Coll. and Res. (2015) 28: 5-53 5 Anuran Fauna of Taiwan and Adjacent Islands Based on Valid Specimen Records Nian-Hong Jang-Liaw1 and Wen-Hao Chou2* 1 Animal Department, Taipei Zoo, 30 Xinguang Rd., Sec. 2, Taipei 11656, Taiwan 2 Department of Biology, National Museum of Natural Science, 1 Kuang-Chien Rd., Taichung 40453, Taiwan (Received June 3, 2015; Accepted June 25, 2015; Published online September 25, 2015) Abstract. A checklist of all species of the order Anura from Taiwan and adjacent islands is provided. In total, distribution data of 33 anuran species are given including three introduced species. The inland distribution information provided is based on specimen collections of four institutes in Taiwan: the Biodiversity Research Center, Academia Sinica, Taipei; National Museum of Natural Science, Taichung; National Taiwan Museum, Taipei; and National Taiwan University Museum, Taipei. Phylogeographic studies of these anurans are also noted if references are available. Keywords: Anura, checklist, distribution, specimen records, Taiwan. INTRODUCTION anuran fauna of Taiwan prior to 2006: Stejneger (1907, 1910), Horikawa (1931), Okada (1934), Taiwan is an island located off the coast of Chen (1969), Lue and Chen (1982), Zhao and southeastern China at latitudes of 21°53'~25°17'N Adler (1993), Chou and Lin (1997b), Yang with tropical to alpine climate zones and high (1998, 2006), and Lue et al. (1999). With the biodiversity (Shao 2006; Kier et al. 2009). On this wide use of different investigative tools, such island, mountainous areas account for about 70% as molecular technology, and good international of the land surface. There are two main mountain communication on anuran research, several ranges, the Central Mountain Range which taxonomic debates on Taiwanese anurans have stretches nearly the entire length of the island, been raised and discussed (Ye and Fei 1994; Chou with elevations exceeding 3000 m (the tallest and Lin 1997a; Tanaka-Ueno et al. 1998; Liu et peak is Jade Mountain at 3952 m), and the Coastal al. 2000; Fu et al. 2005; Che et al. 2007; Jang- Range with a maximum elevation exceeding 1600 Liaw and Lee 2009; Liu et al. 2010; Jang-Liaw m. These mountain ranges and freshwater river and Chou 2011). systems that radiate from them divide Taiwan into Thirty-three species are currently recognized several geographical regions and microclimatic from Taiwan, including 30 native and three areas, providing an excellent opportunity for introduced species. Five of the Taiwanese biogeographical research. native frogs were described in 1978~1995: Anurans, commonly known as toads and Kurixalus idiootocus (Kuramoto and Wang, frogs, are a conspicuous component of the 1987), Rhacophorus arvalis Lue, Lai and world’s vertebrate fauna. Early faunistic studies Chen, 1995, Rha. aurantiventris Lue, Lai and on Taiwanese anurans date from Günther (1864), Chen, 1994, Rha. prasinatus Mou, Risch and followed by several studies that reviewed the Lue, 1983, and Rha. taipeianus Liang and *Corresponding author. E-mail: [email protected] 6 Coll. and Res. (2015) 28: 5-53 Wang, 1978. Furthermore, the first record of In an effort to preserve the biodiversity of the Fejervarya cancrivora (Gravenhorst, 1829) is island, especially some endemic and endangered thought to date to 2005 (Yang 2006). On the species, the Taiwanese government enacted other hand, Rana taiwaniana Otsu, 1973 and the Wildlife Conservation Act in 1989. It is a R. multidenticulata Chou and Lin, 1997 are national policy and differs from the Convention now respectively considered to be synonyms of on International Trade in Endangered Species Odorrana swinhoana (Boulenger, 1903) and R. of Wild Fauna and Flora (CITES); however, sauteri Boulenger, 1909 (Matsui 2005; Jang-Liaw all Appendix I fauna and most Appendix II and Lee 2009; Hsu et al. 2011). In this work, valid fauna listed in the CITES are protected by scientific names basically refer to Frost et al. this law (Hsu and Agoramoorthy 1997). The (2006) and Frost (2014). Wildlife Conservation Act has undergone a Few studies previously investigated series of amendments from 1994 to the present. biogeographic patterns among Taiwanese anurans. Accompanying this act, a “Schedule of Protected Lue et al. (1990) and Lee et al. (1994) used Species” was promulgated in August 1989, which geographic information system data to describe catalogued three classifications of protected the biogeographic distributions of all anurans species: I, endangered species; II, rare and in Taiwan. Their results, however, provided an valuable species; and III, other conservation- incomplete picture of the Taiwanese anuran fauna deserving wildlife. This list has also been because of insufficient taxonomic references modified several times, and 11 anuran species at that time. For example, at that time, Taiwan were included in December 1995: Microhyla frogs of the genus Limnonectes were assigned to butleri Boulenger, 1900, Microhyla heymonsi L. kuhlii (Tschudi, 1838) in several guidebooks Vogt, 1911, Micryletta inornata [synonym of of Taiwanese amphibians; however, L. kuhlii is Micryletta steinegeri (Boulenger, 1909)], R. known to be endemic to Java (McLeod 2010). It taipehensis [synonym of Hylarana taipehensis is necessary to clarify these taxonomic problems (van Denburgh, 1909)], R. taiwaniana [synonym before discussing the natural history of these frogs of Odorrana swinhoana (Boulenger, 1903)], R. on the island. tigerina rugulosa [synonym of Hoplobatrachus On the other hand, several studies focused rugulosus (Osbeck, 1765)], R. guentheri on the phylogeography of Taiwanese anurans [synonym of Hylarana guentheri (Boulenger, using a molecular approach. For example, Yang 1882)], Rha. prasinatus, Rha. taipeianus, Rha. et al. (1994) identified two well-differentiated moltrechti Boulenger, 1908, and Rha. robustus lineages among the Taipei treefrog Rha. [synonym of Buergeria robusta (Boulenger, taipeianus collected from seven populations in 1909)]. The most recent revision of this list Taiwan; Tanaka-Ueno et al. (1998) discussed was completed in August 2008. In this version, the phylogenetic relationships of brown frogs several previously protected frog species were from Taiwan and Japan, which revealed highly removed from the “protected species” list divergent lineages among R. sauteri; Toda et al. because their populations had recovered well, (1998) indicated that Fejervarya limnocharis including Microhyla butleri, Microhyla heymonsi, populations of eastern Taiwan are genetically Micryletta steinegeri, Hop. rugulosus, Hylarana distinct from the remaining Taiwanese guentheri, Bue. robusta, and Rha. moltrechti. populations as inferred from allozyme data; Jang- On the other hand, some threatened species Liaw et al. (2008) proposed four biogeographic were added to the “protected species” list for districts in Taiwan based on the results of the first time, including two rhacophorid frogs phylogenetic analyses of Hylarana latouchii. which were newly described in 1994~1995. Subsequently, the complete phylogeographic The complete “protected species” list for frogs patterns of three Taiwanese frogs, R. sauteri, currently includes Babina okinavana (Boettger, Limnonectes fujianensis, and Buergeria robusta 1895) (II), Hylarana taipehensis (II), Pelophylax revealed by mitochondrial (mt)DNA data were fukienensis (Pope, 1929) (III), Rha. arvalis (II), described chronologically (Jang-Liaw and Lee Rha. aurantiventris (II), Rha. prasinatus (III), and 2009; Jang-Liaw and Chou 2011; Lin et al. 2012). Rha. taipeianus (III). This act prohibits human These studies were helpful in understanding the activities from disturbing these listed animals, formation of biogeography and natural history of including hunting them as food, capturing them anurans in Taiwan and adjacent areas. for pets or the pet trade, and destroying their Jang-Liaw and Chou : Anuran Fauna of Taiwan Based on Specimen Records 7 habitats for any purposes. mm (Lin and Lin 2009). Because of violent Unfortunately, threats to the survival of natural hazards combined with increasing human Taiwanese anuran are still serious. Overuse of activities on this island, the threats to Taiwanese land in both mountainous areas and on the plains anuran populations are become more destructive has caused the rapid disappearance of animal and large-scale. habitats on this island in recent decades (Hsu and Species introduced for commercial purposes Agoramoorthy 1997; Koh et al. 2006). Taiwanese also seriously threaten many native Taiwanese anuran populations (such as Hylarana taipehensis species. There are three introduced frogs in and Rha. arvalis) have been seriously damaged Taiwan now: Lithobates catesbeianus (Shaw, by human activities, such as industrial pollution, 1802), Kaloula pulchra Gray, 1831, and farming and cultivation, deforestation, and urban Polypedates megacephalus Hallowell, 1861. expansion, and both the population size and Lithobates catesbeianus was introduced from distribution range appear to have been reduced in North America as a cultured and pet species (Hsu the past few decades (Chou et al. 1993; Lien et al. and Liang 1970; Hou et al. 2006; Tyler et al. 2007; Chang et al. 2008; Lin et al. 2008). 2007). It is widely distributed in low-elevation In addition to anthropogenic threats to areas throughout Taiwan (Lue et al. 1990; Hou Taiwanese anurans, the main island of Taiwan et al. 2006). It competes with native anuran is subject to frequent earthquakes and typhoons species by preying on them (Wang et al. 2006).