A New Species of Frog Allied to Fejervarya limnocharis from Title the Southern Ryukyus, Japan (Amphibia: Ranidae) Author(s) Matsui, Masafumi; Toda, Mamoru; Ota, Hidetoshi Citation Current Herpetology (2007), 26(2): 65-79 Issue Date 2007-12 URL http://hdl.handle.net/2433/216832 Right © 2007 by The Herpetological Society of Japan Type Journal Article Textversion publisher Kyoto University Current Herpetology 26(2): 65–79, December 2007 2007 by The Herpetological Society of Japan A New Species of Frog Allied to Fejervarya limnocharis from the Southern Ryukyus, Japan (Amphibia: Ranidae) MASAFUMI MATSUI1*, MAMORU TODA2, AND HIDETOSHI OTA3 1 Graduate School of Human and Environmental Studies, Kyoto University, Sakyo-ku, Kyoto 606–8501, JAPAN 2 Department of Zoology, Graduate School of Science, Kyoto University, Sakyo-ku, Kyoto 606–8502, JAPAN 3 Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 903–0213, JAPAN Abstract: The populations of a frog long identified as Fejervarya limnocharis from the Southern Ryukyus (=Sakishima in conventional regional name), Japan, considerably differ genetically and morphologically from the topotypic population of the species from Java. These Southern Ryukyu populations are therefore judged to represent a distinct biological species, which is described here as Fejervarya sakishimensis. This new species differs from F. limnocharis in larger snout-vent length (SVL). Also, it is distinguished from the latter in shorter head and tibia, smaller eye and narrower internarial space, all relative to SVL, and larger ratio of the first toe length to the inner metatarsal tubercle. From F. multistriata, F. sakishimensis differs by relatively larger tympanum, wider head, upper eyelid and anterior and posterior spaces of eyes, and longer forelimb and first toe, besides larger SVL. Furthermore, F. sakishimensis has a larger body, and relatively shorter head, tibia and hindlimb than F. iskandari. Also, this species is differentiated from all other nominate taxa of the F. limnocharis complex by a combination of some morphological characteristics. Key words: Cryptic species; Fejervarya limnocharis; Fejervarya sakishimensis; Ryukyu Archipelago; Yaeyama Islands INTRODUCTION China and Taiwan to southwestern Honshu of Japan mainland (Boulenger, 1920; Kampen, An Asian ranid frog Fejervarya limnocharis 1923; Inger, 1947, 1966). Recent studies, (Gravenhorst, 1829) had long been placed in however, revealed presence of quite a few the genus Rana, and considered as one of the distinct species under this name, and many commonest anurans with an unusually wide new taxa have been described chiefly from geographic distribution, from Sri Lanka and South Asia (Dubois, 1975, 1984; Dutta, 1997). India through Southeast Asia and continental Dutta (1997) called these frogs as the Limnon- ectes limnocharis complex, but the complex * Corresponding author. Tel: +81–75–753–6846; was more recently moved to the resurrected Fax: +81–75–753–2891; genus Fejervarya Bolkay, 1915 (Dubois and E-mail address: [email protected] Ohler, 2000). 66 Current Herpetol. 26(2) 2007 For Southeast and East Asian populations, Wenjiang populations was also not small taxonomic studies of this complex have been (D’=0.250: Toda et al., 1998a). In designat- retarded in comparison with those for the ing the neotype of F. limnocharis, Dubois and South Asian members. However, studies on Ohler (2000) also designated the neotype of systematic aspects other than taxonomy have F. multistriata (Hallowell, 1861) from Hongkong. been intensively made in this region. For Recent Chinese authors assign all Chinese example, genetic studies through allozyme populations of the F. limnocharis complex to electrophoresis made by Nishioka and Sumida this nominate species (Fei et al., 2002), but the (1990) and Toda et al. (1997, 1998a, b) situation is actually not so simple, because revealed the presence of substantial genetic Dubois and Ohler (2000) merely assigned a differentiations among populations within Japan, neotype and gave a description of a single type and among Japanese, Taiwanese, Chinese and specimen. Namely, status of F. multistriata as Southeast Asian populations of the F. limno- a good biological species has never been charis complex. assessed. Most notable finding was the presence of By contrast, the Southern Ryukyu popula- two syntopic, but genetically distinct, popula- tions, as represented by Ishigakijima sample in tions in Java, Indonesia, which is the type Toda et al. (1998a), compose a more compact locality of F. limnocharis (Toda et al., 1998a). entity, and its distinct taxonomic status from Subsequently, Dubois and Ohler (2000) desig- F. limnocharis from Java is less complicated. nated the neotype of F. limnocharis, and by It was Inger (1947) who reported that the comparison of this specimen with representa- population of Rana (now Fejervarya) limno- tive specimens of the two genetic groups in charis from the Ishigakijima Island differs Java, Veith et al. (2001) described F. iskandari morphologically from a population of Okinawa Veith, Kosuch, Ohler & Dubois, 2001 from Island, Central Ryukyus. Subsequent studies this island. This species, corresponding to confirmed and extended Inger’s (1947) view by “Malinping-B” of Toda et al. (1998a), is demonstrating great morphological, as well as remote from the other genetically more acoustic, differences between the Southern uniform populations from Southeast Asia, Ryukyu and the other populations of Japan with which the neotype of F. limnocharis was (i.e., populations from the Central Ryukyus morphologically associated. and Japan Mainland) (Kuramoto, 1979; Maeda Besides these two Southeast Asian forms, and Matsui, 1989). Extensive electrophoretic Toda et al. (1998a) recognized at least two surveys further revealed great differentiation more distinct genetic groups in the F. limno- of the Southern Ryukyu populations from the charis complex, one from China (Hongkong other Japanese populations with substantial and Wenjiang) and the other from Ishigak- genetic distances (Nei’s [1972] genetic distance ijima Island of the Yaeyama Group, Ryukyu [D]=0.276–0.345 in Nishioka and Sumida, Archipelago, Japan. Nevertheless, the taxo- 1990; D’=0.523–0.733 in Toda et al., 1997; nomic status of these two East Asian groups and D or D’ [not specified]=0.310–0.404 in remain uncertain. Sumida et al., 2007). In contrast, the popula- Considering a large genetic distance (Nei’s tions from Japan other than the Southern [1978] genetic distance [D’]=0.287 or 0.440) Ryukyus proved to be electrophoretically from F. limnocharis sensu stricto as defined much closer to some Chinese populations above (corresponding to “Malinping-A” of Toda (D=0.007–0.250: Toda et al., 1997), whose et al. [1998a]), each of these East Asian taxonomic assignment is not easy as noted groups seems to be taxonomically distinct above. from F. limnocharis. However, it is not easy Setting the taxonomic problems of popula- to classify the Chinese populations, because tions from the central Ryukyus, Japan Main- genetic differentiation between Hongkong and land and China aside, taxonomic position of MATSUI ET AL.—A NEW FEJERVARYA FROM RYUKYU 67 the Southern Ryukyu populations can be (FLL); 11) hindlimb length (HLL); 12) tibia determined by their direct comparisons with F. length (TL); 13) foot length (FL); 14) inner limnocharis from Java and other nominate metatarsal tubercle length (IMTL); 15) first species of the F. limnocharis complex. By toe length (1TL); 16) distance between ante- comparing the Southern Ryukyu populations rior corners of eyes (AED); and 17) distance with topotypic specimens of F. limnocharis, between posterior corners of eyes (PED). F. multistriata, and F. iskandari, we have Data for some specimens of the F. limno- confirmed its morphological distinctness from charis complex stored at the Museum National all these species. Combined with known d’Histoire Naturelle, Paris (MNHNP), Natu- genetic uniqueness (see above), the Southern ral History Museum, London (BM), Zoologis- Ryukyu populations can be regarded as a ches Museum, Universität Humboldt, Berlin distinct species of the F. limnocharis complex. (ZMB), and Forschungsinstitut und Naturmu- Maeda and Matsui (1999) already considered seum Senckenberg (SMF) were also incorpo- these populations as specifically distinct and rated. The system for description of toe- referred to them as Rana (Limnonectes) sp., webbing states is that used by Savage (1975). but formal description of the Southern Variation in adult SVL was examined by Ryukyu populations has never been made. analysis-of-variance (ANOVA) with the Tukey The purpose of this study is to provide a range test. For the other characters, we con- description of these unnamed populations. verted each value to a percentage ratio to SVL for comparisons. We first confirmed the MATERIALS AND METHODS absence of significant difference in each of those characters between sexes in Ishigakijima We examined a total of 77 preserved and Iriomotejima samples of F. sp. (both specimens of the F. limnocharis complex from entirely consisting of adults), and then com- East and Southeast Asia stored at the Gradu- bined data for both sexes for comparisons ate School of Human and Environmental among these and other samples. For taxa Studies, Kyoto University (KUHE), and other than the undescribed Southern Ryukyu Department of Zoology, Graduate School of species (F. sp.), subadult specimens were also Science, Kyoto University (KUZ): F. sp.