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A New Species of Zangerlia (Testudines: Nanhsiungchelyidae) from the Upper Cretaceous Redbeds at Bayan Mandahu, Inner Mongolia, and the Relationships of the Genus

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A New Species of Zangerlia (Testudines: Nanhsiungchelyidae) from the Upper Cretaceous Redbeds at Bayan Mandahu, Inner Mongolia, and the Relationships of the Genus A new species of Zangerlia (Testudines: Nanhsiungchelyidae) from the Upper Cretaceous redbeds at Bayan Mandahu, Inner Mongolia, and the relationships of the genus Donald Brinkman and Jiang-Hua Peng Abstract: Zangerlia neimongolensis sp.nov. is described on the basis of material from the Upper Cretaceous redbeds at Bayan Mandahu in Inner Mongolia. Zangerlia neimongolensis is similar to Zangerlia testudinimotpha in the proportions of the carapace and plastron and presence of a knob at the posterior end of the neural series, but differs from it in the arrangement of scutes covering the bridge. The placement of Zangerlia in the Nanhsiungchelyidae is supported by derived features of the bridge peripherals and plastral scutes shared by 2. neimongolensis, Basilemys, and Nanhsiungchelys. These are the presence of ventrally expanded sixth inframarginal scutes, humeral scutes that are narrow at the midline and expanded laterally, pectoral scutes that are wide at the midline and narrow laterally, and large rectangular abdominal scutes. The skull of Zangerlia is more primitive than that of Nanhsiungchelys, the only other member of the family for which a skull is known. It shows extensive emargination of the temporal and cheek regions and the absence of a large, tubular external narial opening. A cladistic analysis of the Trionychoidea using Zangerlia as the representative of the Nanhsiungchelyidae suggests a sister-group relationship between the Nanhsiungchelyidae and Adocidae. RCsumC : Nous dkcrivons Zangerlia neimongolensis n.sp. a partir de vestiges rCcupCrCs dans les couches rouges d'lge CrCtacC tardif, a Bayan Mandahu, Mongolie intCrieure. Les proportions de la carapace et du plastron et la prCsence d'une protubkrance I'extrCmitC postCrieure de la sCrie neurale de 2. neimongolensis sont similaires i ce qu'on observe chez Zangerlia testudinimotpha, mais il y a des diffkrences quant i la disposition des plaques qui recouvrent le dos. Le classement de Zangerlia parmi les NanhsiungchClyidCs est fond6 sur les caractkres dCrivCs des plaques pCriphCriques du dos et les Ccailles du plastron ventral communes i Z. neimongolensis, Basilemys et Nanhsiungchelys. Leur plastron ventral est formCe de six Ccailles inframarginales, d'Ccailles humCrales Ctroites au centre du plastron et s'klargissant For personal use only. lateralement, d'tcailles pectorales larges au centre du plastron se rCtrtcissant latkralement et de larges Ccailles abdominales rectangulaires. Le crsne de Zangerlia est plus primitif que celui de Nanhsiungchelys, seul autre membre de la famille dont le crlne a CtC CtudiC. Le crlne exhibe une Cmargination Ctendue des rCgions temporale et jugale, avec l'absence de grande ouverture tubulaire externe des narines. Une analyse cladistique des TrionychoidCs, utilisant Zangerlia cornrne reprCsentant des NanhsiungchClyidCs, suggCre une relation de groupe-soeur entre les NanhsiungchClyidCs et les AdocidCs. [Traduit par la rCdaction] Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by "National Science Library, Chinese Academy of Sciences" on 04/05/16 Received February 18, 1995. Accepted June 28, 1995. D. Brinkman.' Royal Tyrrell Museum of Palaeontology, P.O. Box 7500, Drurnheller, AB TOJ OYO, Canada. J.-H. Peng. Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica, P.O. Box 643, Beijing 100044, People's Republic of China. ' I Corresponding author (e-mail: [email protected] .ab. ca) . Can. J. Earth Sci. 33: 526-540 (1996). Printed in Canada / Imprim6 au Canada Brinkrnan and Peng 527 Pet$epa~ Zangerlia neimongolend sp.nov. onacrmaeTcR Ha ocHose MaTepaana a3 KpaCHOqBeTHbIX nnaCTOB B Eafi~~MaHAaXY BO BHYTP~HH~~~MoH~o~MH. Zangerlia neimongolensis cxoma c z. testudinimorpha B oTHomeHaa nponopqan KapanaKca a nnacTpoHa a npacyTcTsaR 6yrop~aB san~e~ Koaqe AopcanbHoro pHgar HO OTnaYawrcR pacnonoxeaseM CKYTOB, noKpHsamuax MocTnK. noMeueaae Zangerlia B ~anhsiungchelyidae nOATBepXAaeTCR yHaCnenOBaHHbIMI-3 XapaKTepHCTHKaMH nepa@epasec~ax neperoponoK (cen~)a nnacTpanbHbIx CKYTOB, 064ax An2 Z.neimonqolensis, Basilemys, B Nanhsiungchelys. T~KoBMM~RBnmoTcs npHCyTCTBHe BeHTpaJIbHO BHCTynaKluHX UeCTbIX HH&XLM~~~HH~~~H~IX CKYTOB, nneYeBMX uHTKOB, KOTOpbIe RBnHKlTCR y3KHMI-I B cpen~efinHHaH a pacuapeHHmMa naTepanbso, neKTopanbHMx CKYTOB, KoTopHe mapoKae B cpen~ennaHaa a ys~aenaTepanbao, a 6onbwax npxMoyronbHMx a6no~a~anb~b1xCKYTOB. Yepen Zangerlia 6onee npn~a~a~~anno CpaBHeHPiH3 C Nanhsiunqchel~,eAHHCTBeHHMM ApyraM YneHOM C~M~~CTB~, AnR KOTOpOI'O H3BeCTeH Yepen. OH HeMOHCTPWpYeT CYILleCTBeHHYD BbIeMYaTOCTb BHCOYHHX E-3 WeYHbIX o6nac~efia OTCyTCTBMe 60nbuoro Tpy65~aTor0BHemHerO HO3ApeBOOrO OTBepCTE'iR. AHanE33 KnaAOreHe3a Trionychoidea, nonbsy~cbZangerlia KaK npeAcTaBaTeneM Nanhsiungchelyidae, npeAnonomaTenbHo no~a3a~ae~cecTpaHcKo- For personal use only. rpynnoBoe ~saw~oo~~ome~aeMexqy Nanhsiungchelyidae a Adocidae. [nepeaon BbrnonHeH nnH penaKgm Hayq~o-Mccneno~arenbc~~eXyp~anbr] Introduction of the family. This was followed by Mlynarski (1976), who The Upper Cretaceous redbeds at Bayan Mandahu in Inner included these features in the diagnosis of the Dermatemy- Mongolia have yielded a diverse assemblage of fossil ver- didae. However, the presence of inframarginal scutes is a tebrates that is, in general, similar to that from the Djadokhta primitive character state within the Eucryptodira and dermal Formation in Mongolia (Jerzykiewicz et al. 1993; Dong sculpture varies among taxa included within the Dermatemy- 1993; Dong and Currie 1993). However, turtle remains are didae, so the placement of Zungerlia in the family is only much more abundant in the Bayan Mandahu locality. In con- weakly supported at present. trast to the two fragmentary turtle specimens collected in the Meylan and Gaffney (1989) used derived features to Djadokhta Formation of Mongolia (Gilmore 193I), seven resolve the relationships of many of the turtles placed in substantially complete shells have been collected at Bayan the Dermatemydidae by Mlynarski (1976). The family was Mandahu. Many are associated with postcranial elements, restricted to the genera Dermatemys and Baptemys and was and two include cranial remains. This material is herein placed with the ~inosternidaein the ~inosternoidae.Zunger- described as Zungerlia neimongolensis sp.nov. lia was united with Basilemys and Nanhsiungchelys in the Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by "National Science Library, Chinese Academy of Sciences" on 04/05/16 The genus Zungerlia was erected by Mlynarski (1972) for Nanhsiungchelyidae, and this family was placed with Adocus a large, terrestrial turtle from the "Lower Nemeget Beds" and the ~rion~chiawithin the ~rion~choidea.A sister-group of Mongolia (now the Barun Goyat Formation, Jerzykiewicz relationship between the Nanhsiungchelyidae and Peltochelys and Russell 1991). Mlynarski placed Zungerlia in the Der- plus Trionychia was supported primarily by features from matemydidae, because of the presence of inframarginal scutes the skull of Nanhsiungchelys. and dermal sculpture similar to that of some other members A close relationship between Basilemys and Nanhsiung- Can. J. Earth Sci. Vol. 33, 1996 Fig. 1. Zangerlia neirnongolensis, carapace and plastron. the Trionychoidea. It is concluded that Zungerlia is more (A, B) Type specimen, IVPP 020788-7, in (A) dorsal and closely related to Basilemys than is Nanhsiungchelys. A (B) posterior views. (C) Specimen IVPP 290690-6, in sister-group relationship between Adocus and the Nanhsiung- ventral view. chelyidae is suggested by the new information. Institutional abbreviations IVPP, Institute of Vertebrate Paleontology and Paleoan- thropology, Beijing; NMB, Nei Mongo Bowuguan (= Inner Mongolia Museum), Hohhot, Inner Mongolia; TMP, Royal Tyrrell Museum of Palaeontology, Drurnheller . Taxonomy Order Cryptodira Cope, 1868 Suborder Eucryptodira Gaffney , 1975 Superfamily Trionychoidea Fritzinger 1876 Family Nanhsiungchelyidae Yeh, 1966 Genus Zungerlia Mlynarski, 1972 Diagnosis Shell relatively short and and wide compared with other members of the family; posterior peripherals subvertical in position, not flared outwards; first suprapygal much smaller than second; a knob present at about the position of the first suprapygal and the posterior edge of the shell steeply deflected below this; anterior lobe of plastron subrectangular in shape; posterior lobe short compared with other members of the family, distance between the posterior end of the plastron and posterior end of the carapace greater than the length of the posterior lobe of plastron. Zangerlia neimongolensis sp.nov . (Figs. 1-9) For personal use only. Type specimen IVPP 020788-7, back half of shell, carapace preserved from the seventh neural, and plastron preserved posterior to hyoplastron-hypoplastron suture, skull, neck, girdles and limbs preserved within this portion of carapace, hind limbs and pelvis in articulation except for feet, remainder of skeleton disarticulated. Locality and horizon Inner Mongolia, Nuchidaba locality, about 9 km northeast of chelys was also suggested by Sukhanov and Narmandakh Bayan Mandahu, which lies about 45 km north of Urad (1977), who synonymized the two genera and described the Houqi (Dong 1993; Eberth 1993). All the turtle specimens shell and limbs of a species from Mongolia, "Basilemys" collected from this locality come from a limited area within orientalis. Brinkman and Nicholls (1993) argued that this the Nuchidaba locality referred to as "the gate" (Eberth synonymy is not justified, and that the two Asian
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