Preliminary study on phytogeography of Dipterocarpaceae Blume family in Vietnam

Vietnam Journal of Earth Sciences, 39(4), 337-344, DOI: 10.15625/0866-7187/39/4/10729 337 (VAST) Vietnam Academy of Science and Technology Vietnam Journal of Earth Sciences Preliminary study on phytogeography of Dipterocarpaceae Blume family in Vietnam Vu Anh Tai*, Pham The Vinh, Tran Thi Thuy Van, Le Thi Kim Thoa, Le Duc Hoang, Ngo Thi Bich Hong Institute of Geography (VAST) Received 11 July 2016. Accepted 07 August 2017 ABSTRACT Biogeographically mapping flora of Vietnam requires the studies on the distribution of some important species groups for identifying the typical species composition of each phytochorion. The Dipterocarpaceae family contains taxa originated in tropical Asia and its subfamily of Dipterocarpoideae is proved to have Southeast Asia origin. In Vietnam, this family includes 43 species in 7 genera. In this study, Dipterocarpaceae species from 645 sites in Vietnam are assessed and compared to those in over the world. In Vietnam, this family distributes in tropical and/or slightly passing to subtropical climate but none of its species is naturally found in the Red River and the Mekong River deltas. In the world, the Dipterocarpaceae species found in Vietnam concentrated distributes in Indochina flo- ristic region, corresponding to the originative area of South Myanmar. Statistically, there are 12 endemic species for the Indochinese floristic region and five of them are endemic for four provinces of this region related to Vietnam, re- spectively as follows: South China - 1, North Indochina - 1, South Indochina - 2 and Annam - 1. Additionally, some species distribute in East Asia floristic region of Holarctic Kingdom because of expanding the distribution area from the Indochinese floristic region. All genera of this family in Vietnam were originated in the Indochinese floristic re- gion. Moreover, the floristic data and phytogeographical phylogeny diagram, based on analyses of phytogeography and DNA, would be better to use for finding out the distributional source or the forming time of species or genus, then the phylogenetic diagram. Keywords: Dipterocarps family; Indochina floristic region; North Indochinese floristic province; Annamese floristic province; South Indochinese floristic province; Phytogeographical phylogeny; Endemic species. ©2017 Vietnam Academy of Science and Technology 1. Introduction1 In a purpose of biogeographically mapping Vietnam flora, the taxon distributions are re- quired to find out the typical species composi- tion for each area. Dipterocarps family (Dip- terocarpaceae Blume) (Angiosperm Phylogeny Group, 2009), with its taxon distributing in the *Corresponding author, Email: tai.botany@gmail.com Asia and its sub-family of Dipterocarpoideae, including 475 species of 13 genera, distrib- uting from Seychelles, Sri Lanka, India, Southeast Asia to New Guinea and most of Borneo islands was identified to be originated from Southeast Asia (Myanmar). In Vietnam, there are 43 species and 7 genera of Diptero- carps family, all of them belong to Dipterocar- poideae sub-family (Nguyen Hoang Nghia, 2005). The distributions of these genera were Vu Anh Tai, et al./Vietnam Journal of Earth Sciences 39 (2017) 338 defined as follows (Nguyen Hoang Nghia, 2005): Anisoptera from Bangladesh, Myanmar, to Indochina and New Guinea; Dipterocarpus in Sri-Lanka, India, Myanmar, Thailand, Indo- china, Malay peninsula, Sumatra, Borneo and Philippines; Vatica from Sri-Lanka, India, My- anmar, Thailand, Indochina, Malay peninsula to Hainan and New Guinea; Hopea in Sri- Lanka, India, Myanmar, Thailand, Indochina, Malesia and South of China; Shorea in Sri- Lanka, Myanmar, Thailand, Indochina and most of Malesia; Parashorea from Myanmar, South of China, Thailand, Indochina to Philip- pines and Borneo. Genetic relationships of Asian Dipterocarps species were analyzed in term of nuclear DNA and chloroplast DNA (Dayanandan et al., 1999). Based on DNA analysis, a genetic phylogeny diagram ex- pressing the relationships of the species in genera Anisoptera, Dipterocarpus and Vatica (including species in Vietnam) was intro- duced (Kress et al., 2003). However, there is an unequal between results of analysis of nu- clear DNA and chloroplast DNA as the position of species in phylogeny diagram (Figure 1). Figure 1. Diagram of floristic provinces in the Southeast Asia (Drawing base on Takhtajan (1986) and Averyanov et al., 2003) Takhtajan (1986) had established a global phytogeographical system with the basic unit of the floristic province. According to this system, Vietnam territory is situated in 2 kingdoms: Holarctic Kingdom (Eastern Asiat- ic region, Sikang-Yunnan floristic Province); and Paleotropical Kingdom (Indochina region with 4 floristic provinces as South China, North Indochina, Annam and South Indochi- na). However recent studies in Vietnam are some different from Takhtajan (1986), for ex- ample, the Asian phytogeographical elements Vietnam Journal of Earth Sciences, 39(3), 337-344 339 cover whole the Asia (Le Tran Chan et al., 1999) or the Eastern Asian elements are in all eastern part of Asia on the Pacific coast (Nguyen Nghia Thin, 2004). Both of those elements have been not identified in the Takhtajan’s system (1986); and moreover, the “phytogeographical element” conception has not been defined in this system. Most recent- ly, detailing floristic provinces related to Vietnam territory, Averyanov et al. (2003) have divided the Annamese floristic province in Takhtajan’s system (1986) into 2 sub- provinces: Southern Annam and Central An- nam. In that study, the species, especially en- demic ones, for each province were invento- ried but no species group in relation with the surrounding floristic provinces or regions was mentioned. Thus it is necessary to study phy- togeographical distributions of some taxa for gradually establishing a phytogeographical map of Vietnam flora. In addition, the study on distribution of species, genera could give an idea such as: generating and overlapping points of the distributions of various species or genera, points would be checked with fossil evidence. On the contrary, the taxa distrib- uting far from the overlapping point would be reputed as lately formed species. Based on these analyses, it would be established a phy- togeographical phylogeny diagram, one of the important keys for phytogeographically map- ping Vietnam flora. 2. Materials and Methodology This study is basically synthesized the pub- lications on Dipterocarps species existing in Vietnam and their distributional evidence in Vietnam and over the world, including online data of herbaria over the world and data pub- lished in Ashton P.S., 1982; Kress W.J. et al., 2003; Li X.W. et al., 2007; Nguyen Hoang Nghia, 2005; Nguyen Kim Dao, 2003; Ngu- yen Nga Phi, 2009; Pham Hoang Ho, 2001; Smitinand T.; 1969; 1990). The species are scientifically named by using The Plant List online version 1.1 (The Plant List, 2013). Distribution areas of the Dipterocarps taxa are defined by the interpolation method using 645 pieces of evidence (specimens and published information) of species found in Vietnam. The GIS tools are applied to develop distribution area of all subjects on the world floristic re- gion map (Takhtajan, 1986), in particular within Vietnam territory; the boundaries of floristic provinces are based on the results of Averyanov et al., 2003 and the work of Takhtajan (1986) is added for determining the boundary of Central/Southern Annamese sub- provinces. In this study, a method forming phytogeographical phylogeny diagram is sug- gested to regard the species in principle that the closely related species have close spatial distributions, but the result would only be en- sured by comparing and checking with the phylogenetic diagram. 3. Results 3.1. Distribution of Dipterocarps family in Vietnam The Dipterocarps species naturally distrib- ute in most of Vietnam, except the Red River delta and Mekong River delta. Most of these species are found in the tropical climate with an elevation below 700 m (in the North) or bellow 1000 m (in the South) a.s.l. (Thai Van Trung, 1978). Some species can develop in both tropical and sub-tropical climate with an elevation above 700 m in the North or 1000 m in the South, such as Hopea siamensis, Pen- tacme siamensis, Dipterocarpus obtusifolius and Shorea roxburghi, etc., while some others are present only in narrow band of 500-700 m high in the North or 600-1000 m in the South, belonging to cold tropical climate such as Dipterocarpus grandiflorus, Hopea ferrea, Shorea henryana, Shorea hypochra (Table 1). Vu Anh Tai, et al./Vietnam Journal of Earth Sciences 39 (2017) 340 Table 1. List of species of the Dipterocarps family in Vietnam No Scientific name Vietnamese name Elevation 1 Anisoptera costata Korth. Vên vên <700 2 Anisoptera scaphula (Roxb.) Pierre Kiền kiền nhẵn <800 3 Dipterocarpus alatus Roxb. ex G. Don Dầu rái 200-500 4 Dipterocarpus baudii Korth. Dầu lông 200-800 6 Dipterocarpus condorensis Pierre Dầu cát 200-500 5 Dipterocarpus costatus Gaertn.f. Dầu cát 600-1000 7 Dipterocarpus dyeri Pierre Dầu song nàng <800 8 Dipterocarpus grandiflorus Blanco Dầu giọt tía 500-700 9 Dipterocarpus hasseltii Blume Dầu rái <1000 10 Dipterocarpus intricatus Dyer Dầu trai 500-700 11a Dipterocarpus obtusifolius Teijsm. ex Miq. var. obtusifolius Dầu trà beng 500-1500 11b Dipterocarpus obtusifolius var. subnudus Ryan & Kerr Dầu song nàng < 1300 12 Dipterocarpus retusus Blume Chò nâu 800-1500 13a Dipterocarpus tuberculatus Roxb. var. tuberculatus Dầu đồng 800-1000 13b Dipterocarpus tuberculatus var. grandifolius (Teijsm. ex Miq.) Craib Dầu đồng lá to 800-1000 14 Dipterocarpus turbinatus Gaertn.f. Dầu con quay 30-1100 15 Hopea chinensis (Merr.) Hand.-Mazz. Táu vu 100-1100 16 Hopea cordata J. E. Vidal Sưng đắng 100-1100 17 Hopea ferrea Pierre Chò kiền kiền <700 18 Hopea hainanensis Merr. & Chun Sao hải nam 350-600 19 Hopea helferi (Dyer) Brandis Sao xanh <1000 20 Hopea odorata Roxb. Sao đen 100-750 21 Hopea pierrei Hance Kiền kiền phú quốc <1000 22 Hopea recopei Pierre Chò chay <700 23 Hopea reticulata Tardieu Sao mạng 200-600 24 Hopea siamensis Heim Kiền kiền <1200 25 Hopea sp. Sao đá <700 33 Parashorea chinensis H. Wang Chò chỉ 100-1000 27 Pentacme siamensis (Miq.) Kurz Cà chắc xanh 100-1500 28 Shorea falcata J. E. Vidal Sao lá lệch 100-500 29 Shorea guiso (Blanco) Blume Chăn <600 30 Shorea henryana Pierre Bô bô <900 31 Shorea hypochra Hance Vên vên 700 32 Shorea obtusa Wall. ex Blume Cà chắc 200-1000 26 Shorea roxburghii G. Don Sến cát >1300 34 Shorea stellata (Kurz) Dyer Chò đen <600 35 Shorea thorelii Pierre Trai 50-550 36 Vatica chevalieri (Gagnep.) Smitinand Táu muối 50-300 37 Vatica cinerea King Táu nước <900 38 Vatica diospyroides Symingt. Táu muối <900 39 Vatica mangachopoi Blanco Táu duyên hải <600 40a Vatica odorata (Griff.) Symingt. subsp. odorata Táu mật 300-900 40b Vatica odorata subsp. brevipetiolata Phamh. Táu lá nhỏ 300-800 41 Vatica pauciflora (Korth.) Blume Táu ít hoa <700 42 Vatica philastreana Pierre Làu táu nước <700 43 Vatica subglabra Merr Táu nước 100-900 3.2. Phytogeographically regioning the Dip- terocarps family of Vietnam Analyzing 645 present sites of the Diptero- carps family of Vietnam shows that the distri- bution area of all members of this family in Vietnam covers on 2 floristic kingdoms: Pale- otropis (4 floristic regions, 14 floristic provinc- es) and Holartis (2 floristic regions, 6 floristic provinces) (Table 2). The Dipterocarps of Vietnam, first concentratedly distributed in In- dochina province with 43 species and then Vietnam Journal of Earth Sciences, 39(3), 337-344 341 northwards expanded to Sikang - Yunnan with herein presence of Dipterocarpus retusus, D. turbinatus, Hopea chinensis, Parashorea chinensis (Li et al., 2007), Vatica diospyroides and V. subglabra (Nguyen Kim Dao, 2003) or westwards expanded to Sudano-Zambezian re- gion with the Dipterocarpus turbinatus (Nguyen Kim Dao, 2003) and was southwards limited in the Malesian region with 25 species. Among the floristic provinces, there are 5 provinces cover- ing all Vietnamese territory: Sikang - Yunnan belonging to Holartis, South China, North Indo- china, Annam and South Indochina belonging to the Indochinese region of Paleotropics. Table 2. Distribution of species of Dipterocarps family of Vietnam in floristic units No Kingdom Holarctis: 8 Paleotropis: 43 Region Eastern Asia: 6 (a) India: 9 Indochina: 43 Malesia: 25 (b)Floristic Province (1) (2) (3) (4) (5) (5) (7) (8) (9) (10) (11) (12) (13) (14) (15) (16) (17) (18) Number of species 1 2 2 6 1 5 5 21 5 7 31 33 35 31 25 4 7 6 11 1 1 Anisoptera costata + + + + + + + + + 2 A. scaphula + + + + + + + + 3 Dipterocarpus alatus + + + + + + + + + 4 D. baudii + + + + + + + 5 D. condorensis(*) + 6 D. costatus + + + + + + + + 7 D. dyeri + + + + + 8 D. grandiflorus + + + + + + + + + 9 D. hasseltii + + + + + + + + + 10 D. intricatus(**) + + + + 11 D. obtusifolius + + + + + + + 12 D. retusus + + + + + + + + + + + + + + + 13 D. tuberculatus + + + + + + 14 D. turbinatus + + + + + + + + + + + + + + + 15 Hopea chinensis + + + 16 H. cordata(*) + 17 H. ferrea + + + + + + 18 H. hainanensis + + + + 19 H. helferi + + + + + + 20 H. odorata + + + + + + + 21 H. pierrei + + + + + ? 22 H. recopei(**) + + + + 23 H. reticulata(**) + + +  24 H. siamensis(**) + + 25 Hopea sp. (*)(***) + 26 Parashorea chinensis + + + + 27 Pentacme siamensis + + + + + + 28 Shorea falcata(*) + 29 S. guiso + + + + + + + 30 S. henryana + + + + + + 31 S. hypochra + + + + 32 S. obtusa(**) + + + + + 33 S. roxburghii + + + + + + + 34 S. stellata + + + + + + 35 S. thorelii(**) + + + + + 36 Vatica chevalieri(*) + 37 V. cinerea + + + + + 38 V. diospyroides + + + 39 V. mangachapoi + + + + + + 40 V. odorata + + + + + + + 41 V. pauciflora + + + + + + + + + 42 V. philastreana(**) + + + 43 V. subglabra + + + + +   Vu Anh Tai, et al./Vietnam Journal of Earth Sciences 39 (2017) 342 Note: Floristic region: (a): Irano Turano, (b): Sudano-Zambezian; Floristic provinces: (1): Eastern Himalaya, (2): Khasi-Manipur, (3): Northern Burma, (4): Sikang-Yunnan, (5): Bengal, (6): Malabar, (7): Southern Burma, (8): Andaman, (9): South China, (10): Thailand, (11): North Indochina, (12): Annam, (12): South Indochina, (14): Malay, (15): Kalimanta, (16): Philippian, (17): Sumatra, (18): South Malesia; (*): Endemic for Vietnam; (**): Endemic for Indochina; (***): Hopea sp. (according to Nguyen Hoang Nghia, 2005) None of the Dipterocarps species found in Vietnam is endemic for any floristic province of the Holartis and there is a North - South trend separating these species. The appearance of Dipterocarps species in South China prov- ince seems to be result from a distributional expansion of those in Indochinese floristic re- gion. There are 5 endemic species for 3 prov- inces of the Indochinese floristic region: Dip- terocarpus condorensis for North Indochina; Hopea sp., Vatica chevalieri (Nguyen Hoang Nghia, 2005) for South China; Hopea cordata and Shorea falcata (Smitinand et al., 1990; Nguyen Hoang Nghia, 2005) for Annam; 7 other species are endemic to the Indochinese floristic region: Dipterocarpus intricatus, Hopea recopei, H. reticulata, H. siamensis, Shorea obtusa, S. thorelii and Vatica philas- treana (Smitinand et al., 1990). Based on the comparison between the dis- tributions of Dipterocarps species found in Vietnam and those in other areas, in the Indo- chinese floristic region, the 2 provinces of Thailand and Southern Burma have the close relationship with 3 Indochinese floristic prov- inces covering Vietnam (North Indochina, Annam and South Indochina). Moreover, Ma- lay floristic province in the Malesian floristic region also has the good relationship with the- se 3 provinces. To explain this phenomenon, there may be a reason directly connecting Ma- lay and Indochina Peninsulas. The other flo- ristic provinces of the Malesian and Indochi- nese regions have a small number of Diptero- carps species found in Vietnam. Therefore, together with Thailand, Eastern Burma and Malay, the 3 floristic provinces covering Vietnam as the North Indochina, Annam and South Indochina are core distribution area of the Dipterocarps family through the species discovered in Vietnam. 3.3. Phytogeographic phylogeny diagram of Dipterocarp family in Vietnam A phytogeographical distribution of Dip- terocarps genera found in Vietnam is formed by combining the distributions of their species (Figure 2). Comparing these data with the dis- tribution area of genera in Dipterocarps family of Vietnam published by Smitinand et al., 1990, shows that: Genus Anisoptera has 2 species limitedly distributing in Paleotropis, including the Indo- chinese and Malesian regions and a small part in the Indiana region (Bengal floristic prov- ince, adjacent to Indochinese region). Accord- ing to data of Nguyen Kim Dao (2003), it is new record of genus distribution in Indiana region with species of A. scaphula. Genus Dipterocarpus with 12 species has most distribution area of Dipterocarp family of Vietnam except Sudano-Zambezian region. Its distribution area expands to the Southern of the China, to the Holartis kingdom: Dipteorcarpus retusus distributed in Sikang - Yunnanese floristic province (Li X.W et al., 2007). Genus Hopea with 11 species has relative- ly wide distribution, from the centre of the In- dochinese floristic region: northwards to Sikang - Yunnanese floristic province of Holartis, mostly expanding in Vietnam and somewhere in Yunnan administrative prov- ince of the China; southwards to the Southern Malesian floristic provinces, excluding the Philippine, Sumatra and Kalimantan floristic provinces. In comparison with the data of Smitinand et al., 1990, there is no evidence Vietnam Journal of Earth Sciences, 39(3), 337-344 343 for distribution of this genus in the Indian region. Genus Parashorea has only one species distributing from Indochinese floristic region to Sikang - Yunan province of the Holartis kingdom and to Malay province in the Malesiana floristic region. The study of Smitinand et al., 1990) gave similar result. Genus Pentacme has only one species lim- itedly distributing within floristic provinces of Indochinese region and in Malay floristic province. This is also similar with the data of Smitinand et al., 1990 with the species Shorea siamensis (it is a synonym of Pentacme sia- mensis). Genus Shorea has 8 species, concentrated- ly distributing within the Indochinese and Malesian floristic region except the Kaliman- tan floristic province, expanded to the Indian- an floristic region at Malabar floristic prov- ince with one species as S. roxburghii; and there is no species discovered in the Holartis. This result corresponds to that of Smitinand et al., 1990. Genus Vatica with 8 species concentrated distributes within the Indochinese and Malesi- an floristic regions, expanding to Malabar province of Indian floristic region and to Sikang - Yunnan and Southeastern China provinces of Horiatis kingdom, but there is no evidence found in Kalimantan and Sumatra provinces of Malesian floristic regions. This result is also similar with the data of Smiti- nand et al., 1990. Figure 2. Phytogeographic phylogeny diagram of Dipterocarps family found in Vietnam 4. Conclusions Analyzing geographical distribution of the Dipterocarps family of Vietnam shows that family has concentrately distributed in Indo- chinese floristic region, concurring with origi- native centre in Burma (Myanmar). There are 5 endemic species for 4 floristic provinces, cov- ering whole Vietnam territory, belonging to Indochinese floristic region, while this region has total 12 endemic species of Dipterocarps of Vietnam. In addition, it also suggests that the species ancestors of Dipterocarps in Vietnam appeared in Holarctis due to the distributional Vu Anh Tai, et al./Vietnam Journal of Earth Sciences 39 (2017) 344 expansion from Indochinese floristic region. All genera had originated from the Indochinese floristic region. In comparison with data of Smitinand et al. (1990), the genera distribu- tions are firstly recorded in Indian floristic re- gion with Anisoptera, in the Sikang - Yun- nanese floristic province of Eastern Asian flo- ristic region with the Dipterocarpus. Combining phytogeographical species, genera data and genetic data enables to form the phytogeographical phylogeny diagram, reconstructing the evolution of the Diptero- carps found in Vietnam through the time with natural conditions such as geology, climate, geography, etc. In some particular cases, the phytogeographic phylogeny diagram has cer- tain advantages; it would be replaceable or even better than genetic phylogeny diagram. 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