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Late Pleistocene-Holocene Amphibians Palaeontologia Electronica palaeo-electronica.org Late Pleistocene-Holocene amphibians from Okinawajima Island in the Ryukyu Archipelago, Japan: Reconfirmed faunal endemicity and the Holocene range collapse of forest-dwelling species Yasuyuki Nakamura and Hidetoshi Ota ABSTRACT Amphibian fossils excavated from two Late Pleistocene-Holocene fissures in the southern part of Okinawajima Island in the Ryukyu Archipelago were identified through detailed comparisons with skeletal specimens of extant taxa. The identified species (eight frogs and two newts) were confined to extant elements on the island and were mostly endemic to either the island or the central part of the archipelago including it. Previous Late Pleistocene records of extralimital frogs (two mainland Japanese spe- cies [Glandirana rugosa and Rana tagoi] and one Southeast Asian frog currently natu- ralized in the Ryukyus [Polypedates leucomystax]) from one of the study sites (the Minatogawa Fissure) are therefore considered misidentifications. This may also be the case with Fejervarya kawamurai, a synanthropic frog currently widely distributed in East Asia, including the Central Ryukyus. Of the species obtained from the Late Pleis- tocene deposits at these sites, five frogs (Limnonectes namiyei, Babina holsti, Odor- rana ishikawae, O. narina, and Rana ulma) are forest-dwellers currently confined to the northern forested area of Okinawajima. Additionally, none of these species were detected in the middle Holocene deposits from one of the sites, the Sashiki Fissure. Our findings indicate that the southern limestone area of Okinawajima in the Late Pleistocene harbored a dense and humid forest with diverse lotic habitats, similar to the northern forested area of the island at present, and that these environmental condi- tions were lost by the midpoint of the Holocene Period. Yasuyuki Nakamura. Tropical Biosphere Research Center, University of the Ryukyus, Senbaru 1, Nishihara, Okinawa 903-0213, Japan. [email protected] Hidetoshi Ota. Institute of Natural and Environmental Sciences, University of Hyogo, and the Museum of Nature and Human Activities, Yayoi-gaoka 6, Sanda, Hyogo 669-1546, Japan. [email protected] Keywords: Biogeography; frogs; limestone; newts; paleoenvironment PE Article Number: 18.1.1A Copyright: Palaeontological Association January 2015 Submission: 6 August 2014. Acceptance: 19 December 2014 Nakamura, Yasuyuki and Ota, Hidetoshi. 2015. Late Pleistocene-Holocene amphibians from Okinawajima Island in the Ryukyu Archipelago, Japan: Reconfirmed faunal endemicity and the Holocene range collapse of forest-dwelling species. Palaeontologia Electronica 18.1.1A: 1-26. palaeo-electronica.org/content/2015/935-okinawan-amphibians NAKAMURA & OTA: OKINAWAN AMPHIBIANS INTRODUCTION are currently strictly confined to the northern for- ested hilly area of the island is astonishing (Ota, Original anthropological excavation at the 2003). This is because the habitat requirements of Minatogawa Fissure on Okinawajima Island of the these frogs (dense forest for adults and shallow Ryukyu Archipelago from 1968 to 1974 resulted in streams under forest cover for juvenile growth; the discovery of a number of Late Pleistocene e.g., Utsunomiya et al., 1983) are quite incompati- human bones (Kaifu and Fujita, 2012; and litera- ble with the inherent properties of the Pleistocene ture cited therein) and numerous associated limestone substrate that predominates in the remains of other terrestrial vertebrates (e.g., Hase- southern part of Okinawajima — high porosity gawa, 1980; Kawamura, 1989; Matsuoka, 2000). (Maekado, 1989) and karstic topography, which The amphibian fossil assemblage was reported to can absorb surface freshwater habitats (Nishijima contain a total of 10 or 11 (if “Rhacophorus sp.” et al., 1974; Tachihara, 2003). In fact, the southern was counted) anurans and one newt species part of the island, particularly the limestone areas, (Hasegawa, 1980). However, the assemblage has lack such lotic habitats and therefore the four often been regarded as unusual because the pres- extant frogs and two newts in the region (Fejer- ence of some of the component species exhibited varya kawamurai, Ryukyu Narrow-mouthed Toad an obvious discrepancy with a biogeographers’ Microhyla okinavensis, Ryukyu Kajika Frog Buer- view of the archipelago and the intrinsic character- geria japonica, Okinawa Green Tree Frog Rhaco- istics of the Pleistocene limestone substrate of the phorus viridis viridis, Sword-tailed Newt Cynops southern part of the island, where the fissure is ensicauda, and Anderson’s Crocodile Newt Echi- located (e.g., Ota, 2003). notriton andersoni) are mostly lentic breeders. From the biogeographical viewpoint, the fossil Hasegawa (1980) left several questions unan- occurrence of two ranid species (Wrinkled Frog swered because he simply listed the taxa identified Glandirana rugosa and Tago’s Brown Frog Rana in the Minatogawa Fissure without providing infor- tagoi) known only from mainland Japan and adja- mation to verify his species identification. In addi- cent islets (Maeda and Matsui, 1999; Frost, 2014) tion, the lack of a follow-up study has prevented us remarkably contradict the highly endemic nature of from ascertaining the exact state of the Late Pleis- the current terrestrial vertebrate fauna of the Cen- tocene amphibian fauna of the island as well as its tral Ryukyus (i.e., the Amami and Okinawa Island biogeographical/paleoenvironmental significance. Groups and some nearby islands). This current Here we report on Late Quaternary amphibian fos- state unequivocally reflects the long-standing isola- sils recovered from two limestone fissures located tion of this region from nearby landmasses, partic- in the southern limestone area of Okinawajima. ularly the more northern areas of Japan (e.g., One of the sites is the Minatogawa Fissure, which Okada and Koba, 1931; Ota, 1998; Matsui, 2005; was recently re-excavated in an anthropological Motokawa, 2009). Indeed, the extant indigenous survey. Both of the limestone fissures yielded an amphibian fauna of the Central Ryukyus (11 out of abundance of Late Pleistocene amphibian fossils, 14 species are endemic) does not share its compo- and middle Holocene amphibian remains were dis- nent species with mainland Japan, except for the covered at one of the sites. These fossils are ade- Marsh Frog Fejervarya kawamurai (formerly known quate to document snapshots of Late Quaternary as Rana or Fejervarya limnocharis), a widely dis- local amphibian fauna and provide information tributed dicroglossid frog that ranges from Honshu about changes in the paleoenvironmental condi- (mainland Japan) through the Central Ryukyus and tions in the area. Taiwan to Central-South China (Djong et al., 2011). To explain the biogeographical peculiarity of the MATERIAL AND METHODS Central Ryukyu populations, some researchers argue that the species reached this location Geological Settings through relatively recent oversea dispersals from 2 the Eurasian Continent (Toda et al., 1997; Toda, Okinawajima (ca. 1200 km ) is a narrow 1999) (see Discussion). island oriented on a NE-SW axis with a maximum From a paleoenvironmental perspective, the length of 100 km (Figure 1). The island consists of fossil occurrence of five obligate forest-dwelling two geologically contrastive components: the frog species (Namiye’s Frog Limnonectes namiyei, northern part is comprised mainly of pre-Neogene Holst’s Frog Babina holsti, Ishikawa’s Frog Odor- strata (e.g., Fujita, 1989) and the southern part is rana ishikawae, Okinawa Tip-nosed Frog O. composed of the Plio-Pleistocene Shimajiri Group narina, and Ryukyu Brown Frog Rana ulma) that (siltstone associated with tuff), Pleistocene Chinen 2 PALAEO-ELECTRONICA.ORG FIGURE 1. Maps of the Ryukyu Archipelago (1, 2) and Okinawajima Island (3). The map of Okinawajima shows topography, distribution of the Pleistocene limestone, and study sites. Geological data were obtained from Kizaki (1985). Formation (calcareous sandstone and sandy lime- rhacophorine frog believed to have been intro- stone), and Pleistocene Ryukyu Group limestone duced to the Ryukyus in the mid-20th century (see (e.g., Sagae et al., 2012). The Ryukyu Group dis- Discussion); presumably this frog corresponds to tributed in the southern part is divided into three “Rhacophorus sp.” in Hasegawa (1980). Although sections: the Itoman Formation (Early Pleisto- we were unable to examine the amphibian fossils cene), Naha Formation (Early to Middle Pleisto- discussed by Hasegawa (1980) and Nokariya cene), and Minatogawa Formation (presumably (1983b), we investigated unsorted fossils collected Late Pleistocene) (see Sagae et al., 2012). in the recent (1998–2001) anthropological excava- tion at the fissure (see Hashimoto et al., 2002 for Study Sites details). These fossils were collected by water The amphibian fossils examined here were screening. Part of the fossils were reported to con- derived from two limestone fissures located in the tain one vertebra of Limnonectes namiyei, one southern part of Okinawajima Island (Figure 1). right ilium of Babina holsti, one vertebra of Odor- Minatogawa Fissure (26° 07' 46" N, 127° 45' 34" rana narina, 41 miscellaneous bones of unidenti- E; elevation: 20 m a.s.l). The Minatogawa Fissure fied anurans, and 79 precaudal/caudal vertebrae of in Nagamou, Yaese Town, is located on a bank of unidentified newts (Nohara and Irei, 2002). How- the Yuhi River (0.5 km from the mouth of the river), ever, we did not confirm these records except for
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