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Home , Achalinus, Ateuchosaurus, Ovophis okinavensis, Perochirus, Ryukyu Islands

TROPICS Vol. 10 (1): 5L-62 Issued May 30, 2000

The Curent Geographic Faunal Pattern of and Amphibians of the Ryukyu Archipelago and Adjacent Regions

Hidetoshi Ou Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 903-0213,

ABSTRACT The Ryukyu Archipelago accomrhodates 52 native /subspecies (26 genera belonging to ten families) of tenestrial reptiles and 22native species/subspecies (ten genera belonging to six families) of amphibians. This paper reviews the geographic similarity and diversity pattemJoflhe current herpetofauna of the Ryukyus and adjacent regions, and discusses factors influencing the formation of it in this archipelago. Cluster analyses of similarity indices for both reptiles and amphibians indicate that the Ryukyu herpetofauna is strongly influenced by geohistory ofthis archipelago involving long isolation especially in its north-cenhal part (i.e., southem Tokara, Amami, anA Oiinawa Groupsf Correlation analyses suggest that the faunal diversity on each island is significantly influenced by iis diversity as reflected by the island area (for reptiles) or height (amphibians).

Key words: herpetofauna / isolation / paleogeography / habitat diversity

The Ryukyu Archipelago is an assemblage of old continental islands located in the humid subtropical area of the western Pacific (Takara, 1962l' Kizaki, 1985: Fig. 1). This archipelago accommodates a diverse biota including various reptiles and amphibians (e.g., Ikehara & Kato, 1997). Several previous authors have analysed the Ryukyu herpetofauna (e.g., Inger, L947; Koba, 1962; Kuramoto, 1979) and argued the predominaace of Oriental elements (i.e., taxa having closest relatives in southeastern and ) over Palearctic elements (i.e., those most closely related to taxa in the main-islands of Japan and eastern China). Since then, a number of taxonomic changes have been made for the Ryukyu herpetofauna as a result of discoveries of new taxa, as well as of philosophical and methodological improvements of biological systematics [see Ota (2000) for example]. In this paper, I briefly review the geographic similarity and diversity patterns of the current herpetofauna of the Ryukyu Archipelago and adjacent regions by incorporating information recently added to our knowledge. I then discuss factors influencing the formation of that herpetofauna in this region. 52 H. OtA

Tokara GaP*- -}- rokara Group C \.o Amami GrouP ' Keramauapr -\ -nkinawa Group / t Senkaku \ ' I Group -; \ MiYako GrouP ' A \. "N YaeYamaGrouP tai{ant \ Fig. 1. Map of showing .b \ /. the location of the RYukYu \a Archipelago.

HERPETOEATJNAL DIVERSITY OF TIIE RYUKYU ARCHIPEI,AGO

located The native herpetofauna of the Ryukyus (here defined as an assemblage of continental islands (26 between the Tokara Tectonic Strait and the Strait) consists of 52 species/subspecies belongrng genera belonging to ten families) of terrestrial reptiles and 22 species/subspecies (10 genera to 68'4Vo and to six families) of amphibians (Tables 1 and 2). These numbers, respectively, correspond amphibians so far 36.7Vo ofthe total numbers of native tenestrial species/subspecies of reptiles and the fact known from the whole Japan. These values are surprisingly high, especially when considering that in area the Ryukyu Archipelago constitutes only about Llo of the whole Japan.

GEOGRAPHIC PAITERN OF EAUNAL SIMII,ARITY

pattern of Biogeographical subdivision of the Ryukyu Archipelago. Based on the geographic here could be endemism in the terrestrial reptiles and amphibians, the Ryukyu Archipelago as defined the divided into six areas (Hikida & Ota, 1997; Ota, 1997, 19982 note that these works incorporated Daito Group, an assemblage of purely oceanic islands southeast of the Ryukyu Arc, into the Ryukyu Archipelago): Area I, a few small islands of the Tokara Group south to the Tokara Strait; Area II' northern half of the Amami Group; Area III, the Okinawa Group, including a few islands "politically" Geographic faunal pattern of reptiles and amphibians of the Ryukyus 53

Table 1. Terrestrial reptiles of Japan. Their occurrences in and (southeastern continental China) are also indicated. Data are taken from Hikida & Ota (L997) and Ota (1998), with slight modifications as indicated in foot-notes. Asterisks (*) denote populations of artificial origins. Abbreviations are: MJ, the Main-islands of Japan and adjacent islands (including the lzu Group and the Tsushima Islands); TK, Tokara Group south of the Tokara Tectonic Strait; AM, Amami Group; OK, Okinawa Group; MY, Miyako Group; yy, yaeyama Group; SE, Senkaku Group; DA, Daito Group; oG, ogasawara and Iou Groups; TW, Taiwan; FJ, Fujian (the province of continental China nearest to Taiwan).

Ryukyu Archipelago MJ TK AM OK MY YY SE DA OG TW FJ TESTUDINES Bataguridae Chinemys reevesii +l +* C i s to c le mmy s fl av om a r ginata Ci. f. evelynae Geoemyda japonica Mauremys japonica +l M. mutica M. m. mutica +* M. m. kami +* +* +* Emydidae Trachemys scripta 2" s. elegans +*l +* +* +* Trionychidae Pelodiscus sinensis +1 +* +* +* +* LACERTILIA Agamidae Japalura polygonata J. p. polygonata J. p. ishigakiensis Eublepharidae Goniur osaurus kur oiwae G. k. kuroiwae + G. k. orientalis + G. k. splendens G. k. toyamai + G. k. yamashinge + Gehyra mutilata Gekko japonicus +1 Gk. hokouensis +1 + + Gk. tawaensis +l Gk. yakuensis +l H emidacty lus bow rin gii + + + Hd. frenatus +* +* +* +* +* +* +* H emiphy llodacty lus typus +* +* +{' Lepidodacty lus lugubris +* +{' +* + +{' + Iguanidae Anolis carolinensis +* +* Lacertidae Lacerta vivipara 54 H. Ora

Ryukyu Archipelago MJ TK AM OK MY YY SE DA OG T'TW FJ Takydromus amurensis T. dorsalis T. smaragdinus T. tachydromoides +1 T. toyamai Scincidae At euchos aurus p ellop leurus +t C ry p tob I epharu s b outonii C. b. nigropunctatus Eumeces barbouri E. elegans E. kishinouyei E. latiscutatus +t E. marginatus E. okadae E. stimpsonii Emoia atrocostata Scincella boettgeri S. formosensis +2 S. vandenburghi +

SERPENTES Typhlopidae Ramphotyphlops br aminus +*l +* Achalinus formosanus A. f. chigirai A. spinalis +t A. werneri Amphiesma concelarum Am. ishigakiense Am. pryeri Am. vibakari Am. v. vibakari +1 Am. v. danioense + Calamaria Pavimentata C. p. miyarai C. pfefferi Cyclophiops herminae Cy. semicarinatus Dinodon orientale +t D. rufozonatum D. r. rufozonatum D. r. walli D. semicarinatum Elaphe carinata E. c. carinata E. c. yonaguniensis E. climacophora +t E. conspicillata +t E. quadrivirgata +1 E. taeniura E. t. friesi +* E. t. schmackeri Geographic faunal pattern of reptiles and amphibians of the Ryukyus 55

Ryukyu Archipelago MJ TK AM OK MY YY SE DA OG TW FJ Lycodon ruhstrati L. r. multifasciatus Opi sthot r opis kikuzatoi Pareas iwasakii R. t. tigrinus +l Elapidae Hemibungarus j aponicus H. j. japonicus H. j. boettgeri H. j. subsp. H. macclellandi H. m. iwasakii A gki s tr o do n b I omhoffii +1 A. tsushimaensis + Trimeresurus elegans +* T. flavoviridis + T. mucrosquamatus +{' T. tokarensis okinavensis + 1, Distributed in Kyushu (including the Osumi Group), 2, S.-L. Chen, unpublished data.

regarded as the southern half of the Amami Group; Area IV, the Miyako Group; Area V the yaeyama Group; and Area VI, the Senkaku Group. Of these, Area I accommodates seven species of terrestrial reptiles, including one endemic (Trimeresurus tokarensis), and only a single amphibian species (Buergeria japonica). In Area II, L9 species of terrestrial reptiles and 12 species of amphibians are distributed, of which two reptiles (Goniurosaurw karoiwae splendens and Hemibungarus japonicus japonicus) and three amphibians (Rana amamiensis, Babina subaspera, and Rhacophorus viridis amamiensis\ are endemic to this region. Area III accommodates 3L terrestrial reptiles including six endemic species/subspecies (Geoemyda iaponica, Goniurosaurus k kuroiwae, G. k orientalis, G. k yamashinae, G. k toyamai, and Opisthotropis kifuzatoi), and 15 amphibians including four endemic species/subsp ecies (Rana narina, R. namiyei, B. holsti, and Rhacophorus v. viridis). Of the species/subspecies not strictly endemic to the Area III, 1.1. reptiles (Japalura polygonata polygonata, Ateuchosaurus pellopleurus, Eumeces barbouri, Talcydromus smaragdinus, Dinodon semicarinatwn, Cyclophiops semicarinatus, Amphiesma pryeri, Achalinus wemeri, Hemibungarus japonicus boettgeri, Trimeresurus flavoviridis, and Ovophis okinavensis) and five amphibians (Cynops ensicauda, Tllototriton andersoni, Hyla hallowellii, Rana okinavana, and R ishikawae) are exclusively shared with Area II, or Areas I and IL In Area Iv 19 terrestrial reptiles including three endemic species (Tatqtdronus toyanai, Calatnaria pfefferi, and Amphiesma concelarum) and three amphibians including one endemic subspecies (Bufo gargarizans miyakonis) are distributed. Area V accommodates 27 species/subspecies of terrestrial reptiles and L0 species of amphibians, of which lL reptiles (Cistoclemmys flavomarginata evelynae, Mauremys mutica kami, Eumeces stimpsonii, Talrydromus dorsalis, Calatnaria pavimentata miyarai, Pareas iwasakii, Etaphe carinata yonaguniensii, Amphiesma 56 H. Ore

Table 2. Amphibians of lapan. Their occurrences in Taiwan and Fujian (southeastern continental China) are also indicated. Data were taken from Kuramoto (1996: for Japanese urodelans), Maeda & Matsui (1999: for Japanese anurans), and Zhao & Adler (1993: for Taiwanese and Fijian species), with modifications as indicated in foot-notes. Asterisks (*) denote populations of artificial origins. Abbreviations are: MJ, the Main-islands of Japan and adjacent islands (including the Izu Group and the Tsushima Islands); TK, Tokara Group south of the Tokara Tectonic Strait; AM, Amami Group; OK, Okinawa Group; MY, Miyako Group; YY, Yaeyama Group; SE, Senkaku Group; DA, Daito Group; OG, Ogasawara and Iou Groups; TW, Taiwan; FJ, Fujian (the province of continental China nearest to Taiwan). Ryukyu Archipelago MJ TK AM OK MY YY SE DA OG TW FJ URODELA Hynobiidae Hynobius abei + H. boulengeri +1 H. dunni +l H. hidamontanus + H. kimurae + H. lichenatus + H. naevius +l H. nebulosus +l H. nigrescens + H, okiensis + H. retardatus + H. stejnegeri +l H. takedai + H. tenuis + H. tokyoensis + H. tsuensis + Ony cho dacty lus i aPonicus + S alamandr e I la key s e r lin gii + Cryptobranchidae Andrias iaponicus +1 Salamandridae Cynops ensicauda C. pyrrhogaster +1 Ty lototriton ander s oni ANURA Bufonidae Bufo gargarizans B. g. miyakonis +* +* B. japonicus B. j. iaponicus +l B. i. formosus + B. marinus +* +* +* B. torrenticola Hylidae Hyla hallowellii H. japonica +1 Ranidae Babina holsti B. subaspera Geographic faunal pattern of reptiles and amphibians of the Ryukyus 57

Ryukyu Archipelago MJ TK AM OK MY YY SE DA OG TW FJ Rana amamiensis + R. catesbeiana +* +* +* +* +* R. dybowskii + R, ishikawae + + R. japonica +t R. limnocharis +t + + +* + + R. narina + R. namiyei + R. nigromaculata +r + R. okinavana + + R. ornativentris +l R. pirica + R. porosa R. p. porosa + R. p. brevipoda + R. psaltes + + R. rugosa +1 R. sakuraii + R, supranarina a R. tagoi R. t. tagoi +r R. t, yakushimensis +1 R. tsushimensis + R. utsunomiyaorum + R.sP. + + Rhacophoridae Buergeria buergeri +t B.japonica + + + + + Chrixalus effingeri + + Polypedates leucomystax +. Rhacophorus arboreus + R, owstoni + R. schlegelii +t R. viridis R. v. viridis + R. v. amamiensis + Microhylidae Microhylaornata + + + + + + I, Distributed in Kyushu (including the Osumi Group). ishigakiense' Achalinus fornosanus chigirai, Hemibungarus macclellandi iwasakii, and Trimeresurus elegans) and three amphibians (Rana supranarina, R utsunomiyaorwn, and Rhacophorus owstoni) are endemic to this area. Of the remainder, seven species/subspecies of reptiles (Japalura polygonata ishigakiensis, Eumeces kishinouyei, Scincella boettgeri, Elaphe taeniura schmackeri, Dinodon rufozonatunt walli, Lycodon ruhstrati multifasciatus, and Cyclophiops herminae) and one undescribed Rana (toda et al., 1997; Maeda and Matsui, 1999) are exclusively shared with Area IV. Area VI accommodates six terrestrial reptiles, all common to Taiwan and mostly to continental China. Of these, only two are also shared with other parts of the Ryukyu Archipelago. No amphibians are known fromArea VI. 58 H. Ore

AMPHI BIA REPTILIA

I I 0.0 0.5

Fig.2. UpGMA (Sneath & Sokal, 1973) clusters on the basis of pairwise values of Nomura'Simpson's similarity index for amphibian fauna (left) and fauno (right) in the Ryukyu Archipelago and a{iacent regions.

Faunal shnilarities cmong geographic units recognized in the Ryulcyus and adiacent regions. Analyses using pairwise values of the Nomura-simpson's similarity index (the number of species/subspecies common to two communities in comparison divided by the total species/subspecies number in the smaller community) for indigenous terrestrial reptile fauna and amphibian fauna yielded clusters that are almost identical in topology (Fig. 2). In reptiles, the faunal similarity is greatest between Area VI and Taiwan, followed by Areas I, II and III, and then by Areas IV and V' The Area Vl-Taiwan cluster is first connected with a cluster of southeastertr continental China, and then with the Areas IV-V cluster and the Areas I-I[ cluster in order. Kyushu is most distant from all other geographic units considered in the analysis. The cluster for amphibians, excluding Areas I and VI due to the scarcity or complete absence of representatives (fable 2), shows identical topology with than the reptile cluster except for the initial connection of Taiwan with the Areas IV-V cluster rather with southeastern continental China. Branches in the cluster for amphibians are mostly longer than corresponding branches in the cluster for reptiles.

EACTORS INFLUENCING THE ALPHA DIVERSITY OF REPTILES AND AMPHIBIANS ON ISI.AI\TDS OF THE RYUKYUS

In the theory of island biogeography (MacArthur & Wilson, 1967), the alphd divetsity (i'e', the number of species/subspecies) of fauna on a given island is generally expected to be a function of size and distance from the faunal source of the island. There is, however, apparently little doubt that the Geographic faunal pattern of reptiles and amphibians of the Ryukyus 59

B

o g d C' g C' # ot r- o rts L o o L .o oo J z z5

Area (kma) Hclght (m)

Fig.3. Relationshlps between the number of species/subspecies, and island area 16,; aa6 island height @) in rcptiles (circles) and amphibians (triangles) of the Ryukyu Anchipelago.

diversity of reptiles and amphibians within the Ryukyus does not sirnply correlate with the distance from their hypothetical sources (southeastern continent for most lineages, and Kyushu and eastern cotrtinent for the remainder: Hikida & Ota, 1997; Ota, 1998). I thus limit my investigations here to the influence of island size, as expressed by area ltmz; ana maximum height (m), on the faunal diversity. Tlvelve islands, for which the herpetofauna has been well documented on the basis of field suryeys, were selected from the northern, central, and southern Ryukyus. These are: of the Tokara Group, Amamioshima, and Okierabujima of the Amami Group, Okinawajima, Tokashikijima and Kumejima of the Okinawa Group, Miyakojima of the Miyako Group, Ishigakijima, kiomotejima and Yonagunijima of the Yaeyama Group, and Uotsurijima of the Senkaku Group. Areas and maximum heights of these islands are taken from Takara (1962). Analysis was carried out to test the significance of correlation between the number of species/subspecies and each island dimension, both as log-transformed.

In reptiles, the number of species/subspecies (Y) is significantly conelated with the island area (X) (Y = OJ6TX + 0.783: r = 0.7'1.6, t = 3.24, P < 0.01), whereas it shows no correlation with the island height (r = 0.412, t = 1.43, P > 0.05). The number of amphibian species/subspecies is significantly correlated with both island area (Y = 0.189X + 0.4O2) and heighr (y = 0.896X - l.4g'). Nevertheless, the significance level is much higher with height (r = 0.896, t = 5.33, p < 0.001) than with area (r = 0.603, t=2.OO, P < 0.05) (Fig.3).

GENERAL DISCUSSION

Results of cluster analyses of faunal similarity in reptiles and amphibians largely confirm a geographic pattern assumed by some previous authors (e.g., Koba, 1962; Kuramoto, 1979; Hikida er al., 1989; Hikida & ota, 1997; ota, 1997). such a pattern is also analogous with the phylogeographical pattern illustrated by a number of reptile and amphibian lineages, which could be H. OtA summarized as: (((Area VI, Taiwan, eastern continent)(Area IV, Area V))(Atea I, Area II, Area III))Kyushu. The latter is considered to reflect a spatiotemporal pattern of vicariance in the Ryukyus and adjacent regions, which involves the effective geographic isolation first for Kyushu, and then for Areas I-III, Areas IV-Y and Area Vl-Taiwan-continental China in order (Hikida & Ota, L997; Ota, 1998; Hikida & Motokawa, 1.999; Toda et al., 1999; Yasukawa & Ota, 1999). Resuls of the present faunal analyses indicate that such paleogeographical events have significantly influenced the formation of the cunent faunal pattern. Each branch in the amphibian cluster is longer than the conesponding branch in the reptile cluster. This indicates a lesser similarity in the amphibian fauna between the areas designated above, and suggests that the straits separating those areas have functioned as isolation barriers more effectively for amphibians than for reptiles (but see Toda et al. ll99711 for an example of extensive oversea dispersals by afrogRana limnocharis in the Ryukyus). Results of correlation analysis indicate that in reptiles alplw diversity on an island of the Ryukyus largely depends on its area, but not on its maximum height. This is concordant with results of a few other studies dealing with the reptile fauna in other island groups (e.g., Perry et al',1998i Woinarski et a1.,I999), suggesting that area is a good predictor for the diversity of available to reptiles on an island. On the other hand, present results indicate a stronger correlation of the number of amphibian species/subspecies with maximum height rather than with area of an island. This predicts a stronger association of the maximum height with the diversity of habitats available to amphibians on an island. Apparent greater prevalence on lower islands of the Ryukyu limestone bed (Kizaki' 1985), which easily absorbs the surface water and thus possibly reduces the diversity of inland water habitats (Toyama, t976; Ota, 1983), may explain such an association. Further analyses using data for more islands and additional parameters reflecting the habitat diversity (e.g., vegetation complexity:Petry et al.,1998\ are strongly desired to verify these assumptions.

ACKI\OWLEDGMENTS I wish to express my sincere thanks to Tsutomu Hikida, Masafumi Matsui, Mamoru Toda, Junko Motokawa, Yuichirou Yasukawa, and many other colleagues in Kyoto University and the University of the Ryukyus for stimulating discussions and for provision of pertinent literature, and to Hiroyuki Otsuka, Mitsuru Hotta, and other members of the executive committee for inviting me to the International Symposium, "The Ryukyu Islands: Arena of Adaptive Radiation and Extinction of Island Fauna" (6-7 November 1998, at University), in which I gave a major part of this paper. I am also much indebted to Szu-Lung Chen for his permission to refer to the results of his study on the East Asian Scincellc in advance of their publication, to T. Hikida for helping with preparation of a map of the Ryukyus for Fig. 1, and to H. M. Smith and an anonymous reviewer for critical reading of the manuscript. This research was partially supported by a grant from the Fujiwara Natural History Foundation, Grants-in-Aid from the Japan Ministry of Education, Science, Sports and Culture (C-09839024 and 1183301.3 ro H. Ota), and U.S. National Geographic Society grant (No. 4505-91 to M. Matsui). Geographic faunal pattern of reptiles and amphibians of the Ryukyus 6l

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