PROC. BIOL. SOC. WASH. 105(3), 1992, pp. 426^32 A NEW CHINESE EYED TURTLE OF THE GENUS SACALIA (BATAGURINAE: TESTUDINES) John B. Iverson and William P. McCord Abstract.—A new species of batagurine turtle, Sacalia pseudocellata, is de- scribed from Hainan Island, China. Similar to and apparently sympatric with Sacalia quadriocellata, it differs from that species and S. bealei by having eye- like markings without "pupils" on the posterolateral aspect of the head, a slightly tricarinate (generally unicarinate in S. bealei and S. quadriocellata) and wider carapace, more extensive black plastral pigment, a more extensive plas- tron, longer and narrower gular scutes, and a shorter interfemoral seam. Cistuda bealei (the four-eyed turtle) was Sacalia pseudocellata, new species first described in 1831 byJohn Edward Gray Chinese false-eyed turtle from "China," and it is known to range from Fig. 1A-D southeastern China to at least central Viet- —UF Holotype. 81505, an adult male, nam (Buskirk 1989, Iverson 1992). Al- preserved in alcohol; reported to have been though it has been placed in four different collected between Tungfang [19°03'N, genera by various authors, it was finally re- 108°56'E] and Kancheng [18°51'N, turned to Gray's (1870) monotypic genus 108°37'E; ca. 48 km from Tungfang], west- Sacalia based on biochemical (Merkle 1975) ern Hainan Island, China; purchased from and morphological studies (McDowell local persons living between those cities by 1964). Mr. Oscar Shiu, in the spring of 1988. In 1903 Siebenrock described the western Paratype. —UF 81506, an adult female, geographic variant of Sacalia (then Clem- preserved in alcohol; same data as holotype, mys) bealeias C. b. quadriocellata. This form except purchased in the spring of 1990. was considered a separate species by a few A Diagnosis.— medium-sized species of early authors (e.g., Pope 1935, Bourret 1941); however, Sachsse (1975), Rodel Sacalia (Fig. 1A-D, Table 1) with black- (1985), and Rodel & Praedicow (1988) ar- bordered yellow to olive-green blotches gued that quadriocellata is conspecific with dorsolateral^ on head region, a weakly tri- bealei. Recently, Fu & Zhao (1989) pre- carinate carapace (generally unicarinate in sented morphometric and color pattern data other Sacalia; e.g., S. bealei and S. quad- that supported the recognition ofbealei and riocellata), a carapace of medium width quadriocellata as distinct species. (relatively narrow in other Sacalia), heavily Three-"eyed" turtles from Hainan Island pigmented plastron (lightly pigmented in represent an undescribed species and they other Sacalia), a relatively long, broad plas- show more similarity with Sacalia bealei tron (shorter and narrower in other Saca- and with S. quadriocellata (with which it is lia), a relatively long, narrow gular scute apparently sympatric) than with any other (shorter and wider in other Sacalia), a rel- Chinese batagurine turtles. This distinctive atively long interpectoral seam (shorter in species is tentatively referred to the genus other Sacalia), and a relatively short inter- Sacalia, although confirmation of this as- femoral seam (shorter in other Sacalia). signment must await the availability ofos- Description (based on the types, and a liv- — teological material. ingjuvenile female). Carapace length (CL) VOLUME NUMBER 105, 3 427 -- %.C^ juk : Hk VvB '* ( > Wf,V */v* ^- \ K ^1 *jg , "ii I rfc, ty \ '^^r*':" *j \ 1 2 '^fc^*'. ^^^^^r^^^^^^B • ^^^0^^^ * y^^^v •^ •»•* •*>• «P> 1^^ JH ' * .0 ll^^^^^ Fig. 1. AdultfemaleSacaliapseudocellata(A-D; paratype: UF81506; 176 mmcarapacelength)fromHainan Island, China; typical head patterns ofmale (left) and female (right) Sacalia bealei (E, F) and S. quadriocellata (G, H). 428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Morphometric characters useful in discriminating among species ofthe genus Sacalia. Character abbreviations are interhumeral seam length (IH), interpectoral seam length (IP), interfemoral seam length (IF), maximum gular scute length (GL), maximum gular scute width (GW), maximum carapace width (MCW), and anterior width ofplastral hindlobe (PW3). Sample Sex n Carapace length (mm) IH/IP IF/PW3 M S. pseudocellata 1 151.8 0.434 0.248 F 2 147.8(119-176) 0.255 (0.24-0.27) 0.255 (0.24-0.27) M S. bealei 5 127.2(114-136) 0.609 (0.54-0.68) 0.310(0.29-0.35) F 4 132.0(119-140) 0.521 (0.42-0.69) 0.321 (0.29-0.38) M S. quadriocellata 4 127.0(120-143) 0.677(0.53-0.82) 0.339 (0.27-0.44) F 4 130.6(127-139) 0.596 (0.48-0.70) 0.339 (0.29-0.38) mm to at least 152 in males and at least 176 ' scute on each side ofbridge. Average plas- mm in females; carapace elliptical, weakly tral formula: interpectoral seam (IP) > in- tricarinate (shell height at level of hinge/ terabdominal seam (IAB) > interanal seam » carapace length = CH/CL = 0.36 to 0.39; (IAN) intergular seam (IG) > interfe- X — 0.375), widest at level ofmarginal M7 moral seam (IF) » interhumeral seam (IH). (maximum carapace width: MCW/CL = IG/IAB = 0.53 to 0.67 (X = 0.62); IF/IAN 0.69 to 0.75; X = 0.721), with a smooth to = 0.70 to 0.78 (X = 0.73); IG/IP = 0.55 to slightly serrated posterior margin, and with 0.63 (X= 0.59). Interanal seam present and moderately obvious growth laminae. complete. Plastron yellow or orange, with Ml, M7, M8, or M9 widest, Mil nar- black pigment covering most of each plas- M9 rowest, and tallest. Cervical scute small, tral scute (Fig. 1), although at least the mar- basically rectangular (but widest posterior- gins ofeach scute markedwithyellow; bridge ly), and longer than wide. All vertebrals (V) markedwith a longitudinal black stripe, pri- wider than long; VI contacting (or nearly marily on the pectoral and abdominal scutes. so) seam between Ml and M2; V5 not con- Head narrow; upperjaw not hooked; very tacting M10. Low medial keel most pro- small tubercles present dorsal to angle of nounced on VI, V4 and V5. Carapace dark jaw and anterior to tympanum. Dorsum to rusty brown, with tiny black flecks and uniform yellow to olive green. Dorsolateral streaks anteriorly. head region with black-bordered longitu- Maximum plastron length shorter than dinal yellow to olive-green blotch, inter- carapace length (PL/CL = 0.93 to 0.95), rupted posteriorly, but continuing on neck slightly upturned anteriorly, with no hinge to its base as a separate narrow black-bor- between pectoral and abdominal scutes. dered stripe; blotch may be divided in two Anterior plastral lobe shorter than posterior distinct ocelli (single female; Fig. IB, C), lobe. Plastral forelobe width (across level of hourglass shaped (singlejuvenile female), or junction of humero-pectoral seam and lat- unconstricted (single adult male). Side of eral plastral margin) relatively wide (PW1/ head with similar black-bordered longitu- CL = X = 0.44 to 0.45; 0.445). Plastral dinal blotch immediately behind eye, con- hindlobe width (at lateral junction of ab- tinuous with anterior portion of dorsolat- dominal-femoral seam) relatively wide eral blotch in juvenile and male. Dorsum (PW3/CL = 0.51 to 0.53; X= 0.518). Plas- of neck dark brown to black, with medial tral hindlobe with shallow anal notch. Bridge darkblack-bordered salmon to orange stripe. moderately long (BL/CL = 0.35 to 0.37; X Chin and tomia unmarked yellow; bottom = 0.355); a single small axillary and inguinal of neck yellow anteriorly and orange or VOLUME NUMBER 105, 3 429 — Table 1. Extended. tative specimens (see Specimens examined) ofSacalia bealei and S. quadriocellata from public and private collections were mea- sured (by JBI) and compared following the IP/MCW GW/GL method of McCord & Iverson (1991). Dis- criminant function analysis (Fig. 2) of 13 0.328 1.893 character ratios (MCW/CL, CH/CL, MPL/ 0.335 (0.32-0.35) 1.378(1.27-1.49) CL, PW1/CL, PW3/CL, PW4/CL, GW/CL, 0.238 (0.20-0.27) 2.191 (1.70-2.55) 0.281 (0.26-0.31) 2.269(1.99-2.42) MGL/CL, IH/CL, IP/CL, IAB/CL, IF/CL, 0.212(0.18-0.24) 2.417(2.05-3.02) and IAN/CL) revealed that S. pseudocel- 0.219(0.20-0.23) 2.286(1.88-2.75) lata, S. bealei, and S. quadriocellata are each morphometrically distinct. Specifically, S. pseudocellata is distinguished by its slightly salmon posteriorly, with vague black mot- wider carapace, its much larger plastron tling. Triturating surfaces narrow and un- (both longer and with wider fore- and hind- ridged. lobes), its longer and narrower gular scutes, Anterior surface ofantebrachium covered and its slightly shorter interfemoral seam. with large, imbricate scales, the largest three That analysis also revealed several charac- or four of which are sickle-shaped; largest terratios useful in discriminatingamongthe scales on hindlimb at heel, but much small- three taxa (Table 1, Fig. 3). er than largest forelimb scales. Upper parts Other taxonomic conclusions.—The dis- oflimbs and tail finely scaled. Exposed parts criminant function analysis also demon- oflimbs dark olive green laterally, but im- strated conclusively that the syntypes of maculate orange or salmon ventrally; bot- Cistuda bealei Gray are morphometrically tom ofhindfeet also salmon to orange. Re- identical to specimens from the Chinese cessed areas of skin salmon to orange. Tail mainland from Fujian and eastern Guang- salmon ventrally, anddark olive brown dor- dong Provinces, China and from Hong A sally, with a vague lighter dorsolateral line Kong. restriction of the type locality of on each side. Male with larger, thicker tail Sacalia bealei to the Fuzhou region in Fu- than females; vent at level ofcarapace mar- jian Province, China (see also Pope 1929) gin in male, anterior to it in females. seems justified, since S. quadriocellata ap- Etymology.—In reference to the lack of parently ranges from western Guangdong black "pupils" in the eye-like markings on and eastern Guangxi provinces (Pope 1935) its neck, we name this distinctive species and Hainan Island (Schmidt 1927) west- Sacaliapseudocellata. The unusual mixture ward to Vietnam (Siebenrock 1903) and of Greek {pseudo for false) and Latin (ocel- Laos (Shiu, pers. comm.). lata for eyed) symbolizes the unusual com- The discriminant function analysis also bination ofChinese and American trade ac- demonstrated that S. bealei and S. quadri- tivities that have brought this new species ocellata are distinct morphometrically, but to light. bivariate plots of diagnostic characters re- Other material.—One juvenile female vealed overlap in individual mensural char- (topotypic) alive inthe collection ofWilliam acters (e.g., Fig. 3). However, of over 50 P. McCord (WPM); to be deposited on death wildlife trade specimens ofthe two taxa that UF in the collection. we have seen, none was intermediate in head Distribution. —Known only from the type colorpattern (Fig. 1E-H). Sacalia bealeican locality. be consistently distinguished from S. quad- Comparison with congeners.—Represen- riocellata by the presence ofblack flecks on PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 430 1.1. - - 4- D 3- quadriocellata u 2- D 1- O CM pseudocellata o- A -1- ^A A -2- a bealei -3- -4 -7.5 -5 -2.5 2.5 7.5 10 12.5 15 17.5 DF1 Fig. 2. Discriminant plot ofcombined analysis ofmale (solid symbols) and female (open symbols) Sacalia based on 13 characters (see text). First axis accounts for 91.4% ofvariation; second, 8.6%. Circles are Sacalia pseudocellata; triangles, 5. bealei; and squares, S. quadriocellata. .35 pseucJoceiiata .325 o A A Q.275] ft bealei 1.25 *A A C—L .225 PA D A .2 quadriocellata .175 I I • I • I I I • I • - . .225 .25 .275 .3 .325 .35 .375 .4 .425 .45 IF/PW3 Fig. 3. Bivariate plot of the relationships among species ofthe genus Sacalia based on the characters IF/ PW3 (interfemoral seam length/anterior width ofplastral hindlobe) and IP/MCW (interpectoral seam length/ maximum carapace width). Open symbols are females; solid, males. Species coded as in Fig. 2. VOLUME 105, NUMBER 3 431 the anterior dorsum ofthe head ofthe for- seam usually greater than 25% of mer and the uniform unflecked dorsum in maximum carapace width S. bealei . . the latter (see also Pope 1935, and Fu & b. Dorsum ofhead uniform in color, Zhao 1990). In addition, S. quadriocellata unmarked with black; anteriorpair has four obvious ocelli on the posterior dor- of ocelli on head nearly or as ob- sum of the head, whereas S. bealei nearly vious as posterior pair; interpec- always has much less obvious ocelli, with toral seam usually less than 25% of the posterior pair more obvious than the maximum carapace width anterior pair (Fig. 1G, H). Thus, our data S. quadriocellata support Fu & Zhao (1990) in recognizing S. bealei and S. quadriocellata as separate spe- Acknowledgments cies. Specimens examined.—(See Acknowl- This species would not have been col- edgments for abbreviations) S. pseudocel- lected without the efforts of Oscar Shiu of lata: China, Hainan Island (UF 81505-06; Hong Kong. Curators and collection man- WMP 1, alive). S. bealei: China, mainland agers at the American Museum of Natural (AMNH 35179, 103735; BMNH 1947.3.4.33 History (AMNH), the British Museum of and 1947.3.4.42; MCZ 58086 [formerly Natural History (BMNH), the Museum of AMNH MVZ UF 34198]; 23936; uncat.; Comparative Zoology at Harvard (MCZ), WPM 1-8 alive). S. quadriocellata: China, the Museum of Vertebrate Zoology at WPM mainland (UF uncat.; 1-8, alive); Berkeley (MVZ), and the Florida State Mu- (AMNH China, Hainan Island 28337, seum of Natural History (UF) facilitated (WPM 28339, 28341); and Laos 1-2 alive). specimen loans. D. R. Auth, J. R. Buskirk, M.-O. Rodel, and P. Grossmann provided literature, pertinent information, or other Key to the Genus Sacalia assistance. Support for Iverson was provid- la. Blotches on dorsolateral aspect of ed by Earlham College, the Joseph Moore head without distinct black "pu- Museum ofNatural History, and his family. pils"; plastron mostly black; cara- pace weakly tricarinate; interhu- meral seam less than 45% of Literature Cited interpectoral seam length; interfe- Bourret, R. 1941. Les tortues de l'lndochine.—In- moral seam less than 28% of an- stitutOceanographiquedel'lndochine 38:1-235. terior width ofplastral hindlobe Buskirk, J. R. 1989. NewlocalityrecordsforChinese . non-marine chelonians.—Chinese Herpetolog- S. pseudocellata ical Research 2(2):65-68. b. Ocelli on dorsolateral aspect of Fu,J., &E.Zhao. 1989. ThevalidityofSacaliaquad- head with distinct black "pupils"; riocellata.—Asiatic Herpetological Research 3: plastron with small black blotches, 120-132. streaks, or vermiculations; cara- Gray, J. E. 1831. Synopsis reptilium. I. Cataphracta, pace unicarinate; interhumeral tortoises, crocodiles, and enaliosaurians. Treut- seam greater than 45% ofinterpec- tel, Wurz & Co., London, 85 pp. 1870. Supplement to the catalogue ofshield . toral seam length; interfemoral reptilesin thecollection ofthe British Museum. seam greater than 28% of anterior PartI. Testudinata(tortoises). TaylorandFran- width of plastral hindlobe 2 cis, London, 120 pp. A 2a. Anterior dorsum of head finely Iverson, J. B. 1992. revised checklist with distri- bution maps ofthe turtles ofthe world. Iverson spotted with black; anterior pair of Publishing, Richmond, Indiana, 363 pp. ocelli on head much less obvious McCord, W. P., & J. B. Iverson. 1991. A new box than posterior pair; interpectoral turtle of the genus Cuora (Testudines: Emydi- 432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dae) with taxonomic notesand a key to the spe- eraugenschildkrote, Sacalia bealei (Gray cies.—Herpetologica 47:405-418. 1831).-Sauria 10(4):3-8. McDowell, S. B. 1964. Partition ofthe genus Clem- Sachsse,W. 1975. Chinemysreevesivar. unicolorund mys and related problems in the taxonomy of Clemmys bealei var. quadnoceI'I'ata—Auspra- the aquatic Testudinidae.—Proceedings of the gungen von Sexualdimorphismus der beiden Zoological Society ofLondon 143:239-279. "Nominatformen" (Testudines, Emydidae).— Merkle, D. A. 1975. A taxonomic analysis of the Salamandra ll(l):20-26. Clemmyscomplex (Reptilia: Testudines) utiliz- Schmidt, K. P. 1927. The reptiles ofHainan.-Bul- ing starch gel electrophoresis.—Herpetologica letin ofthe American Museum ofNatural His- 31:162-166. tory 54:395-465. Pope, C. H. 1929. Notes on reptiles from Fukien and Siebenrock, F. 1903. Schildkroten desostlichen Hin- other Chinese provinces.—Bulletin of the terindiens.—SitzungsberichtederAkademieder American Museum ofNatural History 58:335- Wissenschaften in Wien, Mathematisch-Natur- 487. wissenschaftliche Klasse 1 12(1):333—352. . 1935. The reptiles ofChina. Turtles, croco- dilians, snakes, lizards. Natural history ofcen- (JBI) Department of Biology, Earlham tral Asia. Vol. X. American Museum ofNatural College, Richmond, Indiana 47374, U.S.A.; History, New York, 604 pp. (WPM) East Fishkill Animal Hospital, Rodel, M.-O. 1985. ZumVerhaltenvonSacaliabeal- ei(Gray 1831).-Salamandra 21(2/3):123-131. Hopewell Junction, New York 12533, & G. Praedicow. 1988. Die Chinesische Vi- U.S.A. ,