Genetic Resources of Mangifera Species in Thailand

Home , Mangifera, Mangifera foetida, Mangifera odorata

Jpn. J. Trop. Agr. 43 (2) :76-83, 1999

Wichan EIADTHONG*, Keizo YONEMORI*, Akira SUGIUIZA*, Naoki UTSUNOMIYA*2 and Suranant SUBHADRABANDHU*3

* Laboratory of Pomology, Graduate School of Agriculture, Kyoto University, Sakyo-ku, Kyoto 606-8502, Japan *2 Department of Horticulture, Faculty of Agriculture, Kinki University, Nara 631-8505, Japan *3 Department of Horticulture , Faculty of Agriculture, Kasetsart University, Chattujak, Bangkok 10900, Thailand

Abstract The Mangifera species in Thailand were surveyed during the period 1995-1997, and a total of 13 species were identified . Among these species, M foetida, M. odorata and M sylvatica have been cultivated as fruit crops for the local market , as well as M. indica. The other 9 wild species also seemed to have potential values as rootstocks and breeding materials for disease and/or pest resistance, or tolerance to critical environmental conditions . Based on 55 morphological and ecological characters, 13 Mangifera species were classified into two major groups; the first group consists of M indica, M. sylvatica, M oblongifolia, M. collina , and M cochinchinensis, and the second group of M caloneura, M. pentandra, M flava , M. macrocarpa, M gedebe, M griffithii, M. odorata, and M. foetida. All the species in the first group were distributed in a monsoon climate , not in a moist tropical rain forest climate. Since tropical forests are being rapidly destroyed by the expansion of urban areas, a special effort should be made to conserve Mangifera species in Thailand as valuable germplasm for future needs. Key words Mangifera species, morphological character, phylogenetic relationship, species diversity

タイに存在するマンゴー属植物の探索Wichan EIADTHONG*・米森敬三*・杉浦明*・宇都宮直樹*2・Suranant SUBHADRABANDHU*3 *〒606-8502京都市左京区北自川追分町京都大学大学院農学研究科 *2〒631-8505奈良市中町 3327-204近畿大学農学部 *3タイバンコック10900カセサート人学農学部

要約タイに存在するマンゴー属植物の探索を目的として,1995～1997年の3年にわたり,3～7日間の調査を6回実施し ,合計13種のマンゴ属植物のタイでの存在を確認することができた.これらの種のうち,M.foetida, M.odorata , M.sylvaticaはM. indica同様,タイの一部の地域で栽培され,その地域では重要な果樹の1つとして市場で流通していた .その他の野生種についても, マンゴー栽培のための台木あるいは耐病・耐虫性品種の育成のための遺伝資源としてそれぞれの種が潜在的な価値を有しているもの

と思われた.本研究で存在を確認した13種について,55項目の形態的・生態的特性からその類縁性を検討したところ ,M.indica, M.sylvatica, M.oblongifolia, M.collina, M.cochinchinensisのグループとM .caloneura, M.pentandra, M.flava, M.macrocarpa, M.gedebe, M.griffithii, M.odorata, M.foetidaのグループの2つに大別することができた .タイの熱帯モンスーン地域に分布する種はすべて前者のグループに属した.後者のグループに属するほとんどの種は熱帯降雨林地域に自生していた .現在,熱帯林の崩壊が進んでいるが,将来の重要な遺伝資源であると思われるこれらマンゴー属植物の保存・維持は早急な課題であると考えられた. キーワードマンゴー属,形態的特性,系統分類,種の多様性

Islands8, 9). Introduction In Thailand, the most important species in The natural distribution of the genus the genus Mangifera is, needless to say, M. Mangifera is restricted to tropical Asia, indica (common mango). Total production stretching from northern India, Ceylon to amounted to 1.4 million metric tons in Thailand Micronesia. The highest number of species has in 1997, ranking third in the world next to India been found in Borneo, Sumatra, Java and Malay and China5). However, besides common mango, Peninsula. The findings obtained based on horse mango (M. foetida Lour.) and kiwini (M. morphological, phytogeographical, and cytologi- odorata Griff.) are very important species of cal studies indicate that the genus originated fruit crops in Thailand. Banana mango (M. from 2 regions; the monsoon climatic zone sylvatica Roxb.) and M. laurina Bl. are also covering Northeast India, Bangladesh, Myan- cultivated on a small scale in home gardens for mar, Thailand, and Indo-China, and the moist the local market in Thailand. tropical rain forest climatic zone covering the KOSTERMANSand BOMPARD8)described 69 Malay Peninsula, Borneo, and Lesser Sunda species in the genus Mangifera and classified Received June 4, 1998 them into two subgenera with several sections, Accepted Feb. 19, 1999 based on morphological characters. Although EIADTHONGet al.: Genetic Resources of Mangifera Species in Thailand 77

15 Mangifera species have been reported to the collection of specimens for identifying them. occur in Thailand by SMITINAND18),there is no Wild Mangifera species were identified by extensive survey on Mangifera species in comparing the specimens with herbarium Thailand. It is important to analyze the genetic materials maintained at the Herbarium of the resources of Mangifera species in Thailand, Royal Forest Department (BKF), Thailand. since some of them may have potential values for disease and/or pest resistance, or tolerance Phylogenetic analysis based on morpho- to adverse environmental conditions. During logical and ecological characters the period 1995-1997, we had the opportunity to Fifty-five characters (Appendix) were scored 0 survey Mangifera species in Thailand, including or 1 for each species to calculate the coefficient some cultivars of M. indica. The objectives of of JACCARD16).These characters were mainly the present report are 1) to clarify the selected from the report of RHODESet a1.15)and distribution of Mangifera species in Thailand, the lists of KOSTERMANSand BOMPARD8).Most of and 2) to evaluate the potential economic value the characters such as tree size, leaf arrange- of wild and cultivated Mangifera species. ment, and flower morphology were recorded in the field. Trees more than 30 years-old were Materials and Methods selected to score the characters. Then, the tree Germplasm exploration size was estimated by visual observation and Six surveys (3-7 days per survey) were several specimens were taken for averaging carried out during the period 1995-1997 in scores of some characters such as length of several areas of Thailand, where the cultivation of panicle, fruit diameter, fiber length and fruit mango or the presence of wild relatives has been skin hardness, if samples were available. reported8) (Fig. 1). These surveys were focused Mangifera indica cv. Carabao was used as a on the record of morphological characteristics, standard to determine the scores of the ecological habitat, and ethnobotanical informa- characters. However, if the specimens were not tion for the utilization of each species, as well as available for determining some characters, the characters were scored from references describ- ing the characteristics of the species4, 6,18). Phylogenetic analysis was performed by UPGMA cluster analysis with NTSYS program16). Two other genera of Anacardiaceae, Anacardium occidentale (cashew nut) and Bouea macrophylla (gandaria or plum mango), were also collected and scored as the reference of outgroup taxa for phylogenetic analysis. Results and Discussion Mangifera species in Thailand In this study, we identified 13 Mangifera species in Thailand (Table 1). SMITINAND18) reported that 15 Mangifera species occur in Thailand. However, he listed M duperreana Pierre in his report, which was classified by KOSTERMANSand BOMPARD8)as a synonym of M. caloneura Kurz. Therefore, we did not list this species in Table 1, as we followed the classification of KOSTERMANSand BOMPARD8).In addition, SMITINAND18)assigned M oblongifolia Hk.f. and M maingayi Hk.f. to M. odorata and M quadrifida Jack, respectively, although Fig. 1. Survey areas of Mangifera species in KOSTERMANSand BOMPARDB8)classifiedthese two Thailand. as different species. Although SMITINAND18)may 78 Jpn. J. Trop. Agr. 43 (2) 1999

Table 1. Mangifera species reported to be distributed in Thailand.

a) Classification of the species followed that of KOSTERMANSand BOMPARD8)b) SMITINAND18)had assigned it to Moborata Griff. c) (+/-) indicate (presence/absence) of the species. d) Species endemic in Thailand. have identified both M. oblongifolia and M. national parks due to the adequate conservation maingayi in his survey, we did not include these of the forests. Some provinces in the central and two species in his list in Table 1. In contrast, northeastern parts were included in our survey, KOSTERMANSand BOMPARD8)listed 14 Mangifera due to the presence of private gardens and species in Thailand, and MUKHERJEE11)reported germplasm collections of the genus Mangifera that only 8 species occurred in Thailand (Table (Fig. 1). Thailand is characterized mainly by 1). Including those in our survey, a total of 20 two climatic zones; 1) the monsoon climatic Mangifera species have been reported in zone, where the average annual rainfall is Thailand (Table 1). The existence of M. grifthii approximately 1,500 mm and the dry season is Hk.f, found in this survey in Narathiwat was a clearly distinct from the rainy season, and 2) the new record. moist tropical rain forest climatic zone which As for the 13 species collected in our prevails mainly in the southern part of country, survey, four species are under cultivation and where heavy and frequent rainfall is occasionally nine are wild species. Cultivated species include observed and the dry season is not clearly M. indica, M. foetida, M. odorata, and M. distinct from the rainy season. sylvatica. Mangifera indica is one of the major The natural distribution of M. caloneura, M. fruit crops in this country. Mangifera foetida or cochinchinensis Engler, M. collina Kosterm., M. horse mango is also a relatively important fruit oblongifolia and M. sylvatica is observed in the crop in the southern part of Thailand. Mangifera monsoon climatic zone, while M. flava Ervard, odorata and M. sylvatica are cultivated only on a M. foetida, M. gedebe Miq., M. grifthii, M. small scale in home gardens. macrocarpa Bl, and M. odorata are found in the In our survey, we selected areas mainly in moist tropical rain forest climatic zone (Table EIADTHONG et al.: Genetic Resources of Mangifera Species in Thailand 79 80 Jpn. J. Trop. Agr. 43 (2) 1999

Fig. 2. UPGMA cluster analysis based on 55 morphological and ecological characters in 13 Mangifera species and two outgroup taxa.

2). Mangifera odorata and M. sylvatica were from Indian origin. In fact, the species collected found in both climatic zones in our survey, while in the monsoon climatic zone in our survey all M. odorata in Nakon Ratchasima was sampled belonged to the monoembryonic seed type, from germplasm collections at Pak Chong although in some species it was difficult to Research Station of Kasetsart University and M. characterize the seed type due to the absence of sylvatica in Trang was collected from home fruit and/ or references (Table 2). gardens. The native climatic habitat of these M. collina, an endemic species to Thailand8), two species seems to be located in the moist was found and collected in the mountainous tropical rain forest climatic zone and in the areas of Chiang Mai in our survey. Some monsoon climatic zone, respectively. Only M. prominent horticultural characters of each indica and M. pentandra are distributed in both species are listed in Table 2 with the sites of climatic zones, although most cultivars of M collection. indica and some accessions of M pentandra give a higher yield in dry areas than in moist Species relationship in the genus Mangi- areas. fera collected in our survey Seeds with polyembryony among Mangifera UPGMA cluster analysis based on 55 species have been reported only in M. casturi morphological and ecological characters en- Kosterm.8), M. laurina Bl1, 8), M. grifflthii14) and abled to divide the 13 Mangifera species into M. odorata2, 3,14), except for M. indica19).Accord- two major groups (Fig. 2). The first group ing to BOMPARD2,3), M. odorata sometimes consisted of M. indica, M. sylvatica, M. harbours monoembryonic seeds, like M. indica. oblongifolia, M. collina and M. cochinchinensis, Mangifera griffithii also showed monoembryonic and the second group of M caloneura, M. seeds in our survey. It is well known that Indian pentandra, M. flava, M. macrocarpa, M. gedebe, cultivars of M. indica have monoembryonic M. griffithii, M. odorata, and M. foetida. All the seeds while Southeast Asian mango cultivars species in the first group were distributed in the have polyembryonic seeds7). The other Mangif- monsoon climatic zone. Also, the species era species which are adapted to a moist tropical belonging to the subgenus Mangifera and rain forest climate, might have polyembryonic section Mangifera based on the classification of seeds, like M. casturi, M. laurina, M. odorata, KOSTERMANSand BOMPARD8)were placed in the M. grilthii and M indica. MATHEWSand LITZ10) first group except for M flava. Three sub- noted that polyembryonic mango cultivars of M. clusters in the second group were also in indica from Southeast Asia evolved under agreement with the classification of KOSTERMANS constant disease pressure, and were more and BOMPARD8).The species belonging to the disease resistant than monoembryonic ones subgenus Mangifera and section Eurantherae EIADTHONGet al.: Genetic Resources of Mangifera Species in Thailand 81

(M. caloneura and M. pentandra) formed the References first sub-cluster, and M. odorata and M foetida, which both belong to the subgenus Limus and section Perrennis, formed the second sub- 1. BOMPARD, J. M. 1992a Mangifera laurina Bl., cluster. Mangifera gedebe and M. griffithii Mangifera pentandra Hooker f. In : PROSEA : Plant formed the third Sub-cluster, although they Resources of South-East Asia 2 (Edible fruits and nuts). (VERHEU, E. W. M. and R. E. CORONEL eds.) belong to different sections (section Marchandora Prosea Foundation (Bogor) 216-218. and Rawa, respectively) in the same subgenus 2. •\1992b Mangifera odorata Griff. In:

Mangifera. The last two species were distributed PROSEA: Plant Resources of South-East Asia 2 in peat swamp forest where strongly acid soils (Edible fruits and nuts). (VERHEIJ, E. W. M. and R. E. prevail. Only the fourth sub-cluster of M flava CORONEL eds.) Prosea Foundation, (Bogor) 218-220. and M. macrocarpa which belonged to different 3. •\1993 The genus Mangifera rediscovered: subgenera (Mangifera and Limus, respectively) the potential contribution of wild species to mango did not fit into the classification of KOSTERMANS cultivation. Acta Horticulturae 341: 69-77. and BOMPARD8). 4. •\and R. J. SCHNELL 1997 The mango: In addition, two outgroup taxa of Anacar- Taxonomy and systematics. In : The mango: Botany, dium occidentale (cashew nut) and Bouea production and uses. (LITZ, R. E. ed.) CAB macrophylla (gandaria or plum mango) were International (Wallingford) 21-47. 5. FAO 1998 FAOSTAT database 1997 (Mango). Web site placed in a separate position from 13 Mangifera at http://apps.fao.org/lim 500/nph-wrap.pl? Produc- species. MUKHERJEEand CHANDRA13)reported a tion. Crops. Primary&Domain=SUA&servlet=l tentative phylogeny of Anacardiaceae in which 6. Hou, D. 1978 Anacardiaceae. In: Flora Malesiana, the genus Bouea is the most primitive followed ser. 1, (van STEENIS, C. G. G. ed.). Sijthoff & by the genus Anacardium and Mangifera. Noordhoff International Publishers (Netherlands)

MUKHERJEE12)and SINGH17)reported that M. 423-440. sylvatica, M. pentandra and M. caloneura were 7. JAGTIANI, J., H. T CHAN and W. S. SAKAI 1988 Mango. closely related to M. indica compared to other In: Tropical fruit processing. Academic Press Mangifera species, and that the morphological (London) 45-104. characters of these three species were very 8. KOSTERMANS, A. J. G. H and J. M. BOMPARD 1993 The similar to those of M. indica. In our analysis mangoes : Their botany, nomenclature, horticulture based on morphological and ecological charac- and utilization. IBPGR, Academic Press (London) ters, M. indica was separated from M. caloneura pp. 233. 9. MAJUMDER, P. K. and D. K. SHARMA 1985 Mango. In: and M. pentandra, although M. sylvatica showed Fruit of India: Tropical and subtropical. (BOSE T. K. a relatively close relationship with M. indica. ed.) Naya Prokash (Calcutta) 5-123. Recently, new techniques based on molecular 10. MATHEWS, H. and R. E. LITZ 1992 Mango. In biology have become popular in phylogenetic Biotechnology of perennial fruit crops. (HAM- studies of the plant kingdom. We are currently MERSCHLAG, F A. and R. E. LITZ eds.) CAB examining the molecular systematics of Mang- International, (Wallingford) 433-448. ifera species for determining the more precise 11. MUKHERJEE, S. K. 1949 A monograph on the genus relationship of M. indica with other Mangifera Mangifera. Lloydia 22 : 73-136. species and obtaining some indications on the 12. •\1953 Origin, distribution and phylogenet- origin of M. indica. ics affinities of the species of Mangifera. J. Linn. Soc. London (Botany) 55: 65-83.

Acknowledgments 13. •\and D. CHANDRA 1983 An outline of the The authors are grateful to Associate Prof. revision of Indian Anacardiaceae. Bull. Bot. Surv. India 25 : 52-61. Chalongchai BABPRASERTHfor supplying the plant 14. NG, F S. P. 1991 Malayan Forest Record no. 34: materials of Mangifera indica and Mr. Tanit Manual of forest fruits, seeds and seedlings Vol.l. NUYIM for his assistance in the collection of Forest Research Institute Malaysia, Kepong, Kuala some Mangifera species. This research was Lumpur (Malaysia) 14-19. supported, in part, by a Grant-in-Aid for 15. RHODES, A. M., C. CAMPBELL, S. E. MALO, and S. G.

International Cooperative Research Project from CARMER 1970 A numerical taxonomic study of the Japan Society for the Promotion of Science. mango Mangifera indica L. J. Amer. Soc. Hort. Sci. 95: 252-256. 82 Jpn. J. Trop. Agr. 43 (2) 1999

16. ROHLF,E J. 1997 NTSYS-pc (Numerical Taxonomy 18. SMITINAND,T 1978 The genus Mangifera L. In: and Multivariate Analysis System), version 2.01. Mango (Department of Horticulture, ed.) Kasetsart Exeter Software (New York). University (Bangkok) 1-14. (In Thai) 17. SINGH, L. B. 1976 Mango: Mangifera indica 19. VERHEIJ,E. W. M. 1992 Mangifera indica L. In (Anacardiaceae). In: Evolution of crop plants. PROSEA : Plant Resources of South-East Asia 2 (SIMMONDS,NW. ed.) Longman (London) 7-9. (Edible fruits and nuts). (VERHEIJ,E. W. M. and R. E. CORONELeds.) Prosea Foundation, Bogor (Indone- sia) 211-216. EIADTHONGet al.: Genetic Resources of Mangifera Species in Thailand 83

Appendix Morphological and ecological characters scored for phylogenetic analysis.

1.Tree size (m) 0) Small to medium (<30) 1) Large (•†30) 2.Climatic habitat 0) Monsoon climate 1) Moist tropical rain forest climate 3.Distribution area O)Highland 1) Wetland 4.Buttress 0)None 1)Yes 5.Flower perfect 0)Andromonoecious 1) Other 6.Leaf arrangement 0)Alternate 1) Opposite decussate 7.Leaf shape 0)Lanceolate to oblong 1) Other 8.Leaf apex 0)Acute to acuminate 1)Round to emarginate 9.Leaf texture 0)Thin 1)Thick 10.Panicle habit 0)Sub-terminal to terminal 1)Axillary 11.Length of panicle 0)•† 30cm 1)<30cm 12.Panicle color 0)Yellowish green 1)Pink to red 13.Panicle texture 0) Glabrous 1) Glabrate to pubescent 14.Pentamerous flower 0)Yes 1)No 15.Tetramerous frower 0)Yes 1)No 16.Tetra-and pentamerous frower 0)Yes 1)No 17.Sepal and petal color 0) White to yellow 1) Pink to red 18.Sepal shape 0) Elliptic to ovate 1) Linear to lanceolate 19.Petal shape 0) Oblong to ovate 1) Linear to lanceolate 20.Number of fertile stamens 0)1 1) Other 21.Number of staminodes 0) 3-4 1) Other 22.Number of disc lobes 0) 5 1) Other 23.Anther shape 0) Ovoid 1) Ellipsoid 24.Anther type 0) Dorsifixed 1) Basifixed 25.Stamen unequal 0)Yes 1)No 26.Fruit shape 0) Usually oblong-elliptic 1) Round (nearly round) 27.Fruit type 0) Drupe 1) Nut 28.Fleshy receptacle 0)None 1)Yes 29.Fruit diameter(cm) 0) •†5 1)<5 30.Fruit skin color at maturity 0) Pale green to yellow, orange 1) Other (dark purple, red purple, etc.) 31.Presence of blush color on fruit skin 0)No 1)Yes 32.Relative skin thickness 0)Thin 1)Thick 33.Lenticel on fruit skin 0) None or small 1) Large 34.Lenticel color 0)White to green 1) Russet 35.Presence of pulp 0)Yes 1)No 36.Pulp color 0)White to pale yellow 1)Deep yellow to orange 37.Amount of fiber in pulp 0)None to scanty 1)Abundant 38.Fiber length 0) Short 1) Long 39.Relative fiber texture 0) Fine 1) Coarse 40.Aroma strength of fruit 0) None to weak 1) Strong 41.Fruit skin hardness at maturing stage 0) Soft 1) Firm 42.Amount of resin on fruit skin 0)Scanty 1)Abundant 43.Fruit taste; turpentine 0)No 1)Yes 44.Fruit taste; sweet 0)Yes 1)No 45.Relative seed size

per fruit size 0) Small 1) Big 46.Labyrinthine seed 0)No 1)Yes 47.Seed shape 0) Ovoid to oblong 1) Reniform 48.Testa character 0)Free from endocarp 1)Adherent to endocarp 49.Cotyledon color 0)White to pale pink 1) Blue-violet 50.Edible pulp 0)Yes 1)No 51.Edible endosperm 0)Yes 1)No 52.Endosperm shape 0)Straight 1)Reniform 53.Cotyledon character 0)Free 1)Folded or lobed 54.Germination type 0) Heliciopsis type 1) Sloanes type 55.Introduced species in South-East Asia 0)No 1)Yes