A New Subspecies of the Agamid Lizard, Japalura Polygonata (Hallowell, 1861) (Reptilia: Squamata), from Yonagunijima Island of the Yaeyama Group, Ryukyu Archipelago
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Current Herpetology 22 (2): 61-71, December 2003 (C) 2003 by The Herpetological Society of Japan A New Subspecies of the Agamid Lizard, Japalura polygonata (Hallowell, 1861) (Reptilia: Squamata), from Yonagunijima Island of the Yaeyama Group, Ryukyu Archipelago HIDETOSHI OTA Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 903-0213. JAPAN Abstract: Japalura polygonata, occurring in the East Asian islands, is currently divided into three subspecies-J. p. polygonata from the Amami and Okinawa Groups of the central Ryukyus, J. p. ishigakiensis from the Miyako and Yaeyama Groups of the southern Ryukyus, and J. p. xanthostoma from northern Taiwan. A new subspecies is described for this species from Yonagunijima Island of the Yaeyama Group. This subspecies differs from other conspecific subspecies in having distinctly enlarged and irregularly arranged scales on the dorsolateral surface of the body. In other subspecies, the degree of enlargement of such scales is smaller, and they usually form somewhat regular rows in a transverse direction on the flanks, and in a longitu- dinal direction in the paravertebral region. Males of the present subspecies differ from those of other subspecies in having a series of large white spots against a dark grayish tan on the dorsolateral surface of the body, whereas the females are characterized by brilliant green dorsal coloration. Key words: Japalura polygonata; New subspecies; Reptilia; Geographic variation; Yonagunijima Island; Ryukyu Archipelago INTRODUCTION graphic range of the genus (and actually of the family Agamidae as well), being distributed in The agamid genus Japalura consists of 24 northern Taiwan and most islands of the species and two subspecies distributed from Ryukyus south of the Tokara Group. Three northern India and Nepal to the East Asian subspecies are currently recognized for the islands (Ota, 2000a, b; Gau and Hou, 2002: species-J. p. polygonata from the Amami but see Marcey et al., [2000] for the possible and Okinawa Groups of the central Ryukyus, non-monophyly of the western and eastern J. p. ishigakiensis from the Miyako and components, and Schleich and Kastle [2002] Yaeyama Groups of the southern Ryukyus, for a different generic arrangement). Of the and J. p. xanthostoma from northern Taiwan species of Japalura, J. polygonata occupies (Ota, 1991). In the process of their infraspe- the northeasternmost extremity of the geo- cific classification, however, very few island samples have been examined. Hallowell * Tel: +81-98-895-8937; Fax: +81-98-895- (1861), for example, described the species (as 8966;E-mail address:[email protected] Diploderma polygonatum) only on the basis 62 Current Herpetol. 22 (2) 2003 of a single specimen from "Amakarima Island" of the Yaeyama Group, is the westernmost (subsequently restricted to the Kerama Islands island of the Ryukyus. Its herpetofauna, while of the Okinawa Group by Stejneger [1907]). sharing most species and subspecies exclu- Van Denburgh (1912), in describing two sively with other southern Ryukyu islands subspecies, J. p. ishigakiensis and J. p. (Yasukawa et al., 1996; Toda et al., 1997; Ota, miyakensis, examined specimens only from 1998, 2000c; Chen et al., 2001), includes two Ishigakijima and Iriomotejima Islands for endemic snakes (Calamaria pavimentata ishigakiensis, Miyakojima Island for miyak- miyarai and Elaphe carinata yonaguniensis), ensis, and Okinawajima and Amamioshima whose closest relatives occur only in Taiwan Islands for the nominotypical subspecies. and the southern part of the continent Although several subsequent authors referred (Takara, 1962; Ota, 1998, 2000c). The occur- to populations of J. polygonata on some other rence of Japalura polygonata on Yonagunijim islands (e.g., Okada, 1937; Nakamura and Island was first noted in Okada's (1937) Ueno, 1963; Ikehara et al, 1984; Toyama, monograph on the basis of information from 1985), or proposed infraspecific classification "KURODA". Curiously enough, however, different from that proposed by Van Denburgh Okada (1937), while introducing Van Den- (1912) (Nakamura and Ueno, 1963; Matsu- burgh's (1912) subspecific classification of J. moto, 1979; Ota, 1991), none of them investi- polygonata, retained the Yonagunijima popu- gated between-island variation in Japalura lation in the nominotypical subspecies (p. 93). polygonata in detail. Nakamura and Ueno (1963) also assigned the Yonagunijima Island (Fig. 1), a component Yonagunijima population to J. p. polygonata FIG. 1. Map of the southern Ryukyus and adjacent regions showing the location of Yonagunijima Island, the type locality of Japalura polygonata donan subsp. nov. The stippled portions denotes current sea areas that are considered to have constituted additional lands during the Late Pleistocene (i.e., sea areas shallower than 120m: Ota et al., 1993). OTA-NEW JAPALURA FROM YONAGUNIJIMA 63 together with all other Ryukyu populations dimorphism index (SDI) proposed by Gibbons under the assumption that Japalura swinhonis and Lovich (1990) as: Gunther, 1864 (including J. mitsukurii Stej- +x/y, when x>y; or-y/x, when x<y neger,1898: see Ota, 1991) of Taiwan should where x and y denote means for adult females be regarded as a subspecies of J. polygonata, a and adult males, respectively. Differences in view clearly negated on the basis of recent meristic characters among samples were tested morphological and ecological observations by for statistical significance by Kruskal-Wallis Ota (1988a, 1991). Matsumoto (1979), on the test and Dunn's multiple comparison test other hand, implicitly synonymized Van following Zar (1984). Denburgh's (1912) J. p. miyakensis with J. p. In the following sections, catalogue numbers ishigakiensis, because he referred to all of specimens deposited in the Zoological Yaeyama (including Yonagunijima Island) and Collection of the Kyoto University Museum, Miyako populations as J. p. ishigakiensis. and Masami Hinoue's private collection are Recently I had a chance to examine a good preceded by KUZ and HPN, respectively. The series of Japalura specimens from Yona- other institutional acronyms follow Leviton et gunijima Island for the first time. Comparison al. (1985). of these specimens with specimens of J. poly- gonata from many other islands of the Ryukyus Japalura polygonata donan subsp. nov. and Taiwan revealed the presence of slight Fig. 2 but consistent differences between the Yona- gunijima specimens and the remainder. How- Japarula [sic] polygonata polygonata: ever, they exclusively shared an orange [males] Okada, 1937:93 (part). or yellow [females] spot at the center of the Japalura polygonata polygonata: Nakamura gular. Because such a gular spot was not and Ueno, 1963: 103 (part). observed in other species so far studied (Ota, Japalura polygonata ishigakiensis: Matsu- 1991), I tentatively regard this spot as synapo- moto, 1979: 17 (part); Ota, 1983:16 (part); morphy of J. polygonata, and thus describe Toyama, 1985:63 (part); Ikehara et al., 1994: the Yonagunijima population as a new subspe- 275 (part); Ota,1991: 288 (part). cies of this taxon. Holotype KUZ R234, an adult male collected at Mt. MATERIALS AND METHODS Urabudake (24°27'N, 123°00'E, alt. 130m), A total of 56 specimens (42 adult males, 11 Yonagunijima Island, on 17 August 1981 by adult females, and three juveniles) collected H. Ota. from Yonagunijima Island were compared with 354 specimens of J. p. polygonata from Paratypes eight islands of the central Ryukyus, 177 speci- KUZ R102, an adult female collected at mens of J. p. ishigakiensis from three islands Higawa, Yonagunijima Island, on 30 March of the southern Ryukyus, and 148 specimens 1981 by H. Ota; KUZ R231, 238, two adult of J. p. xanthostoma from four localities in females, sampling data same as the holoyupe; northern Taiwan (see Appendix for further KUZ R415, an adult female collected between details). Of the 56 Yonagunijima specimens, Higawa and Arakawabana, Yonagunijima two males and two females were also subjected Island, on 7 March 1982 by H. Ota; KUZ to karyotyping following Ota (1988b). R971, 972, 975, 979, 997, four adult males and Morphological characters were examined one adult female collected between Mt following Ota (1991). Sexual difference in Urabudake and Sonai, Yonagunijima Island, adult snout-vent length (SVL) was examined on 26 March 1983 by T. Hayashi, K. by t-test, and was also expressed by the sexual Kawamura, S. Kobayashi, A. Mori, and H. 64 Current Herpetol. 22 (2) 2003 FIG. 2. Male paratype (A. KUZ R3294) and female paratype (B. KUZ R4515) of Japalura polygonata donan subsp. nov. Ota; KUZ R1382, 1420, two adult females Diagnosis collected at Kubura, Yonagunijima Island, on Adult females largest among the subspecies 27 July 1983 by H. Ota; KUZ R2000-2003, of J. polygonata (55.5-70.5mm in snout-vent 2164, 2876, 3261-3263, 3265, 3269, 3270, length [SVL] [x=63.3]), brilliant green dor- 3281-3283, 3289, 3291-3294, 3982, 4515, sally. Adult males almost as large as females 4517, 4518, 22 adult males, two adult females, (57.7-71.0mm SVL [x=65.6]), with a series and three juveniles collected at Mt. Urabudake of large white spots against dark grayish tan on 15 July 1984 by H. Ota; KUZ R28117, on the lateral surface of body. Middorsal 28118, two adult females collected at Kubura scales 30-44 (x=37.1), fewer than those in on 25 June 1993 by H. Ota; 56152-56156, other subspecies, but with some range overlaps 56158-56167, 15 adult males collected at Mt. (34-47 [x=40.8] in J. p. polygonata; 35-53 Urabudake on 25 March 2003 by H. Ota. [x=44.3] in J. p. ishigakiensis, and 37-48 OTA-NEW JAPAL URA FROM YONAGUNIJIMA 65 [x=42.4] in J. p. xanthostoma); some dorso- irregularly arranged scales, each with a distinct lateral body scales distinctly enlarged, irregu- midline keel; some scales in occipital region larly arranged; orange (male) or yellow with a few additional weak keels, making (female) spot at center of gular in adults; surface rugose; interparietal surrounded by sexual size dimorphisms not evident; buccal eight scales; superciliaries imbricate, each and palatal mucosa light pinkish gray.