NHBSS 051 1J Woodruff Thel
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NAT. NAT. HIST. BULL. SIAM Soc. 51( 1): 97-108 , 2003 THE LOCATION OF THE INDOCHINESE-SUNDAIC BIOGEOGRAPHIC TRANSITION IN PLANTS AND BIRDS David S. Woodrufl ABSTRA Cf Th e biogeographic transition between the In dochinese 組 d Sundaic (Indom a1 ay) bio 凶 1ies 1ies today on 出e 百lai-Malay peninsu1a. A literature review suggests that there may actu a1 1y be be two m 司jor phytogeographic transitions about 500 km ap 副. Most botanists follow V AN STEENIS STEENIS and GOOD and place 出 e transition at 血e Kang 釘 '-Pattani line near the Th ai-M a1 ay border. border. Others p1ace mo 陀 imp 口氏 ance on the transition between seasonal evergreen and mixed moist moist deciduous forests that occurs ne 訂 the Isthmus of Kr a, between Chumpon on the east coast coast and Tavoy on the west coas t. A recent revi 巴w of bird species distributions shows 白at a high1y high1y significant number of their species and subspecies r組 1ge 1imi 臼 occur just north of 血E Isthmus Isthmus of Kr a and are associated with the northern phytogeographic transition but not 白E southern southern one. 百 le transitions in other groups of anima1s are less well documented but gene ra1 1y 1ie 1ie north of Kr a in the northern peninsu1a. A review of 由e c1 irnato10gic a1 and p a1 eogeographic factors factors a1 1eged to co 口紅'01 血 ese transitions shows 伽 t we c創 mot adequate1y account for 自由 cu 町 ent positions.τ 'h e possible ro1e of 甘ans-peninsu1ar seaways as barriers permitting the divergence divergence of 白e In dochinese and Sundaic biotas is discussed and the directions for future research research to test various hypotheses 紅 e outlined. Keywords: Keywords: zoogeography ,phytogeography ,pa1 eogeography ,Te 昨iary seaways ,Tha i1 and ,Ma- 1aysia , Isthmus of Kra , Sunda1and INTRODUCTION Th e biotas of two different zoogeographic subregions and phytogeographic provinces meet in 官 lailand and 血is biogeographic provincial boundary begs bo 出 historical and biological biological expl 組 ation. ALFRED RUSSEL WALLACE (1 876) , the founder of zoogeography , recognized a major faunal transition in Thailand but the phenomenon has received little scientific scientific attention since then. He put the boundary between the northem Indochinese Subregion and the southem In do-Malayan Subregion in 出e Tenasserim Range at the northem end of 血, e Thai 司 Malay peninsula. Th e corresponding phytogeographic transition sep 訂 ates the the Continental SE Asiatic Fl oristic Pr ovince and the Malayan Fl oristic Province of the Ind o- Malayan Subkingdom (GOOD ,1964). 百出 transition is typically shown ne 紅 the Thai-Malay border 900 km further south. IEco10gy ,Behavior ,Evo1ution Section. Division of Bio10gica1 Sciences ,University of California , San Diego ,La Jolla Jolla CA 92093 ・0116 U.S.A. Address Address for editori a1 correspondence: Professor David S. Woodruff Division of Bio10gica1 Sci 巴nces ,University of of C a1 ifornia , San Diego ,La Jolla CA 92093 ・0116 U.S.A. Te1ephone: 1・858-534-2375 FAX: 1-858-534 ・7108 E-ma i1: [email protected] Received Received 31 October 2002; a∞ epted 7 May 2003. 97 97 BIOGEOGRAPHIC TRANSITION 99 PosmON OF THE BIOGEOGRAPHIC TRANSmON Although Although there is general agreement that a major biogeographic transition occ 町 s on the the Th ai-Malay pen 泊sula there 釘 'e diverse opinions 出 to its location and breadth. 百le position position of biotic 甘ansitions may be associated with present or former physical and environmental environmental barriers to dispersal ,and with the effects of dispersal , competition and invasibility ,and obligate co ・dis 凶bution (BROWN & LοMOL 刑 0 ,1998). 百lepu 中ose of 出is review review is to draw attention to 出e problems associated with 白is particular biogeographic transition transition and to 血us con 凶bute to their solution. A1 though Th ailand sits centrally 泊血e Oriental Zoogeographic Region the faunas of northem and southem 官 lailand are clearly different at 血e species leve l. Herein ,1 will use 出e terms In dochinese 組 d Sundaic for the northem and southem biotas ,respectively. 百le zoogeographic zoogeographic transition was placed at the northem end of the peninsula , at about 14 明 by W ALLACE (1 876). One hundred years later WELLS (1 976) provid 吋血 e frrst species level analysis analysis of the transition and , based primarily on 出e dis 位ibutions of Sundaic birds ,he concluded concluded that the transition lay just no 目h ofthe Isthmus of Kr a at 10 0 30'N. More recently , using using bird distributional records extending 合om Chang Rai to Singapore ,my colleagues and 1 have refined 出is result (HuG 田 S EJ AL. , 2003; ROUND EJ AL. , 2003). Our statistical analyses analyses revealed a significantωmover in bird species assemblages between 11 0 and 13 0 N on 血e northem 百lai-Malay peninsula. We found 血at 152 species , or half the forest- associated associated species ,have range limits in 血is area (Fig. 1). 0 町 fmding 血at the avian transition transition lies a little further no 巾 of 血e latiωde described by WELLS 泊 1976 is , in fact , what he would have predicted and has subsequently found (WELLS , 1999: xxi). Comparable documentation of the magnitude and location of the zoogeographic transition transition in other phyla are st 出 lacking but , as a broad generalization ,manlmals , amphibians and butterflies appear to similar exhibit pattems (LEKAGUL EJ AL. , 1970; CORBET & PENDLEBURY , 1992; INGER , 1999). PAUWELS EJ AL. (2002 , 2003) have surveyed reptiles in two provinces 200 km north and south of 血, e Is 血mus of Kr a,respectively. Th ey found 108 of of 152 snake species may have range limits in 血e northem peninsula but estimate 白紙出eir species species lists are only 55 -6 5% complete and 血ink 血e In dochinese-Sundaic 仕組sition lies furtherno 油田紅Kanchanaburi at about 14~. 百le geographic ranges of manlmals described by LEKAGUL & M cN EELY (1 988) and CORBET & HILL (1 992) show 伽.t numerous species and subspecies boundaries lie ne 釘 the Isthmus of Kr a. C HAl MANEE (1997) has analyzed 出e extant rodent fauna and found 血at , of 35 species of murines , 16 species 釘 'e re 抑制ed to 出, e no 叫 1 and 5 species to the south of the Isthmus; 白, e remaining 14 species occur in bo 血In dochinese and Sundaic subregions. For 16 species of sciurines and 10 species of petaurist 泊es ,白 e counts 釘 e 2:8:6 ,and 4:2:4 ,respectively. Of these 61 species of rodents , 61 61 % have range limits associated with the Isthmus of Kr a. Pleistocene fossils ,on the other hand ,show 血at individual species shi 負ed th 出 r加 ges 40 0- 500 km north and south of today's today's and 白紙出e early Pleistocene transition lay 500 km south of Kr a. TOUGARD (2001) reviewed reviewed the Pleistocene and recent distribution records of large mammals (1 29 extant sp 回 ies of primates ,c紅 nivores ,proboscidi 組 s,perissodactyls 佃 d artiodactyls) 泊 southeast Asia Asia and also concluded 出at ,al 出ough today's provincial boundary lies at 出e Isthmus of Kr a,血 e transition shifted further south to northem pen 泊sul 訂 Malaysia during Pleistocene hypothermals. hypothermals. Amphibian and butterfly species also show tumover at the isthmus; 泊 amphibians amphibians more In dochinese species extend south 出an Sundaic species extend north of 100 100 DA Vl D S. W ∞ORUFF the the transition (INGER ,1966) ,and the reverse is reported in butterflies (CORBET & PENDLEBURY , 1992). Th ere are app 釘 ently two phytogeographic transitions on 血e pen 凶sula of which the best best known lies near 血e 百凶ーMalay border ,500 km south of the Is 伽 nus of Kr a. 百lis southem 住'ansition ,between perhumid evergreen rainforest and wet seasonal evergreen r副 orest , is widely portrayed 部出em 吋or In dochinese-Sundaic plant boundary (v AN Sτ 宙開s, 1950; GOOD , 1964; KENG , 1970; WHITMORE , 1984; UDVARDY , 1969; ASHTON , 1992; 1992; R1 CHARDs , 1996; MORLEY , 2000). W natA MANAYA 阻 EJ' AL. (2002) also followed VAN S百 ENIS (1 950) in their regional conservation habitat-mapping project and put 出e boundary boundary between the In dochina Bioregion and the Sunda Shelf and Philippines Bioregion ne 紅白eτ 'h ai-Malay border. Th ere is also a second transition ,between se 部 onal evergreen rainforest rainforest and mixed moist deciduous forest , north of 由e Isthmus of Kr a. 百lis northem 住佃sition is widely shown on 1紅 'ge-scale vegetation maps but not fonnally documented 祖 師 litera 知re.T 盛岡・AANJAN (1 986) 叩 d COLLINS EJ' AL. (1991) pla 田 d 恥 boundary between the the In do ・Chinese and Malesian floristic regions at the Kra is 由mus. As birds are typically co-distributed co-distributed with pl 制 communities (MAcARn 町 R&MAcARn 町 R,196 1) the relationship between the avian 凶 nsition and the two ph: 戸ogeographic 回 nsitions is puzzl 泊g. Th e birds appe 紅 to ignore the southem transition and their relationship wi 白血e northem transition is is recognized but not yet well documented (HuG 田 S ET AL. , 2003; ROUND EJ' AL. , 2003). Our understanding of the position of the southem 住ansition has been modified several times times during the last cen 飽ry. R1D LEY (1 911a , b) was among 也 e frrst to note 白紙, on 也e west west coast , the flora of what was then called Lower Siam was quite different 仕om that of Malaya. Malaya. He con 回 sted the heath floras of Satun (百則 land) and nearby Pahang (Malaysia) and listed many Malay genera 出at are not found north of Al or Se 町 (Malaysia).