The Mango, 2nd Edition

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The Mango, 2nd Edition

Botany, Production and Uses

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The Mango, 2nd Edition

Botany, Production and Uses

Edited by

Richard E. Litz

Tropical Research and Education Center and

Center for Tropical Agriculture University of Florida 18905 SW 280 Street

Homestead, FL 33031-3314

USA

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Library of Congress Cataloging-in-Publication Data The mango : botany, production and uses/edited by Richard E. Litz. -- 2nd ed.

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Includees bibliographical references and index.

ISBN 978-1-84593-489-7 (alk. paper) 1. Mango. I. Litz, Richard E.

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Contributors

N. Alkan, Department of Technology and Storage of Agricultural Products, Agricultural Research Organization (ARO), The Volcani Center, PO Box 6, Bet Dagan 50250, Israel.

M. Aluja, Instituto de Ecología AC, Km 2.5 Antigua Carretera a Coatepec No. 357, Congregación El Haya, Apartado Postal 63, CP 9100 Xalapa, Veracruz, Mexico. E-mail: [email protected] I.S.E. Bally, Horticulture and Forestry Sciences, Department of Primary Industries and Fisheries, 28

Peters Street (PO Box 1054), Mareeba, QLD 48890, Australia. E-mail: [email protected] J.M. Bompard, Les Mazes, 34160 Montaud, France. E-mail: [email protected]

J.K. Brecht, 1143 Fiﬁ eld Hall, Department of Horticultural Sciences, University of Florida, Gaines- ville, FL 32611-0690, USA. E-mail: [email protected]ﬂ .edu

R.J. Campbell, Fairchild Tropical Botanic Garden, 10901 Old Cutler Rd, Miami, FL 33156-4296, USA. E-mail: [email protected]

J.H. Crane, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA. E-mail: [email protected]ﬂ .edu

T.L. Davenport, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA. E-mail: [email protected]ﬂ .edu

E.A. Evans, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA. E-mail: [email protected]ﬂ .edu

S. Freeman, Department of Plant Pathology and Weed Research, Agricultural Research Organization (ARO), The Volcani Center, Bet Dagan 50250, Israel. E-mail: [email protected]

V. Galán Saúco, Departamento de Fruticultura Tropical, Instituto Canario de Investigaciones Agrarias, Apartado Correos 60, 38200 La Laguna, Tenerife, Canary Islands, Spain. E-mail: [email protected]

M.A. Gómez-Lim, Centro de Investigacion y de Estudios Avanzados del IPN (CINVESTAV), Apar- tado Postal 629, Irapuato GTO, Mexico 36500. E-mail: [email protected]

P.J. Hofman, Department of Primary Industries and Fisheries, PO Box 5083, SCMS Nambour, QLD 4560, Australia. E-mail: [email protected]

C.P.A. Iyer, Indian Institute of Horticultural Research, Hessaraghatta Lake Post, Bangalore 560089, India. E-mail: [email protected]

G.I. Johnson, Horticulture 4 Development, PO Box 412, Jamison, ACT 2614, Australia. E-mail: greg.

[email protected]

R.J. Knight, Jr, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA. E-mail: [email protected]ﬂ .edu

vii

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viii Contributors

I. Kobiler, Department of Technology and Storage of Agricultural Products, Agricultural Research Organiza- tion (ARO), The Volcani Center, PO Box 6, Bet Dagan 50250, Israel. E-mail: [email protected] U. Lavi, Department of Horticulture, Agricultural Research Organization (ARO), The Volcani Center,

PO Box 6, Bet Dagan 50250, Israel. E-mail: [email protected]

T.-S. Lin, 111 Room, No. 4 Hall, Department of Horticulture, National Taiwan University, No. 1 Sec.

4 Roosevelt Road, 106 Taipei, Taiwan. E-mail: [email protected]

R.E. Litz, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Home- stead, FL 33031-3314, USA. E-mail: relitz@uﬂ .edu

P. Lu, EWL Sciences, PO Box 39443, Winnellie, NT 0821, Australia. E-mail: ping.lu@ewlsciences.

com.au

I. Maguire, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA. E-mail: imaguire@uﬂ .edu

O.J. Mendoza, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA.

I. Miyara, Department of Technology and Storage of Agricultural Products, Agricultural Research Organization (ARO), The Volcani Center, PO Box 6, Bet Dagan 50250, Israel.

S.K. Mukherjee (deceased), Department of Agriculture, Calcutta University, 35 Ballygunge Circu- lar Road, Calcutta 700 019, India.

M.T. Ombico, Fruit and Vegetable Laboratory, Food Science Cluster, College of Agriculture, Univer- sity of the Philippines Los Baños, Laguna, 4031, Philippines.

J.E. Peña, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Home- stead, FL 33031-3314, USA. E-mail: [email protected]ﬂ .edu

R.C. Ploetz, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA. E-mail: [email protected]ﬂ .edu

D. Prusky, Department of Technology and Storage of Agricultural Products, Agricultural Research Organiza- tion (ARO), The Volcani Center, PO Box 6, Bet Dagan 50250, Israel. E-mail: [email protected] A.C. de Queiroz Pinto, Private Consultant Tropical Fruits, SHCGN 706 Bloco P Casa 13, 70740-

716, Brasilia-DF, Brazil. E-mail: [email protected]

S. Ram (deceased), Department of Horticulture, GB Pant University of Agriculture and Technology, Pantnagar 263 145, India.

L.C. Raymundo, Fruit and Vegetable Laboratory, Food Science Cluster, College of Agriculture, Uni- versity of the Philippines Los Baños, Laguna, 4031, Philippines. E-mail: [email protected] S. Salazar-García, Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Campo

Experimental Santiago Ixcuintla, Km. 6 Carret. Intnal. Tepic-Mazatlán, Apartado Postal 100, San- tiago Ixcuintla, Nayarit 63300, Mexico. E-mail: [email protected]

B. Schaffer, Tropical Research and Education Center, University of Florida, 18905 SW 280 Street, Homestead, FL 33031-3314, USA. E-mail: [email protected]ﬂ .edu

R.J. Schnell, United States Department of Agriculture (USDA) Agriculture Research Service (ARS), Subtropical Horticultural Research Unit/National Germplasm Repository, 13601 Old Cutler Road, Miami, FL 33158, USA. E-mail: [email protected]

Z.-H. Shü, Department of Biological Science and Technology, Meiho Institute of Technology, 23 Ping Kuang Road, Neipu, Pingtung, 91202, Taiwan. E-mail: [email protected]

L. Urban, Directeur de l’UR Génétique et écophysiologie de la qualité des agrumes (GEQA), Centre INRA de Corse, 20230 San Giuliano, France. E-mail: [email protected]

T.M. de Villa, Fruit and Vegetable Laboratory, Food Science Cluster, College of Agriculture, Univer- sity of the Philippines Los Baños, Laguna, 4031, Philippines.

A.W. Whiley, Sunshine Horticultural Services Pty Ltd, 287 Dulong Road, Nambour, QLD 4560, Australia. E-mail: [email protected]

M. Wysoki, Department of Entomology, Institute of Plant Protection, The Volcani Center, Bet Dagan 50250, Israel. E-mail: [email protected]

E.M. Yahia, Facultad de Química, Universidad Autónoma de Querétaro, Querétaro, 76190 Qro, Mexico.

E-mail: [email protected]

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ix

Preface

The ﬁ rst edition of The Mango: Botany, Production and Uses appeared in 1997, and went into an unprecedented second printing in the following year. Despite the worldwide importance of the mango, this was the ﬁ rst book that was devoted solely to this fruit crop species since the publication of The Mango by Gangolly et al. in 1957 and The Mango: Botany, Cultivation and Utilization by L.B.

Singh in 1960. The appearance of The Mangoes: their Botany, Nomenclature, Hor- ticulture and Utilization by Kostermans and Bompard in 1993 had provided a much-needed taxonomic and systematic revision of mango and the related Mangifera species; the Kostermans and Bompard book also stimulated interest in the Mangifera spp. germplasm for breeding and rootstock development.

The Mango: Botany, Production and Uses (Litz, 1997) provided a fresh per- spective of the mango. The authors represented several countries, including India, Australia, Israel, the UK, France, USA, Mexico, Pakistan and South Africa, and reﬂ ected the expansion of mango production outside its traditional areas of cultivation during the mid-20th century and the development of new technologies in these new lands. The worldview of the ﬁ rst edition was unique, and the authors were at the forefront of the advance of science in support of mango production. I wish to particularly acknowledge L.A.

Milne (South Africa), R.V. Mosqueda-Vazquez (Mexico), S.K. Mukherjee (India) and S. Ram (India), who contributed to the ﬁ rst edition, and who have passed away since then.

Since 1997, other mango books have appeared: El Cultivo del Mango by V.

Galán Saúco in 1999 (Spain), Mango Cultivation edited by R.P. Srivastava in 1998 (India), A Cultura da Mangueira edited by P.J. de Carvalho Genu and A.C. de Queiroz Pinto in 2002 (Brazil) and El Mango by E. Yahia Kazuz, J. de J. Ornelas Paz and R. Ariza Flores in 2006 (Mexico). These books have generally targeted audiences in speciﬁ c mango-producing countries. Drs Galán Saúco, Pinto and Yahia are also contributors to the second edition of The Mango: Botany, Production and Uses.

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x Preface

Much has happened in the decade following the appearance of the ﬁ rst edition of The Mango: Botany, Production and Uses. China has emerged as the second largest producer of mango fruit; India’s production is now less than half of the world total. Fresh mangoes are now consumed worldwide and are available year-round in the European Union (EU), North America and Japan.

The availability of fruit of a range of mango cultivars is increasing. Mango products, including fruit nectars, leather, dried fruit slices, preserves, yogurt, etc. have become widely popular outside the tropics.

The authorship of the second edition of The Mango: Botany, Production and Uses represents the USA, Mexico, Brazil, Australia, the Philippines, Tai- wan, India, Israel, France and Spain, and includes leading authorities in each ﬁ eld. The subject matter of this book ranges from the most basic to the applied, and is designed to be a compendium that will remain highly relevant for researchers and growers for many years.

I would like to express my appreciation and thanks to all of the authors for their persistence during the 3-year gestation period. I would like to express my gratitude to Ian Maguire of the Tropical Research and Education Center of the University of Florida for his photographic assistance. Financial assistance provided by Dr Yungcong Li, also of the Tropical Research and Education Center, for reproduction of colour plates is gratefully acknowledged. Special thanks to Pamela A. Moon, Guillermo Padilla and Irene Perea who tolerated me while I worked on this project.

Richard E. Litz

References

de Carvalho Genu, P.J. and de Queiroz Pinto, A.C. (eds) (2002) A Cultura da Mangueira.

EMBRAPA Informacao Tecnologica, Brasilia DF.

Galán Saúco, V. (1999) El Cultivo del Mango. Mundi-Prensa, Madrid.

Gangolly, S.R., Singh, R., Katyal, S.L. and Singh, D. (1957) The Mango. Indian Council for Agricultural Research, New Delhi, India.

Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mangoes: their Botany, Nomencla- ture, Horticulture and Utilization. Academic Press, London.

Litz, R.E. (ed.) (1997) The Mango: Botany, Production and Uses. CAB International, Wallingford, UK.

Singh, L.B. (1960) The Mango: Botany, Cultivation and Utilization. Leonard Hill, London.

Srivastava, R.P. (1998) Mango Cultivation. International Book Distributing Co, Lucknow, India.

Yahia Kazuz, E., de J. Ornelas Paz, J. and Ariza Flores, R. (2006) El Mango. Editorial Trillas, S.A. de C.V., Mexico.

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xi

Acknowledgements

The assistance provided by Ian Maguire is gratefully acknowledged. Artwork for the covers was provided by Ian. Special thanks to Campbell, Penelope and Anna.

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(ed. R.E. Litz) 1

1 Introduction: Botany and Importance

S.K. Mukherjee

¹

and R.E. Litz

²

1Calcutta University, Calcutta, India

2University of Florida, Florida, USA

1.1 Introduction 1

1.2 Description of Mango 2

The tree 2

Flowers 2

The fruit 3

The seeds and polyembryony 4

1.3 History of Cultivation 5

Origin of Mangifera indica 5

Domestication of mango 9

Distribution 10

1.4 Germplasm Conservation 11

Genetic erosion 11

Collection and documentation of Mangifera germplasm 12

Relevance of germplasm resources to mango improvement 12

1.5 Importance of Mango 12

Cultivars 12

1.6 Production and Uses 14

1.1 Introduction

Mango has become a major fruit crop of the tropics and subtropics, particularly in Asia, where the mango has always been the most important fruit crop and where it has been considered the ‘king of fruits’ (Purseglove, 1972). A generation ago, the Green Revolution culminated, creating surpluses of sta- ple and horticultural crops in many developing countries. The Green Revo- lution was the result of nearly a century of effort of applying Mendelian genetics to crop improvement (i.e. conventional breeding) together with the optimization of agronomic and horticultural practices and the successful management of insect pests and diseases. However, improvement of tree

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S.K. Mukherjee and R.E. Litz 2

crops has lagged far behind fi eld crops for several reasons: their heterogene- ity, polyploidy, lengthy juvenile period, time required for evaluation of trees in the fi eld, and the relatively high cost of maintaining tree plantings. For the most part, fruit cultivars continue to be ancient selections, many of which have serious problems, including alternate bearing, lack of disease resistance, low yields, etc. The rapid growth of mango production in recent years has been due to its expansion into new growing regions of the New World, China and parts of Africa; the planting of regular bearing selections; and the adop- tion of modern fi eld practices, which include irrigation management, control of fl owering, etc. Agricultural practices are currently undergoing another revolution, as integrated pest and disease management replaces the earlier reliance on agrichemicals, and emerging fi elds within biotechnology begin to impact cultivar development.

1.2 Description of Mango

The tree

The mango tree is believed to have evolved as a canopy layer or emergent species of the tropical rainforest of South and South-east Asia (Kaur et al., 1980; Bompard, Chapter 2, this volume). Mature trees can attain a height of 40 m or more, and can survive for several hundred years. Mango trees that have been domesticated by selection from openly pollinated seedling popu- lations show variation in tree architecture (i.e. shape and size). The tree is an arborescent evergreen. Leaves are simple and alternate, with petioles that range in length from 1 to 12.5 cm. Leaf morphology is highly variable, depending on the cultivar: leaves can be lanceolate, oblong, ovate and intermediate types involving these forms. Leaf length ranges from 12 to 38 cm and width can be between 2–13 cm. Young leaves are copper-coloured, changing gradually to light and then dark green with age. The leaves are spirally arranged in whorls and are produced in ﬂ ushes. The canopy is normally oval, elongated or dome shaped. The juvenile period of seedling trees can range from 3 to 7 years. The root system consists of a long, vigorous taproot and abundant surface feeder roots.

Flowers

Mango fl owers are borne on terminal pyramidal panicles, and are glabrous or pubescent; the infl orescence is rigid and erect, up to 30 cm long, and is widely branched, usually tertiary, although the fi nal branch is always cymose.

The infl orescence is usually densely fl owered with hundreds of small fl owers, which are 5–10 mm in diameter. The fl owers are either monoecious or polygamous, and both monoecious and polygamous fl owers are borne within a single infl orescence (Plate 1). The pistil aborts in male fl owers. The ratio of monoecious to polygamous fl owers is strongly infl uenced by

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Introduction: Botany and Importance 3

environmental and cultural factors. The fl owers have four or fi ve sepals and petals that are ovate to ovoid to lanceolate and also thinly pubescent. The fl oral disc also is four- or fi ve-lobed, fl eshy and large and located above the base of the petals. There are fi ve large, fl eshy stamens, only one or two of them being fertile; the remaining stamens are sterile staminodes that are sur- mounted by a small gland. In addition, two or three smaller fi laments arise from the lobes of the nectaries. The stamens are central. The ovule is anatro- pous and pendulous. It is believed that the fl owers are cross-pollinated by fl ies (see Davenport, Chapter 5, this volume).

Mukherjee (1951a, 1953) investigated the pollen morphology of mango and 12 other Mangifera species. Their pollen grains were tricolpate of almost the same size. Mondal et al. (1982, cited in Kostermans and Bom- pard, 1993) attempted to correlate pollen morphology with taxonomic relationships of 17 Mangifera species based upon different characteristics of the exine and sporoderm. They demonstrated that all of the species of section II (subgenus Limus) possess coarse exine; whereas there was no clear correlation with pollen type in species within section I (subgenus Mangifera).

The fruit

Description

The mango fruit is a large, fl eshy drupe, containing an edible mesocarp of varying thickness. The mesocarp is resinous and highly variable with respect to shape, size, colour, presence of fi bre and fl avour. The fl avour ranges from turpentine to sweet. The exocarp is thick and glandular. There is a characteristic beak that develops laterally on the proximal end of the fruit. A sinus is always present above the beak. Fruit shape varies, including elongate, oblong and ovate or intermediate forms involving two of these shapes. Fruit length can range from 2.5 to > 30 cm, depending on the cultivar. The endocarp is woody, thick and fi brous; the fi bres in the mesocarp arise from the endocarp.

The mango fruit is climacteric (see Brecht and Yahia, Chapter 14, this volume), and increased ethylene production occurs during ripening. Chloro- phyll, carotenes, anthocyanins and xanthophylls are all present in the fruit.

The skin is generally a mixture of green, red and yellow pigments, although fruit colour at maturity is genotype dependent. During ripening the chloroplasts in the peel become chromoplasts, which contain yellow and red pigments (Krishnamurthy and Subramanyam, 1970; Akamine and Goo, 1973;

Salunkhe and Desai, 1984; Mitra and Baldwin, 1997). Peel colour obviously is cultivar dependent (see Knight et al., Chapter 3, this volume). Fruit of ‘Bom- bay Green’ is green; ‘Carabao’, ‘Manila’, ‘Mulgoa’ and ‘Arumanis’ are greenish- yellow; ‘Dashehari’ and ‘Alphonso’ are yellow; and ‘Haden’, ‘Keitt’ and

‘Tommy Atkins’ have a red blush. The red blush is due to the presence of anthocyanins (Lizada, 1991). The pulp carotenoids in ripe fruit also vary with respect to cultivar (Mitra and Baldwin, 1997).

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Flavour

Flavour of the mango mesocarp is a function of carbohydrates, organic acids, lactones, monoterpene hydrocarbons and fatty acids (Mitra and Baldwin, 1997). During fruit maturation, starch that accumulates in the chloroplasts is hydrolysed to sucrose, glucose and fructose (Medlicott et al., 1986; Selvaraj et al., 1989; S. Kumar et al., 1994); sucrose is present in slightly higher concen- trations than either fructose or glucose. Organic acid content decreases during ripening (Krishnamurthy and Subramanyam, 1970). The dominant organic acid is citric acid, but glycolic acid, malic acid, tartaric acid and oxalic acids are also present (Sarker and Muhsi, 1981; Medlicott and Thompson, 1985). The peach-like ﬂ avour of mangoes is attributed to the presence of lactones (Lakshminarayana, 1980; Wilson et al., 1990).

Nutrition

Mango fruit contain amino acids, carbohydrates, fatty acids, minerals, organic acids, proteins and vitamins. During the ripening process, the fruit are initially acidic, astringent and rich in ascorbic acid (vitamin C). Ripe mangoes contain moderate levels of vitamin C, but are fairly rich in provitamin A and vitamins B₁ and B₂. Perry and Zilva (1932) determined the vitamin A, C and D content of the fruit of three Indian mango cultivars, and found that the pulp of mangoes is a concentrated source of vitamin C. The pulp of mango fruit contains as much vitamin A as butter, although vitamin D is not present in a signiﬁ cant quantity. Fruit acidity is primarily due to the presence of malic and citric acids. In addition, oxalic, malonic, succinic, pyruvic, adipic, galac- turonic, glucuronic, tartaric, glycolic and mucic acids are also present (Jain et al., 1959; Fang, 1965). Acidity is cultivar related; for example, immature Florida cultivars have low acidity (0.5–1.0%) in comparison with ‘Alphonso’ (3%).

During ripening, acidity decreases to 0.1–0.2%. Following fruit set, starch accumulates in the mesocarp. Free sugars, including glucose, fructose and sucrose, generally increase during ripening; however, the sucrose content increases three- to fourfold due to the hydrolysis of starch. Sucrose is the principal sugar of ripe mangoes. The sucrose content of ripe fruit of three Indian cultivars, ‘Alphonso’, ‘Pairie’ and ‘Totapuri’, ranges from 11 to 20%

representing 15 to 20% of the total soluble solids (Popenoe, 1932).

The seeds and polyembryony

Mango seeds are solitary, large and ﬂ at, ovoid oblong and surrounded by the ﬁ brous endocarp at maturity. The testa and tegumen are thin and papery.

Embryos are dicotyledonous. Seeds of monoembryonic mango types contain a single zygotic embryo, whose cotyledons can be unequal in size or lobed in shape. The seeds of polyembryonic mango types contain one or more embryos (Plate 2); usually one embryo is zygotic, whereas the remaining embryos are derived directly from the nucellus, a maternal tissue. Nucellar embryos apparently lack a suspensor. Polyembryony has also been reported in Mangifera casturi, M. laurina and M. odorata (Bompard, 1993). Certain

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polyembryonic cultivars reportedly can produce seeds with adventitious nucellar embryos only, for example ‘Strawberry’ (Juliano, 1934), ‘Carabao’

and ‘Pico’ (Juliano and Cuevas, 1932) and ‘Olour’ and ‘Cambodiana’

(Maheshwari et al., 1955). Early studies suggested that polyembryony appeared to be a polygenic trait (Juliano, 1934; Sturrock, 1968), segregating as a recessive character in the progeny of controlled crosses. Recent studies, however, have demonstrated that the polyembryony trait is inherited as a dominant character (Aron et al., 1998). Several studies have shown that nucellar seedlings can be distinguished from the single zygotic seedling of polyembryonic seeds by isozymes (Schnell and Knight, 1992; Degani et al., 1993) and DNA markers, for example single sequence repeats (SSRs) (Eiadthong et al., 1999a), ampliﬁ ed fragment length polymorphisms (AFLPs) (Kashkush et al., 2001) and inter-simple-sequence-repeats (ISSRs) (Gonzalez et al., 2002). Mango seeds are considered to be recalcitrant, and cannot survive for more than a few days or weeks at ambient temperatures (Parisot, 1988). This important characteristic of mango seeds would have inhibited the long distance dis- persal of mango by seed until recent times.

1.3 History of Cultivation

Origin of Mangifera indica

The largest number of Mangiferaspecies occurs in the Malay Peninsula, the Indonesian archipelago, Thailand, Indochina and the Philippines (Mukher- jee, 1985; Bompard, 1989; see Bompard, Chapter 2, this volume). The most recent classifi cation of Mangifera species was based upon fl oral morphology (Kostermans and Bompard, 1993) and included 69 species, most of which are included in two subgenera Mangifera and Limus with another 11 species occupying an uncertain position (Table 1.1). Eiadthong et al. (1999b) described the phylogenetic relationships among Mangifera species using genomic restriction fragment length polymorphisms (RFLPs) and amplifi cation of chloroplast DNA (cpDNA), and suggested that the Mangifera species should be classifi ed using molecular data. In the next few years, it is likely that molecular biology will have a major impact on phylogenetic studies involving mango and its relatives.

Mangiferaspecies with a single fertile stamen are distributed in north- eastern India, Myanmar, Thailand and the Malay Peninsula. Many of the mango relatives have small fruits with thin, acidic ﬂ esh, large seeds, abundant ﬁ bre and astringent resinous substances that are localized near the skin.

In addition to M. indica, edible fruit is produced by at least 26 other species in the genus, primarily species found in South-east Asia (Gruezo, 1992).

Mangifera caesia, known as ‘binjai’ or ‘kemang’ in South-east Asia, is cultivated in Java, where it bears fruit in the mango off-season (Bompard, 1992a).

Mangifera foetida is less commonly cultivated due to its highly astringent fruit; however, the fruit is widely used for pickling and as a substitute for tamarind (Bompard, 1992b). Mangifera kemang and M. altissima are consumed

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S.K. Mukherjee and R.E. Litz6 Table 1.1. Classiﬁ cation of Mangifera species according to Kostermans and Bompard (1993).

Genus Subgenus Section Species

Mangifera Mangifera Marchandora Pierre M. gebede Miq Euantherae Pierre M. caloneura Kurz

M. cochinchinensis Engler M. pentandra Hooker f.

Rawa Kosterm. M. andamanica King M. minutifolia Evard.

M. gracilepes M. nicobarica Kosterm.

M. grifﬁ thii Hooker f. M. paludosa Kosterm.

M. merrillii Mukherji M. parvifolia Boerl. & Koorders M. microphylla Griff. ex Hooker f.

Mangifera Ding Hou M. altissima Blanco. M. mucronulata Bl.

M. applanata Kosterm. M. oblongifolia Hooker f.

M. austro-indica Kosterm. M. orophila Kosterm.

M. austro-yunnanensis Hu M. pedicellata Kosterm.

M. casturi Kosterm. M. pseudo-indica Kosterm.

M. collina Kosterm. M. quadriﬁ da Jack M. dewildei Kosterm. M. rigida Bl.

M. dongnaiensis Pierre M. rubropetala Kosterm.

M. ﬂ ava Evard. M. rufocostata Kosterm.

M. indica L. M. similis Bl.

M. lalijiwa Kosterm. M. sulauesiana Kosterm.

M. laurina Bl. M. sumbawaensis Kosterm.

M. linearifolia (Mukherji) Kosterm. M. sylvatica Roxb.

M. longipetiolata King M. swintonioides Kosterm.

M. magniﬁ ca Kochummen M. timorensis Bl.

M. minor Bl. M. torquenda Kosterm.

M. monandra Merr. M. zeylanica (Bl.) Hooker f.

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Introduction: Botany and Importance7 Limus (Marchand)

Kosterm.

M. blommesteinii Kosterm. M. leschenaultii Marchand

M. caesia Jack M. macrocarpa Bl.

M. decandra Ding Hou M. odorata Griff.

M. foetida Lour. M. pajang Kosterm.

M. kemanga Bl. M. superba Hooker f.

M. lagenifera Griff.

Species of uncertain position

M. acutigemma Kosterm. M. persiciformis Wu & Ming M. bompardii Kosterm. M. subsessifolia Kosterm.

M. bullata Kosterm. M. taipa Buch.-Hamilton M. campospermoides

Kosterm.

M. transversalis Kosterm.

M. hiemalis Liang Jian Ying M. utana Utana M. maingayii Hooker f.

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as fresh fruit or used green as a salad (Angeles, 1992; Bompard, 1992a).

Mangifera pajang has the largest fruit in the genus, and is an attractive fruit.

Mangifera odorata is grown in the Philippines and Indonesia, and has occasionally been used as a rootstock for mango (Ochse, 1931; Bompard, 1992c).

Mangifera odorata is widely grown in the humid lowlands of South-east Asia in areas that are unsuitable for mango as a mango substitute. Mangifera lau- rinaand M. pentandra are appreciated as salad ingredients (Bompard, 1992d).

In addition, M. grifﬁ thii, M. minor, M. monandra, M. quadriﬁ da and M. similis have palatable fruit that are considered to have great potential (Gruezo, 1992). All mango cultivars belong to the species M. indica.

According to De Candolle (1884), ‘It is impossible to doubt that it (the mango) is a native of south Asia or of the Malay Archipelago, when we see the multitude of varieties cultivated in those countries, the number of ancient names, in particular a Sanskrit name, its abundance in the gardens of Bengal, of Deccan peninsula, and of Ceylon even in Rheede’s time (i.e. 1683).’

Although the centre of origin and diversity of the genus Mangifera is now ﬁ rmly established as being in South-east Asia, the origin of M. indica has been a matter of speculation for many years. The fossil record provides few clues, as only a single fossil bearing the imprint of a leaf of M. pentandra has ever been found (Seward, 1912). Mangifera indica is believed to have ﬁ rst appeared during the Quatenary period (Mukherjee, 1951b). Blume (1850) considered that mango might have originated from several related species, primarily located in the Malay archipelago.

On the basis of ancient accounts of travellers and the written historical record, it was believed for many years that mango must have originated in India and spread outwards from there to South-east Asia and thence to the New World and Africa. Because north-eastern India is at the northernmost edge of the distribution of the Mangiferaspecies, Hooker (1876) suggested that mango might have been naturalized in India. The historical record provides a sometimes conﬂ icting account of the distribution of mango. Miquel (1859) did not record it as being wild in the Indonesian archipelago. Accord- ing to Rumphius (1741), the mango was introduced into certain islands of the Indonesian archipelago within recent times; however, the mango was in cultivation in Java at least as early as ad 900–1100, when the temple at Borobo- dur was built and faced with carvings of the Buddha in contemplation under a mango tree (Plate 3). Based upon taxonomic and recent molecular evidence, it is now apparent that the mango probably evolved within a large area including north-western Myanmar, Bangladesh and north-eastern India (see Bompard, Chapter 2, this volume).

Polyembryonic and monoembryonic M. indica

Within M. indica, there are two distinct types that can be distinguished on the basis of their mode of reproduction and their respective centres of diversity:

a subtropical group with monoembryonic seed (Indian type) and a tropical group with polyembryonic seed (South-east Asian). A few polyembryonic cultivars occur along the west coast of India; however, they may have been introduced into Goa from South-east Asia, perhaps by the Portuguese from

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their colonies of Malacca in the Malay Peninsula or Timor in the Indonesian archipelago. Kumar et al. (2001) estimated the genetic relatedness among ten polyembryonic and monoembryonic cultivars from the west coast of southern India using genomic and chloroplast DNA RFLP analysis. The cultivars could be grouped on the basis of embryo type (i.e. monoembryonic and polyembryonic) and had distinctly different genetic backgrounds. They con- cluded that polyembryonic mangoes could not have originated in India, and must have been introduced, probably from South-east Asia.

Domestication of mango

Historical record

It is probable that mango cultivation originated in India, where De Candolle (1884) estimated that mango cultivation appeared to have begun at least 4000 years ago. In the early period of domestication, mango trees probably yielded small fruit with thin ﬂ esh. Such fruit can be found today in north-eastern India and in the Andaman Islands (Anonymous, 1992). Folk selections of superior seedlings over many hundreds of years would have resulted in larger fruit with thicker ﬂ esh. Mukherjee (1950a, b) described many of these primitive selections from Orissa in north-eastern India; they demonstrated great variation in fruit shape and size.

The mango is a very important cultural and religious symbol of India.

Buddhist pilgrims Fa-Hien and Sung-Yun mentioned in their travel notes that the Gautama Buddha was presented with a mango grove by Amradarika (c.500 bc) as a place for meditation (Popenoe, 1932). According to Burns and Prayag (1921), a mango tree is depicted in friezes on the stupa of Bharut, which was constructed c.100 bc. Other travellers to India, including the Chi- nese Hwen T’sung (ad 632–645), the Arabs Ibn Hankal (ad 902–968) and Ibn Batuta (ad 1325–1349) and the Portuguese Lurdovei de Varthema (ad 1503–

1508), all described the mango. The Indian subcontinent was the birthplace of some of the earliest highly developed civilizations, and over the centuries, India exerted strong cultural, religious and commercial inﬂ uence over South and South-east Asia. In successive waves, Hinduism, Buddhism and Islam were introduced into South-east Asia from India. To this day, many commonly used words in Indonesia are derived from both Sanskrit and Tamil.

One of the most widely used words for mango in Malaysia and Java (Indone- sia) is ‘mangga’, which is derived from the Tamil ‘manga’. Traders and monks from India possibly introduced superior selections of mango into South-east Asia; however, vegetative propagation was unknown in India until after the arrival of the Portuguese in Goa in the 15th century. Moreover, the most important mango selections of Thailand, Cambodia, Vietnam, Malaysia, Indo- nesia and the Philippines historically have all been of the polyembryonic type, and have traditionally been seed propagated. Until the establishment of Portuguese enclaves on the coast of India beginning in the late 15th century, mango cultivars did not exist in India, as there was no known method for vegetatively propagating superior selections (see Iyer and Schnell,

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Chapter 4, this volume). However, under the Moghul emperor Akbar (1556–

1605), the best selections of seedling mangoes were propagated by approach grafting and were planted in large orchards. The ‘Lakh Bagh’, a mango orchard of 100,000 trees, was planted near Darbhanga in Bihar. Perhaps noth- ing more eloquently attests to the importance of this fruit and the esteem in which it was held than this vast mango orchard. The Ain-i-Akbari, an ency- clopedic work that was written during the reign of Akbar, contains a lengthy account of the mango, and includes information about the quality of the fruit and varietal characteristics. There was evidently a strong body of information about mango cultivation that had accumulated up to that time. Most of the mango cultivars of India had their origin in those years, and have been maintained under cultivation for over 400 years by vegetative propagation.

‘Alphonso’, ‘Dashehari’, ‘Langra’, ‘Rani Pasand’, ‘Safdar Pasand’ and other mango cultivars were selected during that time. Relics of orchards from the time of Akbar are found in different parts of India, and it has been suggested that they could still provide valuable material for selection of superior mango cultivars.

Distribution

Spreading from the centres of domestication

The global spread of mangoes and their cultivation outside their original centres of domestication probably did not occur until the beginning of the European voyages of discovery and colonialization in the 15th and 16th centuries. Because mango seeds are recalcitrant, and cannot survive for more than a few days or weeks, mango germplasm in the early days must have been transported as ripe fruit, seedlings or, later on, as grafted plants. It is believed that the Portuguese transported the mango from their colonies in India to their African colonies, although Purseglove (1972) suggested that it might also have been introduced to Africa via Persia and Arabia in the 10th century by Arab traders. The Portuguese later introduced the mango into Brazil from their African colonies of Mozambique and Angola. Spaniards, who encountered a mango-growing civilization in the Philippines after Magellan’s passage across the Pacifi c Ocean, introduced polyembryonic mango types to their New World colonies through the Pacifi c trading ports of Mexico and Panama. The most important, traditional mango cultivar in Mexico remains the ‘Manila’, refl ecting its Philippine origin. ‘Carabao’ and

‘Manila’ are probably identical. The mango was introduced to the West Indies in the mid- to late 18th century, probably from Brazil. The ﬁ rst introductions of mango into Florida (USA) occurred in 1861, and involved the ‘No. 11’

polyembryonic seedling from Cuba. Seven years later, another polyembryonic selection, ‘Peach’ was introduced into the state (Knight and Schnell, 1993). Many of the early introductions into Florida proved to be unproductive, although ‘Mulgoba’ was planted on a small commercial scale (this cultivar is referred to as ‘Mulgoa’ in India, ‘Mulgoba’ in the USA and ‘Malgoa’ in Malaysia).

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Secondary ‘centres of diversity’

In 1910, a seedling of ‘Mulgoba’ came into production in Florida. Its fruit had a highly attractive red blush, and appeared to bear more heavily than its parent(s) (Wolfe, 1962). This selection was named ‘Haden’. Although ‘Haden’

was not superior with respect to fruit quality in comparison to the imported germplasm from India, its genetic base was much wider. During the 20th century, more introductions of mango germplasm into Florida occurred from South-east Asia (the Philippines, Cambodia), India and elsewhere. It was at one time believed that these introductions of mango germplasm created a secondary centre of diversity of the species (Knight and Schnell, 1993).

‘Eldon’, ‘Glenn’, ‘Lippens’, ‘Osteen’, ‘Parvin’, ‘Smith’, ‘Springfels’, ‘Tommy Atkins’ and ‘Zill’ are progeny of ‘Haden’. ‘Saigon’ seedlings were selections made from ‘Cambodiana’, a polyembryonic introduction from Indochina.

From ‘Saigon’ seedlings, ‘Alice’, ‘Herman’ and ‘Florigon’ were selected.

Based upon more recent genetic analysis involving microsatellite markers, it is now estimated that the majority of Florida cultivars are descended from only four monoembryonic Indian mango cultivar accessions, i.e., ‘Mulgoba’,

‘Sandersha’, ‘Amini’ and ‘Bombay’, together with the polyembryonic ‘Tur- pentine’ from the West Indies (Schnell et al., 2006). The Florida mango cultivars have been found to be highly adaptable to many agroecological areas and bear regularly, whereas many of the outstanding Indian cultivars have been unproductive outside their centre of domestication, and are alternate bearing. These selections also have a highly attractive red blush at maturity, fi rm fl esh, a high fl esh to seed ratio and a regular bearing habit. Some of the Florida cultivars, for example ‘Tommy Atkins’, ‘Keitt’, etc. are also moder- ately resistant to anthracnose, the most important production and postharvest problem of mango in many areas. In the latter half of the 20th century, plantings of Florida cultivars have been established in many countries and now form the basis of international trade of mangoes.

Current distribution

The mango is cultivated commercially throughout the tropics and in many subtropical areas. It is grown at the equator and at a latitude of 35–37q in southern Spain. According to Knight and Schnell (1993), ‘The process that began in Florida – introduction of superior germplasm from abroad followed by selection of improved cultivars adapted to local conditions – is now underway in many areas.’

1.4 Germplasm Conservation

Genetic erosion

The Mangiferaspecies have their centre of diversity and origin in South-east Asia, a region that has experienced great economic development in recent years. Vast wooded areas have been completely or partially deforested either for expanding agriculture or for removal of tropical hardwoods for export.

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This has caused great genetic erosion within many species and genera. The Mangiferaspecies, like many other tropical fruit trees, are canopy and emergent trees of the tropical rainforest (Kaur et al., 1980). These trees are widely scattered in the tropical rainforest, ﬂ ower erratically and reproduce from large seeds that deteriorate rapidly. As such, they are particularly vulnerable and in danger of extinction.

Collection and documentation of Mangifera germplasm

The International Plant Genetic Resources Institute (IPGRI), formerly known as the International Board for Plant Genetic Resources (IBPGR), commis- sioned an ecogeographical study of known Mangiferagenetic resources (Muk- herjee, 1985). Based upon this documentation, a joint IBPGR-International Union for the Conservation of Nature (IUCN)-World Wildlife Fund (WWF) project was initiated to collect wild mangoes on the island of Borneo and in the Malay Peninsula (Bompard, 1989), the regions that held the highest con- centrations of Mangiferaspecies. Kostermans and Bompard (1993), in the lat- est revision of the taxonomy of Mangifera, recognized 69 species, many of which were collected during the course of this project (Table 1.1). Because of the loss of natural habitat, the establishment of in situ and ex situ germplasm collections of Mangiferaspecies was considered to be imperative.

Relevance of germplasm resources to mango improvement

The genetic improvement of mango hitherto has depended on the utilization of the genetic variability found within a single species, M. indica. According to Mukherjee (1985), ‘A concerted sampling strategy should be devised for ex situsamples to meet urgent needs for use in research for improvement of the crop through breeding or as rootstocks. Sources of resistance to mango mal- formation, anthracnose, powdery mildew, gall midge are urgently needed.’

1.5 Importance of Mango

Cultivars

A partial list of the principal mango cultivars has been provided in Table 1.2.

This list includes many cultivars that were identifi ed in a survey of world mango production compiled by Watson and Winston (1984). The distribution of mango cultivars outside their centres of domestication can be attributed primarily to three historical events: (i) the movement of Indian varieties (monoembryonic) along the trade routes of the Portuguese to Africa and South America; (ii) the spread of South-east Asian varieties (polyembryonic) across the Pacifi c Ocean to Central and South America by the Spaniards; and (iii) the identifi cation of improved mango cultivars initially in Florida and

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Table 1.2. Most important mango cultivars in major producing countries.

Continent Country Cultivars

Africa Cote d’Ivoire ‘Amelie’, ‘Kent’

Egypt ‘Alphonso’, ‘Bullock’s Heart’, ‘Hindi be Sennara’, ‘Langra’, ‘Mabrouka’, ‘Pairie’, ‘Taimour’, ‘Zebda’

Kenya ‘Boubo’, ‘Ngowe’, ‘Batawi’

Mali ‘Amelie’, ‘Kent’

South Africa ‘Fascell’, ‘Haden’, ‘Keitt’, ‘Kent’, ‘Sensation’, ‘Tommy Atkins’, ‘Zill’

Asia Bangladesh ‘Aswina’, ‘Fazli’, ‘Gopal Bhog’, ‘Himsagar’, ‘Khirsapati’, ‘Langra’

China ‘Gui Fei’, ‘Tainong No. 1’, ‘Keitt’, ‘Sensation,’ ‘Zill’, ‘Zihua’, ‘Jin Huang’

India ‘Alphonso’, ‘Banganapalli’, ‘Bombay’, ‘Bombay Green’,

‘Chausa’, ‘Dashehari’, ‘Fazli’, ‘Fernandian’,

‘Himsagar’, ‘Kesar’, ‘Kishen Bhog’, ‘Langra’, ‘Mallika’,

‘Mankurad’, ‘Mulgoa’, ‘Neelum’, ‘Pairi’, ‘Samar Behisht’, ‘Suvarnarekha’, ‘Totapuri’, ‘Vanraj’, ‘Zardalu’

Indonesia ‘Arumanis’, ‘Dodol’, ‘Gedong’, ‘Golek’, ^'Madu’,^'Manalagi’

Israel ‘Haden’, ‘Tommy Atkins’, ‘Keitt’, ‘Maya’, ‘Nimrod’, ‘Kent’, ‘Palmer’

Malaysia ‘Apple Rumani’, ‘Arumanis’, ‘Golek’, ‘Kuala Selangor 2’, ‘Malgoa’

Myanmar ‘Aug Din’, ‘Ma Chit Su’, ‘Sein Ta Lone’, ^'Shwe Hin Tha’

Pakistan ‘Anwar Ratol’, ‘Began Pali’, ‘Chausa’, ‘Dashehari’,

‘Gulab Khas’, ‘Langra’, ‘Siroli’, ‘Sindhri’,

‘Suvarnarekha’, ‘Zafran’

The Philippines ‘Carabao’, ‘Manila Super’, ‘Pico’

Taiwan ‘Irwin’, ‘Jin-hwung’, ‘Keitt’, ‘Tommy Atkins’, ‘Tainong No. 1’, ‘Tsar-swain’

Thailand ‘Nam Doc Mai’, ‘Ngar Charn’, ‘Ok Rong’, ‘Keow Savoey’, ‘Pimsen Mum’

Australia ‘Calypso’, ‘Kensington Pride’

North and Central America

Costa Rica ‘Haden’, ‘Irwin’, ‘Keitt’, ‘Mora’, ‘Tommy Atkins’

Dominican Republic

‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’

Guatemala ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’

Haiti ‘Francine’, ‘Madame Francis’

Mexico ‘Ataulfo’, ‘Haden’, ‘Keitt’, ‘Kent’, ‘Manila’, ‘Palmer’, ‘Sensation’, ‘Tommy Atkins’, ‘Van Dyke’

USA ‘Keitt’, ‘Kent’, ‘Tommy Atkins’

South America

Brazil ‘Bourbon’, ‘Coite’, ‘Coquinho’, ‘Coracao’, ‘Espada’,

‘Haden’, ‘Itamaraca’, ‘Keitt’, ‘Mamao’, ‘Palmer’, ‘Rosa’,

‘Tommy Atkins’, ‘Uba’, ‘Van Dyke’

Colombia ‘Vallenato’

Ecuador ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’

Peru ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’

Venezuela ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’

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later in other new mango-producing areas, as a result of open and controlled pollination among local and introduced mango germplasm from India and South-east Asia.

Further information about many of the mango cultivars, including their fruit characters, is available in Knight et al. (Chapter 3, this volume), and in publications by Burns and Prayag (1921) for mangoes of Maharashtra, Naik and Gangolly (1950) for south Indian mangoes, Singh and Singh (1956) for Uttar Pradesh mangoes, Mukherjee (1948) for Bengal mangoes and Camp- bell (1992) for Florida mangoes.

Because many clonally propagated mango cultivars have unique local and/or regional names, there is considerable confusion in nomenclature. The Indian Agricultural Research Institute (IARI), New Delhi, has been recognized by the International Society for Horticultural Science (ISHS) as the International Registration Authority for Mango, whose mission is to consoli- date superﬂ uous names of mango cultivars. The potential for molecular, for example randomly ampliﬁ ed polymorphic DNA (RAPD), markers, to resolve much of this confusion has been demonstrated by Schnell and Knight (1992), Degani et al. (1993), Schnell et al. (1995), Eiadthong et al. (1999a), Kashkush et al.

(2001) and Gonzalez et al. (2002) (see Bompard, Chapter 2 and Iyer and Schnell, Chapter 4, this volume).

There is little variation among seedlings derived from polyembryonic mangoes. None the less, a certain amount of variability does occur, probably as a result of somatic mutation. Thus, in Indonesia there are several ‘Aru- manis’ selections that are denoted numerically, for example ‘Arumanis 1’,

‘Arumanis 2’, etc. In addition, although Philippine mango cultivars are distinguished by different names, for example ‘Carabao’, ‘Manila’, ‘Philippine’, etc., the differences among them are quite subtle.

1.6 Production and Uses

The mango is the most important fruit of Asia, and currently ranks ﬁ fth in total production (in metric tonnes) among major fruit crops worldwide, after Musa(bananas and plantains) (105,815,354 t), Citrus(all types) (105,440,168 t), grapes (65,584,233 t) and apples (59,444,377 t) (FAOSTAT, 2006). According to the Food and Agriculture Organization of the United Nations (FAO) database (FAOSTAT, 2006), world mango production has increased from 16,903,407 t in 1990 to 28,221,510 t in 2005. Much of this new production has occurred outside the traditional centres of mango culture of South and South-east Asia. In 1990, India produced approximately 51% of the world’s mangoes, but by 2005, India’s share had declined to approximately 38%, despite the substantial increase in mango production since 1990 (from 8,645,405 to 10,800,000 t between 1990 and 2005). The current leading producing nations after India include (in metric tonnes) China (3,450,000), Thailand (1,800,000), Pakistan (1,673,900), Mexico (1,600,000), Indonesia (1,478,204), Brazil (1,000,000) and the Philippines (950,000). Although world production has increased by 67% between 1990 and 2005, mango exports have increased almost sixfold

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from 158,030 to 907,782 t, with total export value estimated to be US$583,763,000 (FAOSTAT, 2006). The major exporting countries are (in metric tonnes) Mexico (212,505), India (156,222) and Brazil (111,181). As a result, mangoes are widely available as fresh fruit and as processed products (i.e.

dried fruit, dairy products, juice, pickles, etc.).

Mangoes are an important component of the diet in many less developed countries in the subtropics and tropics. In regions of the world that have experienced low living standards and serious nutritional defi ciencies, their attractiveness and fl avour have also enhanced the quality of life. Surplus production has increasingly been processed and fruit of certain cultivars is destined for export as fresh fruit. Approximately 1% of mango production is utilized for processing for juice, nectars, preserves (including chutney), fruit leather, dried fruit slices, frozen pulp and as a fl avouring for baked goods, ice cream, yoghurt, etc. (see Raymundo et al., Chapter 17, this volume). No part of the fruit is wasted. In India and the subcontinent, the seed is used for extraction of starch ‘amchur’, and the peels (skin) have been used as a source of anacardic acid. Mango wood is a low quality timber, and the bark of the tree is an important source of tannins for curing leather.

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(ed. R.E. Litz) 19

2 Taxonomy and Systematics

J.M. Bompard

Les Mazes, Montaud, France

2.1 Introduction 19

2.2 The Genus Mangifera L. 20

Distribution 20

Ecology and habitat 20

2.3 Taxonomy and Systematics 22

Taxonomic history 22

2.4 Phytogeography 28

Species distribution 28

Subgenera and section distribution 29

2.5 Interspeciﬁ c Molecular Characterization 30

2.6 Region of Origin of the Genus 31

2.7 Origin of the Common Mango 32

The common mango in South-east Asia 32

2.8 Conclusion 35

Potential contribution of wild species to mango cultivation 35

Source of rootstock 35

Hybridization 36

Potential of wild species 36

2.1 Introduction

The genus Mangifera is one of the 73 genera (c.850 species) belonging to the family of Anacardiaceae, in the order of Sapindales. Anacardiaceae is a family of mainly tropical species, with a few representatives in temperate regions.

Malesia, which is the phytogeographic region extending from the Malay Peninsula south of the Kangar-Pattani line to the Bismarck Archipelago east of New Guinea (Whitmore, 1975) contains more species in the Anacardiaceae than any other area. Within Malesia occurrence is mainly in Western Malesia (Ding Hou, 1978b).

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J.M. Bompard 20

Apart from mango, several other cultivated fruit trees belong to the family, for example the ambarella or Otaheite apple (Spondias dulcis Forst.) probably from Melanesia, and the yellow and purple mombins (Spondias mombin L. and S. purpurea L., respectively) from tropical America, the Boueaspecies from IndoMalesia, dragon plums (Dracontomelum spp.) from IndoMalesia and the Paciﬁ c region, kafﬁ r plum (Harpephyllum caffrum Bernh. ex K. Krause) and the marula plum (Sclerocarya caffra Sond.) of southern Africa. The cashew (Anacardium occidentale L.) is from tropical America and the pistachio (Pistacia vera L.) from Iran and Central Asia. Anacardiaceous species also yield other valuable products: wood (several genera), gums and resins (Pistacia spp.), varnishes (Rhus spp. and Melanorrhoea spp., ‘lacquer trees’) and tanning materials (Rhusspp. and Schinopsisspp.). It is also a family well known for the dermal irritation produced by some of its members, such as the poison ivies and oaks (Rhusspp.) in North America, rengas (Glutaspp.) in South- east Asia and other species including some Mangifera species whose resinous sap may induce a mild to strong allergic reaction.

2.2 The Genus Mangifera L.

Distribution

The range of natural distribution of the 69 Mangifera species is mainly restricted to tropical Asia, and extends as far north as 27° latitude and as far east as the Carolines Islands. Wild mangoes occur in India, Sri Lanka, Ban- gladesh, Myanmar, Sikkim, Thailand, Kampuchea, Vietnam, Laos, southern China, Malaysia, Singapore, Indonesia, Brunei, the Philippines, Papua New Guinea and the Solomon and Carolines Islands. The highest species diversity, c.29 species, occurs in western Malesia, especially in peninsular Malay- sia and in Borneo and Sumatra, which represent the heart of the distribution range of the genus (Fig. 2.1).

Ecology and habitat

The majority of Mangifera species occur as a rule as scattered individuals in tropical lowland rainforests on well-drained soils. The species are distributed mostly below 300 m, but can occur up to c.1000 m above sea level, on well-drained soils (44 species), in periodically inundated areas (ten species) and in certain types of swamp forest (i.e. M. gedebe, M. grifﬁ thii and M. parvi- folia). Three species are mainly found in sub-montane forests above 1000 m and occasionally up to 1700 m above sea level (M. bompardii, M. dongnaiensis and M. orophila). There are also species that are adapted to seasonally dry climates in deciduous or semi-deciduous forests (e.g. M. caloneura, M. collina, M. timorensis and M. zeylanica). A few species occur north of the Tropic of Cancer, for example M. austro-yunnanensis and M. persiciformi