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Pacific Science (1993), vol. 47, no. 3: 248-255 © 1993 by University of Hawaii Press. All rights reserved

The Herpetofauna of the Senkaku Group, Ryukyu Archipelago!

HIDETOSHI OTA,2 NORIAKI SAKAGUCHI,3 SADAO IKEHARA,4 AND TsuTOMU HIKIDA 5

ABSTRACT: The herpetofauna of the Senkaku Group, western Ryukyus, was reviewed on the basis of recent fieldwork, as well as museum specimens and literature records. As a result, six species of reptiles were recorded from the islands. They are Gekko hokouensis Pope, Eumeces elegans Boulenger, Scincella sp., Ramphotyphlops braminus (Daudin), Elaphe carinata carinata (Gunther), and Dinodon ruJozonatus ruJozonatus (Cantor). No amphibian species were recorded. The herpetofauna of the Senkaku Group is distinct from that of other parts of the Ryukyu Archipelago and is more similar to that of and eastern continental . These conclusions conform with paleogeographical evidence indicating that most islands of the Senkaku Group and Taiwan were connected to the eastern margin of the continent during the most recent glacial period, when the remaining Ryukyu Islands were never connected by dry land with the continent.

THE SENKAKU GROUP consists of eight un­ absence ofregular transportation media from inhabited islets, ca. 150 km NNW of the the other islands of the Ryukyu Archipelago. Yaeyama Group and ca. 160 km ENE of Moreover, although a comprehensive biologi­ Taiwan (Figure 1). Unlike the other Ryukyu cal survey was made in 1979 by S. Ikehara Islands, the Senkaku Group lies on the north­ and other colleagues, the results remain western side of the (i.e., the unpublished. southeastern margin ofthe East China Conti­ An opportunity permitted N.S. and S.l. to nental Shelf), thus having a unique geohistory visit and to survey the herpetofauna of the (see Discussion for further details) and a four major islands ofthe Senkaku Group (see fauna and flora distinct from that ofthe other Figure 1) in May 1991. Here we review the Ryukyus. herpetofauna ofthis island group on the basis Relatively few and mostly brief surveys of that collection as well as available museum have been made of the fauna and flora of the specimens and literature reports. Senkaku Group (Kuroiwa 1900, Miyajima 1900, 1901, Masaki 1941, Takara 1954, Ike­ Descriptions ojthe Islands hara and Shimojana 1971). This neglect is chiefly due to the group's isolation and the Uotsurijima (25 0 45' N, 123 0 28' E) is the largest island of the Senkaku Group, ca. 4.3 2 1 Field trips of H.O. and T.H. for the collection of km and 362.0 m maximum elevation. This comparative specimens were supported by a U.S. Na- island is composed mostly of sandstone beds -"ti-"!on'!

Kobisho Uotsurijima o Kita-kojima '" Minami-i \) kojima d 5km I Sekibisho,.

Senkaku Group Taiwan

Ryukyu Arch ipelago

0 25N ~ ~.h Yaeyama Gr~p d o Miyako Group 0 Q~'o .. 0 50 km 11m 123 <::> 124 125E - - -

FIGURE I. Map of southern Ryukyus and Taiwan, showing the location of the Senkaku Group.

Scaevola taccada [Gaertner] Roxburgh and posed of igneous rocks obviously from a lava Argusia argentina [Linnaeus fil.] Rein) domi- flow (Takara 1962, Kizaki 1985). The bare nate (Niiro 1964). rock and a few crater remnants led Kizaki Kita-kojima (25 0 42' N, 123 0 33' E) lies (1985) to suggest that this island first emerged about 5 km E ofUotsurijima. It is 0.3 km2 and as a volcano during the Pleistocene. There are 124.5 m maximum elevation, and consists no permanent springs or streams on these chiefly of a sandstone bed with an upheaved latter three islands. They also lack large coral reef zone along the northern half of the woody plant species (Masaki 1941, Niiro ----G0ast-(+aKaFa-1-ge2-,Ki~a·Ki-1-98_;)_)~Minami-I-964,Kizak-i-l-98- 1 kojima (25 0 42' N, 123 0 34' E) is about 200 m SE of Kita-kojima. It is 0.5 km2 and 146.9 m maximum elevation, and consists oftwo areas METHODS of sandstone beds and a mass of upheaved coral reefjoining them (Takara 1962, Kizaki The itinerary ofN.S. and S.1. was as follows: 1985). Kobisho (25 0 56' N, 123 0 41' E) is about Kita-kojima, daytime of 24 May; Minami­ 27 km NE ofUotsurijima and is 1.1 km2 and kojima, evening of 24 May to noon of 25 118 m maximum elevation. Kobisho is com- May; Kobisho, evening of25 May to morning 250 PACIFIC SCIENCE, Volume 47, July 1993 of27 May; and Uotsurijima, noon of27 May Eumeces elegans Boulenger to morning of 29 May. Each island was surveyed in as many environments as possible, Four specimens (Minami-kojima: I, KUZ from the bare rocky zone near the coast inland 18077; Kobisho: 3, KUZ 21165-21167) were to grassland and the forest (except for Kita­ collected. On Minami-kojima, this skink lived around rocks and forests near the coast kojima, which has no forest at all). The survey th~ took place during both day and night, except whereas on Kobisho it was found around on Kita-kojima. border between the inland bare rocks and Morphological features of the specimens shrubs near the coast. Several individuals were examined after preservation. In the were also observed near the coast, but not in following section, zoological collections of the forest, on Uotsurijima. Five specimens Ky?to l!niversity (Department of Zoology), from Uotsurijima and five from Minami­ Umverslty of the Ryukyus (Department of kojima (from surveys in 1971 and 1979) are in Biology), and Okinawa Prefectural Museum URB (uncatalogued), OPM (UCI-4), and are referred as KUZ, URB, and OPM, KUZ (18921-18924). respectively. All specimens are characteristic of E. ele­ gans: SVL of adults (n = 7) 67.0-75.2 mm (x = 71.0), enlarged and irregularly arranged postfemoral scales and a pair ofkeeled posta­ SPECIES ACCOUNTS nal scales (Hikida 1989) present, and no post­ nasals. Six specimens have a juvenile color Gekko hokouensis Pope pattern of five light lines against the dark Fourteen specimens (Uotsurijima: 7, KUZ ground color; the dorsolateral line wholly 21161 and six uncatalogued specimens; Mi- enters the ear opening. The others exhibit nami-kojima: 2, KUZ 21162 and one uncata- adult coloration and lack such lines. logued specimen; Kobisho: 5, KUZ 21164 Eumeces elegans occurs widely in Taiwan and four uncatalogued specimens) were found and southeastern continental China (Ota mainly in crevices of rocks near the coast 1991) but not elsewhere in the Ryukyu Archi­ during the day. On Uotsurijima, several indi- pelago (Toyama 1989). No variation exists viduals occurred beneath the bark of dead between Taiwanese and continental popula­ trees. This gecko was active on rock surfaces tions or among continental populations (Tay­ ~nd tree trunks at night. On Minami-kojima, lor 1935, Hikida 1986). There is, however a It was found on drifts of trees near the coast. slight difference in the number of scale ro~s Four additional specimens from Uotsurijima at midbody between the Senkaku sample and (from surveys in 1971 and 1979) are in URB samples from Taiwan and the continent· most (REPI24-127). Two of these specimens were (nine) of the former have 28 rows (x ~ 27.7, referred to Gekko japonicus (Dumeril & 26-29, n = 13), whereas most from Taiwan Bibron) by Ikehara and Shimojana (1971). (106/150) and the continent (45/61) possess 26 All specimens (n = 18) have the usual char- rows. acteristics of G. hokouensis (e.g., snout-vent . length [SVL] 44.9-58.8 mm (x = 54.8), entire Scmcella sp. _subdigital scansors, single cloacal spur, dorsal This small, diurnal brown skink lives on the tubercles, alrd-a.bsence~oftubercles-on-limbs-f()res-t-f1()()r-()n-H0tsufijimaT-h0w~ver,-no ­ rOta 1989]). ThiS species occurs throughout was captured. The lizard was not seen on the the ~yukyu Arc?ipelago, Taiwan, and.ea.stern other islands visited and no specimens now contmer:ttal Chma (Ota 1991).. Prehml!1ary occur in museum collections. Although we comp~nsons showed no recogmzable dlffer- were unable to examine a specimen of Scin- ences m .external characters between the pres- cella sp. from the Senkaku Group, Toyama ent specimens and those fr?m other Ryukyu (personal communication) once examined Islands (n = 101) and Taiwan. ~n = 73; see two specimens from Uotsurijima in 1971 and Kohno and Ota (1991) for 10cahtIes). confirmed the identity of this population. Herpetofauna of the Senkaku Group, Western Ryukyus-OTA ET AL. 251 Takara (1954) first reported Leiolopisma (1962) are similar to those of Taiwanese and laterale (Say) (i.e., Scincella sp. sensu Mittle­ continental samples described by Maki (1931) man [I950, 1952]) from the forest of Uotsuri­ and Pope (1935), respectively. They are, how­ jima. While allocating Leiolopisma to sub­ ever, distinct from the Yonagunijima popula­ generic status of the widely distributed genus tion in having 23 midbody scale rows; the Lygosoma, Nakamura and Ueno (1963) com­ latter population, usually regarded as an en­ bined the popul~tion of Uotsurijima with demic subspecies, E. c. yonaguniensis Takara, those ofthe Yaeyama and the Miyako groups possesses 25 midbody scale rows (Takara into a single subspecies, Lygosoma (Leio­ 1962, Nakamura and Ueno 1963). lopisma) reevesii boellgeri (Van Denburgh). Ikehara and Shimojana (1971) reported that Dinodon rujozonatus rujozonatus (Cantor) this skink also occurs on Kobisho. None of those authors, however, described the mor­ An adult female (KUZ 18132: SVL = 51.1 phology of Senkaku specimens or compared mm) was collected on the forest floor during them with individuals from neighboring local­ the day on Uotsurijima. Another adult female ities, such as the Yaeyama Group, Taiwan, (URB REPI28: SVL = 55.4 mm) was col­ and eastern continental China. lected from this island during the 1979 survey. Dinodon rujozonatus occurs widely in the Ramphotyphlops braminus (Daudin) eastern part ofthe continent, Taiwan, and the Miyako and Yaeyama groups ofthe southern This tiny, secretive snake was not found Ryukyus (Takara 1962, Ota 1991). Based on during the survey reported here. As has been lower counts of ventrals and transverse dark­ reported already, one specimen (OPM H0546: light body bands and absence of reddish SVL = 91.6 mm), collected from Uotsurijima coloration in light bands, populations of the during the 1979 survey, showed no differences Yaeyama and the Miyako groups are re­ from specimens from Taiwan or other Ryu­ garded as a distinct subspecies, D. r. walli kyu Islands (Ota et al. 1991). Stejneger, rather than the nominotypical one from Taiwan and the continent (Stejneger Elaphe carinata carinata (Gunther) 1907, Maki 1931, Nakamura and Ueno 1963). Both of the specimens from Uotsurijima Two individuals were observed at the bor­ showed slight but distinct reddish coloration der between the forest and grassland during in the light bands on body and tail. This the day on Uotsurijima. One, an adult male coloration and the relative similarities of (KUZ 18076), was captured. None was ob­ meristic characters (Table I) suggest that the served on the other islands. Two specimens Uotsurijima population is more closely re­ (URB, uncatalogued) were collected in 1979. lated to the Taiwanese and continental popu­ Takara (1954) and Ikehara and Shimojana lations than to those of other parts of the (1971) reported this snake from Minami­ southern Ryukyus. kojima and Kita-kojima. Moreover, Takara (1962) stated that E. c. carinata also occurs on Kobisho; however, he provided no evidence DISCUSSION ofspecimens to substantiate the record. Thus, erification-is-needed-fQ.f-K-Qhish0'~.------ifir----d-,- ncon rme ecor s Elaphe carinata occurs widely in eastern and southeastern China, including Taiwan Takara (1954) reported Hemidactylus jre­ and northern Indochina (Ota 1991). Within natus Dumhil & Bibron from the forest on the Ryukyus, it is confined to Yonagunijima Uotsurijima. No individuals were observed of the Yaeyama Group and the Senkaku and no vocalizations were heard there during Group (Takara 1962). Meristic characters of recent fieldwork (i.e., 1971 and 1979 surveys three Senkaku specimens (SVL: 1250-1421 [Ikehara, unpublished data], and the survey mm,.x = 1363) and those described by Takara reported here). No specimens of this gecko 252 PACIFIC SCIENCE, Volume 47, July 1993

TABLE 1 MERISTIC CHARACTERS OF Dinodon ru!ozonatus FROM SEVERAL LOCALITIES

SUBSPECIES LOCALITY n VT SC BB TB rulo::onatus Uotsurijima 2 193.5 79.0* 55.0 22.0* (192-197) (79) (51-59) (22) Taiwan 14 196.2 84.3 54.1 22.8 (182-204) (79-89) (.50-60) (16-29) Continent 17 201.0 72.9 59.5 18.1 (185-213) (64-88) (54-67) (15-21) waW Yaeyama Group 89 188.8 81.2 30.0 16.4 (176-202) (68-89) (21-47) (6-26) Miyako Group 8 182.9 82.8 25.1 16.3 (180-190) (78-86) (20-30) (14-18)

NOTE: Data presented as means, followed by ranges in parentheses. Data for the continental sample are those provided by Pope (1935). Other data are from the study reported here. VT, ventrals; SC, subcauda1s; BB, body bands; TB, tail bands. * Examined for only one individual having an undamaged tail tip.

were found among previous collections either. no E. marginatus among the Senkaku Eu­ Thus, we suspect that Takara's (1954) record meces; all were E. elegans. The two species are was either a misidentification of specimens or very similar, and their distributions in the specimens with wrong locality records and Ryukyus and Taiwan have often been wrong­ that H.frenatus does not actually occur in the ly documented as a result ofmisidentifications Senkaku Group. (see Hikida [1989] for review). Thus, we sus­ Populations of Gekko in the Ryukyus, Tai­ pect that records of E. marginatus from the wan, and eastern continental China were long Senkaku Group are also incorrect. considered a single species, G. japonicus. Re- cently, it was discovered that there are actu- Herpetofaunal Characteristics ofthe Senkaku ally two species, G. japonicus and G. ho- Group and Their Historical Significances kouensis, and that most, ifnot all, populations in the Ryukyus and Taiwan and part of those Six species and subspecies of reptiles occur in continental China belong to the latter in the Senkaku Group, but no amphibians. Of species (Zhou et al. 1982, Ota 1986, 1989, the six reptiles, G. hokouensis and R. braminus 1991, Ota et al. 1989). This conclusion re- occur throughout most of subtropical East quires reexamination of all other East Asian Asia, including the other island groups of the populations referred to G. japonicus (sensu Ryukyu Archipelago, Taiwan, and the conti­ lato: e.g., Shibata 1989, Hikida et al. 1992). nent, and thus offer no data on the affinities of Ikehara and Shimojana (1971) reported G. the Senkaku herpetofauna. Without a specific japonicus from Uotsurijima, Minami-kojima, identification, the Senkaku Scincella is simi­ and Kobisho, but our results and the above- larly noninformative. The remaining three mentioned changes suggest that the occur- taxa, E. elegans, E. c. carinata, and D. r. -0f-fr.-japonieus-Esensu-st-fietej-i-n-thl:l -r-ujozonatus,---dQ-nQt-Qccur-in-the-resLoL.th _ Senkaku Group is unlikely. Ryukyu Archipelago and are shared exclu- Several previous authors have stated that sively with Taiwan and the continent, demon­ Eumeces marginatus (Hallowell), a skink strating a faunal affinity of the Senkaku known from the central Ryukyus, also occurs Group with eastern China. in the Senkaku Group (Miyajima 1901, Ta- Recent geohistorical studies reveal that kara 1954, Ikehara and Shimojana 1971). land area was larger globally during the late None of those authors, however, provided Pleistocene (15,000 to 18,000 yr ago) than now adequate data to confirm the identity of because of lower sea level during continental skinks observed and/or collected. We found glaciation. Estimates ofthe sea level drop vary Herpetofauna of the Senkaku Group, Western Ryukyus-OTA ET AL. 253

o 100 krn 125 130E -

fIGURE 2~rEast1\Sia,sfiOWmg the current land areas (darkened portions) and minimally estimated addi tionalland areas exposed above sea level during the late Pleistocene (currently submerged by the sea but as deep as, or shallower than, 120 m: stippled portions). Note that islands of the Senkaku Group (indicated with an arrow) were attached, or adjacent, to the continent during the late Pleistocene. from 120 to 140 m (Donn et al. 1962, Kizaki within the last 20,000 yr are supposed to be and Oshiro 1980, Hopkins 1982). On the other negligibly small in (Kizaki and hand, geomorphological changes such as up­ Oshiro 1980, Ujiie 1991). Therefore, the con­ lifting or subsidence by tectonic movement figuration of land areas during the late Pleis- 254 PACIFIC SCIENCE, Volume 47, July 1993 tocene can largely be reconstructed simply Zug, and one anonymous reviewer for con­ by drawing lines around areas above 120 or structive criticism on early versions of the 140 m below the current sea level. manuscript, and to M. Toyama for informa­ The minimum estimate ofland areas during tion regarding specimens ofScincella sp. from the late Pleistocene are depicted in Figure 2, Uotsurijima. Special thanks are due Y. Chi­ drawn on the basis of depth charts of related gira for checking the entire OPM collection regions by the Maritime Safety Agency (1978). for pertinent specimens. This map shows that Uotsurijima, Kita­ kojima, Minami-kojima, and Taiwan were fused as an eastward extension of the conti­ LITERATURE CITED nent during the last glacial period. Kobisho may have been separated from the continent DONN, W. L., W. L. FARRAND, and M. EWING. by a narrow strait, because current depths are 1962. Pleistocene ice volumes and sea level as deep as or deeper than 140 m. Even so, it lowering. J. Geol. 70:206-214. was much closer to the continent than to other HIKIDA, T. 1986. Geographic variation of parts of the Ryukyus, which would have been Eumeces elegans. Acta Herpetol. Sin. 5: 65­ not much different from current configura­ 66. tions (Figure 2). The Senkaku Group seems .---. 1989. The Ryukyu blue-tailed skink, to have been isolated from the continent and Eumeces marginatus, not distributed in Tai­ Taiwan only recently (i.e., less than 15,000 yr wan. J. Taiwan Mus. 42:81-88. ago) with rising sea level. HIKIDA, T., H. OTA, and M. TOYAMA. 1992. The herpetofauna of the Senkaku Group, Herpetofauna of an encounter zone of ori­ with its similarity to that of Taiwan and the ental and palearctic elements: Amphibians continent, can be explained by the above­ and reptiles of the Tokara Group and mentioned paleogeographical scenario. Obvi­ adjacent islands in the northern Ryukyus, ously, those species exclusively shared with Japan. BioI. Mag. Okinawa (30): 29-43. Taiwan and the continent could have mi­ HOPKINS, D. M. 1982. Aspects of the paleo­ grated to Uotsurijima, Minami-kojima, and ecology of Beringia during the late Pleisto­ Kita-kojima through the area exposed above cene. Pages 3-28 in D. M. Hopkins, J. the sea during the late Pleistocene. Those on V. Matthews, Jr., C. E. Schweger, and S. B. Kobisho might have had to cross a narrow Young, eds. Paleoecology ofBeringia. Aca­ strait by rafting, but the herpetofaunas of demic Press, New York. other parts ofthe Ryukyus are also considered IKEHARA, S., and M. SHIMOJANA. 1971. The as having partially dispersed by rafting (e.g., terrestrial animals of . Kohno and Ota 1991, Hikida et al. 1992). The Pages 85-138 in S. Ikehara, ed. Survey significance of the apparent differentiation report on Senkaku Islands. University of of E. elegans and the taxonomic status of the Ryukyus, Naha (in Japanese with En­ Scincella sp. in the Senkaku Group require glish summary). future studies. KIZAKI, K., ed. 1985. Geological synopsis of the Ryukyu Arch. Okinawa-Times, Naha (in Japanese). ACKNOWLEDGMENTS KIZAKI, K., and I. OSHIRO. 1980. The origin =':::...:~==~~-- R_y-uk-yu-Islands.-Eages-8=.3J_in-K. We wish to express our sincere thanks to K. Kizaki, ed. Natural history of Ryukyu. Kurihara, the registered owner of the Sen- Tsukiji-shokan, (in Japanese). kaku Group, for permission to land and KOHNO, H., and H. OTA. 1991. Reptiles in a survey there, and to the Okinawa Branch of seabird colony: Herpetofauna of Nakano- Nippon Hoso Kyokai (NHK) for financial kamishima Island of the Yaeyama Group, support. M. Yokota and Y. Niijima provided Ryukyu Archipelago. Is. Stud. Okinawa valuable company during the fieldwork. We (9): 73-89. are also much indebted to H. M. Smith, G. R. KUROIWA, H. 1900. Report from the expe- Herpetofauna of the Senkaku Group, Western Ryukyus-OTA ET AL. 255 dition to the Senkaku Group. Chigaku- ofEast Asia and Saipan. Amphib.-Reptilia Zasshi 12: 476-483,528-543 (in Japanese). 12: 181-193. MAKI, M. 1931. A monograph of the snakes OTA, H., K.-Y. LUE, S.-H. CHEN, and W. C. of Japan. Dai-ichi Shoboo, Tokyo. BROWN. 1989. Taxonomic status of the MARITIME SAFETY AGENCY, ed. 1978. Depth Taiwanese Gekko, with comments on the charts of . Maritime Safety synonymy of Luperosaurus amissus Taylor. Agency, Tokyo. J. Herpetol. 23: 76-78. MASAKI,T.194I.SurveyoftheSenkaku.Coll. POPE, C. H. 1935. 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