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MANGIFERA OF

Fitmawati Ibna Hayati

With Contribution by Eko Prasetyo Adi Zulkifli

Photographs by Fitmawati Salpa Hartanto

UR PRESS 2018 iii

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ACKNOWLEDGMENT

First of all, We would like to thank to Allah, God of the ‗Alameen (the whole universe), for all of His blessings. It was Allah who has opened a broad way for us so that We can completed the manuscript of of Sumatra.

The present results would not have been possible without the generous support of our colleagues in Biology Department of Universitas Riau, our biology students who have done the research related to Mangifera, and HIKOM. The Indonesian Minister of Research Technology and Higher Education provided financial grant and I am much grateful to DIKTI for unstinting support.

I am indebted to Prof. Mien A. Rifai, Prof. Elizabet Wijaya and Prof. Alex Hartana who gave me guidance, support, and encouragement to finish this manuscript.

It is a pleasure to warmly thank my exploration companions, Anggi, Uyun, Fransiska, Anto, Suci, Puji, Roslina, Masruroh, Ibna, Sandi, Erwina, Esi, Miladia, Nurul, Rissan, Syaiful fortheir time in the field; my greatest debt goes to Salpa, Adi, Syafroni who accompanied me on all trips,tirelessly organising the camp, the meals, cutting the plot boundaries, climbing trees for proper botanical collection, and helping in pre-identifying the samples.

We would also like to express our sincere and deep gratitude to our supervisors, Prof. Mien, for his guidance and help, huge understansing, patience, kindness, immense support, and encouragement throughout this manuscript,.

Numerous people have helped us during our research and the writing up. We would like to thank them all: Nery Sofyanti, Anggi Swita, Anto, Fransiska Warni, Suci Rochyati who have helped me exploring in Central Sumatra, Adi Zulkifli vi

Salpa Hartanto and Rissan Suriatno, who have helped me exploring mango in eastern Sumatra until mapping distribution, Ibna Hayati, Erwina Juliantari, Sandi Pratiwi Harahap, Syaiful Ramadhan, Syafroni Pranata who have helped me exploring mango in southern Sumatra. For molecular team: Roslina, Masruroh, Ibna, Sandi, Wiwin, Esi, Miladia, Nurul.

PREFACE

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Diversity Mango (Mangifera) in is an important and interesting aspect to be studied for mango improvement program relies heavily on the genetic diversity available. Exploration and surveys of mango genetic diversity in Indonesia have been conducted in Java (Fitmawati 2003, Sulistiowati (1989), West Kalimantan and South Kalimantan (Kostermans and Bompart 1989), South Sulawesi (Wirawan 1999), Southeast Sulawesi (Fitmawati 2005) exploration and surveying the diversity of mango species in eastern and southern Sumatra has never been done.It is very important to do exploration to reveal the biological wealth of Sumatran mangoes in order to sow the resources of germplasm of wild mangoes in Sumatra that are racing with the rapid transfer of land function to the area of oil palm and rubber plantation.

Mango has become a major 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 ‘ (Purseglove, 1972). However, improvement of tree crops has lagged far behind field crops for several reasons: their heterogeneity, 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 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 adoption of modern fi eld practices, which include irrigation management, control of flowering, etc. Agricultural practices are currently undergoing another revolution, as integrated pest and disease management replaces viii the earlier reliance on agrichemicals, and emerging fields within biotechnology begin to impact development.

The Mangifera species 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. This has caused great genetic erosion within many species and genera. The Mangifera species, 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, erratically and reproduce from large seeds that deteriorate rapidly. As such, they are particularly vulnerable and in danger of extinction.

Mango ( L.) has attained the status of most important fruit crop in tropics and subtropics, particularly in Asia and considered as ‗king of fruits‘. India occupies first place in mango production of the world and accounts for almost half of the global production and area. India being primary and secondary centre of domestication of Mangifera indica, substantial contribution of mango industry in economy, export, livelihood support is well known. Enormous genetic diversity exists in the country. Considering the importance of mango national database on mango has been developed with the support of DBT (Department of Biotechnology, Government of India, New Delhi) in network mode involving ten research institutes/universities viz., Central Institute for Subtropical Horticulture (CISH), Lucknow; Indian institute of Horticultural Research (IIHR), Bengaluru; Horticultural College & Research Institute for Women (HCRIW) (Tamil Nadu Agricultural University), Trichy; Bidhan Chandra Krishi Viswavidyalaya (BCKV), Nadia; National Research Centre for Litchi (NRCL), Muzaffarpur; Central Horticultural Experiment ix

Station (CHES), Bhubaneswar; Fruit Research Station (FRS) (Andhra Pradesh Horticultural University), Sangareddy; Regional Fruit Research Station (RFRS) (Dr. Balasaheb Sawant Konkan Krishi Vidyapeeth), Vengurle; Agriculture Experimental Station (AES) (Navsari Agriculture University), Paria and ICAR Research Complex for NEH Region (ICAR RC), Barapani.

As the information on morphological characterization of regional farmer‘s variety and eco-geographical adaptations, package of practices, traditional knowledge, geographical indicators related to different varieties specific to mango growing areas of India, thus the involvement of these institutions for collection of information at district level resulted a huge data set representing country level information, which further reorganized for the development of an interactive and informative national network database meeting the strong need of the country. All centres collected geographical location specific information and final integration of the digitized information developed at Central Institute for Subtropical Horticulture (CISH), Lucknow as National Mango Database.

Mango database providing information on genetic, phenotypic, molecular markers, proteins, endogenous phytochemicals, usage, packages of practices, traditional knowledge, nutritional and medicinal value, production, harvesting and postharvest technologies, on farm conservation sites, gene bank status, IPR issues, diseases and pests, geographical indicators, bibliographic records and GIS applications for georeference database and several other important aspects on the crop. The national database on various aspects of mango germplasm would be highly useful for biotechnologists, students, researchers, farmers, policy makers and mango lovers. From IPR point of view, utilization of germplasm in breeding programs and other biotechnological researches on mango will be highly useful. x

Genetic resources molecular characterization and bioinformatics resource information including nucleic acid, ESTs, proteins, chemicals of different Mangifera collected by CISH, Lucknow. CISH, Lucknow also generated information on GIS based mapping of thematic areas like genetic diversity, molecular information and varietal climate suitability, database useful for DUS testing, IPR issues including patents on mango.

Development and maintenance of national database containing information on various aspects related to mango will result in proper utilization of genetic resources in improvement programmes and germplasm conservation, which will also provide sustainable management tools. Mango database also delivers information on various aspects like improvement, production technology, crop protection, biotechnology, genetics, bibliographic citations and related issues affecting livelihoods and nutritional security for the benefit of researchers, farmers, local communities and tropical fruit tree user groups.

Information on mango accessions available in different gene banks was augmented and updated for providing information of the accessions from various parts of the country. Existing documentation available on mango have been converted into digital format. Information on the farmer‘s varieties and landraces, available in the literatures were digitized for the updates of database. For this purpose, descriptions made by research workers from different parts of the country were collected. Information on the important eco-geographical areas rich in genetic diversity presented could also be retrieved from the database. The database is also equipped with characterization information on different mango varieties along with extant farmer‘s varieties, information on different mango diseases & pest, symptoms, integrated management, products, DUS testing of mango varieties, identified geographical indications, database on the chemical profile of mango, secured patents and other IPR issues. A lot of germplasm has been conserved and characterized in different field gene banks in xi different parts of the country. From IPR point of view, utilization of germplasm in breeding programme and other biotechnological research, database development on mango genetic resources will be highly useful to the research worker involved in these fields. Development of national database will add a value to the national intellectual property and serve information source for the students, scientists, researchers, policy makers, exporters and farmers.

Contributors xii

Exploration Team Molecular Team Salpa Hartanto Roslina Fauziah Adi Zulkifli Masruroh Syafroni Pranata Ibna Hayati All of the team Erwina Juliantari Sandi Pratiwi Harahap Morphology Team Esi Resida Suci Rochyati Nurul Rafidah Anggi Swita Miladia Rahmawati Ningrum Fransiska Warni Pakpahan Rissan Suriatno Photography Team Syaiful Ramadhan Salpa Hartanto Anto Fitmawati

Anatomical Team Mapping Team Puji Astuti Eko Prasetya Adi Zulkifli Supervisi Team Nery Sofiyanti Herman Fitra Suzanti Radith Mahatma

Suci Rochyati Biologi 2005 Morphology Team, Exploration Team

Salpa Hartanto xi

Biologi 2008 Photography, Exploration Team

Anggi Swita Biologi 2009 Morphology Team, Exploration Team

Fransiska Warni Pakpahan Biologi 2009 Morphology Team, Exploration Team

Anto Biologi 2009 Morphology Team, Exploration Team

Puji Astuti Biologi 2009 Anatomy Team, Exploration Team

Adi Zulkifli Biologi 2010 Mapping Distribution, Exploration Team

Roslina Fauziah Biologi 2011 Molecular Team

Masruroh Biologi 2011 Molecular Team

Ibna Hayati Biologi 2012 Molecular Team, Exploration Team

Erwina Juliantari Biologi 2012 Molecular Team, Exploration Team

Sandi Pratiwi Harahap Biologi 2012 Molecular Team, Exploration Team xii

Syafroni Pranata Biologi 2012 Photography, Exploration Team

Esi Resida Biologi 2013 Molecular Team, Exploration Team

Nurul Rafidah Biologi 2013 Molecular Team, Exploration Team

Miladia Rahmawati Ningrum Biologi 2013 Molecular Team, Exploration Team

Syaiful Ramadhan Biologi 2013 Morpholgy Team, Exploration Team

Rissan Suriatno Biologi 2013 Morpholgy Team, Exploration Team

Contents xiii

Preface

Chapter 1 Mangifera Introduction The Mangifera Habit Distribution Ecology The Tree Inflorescence The Fruit Seeds The

Chapter 2 The Sumatra Introduction Geographical Features Climate Vegetation Rain fall Volume Altitude

Chapter 3 The Species Introduction Distribution Pattern 1. Mangifera foetida Lour. 2. Mangifera odorata Griff. 3. Mangifera laurina Bl. 4. Mangifera indica L. 5. Mangifera zeylanica (Bl.) Hooker f. 6. Mangifera sumatrana Miq. 7. Mangifera magnifica Kochummen. 8. Mangifera quadrifida Jack. 9. Mangifera torquenda Kosterm. 10. Mangifera kemanga Bl. xiv

11. Mangifera lalijiwa Kosterm. 12. Mangifera griffithii Hooker f.

Chapter 4 Phylogenetic Analysis What the common ancestor of Mangifera is What the suitable DNA Barcode of Mangifera is What the ITS Sequence can tells What the trnL-F Intergenic Spacer can informs What the rbcL Gene can manifests What the matK Gene can says

Chapter 5 Conservation What Mangifera should be conserved Conservation Status of Mangifera in Sumatra

Mangifera 15

CHAPTER 1 Introduction

What Mangifera is

he genus Mangifera is one of the 73 genera belonging to family, in the order. There were 69 species described by Kostermans and T Bompard (1993) in the latest classification of Mangifera. The natural habitat of Mangifera is restricted to tropical Asia. The highest species diversity found in the western part of Malesia (particularly Malay Peninsula, Sumatra and which represent the heart of the distribution range of the genus).

Mangifera species are rarely gregarious, usually living alone and widely distributed in natural habitat. Almost all the species of Fig. 1.Tree of M. kemanga during Mangifera occur as exploration in scattered individuals in wet tropical lowland Southern rainforests Sumatra (44 species on well-drained soils, 10 species in periodically Mangifera 16 inundated areas, 3 species in swamp forest). A few are found in dry, deciduous forests (3 species), savannahs and also mountain forest. The greater part of the species grow in lowland up to 700 m altitude, even up to 1700 m above sea level for sub-montane forests species (7 species). For lowland species, the higher the altitude, the more difficult to bear fruit.

Mangifera did not have the fixed season for flowering and fruiting. Flowering in Mangifera is unique, usually founded only one part of the tree flowers during one season, while the other part is waiting for the other season. Several species tend to have on/off season that become a notable constraint in cultivation.

From millions of flowers that Mangifera has in a tree, only ten fruits were produced. Fortunately not all the species undergone this circumtances. Some wild species tend to have flowers resistance against wet climate such as Mangifera in Sumatra (Fitmawati et al. 2013). This also become one of the substantial properties for cultivation. Pollination becomes the most mandatory thing for setting Mangifera fruit. When the rainy season starts, the flowering occur and when the rainy season ends,the ripening fruit appears. It takes 80-90 days from ripening since fertilization. Some species has tendency to fail in setting fruit because of high rainfall but not for Mangifera species in Sumatra. Mangifera 17

The dispersion of Mangifera was assisted by water (one species) and animals (the rest species). Based on their mode of reproduction, Mangifera divided into two centres of diversity. A subtropical group with monoembryonic seed (Indian type) and a tropical group with polyembryonic seed (Southeast Asian). Fitmawati and Hartana (2010) reported that the polyembrionic species of Mangifera origin from Celebes. It is assumed that the common ancestor of Mangifera is the polyembryonic one (Fitmawati & Hartana

2010). Fig. 2.Fruit candidates from million flowers of M. indica

How unique Sumatran Mangifera are

Mangifera is one of the most important genus which commercially succesfull in fruit production in the world. Mangifera growth in low rain volume required dry climate for four months to stimulate the inflorensence. The characteristics of Mangifera species in Sumatra were tolerant to high rainfall, capable of fruiting out of season, high production and flowers resist against wet climate. The species with these traits had a germplasm resources potential (Fitmawati et al. 2013). Mangifera 18

Exploration on Mangifera species has been conducted by Fitmawati et al. (2013), (2015) and (2017) in 8 provinces in Greater Sumatra Island. Twelve of Mangifera species which typical in Sumatra were obtained. Mangifera species in Sumatra were divided into three categories such as: wild types, semi cultivated types and cultivated types (Fitmawati et al. 2015). Due to high frequency of forest and land fires in Sumatra, specific types of Mangifera Sumatra were threatened in natural habitat, therefore wild germplasm resources must be conserved before became extinct in the wild.

What Mangifera looks like

Tree. Emergent tree of Mangifera in tropical rainforest of Sumatra can attain a height of 40 m or more and survive for hundred years. Tree architectures (crown shape and size) are highly variable. They can be supporting data to determine the species. Tree architecture was correlated to the starting point of branch growth. The excurrent branching habit resulting in conoid shapes and the decurrent branching habit resulting in round shapes.

Mangifera 19

Leaves. Leaves are simple and alternate, with high variation in petioles length. morphology is highly variable depending on the cultivar or infrageneric level. High morphological plasticity of leaves become one of the most difficult task to delimitate species in the genus even in the infraspecific level. Leaves can be lanceolate, oblong, ovate and intermediate types involving these forms.The leaf shape changes and the size diminishes drastically towards the inflorescences.

The reticulation of leaves is often a distinctive character. From completely smooth-leaved species without a trace of reticulation, the leaves being rigidly coriaceous leaves; distinctly conspicuously raised, rather lax reticulation; the greater part of the species has dense reticulation with different pattern from usual, anastomosing kind, to the non- anastomosing meander Fig. 3.Stiffly coriaceous leaves of M. magnifica type, to the line like. from Kampar Thick leaves tend to have less discernible reticulation than the thinner leaves.

Mangifera 20

Generally, leaf texture of Mangifera divided into two large group namely coriaceous and chartaceous type. The species with stiffly, woody coriaceous, having fibres and smell of turpentine after breaking is Mangifera magnifica. Coriaceous type tend to be more primitive than chartaceous. However, molecular data support M. magnifica as the modern species of Mangifera in Sumatra based on the longest genetic distance from neighbor joining method (Fitmawati et al. 2016a). Some species showed aberrant of leaf morphology. In M. quadrifida, young leaves has different colour. The upper surface has greenish while lower surface has dark pink colour. Young leaves color changing gradually to light and then dark with age. In M. macrocarpa, seedling and 3 m sapling leaves look like grass, long and linear with numerous ribs.

Fig. 4. Young leaves colour variation in Mangifera species. From reddish, brownish, yellowish, greenish to pink Mangifera 21

Fossil.The oldest Mangifera-like leaf fossil was from the upper Paleocene of northeast India (Mehrotra et al. 1998). There were many leaf fossils of Mangifera reported since 1912. However, all of them need corroboration. Accuracy of fossil identification was still doubted. Sawangchote et al. (2009) founded two leaf fossil from late Oligocene or early Miocene deposits in northern which were believed to be an area with long history of evolution and diversification of Mangifera, particularly M. indica. Fossil wood of Mangifera is also known from the middle Miocene of Assam, middle to upper Miocene of West Bengal, upper Miocene-Pliocene of Arunachal Pradesh and Pondicherry, Pliocene of Rajasthan, and the Neogene of Sumatra and Borneo (Mehrotra et al. 1998).

Inflorescence. All species of Mangifera have panicle inflorescence. The trees often do partial flowering in one year while another part in the following year. This is one of the adaptation mechanisms to divide the task as food suplier and fruit producer.

Inflorescence candidate are enclosed by a number of prophylls (bud scales). These bud scales are either small and a few in number or very large and numerous, leaving a collar of large scar on the branchlet (deciduous species). The panicles represent a continuation of a leaf branch, the leaves being Mangifera 22

reduced to bracts that drop usually at an early stage, the branching following the same pattern as that of the leafy branches. The leaves below the inflorescence are, as a rule, much shorter and often with different apex and Fig. 5.Bud scales of M. kemanga base as compared with the normal leaves. It is recommended therefore that collectors next to a flowering branch, collect also a vegetative branch with the normally sized leaves. The panicles are constantly pyramidal or conical in outline.

The panicles cannot be used for deduction of phylogenetical trends, as there is no indication how long ago they developed and an inflorescence may be looked at as primitive, but may be at the same time very young in history. Different panicles were formed different lines and cannot be co-ordinated. Fig. 6.Inflorescence of M. indica Mangifera 23

The colours of the inflorescence are very characteristic, they are different in the main peduncle and the branchlets and vary from green (usually pale green) with or without patches of of the main peduncle to pure white of the entire inflorescence, to more or less darker to violet red of the branches.

Flower. Delimitations of species in Mangifera are mainly based on flower characters. Short inflorescence has only few flowers while the large panicles have ten to thousands of flowers. The flower start opening from the base, the terminal flower of each cyme flowering the last. There are two kinds of flowers: male ones (have one or more and often and a completely abortive or a much-reduced pistil) and larger hermaphrodite flowers (one or more fertile , staminodes or none and a well-developed pistil). The ratio of male to hermaphrodite flowers is strongly influenced by environmental and cultural factors.

The flowers are 4- or 5-merous, sometimes both of them occur in the same specimens. In other species the numbers are constant. In cultivars the numbers of may increase to 6 or more. Size in flowers, , petals and reproductive organs are highly variable. Colour gradation of flowers from the first time appeared until anthesis is also highly variable. These characters Mangifera 24 can be used for specific delimitation. The wild species tend to have larger than the cultivated ones. The cultivated species tend to have more flowers than the wild ones, even precisely 3000 flowers per panicle.

The flowers have four or five sepals and petals that are ovate to ovoid to lanceolate and also thinly pubescent. The floral disc also is four- or five-lobed, fleshy and large and located above the base of the petals. There are five large, fleshy stamens, only one or two of them being fertile; the remaining stamens are sterile staminodes that are surmounted by a small gland. In addition, two or three smaller filaments arise from the lobes of the nectaries. The stamens are central. The ovule is anatropous and pendulous. It is believed that the flowers are cross- pollinated

Fig. 7.Flowers of M. foetida The number of fertile stamens varies from 1 to 10. In the one-stamen flower, the 4 abortive stamens are either present as small staminodes, that may be subulate without anthers or bearing sterile anthers. The reduction is not equal, but increases Mangifera 25 in the staminodes farthest from the fertile one. A common number is 5 fertile stamens (M. pentandra), but irregularities are found here in the number of fertile stamens. The 10-stamen species (M. decandra) are stable in their stamen number. This is considered the most primitive situation (but whether arisen early or not, is of course unknown). The filaments are white, red or purplish. The anther is dorsifix, two-locular and longitudinally, introrsely dehiscent. Often they are dark violet. The amount of pollen is very small, there are often two kinds of pollen grains, small ones and big (diploid) ones.The stamens are attached below or on the disc. In some species the petals are basally slightly connate. The stipe-like disc may be even almost lacking.The fertile stamen has its anther bent and almost touching the stigma, suggesting selfing.

Fig. 8.Narrower disc (above) and broader disc (below) of the Mangifera flower

The is seated on a disc, that can be either broader than the ovary, cushion like, relatively large, papillose and Mangifera 26 divided into 4-5 lobes, closely packed or narrow, stalk-like (Fig. 8). The superior ovary has special colour (bright yellow in M. foetida, green or white in others), is always glabrous; it is seated on the somewhat concave top of the disc, it is asymmetric and bears one style with a small, inconspicuous stigma, either on the centre of the ovary (rare) or asymmetrical to the opposite side of the fertile stamen.The disc acts as a nectary, the amount of nectar varies, may be sometimes high. The ovule is anatropous, single, the ovary one-locular, but according to Sharma (1954) the original state was a three-locular ovary.

Shape, size, flavour, pulp, terpene content, fibres content, water content of fruit are highlt variable. These are possible resulted from the cross-breeding in infrageneric level. During the development of the ovary, the apex of the ovary grows and pushes the style more sideways and lower down.

Fruit.The fruit in most cases is the typical mango fruit, kidney-shaped, the base rounded, one side with a distinct sinus above the distinctive remainder of the style base, the so-called beak or nose. The apical part of the fruit is as a rule oblique and may be rounded or pointed, the base may be asymmetrical.

Mangifera 27

Fig. 9. Shape variation of Mangifera from Rokan Hulu regency

The skin is thick or thin, smooth or roughish-scurfy, green at maturity to whitish to green, flushed with yellow and red. Rough-skinned ones are easily distinguishable from the glossy, smooth skinned ones. The skin sometimes contains anacardol acid, which cause severe blisters on skin and mucous membranes. The resulting wounds take considerable time to heal. This acid is also often found in the bark, wood, and flowers and sensitive people are affected by pollen falling from the flowers. Aberrantly shaped fruits are found in M. gedebe, here they are the flattish, discus-like and in the Deciduae where they may Mangifera 28 be pear-shaped.The black-coloured mangoes belong mostly to the Rawa section. They were already mentioned by Rumphius in 1741, but wrongly interpreted by later botanists.Sometimes the beak disappeared completely and the fruit may not be compressed, but almost globose (e.g. M. foetida, M. magnifica).

The pulp is either white or yellow or orange (Fig. 8). That of the deciduous group is white, in M. pajang yellow and quite different from that of the other species in taste, smell and consistency, orange pulp is often correlated with black-skinned mangoes.The pulp contains fibres in different quantities and of different length. These fibres are attached to the endocarp. In some species the fibres are completely absent (M. magnifica), the endocarp (the ‗seed‘) looking glossy and smooth like a billiard ball.

The number of fibres can be enormous; in M. decandra they form, after the pulp has been washed out, a curly mass of very long and very dense fibres (like a medusa head). In the cultivars of M. indica, one of the main

Fig. 10.Some colour variation in pulp of Mangifera Mangifera 29 aims has been to get fibreless varieties (the commercial ones). Unfortunately they lose their typical mango aroma and taste and smell become flat.At maturity the pulp may be either solid (like a peach) or becomes more or less fluid (M. laurina, M. zeylanica), the the fruit can be sucked out.

If the fibres are short (M. torquenda) or absent (M. magnifica), the fruits are sometimes opened by making a cross cutting and twisting the two halves in opposite directions, they easily separate from the endocarp. These are called: putaran (M. torquenda in IndonesianBorneo; M . magnifica in Central Sumatra; the latter species is also called kemang pulas or kemang putar in W. Malesia; kemang= M. kemanga, putar or pulas = to turn or twist). Seeds. The seed or stone consists of a rather woody or leathery endocarp, at the outside with longitudinal curved incised lines.There are two seed coats, both very thin. In M. gedebe the inner seed coat penetrates the cotyledons, which become labyrinthine.The seeds are solitary, large and flat, ovoid oblong and surrounded by the fibrous endocarp at maturity. The testa and tegumen are thin and papery.Embryos are dicotyledonous. The cotyledons are usually equal, convex-flat, but in M. pajang they are equal and one partly enveloping the other.The cotyledons may be very flattened or not, their surface is smooth. Mangifera 30

Fig. 11. The seeds and stones of M. magnifica from Kampar

Seeds of monoembryonic mango types containa single zygotic embryo, whose cotyledons can be unequal in size or lobed inshape. The seeds of polyembryonic mango types contain one or more embryos; usually one embryo is zygotic, whereas the remaining embryos are derived directly from the nucellus, a maternal tissue. Nucellarembryos apparently lack a suspensor. Polyembryony has also been reported in Mangifera casturi, M. laurina and M. odorata (Bompard, 1993). Certain polyembryonic cultivars reportedly can produce seeds with adventitious nucellar embryos only. Early studies suggested that polyembryonyappeared to be a polygenic trait segregating as a recessive character in the progeny ofcontrolled crosses. Mango seeds are considered to be recalcitrant, and cannot survive for more than a few days or weeks at ambient temperatures. This important characteristic of mango seeds would have inhibited the long distance dispersal of mango by seed until recent times. Mangifera 31

One of the best mango cultivar from Indonesia is Arumanis cv. This cultivar is a polyembriony mango indigenous from Indonesia. The characteristics of this cultivar are sweet, soft fiber, low water content, scented fragrance and color of yellow-orange fruit flesh (idiotype), with quality standards that become favoriteof international customers (Fitmawati et al. 2009). The major problem in conventional plant breeding mango are the small number of seedlings obtained, the complex nature of panicula flowers, long life cycle, the high heterozygosity pollinate and low success rates because of the nature of self-sterile. The result is an excessive loss of quality fruit, due to pushing interest rates to be able to produce fruit lend more difficult than the mango crop. Besides the high heterozygosity mango seedlings caused difficulty obtained a uniform zygotic.

Arumanis cv. reproduces through polyembryoni mechanisms, their seeds develop with fertilization called zygotic embryo and the others without fertilization called apomixes (non-zygotic). Apomicts progenies may have the same genotype as their mother plant. In morphological characteristics, the difference vigority charactereristic are founded between zygotic and non zygotic progeny. DNA banding pattern polyembriony parent and progeny (zygotic and non-zygotic) shows difference bands. The specific bands are only owned by the parent and progeny of non-zygotic (Fitmawati et al. 2012). Mangifera 32

Fig. 12. Polyembrionic seedlings between zygotic and non-zygotic seedlings in Arumanis cv.

Pollen. Mukherjee (1953) investigated the pollen morphology of mangoand 12 other Mangifera species. Their pollen grains were tricolpate ofalmost the same size. Mondal et al. (1982, cited in Kostermans and Bompard, 1993) attempted to correlate pollen morphology with taxonomicrelationships of 17 Mangifera species based upon different characteristicsof the exine and sporoderm. They demonstrated that all of the species ofsection II (subgenus Limus) possess coarse exine; whereas there was noclear correlation with pollen type in species within section I (subgenusMangifera).

Mangifera 33

CHAPTER 2 The Sumatra

Where the Sumatra is umatra and the islands off its coast are part of the Republic Indonesia and have been since the modern nation was established in 1945. The island were divided into ten provinces (Fig. 13).Culturally, the Speople of northern Sumatra have strong affinities with those of the Malay Peninsula, with regular migrations back and forth occurring over the millenia. Oil, agriculture and logging are the mainstays of the economy.Curiously, the flora of Sumatra has been neglected in comparison to that of other islands inthe Malayan Archipelago, a fact often acknowledged by the taxonomists themselves reported by Laumonier (1997). Mangifera 34

Fig. 13. Map of Sumatra (after Anonim 2011) The Species 30

Geographical Features Sumatra is roughly located between 5° N and 5° S of the equator and between 95° E and 105° E of Greenwich. The West coast of Sumatra is bordered by the vast, undisturbed Indian Ocean. On the East coast of Sumatra the strait of Malaca separates the island from the Malaysian peninsula. In the North this strait is more than 300 km wide but it narrows down to less than 50 km where Tebing tinggi faces . Further Souththe East coast is bordered directly by the China sea.

Sumatra is a relatively narrow island which lies across the equator at the western boundary of the Malay Archipelago. With an area of 475,000 km2, it is the fifth largest island in the world and the third largest in Malesia. It is more than 1700 km long, but only 350 km at its widest point (Fig. B2), with a north-west to south-east orientation. The west coast of the island is dominated by the Bukit Barisan, a mountain range formed largerly from the uplift of sedimentary rocks and punctuated by a number of volcanoes. The Bukit Barisan has more then 130 peaks which exceed 2000 m in height and nine of these are higher than 3000 m. At 3804 m, Gunung Kerinci is the highest mountain in Sumatra and the second highest in the Sunda region. To the east of the Barisan range lie vast areas of low plains which extend to the Strait of Melaka and the Riau Archipelago. The western side of the island is mostly mountainous with lowland areas confined to a narrow coastal strip, whereas the eastern side, except in the northernmost province of Aceh, is low lying. The Tigapuluh mountains of Riau and Jambi are the only eastern The Species 31

mountain range of note, but no peaks exceed 1000 m in height.

Sumatra has a very complex geological history which is reflected by the wide variety of rock and soil types found throughout the island. Much of the highlands are of sedimentary or volcanic origin, with significant area of limestone and sandstone throughout the Bukit Barisan, while the lowlands are comprised mainly of alluvial soils, peat and sandstone (Laumonier 1997). Several of South- east Asia‘s largest and highest lakes occur in Sumatra. Danau Toba is the biggest with an area of 800 km2, whereas Danau Tujuh, at 1994 m asl, is the highest. Both of these lakes were formed by cataclysmic volcanic eruptions and it has been suggested that the pyroclastic explosion approximately 200,000 years ago which led to the formation of Danau Toba was the largest known to have occurred throughout the geological history of the Earth (Sparks et al. 1997). Pulau Samosir, which lies near the west coast of Danau Toba, was formed by the collapse and subsequent uplift of the mountains on the western periphery of the now-annihilated volcano.

In comparison with the other parts of the Indonesian arhipelago, Sumatra and neighbouring Java are geologically young, with most of Sumatra‘s granite massifs dating from the Cretaceous period. Other parts of the island are dominated by tertiary formations. Of the volcanoes, more than 30 are currently active. Many of the older volcanoes produced acidic lavas, whereas those produced in the Quartenary period are intermediate or basic (Laumonier The Species 32

1997). Although it has been difficult to find significant correlations between the different types of rock formations and the phytogeography of the island, it is noteworthy that few Mangifera occur on or over basic soils. Mangifera are scarce in the southern parts of the island (only one highland species), where the majority of the volcanes are relatively young.

Climatic Conditions The climate of Sumatra is characterized by abundant rainfall moderately well distributed over the year. The typical dry and wet seasons of Java and East Indonesia are only weakly developed. Due to a massive mountain range across the length of Sumatra facing the monsoon winds, strong winds may develop through mountain depressions. The word "Sumatra" means storm (angin ribut) in the local dialect. A closer look at the rainfall distribution and the wind systems on Sumatra reveal great variations in both intensity of rainfall and period of occurrence of rainy seasons. This is mainly due to its geographic location and the physiography of the island.

Near the equator air pressure is low and varied little over great distances. As a consequences surface sea winds to be light and evaporated water to rise almost vertically. When the humid air cools off, the air become saturated and heavy rainfall occurs. The energy released forces the air to move to higher altitudes before it finally spreads in northern and southern direction. Then it hits the jet streams before is forced to descend. When it reaches the surface of the earth it is warmed up and blown back over the oceans towards the The Species 33

equator. This is the classic explanatory model for the genesis of trade wind illustrated by Oldeman et al. (1979).

Due to its geographic position and its physiographic characteristics the climate of Sumatra, in particular the prevailing winds and the rainfall distribution show great variations. Temperature on the other hand depends primarily on the elevation above sea level, but monthly variations are relatively small. Although air pressure shows only small annual fluctuations it is determining factor for wind direction and wind force. Barisan mountains represent a considerable obstacle for the monsoon that make differences on the amount of rainfall in east coast dan west coast of Sumatra. The insular and equatorial character of Sumatra, the convection phenomena, local storms, rising of hot air, altitude, all interact in determining regional rainfall patterns.

The rainfall in Sumatra may vary over short distances from nearly 6000 mm per year on the west slope of the Bukit Barisan to less than1500 mm in secluded valleys on the east side of this mountain chain. Rainfall distribution varies from the typical Java type with pronounced dry seasons around July in South Sumatra to an equal distribution with two peaks of rainover large areas in Riau and Jambi (equatorial type), while in North Sumatra the Asiatic type prevails with pronounced dry seasons around February. Most parts of Sumatra receive annual rainfall more than 2500 mm per year. Because of its geographical locationand the presence of the mountain range over the whole length of the island, Oldeman et al. (1979) divided The Species 34

Sumatra in a number of zones with typical rainfallcharacteristics.

Rainfall is abundant throughout the year along the entire length of the Westcoast. Because the close vicinity of the mountains to the coast sea winds govern the wind direction and monsoonal effects are minimal except in thenorthern parts, where February is much drier and in the southern part whereJune is the driest month.

However, the Bukit Barisan presents an obstacle to the passage of the monsoons. The Bukit Barisan encloses a number of highland plains particularly in North and West Sumatra. These plains are East of the watershed and often shelteredon its eastern side by a secondary mountain range. Sea winds can penetrate these sheltered locations only through depressions.Therefore rainfall is strongly reduced compared to the western slopes. Since the plains arealso sheltered on the East side, the monsoon influence is small.

In contrast to the western slopesof the Bukit Barisan, the eastern slopes are not directly along the East coastexcept in North Sumatra. Here extremely heavy rainfall is recorded throughout the year. However, Southern Medan rainfall intensity along theeastern slopes gradually diminishes mainly because of the sheltering effectof the Malaysian peninsula and the narrowing of Strait Malaca.

The Bukit Barisan blocks the circulation of humid air from the Indian Ocean. The Malaysian peninsula serves as abarrier for humid air from the China Sea along the The Species 35

Northeast coast. As aresult the entire East coast and particularly the Northeast coast receives considerably less rainfall throughout the year compared to the West coast. Because of the absence of a mountain-valley wind system the dry season is however not pronounced as along the North coast of Java.

Mean temperatures in lowland areas generally vary little throughout the year in Sumatra, ranging from minima of 22oC to maxima of 34oC (Laumonier 1997). In highland areas the temperatures are correspondingly cooler with frosts occurring above 2500 m asl. Most highland areas experience marked diurnal variation in temperature, with cool nights and relatively warm days.

Vegetation Sumatra has a wide variety of vegetation types which are described in detail by Laumonier (1997). This variety is a result of several (often inter-related) factors, including the diversity of soil types, altitude, volcanic activity and climatic effects. Nearly all of the lowland forests of Sumatra have now been cleared or disturbed to some extent (Laumonier 1997). Before this, the lowland forests mainly comprised tropical lowland evergreen rain forests and peat swamp forests, which are dominated by trees of the dipterocarp family. Lowland health forests are not common on Sumatra itself, but occur sporadically along the east coast (Whitmore 1984). They are extensive on the islands of Bangka and Belitung in Riau province. Low hills, up to 1000 m asl, generally support lowland evergreen rain forests, but from 1000-1500 m they are usually replaced by The Species 36

lower montane forests, in which oaks and laurels are more prominent. Above 2000 m, the lower montane forests are replaced by upper montane formations, in which the Coniferae, Myrtaceae and Ericaceae are well represented. The presence and structure of the summit vegetation of Sumatran volcanoes is often determined by the amount of time which has elapsed since the last eruption.

Laumonier (1997) notes that in 1985, roughly 33% of Sumatra was covered with undisturbed forest, whereas 29% comprised secondary forest types. Almost 24% was under agriculture and about 13% had been logged.

Laumonier (1997) reported that there were approximately 33.38% coverage of the major types of undisturbed vegetation on the island of Sumatra ie: Lowlands (7.25%), low hills (6.61%), lower montane (6.71%), upper montane (1.32%), limestone (0.91), pine forest (0.32%), fresh water swamps (2.05%), peat swamps (7.70%), and mangroves (0.47%). Of the 12 Sumatran Mangifera species described, one occurs only in hills of North Sumatra, whereas eleven others occur in lowland area up to 1800 m asl.

Rainfall volume Sumatra is always considered an equatorial island with abundant rain evenly distributed throughout the year, with fairly uniform mean annual temperatures. Laumonier (1997) divided Sumatra in to five bioclimates and rainfall regimes, ie:

The Species 37

Subhumid bioclimates (1000

Humid bioclimates (1500

the interior of Banda Aceh to Palembang, and from the depression west of Jambi (Tebo Kwantan) to the plains east of Lampung. To these must be added those regions on the periphery of the drier areas of the Barisan mentioned above. The average monthly rainfall may fall below 60 mm, for a month at most. The mean annual number of rainy days varies between 120-150.

Superhumid (2500 < P < 3000mm/year). Superhumid bioclimates cover 29% of the surface area and cover the hill region east of the Barisan range, the swampy lowlands to the south of Bengkulu, and the Ketaun river region. The average number of rainy days varies between 140-170 per year. Most of the weather stations at medium altitudes show this type of bioclimate (Merapi, Kayu-Aro, Dolok Sinabung).

Hyperhumid bioclimates (P> 3000mm/year). The rest of the island is under hyperhumid bioclimates. The eastern piedmonts and most of the western piedmonts have a rainfall exceeding 3000 mm/year. The average annual number of rainy days varies between 180-220.

Latitudinal and altitudinal gradients. Rainfall does not necessarily increase with altitude. The wettest regions are usually the piedmonts. Although there are few rainfall records for very high altitudes, it is clear however the summits of Sumatra are far from dry, as can be ascertained when climbing volcanoes. Singgalang at 2877 m has a rainfall of 3500 mm/year. Overall, mean annual rainfall at low altitudes increases from the south to the north, the The Species 39

length of the west coast (Bengkulu 3300 mm/year, Padang 4700 mm/year, Sibolga 4900 mm/year). In the east, there is also an increase in the rainfall from the south to the north as far as the Malacca Straits. Rainfall then decreases in the north-east where the land is protected from the west and south-west winds by the Barisan Range and from the north, north-east wind by the Malay peninsula.

Altitude Altitudinally, 1000 m is generally recognised as an important,(albeit approximate) boundary which differentiates a particular climatic zone, for which absolute temperature minima become real limitations for certain animal and plant organisms. It is obvious these boundaries may vary, which inevitably, posses the question of their existence.

The present phytocrenological studies in Sumatra have allowed to work out an altitudinal classification for the island. An analysis based on the presence-absence of 1500 liana and tree species in our floristic releves has been undertaken (Laumonier 1990). Laumonier (1997) proposed altitudinal zonation for Sumatra vegetation (Table 1). Altitudinal variations observed in the distributionof flora and vegetation types should also be studied in detail.

The Species 40

Table 1. Altitudinal zonation of the Sumatran vegetation (Laumomier 1997) 0 – 150 m Lowlands 150 – 400 m Low elevation hills 400/500 – 800/900 m Medium elevation hills 800/900 – 1300/1400 m Submontane 1300/1400 – 1800/1900 m Lower montane 1800/1900 – 2400/2500 m Montane >2500 m Tropical uppermontane and subalpine

CHAPTER 3 The Species The Species 41

Mangifera Exploration in Sumatra arly eploration of Mangifera in Sumatra has been conducted by Fitmawati since 2012. Fitmawati et al. (2013) reported that there were 929 collection numbers of 10 species found in Central Sumatra E (consists of West Sumatra, Riau and Jambi Provinces). Fitmawati et al. (2015) reported that there were 697 collection numbers of 7 species found in Eastern Sumatra (consists of Aceh and North Sumatra Provinces). Fitmawati (2016) reported that there were 930 collection numbers of 10 species found in Southern Sumatra (consists of Bengkulu, South Sumatra and Lampung Provinces). With total 2,556 collection numbers of 12 species (with many cultivars of each species) of Mangifera found in Sumatra Greater Island. However, the number of species are reduced significantly differ greatlywith the previous record reported by Kostermans and Bompard (1993) that there were approximately 34 species foundFig. 14 in. Dr. Sumatra Fit during Island exploration before (Table 2). However, habitat loss of Mangifera due to the large scale opening of land clearing and conversion of forest for things such as human The Species 42

settlements and palm-oil plantation was not something suprising. Since 2012, Fitmawati et al. has conducted Mangifera exploration on mainland of Sumatera covering eight provinces. many types of Mangifera scattered in previous documentation not found in this exploration. The results of this exploration do not get many types of Mangifera ever recorded in the previous documentation. However, the types of mango cultivation are quite common. Of the 34 species of Mangifera ever recorded found in Sumatra, only about 12 species are still to be found in Sumatra (Table 3). Mangifera species founded in each of provinces in Sumatra based on geomorphology are presented in (Table 4). Table 2. Record of Mangifera in Sumatra (Kostermans and Bompard 1993) Locality Species 1 2 3 4 5 6 7 8 9 10 M. gedebe Miq. - - √ - - - √ √ - - M. pentandra Hooker f. ------√ M. parvifolia Boerl. & Koorders - √ √ √ - - - √ - - M. paludosa Koterm., spec., nov. - √ √ - - √ √ - M. griffithii Hooker f. - √ √ √ √ √ - √ √ √ M. gracilipes Hooker f. - - √ ------M. microphylla Griff. ex Hooker f ------√ - - - M. similis Bl. ------√ - - M. torquenda Kosterm. - √ - - - √ - √ - - M. applanata Kosterm., spec., nov. - - - √ ------M. longipetiolata King - √ - √ - - - √ - - M. quadrifida Jack √ √ - √ - - - √ √ - M. magnifica Kochummen - √ √ √ - - - √ √ - M. swintonioides Kosterm., spec., nov. ------√ - M. dewildei Kosterm., spec., nov. - √ ------M. indica L. - √ - - - - √ √ - - M. rubropetala Kosterm., spec., nov. - - - √ ------M. rigida Bl. √ √ ------M. oblongifolia Hooker f. - √ - √ - - - √ - - M. rufocostata Kosterm., spec., nov. - - - √ - - - √ - - M. laurina Bl. - √ - √ - - - √ - - The Species 43

M. lalijiwa Kosterm., spec., nov. - - √ ------M. orophila Kosterm., spec., nov. √ ------M. lagenifera Griff. - √ ------M. decandra Ding Hou - √ √ - √ - √ - - M. caesia Jack - √ √ - √ - √ √ - - M. kemanga Bl. - - - √ - - - √ - - M. macrocarpa Bl. - √ - - - √ - - √ M. foetida Lour. - √ - √ √ - √ - - - M. leschenaultii Marchand √ √ - √ - √ √ - - - M. odorata Griff. - √ - - - - √ √ - - M. bompardii Kosterm., spec., nov. √ √ ------M. bullata Kosterm., spec., nov. - √ - √ ------M. subsessifolia Kosterm., spec., nov. - - - √ √ - - - - - 1:Aceh, 2:North Sumatra, 3:Riau, 4:West Sumatra, 5:Jambi, 6:Bengkulu, 7:Lampung, 8:South Sumatra, 9:Bangka Belitung, 10:Riau Archipelago

Table 3. Record of Mangifera by Fitmawati et al. (2012-2016) Distribution/Locality Species name A NS R WS J B SS L M. kemanga Bl. - - √ - √ √ √ √ M. foetida Lour. √ √ √ √ √ √ √ √ M. odorata Griff. √ √ √ √ √ - - - M. torquenda Kosterm. - - √ √ √ - - - M. quadrifida Jack. √ √ √ √ √ √ √ √ M. sumatrana Miq. - - √ √ √ √ √ √ M. indica L. √ √ √ √ √ √ √ √ M. laurina Bl. - - √ √ √ √ √ √ M. zeylanica (Bl.) Hooker f. - - √ √ √ - - - M. magnifica Kochummen. - - √ - - - - - M. griffithii Hooker f. - √ ------M. lalijiwa Kosterm. - - √ - - √ √ √ Anacardium occidentale L. - - √ √ √ - - - macrophylla Griff. - - √ √ √ - - -

The Species 44

Table 4. Collection of Mangifera Germplasm based on Geomorphology in Sumatra since 2012 Region Latitude Longitude Geomorphology Collections Aceh 2o – 6o LU 95o – 98o BT Lowlands M. indica, M. Zeylanica Northern Mountains M. laurina, M. Odorata Central Mountains, or GayoM. laurina, M. foetida, M. odorata South Mountains, or Alas M. foetida, M indica, M. quadrifida North Sumatra 1o – 4o LU 98o – 100o BT Lowlands M. indica, M. Laurina Fold region Mangifera sp., M. indica, M. foetida, M. odorata Mountains M. laurina, Mangifera sp. 2 West Sumatra 0o 54‘ – 3o 30‘ LU 98o 36‘ – 101o 53‘ BT Volcanic Mountains M. laurina, M. Foetida Tertiary Hilly M. odorata, M. laurina, Lowland M. indica, M. laurina, M. zeylanica Riau 01o 05‘ – 02o 50‘ LU 100o – 105o 05‘ BT Swamps - Lowlands M. indica, M. laurina, M. zeylanica Hills M. kemanga, M. quadrifida, Mangifera sp. 1, M. torquenda, M. foetida Jambi 0o 45‘ LU – 2o 45‘ LS 101o 10‘ – 104o 55‘ BT Swamps - Lowlands M. indica, M. laurina, M. zeylanica, M. kemanga, M. quadrifida Volcanics M. foetida, M. Laurina Bengkulu 2o 16‘ – 3o 31‘ LS 101o 10‘ – 103o 41‘ BT Lowland (West coast) M. indica, M. laurina, M. zeylanica Lowland (East coast) M. foetida, M. laurina, M. odorata Jalur Semangko M. foetida, M. indica, M. quadrifida Bukit Barisan East side M. indica, M. laurina, M. odorata The Species 45

South Sumatra 1o – 4o LS 102o – 106‘ BT Swamps - Folds inland M. foetida, M. Laurina Volcanics M. foetida, M. Laurina Outside the crease M. indica, M. Laurina Lampung 50o 20‘ – 50o 30‘ LS 105o 28‘ – 105o 37‘ BT Lowlands M. indica, M. laurina, M. zeylanica Swamps - Volcanics M. foetida, M. Laurina Semangko breaks M. odorata Mountains M. foetida, M. Laurina

The Species 46

Distribution Pattern Distribution of plant species can be affected by rocks, topography and climate. Based on topographic maps, Sumatra Island is generally divided into five vertical regions, entail hilly in western part, montane in central west part, curly-wavy in middle east part, flat-wavy in east part and flat in the easternmost part of Sumatra. The majority of Mangifera species occur as a rule as scattered individuals in tropical lowland rainforests on well-drained soils. The species mostly distributed below 300 m, but can occur up to c.1000 m above sea level, on well-drained soils (44 species), in periodically inundated areas (10 species) and in certain types of swamp forest (i.e. M. gedebe, M. griffithii and M. parvifolia). 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 Center, for example M. austro-yunnanensis and M. persiciformis in China, M sylvatica Roxb. in Sikkim and southern China, at altitudes of 600–1900 m above sea level; apparently wild M. indica can also be found outside the tropics.

The majority of wild mangoes occur as scattered individuals at very low densities in lowland forests on well- drained soils. Some of these are very rare; there are normally one to three trees above 40 cm in diameter/10 ha. Only a few species (M. gedebe, M. griffithii and M. parvifolia) are gregarious in certain types of swamp forest. Most species are The Species 47 evergreen although a few are deciduous in the rainforests following a dry period, and stand bare for a short time before flushes of new leaves appear. A deciduous habit that is not linked to a seasonal climate also occurs in other genera of Anacardiaceae (Ding Hou, 1978b).

Generally the type of Mangifera widely spread in the area with an altitude range between 5-1418 m asl (Fitmawati et al. 2015). In nature, Mangifera divided into three groups: wild group including M. foetida, M. kemanga, M. torquenda, M. quadrifida, M. griffithii, Mangifera sp., semi-cultivated group entailingM. sumatrana, M. odorata, M. laurina, and cultivated group involving M. indica, M. lalijiwa and d M. zeylanica. These groups have different distribution.

The wild depend on the existence of natural dispersal agent and ecological compatibility hence these species generally found in undisturbed forest. Forest conversion significantly decrease the habitat extent of wild Mangifera in Sumatra forest.

The semi-cultivated are considered minor fruit and generally located in the edge of forest, These species are vulnerable to land clearing, plantation and land conversion. While the cultivated commonly found in the yard or plantation area. These species are depend on anthropocentric activity. The cultivated species are widely distributed near the residential area, vice versa for the wild species because of shortcoming in economy.

The Species 48

Altitude determined the existence of plants. Vegetation are affected by climate change, altitude and temperature which are correlated to each other (Ewusie 1990). Anwar et al. (1987) stated temperature decreaserates approximately 0,60C in each 100 m increasing altitude. However it can be different depend on area, season, time and humidity. These factor may affect the difference distribution of each of Mangifera species.

Fig. 15. Map distribution of Mangifera in Sumatra based on MaxEnt model The Species 49

Analysis of global climate suitability map of Mangiferain Sumatra generated using MaxEnt model is depicted in Fig.12. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training. The red color indicates areas with a high probability of occurrence, the and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for Mangifera. In fact, this world wide climate suitability map can be used in the countries that lack of precise coordinates of Mangifera occurrences and generate a preliminary climate suitability map of Mangifera because it may be too late to wait for the precise coordinates of Mangifera occurrences to generate a perfect climate suitability map.

Key to the Species 1. Flower disc narrow, thin, short, stipe-like ...... 2 Flower disc broad, thick, cushion-like ...... 3

2. Panicles blood-red, brick-red or dark red ...... 4 Panicles white purplish or white-violet ..... 1. M. kemanga

3. Flowers tetramerous ...... 5 Flowers pentamerous ...... 8

4. Perianthium pink or red, reflexed ...... 2. M. foetida Perianthium white, curved-reflexed...... 3. M. odorata

5. Stone completely bald, glossy white ...... 4. M. magnifica Stone with thin or short fibres, or hard or leathery ...... 6

6. Panicles axillary ...... 5. M. griffithii The Species 50

Panicles pseudo-terminal ...... 7

7. Anther purple blackish ...... 6. M. torquenda Anther black ...... 7. M. quadrifida

8. Panicles glomerulate ...... 9 Panicles non glomerulate ...... 10

9. Fruits green, ovate-oblong ...... 8. M. indica Fruits yellow orange, cordate ...... 9. M. zeylanica

10. Panicles conical ...... 10. M. laurina Panicles pyramidal ...... 11

11. Crown shape semi-circular ...... 11. M. sumatrana Crown shape spherical ...... 12. M. lalijiwa

The Species 51

1 Mangifera foetida Lour. Trees: Tree, 20 – 40 m in tall, 50 – 90 cm in diameter, growth habit spreading, bark grey to whitish grey, crown broadly pyramidal. The shoot brownish red. Leaves: dark green, scattered, semi- drooping in branch, stiffly coriaceous, crackling when folded, elliptic-oblong with blunt or emarginate apex, base acute; both surfaces very smooth, midrib 19-20 cm in length, prominent, nerves 19-21 pairs, prominent, areola reticulation dense, three branches. Petiole 3.5-4 cm in length, 0.3-0.4 cm in diameter. Panicles: pseudo- terminal, horizontal, conical, dark red, non glomerulate, no hairy in primary and secondary branch, flower densityintermediate. Flowers 5, dark pink with light purple tinge, after anthesis dark yellow with brick red. Bracts reddish green, 5, broad triangular, apex acuminate, base deepenly curved and no hairy in dorsal and ventral. Sepals 5, dark red, broad ovate, apex acute, and no hairy in dorsal and ventral. Petals 5, dark pink, reflexed outwards, oblong, apex blunt, no hairy, ridges 3-4. Disc narrow, reduced or absent. Stamen fertile 2-3, staminodes 3-5, filaments adnate to the base, anther blackish purple. Ovary rather round, lateral-frontal, stylus slightly to the side and curved. Fruits: yellowish green, oblong, strong fragrant, with many dirty spots and dots, apex blunt, beak perceptible, sinus shallow, slope of fruit central The Species 52 shoulder rising and then rounded, base flat fruit stalk insertion oblique, shallow, neck slightly prominent, fruit skin rough, waxy, and dense lenticels density. Pulp orangish yellow, soft, very fibrous, sweet. Seed oblong, texture coarse and adherence strong. Monoembriony.

Leaf Anatomy: Anomocytic stomata type. Stratified epidermis. Stratified palisade mesophyll. Upper midrib of M. foetida has convex shape and lower midrib also has convex and elongated shape.

Vernacular names. Bacang (General),Manchang (Aceh), Ambachang (Batak,Minangkabau), Membacang (Palembang).

Ecology. A species of the wet, evergreen primary lowland forest. It is adapted to areas with abundant rainfall, evenly distributed over the year (up to 1000-1500 m asl, generally in cultivation).

The Species 53

Fig. 16.a. Anomocytic stomata; b. Cross section of Embacang leaf; c. Midrib of Embacang

Uses. The durability of timber is short, rarely used, pale or yellow, coarse. The irritating sap is used as a lotion for ulcers. It is also used to deepen tattoo scars.

The unripe fruit is seldom used because of the sap, but sometimes is used for rujak or (pickles) or lalab, by soaking the fruit in lime water for a while.

The grated seed is mixed with oncom, sugar, terasi and salt gives the sambel pelok limus. Pelok means mango seed in Java.

The fruit is edible. However, young fruits extrude sap that can cause blisters. Mature fruits have a strong turpentine smell is sweet and tasty. People in Europe may mix it in fruit cocktail. The fruit should be peeled thickly to prevent contact with irritating sap which causes blisters on lips and tongue.

Protease is one of the most important industrial enzymes with a multitude of applications in both food and non-food sectors. Although most commercial proteases are microbial proteases, the potential of non-conventional protease sources, especially plants, should not be overlooked.

M. foetida fruit, known to produce latex with a blistering effect upon contact with human skin, was chosen as a source of protease. The latex is notorious for its skin-blistering effect, The Species 54 suggesting the presence of strong proteolytic activity. Sap of Mangifera foetida Lour (Ahmad 2012) and the pulp also showed proteolytic activity (Silaban 2009; Katzer 2007; Byun 2005).

Fig. 17.M. foetida Lour cv. Embacang Bulat

Notes on Distribution. Sumatra—throughout the island (Fig. 18). Analysis of global climate suitability map of M. foetida in Sumatra generated using MaxEnt model is depicted in Fig.18. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training. The image uses colors to indicate predicted probability that conditions are suitable, with red indicating high probability of suitable conditions for the species, green indicating conditions typical of those where the species is found, and lighter shades of blue indicating low predicted probability of suitable conditions. For M. foetida, it can be The Species 55 seen that suitable conditions are predicted to be highly probable through most of lowland Sumatra and the west coast of Sumatra. This one of the best known and most commonly cultivated mango species with magnificent red flowers.

Fig. 18.Distribution of M. foetida in Sumatra based on MaxEnt model

M. foetida is widely spread almost throughout the mainland of Sumatra also known as cosmopolitan species. It occurs chiefly in primary lowland forest in the wet tropics. Its preferences to lowland, wet and tropical region was make this species as one of the most abundant species in Sumatra. It can be found in area with altitude 0 – 1200 m asl. It has strong adaptation to live in area with abundant rainfall, evenly The Species 56 distributed over the year, though this species is not origin from Sumatra.

M. foetida widely dispersed on soil types of distropepts, tropohemists, tropaquepsts, rendolts, tropudultst and distrandepts. M. foetida is scattered in various types of soils ranging from moist/soft textured soil to fairly dry areas, as a result the existence of this mango grows in many areas of rice fields, dry agricultural land and shrubs area. Fitmawati et al. 2015 reported M. foetidais distributed in dryland farming, shrubs, mixed dryland agriculture, primary dryland forest, secondary dryland forest and open land.

M. foetida has wide range of taste from very sour to very sweet. This may be affected by monoembriony and zygotic embryo. The ancient species of M. foetida could not found any where now a days. It is because this species is minor fruit. The rest of old trees which still exist are in danger. Old trees are potential to take for the timber while the fruit is minor economically.

Because of the minority of this fruit economically and in cultivation, M. foetida are threatened. Based on the research conducted since 2012, it is founded many cultivars with varied fruit shape morphology in Sumatra. For instance, there was Macang kabau with ±1 kg weigh which found in Sijunjung region, Macang lado which became favourite fruit in Muara labuh district because of its small size, soft flavor, sweet taste and orange pulp.Researchers also found an unique shape of fruit of Macang susu from Southern Sumatra which have mamae-like beak of fruit (Fig. 18). The Species 57

Fig. 19. Fruit Sampel Collection of M. foetida in Sumatra from left to right (ESK08, ESK17, ESK25, ESK45, CS1, CS2, CS3, CS4, CS5, SS1) ES=Eastern Sumatra; CS=Central Sumatra; SS=Southern Sumatra

Fig. 20.Fruits of Embacang Kebo from Central Sumatra

Mangifera foetida was showed a closely related to M. odorata. This theory about M. odorata is a hybrid from M. indica and M. foetida stated by Hou 1978 and a further study The Species 58 conducted by Teo et al. (2002) and Yonemori et al. (2002) highly supported Hou 1978 statement. This findings is contrary with Kostermans and Bompard (1993) that said the reticulation of M. odorata was completely different from M. indica and M. foetida and also the flower was not an intermediate of both Mangifera. This research found that the grouping of M. foetida is tend to has close relationship with M. odorata. Therefore, it is believed that M. foetida is one of the ancestor of M. odorata.

The Species 59

Trees: Tree, up to 30 m tall and 80 cm indiameter. Crown spherical, straight bole; bark grey, growth habit spreading, the shoot light red brick. Leaves: scattered, dark green, coriaceous, oblong, apex acuminate, base acute, both surfaces rough; midrib 25-26 cm in length, prominent, nerves 19-23 pairs, prominent below, areola reticulation dense, prominent and three branches; petiole 4-4.5 cm in length, 0.3-0.4 in diameter. Panicles: pseudo-terminal, semi-erect, dark red, non glomerulate, no hairy in primary and secondary branchs, intermediate in flower density. Flowers white with red and yellow tinge, 5-merous, dark red with brown tinge after anthesis. Sepals greenish red, 5, narrow ovate, hairy in dorsal and ventral. Petals 5, white, curved-reflexed outwards, elliptic, apex acute, no hairy. Ridges 4-5. Fertile stamen one, anther purplish red, staminodes 4-5, filament adnate to base. Ovary rather round, lateral-frontal. Stylus slightly to the side and curved. Fruits: yellowish green, ovate-oblong, strong fragrant, apex blunt, beak perceptible, sinus absent, slope of fruit central shoulder ending in a long curve, base acuminate-rounded, fruit stalk insertion oblique, neck prominence absent, fruit skin rough, waxy, medium of lenticels density. Pulp light orange, soft, adherence strong, fibres high density, juicy and sweet. Seed reniform, fibres in stone coarse, adherence strong, , veins on stone level with surface. Polyembriony. Leaf anatomy. Anomocytic stomata type. Simple epidermis. Stratified The Species 60 palisade mesophyll. Upper midrib of M. odoratahas convex shape while lower midrib has concave shape.

Vernacular names.Kweni, kwini, kuweni (general);Mbem, keweni (S. Sumatra)

Ecology. Up to 1000 m alt (often 800 m) in ever wet tropical rainforest

Fig. 21. M. odorata

The Species 61

Fig. 22.a. Anomocytic stomata; b. Cross section of Kweni leaf; c. Midrib of Kweni

Uses. Fruit is edible, but has to be peeled thickly because of the irritating sap, the young fruits are used for pickling and are soaked for some time in lime water. A flour is made from the seeds which in Java is used for making the delicacy jenang or dodol.

The species is cultivated everywhere in South-east Asia. Its merits as a table fruit are debatable, for some the pungent odour is pleasant, for others offensive. The consistency of the fruit makes it worthwhile to experiment for improving it.

Medicinal uses. A leaf poultice is applied externally against hysteria, epilepsy. Also the bark is used in the form of a The Species 62 compact cosmetic-like mixture against hysterio-epilepsy in Malaya.

The wood similar to that of M. indica and of poor quality.

In Malay region in Sumatra especially in Riau province, M. odorata became the main ingredients to make Es Laksamana Mengamuk. Mengamuk means rampage. It is because the sour taste from kweni (M. odorata) makes the tounge rampage .

Fig. 23. Es Laksamana Mengamuk from Riau

Notes on Distribution. Sumatra—throughout the island (Fig. 24). Analysis of global climate suitability map of M. odorata in Sumatra generated using MaxEnt model is depicted in Fig. 24. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training. The image uses colors to indicate predicted The Species 63 probability that conditions are suitable, with red indicating high probability of suitable conditions for the species, green indicating conditions typical of those where the species is found, and lighter shades of blue indicating low predicted probability of suitable conditions. For M. odorata,it can be seen clearly that this Mangiferasuitable conditions and predicted to be highly probable through most of lowland Sumatra and the west coast of Sumatra.

Fig. 24. Distribution of M. odorata in Sumatra based on MaxEnt model

M. odorata is a specific species for Sumatra. It was distributed from 0 - 800 meters asl altitude (Fitmawati et al. 2015). M. odorata thrives below 1000 m in tropical areas with a fairly heavy rainfall that is equally distributed throughout the year, although it grows even with a moderate rainfall (1200 The Species 64 mm) provided there are no prolonged dry periods. If M. odoratagrown on the 800 meters asl altitude, it could decrease the ability of fruiting.

M. odorata is spread on the soil type oftropudalts, distransdepts, paleudults, distropepts, paleusults, tropaquepts and tropudults. It can be found along the east and west coasts of Sumatra island. M. odoratais distributed in open agricultural land, secondary dryland forest, shrubs, mixed dryland agriculture, primary dryland forest, and settlements (Fitmawati et al. 2015). It was not much varied because of the crossbreeding. This species were potentially developed for mango prodcution in Sumatra. This species are adaptable and has worthy value in agriculture. The Species 65

Trees: Tree, 60-80 cm in tall, 90-120 cm in diameter, bark brownish white, growth habit spreading, the shoot brownish red, crown semi-circular. Leaves: scattered, semi-erect, dark green, chartaceous, oblong- lanceolate, apex acuminate, base acute, both surfaces smooth; midrib 18.5-19 cm in length, prominent, nerves 29-31 pairs, prominent below, areola reticulation intermediate, slightly prominent. Petiole 3.5-4 cm in length, 0.2-0.35 cm in diameter, base slightly curved, slightly submerged. Panicles: terminal, semi-erect, greenish yellow, conical, non glomerulate, no hairy in primary and secondary branches, medium flower density. Flowers white with light yellow tinge, 5-merous, after anthesis pale yellow with brownish tinge; bracts light green, 5, narrow ovate, apex acuminate, base flat and smooth hairy high in dorsal and ventral. Sepals 5, broad triangular, apex acute, smooth hairy sparse in dorsal and ventral. Petals 5, reflexed outwards, oblong, apex blunt, no hairy; ridges 5, thicken. Disc swollen, broader than ovary. Stamen fertile one, staminodes 4-5, filaments adnate to the base, anther red. Ovary rather round, frontal, stylus slightly to the side and slightly curved. Fruits: orange yellow, oval-oblong, apex pointed, beak pointed, sinus slight, slope of fruit ventral shoulder ending in a long curve, base acuminate rounded, fruit stalk insertion vertical, shallow, neck prominence slightly prominent, fruit skin smooth, waxy and sparse lenticel density; pulp orange yellow, soft, adherence The Species 66 weak, fibres low, juicy and sweet. Stones oblong, soft fibres, adherence medium, veins on stone depressed with two branches. Polyembriony.Leaf anatomy. Anomocytic stomata type. Simple epidermis. Stratified palisade mesophyll. Upper midrib of M. laurina has convex shape while lower midrib has concave shape.

Vernacular names. Mangga parih, Mangga aer; mempelam, polam, pelam, pogang; mangga apung (South Sumatra)

Ecology. Lowland wet, evergreen tropical forest, up to 1700 m alt.

Uses. Semi-fluid pulp at maturity fits for making juices, but the taste is too bland.

Fig. 25.Flowers of M. laurina

The Species 67

Fig. 26.a. Anomocytic stomata; b. Cross section of M. laurina leaf; c. Midrib of M. laurina

Notes on Distribution. Sumatra—throughout the island (Fig. 27). Analysis of global climate suitability map of M. laurinain Sumatra generated using MaxEnt model is depicted in Figure 27. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. laurina. In fact, this worldwide climate suitability map can be used in the countries that lack of precise coordinates of M. Laurina occurrences and generate a preliminary climate suitability map of M. laurina because it may be too late to wait for the precise coordinates of The Species 68

M. laurina occurrences to generate a perfect climate suitability map.

Fig. 27.Distribution of M. laurina in Sumatra based on MaxEnt model

M. laurina is believed to originate from south-east Asia. Based on the research conducted in East Sumatra, one sub- species of M. laurina known as mangga udang which is the introduced species in Toba lake. Itbecame the glory mango from this area. M. laurina was found from 0 – 1700 m asl. The sub-species mangga udang was distributed from the lowland area to area with 1900 m asl. Both of this M. laurina wild and cultivated still can be found in the forest of North Sumatra. It was proved that M. laurina is the original species from Indonesia.

The Species 69

M. laurina is widely distributed in soil types of fluvaquents, distropepts, tropudults, paleodults, distrandepts, tropochemists, tropaquepts, tropodultst, paleudultst, and tropaquepsts. M. laurina is also cosmopolitan plant because it can be spread on various soil types and various natural conditions.M. laurinais spread in shrubs, primary dryland forests, secondary dryland forests, industrial plantations, plantations, settlements, dryland farming, mixed dryland agriculture and transmigration areas (Fitmawati et al. 2015).

Fig. 28.Some of the fruit collection of M. laurina from Sumatra

In non-flowering herbarium, M. laurina has similar leaves with M. sumatrana, M. indica and M. lalijiwa. In flower, M. indica can be easily seperated by its flowers in glomerules and being densely pubescent (in M laurina a lax, non-glomerulate, almost glabrous inflorescence). M. indica The Species 70 and M. lalijiwa both havee larger flowers with broader and thicker petals.

The fruit of M. laurina is small, mango shaped, yellow green, the pulp is almost fulid and sukced out (hence the local name: watery mango; mangga aer) and not firm as in M. indica. M. lalijiwa often remains very dark green at maturity with thin, orange mesocarp and an enormous stone, the inflorescence are lax and glabrous.

It is grown, as it is well adapted to ever wet climate. The inflorescence show no sign of attack by anthracnose whereas M. indica in the same area is heavily attacked. It can stand a very wet soil with common flooding periods.

The relationship between M .laurina Bl. and M. sumatrana Miq. Is remain debatable. Mangifera sumatrana Miq., an unique species found in Sumatra was treated as synonym of M. laurina Bl. based on morphological characters in the latest classification by Kostermans and Bompard (1993). Prior them, this species was treated as synonym of M. longipes Griff. by Mukherji and Ding Hou. Phylogenetic analysis based on morphological characters by Fitmawati et al. 2013 also showed M. sumatrana Miq. has a close relationship with M. indica L. and M. laurina Bl. However, based on molecular analysis using ITS sequence by Fitmawati et al. 2016a was obtained the results M. sumatrana Miq. did not form a clade with both of M. indica L. and M. laurina Bl. Corresponding to the tree of MP/ML/NJ/Bayesian analysis using ITS sequence, M. sumatrana Miq. is not synonym of M. laurina Bl.

The Species 71

Trees: Tree, 5-10 cm in tall, 20-60 cm in diameter, bark brownish grey, growth habit erect, the shoot brownish green and crown spherical. Leaves: green, scattered in semi- drooping branch, chartaceous, oblong-lanceolate, apex acuminate, base acute, both surfaces smooth; midrib 15.1- 15.5 cm in length, prominent, nerves 21-24 pairs, prominent below, medium areola reticulation, slightly prominent, two branches; petiole 3.2-3.5 cm in length, 0.28-0.3 cm in diameter. Panicles: terminal, horizontal, reddish green,broadly pyramidal, glomerulate, hairy high and smooth in primary and secondary branches, high flower density. Flowers: pale yellow with light yellow tinge, 5-merous, after anthesis pale yellow with orange-yellow tinge; bracts light green, 5, broad ovate, apex blunt, curved and dense hairy and smooth in dorsal and ventral. Sepals 5, light green, broad ovate, apex acute, hairy in dorsal and ventral. Petals 5, pale yellow, erect-curved outwards, oblong-lanceolate, apex acute, sparse hairy in the base, ridges 5. Disc swollen, broader than ovary with 5-lobed. Stamen fertile one, staminodes 4-5, filaments adnate to the base. Anther purplish. Ovary rather round, frontal. Stylus slightly to the side and slightly curved. Fruits: green, ovate- oblong, apex obtuse, fruit skin smooth, waxy, density of lenticels on fruit skin sparse, beak perceptible, sinus shallow, slope of fruit central shoulder ending in a long curve, fruit stalk insertion vertical, neck prominence absent. Pulp light orange, The Species 72 texture soft, adherence medium, quantity of fibre medium, juicy and sweet. Stones oblong, fibre texture soft, adherence of fibre to stone medium, veins on stone level with surfaces and pattern of stone venation forked. Monoembriony- Polyembriony. Leaf anatomy. Anomocytic stomata type. Simple epidermis. Simple palisade mesophyll. Upper midrib of M. indica has convex shape while lower midrib has concave shape.

Ecology. A tropical tree, requesting a mean shade temperature of about 80o F, It needs abundant moisture in the atmosphere (optimal rainfall 75-200 mm), which must alternate during the flowering and fruit setting period with a rainless period. As all Mangifera species, it grows well in vegetatively, even under poor conditions, but the fruit setting is very irregular; there are completely flowerless years and ―on‖ years (good crop) are only every 3-4 years. A high C/N ration is necessary for flower initiation. Rain, fog and cloudy weather at flowering time reduce fruit setting and increase diseases (anthracnose); heavily rainfall during fruit setting also reduces the crop.

It is more or less indifferent to soil types and can grow well on poor soils, provided that they are well drained, fairly deep and not too alkaline. On shallow soils the trees are stunted. A pH of 5.5-7.5 is preferred. Luxurious growth in rich soils produces less fruit.

Although it is a monsoon tree, it is non-deciduous and grows flush-wise after the terminal buds (which are protected by a small number of bud scales) have undergone a rest period.

The Species 73

Fig. 29.a. Anomocytic stomata; b. Cross section of M. indica leaf; c. Midrib of M. indica

Fig. 30.M. indica from Souther Sumatra The Species 74

Pollination and fruit set. Flowers open early in the morning over a period of 2-3 weeks, usually between 8 and 12 am and complete their anthesis in the afternoon, but bud also begin to open at night and are fully open in the morning. Anther dehiscence takes place after the flowers have opened, the anthers are then bluish due to the pollen. Abnormal meteorological conditions may delay dehiscence and anthesis considerably, both temperature and humidity are effective.

Stigmas are immediately receptive and this may continue up to 3 days after flower dehiscence. Nectar is secreted by the disc. Pollination is essential for fruit settin, even for apomictic embryos. Bagged panicles usually do not set fruit. A proportion (65-85%) of the bisexual flowers remain unpollinated and only 0.1-0.25% reach the mature fruit stage. Flowers are self-compatible.

Pollinators are mainly house flies, but also bats, wasps, butterflies, beetles, ants, thrips and etc., in addition to self pollination. After anthesis the flowers turn pinkish.

Only one of 1000 bisexual flowers sets fruit, a wasteful procedure, quite common in tropical trees. One of the most pressing problems is alternative bearing. Even an Indian cv cropped well in 8 out of 27 years. A high N ratio is required for flower induction as is the production of new growth during an ―off‖ year. Alternative flowering and non-flowering years (off and on years) are a common feature in Mangifera species. Years of non-flowering may extend to 10 years and more.

The Species 75

For acceleration, the flowering is stimulated by smudging (smoky fires, often in special oven); the effective agent is the carbon dioxide, not the heat. Results are also obtained with foliar spraying with chlorethan phosponic acid, where the effecting agent is acetylene and perhaps with potassium nitrate. Pruning may also be effective. It takes 2-5 months from fertilization to mature fruit; fruit drops at all stages. In general, pollination is inadequate.

Cultivation. The quality of soil is less important than the climatic conditions, the amount of rainfall, the time in which it occurs and the presence of the dry season. Rain and even damp cloudy weather during the blooming period may result in partial or complete loss of crop. Moist weather favours the spreas of anthracnose and blossom blight. Some cultivars produce several crops a year, but most flower only once in a season. The ideal region has a rainfall of 75-125 cm, with a very small proportion of spraying with Bordeaux mixture during flowering.

In planting budded young trees in orchards, they should be set at least 9-10.5 m apart. The greatest care should be given in application of fertilizers. Double grafting is perhaps a way to diminish the tree size.

Uses. The plant is used in ceremonies, customs and rituals. It contains no alkaloids. Powdered, it is used as a fumigant against mosquitoes.The roots are used against diarrhoea and leucorrhoea (tannic acid). Root bark is diuretic, astringent, haemostatic and anti-rheumatic in hot baths and hot dressing; it is used in a wash for biennorrhoea and leucorrhoea. The Species 76

The resin is a remedy for aphthae and dysentry. Mized with oil and lime juice, it is useful for cutaneous disease. The bark can used as a dye for silk, colour yellowish shade of brown.

Leaf, bark and fruit inhibit growth of Micrococcus pyogenes, Staphylococcus aureus and Escherichia coli. They are negatives as antibiotics and negative against malaria. Leaves are used in times of shortage as cattle fodder; prolonged feeding causes death. The urine of leaf-fed cows contains a yellow substance (mangiferin), known as Indian Yellow, which is excreted as the magnesium salt of eurianthime acid and used to turn blue syed matting and cloth (indigo) to green. This cruel, barbaric method of obtaining Indian Yellow has been abolished.

Leaves are used against skin diseases, against dysentry. Smoke of the leaves causes dermatitis. They are also used for asthma and coughs. Leaves and twigs contain anacardol and hydrocyanic acid. Ashes are used in China for burns and scalds. In China, the leaves are made into a tea and also used as a bath to treat fever and cold. Charred and pulverized leaves are used as a plaster to remove warts and also act as a styptic.

Notes on Distribution. Sumatra—throughout the island (Fig. 31). Analysis of global climate suitability map of M. indica in Sumatra generated using MaxEnt model is depicted in Figure 31. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the The Species 77 blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. indica. In fact, this worldwide climate suitability map can be used in the countries that lack precise coordinates of M. indica occurrences and generate a preliminary climate suitability map of M. indica because it may be too late to wait for the precise coordinates of M. indica occurrences to generate a perfect climate suitability map.

Fig. 31. Distribution of M. indica in Sumatra based on MaxEnt model

M. indica is one of the cultivation mango type which success commercialized in the whole world. No body in the world knows this kind of mango. The biggest project for mango The Species 78 breeding is come from this species. It grows up to an altitude of 1200 m, but the fruit setting occurs only up to 900 m.

Fig. 32.M. indica collection 26 from Eastern Sumatra

Fig. 33.M. indica collection 28 from Eastern Sumatra

Fig. 34.Several M. indica collections from Central Sumatra The Species 79

Trees: Tree, 10-15 m in tall and 50-60 cm in diameter, growth habit erect, bark brownish white, the shoot brownish red, and crown semi-circular. Leaves: dark green, scattered, horizontal in branch, chartaceous, ovate-oblong, apex acuminate, base acute, both surfaces smooth, midrib 21-23 cm in length, prominent, nerves 19-22 pairs, areola reticulation medium, slightly prominent, two branches. Petiole 1.9-2.4 cm in length, 0.19-0.23 in diameter. Panicles: pseudo-terminal, horizontal, dark pink, pyramidal, glomerulate, sparse hairy and smooth in primary and secondary branches, high flower density. Flowers pale yellow with light yellow tinge, 5-merous, after anthesis pale yellow with orangish yellow tinge. Bract yellowish green, 5, broadly triangular, acuminate, even and hairy, both dorsal and ventral medium hairy and smooth. Sepals pale yellow, 5, narrowly ovate, acute and hairy and smooth. Petals pale yellow, 5, curved outwards, elliptic, apex blunt, no hairy, ridge 3. Disc swollen, broader than ovary. Stamen fertile one, staminodes 4-5, filaments adnate to the base. Ovary rather round, lateral-frontal. Stylus slightly to the side and curved. Fruits: orange-yellow, oval-oblong, apex flat, skin surface texture smooth, waxy, density of lenticels on fruit skin sparse, beak pointed, sinus absent, slope of fruit central shoulder ending in a long curve, fruit stalk insertion vertical, neck prominence slightly prominent. Pulp orannge-yellow, texture soft, adherence weak, quantity of fibre low, juicy and The Species 80 sweet. Stones oblong, fibre texture soft, adherence of fibre to stone medium, veins on stone depressed and pattern of stone venation forked. Polyembriony.Leaf anatomy. Anomocytic stomata type. Simple epidermis. Stratified palisade mesophyll. Upper midrib of M. zeylanica has convex shape while lower midrib has concave shape.

Vernacular names. Mangga apel, mangga kelapa (Sumatra)

Ecology. Both in the wet and dry zones (but in water rich spots), very scattered, up to 800 m alt. M. zeylanica is one of the cultivation mango type which success commercialized in the whole world.

Fig. 35.a. Anomocytic stomata; b. Cross section of M. zeylanica leaf; c. Midrib of M. zeylanica

The Species 81

Uses. The fruit has an excellent taste. The fruit is fully ripe when it is dropped. The tree is not cultivated, but could be used to improve varieties of M. indica. It was tried out as rootstock. It also has been successfully crossed with M. odorata.

Fig. 36.M. zeylanica cv. kelapa

Fig. 37.M. zeylanica cv. red apple The Species 82

Fig. 38. Fruits of M. zeylanica from Central Sumatra

Notes on Distribution. Sumatra—certain place (Fig. 39).

Analysis of global climate suitability map of M. zeylanica in Sumatra generated using MaxEnt model is depicted in Fig.39.Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. zeylanica. In fact, this worldwide climate suitability map can be used in the countries that lack precise coordinates of M. zeylanica occurrences and generate a preliminary climate The Species 83 suitability map of M. zeylanica because it may be too late to wait for the precise coordinates of M. zeylanica occurrences to generate a perfect climate suitability map. M. zeylanica has high probability of occurrence in the west coast of southern Sumatra.

Fig. 39 Distribution of M. zeylanica in Sumatra based on MaxEnt model

The Species 84

The Species 85

Fig. 40. Distribution of rare Mangifera based on land cover

M. griffithii, M. magnifica, M. kemanga, M. sumatrana, M. torquenda, M. quadrifida and M. lalijiwa are very rare found in Sumatera. These species have sparse distribution throughout Sumatra. The land conversion has led to the declines of population of Mangifera Even, M. magnifica was only found in Kampar District, Riau while M. griffithii was only found in Sipirok, North Sumatra. Beside that, because of the minority of the fruit of these rare species economically and in cultivation, they became very difficult to find in Sumatra. Based on existing data, it can be predicted that the existence of Mangifera species is no longer in natural forests but has been widely cultivated in the community, especially in areas such as settlements, rice fields and others.

The Species 86

Trees: Tree, up to 40 m tall and 100-140 cm in diam, growth habit spreading, bark brownish white with cream sap, the shoot brownish yellow and crown semi-circular. Leaves: dark green, scattered, semi- drooping in branch, chartaceous, oblong-ovate, apex acuminate, base acute, both surfaces smooth, midrib 13.7-14.2 in length, above and below midrib prominent, nerves 21-23 pairs, areola reticulation dense, slightly prominent, two branches. Petiole 2.8-3 cm in length, 0.19-0.22 in diameter. Panicles: terminal, semi-erect, yellowish cream, pyramidal, non glomerulate, medium flower density. Flowers pale yellow with light yellow tinge, 5-merous, after anthesis pale yellow with orangish yellow tinge. Bract yellowish green, 5, broadly triangular, acuminate, even and hairy, both dorsal and ventral smooth. Sepals light green, 5, broad ovate, acute and hairy and smooth. Petals pale yellow, 5, curved-reflexed outwards, elliptic, apex blunt, no hairy, ridge 5. Disc swollen, broader than ovary. Stamen fertile one, staminodes 4-5, filaments adnate to the base. Ovary rather round, lateral-frontal. Stylus slightly to the side and curved. Fruits: orange, roundish, apex round, skin surface texture smooth, thickness 0.2-0.3 cm, non-waxy, density of lenticels on fruit skin sparse, beak pointed, sinus shallow, slope of fruit central shoulder rising and then rounded, fruit stalk insertion oblique, neck prominence absent. Pulp orannge-yellow, texture soft, adherence intermediate, quantity of fibre low, juicy and The Species 87 sweet. Stones oblong, fibre texture rough, adherence of fibre to stone weak, veins on stone depressed and pattern of stone venation forked. Monoembryony.Leaf Anatomy: Anomocytic stomata type. Simple epidermis. Simple palisade mesophyll. Upper midrib of M. sumatrana has convex and lower midrib has concave shape.

Fig. 40. a. Anomocytic stomata; b. Cross section of Paoh leaf; c. Midrib of Paoh

M. sumatrana is the specific species which only found in lowland tropical rainforest in Sumatra. The rare species in Sumatra were potentially developed for mango breeding. The wild species of mother land in Sumatra provide unique character to support mango agriculture in Sumatra. Mostly, Sumatran species of Mangifera did not need long time to stimulate flowering stage.

The Species 88

Notes on Distribution. Sumatra—certain places (Fig. 41). Analysis of global climate suitability map of M. sumatranain Sumatra generated using MaxEnt model is depicted in Fig.41. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. sumatrana. In fact, this worldwide climate suitability map can be used in the countries that lack precise coordinates of M. sumatrana occurrences and generate a preliminary climate suitability map of M. sumatrana because it may be too late to wait for the precise coordinates of M. sumatrana occurrences to generate a perfect climate suitability map.

Fig. 41 Distribution of M. sumatranain Sumatra based on MaxEnt model The Species 89

Trees: Tree, up to 40 m tall and 120 cm in diameter; Crown broadly pyramidal, growth habit spreading, bark greyish brown. Leaves: dark green, scattered, horizontal,thickly and rigidly coriaceous, elliptic to ovate-elliptic, apex acute, base acute, both surfaces smooth; midrib 30-33 cm in length, prominent, nerves 19-23 pairs, prominent, 8-9.5 cm in length, areola reticulation sparse, slightly prominent with three branches; petiole 4-5 cm in length, 0.4-0.7 cm in diameter, base curved, submerged, 2.5-3 cm in length and 0.65-0.77 cm in diameter. Big pulvinus. Panicles: lax (freshly), terminal, semi-erect, greenish white, pyramidal, 26.3-28 cm in length, non glomerulate, no hairy in primary and secondary branches. Flowers white with light yellow tinge, prominent, small, fragrant, 4-merous. Sepals 4, yellowish white or with red tinge (fresh), ovate, 3-3.5 mm in length, 1.5-2 mm in width, base blunt-acute. Petals 4, white, curved-reflexed, oblong, 7-7.5 mm in length, 2-2.5 mm in width, base acute, no hairy; ridge 3-4, thicken. Disc thick, large, cushion-like, obscurely lobed, concave above, 1 mm high. Fertile stamen 1, staminodes 3-5, filament adnate to base, anther black. Ovary rounded, lateral; stylus slightly to the side and curved. Fruits: obovoid, yellowish green, 11.42-12.46 cm in length, 7-13.8 cm in diam., 450.6-480 gr in weight, 7.4-8 cm in thickness, apex obtuse, sinus absent, slope of fruit central shoulder rising and then rounded, base acuminate-ovate, fruit stalk insertion oblique, The Species 90 neck prominence slightly prominent, fruit skin rough, waxy and lenticels density sparse; pulp white, soft, adherence weak, fibres low, juicy and sweet. Stones oblong, white, smooth fibres, 9.5-10.5 cm in length, 4.7-5.6 cm in width, 38.9-42 gr in weight, 2.4-3 cm in thickness, adherence weak, veins on stone level with surface. Monoembriony. Leaf Anatomy: Cyclocytic stomata type. Stratified epidermis. Stratified palisade mesophyll. Upper midrib of M. magnifica has concave, very rounded and bigger shape than the others.

Ecology. In low land and up to 1300 m in wet evergreen tropical climate on drained soils.

Fig. 42Habitus of Putaran in Kampar District, Riau

The Species 91

Fig. 43a. Cyclocytic stomata; b. Cross section of Putaran leaf; c. Midrib of Putaran

Uses. The flesh is generally sour. It serves mainly in the preparation of sambel or rujak. The timber is also used and mixed with white meranti.

Distribution. Widely spread in Western Malesia (except Java), not in Celebes or the Moluccas. Sumatra—presently known from Kampar district, Riau.

Notes.This species was established by Kochummen (1983). At first, Researchers thought it was a new species found in Kampar district. It was because the fruit shape different from M. magnifica was described by Kochummen. A confirmation were made with another researcher in Singapore related to similar habitus of M. magnifica and this species (local people The Species 92 called it Putaran). Occasionally cultivated in some areas such as Western Borneo and Riau.M. magnifica known as Putaran was only found in plantation area and the riverside of Kampar district of Riau. Furthermore, approximately ten trees of this species were found in Sumatra and it restricted to Kampar district of Riau. Its sour taste of fruit was used in cooking.

Sapling leaves may be up to 30 cm long with 5-cm long petioles and are more elongate with acute base. The leaves resembles somewhat those of M. foetida, but can at once be distingushed by the smooth reticulation on both surfaces (in M. foetida entirely smooth) and the rounded leaf base.

The flowers are entirely different from those of M. foetida and resemble those of M. quadrifida but there are differences in colur and shape. The fruit is very different from the black, subglobose one of M. quadrifida with many fibres. It resembles more that of M. foetida, except for the pulp.

Next to M. foetida and the sclerophilous variety of M. griffithii, it has the most leathery leaves of all Mangifera species. The fibreless pulp is remarkable.

The Species 93

Trees: Tree, 25-40 m in tall, 100-150 cm in diameter. Crown oblong, bark yellowish brown, growth habit spreading, the shoot above greenish yellow and below dark pink. Leaves: scatteres, horizontal to semi- drooping, green, stiffly coriaceous, spatulate, apex acute, base acute, upper surface smooth, below rough. Midrib 24-26 cm in length, prominent, nerves 19-21 pairs, prominent, areola reticulation intermediate, prominent. Petiole 6-7 cm in length, 0.16-0.2 cm in diameter, base curved, submerged. Small pulvinus in the end of twigs. Panicles: pseudo-terminal, pulvinus, semi-erect, pyramidal, light green, 24-27 cm in length, non glomerulate, no hairy in primary and secondary inflorescence and sparse flower density. Flowers white with light yellow tinge and prominent, 3-4 –merous, after anthesis brownish white, with brownish tinge. Bracts light green, 3-4, narrow triangular, apex acuminate, no hairy. Sepals light yellow, 3-4, broadly ovate, apex blunt, no hairy in dorsal and ventral. Petals 3-4, curved outwards, elliptic, apex acute, no hairy, ridge 4-5. Disc swollen, broader than ovary. Stamen fertile one, staminodes 3- 4, filaments adnate to the base, anther blackish. Ovary rather round, lateral-frontal, stylus slightly to the side and curved. Fruits: purplish green, cordate, apex acute, beak perceptible, sinus shallow, slope of fruit ventral shoulder ending in a long curve, base slightly flat, fruit stalk insertion oblique, neck prominence absent, fruit skin smooth, waxy, sparse lenticels The Species 94 density. The inner skin purple. Pulp orange yellow, texture soft, adherence intermediate, fibre quantity high, juicy and sweet. Stones oblong, texture coarse, adherence strong, veins on stone level with surface. Endocarp purple. Polyembriony.Leaf anatomy: Anomocytic stomata type. Simple epidermis. Stratified palisade mesophyll. Upper midrib of M. quadrifida has convex while lower midrib has concave shape.

Vernacular names. Asam kumbang (Riau), langong, longung, longuang (West Sumatra), longgun (N. Sumatra)

Ecology. Ever wet lowland tropical rainforest, in Sumatra and the Malay Peninsula mostly the form with acutish leaves and larger fruit, in Borneo mostly spathulate-leaved form with smaller fruit. Also in Sulawesi, the Lesser Sunda Islands and the Moluccas.

Fig. 44 a. Anomocytic stomata; b. Cross section of Asam kumbang leaf; c. Midrib of Asam kumbang

The Species 95

Fig. 45Inflorescence of M. quadrifida

Fig. 46The fruit of M. quadrifida The Species 96

Distribution. Sumatra—restricted to Aceh, Riau, South Sumatra (Fig. 47 ).

Analysis of global climate suitability map of M. quadrifidain Sumatra generated using MaxEnt model is depicted in Figure 47. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. quadrifida. In fact, this worldwide climate suitability map can be used in the countries that lack precise coordinates of M. quadrifidaoccurrences and generate a preliminary climate suitability map of M. quadrifida because it may be too late to wait for the precise coordinates of M. quadrifida occurrences to generate a perfect climate suitability map. The Species 97

Fig. 47 Distribution of M. quadrifida in Sumatra based on MaxEnt model

M. quadrifida still can be found in Eastern Sumatra, Riau and South Sumatra. It has unique characteristic such as black reddish fruit and purple pulp. It similar to other rare species that the timber is attracted for people but the fruit is not, therefore it became one of the threaten factor for the rareness of this species. The Species 98

Trees: Tree, up to 40 m tall and 90-100 cm in diameter, bark reddish brown, growth habit spreading, crown jorong ke atas broadly pyramidal and the shoot light green. Leaves: green, scattered, semi-erect in branch, rigidly coriaceous, ovate, apex acute-acuminate, base acute, above surface rough, below surface smooth, midrib prominent, nerves 20-24 pairs, areola reticulation dense, prominent. Petiole 12-14 cm in length and 0.27-0.3 cm in diameter. Panicles: pseudo-terminal, horizontal, greenish yellow, conical, 25-27 cm in length, glomerulate, hairy and smooth in primary and secondary inflorescence, dense flower density. Flowers white, 4-merous, after anthesis cr eam white. Fruits: yellowish green, ovate-oblong, strong smell, apex round, beak percetible, sinus absent, slope of fruit central shoulder ending in a long curve, base flat, fruit stalk insertion vertical, neck prominence absent, fruit skin texture rough, waxy, density of lenticels on fruit skin dense. Pulp white, texture soft, adherence intermediate, quantity of fibre low, juicy and sweet. Seeds reniform, fibre in stones rough, adherence weak. Polyembriony.Leaf anatomy: Cyclocytic stomata type. Simple epidermis. Stratified palisade mesophyll. Upper midrib of M. torquenda has convex while lower midrib has concave shape.

Vernacular names. Taias, tayas,tais, tajas (Sumatr, Malay) The Species 99

Ecology. Wet, evergreen tropical climate, along rivers, inundated soils; 100 – 800 m alt.

Fig. 48 a. Cyclocytic stomata; b. Cross section of Tayeh leaf; c. Midrib of Tayeh Notes on Distribution. Sumatra—presently known from Riau, W. Sumatra, Jambi, S. Sumatra (Fig. 49)

Analysis of global climate suitability map of M. torquenda in Sumatra generated using MaxEnt model is depicted in Fig.49. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. torquenda. In fact, this worldwide climate suitability map can The Species 100 be used in the countries that lack precise coordinates of M. torquendaoccurrences and generate a preliminary climate suitability map of M. torquenda because it may be too late to wait for the precise coordinates of M. torquenda occurrences to generate a perfect climate suitability map.

Fig. 49 Distribution of M. torquendain Sumatra based on MaxEnt model

Based on the soil type and land altitude, M. torquendra was scattered in lowland areas at an altitude of 6-12 m asl. This type of mango is distributed in soil types of tropudalts, tropohernists, tropudults, distransdepts and paleudults. Based on the slope map, M. torquenda is widely distributed on the ground with a slope of 0-15% and>40%. M. torquenda is the wild species of Mangifera with white flower and rounded fruit The Species 101 characters. This species is very rare found in Sumatra because of the minor fruit. Nowadays M. torquenda is widely dispersed in transmigration areas, mixed dryland farming, open land and plantations. M. kemanga is widespread in mixed dryland farming areas, plantations, primary dryland forests, plantations and open land (Fitmawati et al. 2015). The Species 102

Trees: Deciduous, large tree, up to 30 m tall. Bark rather smooth, grey to pinkish grey, in old trees superficially fissured. Crown rather open with many small subcrowns and long primary branches. Leaves: appearing flush-wise after the tree has been bare for a short period. The leaf buds are very large and surrounded by numerous, large, outside densely, minutely silky puberulous caducous bud scales, leaving large collars of flat scars. Twigs very stout, angular smooth. Leaves crowded at the apices of the twigs, stiffly coriaceius, subsessile, oblanceolate to obovate-oblong or cuneate-oblong, 20-40 cm long, 7-10 cm wide, acute or shortly, abruptly and obtusely acuminate (in saplings long- acuminate), the edges subundulate, gradually narrowed from below the middle to the base, glabrous, no reticulation on both surfaces; above midrib very broad, flattened, prominulous, nerves filiform; below midrib stout, prominent, broad; nerves 20-30 pairs, rather patent, thin, prominulous. Petiole very broad, the leaf margins narrowly decurrent (winged petiole), up to 7.5 mm long but usually absent. Panicles: subterminal, pyramidal, up to 75 cm long, white violet (bluish red in sicco); main peduncle stout, covered by minute white hairs, intermixed with a few longer, brown ones; branches angled, suberect to erect; the flowers glomerulate. Bracteoles broadly ovate to ovate-lanceolate, concave, pubescent, caducous. Flowers 6-9 mm long, pinkish –purple. Pedicel very short, The Species 103 thickish, articulate. Sepals 5, erect, linear-lanceolate, 3-4 mm, thick, concave, pubescent outside, glabrous inside. Petals 5, less than 2-2.5 mm times as long as the sepals, erect, tube- shaped, linear-lanceolate, concave, thick, the edges thickened and undulate, glabrous, with a thick, medial single ridge at the inside, which is adnate to the disc. Fertile stamen oner, shorter than the petals; filament long, exert, lower part white, upper part violet; anther short, ovate. Staminodes 3-4, adnate to the disc, tooth-like, short, whitish green. Disc narrow, embracing the base of the subglobose ovary, imperfectly 5-lobed. Style sublateral, 7 mm, exerted, dark violet; stigma minute, terminal. Pistil absent in male flowers. Fruits: at maturity yellowish brown, roughish, pear-shaped or ellipsoid, not compressed, gibbose at one side of the base, obliquely emarginate at the apex. Pulp copious, white, very juicy with strong flavour (to some agreeable, to others offensive) very fibrous. Stone lanceolate, slightly compressed, ellipsoid, white, fibro- coriaceous. Seed erect. (Kostermans and Bompard 1993)

Vernacular names. Indonesia: Kemang (Sundanese, W. Java), Kemang (Malay, Sumatra), Palong (Kutai, East Kalimantan); : Kemang, Kemanga (Peninsular), Beluno (Sabah)

Ecology. A species of the wet topical lowlands, generally below 400 m (rarely up to 800 m). It requires a rainfall which is evenly distributed through the year. Rather rare in forests, being found more frequently in periodically inundated areas and marshes.Commonly cultivated on periodically inundated riverbanks.

The Species 104

Conservation status. Not assessed. However this species is one of the rare species in Sumatra.

Use. For Malay people and Sundanese kemang fruit is the favorite one. Ripe Kemang fruit is eaten fresh, pickled, or used for home-made juice with sugar, ice and a pinch of coffee powder. Unripe fruit used to make ‗rujak‘, a dish of sliced green fruit covered with a spicy sauce. Occasionally a dish is made from fresh, grated seeds, with fermented soya beans and spices. The fruit is very popular in West Java and can be found in supermarkets. The Sundanese in western Java commonly eat the young leaves in ‗lalap‘, a dish of fresh leafy vegetables. Unlike the related Mangifera caesia, the sap of this species does not cause skin irritations or blisters. Its timber is pinkish and better than M. caesia ‗binjai‘, being more compact and firmer.

Notes. M. kemanga is closely related to M. caesia, with which it is united by some botanists (Flora Malesiana). However, it should be considered as a species distinct from M. caesia. It can be recognized by its subsessile leaves with narrowly decurrent margins; its panicles which are longer (up to 75 cm long), more open and contain fewer flowers; its fruits which are dull greenish-brown to yellowish-brown at maturity, gibbous at the base and scurfy with pale creamy-white flesh (Fig. 50). The Species 105

Fig. 50The fruit of M. kemanga

Agronomy. It was propagated from seed, an indication that the agronomy of the crops has received little attention. Presumably grafting on seedling stock would be possible, in particular through inarching potted, decapitated root stocks into twigs of mother trees. Grafting on M. indica has been attempted without success, probably because this species is not closely related. Mature trees require much space, 12-16 m each way. Ripe fruit must be handled with care as it is soft and juicy. The Species 106

Fig. 51The leaves of M. kemanga

Distribution. Sumatra—presently known from Riau, Jambi, S. Sumatra (Fig. 52). Analysis of global climate suitability map of M. kemanga in Sumatra generated using MaxEnt model is depicted in Fig. 52. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. kemanga. In fact, this worldwide climate suitability map can be used in the countries that lack precise coordinates of M. kemanga occurrences and generate a preliminary climate suitability map of M. laurina because it may be too late to wait for the precise coordinates of M. kemanga occurrences to generate a perfect climate The Species 107 suitability map. For M. kemanga, It can be seen that suitable conditions are predicted to be highly probable through most of wet lowland areas of Sumatra.

Fig. 52 Distribution of M. kemangain Sumatra based on MaxEnt model

Kemang is indigenous to Peninsular Malaysia, Sumatra and Borneo. It is commonly cultivated in western Java, especially near Bogor, but less frequently in Borneo where binjai dominates.

M. kemanga was scattered in lowland areas at an altitude of 4-27 m asl. M. kemanga can be found in soil types of tropudalts, paleudults, tropaquepts, troportents, fluvaquents and tropudults with slope 0-25% and> 40%. M. kemanga is one of the most endangered species in Sumatra. Nowadays, The Species 108 this species only found in riverside along the South Sumatra region (Bengkulu, Lampung and South Sumatra) until Jambi along the Batang hari riverside. Based on local wisdom which contains conservation value, Kemang trees which located in the riverside are believed to prevent the landslide and flood therefore the rest of M. kemanga trees in Sumatra were big and old tree which have never been cut down.

The Species 109

Trees: Tree, up to 40 m tall and 80 cm dbh, growth habit spreading, crown spherical, bark grey, similar to that M. indica. Leaves: very dense,extremely dark green, semi- erect, scattered and subverticillate, chartaceous, ovate-elliptic, obscurely acuminate with sharp tip, base shortly cuneate, both surfaces very densely, very minutely reticulate-subareolate; midrib prominulous both sides, distinct, nerves above rather obscure to distinct, below slender, erect- patent, 16-18 pairs, slender, prominulous. Petioles long, both side convex, the base slightly thickened. Panicles: pseudo- terminal, pyramidal, lax, up to 40 cm long. Main peduncle not very thick to thickish, light green, smooth, secondary branchlets short, very thin. Flowers not glomerulate, 5-merous, large for the genus, white. Sepals 5, erect, the tips slightly cut, narrowly ovate, acute, thin. Petals 5, ovate, not narrowed at the base, thin, acutish. Ridges 5-7, faint, red, the tips darker red and slightly free from the surface, the entire like a hand and fingers upright. Fertile stamen one, almost central. Disc large, cushion-like. Ovary with short excentric style. Fruits: small, mango-shaped, broadly obovoid, fruit skin yellowish green, apex obtuse, base oblique, neck prominence slightly prominent, glossy, yellowish at maturity. Pulp thin, rather dry, soft, pale yellow to orange with rather vague sweet acid taste. Stone very large, longitudinally furrowed, no or few short fibres. Seeds oblong, apex obtuse, base obtuse. Monoembriony. The Species 110

Ecology. In the wild very rare. Also cultivated. The fruit appears in great quantities in local markets. Distribution. Sumatra—presently known fromSouth Sumatra. (Fig. 53). Analysis of global climate suitability map of M. lalijiwa in Sumatra generated using MaxEnt model is depicted in Fig.53. Warmer colors show areas where better predicted conditions exist. White dots show the presence locations used for training, while violet dots show test locations. The red color indicates areas with a high probability of occurrence, the blue and green represent moderate probability, the yellow color represents low probability of occurrence and the white indicates areas not suitable for M. lalijiwa. In fact, this worldwide climate suitability map can be used in the countries that lack precise coordinates of M. lalijiwaoccurrences and generate a preliminary climate suitability map of M. lalijiwa because it may be too late to wait for the precise coordinates of M. lalijiwa occurrences to generate a perfect climate suitability map.

The Species 111

Fig. 53 Distribution of M. lalijiwain Sumatra based on MaxEnt model

Phylogenetic Analysis 103

Trees: Tree, up to 40 m tall and 80 cm dbh, growth habit spreading, crown spherical, bark grey, similar to that M. indica. Leaves: very dense,extremely dark green, semi- erect, scattered and subverticillate, chartaceous, ovate-elliptic, obscurely acuminate with sharp tip, base shortly cuneate, both surfaces very densely, very minutely reticulate-subareolate; midrib prominulous both sides, distinct, nerves above rather obscure to distinct, below slender, erect-patent, 16-18 pairs, slender, prominulous. Petioles long, both side convex, the base slightly thickened.Panicles: pseudo-terminal, pyramidal, lax, up to 40 cm long. Main peduncle not very thick to thickish, light green, smooth, secondary branchlets short, very thin. Flowers not glomerulate, 5-merous, large for the genus, white. Sepals 5, erect, the tips slightly cut, narrowly ovate, acute, thin. Petals 5, ovate, not narrowed at the base, thin, acutish. Ridges 5-7, faint, red, the tips darker red and slightly free from the surface, the entire like a hand and fingers upright. Fertile stamen one, almost central. Disc large, cushion-like. Ovary with short excentric style. Fruits: small, mango-shaped, broadly obovoid, fruit skin yellowish green, apex obtuse, base oblique, neck prominence slightly prominent, glossy, yellowish at maturity. Pulp thin, rather dry, soft, pale yellow to orange with rather vague sweet acid taste. Stone very large, longitudinally furrowed, no or few short fibres. Seeds oblong, apex obtuse, base obtuse. Monoembriony.

Vernacular names. Rawa-rawa or rawo-rawo (Sumatra, Palembang) Phylogenetic Analysis 104

Ecology. A species, preferring temporarily inundated areas, on river levees, marshy places behind the leves, lowland, ever wet tropical climate.

Distribution. Western Malesia: Malay Peninsula and Borneo, also cultivated in Sumatra; not in Java, also not cultivated; not in Celebes.

Phylogenetic Analysis 105

CHAPTER 4 Phylogenetic Analysis

What the common ancestor of Mangifera is he family Anacardiaceae Lindley 1830 is closely related to Burseraceae. Cronquist (1968) included the family in the Order Sapindales Benth. & Hooker (1862); the order encompassing 15 families, the largest one being Sapindaceae and Rutaceae. Takhtajan (1969) included the family in the T Order Rutales, keeping separate the Order Sapindales. Both may be right or wrong.

Kostermans and Bompard (1993) suggested that there was no way (not even with cladisctics) to find out the ancestry of the genus and the species of Mangifera. The two subgenera, Mangifera and Limus, may have originated from two different ancestors.

Primitive characters are considered: 10 stamens in two whorls, implying that the greater part of the species is not primitive, and the one or more-celled condition of the pistil. Mangifera‘s ovary seems to have been originally three-locular (Sharma 1954), its one-locular character state is considered an advanced one. Previous findings suggest that, allinflorescences show exactly the same pattern.As the genus is entirely Asiatic, it may be compared with the entirely Asian Dipterocarpaceae, that are assumed to be of Gondwanian origin.

Phylogenetic Analysis 106

In order to quantify the broad relationship among the Mangifera species of Sumatra, a cladistic analysis was performed using 4 molecular markers. A secondary aim to this analysis was to re-examine Kostermans and Bompard‘s groupings, insofar as they are applicable to the species from this region. The analysis was performed using MEGA6 with the parsimony method. Strict consensus trees derived from the most parsimoniuos cladogram generated by this analysis are presented in Fig. 54 (ITS sequence), Fig. 55 (trnL-F IGS), Fig. 56 (rbcL gene) and Fig. 57 (matK gene). Important clades are numbered and discussed in the next followed. Details of the methods used in the analyses are presented in Appendix A. Cladistics allow researchers to make phylogenetic inferences from numerical data about the organisms they wish to compare. While this is of considerable interest in this analysis, the primary objective is to examine the superficial similarities among the species. The evolution of (and within) Mangifera remains nebulous and as the data in this analysis include genetic information should be considered definitive of evolutionary relationship in the genus as a whole.

What the suitable DNA Barcode of Mangifera is he use of morphological characters has long been used in many phylogenetic studies but the morphological character plasticity makes it difficult to define evolutionary relationships. As T a results informative molecular-based DNA sequencing approaches are profoundly required to support and reinforce morphological character Phylogenetic Analysis 107

data. The development of molecular biology techniques of this century facilitates the assessment of genetic kinship of a taxa. Mango research on DNA barcoding has not been conduct for mango identification, especially from Sumatra.

Based on Barcode of Life Database (BOLD) The standard barcode DNA sequence data system is rbcL and matK for coding region, this sequence is conservative and slightly varied especially at the species or infraspesies level. Therefore, the use of non coding markers may support information for more obvious separation of species and infraspecies.

Chloroplast DNA (cpDNA) has been used extensively in the study of phylogeny at several taxonomic levels. The results of PCR direct sequence are widely used in evolutionary and plant systematic studies. The rbcl gene encoding a large ribulose- 1.5-bisphosphate carboxylase / oxygenase (RUBISCO) unit has been traced to a number of plants.

MaturaseK (matK) is a candidate of barcode genes in angiosperm. This gene provides a potential area in flowering plants. This gene is often chosen because it provides a single copy and has variations in DNA nucleotide substitution and also widely used in the phylogenetic studies of different plant species. Furthermore, the most widely used chloroplast (CpDNA) marker is the trnL-F intergenic spacer, part of the noncoding cpDNA genome, this region is more variable than the coding region, making it more suitable for exposing the evolutionary relationship to the lower taxa ( Bayer et al., 2000). Several studies in non-coding areas of chloroplasts exhibit higher variation and frequent mutations (Baldwin et al. 1995) in the form of transversions, transitions, insertions, and deletions. Phylogenetic Analysis 108

The intergenic spacer region between trnL (UAA) 3 'exon and the trnF gene (GAA) also has the potential for phylogenetic studies (Soltis et al., 1998). This area of DNA is easily amplified and sequenced, its size is relatively small 120-350 bp and single copy genes, making it relatively easy to test the entire genome. Regional sequences are more informative at the genetic and species levels

The ITS (Internal Transcribed Spacer) area is part of the core ribosomal DNA (nrDNA) that encodes the ribosomal RNA (rRNA gene). Each nrDNA unit in a series of chromosomes has an encoding region of 18S, 5.8S, and 26S flanking the ITS region (Baldwin et al., 1995). In the last two decades, the use of the ITS sequence has proved effective in some cases to reconstruct phylogenetic relationships at the species level and cultivar

Internal Transcribed spacer (ITS) of nrDNA has been used for molecular markers at specific level of Angiospermae (Baldwin et al. 1995; Yonemori et al. 2002). Sequences of ITS were also useful because it has conserve region, short size (±700 bp), high evolution rate, informative and universality (Baldwin et al. 1995). Molecular approach has benefit to find the best phylogenetic tree model which useful in conservation and cultivation strategies.

109 Phylogenetic Analysis

Table 4Sources of Mangifera ITS sequences and their geographical origin Species Genbank Acc. no. (ITS) Geographic origin Reference M. kemanga acc.1 KX347955 Central Sumatra M. kemanga acc.2 MF678503 Southern Sumatra M. kemanga acc.3 MF990368 Southern Sumatra M. foetida acc.1 KX347956 Central Sumatra M. foetida acc.2 MF678499 Eastern Sumatra M. foetida acc.3 MF678500 Eastern Sumatra M. foetida acc.4 MF678501 Southern Sumatra M. foetida acc.5 MF678505 Southern Sumatra M. foetida acc.6 MF678506 Southern Sumatra M. odorata acc.1 KX347957 Central Sumatra M. odorata acc.2 MF678496 Eastern Sumatra M. odorata acc.3 MF678497 Eastern Sumatra M. odorata acc.4 MF678507 Southern Sumatra M. laurina acc.1 KX347963 Central Sumatra M. laurina acc.2 MF678495 Eastern Sumatra M. laurina acc.3 MF678498 Eastern Sumatra M. laurina acc.4 MF678508 Southern Sumatra M. laurina acc.5 MF990367 Southern Sumatra M. indica acc.1 KX347960 Central Sumatra 110 Phylogenetic Analysis

M. indica acc.2 MF678502 Eastern Sumatra M. indica acc.3 MF678509 South Sumatra M. indica acc.4 MF678510 Southern Sumatra M. zeylanica acc.1 KX347962 Central Sumatra M. zeylanica acc.2 MF990364 Southern Sumatra M. lalijiwa MF678504 Southern Sumatra M. casturi MF678493 Central Sumatra M. sumatrana acc.1 KX347961 Central Sumatra M. sumatrana acc.2 MF990366 Southern Sumatra M. quadrifida acc.1 KX347959 Central Sumatra M. quadrifida acc.2 MF678511 Southern Sumatra M. torquenda acc.1 KX347958 Central Sumatra M. torquenda acc.2 MF990365 Southern Sumatra Mangifera sp. 1 KX347964 Central Sumatra Mangifera sp. 2 MF678494 Eastern Sumatra Genbank sequences BoueamacrophyllaGriff. AB071691 Narathiwat [4] Anacardiumoccidentale AB071690 Nakhon Si Thammarat [4]

Phylogenetic Analysis 111

Table 5Sources of Mangifera trnL-F IGS sequences and their geographical origin Species Genbank Acc. no. (ITS) Geographic origin Reference M. kemangaacc.1 KY392620 Central Sumatra MF919592 M. kemanga acc.2 MF919593 Southern Sumatra M. kemanga acc.3 MF919594 Southern Sumatra M. foetidaacc.1 KY392618 Central Sumatra M. foetidaacc.2 KY392613 Eastern Sumatra M. foetida acc.3 KY392608 Eastern Sumatra M. foetidaacc.4 MF945597 Southern Sumatra M. foetida acc.5 MF997585 Southern Sumatra M. foetida acc.6 MF997584 Southern Sumatra M. odorataacc.1 KY392623 Central Sumatra M. odorata acc.2 KY392610 Eastern Sumatra M. odorataacc.3 KY392615 Eastern Sumatra M. odorata acc.4 MF945595 Southern Sumatra M. odorataacc.5 MF945596 Southern Sumatra M. laurinaacc.1 KY392621 Central Sumatra M. laurinaacc.2 KY392612 Eastern Sumatra M. laurinaacc.3 KY392609 Eastern Sumatra M. laurina acc.4 MF997588 Southern Sumatra M. indicaacc.1 KY392619 Central Sumatra Phylogenetic Analysis 112

M. indicaacc.2 KY392616 Eastern Sumatra M. indica acc.3 MF997586 Southern Sumatra M. zeylanicaacc.1 KY392611 Eastern Sumatra M. zeylanicaacc.2 MF997591 Southern Sumatra M. lalijiwa MF997587 Southern Sumatra M. sumatrana MF997590 Southern Sumatra M. quadrifidaacc.1 KY392614 Eastern Sumatra M. quadrifidaacc.2 MF997589 Southern Sumatra Mangifera sp. 1 KY392622 Central Sumatra Mangifera sp. 2 KY392607 Eastern Sumatra Genbank sequences BoueamacrophyllaGriff. KY392617 Central Sumatra Bouea

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What The ITS Sequence can tells

Parsimony analysis of ITS sequence revealed the monophyletic group of Mangifera in Sumatra with bootstrap value 100% (Fig. 54). M. quadrifida acc.1 from Central Sumatra separate from the rest of Mangifera accessions and considered as the root species of Mangifera.

M. casturi from Central Sumatraand Mangifera sp. 2 from Eastern Sumatra are closely related to M. griffithii from Thailand (we assume that Mangifera sp. 2 are M. griffithii). They are forming a clade with bootstrap value 53%.

Mangifera sp. 1 is the sister group from a clade consists of M. sumatrana acc.1, M. kemanga acc.3, M. torquenda acc.1, M. torquenda acc.2, and M. quadrifida acc.2. Together with Mangifera sp. 1, all of them are forming a monophyletic group with bootstrap value 99%.

Most of the accession from subgenus Limus (M. kemanga acc.2, all of the accessions of M. foetida and M. odorata) are forming a clade together with M. laurina acc.5 with bootstrap value 90%.

All of the accessions of M. zeylanica, M. laurina, M. indica are forming a clade together with M. kemanga acc.1 and M. sumatrana acc.2. This clade mostly from subgenus Mangifera except for last two accessions mentioned.

Generally,grouping based on ITS sequence is more reliable and natural than other molecular markers. However, there were Phylogenetic Analysis 114 several accessions that confound the cladogram. Notto mention they are from the same species. There were four confusing fact obtained from this analysis. They scattered in the tree and not assemble in a clade. First fact, M. kemanga acc. 1 from Central Sumatra, M. kemanga acc. 2 and M. kemanga acc. 3 from Southern Sumatra. Second fact, M. quadrifida acc. 1 from Central Sumatra and M. quadrifida acc.2 from Southern Sumatra. Third fact, M. sumatrana acc. 1 from Central Sumatra and M. sumatrana acc.2 from Southern Sumatra. Four fact, M. laurina acc.5 from Southern Sumatra. Based on this ITS sequence, all of them are not form a clade in the name of same species. These facts confound the tree the most.

The ITS sequence is nearly perfect to be a potential DNA barcode for the Mangifera if there were no minor errors for some deviant accessions from the tree. However researhers cannot deny that it is to be blame due the erroneous during work in laboratory. This is understandable. So that ITS is recommended as one of the best marker for Mangifera in infraspecific level.

What The trnL-F Intergenic Spacer can informs Parsimony analysis of trnL-F Intergenic Spacer sequence revealed the monophyletic group of Mangifera in Sumatra with bootstrap value 89% (Fig. 55). At least, there were four main subclades are important to discussed.

Subclade I consists of M. indica acc.1, M. indica acc.2, M. laurina acc.3 and M. zeylanica acc.1. Subclade II comprised of Phylogenetic Analysis 115

Mangifera sp. 2, M. foetida acc.2 and acc.3, M. odorata acc.1, acc.2 and acc.3, M. quadrifida acc.1 and Mangifera sp. 1.

All of accessions of M. kemanga form a clade with bootstrap value 71%. M. odorata acc.1, M. laurina acc.1 and acc.2 form a clade with bootstrap value 100%. All of them together with M. odorata acc.4 form a larger monophyletic group in subclade III.Subclade IV contained 7 different species such as M. quadrifida acc.2, M. lalijiwa, M. foetida acc.6, M. sumatrana, M. zeylanica acc.2, M. laurina acc.4, M. foetida acc.5 and M. indica acc.3.

Comparing to the classification based on morphological characters, this marker could not differentiate species between subgenus Limus and Mangifera. All of the accessions are scattered all over the cladogram.

Generally, the grouping based on trnL-F IGS sequence is more varied than ITS sequence. From the cladogram we could say that the trnL-F IGS could not gather the accession from the same species in a clade, except for M. kemanga case. It means trnL-F did not suitable for using in lower taxa level such as infraspecific level.

What The rbcL Gene can manifests Parsimony analysis of rbcL sequence revealed the monophyletic group of Mangifera in Sumatra with bootstrap value less than 50% (Fig. 56). There were seven subclades on the cladogram. All of the accession from same species scattered over the cladogram. The grouping based on rbcL gene did not able to strictly separate the subgenus Mangifera. Even the same species Phylogenetic Analysis 116 did not gather in a clade. As we know rbcL gene has many conservative gene with low evolution rate therefore this marker could only separate the genus level in case of Mangifera.

What The matK Gene can says

Parsimony analysis of matK sequence revealed the monophyletic group of Mangifera in Sumatra with bootstrap value less than 50% (Fig. 57). There were three subclades formed. Subclade I only consists of Mangifera sp. 1 from Central Sumatra. Subclade II contained species of M. sumatrana acc.2, M. odorata acc.2, M. indica acc.2 and M. zeylanica. Subclade III comprised of M. sumatrana acc.1, M. quadrifida acc.2, M. foetida acc.1 and acc.2, M. laurina, M. indica acc.1, M. odorata acc.1, and M. quadrifida acc.1

The grouping based on rbcL gene did not able to strictly separate the subgenus Mangifera. Even the same species did not gather in a clade. matK gene has many conservative gene with low evolution rate therefore this marker could only separate the genus level in case of Mangifera.

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Fig. 54 The 50% majority-rule consensus tree derived from the maximum parsimony analysis of ITS sequences for Mangifera and outgroup taxa. Numbers below branches are bootstrap value

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Fig. 55 The 50% majority-rule consensus tree derived from the maximum parsimony analysis of trnL-F Intergenis Spacer sequences for Mangifera and outgroup taxa. Numbers below branches are bootstrap value Phylogenetic Analysis 119

Fig. 56 The 50% majority-rule consensus tree derived from the maximum parsimony analysis of rbcL sequences for Mangifera and outgroup taxa. Numbers below branches are bootstrap value

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Fig. 57The 50% majority-rule consensus tree derived from the maximum parsimony analysis of matK sequences for Mangifera and outgroup taxa. Numbers below branches are bootstrap value

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Three potential DNA barcodeswere use entailing ; Internal Transcribed Spacer, rbcL gene, and trnL-F Intergenic Spacer. Each of markers nearly gives the similar clade pattern. However, for lower taxa level ITS is better than tnrL-F IGS than rbcL.

Results of alligned sequence of entire ITS revealed ITS region was flanking conserve 5.8S region (coding region) encoded ribosomal RNA which is important in protein synthesis (add reference). Mutation rate of conserve gene is slower than non coding region. ITS region as non coding region has more variation and higher mutation rate than coding region. Non coding region (intron) has role in gene expression regulation which adaptable with niche/habitat. Most of this non coding region could be observe through phenotypic characters.

Based on this study many differentiations were found between classification based on morphological characteristics by Kostermans and Bompard (1993) and molecular study .This results could be use as strong basis to develop a new system of classification. Classification based on DNA sequence is assumed to produce nature and accurate classification because DNA is a basic unit of information that encode organism.

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CHAPTER 5 Conservation

What Mangifera should be conserved he major threats to wild Mangifera in Sumatra are habitat destruction and removal of plants by collectors. For most species, loss of habitat is the more pressing concern, but few seem to be seriously imperilled by T either threat at present. This is because the majority of species occur in remote or protected locations which have not been disturbed by humans, or flourish in secondary habitats and may actually benefit from certain kinds of disturbance. Assuming the remote or protected area remain undisturbed, the species that occur within them are unlikely to be threatened in the foreseeable future. Despite this, a small number of species appear to be threatened, while very little is known about several others. In this chapter, the conservation status of all Mangifera species from Sumatra is evaluated and discussed.

Assesing The Conservation Status of Mangifera in Sumatra Since the last publication by Kostermans and Bompard (1993) has showed that there were approximately 34 species of Mangifera ever found in Sumatra. However, since the last exploration by Fitmawati in 2016, there were only 12 species remain. It was agreed with the data that these species were threatened by natural habitat loss. 123

Margono et al. (2014) showed that only 13.4 Mha or 28.3% of the land area of Sumatra remains covered by primary forests in 2012. Of the various forest types, Mangifera are abundant in lowland. See Table 6. Within Indonesia, Sumatra Island stands out due to the intensive forest clearing that has resulted in the conversion of 70% of the island‘s forested area through 2010 (Margono et al . 2012). So far only a few species are in real danger of extinction see Table 7.

Table 6. Primary forest extent in Sumatra in 2012, after Margono et al. (2014) Landforms Mha Percent coverage Lowland 4.5 9.4% Wetland 2.7 5.7% Upland 3.7 7.8% Montane 2.5 5.3% Total 13.4 28.3%

Table 7. IUCN status of Mangifera M. zeylanica WCMC 1998 Vulnerable A1c M. rubropetala WCMC 1998 Extinct in the Wild M. magnifica WCMC 1998 Lower Risk/Least Concern M. odorata WCMC 1998 Vulnerable M. foetida WCMC 1998 Lower Risk/Least Concern M. quadrifida WCMC 2016 Least Concern M. similis WCMC 1998 Vulnerable A1c M. superba WCMC 1998 Endangered M. casturi WCMC 2016 Extinct in the Wild 124

M. caesia WCMC 1998 Lower Risk/Least Concern

Conservation of Mangifera species should be both ex situ and in situ. Although in situ conservation has more advantages, as the trees remain in their natural niche, its drawback is that the nature reserves in Asia, especially in Indonesia, are not safe. The reserves exist on paper but it is practically impossible to protect them from a continuously exploding hungry population, looking for new arable land and trees to cut for income. Even officially they are sometimes logged. Nature reserves created on logged and partly burnt areas, like Bukit Suharto in E. Kalimantan, do not make sense; such areas take a thousand years to become re- afforested naturally, owing to lack of propagules.

However, as there is no alternative, it is advisable to introduce endangered species from elsewhere into such reserves, with the faint hope that they will be safe against man. It should be stressed that the local population is not responsible for the vegetation destruction in Asia, the main felon being the foreigner who cuts the timber to make huge profits, irrespective of the future of the local inhabitans. Proposals have been put forward in developed countries to stop the vegetation demise by refusing such timer imports, but in vain; money is more important than human beings.

The alternative ex situ conservation, may be executed in several differents ways. Endangered species could be transplanted in botanic gardens (they are too small and too expensive for undeveloped countries), in gene banks (often not viable without foreign support), migrated to foreign countries of 125 better economic standing, but the best way is to bring them to fruit farm nurseries in the tropics (goos examples are in Thailand and tropical Australia). Gene banks are created by the endeavours of IBPGR, Rome.

So far, we have discovered that the most of the Mangifera species are not rare, but rather occur as single individuals spread over large areas. The danger is that these so-called wild species (for the most part, cultivated in restricted places on a small scale) are becoming unpopular with the influx of alien transmigrants, who do not know the species and thus are not interested in their maintenance and propagation.It is rare nowadays to find that any of the younger population know the useful trees, formerly grown by their ancestors.

Mangifera species, like most other tropical trees, are very sensitive to their niches and small differences in climate or soil may hamper growth or fruit setting. Hence collections should not be established in one place, but on the different islands of Malesia and the different countries of mainland S.E. Asia. In Malaysia, the Forest have started to establish collections of wild fruit trees. In Indonesia, remnants of colonial horticultural fruit gardens still exist and new collections have been established, but the development is stagnating or has come to a standstill.

The result is still that very little known of the Mangifera species even in the Malay Peninsula. Some have been collected only once in the last 150 years. Horticulturists in Java are ignorant of species outside Java.

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