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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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 longitudinal 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

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. overlapping one third to half with successor; scales around eye very small; scales on ventral Etymology surface of head obliquely arranged, most of The subspecific epithet, "donan", refers to them more or less keeled medially; but a few an old vernacular name of Yonagunijima not keeled; longitudinal gular fold very slight; Island. "Donan" is not in official use at slender, hair-like sense organs present on present. However, this term is still commonly several head scales; longitudinal middorsal used by the inhabitants of this island. "Donan" scale row consisting of 40 enlarged, strongly also refers to a kind of special liquor of keeled scales, anterior eight especially enlarged, Yonagunijima, which added a great joy to my and much compressed, each almost as broad fieldwork there. as high, forming a nuchal crest; scales adjacent to middorsal row smaller, directed backward Description of holotype and slightly upward; most scales in dorsolat- Measurements (in mm): SVL 69.4; head eral and lateral regions of body weakly keeled, length 21.8; snout-eye length 9.0; interorbital as large as or smaller than those on venter, distance 10.9; fore-limb length 33.6; axilla to intermixed with much enlarged, strongly keeled, groin distance 31.8; hindlimb length 55.4; tibia irregularly arranged scales; scales on ventral length 18.1; toe IV length 13.4; tail length surfaces of body and limbs moderately keeled 178.1. and regularly arranged longitudinally; slight Snout tapering, roundish at tip; rostral oblique fold on each shoulder; limbs relatively quadrangular, about three and a halh times short, toe IV (exlusive of claw) reaching to as wide as long, covering anterior tip of snout; middle of eye when hindlimb adpressed for- seven scales, in contact with rostral; nasal ward; scales on dorsal surface of limb larger separated from rostral by one scale, contacted than those on ventral surface; finger and toe I by this intervening scale, first supralabial, and shortest, finger IV almost as long as finger III, six other scales; supralabials not keeled, seven toe IV distincly longer than toe III; subdigital on left, six on right, second separated from scales 9-9, 13-13, 18-19, 19-19, 12-12 on nasal by one scale; a longitudinal row of left-right fingers I, II, III, IV, Y, 8-7, 13-13, keeled and moderately enlarged scales, each 20-20, 23-25, 16-16 on left-right toes I, II, III, larger than half the size of supralabials, behind IV, V; tail slightly compressed, oval in cross nasal to imaginary vertical line from posterior section, with slightly enlarged median scales margin of eye; one row of much smaller on dorsum; subcaudal scales strongly keeled, scales intervening between preceding row and distinctly larger than scales on dorsum. supralabials; mental pentagonal, surrounded by four scales; infralabials slightly keeled, Color in life seven on both sides; a row of enlarged scales Dorsal ground color dark grayish tan; on anterior part of lateroventral corner of infraorbital region paler than surroundings but mandible, first ones on both sides and second not white; otherwise, no distinct markings on one on right side contacting first infralabials, head; each side of body with five quadrangular the others separated from labial series by white spots forming a longitudinal row; second one or two scales; tympanum completely and third spots partially fused on left side, concealed beneath skin; dorsal and lateral separated on right side; ground color of limbs surfaces of head covered with enlarged and and tail slightly lighter than that of trunk, 66 Current Herpetol. 22 (2) 2003

several indistinct dark annuli on limbs, 15 dark Variation annuli on tail. Variations in some meristic characters and Ventral surface of head light gray, without SVL are given in Tables 1 and 2, respectively. markings except for one large orange spot in There are no significant differences in those midst of gular region; ventral surfaces of meristic characters between the sexes (P>0.05). remaining part of body pale gray. Buccal- Unlike in J. p. polygonata and J. p. ishigakiensis palatal mucosa light pinkish gray. where sexual size dimorphism is evident with males being larger, J. p. donan, as well as J. p. Color in ethanol xanthostoma, does not show significant differ- Dorsal ground color turned slightly paler; the ences in adult SVL between sexes (Table 2). orange spot on gular faded to invisible; the Live coloration of females remarkably remaining color patterns similar to those in life. differs from that of males. Dorsal surface of

TABLE 1. Variation in meristic characters (x±SD, followed by ranges in parentheses) among island samples of Japalura polygonata examined. Abbreviations are: SER, number of scales contacting rostral; IOS, number of scales touching an imaginary line drawn between outer edges of left and right supraciliaries; DC, middorsal crest-like scales from just posterior to occipital granules to just above anterior margin of vent; T4S, slightly enlarged scales beneath toe IV, counted from branching point with toe III to base of claw. OTA-NEW JAPALURA FROM YONAGUNIJIMA 67

TABLE 2. Variation in adult male and female SVLs (mm) in island samples of Japalura polygonata examined. Significance levels of difference in adult SVL between sexes from the same islands are as follows: +++, P<0.001; ++, P<0.01; -, P>0.05. SDI, sexual dimorphism index (Gibbons and Lovich, 1990: see text).

body of females is brilliant green, with no Distribution and habitat white spots but with several obscure dark cross Japalura polygonata donan is endemic to bands around the middorsal region. In females, Yonagunijima Island of the Yaeyama Group, there is a somewhat obscure yellow spot in Ryukyu Archipelago, Japan. Most specimens midst of gular region. were found in the marginal portions of well- recovered secondary forests consisting of Karyotype evergreen broad-leaf trees. Like other subspecies of J. polygonata (Ota, 1991), J. p. donan has a karyotype consisting Ecological notes of 2N=46 uniarmed chromosomes in a graded All three females collected in March had series. ovarian follicles (3-5mm in diameter) only, whereas the six females collected between mid July and mid Aunust exhibited flaccid. empty 68 Current Herpetol. 22 (2) 2003 oviducts and poorly developed ovarian folli- [x=42.4] in J. p. xanthostoma: Tables 1 and cles (<3mm in diameter). Of the remaining 3). The males of J. p. donan differs from the two females, both collected in the late June, males of other subspecies in having a series of one had two and one large follicles (7mm in large white spots against dark grayish tan on diameter) in left and right ovaries, respectively. the dorsolateral surface of the body. In other The other had one elliptic egg (15.8×7.5mm) subspecies, the dorsolateral surface of the in each oviduct. It is thus probable that in this body has a continuous white or yellow band subspecies ovulation and oviposition occur instead. The adult females of J. p. donan are chiefly in late June or early July. the largest among the females of J. polygonata One colubrid snake, Dinodon rufozona- populations (55.5-70.5mm in SVL, [x= tum walli, collected at Mt. Urabudake in 63.3mm]), and are almost as large as the adult August 1981, had two slightly digested adult males of the same subspecies (57.7-71.0mm male J. p. donan in its stomach. SVL [x=65.6]). Thus, the male large sexual size dimorphism is not evident in this subspecies unlike in other subspecies exclusive of J. s. DISCUSSION xanthostoma (Table 2). The females J. p. Japalura polygonata donan shares an donan are also characterized by the brilliant orange (males) or yellow (females) spot in the green coloration on the dorsum of the body. center of the gular exclusively with the other The buccal and palatal mucosa of the present conspecific subspecies. However, it differs subspecies is not yellowish as in J. p. xanthos- from the latter in having distinctly enlarged toma. and irregularly arranged scales on the dorso- Taxonomic treatment of allopatric, slightly lateral surface of the body. In other subspecies, but diagnosablely diverged entities like the the degree of enlargement of such scales is Yonagunijima population of J. polygonata is a smaller, and they usually form somewhat matter of serious debate among taxonomists, regular rows in a transverse direction on the depending on which school they belong to flanks, and in a longitudinal direction in the (Frost and Hillis, 1990; Grismer et al., 1994). paravertebral region. In the present subspe- In the present case, the population is clearly cies, the middorsal scales, ranging from 30 to diagnosable (see above). Moreover, from the 44 (x=37.1), are fewer than in other subspe- geographic pattern of variation in J. polygo- cies, although there are some range overlaps nata as a whole (Table 3), naming of the (34-47 [x=40.8] in J. p. polygonata; 35-53 Yonagunijima population is not an arbitrary [x=44.3] in J. p. ishigakiensis, and 37-48 slicing of a geographically clinal entity, a

TABLE3. Comparisons of island samples of Japalura polygonata by Kruskal-Wallis test (KWT) and Dunn's multiple comparison test. Abbreviations of island samples are: KK, Kikaijima; AO, Amamioshima; KR, Kakeromajima; TN, Tokunoshima; IH, Iheyajima, OK, Okinawajima; TK, Tokashikijima; KM, Kumejima; MY, Miyakojima; IS, Ishigakijima; IR, Iriomotejima; YN, Yonagunijima; TW, Taiwan. See Table 1 for character abbreviations. Significance levels for KWT are: +++, P<0.001; +, P<0.05. Samples sharing same superscript letters showed no significant differences in Dunn's multiple comparison test (P>0.05). OTA-NEW JAPALURA FROM YONAGUNIJIMA 69 practice that can not be justified by the part of the comparative materials used here, standards of modern taxonomy (e.g., Frost to Yoshinori Chigira, Masanao Toyama and Hillis, 1990). Also, one may argue that (OPM), Yasuhiko Shibata, Kiyotaka Hatooka the Yonagunijima population deserves the (OMNH), and Richard G. Zweifel (AMNH) status of a full species rather than a subspecies for the loan of samples in their care, to of J. polygonata. However, the presence of Hidetoshi Nagamasu, Hiroshi Shinji, Naofumi less diverged populations on both sides of Nakagawa, Yutaka Onishi, Akira Mori, Teru- Yonagunijima Island that are currently treated take Hayashi, Koichi Kawamura, Shuji as two conspecific subspecies (i. e., J. p. Kobayashi, Chia-Hsiang Wang, Jun-Tsong ishigakiensis of the remaining Yaeyama Lin, Szu-Lung Chen, Gaus Shang, and Mika Group and Miyako Group islands, and J. p. Ota for helping with field sampling, and to xanthostoma from Taiwan: Ota, 1991) makes Gen Masunaga for helping with manuscript it safer at present to retain the Yonagunijima preparation. Special thanks are due Motoo population also at the subspecihc status. Future Tasumi and Toshitaka Hidaka (formerly of the detailed investigations of historical relation- Graduate School of Science, Kyoto Univer- ships at the population level and captive sity) for their thoughtful care and help with hybridization experiments may lead to eleva- various aspects of my study when I was in their tion of J. p. donan to the full species status. laboratory as a graduate student. Yonagunijima Island is separated from other My field sampling was partially supported neighboring islands and the continent by by grants-in-aid from the Japan Ministry of straits deeper than 500m (Maritime Safety Education, Science, Sports and Culture Agency, 1978: Fig. 1). This suggests that the (JMESSC) (A-63790257 and A-05740523 to island has been isolated for a long period of me, and B-11480152 to Masako Izawa), and time (e.g., Ota et al., 1993). Japalura polygo- also by grants from the Fujiwara Natural nata donan and the two other reptiles endemic History Foundation (to me), the U. S. to Yonagunijima Island (i.e., Calamaria pavi- National Geographic Society (no. 4505-91: to mentata miyarai and Elaphe carinata yona- Masafumi Matsui), and the Ministry of guniensis) are likely to have diverged from Environment, Japan (Global Environment their relatives in Taiwan and the southern Research Programme: project leader, Koichi Ryukyus through such an isolation. Goka). Laboratory work was also supported in part by a JMESSC grant-in-aid (C-133415 to me). ACKNOWLEDGMENTS

I thank Tsutomu Hikida (Graduate School LITERATURE CITED of Science, Kyoto University), and Masafumi Matsui (Graduate School of Human and CHEN, S.-L., H. OTA, AND T. HIKIDA. 2001. Environmental Studies, Kyoto University) for Geographic variation in the two smooth skinks, much invaluable advice throughout my study Scincella boettgeri and S. formosensis (Squa- on the systematics and biogeography of the mata: Scincidae), in the subtropical East Asian East Asian Japalura, from which this paper is islands. Zool. Sci. 18: 115-130. derived. I am also much indebted to Masanao FROST,D. R. AND D. M. HILLIS. 1990. Species in Toyama, Satoshi Tanaka, Mamoru Toda, concept and practice: herpetological applica- Masanao Honda, and Masaharu and Junko tions. Herpetologica 46: 87-104. Motokawa for constructive discussion on GAU, Z. -F. AND M. HOU. 2002. Description of a relevant topics, and Ronald I. Crombie for new Japalura species from western Sichuan provision of pertinent literature. I am also Province, China. Sichuan J. Zool. 21: 3-5. much indebted to Tsutomu Hikida, Satoshi GIBBONS,J. W, AND J. E. LOVICH.1990. Sexual Tanaka, and Masami Hinoue for provision of dimorphism in turtles with emphasis on the slider 70 Current Herpetol. 22 (2) 2003

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YASUKAWA,Y., H. OTA, ANDJ. B. IVERSON.1996. 2338, 2393-2395, 2409-2420, 3962, 4059- Geographic variation and sexual size dimor- 4062, 4067-4076. phism in Mauremys mutica (Cantor, 1842) J. p. ishigakiensis. Ryukyu Archipelago: (Reptilia: Bataguridae), with description of a new Yaeyama Group: Ishigakijima Is. (type loc.): subspecies from the southern Ryukyus, Japan. KUZ R3-5, 225, 227, 702-714, 719-722, 903, Zool. Sci. 13: 303-317. 905, 906, 934-936, 1023, 1373-1379, 1386, ZAR, J. H. 1984. Biostatistical Analyses. Prentice 1474-1480, 2008, 2205-2207, 2218, 2219, Hall, Englewood Cliffs, New Jersey. 2222-2224, 2226, 2877, 2878, 2880-2883, 2885, 2897, 2928, 3052, 3279, 3280, 3290, 3977, 4519. OMNH R772, 786, 787, 1653, APPENDIX 1696-1699, OPM H0054, 0093; Yeyama Comparative specimens examined Group: Iriomotejima Is.: KUZ R2196-2204, Japalura polygonata polygonata. Ryukyu 2356-2359, 2390-2392, 3274, 3275, 3947, Archipelago: Okinawa Group: Kerama Islands 3948, 3959-3961, 3978, 3979, 4102, 4103, (type loc.): Tokashikijima Is.: KUZ R2300- 4365, 4366, 4521, 4522, 4524-4537, 4540, 2306, 2360-2369, 4082, 4085, 4087-4101, OPM H0025, 0051, 0052, 0074, 0081, 0085, 4494-4514, 4568, 4569, 4574-4576, OPM 0183-0186; Ryukyu Archipelago: Miyako H0187, 0188, 0437; Okinawa Group, Kume- Group: Miyakojima Is. (tyoe loc. of J. p. jima Is.: KUZ R2241, 2333, 2334, 2421-2424, miyakensis: KUZ R2150-2158, 2183, 2339- 4077, 4444-4493, 4578-4582, OPM H0082, 2348, 4080, 20 uncatalogued specimens. 0451; Okinawa Group: Okinawajima Island: J. p. xanthostoma. Taiwan: Taipei: Waishu- AMNH 21167-21169, 21172, HPN 42, 51, 55, angchi (type loc.): KUZ R9855 (holotype), 56, 59, 60, 69-71, 88-91, 383, 439, KUZ KUZ R1600-1602, 1609, 1610, 1649, 1657, R1146-1200, 1355-1358, 1380, 1381, 1383- 2058, 2059, 2715, 2809, 2919-2923, 2933, 1385, 1423, 1483, 1485, OMNH R93-98, 733, 2978, 2979, 3036, 3056, 3219, 3964, 4360, 734, 736, 737, 744, 759-762, 897, 898, 1393, 4361, 6707-6710, 7519-7534, 7537-7545, 1394, 1451-1453, 1455-1458, 1460, 1618, 7846, 7851-7855, 7861-7871, 7880-7882, 1704); Okinawa Group: Iheyajima Is.: KUZ 7884, 7901, 7922, 7941, 7942, 7952, 7961, R2267, 2307; 2308, 2370-2378, 3950-3958, 7973, 7991, 7992, 8009, 8010, 8019, 8020, 4083, 4086; 4570-4573, OPM HJ0029; Ryukyu 8066, 8067, 8099, 8420, 8446-8452, 9775- Archipelago: Amami Group: Tokunoshima 9780, 9842-9854, 9856-9861, 13010-13019; Is.: KUZ R2159-2162, 2425-2429, 2940, Taipei: Tansui: KUZ R8424-8431; Taipei: 2943-2945, 3000, 3014, 3057, 3102, 3224, Peitou: AMNH 77093; Taiwan: Ilan: Tungao: 3225, 3286, 3457, 3465, 3965, 4003-4031, KUZ R6914, 6915, 6943, 6944, 6947 (all 4033; Amami Group: Kakeromajima Is.: KUZ paratypes). R2396-2408, 3984-3999, 4063-4066; Amami Group: Amaioshima Is.: KUZ R2163, 2379- 2389, 2968-2971, OPM H0028; Amami Group: Accepted: 22 November 2003 Kikaijima Is.: KUZ R2266, 2321, 2335, 2337,