The common mango apparently originated in regions on the western border of the secondary centre of diversifi cation mentioned above. Truly wild mango trees have been recorded in Bangladesh (Chittagong Hill tract, c.23°N), north-eastern India (‘undoubtedly indigenous in the evergreen tracts of valley of Assam’ according to Kanjilal et al., 1937), and in Myanmar where it was reported as ‘not unfrequent in the tropical and lower mixed forests all over Burma from Arracan and Pegu down to Tenasserim’ (Kurz, 1877). It would be desirable to assess its affi nity with the species of the section Euantherae, as well as with species of other sections of the subgenus Mangifera that occur in the same area and region. It is also believed to be wild ‘in the sub-Himalayan tract, in deep gorges of the Baraitch and Gonda hills in Oudh, and the outer hills in Kamaon and Garhwal’ (Brandis, 1874). The common mango has been grown and disseminated for such a long time in India that semi-wild trees can be found in the forests throughout the subcontinent. The fruits of wild trees are said to be small and of poor quality. Watt (1891) mentioned two so-called ‘almost unaltered wild varieties’ existed under cultivation in Tirhoot,
‘one originating from Kangra, a very variable one, and the other from Sikkim which was evidently the progenitor of the varieties cultivated in Malda’.
The common mango in South-east Asia
The Linnean binomial (Mangifera indica) indicates in this instance the place where the common mango was selected and improved, and not necessarily its place of its origin. It has been traditionally accepted that mango was domesti-cated several millennia ago in India (see Mukherjee and Litz, Chapter 1, this volume); however, it cannot be excluded that domestication occurred inde-pendently in several areas, possibly in the south-western and south-eastern regions of its centre of origin, or later differentiated in those two regions. This hypothesis would account for the differences that exist between the local polyembryonic cultivars of Myanmar, Thailand, Indochina and Indonesia, and the monoembryonic Indian cultivars. Note that polyembryony occurs
also in the cultivated M. casturi, M. laurina and M. odorata. Aron et al. (1998) have demonstrated that polyembryony in mango is under the control of a single dominant gene.
According to Juliano (1937), Bijhouwer suggested that there were two main centres of domestication of mango, ‘one in India with monoembryonic mangoes, the other in the Saigon area, Indonesia and the Philippines with polyembryonic mangoes’. The ‘Saigon’ area must in fact be extended to southern Vietnam, other parts of Indochina, Thailand and Myanmar, which were recognized by Valmayor (1962) as homes of polyembryonic mangoes.
Notwithstanding, the origin of polyembryonic mangoes is probably better placed in Myanmar, and possibly the eastern part of Assam. According to Brandis (1874), ‘in Burma, the mango is not generally grafted, and seeds of a good kind, as a rule, produce fruit of a similar description’. There are only a few polyembryonic mango cultivars in India. They are restricted to the south-western coastal region, and geographically isolated from the polyembryonic mangoes of Myanmar and South-east Asia. Analysis of genetic relatedness using RAPD markers among polyembryonic and monoembryonic cultivars grown in the west coast of southern India suggest that the polyembryonic types are unlikely to have originated from India and might have been intro-duced from South-east Asia (Ravishankar et al., 2004).
Indian Buddhist monks might have introduced the common polyembry-onic mango to South-east Asia, fi rst along land trade routes through Myan-mar, where they might have found better races, and from there into insular South-east Asia. It is well established that some local names of the common mango currently used in parts of Indonesia are of Sanskrit origin (‘ampelam’
and its cognates), and are sometimes used to designate M. laurina, which is a truly native species. Vernacular names do not always travel with a plant, and even if they did so in the case of the common mango, it is very unlikely that these introductions were the fi rst ones and that they came obligatorily from India. In the absence of a comprehensive classifi cation of the innumerable South-east Asian cultivated forms of the common and wild mangoes, includ-ing the countless primitive races, we have to rely on linclud-inguistics and the rich history and prehistory of this region.
Vernacular names
The different local names of the common mango in Indonesia (‘pauh’, ‘ampe-lam’ and its variants, and ‘mangga’) bear evidence of a long history of con-tacts with mainland Asia and India, and point to possible introduction at different times from different places. In some parts of Indonesia, the vernacu-lar names ‘paoh’ or ‘pauh’ refer either to primitive races of the common mango, or to native species, as a rule the ones most closely resembling the common mango, for example: ‘pauh asal’ (= native mango) for M. pentandra in peninsular Malaysia; ‘pahohutan‘ or ‘pahutan’ (= forest mango) for M.
altissima in the Philippines; and ‘pao pong’ (= forest mango) for M. minor in Flores, Lesser Sunda Islands. ‘Pau’ is a word belonging to Austronesian lan-guages, nowadays spoken over a very wide area from Madagascar to the Easter Islands by people who originate from mainland Asia. These languages
are still spoken by certain minority populations in Vietnam, Cambodia and the Mergui Archipelago off the coast of Myanmar (Bellwood et al., 1995). In Cambodia, which was occupied by the Chams from about the 3rd to the 15th century ad, ‘pa:uh’ is a Chamic word. ‘Sva:y’, used by the Khmers (as in
‘sva:y srok’ meaning mango of the village (M. indica), and ‘sva:y prey’, wild mango (i.e. M. caloneura) as attested in pre-Angkorian Khmer inscriptions dating from the 6th to the 8th century ad (Pou and Martin, 1981)) is of Austro-Asiatic origin. ‘Sva:y’ has cognates in south Vietnam (‘xoay’) and in Asian languages spoken by aboriginal people in peninsular Malaysia. ‘Wai’, another cognate, is a vernacular name of M. minor in several parts of New Guinea.
Pawley and Ross (1995) proposed ‘wai’ and ‘pau(q)’ as the reconstructed Proto-Oceanic terms referring, respectively, to a generic name for mango, and a species that is probably M. indica.
Nowadays, these two words are generic terms for mango fruits that rather closely resemble M. indica. In the same way, ‘thayet’ which is the com-mon vernacular name referring to M. indica in Myanmar (‘sinnin thayet’ and
‘taw-thayet’ for M. caloneura and M. sylvatica, respectively), or ‘mamuang’ in Thai languages are probably generic names.
Obviously, linguistic evidence alone provided by these vernacular names is not suffi cient to prove the time and place of an introduction. None the less, it is signifi cant that in mainland South-east Asia none of the vernacular names of the common mango exhibits signs of an Indian infl uence, moreover, cog-nates of these names are also applied to primitive races in some parts of insular South-east Asia.
Evidence of early trade in South-east Asia
The history of plant domestication in mainland South-east Asia has undoubt-edly involved introduction of plants by people migrating from the mainland into insular South-east Asia. In more recent times, there is evidence of con-tacts and sea trade since at least the fi rst centuries ad between mainland and insular South-east Asia to indicate that there have been numerous opportuni-ties for introduction of the common mango from different places at different times prior to the 4th century (before the Indianization of early South-east Asian states) into present-day Malaysia and Indonesia.
Recent studies based on archaeological evidence stress the long unrecog-nized importance of South-east Asian trade (emanating from South-east Asia) between ports established along the Java Sea, those of mainland Asia, and India, back to the 1st century ad, and possibly earlier (Walker and San-toso, 1984). Trade routes connected the developing population centres of the mainland, such as the earliest known South-east Asian political entity, Funan, an advanced agrarian society located on the southern Vietnam coast, which became infl uenced by the Indians and reached the zenith of its commercial prosperity in the middle of the 3rd century (Hall, 1985). Increasingly, king-doms organized according to the Indian concept of royalty were established in the Indonesian archipelago, for example Kutai in East Kalimantan (4th century) and Central Java (8th to 9th century), the latter being famous for the Buddhist temple at Borobudur, where sculptures depict the mango tree.
It is highly probable that the eventual introductions of superior cultivars of polyembryonic mangoes from the south-west coast of India, ‘between the 6th and 14th century, the height of classical South-east Asian civilization and also the golden age of early south Indian civilization’ (Hall, 1985), were not the fi rst ones.
During the 16th and 17th centuries, the Portuguese and Spaniards con-tributed to the widest distribution of superior varieties in the archipelago, espe-cially to the east. The name mango itself derives from the Tamil ‘man-kay’ or
‘man-ga’ (see Mukherjee and Litz, Chapter 1, this volume), which the Portu-guese adapted as ‘manga’ and ‘mangueira’ when they colonized west India.
Superior Philippine cultivars originated through introduction of culti-vars from Indonesia, for example ‘Dodol’ into Mindanao, and from Indo-china, for example ‘Carabao’ and ‘Pico’ in Luzon, the Visayas and northern Mindanao (Wester, 1920; Bondad et al., 1984). However, these introductions dating from the fi rst half of the 17th century were also preceded by the intro-duction of primitive races of the common mango as well as other species into the Sulu Archipelago and Mindanao through contacts with north Borneo, as attested by their local names quoted by Wester (1920), that is mampalam (M. indica, and possibly also M. laurina), baonoh (M. caesia) and wannih (M.
odorata).
The South-east Asian M. indica germplasm includes many races that defy classifi cation. Natural cross-pollination has undoubtedly occurred with native species, such as M. laurina, which was also brought into cultivation in several areas before the introduction of M. indica.
2.8 Conclusion
Potential contribution of wild species to mango cultivation
To date, the improvement and breeding of common mango has depended on the use of genetic variability within a single species, M. indica. Mukherjee (1957) observed that ‘similarity in chromosome number and pollen morphol-ogy in different species suggests close compatibility during hybridization and stock-scion relationship if other species are used as stock for the com-mon mango’. Biotechnology opens new perspectives for mango improve-ment (Litz, 2004). As noted by Litz et al. (Chapter 18, this volume), the transformation of mango with genes from other species could address a number of plant breeding objectives.
Source of rootstock
Grafting experiments between M. indica and other species are reported in the literature, for example budding of M. indica on M. foetida and M. odorata in Java (Ochse and Bakhuizen, 1931), M. odorata on M. indica in the Philippines (Wester, 1920), and M. indica on M. zeylanica in Sri Lanka (Gunaratman, 1946).
Mangifera indica ‘Madu’ in Java, and M. laurina in Sabah have been used as rootstocks for M. casturi. Trials of grafted M. caesia on M. indica (Wester, 1920) and M. indica on M. kemanga or M. caesia (Ochse and Bakhuizen, 1931) were unsuccessful, as these two species have distinct bark features and only remote affi nity with the common mango. Better compatibility can be expected using species more closely related to the common mango within the subgenus Mangifera. In West Kalimantan, M. laurina is occasionally used as a rootstock for the common mango on periodically inundated riverbanks. It has been tried as a rootstock by the Department of Agriculture in Sabah (Lamb, 1987).
Campbell (2004) reported that M. casturi, M. griffi thii, M. laurina, M. odorata, M. pentandra and M. zeylanica grafted on M. indica had a high percentage of success.
Several species that can grow in permanently inundated areas (i.e. M.
gedebe, M. quadrifi da, M. griffi thii and other species of the section Rawa) repre-sent a potential source of rootstock for the development of mango cultivation on poorly drained soils or in areas liable to prolonged fl ood. Other species may be a source of dwarfi ng rootstocks.
Hybridization
From our observations in Borneo, natural interspecifi c hybridization involv-ing various cultivated Mangifera species can occasionally occur. Suspected hybrids were observed between wild M. gedebe and cultivated M. laurina in the lakes area along the Mahakam River in East Kalimantan, where impor-tant populations of M. gedebe occur; between cultivated M. foetida and M. pajang, two species showing close affi nity, in different areas of Kalimantan where both species are grown together; and between closely related M. caesia and M. kemanga in cultivation. A hybrid origin has been suggested for M. odorata (M. indica × M. foetida), which is unknown in the wild (Ding Hou, 1978a).
Based on AFLP analysis, Teo et al. (2002) and Kiew et al. (2003) have con-fi rmed that M. odorata is a hybrid between M. foetida and M. indica. The index of similarity showed that M. odorata is closer to M. foetida (76% similarity) than it is to M. indica (66%). Yamanaka et al. (2006) showed a high genetic similarity among 11 landraces of M. odorata from the Malaysian Agricultural Research and Development Institute (MARDI) gene bank. Higher variability can be expected from Sumatra and Java samples.
Existing information about experimental interspecifi c hybridization is scarce. According to Mukherjee et al. (1968), successful crosses between M. odorata and M. zeylanica were made in India.
Potential of wild species
There is little doubt that wild mangoes are potentially valuable in breeding programmes. Some species have important horticultural implications as they demonstrate many desirable characteristics (Bompard, 1993). Fairchild (1948)
noted that crosses between the common mango and related fi ve-stamen spe-cies of the section Euantherae might produce hybrids with better pollinating quality. Mangifera pentandra, which is grown in peninsular Malaysia and Sabah, is a prolifi c bearer, due to its high proportion of hermaphrodite to male fl owers.
Stress resistance
In the Malesian rainforests, wild mangoes thrive well under an ever-humid climate, without a prolonged dry season, i.e. is in areas with an annual rain-fall > 4000 mm and no monthly mean < 100 mm and where the common mango cannot be grown satisfactorily. Species, occurring in subtropical areas, including primitive races of the common mango, or in high altitude tropical forests, should be evaluated for cold tolerance, opening up the possibilities for mango production in subtropical and Mediterranean areas. Mangifera lau-rina and other species related to the common mango that grow in the rainfor-est (e.g. M. minor in New Guinea) are apparently immune to anthracnose.
Sharma and Choudhury (1976) also observed that trees of an unknown wild race found in the Tripura State (north-eastern India) were free from mango malformation.
Potential new fruits
Extensive, yet largely unrecorded variability also exists among the non-indica species under cultivation. Sadly, this gene pool is barely represented in exist-ing collections, and is rapidly vanishexist-ing. An increasexist-ing number of horticul-turists are demonstrating a keen interest in the wild relatives of the mango.
It is hoped that local peoples who have contributed to the recognition and maintenance of these species can benefi t from future innovative mango breeding.
Since early times, local peoples have planted seeds collected from trees that were observed to produce better quality fruits in the forests around their settlements. In areas now completely devoid of lowland primary forest, espe-cially in Sumatra and Borneo, the only wild relatives still found are those which have been integrated into indigenous agroforests which represent gene banks for an amazing diversity of fruit trees. A tenuous but constant selection pressure over many centuries has resulted in improved selections of several species. Today, some of these selections hold economic importance for their intrinsic characteristics. In Malesia, forms of M. odorata and M. foetida with sweeter and less fi brous fl esh have been identifi ed. The ‘wani’, a form of M. caesia from Bali and Borneo, has green-skinned fruit with milky white soft fl esh and a sweet taste quite different from the fruit of common forms of M.
caesia. In addition, there are many interesting selections of M. casturi, M. grif-fi thii and M. torquenda.
Further improvement of these wild mangoes is especially desirable owing to their local economic importance in the wet tropical regions. Use of vegetative propagation methods must be encouraged. With proper selection, there is every reason to believe that other Mangifera species can become valu-able commercial fruits.
Acknowledgement
Thanks are due to Dr Dawn Frame who assisted in correcting the text.
Note
1Surveys were carried out in Kalimantan in cooperation with the Indonesian Institute of Science (LIPI) and the Indonesian Commission on Germplasm, and in Malaysia with the Forest Research Institute of Malaysia (FRIM).
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