12 days after oral anthesis have been reported to have ascorbic
acid levels of 2736 g/kg pulp (Fig. 12.7). This increased to a maximum of
Chapter 12
Ascorbic Acid (g/kg)
338

Fig. 12.7. Acerola fruit growth of clone 269-2 and ascorbic acid content of clone
5A-28 at different stages of development. (After Miyashita et al., 1964; Nakasone
et al., 1966.)
slightly over 40 g/kg at 15 days after anthesis. After attaining the maximum,
the ascorbic acid content rapidly declined to 2426 g/kg pulp by day 24, when
the fruit was fully ripe (Nakasone et al., 1966).
Cytogenetics and breeding
Information on the cytogenetics and genetics of acerola is scarce. Miyashita
(1963) reported 2n = 20 for a clone. The breeding objectives are to select
desirable cultivars and develop cultural practices and processing methods, and
to understand the biochemical aspects of the fruit. Further selection criteria
emphasize tree type, fruit yield, ascorbic acid content, fruit size, juice yield,
rmness and ease of propagation from cuttings. In Florida, cold tolerance is
also considered. Approaches involve planting large seedling populations with
wide diversity.
Cultivars
The pH of the juice from acid types ranges from 3.1 to 3.3, while that of the
sweet types ranges from 3.4 to 3.6 (Table 12.5). Acid and sweet types contain
different ranges of ascorbic acid: sweet types have less than 20 g/kg and acid
types up to 29 g/kg. Malic acid makes up 2550% of the total acids of tart
clones and is possibly absent from semi-sweet clones.
The acid types generally give better yields and have more desirable tree
forms. The sweet cultivars are generally developed for home planting and
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Table 12.5 Characteristics of sweet and acid acerola cultivars evaluated in Hawaii
(Nakasone et al., 1968). The yields are for 2-year-old trees.
Cultivar
Fruit mass
(g)
Juice pH
Sweet
Manoa Sweet
Tropical Ruby
Hawaiian Queen
Acid
Maunawili
J.H. Beaumont
C.F. Rehnborg
F. Haley
Red Jumbo
Total
soluble
solids (%)
Ascorbic
acid
(mg/100 g)
Fruit yield
(kg/tree)
6.2
7.2
6.1
3.6
3.5
3.5
10.0
9.0
9.6
1537
1437
1577
7.8
5.3
5.1
4.3
6.9
6.2
4.5
9.3
3.1
3.3
3.3
3.2
3.1
8.9
11.0
8.3
8.2
8.7
2010
2820
1717
2663
2183
4.0
23.5
32.0
21.2
14.1
potential use in baby foods and products requiring minimum acidity. Puerto
Ricos selection B-17, an acid-fruited type, produces a fruit with an average
weight of 912 g and yields of 8.112.1 t/ha from 4-year-old trees. Juice yield
is as high as 73% by fruit mass, with vitamin C content from 13.322.5 g/l
juice (Arostegui et al., 1954). Florida Sweet has upright, open-type growth
and some cold tolerance and is readily propagated by cuttings. It produces
rm fruit of 14 g. Brazil has introduced some new varieties in recent years:
CPATSA 2.3, CPATSA 4.1, CPATSA 6.1, CPATSA 9.1 and CPATSA 14.3.
These varieties are capable of producing more than 100 kg/tree/year. They
have bright and attractive red-colored fruit that weigh 810 g each. The pulp
contains more than 2000 mg/100 g vitamin C (Gonzaga-Neto and Soares,
1994). Several clones have been selected with high yields and large fruit in the
northern Brazilian state of Cear. These include BRS 236 (Cereja), which has
a very high vitamin C content, and BRS 237 (Roxinha), which has redpurple
pulp, high consumer desirability and a good shape that demands less pruning
(de Paiva et al., 2003).
Cultural practices
Propagation
Seed propagation is used primarily to produce seedlings for selection purposes
or rootstock, or if grafting is to be used for clonal propagation. Because of
their high heterozygosity, seedlings are not recommended for production
purposes. Large quantities of pyrenes are necessary, because as many as
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Chapter 12
5060% are devoid of viable seeds. Germination usually occurs within

2030 days and is epigean (de Jesus et al., 1998). Seedlings 45 cm tall are
transplanted into individual containers, 78 cm in diameter, for further
growth. Six- to 8-month-old seedlings, well-hardened under direct sunlight,
can be eld-transplanted.
Propagation by cuttings allows large quantities of clonal plants to be
obtained within a relatively short time. Cuttings 2025 cm long and 0.64
1.3 cm in diameter, with some leaves retained, are treated at the base with
a root-inducing hormone and stuck in a medium such as vermiculite, sand
or a mixture of porous material under mist. Cuttings root in 6 weeks, with
differing rooting abilities among clones. Leafy terminals from short laterals are
superior to other cuttings. Grafting on seedling rootstock is recommended only
if the rootstock can provide special advantages, such as disease or nematode
resistance; otherwise it is costly and time-consuming and plants propagated by
cuttings should be used instead. Side-veneer grafting with a terminal scion is
the best method and the branch where the scion will be taken from does not
need any special previous preparation. Grafting on related species, looking for
resistance, has not been successful (Alix and Duarte, 1999).
Field preparation follows conventional practices. Field transplanting can be
done at any time of the year where irrigation is available. In rain-fed areas,
transplanting may be restricted to the beginning of or during the rainy
season. Approximately 15 g of a complete fertilizer, such as triple-16, should
be placed at the bottom of each hole. A postplanting application of 710
g fertilizer is spread and mixed with the surface soil and irrigated. Spacing
depends on the growth habits of the clones selected for planting. Clones with
compact, upright growth may be planted in a hedgerow system with about
2.44 m within rows and 4.6 m between rows. Conventional orchard spacing
is 3.7 u 4.6 m to 5.5 u 5.5 m with tree densities of 478 and 332 trees/ha,
respectively.
Irrigation
Where rainfall is well distributed throughout the year, irrigation may not be
necessary. However, in most tropical areas with a well-dened dry season,
irrigation is necessary to maintain the best levels of production and fruit size.
Pruning
Acerola height can be readily controlled by pruning. Pruning begins at the
nursery stage in order to train young plants into a manageable tree type.
Plants trained to a single trunk system, with scaffold branches produced
about 6090 cm above ground, appear to be a desirable type in terms of
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341
weed control and harvesting. Bearing trees benet from light pruning when
fruiting is complete. Some cultivars have rank growth and long, widely
spread, multiple leaders that should be cut back to encourage more axillary
branching. Others produce thick, compact growth, which may require some
thinning each year. Pruning performed during the non-owering period
leads to a long period between pruning of primary branches and production
of owers on secondary branches, compared with pruning in the owering
period. Flower-bud production appears to occur between 15 and 18 days after
pruning (Miyashita et al., 1964).
Fertilization
Nitrogen deciency has the strongest inuence on growth and yield.
Deciencies in phosphorus, boron, sulfur and iron result in some growth
depression, with signicant reductions in fruit yield. Omission of potassium
reduces tree growth, with no effect on yield in the reported test; deciencies
in magnesium and manganese are also without apparent effect. The owering
cycles of acerola trees occur almost monthly to bimonthly, so several
applications of fertilizer are necessary each year. Recommendations in Brazil
for the rst year are 400500 g simple superphosphate, 300400 g KCl and
200 g calcareous dolomite per tree, to be added in the planting hole, plus 10
20 l cured manure, especially in light-textured soils where nematodes could be
a problem (Gonzaga-Neto and Soares, 1994). For the second and third years,
400 g calcium nitrate or ammonium sulfate, 400 g simple superphosphate
and 200 g KCl supplemented with 1520 kg of cured manure per tree per year
are recommended, and in the fruit bearing phase 6001000 g ammonium
sulfate or calcium nitrate, 600900 g calcium superphosphate and 200 g
KCl/tree/year should be applied (Simo, 1971).
Pest management
The acerola is relatively free from serious fungal diseases. In Hawaii,
Cercospora leaf spot (C. bunchosiae) occasionally poses a serious problem of
defoliation. Incidence of this disease is low where rainfall is low to medium.
Approximately 23 different pests on acerola have been listed in the Caribbean
region, but none has been found to be serious. Sucking insects, particularly
scale insects, pose a threat. Fruit ies are a constant threat if fresh fruit is the
preferred use. Some species of birds can cause serious fruit loss. Root-knot
nematode (Meloidogyne incognita) is one of the limiting factors to production in
Florida and Puerto Rico.
Weed management
A considerable amount of weed control can be achieved by heavy mulching
with organic material or by the use of polyethylene sheets.
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Chapter 12

The acerola fruit is thin-skinned, delicate and highly susceptible to bruising.
Whether the fruit are to be eaten out of hand or used for processing or
extraction of vitamin C, they should not be left on the tree as they abscise and
fall on ripening. Half to fully ripe fruit can be used for any purpose. Maturegreen fruit are suitable for vitamin C extraction, inasmuch as these fruit
have a much higher vitamin C content (Fig. 12.7). It has also been shown
that there is little or no increase in fruit size after 18 days from anthesis (Fig.
12.7). Since acerola fruit mature and ripen in 2225 days, harvesting should
be done every other day. If all ripe fruit and those just changing color are
picked then the harvesting cycle may be stretched to every 3 days without loss
of fruit. Acerola is harvested by hand, although shaker harvesters have been
evaluated.
Commercial yields can be expected from 2-year-old trees, with some clones
in Hawaii producing as much as 2332 kg/tree/year. Older trees give higher
yields and will continue to produce for 1520 years. An orchard of 200 trees
should produce 27224082 kg/year.
The fruit are highly perishable after harvest and losses are mainly caused by
physical injuries. The skin is also very thin, which lends to rapid moisture loss
if temperatures are high and relative humidity is low. Harvested fruit should
be stored in the shade, as fruit left exposed to sun in the eld can lose as much
as 25% of their vitamin C over an 8-h period. Fruit should the processed
without delay or refrigerated immediately (Alves et al., 1995).
Utilization
Besides eating out of hand, the fruit can be juiced for punch and jellies, and
used in the preparation of gelatin desserts, salads and sherbets, and for
fortifying other juices that are low in vitamin C (particularly as a supplement
to baby fruit juices). The juice can also be used to prevent the oxidation of fruit
in salads and fruit cups, simultaneously enhancing the vitamin C content.
Processing the fruit into juice results in little loss of vitamin C (Mohammed,
2011). Hot- and cold-pressed frozen juice can retain about 85% of the
original vitamin C content after 8 months of freezer storage. Depending on
the size of the fruit and concentration of ascorbic acid, one or two fruit can
provide a persons recommended daily allowance of vitamin C. Acerola, like
most fruit, is considered a fair source of provitamin A (408 IU vitamin A),
but is low in calcium, phosphorus and iron, and the B vitamins thiamine,
riboavin and niacin (Table 12.2).
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NARANJILLA
Introduction
Naranjilla, Solanum quitoense Lam. (syn. Solanum angulatum R. & P.), belongs
to the Solanaceae family. Its name means little orange and is derived from its
round shape and orange color. In Spanish-speaking countries it is called lulo
(Colombia, Peru), naranjilla (Ecuador and most other counties), naranjilla
de Quito and naranjilla de Castilla. In English, it is called naranjilla or Quitos
orange; in French, morelle de Quito; and in Dutch, gele terong.
Related species include the cocona (Solanum sessiliorum, formerly S. topiro
Humb. & Bonpl. and mistakenly called S. hyporhodium) (Martin et al., 1987);
the pepino or melon shrub (S. muricatum Alt.); and the tree tomato, named
tamarillo in New Zealand to make it more appealing to the export markets
(S. betaceum, formerly Cyphomandra betacea Sendt.), which is a native of the
Andean highlands, producing in the higher tropics and warm subtropics.
Physalis peruviana, the cape gooseberry, is another member of this family that
bears edible fruit and has become popular.
Naranjilla is native to the higher parts (12002200 m) of the humid oriental
foothills of the Andes, from southern Colombia to Ecuador and Peru. Some
types are found in the central and northern Andes of Venezuela and Central
America, and also in areas close to the Amazonia. Naranjilla is better adapted
than cocona to higher and cooler zones.
The fruit is much appreciated and is used for preparing beverages, jams
and other sweet preparations. The main producers are Ecuador and Colombia,
since the fruit is very popular in these countries. About 10,000 ha is planted
in Ecuador, while in 2009 Colombia planted 6490 ha, producing 53,160 t of
fruit destined for juice and concentrate production (Ministerio de Agricultura
and Desarrollo Rural de Colombia, 2009). Small plantings are found in Peru,
Panama, Costa Rica and Guatemala. Naranjilla is fairly unknown in the rest of
the world.
Ecology
Soil
Naranjilla requires light, fertile, well-drained soils (Martin et al., 1987) that
are rich in organic matter (ideally >10%). It will grow well in loamy soils or
sandy and clay loams (Gallozzi and Duarte, 2007). As with many Solanaceae,
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Chapter 12
naranjilla should not be cultivated in soil where it or other members of

this family have been previously grown, because of root rot and nematode
problems. It is preferably rotated with a non-susceptible crop for a couple of
years to reduce the inoculum levels. Excess soil moisture that favors rootinfecting fungi should also be avoided.
Climate
RAINFALL Moist tropical highlands are the best place for naranjilla, as it
requires annual rainfall of 10002800 mm with an optimum of 2500 mm.

With irrigation, it grows well in dry areas such as the Peruvian coast.
In the oriental parts of the Andes this crop does well at
elevations of 16002400 m with the optimum being 1800m; the acid type
can be grown above 2000 m. At higher latitudes the altitude can be lower.
The temperature range for this crop is 1624C, with an optimal temperature
of 1718C. It therefore does well on the central coast of Peru, which has a
subtropical climate with cool winters and not excessively hot summers. This
species requires cooler locations than cocona, although both can be grown in
areas where the climatic conditions for both species overlap.
TEMPERATURE
Naranjilla tolerates certain amounts shade. In

many cases, especially in the Amazonia, it is planted under partial shade given
by trees that are left after clearing the forest. The plant can also grow and
adequately produce under full sun. No specic information is available about
photoperiodic requirements, but since it owers almost continuously there
appear to be none.
WIND There is not much information on the wind tolerance of naranjilla.

Being a rather small plant with a well-anchored root system, it should not be
very susceptible to uprooting by winds. However, its large and soft leaves will
probably be severely damaged.
Plant
Naranjilla is a spreading herbaceous shrub of up to 23 m. The stems are
thick, cylindrical and pubescent, and become woody with age. The alternate
leaves are oblongovate and up to 60 cm long and 3545 cm wide, woolly and
soft (Fig. 12.8). Spines are present in the stems, petioles, midrib and lateral
veins, as well as in the upper and lower parts of the leaf blade. In some types,
however, the leaves are spineless. Young leaves, stems and petioles have a coat
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345
Fig. 12.8. Naranjilla leaf (A), ower (B) and fruit (C). (Used with permission from
Dr. Jorge Leon 2000. Botnica de los Cultivos Tropicales, Agroamerica del IICA,
Costa Rica.)
of purple stellate hairs. The primary root (being of sexual origin) is pivotal but
the secondary root system is fairly supercial, not going below 5060 cm.
Flowers
The pentamerous fragrant owers (34 cm in diameter) appear in short
axillary clusters of ve to 10 (sometimes 12) owers at each axil that open
sequentially. The ve sepals form a greenish persistent calyx. The ve petals
are creamy to white on the upper surface and purple and hairy on the
lower surface, and form a star-like gure (Figure 12.8). The ower has ve
prominent yellow stamens and the stigma is also yellow. Flower buds are
covered with purple hairs before opening. Once the plants starts to ower it
will do so almost continuously with certain uctuations, which means that
harvesting will follow the same pattern. A plant can produce around 1000
owers during its lifetime, with 510% of them setting fruit.
Naranjilla owers are hermaphrodite and can be self-fertile (Martin et al.,
1987) or show allogamy, depending on insects for pollination. Flowers
can have short, medium or long styles, and the last two types can be crosspollinated. Pollination is performed by insects such as honey-bees and bumblebees.
346
Chapter 12
Fruit
The fruit is a globose to ovoid berry (Fig. 12.8) that is produced in clusters
of three to six (Martin et al., 1987). It can measure 410 cm in diameter.
When mature, the external color can be yellowish-orange to deep orange
and the peel is covered by a brown hairy coat that protects the fruit until fully
ripe. These hairs can be fairly hard and are rubbed off after harvest to show
the bright orange color of the smooth, leathery and thick peel. The vepointed calyx is persistent. Internally the fruit resembles a tomato, with four
compartments separated by membranous partitions. These are lled with
a translucent, sticky, juicy green or yellowish-green pulp, which has a very
pleasant acid avor and contains numerous (around 1000) pale-buff, thin,
at, greenish-yellow seeds, measuring about 3 mm. Because of its sequential
and continuous owering habit, the plant will simultaneously have owers
and small, medium and ripening fruit. Fruit mature in 5060 days (Martin
et al., 1987). At processing, the whole pulp is used leaving only the peel. This
differs from cocona, which has a hard hoof and partitions enclosing the juicy
pulp with the seeds and has a different avor and texture. Cocona is used for
other purposes.
Naranjilla (2n = 22) apparently has no wild ancestors. The plant seems to
have been domesticated fairly recently since it shows little variation for a
domesticated plant (Barbeau, 1990; Heiser, 2008). A form with small spines is
grown in Colombia, while unarmed forms are grown in Ecuador.
Breeding, selection and evaluation
Little has been done toward improving this plant, although modest efforts
have been made in Colombia, Ecuador and at Indiana University in the USA.
Interspecic hybrids have been shown to be very useful. An interesting subject
is to create naranjillas for higher altitudes by crossing them with S. felinum
and S. vestissimum. The possibility of doubling the number of chromosomes of
the nearly sterile hybrids with cocona, S. sessiliorum, offers possibilities. Such
polyploids are soon to be tested in Ecuador (Heiser, 2008).
Cultivars
According to Heiser (2008), naranjilla could become more widely appreciated
if more efforts are made to improve it. It has many disease, insect and
nematode problems that signicantly reduce yields or force producers to
handle it as an annual crop, because after the rst year the plants start
deteriorating and dying. A nematode-resistant cultivar developed from hybrids
with the wild S. hirtum was released in Colombia in 1998, but only about 5%
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347
of the Colombian area is currently planted with it. Another factor responsible
for the low production of naranjillas is that the plant has rather strict climatic
requirements, particularly with respect to rainfall.
In Colombia, most cultivars belong to the botanical var. septentrionale and
var. quitoense. The former has spines in the branches and leaves, while the
latter is spineless with less-acid fruit (Casierra-Posada et al., 2004). In Ecuador,
naranjillas have been distinguished not as dened varieties but rather by
groups: Naranjilla Agria (acid naranjilla) is a vigorous, insect-resistant plant
with round, yellow fruit that are slightly compressed at the poles and have a peel
that is resistant to transport. The acid pulp is very appreciated in the market
and is used in ice cream, refreshments and prepared foods. Naranjilla Dulce
(sweet naranjilla) is more susceptible to insect attacks and has large, round,
orangered fruit with a thicker peel that is less resistant to transport, handling
and storage. The fruit is not as popular in the market as the acid form, but is
used in sweets, refreshments and gelatins.
At present, two articial interspecic hybrids of S. quitoense u S. sessiliorum
released by INIAP are widely grown in Ecuador. These hybrids are more vigorous
and productive than the normal naranjilla and enough fruit is produced to
allow some exports. Only a small percentage of the crop in Ecuador is from the
original naranjilla with its very low yields. About a third comes from the Puyo
hybrid, a cross of S. quitoense with a small-fruited, wild variety of S. sessiliorum.
This hybrid produces small fruit and has to be sprayed with a low concentration
of 2,4-D at owering to increase fruit size. The remaining production comes
from another hybrid named Palora, a cross of naranjilla S. quitoense (Baeza
Roja) with a large-fruited cocona S. sessiliorum (Morona Grande). The
Palora hybrid is tolerant to most naranjilla diseases. This is an interesting new
option for the naranjilla growers of Ecuador. It is easy to propagate by cuttings,
and tolerant to anthracnosis and neck-rot (Phytophthora). It can be attacked
by Neoleucinodes elegantalis, but tolerates other insects (Rodrguez et al., 1994).
Both hybrids have orange instead of green juice, which has created a certain
reduction in acceptance despite their larger fruit. Fruit from the hybrids are sold
for less than those of the true naranjilla, but this is compensated for by much
larger yields.
Cultural practices
Propagation
SEXUAL Naranjilla, except for the two above-mentioned naranjillacocona
hybrids, which are sterile, is normally propagated by seeds (Martin et al.,
1987). This method is fairly safe in maintaining genetic uniformity since seeds
are obtained from the best fruit (i.e. large size and desired form) from healthy
and productive plants. Special care should be taken to ensure the mother plant
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Chapter 12
is free of Corynebacterium, which can be transmitted by seed. Propagation is

not difficult as the seeds have no dormancy; the pulp from mature fruit is put
in a blender with water and blended for a short while. This mixture is passed
through a sieve and the seeds are separated by otation, eliminating all pulp
residues. After cleaning, the seeds are dried in thin layers in the shade and are
then ready to be sown. Seeds should be handled like any vegetable seed. In the
past, small seedlings were transferred to bags to become larger; currently, seeds
are germinated in 60- to 72-cell trays and transplanted directly to the eld
when they reach 15 cm, provided irrigation is available or the rainy season
has started and ideally on a day when rain is expected. If this is not the case
then larger bags will be used to take taller plants to the eld. Fruit production
starts 912 months from seed sowing (Martin et al., 1987), following about
23 months in propagation trays and 78 months in the eld.
ASEXUAL Cuttings can be made by taking lateral shoots from adult plants
in their second year or 25 cm terminals of branches that are not excessively

woody, with three to four buds. Cuttings are placed in nursery bags after
disinfection. The resulting plants will be less vigorous than those from seedlings.
They lack pivotal roots, leading to a more supercial root system. They will start
producing sooner, but the lifecycle will be shorter. Grafting has been tried on the
nematode-resistant stock S. torvum with success (Martin et al., 1987), with cleft
grafting using a terminal scion. In Florida, S. macranthum has also been used as
a stock for cleft grafting when plants reach 60 cm. Other grafts have also been
successful (Morton, 1987).
Land preparation is carried out as for planting a vegetable crop. Transplanting
can be done directly from trays containing small (1015 cm) seedlings. If this
is the case then soil preparation and irrigation must be fairly adequate. Larger
plants (2030 cm) in bags can be used in conditions that are less adequate.
Plants should be set 1.82.5 m apart under appropriate growing conditions.
In the Amazonia, spacing of 2.02.5 m in triangles is usually used (Villachica
et al., 1996). In every new situation an evaluation must be conducted to
establish the best distance for that particular area. Naranjilla plants usually
live for 34 years, although this decreases to around 1215 months if the
fungal or nematode presence in the soil is too high. In such situations the
plants will start to die progressively, starting as fruit begin to ripen (as this
creates a stress situation).
Irrigation
Naranjilla is most commonly grown without irrigation, depending on rainfall.
Irrigation using furrows or drippers is needed in places with a long dry season
or little or no rain. The plant is very succulent and does not tolerate long
periods of drought.
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349
Pruning
The recommended pruning practices include: (i) formation pruning, which
consists of eliminating all basal sprouts of the main stem below 4050
cm; (ii) maintenance pruning, which consists of eliminating the lower and
old yellowish leaves to improve air movement and allow for easier weeding
(normally performed after 1824 months in the eld); and (iii) renovation
pruning, which is conducted if there is a heavy anthracnose or insect attack
that damages too many fruit and there are no fruit in the basal part. In
this case, cutting the plants back to 80100 cm to stimulate new growth is
advisable.
Fertilization
Naranjilla prefers rich soils with high organic matter content. A soil analysis
should be performed 23 months before planting to determine the proper
amounts of fertilizer to be applied, starting at transplanting. A general
recommendation is to apply nitrogen, phosphorus and potassium at dosages
of 120 kg/ha nitrogen, P2O5 and K2O. If fertigation is not available then the
fertilizer should be applied in a circular band around the stem and at 20 cm
from it when the soil is moist. Nitrogen and potassium should be split into
two or three applications during the rainy season, while phosphorus can be
applied all at once with the rst application at the start of the rainy season.
The addition of manure to improve organic matter and biological activity in
the soil is very effective.
Pest management
DISEASES The main diseases of naranjilla include damping-off, anthracnose
and leaf spots. Damping-off, caused by Rhizoctonia solani, is mainly a seedbed problem; recommended sanitary measures should be used. Anthracnose
caused by Colletotrichum gloeosporioides can damage foliage and fruit under
very humid and rainy conditions. Leaf spots caused either by Septoria solanicola
or Cercospora can be a problem, as can Fusarium wilt and potato blight caused
by Phytophthora infestans. Some viral problems can exist, but they are not
important at this time (Rodrguez et al., 1994; Gallozzi and Duarte, 2007).
Bacterial rot caused by Pseudomonas solanacearum can be a serious problem,
and plants infected with bacteria should be uprooted and burned far away from
the eld.
INSECTS The main insects are the fruit-eating worm Neoleucinodes elegantalis;
the stem-base borer Faustinus apicalis; the stem borer Alcidion; the leaf worm
Machanitis; and aphids Myzus persicae (Rodrguez et al., 1994; Gallozzi and
Duarte, 2007). There are reports about fruit ies attacking the fruit (Villachica
et al., 1996), and the white or West Indian scale Pseudaulacaspis pentagona can
350
Chapter 12
be a serious problem in Central America. In addition, the root-knot nematode

Meloidogyne incognita can severely reduce production. Grafting on the relative S.
macranthum has been attempted to reduce this problem, resulting in improved
yields and plants living beyond 3 years. Other nematodes such as Rotylenchulus
reniformis and Helicotylenchus can also be a problem, but are minimized by the
use of abundant organic matter and crop rotation.
Weed management
Under rainfall and irrigation, weed control during the dry season is required
to reduce competition. At the same time, however, a certain number of weeds
can be left to maintain the microclimatic, as long as they are not hosts of
Neoleucinodes species or any important fungal disease. If weeding is performed
with a machete or hoe then the tools should be disinfected between every plant
to avoid disease transmission, particularly if bacterial diseases are present.
Orchard protection
There are generally no strong winds in the areas where this crop is grown, and
protection from neighboring trees is frequently found.

Plants can be harvested for 13 years after 912 months in the eld. An
average production might be 135 fruit/plant/year or 1015 t/ha/year
(Villachica et al., 1996). Yields are much lower in most areas where naranjillas
have been planted for a long time because of disease unless the naranjilla u
cocona hybrids are used, which allow yields of around 18 t/ha/year instead
of 45 t/ha/year. Soil-borne diseases and nematodes are the main reasons for
low yields. Colombia reports yields of 714 t/ha/year (Gallozzi and Duarte,
2007).
Harvesting
Naranjilla plants will start production 78 months after transplanting. For
local markets, fruit are collected when half or two thirds of the fruit color is
orange, while for distant markets fruit are harvested when the orange color
starts to appear. Harvesting as the fruit start to turn orange is not ideal as the
nal fruit avor is poorer, but it does increase postharvest life. A sharp knife
is usually used for harvesting, but some people twist and then pull the fruit
to break its very short peduncle. The fruit should be harvested early in the
morning since the peel hairs will be moist and will not oat in the air; later
they dry and are dispersed, and can cause a very unpleasant irritation in some
peoples eyes, nose and skin. Harvested fruit should be placed into bags or
boxes and left in the shade while in the eld. Harvesting is usually conducted
once a week.
American Fruit
351
The harvested fruit are classied by color, size, type and degree of damage.
The fruit must be cleaned to remove dust and the hairs from the peel. This
can be done by putting the fruit in jute bags and moving them so the fruit
rub against each other or the bag. Alternatively, the fruit can be washed
using a disinfectant (chlorine, ozone or a specic disinfectant) and rubbed
mechanically or by hand with a cloth to eliminate the fuzz. In some cases, the
fruit are waxed or coated using sucrose-derived waxes to reduce water loss
and give an attractive gloss. After washing and/or coating, the fruit are left to
dry.
Naranjilla fruit are climacteric. If harvested at the half-mature stage
(yelloworange color) then they can be held in good condition for 8 days at
room temperature, or stored for 12 months at 710C and 7080% relative
humidity. Coatings and PVC lms can create a modied atmosphere and
prolong the postharvest life. Naranjilla fruit stored at 515C retain their
appearance and quality for 3 weeks (Daz and Manzano, 2002). Fruit harvested
almost fully colored or taken out of cool storage should be held at 2125C to
develop their full color, aroma and avor.
Utilization
Naranjilla is used mainly for preparing refreshing drinks. The fruit contains
a fair amount of sugar, ascorbic acid and niacin (Table 12.2). The ripe fruit
is usually placed in boiling water for a short time to soften the pulp, after
which it is split into two by cutting at the equatorial line and squeezed so that
only the peel remains. The balance is blended with chilled water and sugar,
strained to eliminate the seeds and served. The peel and seeds constitute
around 50% of the total fruit weight. The juice oxidizes readily and turns
brown. Even canned juice with 0.1% ascorbic acid added and pasteurized at
92C for 75 seconds will eventually lose its characteristic color and avor.
Today, naranjilla pulp is also used in yoghurt and ice cream. In addition, it
is presented as an instant soluble powder for preparing juice. Frozen pulp is
exported, especially to the USA, as concentrate with 34% total soluble solids.
It is quickly frozen after using a rotary evaporator at 35C. Prepared juice is
also being exported in this form as it retains its appearance for a long time.
352
Chapter 12
FURTHER READING
Mamey sapote
Alia-Tejacal, I., Villanueva-Arce, R., Pelayo-Zaldvar, C., Colinas-Len, M.T., LpezMartnez, V. and Bautista-Baos, S. (2007) Postharvest physiology and technology
of sapote mamey fruit (Pouteria sapota (Jacq.) H.E. Moore & Stearn). Postharvest
Biology and Technology 45, 285297.
Crane, J.H. (2008) Pouteria sapota mamey sapote a. In: Janick, J. and Paull, R.E. (eds)
Yahia, E.M. and Gutierrez-Orozco, F. (2011) Mamey sapote (Pouteria sapota Jacq. H.E.
Moore & Stearn). In: Yahia, E.M. (ed.) Postharvest Biology and Technology of Tropical
and Subtropical Fruits, Volume 3. Cocona to Mango. Woodhead Publishing Ltd,
Cambridge, pp. 482491.
Chiku or sapodilla
Paull, R.E. (2008) Manilkara zapota, Chiku. In: Janick, J. and Paull, R.E. (eds) Encyclopedia
Yahia, E.M. and Gutierrez-Orozco, F. (2011) Sapodilla (Manilkara achras (Mill.) Fosb., syn
Achras sapota L). In: Yahia, E.M. (ed.) Postharvest Biology and Technology of Tropical
and Subtropical Fruits, Volume 4. Mangosteen to White Sapote. Woodhead Publishing
Ltd, Cambridge, pp. 351362.
Pitaya
Bauer, R. (2003) A synopsis of the tribe Hylocereeae F. Buxb. Cactaceae Systematics
Initiatives 17, 363
Le Bellec, F. and Vaillant, F. (2011) Pitahaya (pitaya) (Hylocereus spp.). In: Yahia, E.M.
Acerola
Alves, R.E., Bezerra, M.A., Rodrguez de Miranda, F. and Silva, H. (2007) In: Crisstomo,
L.A. and Naumov, A. (managing eds) and Johnston, A.E. (ed.) Fertilizing for High
Yield and Quality Tropical Fruits of Brazil. International Potash Institute, Bulletin 18,
IPI, Horgen, Switzerland, pp 1330.
Duarte, O. and Paull, R. (2008) Acerola (Malpighia emarginata DC). In: Janick, J. and
pp. 461466.
Gonzaga-Neto, L. and Soares, J.M. (1994) Acerola para Exportacao. Ministero
American Fruit
353
da Agricultura, do abastecimiento e da Reforma Agraria, Secretaria de

Desenvolvimento Rural, FRUPEX, Brasilia, Brazil.
Naranjilla
Heiser, C. (2001) Interspecic hybridization and the improvement of the naranjilla,
Solanum quitoense. In: van den Berg, R., Barendse, G., Van der Weerden, G. and
Mariani, C. (eds) Solanaceae V; Advances in Taxonomy and Utilization. Nijmegen
University Press, Nijmegen, The Netherlands, pp. 307310.
Heiser, C. (2008) Solanaceae, naranjilla. In: Janick, J. and Paull, R.E. (eds) Encyclopedia
of Fruit and Nuts. CAB International, Wallingford, UK, p. 858.
National Research Council (1989) Lost Crops of the Incas: Little-known Plants of the Andes
with Promise of Worldwide Cultivation. National Academy Press, Washington, DC.
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INDEX
Acerola
climate 335336
clonal propagation 339340
commercial use 342
cultivar development 338339
cytogenetics and breeding 338
fertilization 341
owers 336, 337
fruit 337338
genera and species 334
harvesting and postharvest handling 342
irrigation 340
Malpighiaceae 334
origin and distribution 334335
pest management 341
pollination 336337
pruning 340341
soil 335
transplanting and plant spacing 340
tree 336, 337
weed management 341
Ackee
achin 249250
climate 242
cytogenetics and breeding 245
fertilization 246
owers 243, 244
fruit 244
irrigation 246
JVS 248249
nutritional value 247248
pest management 246247
pollination 244
pruning 246
Sapindaceae 241
seed and aril toxic compound 248, 249
soil 242
tree 243
weed management 247
Ambarella
Anacardiaceae 289
climate 290291
fertilization 294
owers 291, 292
fruit 292
irrigation 293294
medicinal use 296
orchard protection 295
pruning 294
soil 290
tree 291, 292
weed management 295
Anastrepha striata 112
Anastrepha sp. 311312
Annona
A. muricata 5
A. squamosa 5, 1011
Artocarpus
A. altilis 26
A. camansi 26
A. heterophyllus see Jackfruit
A. integer see Chempedak
A. mariannensis 26
A. odoratissimus see Marang
363
364
Index
Averrhoa
A. bilimbi see Bilimbi
A. carambola see Carambola
Bephratelloides 1617
Bilimbi
climate 54
clonal propagation and nursery
management 63
eld preparation 64
owers 5759
fruit 5960
irrigation 6465
light and photoperiod 56
pollination 59
pruning and training 6566
soil 54
temperature 5556
transplanting and spacing 64
weed management 68
wind 56
Blighia sapida see Ackee
Bombacaceae 75
Botryodiplodia theobromae 15, 131
Breadfruit
climate 26
management 31
edible portion, composition 33, 34
fertilization 32
eld preparation 31
owers 27, 28
fruit 28, 29
growth of 2830
latex, use 33
methanol spray, vegetative growth 28
pest management 32
pollination 28
pruning 31
seedless cultivars 29, 30
soil 26
tree 27
triploidy 29, 31
weed management 32
Calandra linearis 231

Capnodium citri 286287
Carambola
climate 54
management 6364
disease tolerance and fruit 61, 62
fertilization 67
eld preparation 64
owers 5657
ower style length 57, 58
fruit 5960
genetics and breeding 60
irrigation 6465
pollination 59
selection and evaluation 61
soil 54
soil moisture availability 54, 55
soluble solids and titratable acidity 61, 63
sugar-to-acid ratio 71
temperature 5455
transplanting and spacing 64, 65
tree 56, 57
weed management 68
wind 56
yield vs. irrigation 54, 55
Carpophilus 6
Caryedon serratus 231
Cephaleuros virescens 150
Ceratitis capitata 17
Cerconota moth 16
Cercospora averrhoa 67
Chempedak
climate 35
management 39
fertilization 40
eld preparation 39
owers 36
fruit 37
irrigation 39
pollination 36
pruning 40
Index
soil 35
tree 3536
weed management 40
world production and nutritional value 42
Coconut
breeding program 201
dwarf and tall characteristics 199200
ecology 192193
fertilization 203204
fronds 195
fruit 197198
harvesting 206207
inorescence and owers 195196
irrigation 203
lightning 194
macapuno 199200
marketing 208209
Nam Hom 200
pollination 197
postharvest handling 207208
pruning 203
radiation 193
rainfall 193, 194
soil 193
stem and roots 194195
temperature 193
water volume and total soluble solid 198,
199
weed management 206
wind 193
Cocos nucifera see Coconut
Colletotrichum gloeosporioides 15, 110111
Colletotrichum spp. 67
Conotrachelus psidii 112
Corticium salmonicolor 40, 268
Dacus dorsalis 17, 68
Dolabra nepheliae 150
Durian
breeding 8081
climate 75
management 82
edible portion, composition 87
fertilization 8384
eld preparation 82
365
owering cycle 76, 78

owers 76, 77
fruit 77, 79
irrigation 8283
leaf ushing 80
marketing, fruit 86
pest management 84
pollination 78
pruning 83
sigmoid growth pattern 7980
soil 75
tree 76
utilization 86
weed management 84
world production 86
Elsino fawcettii 286287
Eriophyes sandorici 268
Erwinia carotovora 46
Eugenia
E. brasiliensis 92
E. uniora 92
Feijoa sellowiana 93
Fusarium 181
F. decemcellulare 67
F. oxysporum 110111
Garcinia mangostana see Mangosteen
Guava
asexual seed germination 103104
Beaumont and Kahua Kula 101
breeding 100
carne 116117
conventional processing 117
cultivars development 100101
cytogenetics and genetics 99100
Eugenia 92
Feijoa 93
eld preparation 104
owers 9596
fruit 99
fruit-detachment force 114, 115
fruit harvesting 113114
fruit thinning and bagging 109110
gibberellic acid-lanolin mixture 9798
366
Index
Guava continued
insect and nematode pest 111112
irrigation 105
light 95
Myrciaria 92
Myrtaceae 91
natural pollination 9697
open-center at-pruning method 110
pruning
continuous light pruning 107
GA9-EX39 106
gibberellic acid, ethephon 106107
leaf production pattern 105106
orchard management 105
Psidium 9192
rainfall 9394
Roja Enana Cubana 101
sexual seed germination 103
soil 93
Syzygium 92
Tai-kuo-bai 101
temperature 9495
tree 95
weed management 113
wind 95
world production 115116
Hylocereus
H. costaricensis 329330
H. undatus 329330
see also Pitaya
Jackfruit
climate 35
management 3839
fertilization 40
eld preparation 39
owers 36
fruit 37
pest management 40
pollination 36
pruning 40
soil 35

tree 3536
watering and grass mulching effect 39
weed management 40
world production and nutritional value 42,
43
Jamaican vomiting sickness syndrome (JVS)
248249
Langsat and duku
climate 256
management 259260
dysentery and malaria treatment 263
eld-preparation 260
owers 256, 258
fruit 257258
genetics and cytogenetics 258259
irrigation 260
longkong 259
Meliaceae 255
pollination 256257
pruning and thinning 260
selection and evaluation, cultivar 259
soil 255
tree 256, 258
weed management 262
Lansium domesticum Jack see Langsat and
duku
Malpighia emarginata see Acerola
Mamey sapote
asexual propagation 308, 310
climate 304305
cross-pollination 306307
cultivars evaluation and adaptability 308,
309
fertilization 311
eld preparation 310
owers 305306
fruit 306, 307
harvesting and postharvest handling
312
Index
irrigation 310
medicinal use 313
nutritional value 312314
pruning 310311
Sapotaceae 303
selection and evaluation, cultivars 308
sexual propagation 308
soil 304
tree 305, 306
weed management 312
Mangosteen
aril acidity and total soluble solids 133
climate 124
management 129
Clusiaceae 123
fertilization 131
eld preparation 129
owers 125126
fruit 127128
harvest index stage 132133
irrigation 130
marketing 134
paclobutrazol 126
pest management 131
phenology steps, owering induction 126,
127
pollination 126
pruning 130
shoot/root ratio 125, 126
soil 124
tree 124125
weed management 131
world production and use 134135
Marang
management 45
ecology 44
fertilization 46
leaf and fruit 44, 45
pest management 46
pruning 46
weed management 46
Marula 232233
asexual seed propagation 238
climate 235236
ecology 235
owers 236, 237
fruit 237238
239
irrigation 239
medicinal use 241
Mukumbi 240
pest 239
pollination 236
sexual seed propagation 238
soil 235
tree 236
Meloidogyne sp. 182
Membracidae spp. 68
Naranjilla
asexual propagation 348
breeding, selection and evaluation
346
climate 344
cultivars 346347
fertilization 349
owers 345
fruit 345, 346
350351
herbaceous shrub 344345
irrigation 348
pollination 345
pruning 349
refreshing drink preparation 351
soil 343344
Solanaceae 343
weed management 350
367
368
Nephelium
N. intermediam 147
N. lappaceum see Rambutan
N. ramboutan-ake see Pulasan
Palms
coconut see Coconut
Palmae 191
salak see Salak
Panonychus citri 68
Passion fruit and giant passion fruit
breeding and selection 172173
commercial orchards, yield 182
diseases of 180182
ecology 164
owers 167169
fruit 170171
fruit processing 186
hand-harvesting 183184
hand pollination 176177
irrigation 178
light 165166
nutritional value and vitamins 186
Passioraceae 161
passiorine 186
P. edulis Sims vs. P. edulis f. avicarpa 161
pest management 182
photoperiod 166
pollination 169170
rainfall 164165
soil 164
temperature 165
vine 166167
Pellicularia koleroga 131
Phomopsis annonacearum 15
Phyllophaga bruneri 311
Phytophthora 181
P. palmivora 84
P. phaseoli 268
Pitaya
Cactaceae 324325
climate 327
Index

fertilization 332
eld preparation 331
owers 328329
fruit 325, 329
333
irrigation 331
medicinal use 334
pest management 332
plant 327328
pollination 329
pruning 332
soil 326
weed management 332
Psidium
P. friedrichsthalianum 9192
P. guajava see Guava
P. guineense 92
P. littorale 92
Pulasan
ecology 152153
owers 154
fruit 155
genera and species 139, 152
nutrition and fertilization 156
pest and weed management 156
pollen germination test 154155
Sapindaceae 139
tree 153154
Pummelo
breeding 284
climate 282
management 285
fertilization 286
eld preparation 285
owers 282
fruit 283284
Index

287288
irrigation 285286
nutritional value 288, 289
pharmaceutical use 288
pollination 282283
pruning and fruit thinning 286
Rutaceae 280
soil 281
tree 282, 283
weed management 287
Rambutan
breeding and crop improvement 145146
erratic bearing 150
eld preparation 147
owers 141, 143
fruit 144145
fruiting cycle 140142
irrigation 148
light intensity and photoperiod 141
pollination 143144
pruning 149
rainfall and moisture 140141
Sapindaceae 139
soil 140
temperature 141
tree 141, 143
weed management 151
wind 141
Rhizopus artocarpi 40
Rollinia
Annonaceae 1
fertilization 13
eld preparation 11
ower 5, 6
growth regulator 7
369
hand pollination 7
irrigation 1112
peak harvesting season 19
protogynous ower, pollination 7
pruning 1213
pulp 20
rainfall 3
soil 2
syncarp fruit 6, 8
temperature 3
tree 5
weed management 18
wind 4
Salacca zalacca see Salak
Salak
fertilization and irrigation 214215
eld preparation and spacing 214
fruit 212
inorescence and owers 211212
pest management 215
pollination 212
pruning and fruit thinning 215
radiation 210
rainfall 210
soil 210
stem and leaves 210211
temperature 210
vegetative propagation 214
weed management 216
Sandoricum koetjape see Santol
Santol
Bangkok 267
climate 264265
management 267
fertilization 268
eld preparation 267
owers 265, 266
fruit 265, 266
irrigation 268
medicinal use 269270
Meliaceae 263
Orchard protection 269
370
Santol continued
pest management 268, 269
pollination 266
pruning 268
soil 264
tree 265
weed management 268
Sapodilla
climate 315316
cultivars development 319, 320
cultivars selection and evaluation criteria
318319
eld preparation 321
owers 316, 317
fruit 316318
323324
irrigation 321
medicinal and ornamental use 324
origin and distribution 313, 315
pest management 322
pollination 316
pruning 321
soil 315
tree 316, 317
vegetative propagation 320321
weed management 322
Sclerocarya birrea see Marula
Sclerotium rolfsii 84
Selenothrips rubrocinctus 182
Solanum quitoense see Naranjilla
Soursop
Annonaceae 1
anthracnose 15
black canker and diplodia rot 15
breeding 8, 10
diseases of 16
fertilization 13
eld preparation 11
ower 4, 5
fruit growth 8, 9
Index
fruit ripening, compositional change 20

genetics 8
growth regulator 7
insect pests 1618
irrigation 1112
leaf nutrient concentration 14, 15
NPK ratio 1314
peak harvesting season 1819
pollination, phases 56
pruning 1213
pulp 20, 21
rainfall 3
soil 2
syncarp fruit 8, 9
temperature 3
tree 4
weed management 18
wind 4
Syzygium
S. pycnanthum 275
S. samarangense see Wax apple
Tamarind
breeding 227
climate 224225
commercial orchards, cultivar development
228
cultivar selection and evaluation 228
ecology 224
Fabaceae 223
fertilization 230
eld preparation 229
owers 225, 226
fruit 226, 227
231232
irrigation 229
pollination 226227
pruning 230
soil 224
tree 225
Index
weed management 231

Tamarindus indica see Tamarind
Timocratica albella 112
Uroporus spp. 6
Wax apple
bald cut 274
climate 271272
management 275
cultivars, fruit characteristics 275, 276
fertilization 277
eld preparation 277
ower forcing
improved 274
simple 274
owers 272, 273
fruit 275
371

278279
irrigation and pruning 277
Myrtaceae 270271
nutritional value 279, 280
pollination 273274
shading and bald cut 275
soil 271
tree 272
weed management 278
Xanthomonas citri 286
Zignoella garcineae 131

Tropical Fruit

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CROP PRODUCTION SCIENCE IN HORTICULTURE SERIES

Series Editor: Jeff Atherton, Professor of Tropical Horticulture,

Ornamental Bulbs, Corms and Tubers A.R. Rees

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CABI is a trading name of CAB International

Tel: +44 (0)1491 832111

Commissioning editor: Sarah Hulbert

Volume I presented the general aspects of tropical fruit production and

The authors gratefully acknowledge the support, information, and ideas

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Important genera and species

Paull and Duarte 2012. Tropical Fruits, 2nd Edition, Volume II

mulato (Colombia), cachiman cochon or cachiman montagne (Guadalupe),

Area of origin and distribution

Annonas: Soursop and Rollinia

Fig. 1.1. The leaves and ower of soursop.

Annonas: Soursop and Rollinia

Rollinia generally reaches 610 m in height, but can grow as high as 20

Pollination and fruit set

Fig. 1.2. Rollinia owers (A) and fruit (B).

Annonas: Soursop and Rollinia

Problems with breeding

Annonas: Soursop and Rollinia

Fig. 1.3. Fruit of soursop, sweetsop and atemoya.

Time from Anthesis (weeks)

in 2 years and full production in 56 years. This juvenile period is extremely

Annonas: Soursop and Rollinia

Transplanting and spacing

moisture is required to encourage vegetative growth, however, since owering

Annonas: Soursop and Rollinia

Another recommendation suggests that non-irrigated bearing plants should be

Annonas: Soursop and Rollinia

Table 1.3. Major diseases of soursop.

Flowers, fruit, leaves,

Armillaria root rot Armillaria luteobubalina

Roots, base of trees,

Leaf scorch, twig

Annonas: Soursop and Rollinia

Table 1.4. Major insect pests of soursop.

America. This wasp is considered to be the most important pest in Florida.

HARVESTING AND POSTHARVEST HANDLING

Table 1.5. Peak harvesting seasons for soursop.

Annonas: Soursop and Rollinia

Differences in recovery percentages are caused by differences in equipment,

Compositional changes during fruit ripening

Annonas: Soursop and Rollinia

Table 1.6. Composition of 100 g edible portion of soursop

Annonas: Soursop and Rollinia

seedless cultivars have different common names. Seeded breadfruits are

Breadfruit, Jackfruit, Chempedak and Marang

Breadfruit, Jackfruit, Chempedak and Marang

Total Sugars (g)

White esh, seedless,

Meinpadahk Micronesia Oval to

Breadfruit, Jackfruit, Chempedak and Marang

Harvesting and postharvest handling

Breadfruit, Jackfruit, Chempedak and Marang

JACKFRUIT AND CHEMPEDAK

Table 2.2. Composition of 100 g edible portion of breadfruit, jackfruit, chempedak

Jackfruit is also known as jacquier (French), nangka (Javanese and Malay),

Breadfruit, Jackfruit, Chempedak and Marang

when ripe. In addition, the embryo radicle is immersed while in jackfruit it is

Breadfruit, Jackfruit, Chempedak and Marang

Length and Girth (cm)