Pleistocene lizards (Squamata, Reptilia) from the karst caves in Chongzuo, Guangxi, southern China

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Pleistocene lizards (Squamata, Reptilia) from the karst caves inChongzuo, Guangxi, southern China

Jim I. Mead a,*, David Moscato a, Yuan Wang a,b, Changzhu Jin b, Yaling Yan b

aDepartment of Geosciences and Don Sundquist Center of Excellence in Paleontology, East Tennessee State University, Johnson City, TN 37614, USAbKey Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology,Chinese Academy of Sciences, Beijing 100044, China

a r t i c l e i n f o

Article history:Available online xxx

Keywords:LizardsGekkoChongzuoGuangxiPleistoceneChina

* Corresponding author. Department of GeosciencesExcellence in Paleontology, East Tennessee State U70357, Johnson City, TN 37614, USA.

E-mail address: [email protected] (J.I. Mead).

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a b s t r a c t

Here is provided the first description of Pleistocene lizards recovered from five caves (Baikong, Juyuan,Queque, Sanhe, and Zhiren) in Chongzuo, Zuo River area of Guangxi Zhuang Autonomous Region,southern China. Lizard remains reported here come from faunas that include Gigantopithecus blacki orHomo sapiens dating from the Early to Late Pleistocene with age-range estimates from 2.0 Ma to 111 ka.The recovery of agamids (Agamidae), skinks (Scincidae), a large species of Gekko, and smaller forms ofgekkonids (Gekkonidae) is of interest in that the diversity of lizards for the caves is high given theextremely small sample size from the deposits. The description of cf. Draco reported here is the firstindication of the possible occurrence of this genus in the Pleistocene record. Southern China and theadjacent territories in Vietnam and Laos have a diverse lizard fauna today. Cave deposits with establishedchronologies in the local karst regions such as recovered in Chongzuo, can help record the diversitywithin the local lizard community through time. Although the fossil lizards reported here appear torepresent mammalian prey remains (highly fragmented), caves that contain owl roost pellet depositsmight represent a more diverse squamate community with the additional advantage of having speci-mens that are not as fragmented. Because a Pleistocene-age lizard record does occur in the caves re-ported here, in all likelihood equivalent deposits should occur in the numerous caves throughoutsouthern China and Southeast Asia.

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1. Introduction

Published accounts of fossil squamates (lizards and snakes;Reptilia) from Asia are not abundant. Asian squamates from outsideof China are known from predominantly Paleogene-age sedimen-tary units (e.g., Averianov and Danilov, 1996; Prasad and Bajpai,2007; Head et al., 2013). Accounts from within China center onEocene and some Paleocene-age deposits (Chow, 1957; von Huene,1959; Young, 1961; Dong, 1965; Chow et al., 1973; Hou, 1974, 1976;Li, 1991a,b; Smith and Schaal, 2011). Squamates from Neogene andQuaternary deposits are less well understood, with a few reportsrepresenting the Miocene (Sun, 1961; Li et al., 1983), Pliocene(Patnaik and Schleich, 1998), and Pleistocene (Li et al., 2004; Joshiand Kotlia, 2010).

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The Pleistocene karst caves in Chongzuo along the Zuo Rivervalley of the Guangxi Zhuang Autonomous Region (Guangxi ZAR) ofsouthern China are known to contain a rich fossil record includingintriguing finds of the primates Gigantopithecus blacki and Homosapiens as well as a diverse associated vertebrate fauna dominatedby mammalian remains (Jin et al., 2008, 2009a,b; this volume; Liuet al., 2010) (Fig. 1). Numerous caves received multi-year excava-tions of their sedimentary deposits for the analysis of these faunalremains. Chronologies for these caves and their contents have beenestablished through detailed assessment of the mammalian re-mains (biochronological assignments) and in some cases paleo-magnetic and U-series radioisotopic analyses (Jin et al., 2009b; thisvolume; Liu et al., 2010; Sun et al., this volume). The vertebratefossils that receive the most detailed attention include taxonomicanalyses of primates (Zhao et al., 2008; Zhang et al., 2010; Harrisonet al., this volume), artiodactyls (Dong et al., 2011, 2013, thisvolume), rhinos (Yan et al., this volume), proboscideans (Wanget al., 2013), and small mammals (Jin et al., 2008, 2010; Wanget al., 2009, 2010). Vertebrates that largely have been neglected

amata, Reptilia) from the karst caves in Chongzuo, Guangxi, southern.2014.03.047

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J.I. Mead et al. / Quaternary International xxx (2014) 1e62

are the lizards and snakes. Here we provide the first description ofPleistocene lizards recovered from five caves in Chongzuo (Fig. 1;Table 1).

Table 1List of caves with lizard (Squamata) remains in the Chongzuo karst region, Guangxi,southern China e see text for clarification of identifications. Locality includes cavename, mountain in which cave is positioned, and IVPP (V) collection number. Ab-breviations: ka, thousands of years before present; Ma, megannum.

Locality Assigned age; Ma/ka; reference Lizard species

Zhiren Cave,Mulan Mtn.,V19020

Early Late Pleistocene; 111 ka;Jin et al., 2009b; Liu et al., 2010

Gekko sp., large

Queque Cave,Wuming Mtn.,V19019

Late Early Pleistocene; 1.0 Ma;Jin et al., and Sun et al., this volume

cf. Gekkonidae;cf. Sphenomorphini

Sanhe Cave,Wuming Mtn.,V19018

Middle Early Pleistocene; 1.2 Ma;Jin et al., 2009a; Jin et al., andSun et al., this volume

cf. Draco;Large agamid;Gekkonidae;cf. Plestiodon

Juyuan Cave,Boyue Mtn.,V 19017

Early Early Pleistocene; 1.8 Ma;Jin et al., and Sun et al., this volume

Lizard

Baikong Cave,Liyu Mtn., V19016

Early Early Pleistocene; 2.0 Ma;Jin et al., and Sun et al., this volume

Gekkonidae

1.1. Herpetofauna

China has one of the most taxonomically diverse living herpe-tofaunas (amphibians and reptiles) of any country (Zhao and Adler,1993; Zhao et al., 1999). Although possibly fairly well understood inits overall biological context, the details about the various taxo-nomic clades are still in their infancy because new species are stillbeing discovered and described in China. To better understand theliving and potential fossil lizard fauna of China it was found that thepublished herpetology from the surrounding region provides inmany cases greater details. The following references were con-sulted for Malaysia (Grismer, 2011a,b), Vietnam (Bobrov andSemenov, 2008; Nguyen et al., 2009), the Philippines (Gaulke,2011), and northern Eurasia (Szczerbak, 2003; Ananjeva et al.,2006).

As various groups of living herpetofaunas are collected andstudied, and voucher specimens are curated into national andregional museums, rarely are these specimens, or subsets of them,prepared for the study of their skeletal anatomy. This aspect of acomparative collection becomes of paramount importance whenanalyzing the latest Neogene and Quaternary-age herpetofaunasrecovered from fossil localities such those described here. As pre-sented by Bell andMead (2014) few collections curate an adequate-enough skeleton collection (ideally an ontogenetic series of artic-ulated and disarticulated specimens) of extant amphibians andreptiles to produce an acceptable analysis and identification ofrecovered herpetofaunas. That said, the identifications of fossilsdescribed here were attempted with the aid of the skeletal collec-tion housed in the Vertebrate Paleontology Laboratory, Departmentof Geosciences, East Tennessee State University, and from de-scriptions in the literature. Skeletal nomenclature follows Evans(2008) along with others as presented.

2. Geological and faunal settings

The caves in Chongzuo of Guangxi, located approximately120 km northeast of the China-Vietnam boundary and nearby theChongzuo Eco-Park (22� 16.49300 N, 107� 30.66300 E), are formed inthe karst limestone peaks known throughout the region. Due touplift and erosion during the late Neogene (Pliocene) and

Please cite this article in press as: Mead, J.I., et al., Pleistocene lizards (SquChina, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint

Pleistocene, cave entrances and their deposits formed along sixmajor elevations within the valley, with the oldest at the highestpositions on the peaks (Jin et al., 2009a,b; this volume). At present,lizard fossils are recovered from only a few cave deposits (Fig. 1;Table 1): Baikong Cave (Liyu Mountain), Juyuan Cave (BoyueMountain), Queque and Sanhe caves (Wuming Mountain), andZhiren Cave (Mulan Mountain). Lizard remains reported here comefrom cave faunas that include specimens of G. blacki or H. sapiensdating from the Early to Late Pleistocene with age-range estimatesfrom 2.0 Ma to 111 ka (for details see references listed in Table 1).

3. Lizard faunas

3.1. Taphonomy

Squamate remains (vertebrae and toothed elements) wererecovered by D. Moscato from sediments already pre-sorted andsieved by field and lab personnel. Toothed elements (dentary andmaxilla) were the predominant lizard remains recognized andrecovered from the caves. Often other skeletal elements (such asthe frontal, prefrontal, parietal, and quadrate) have more identi-fying diagnostic characters; these bones were not recovered fromthe deposits. Most of the skeletal elements retrieved are heavilyfragmented. Many of the specimens from the caves show evidenceof having been chewed by small carnivorans. Some of thedestruction observed on the fossils is likely taphonomic andhappened as a course of being incorporated into a cave deposit viafluvial or colluvial processes. Some of the broken edges appearrecent and could be due to screen-washing methodology. Althoughnot implemented for this field excavation, a procedure that uses�1.0 mmmesh sieves and a gentle stream of water or a water-bathsoaking with slight agitation typically produces many specimens ofoften good to excellent preservation, even from minute taxa.Identifications provided below are preliminary (‘cf.’) and at themost rudimentary level due to the preservation of specimens anddue to a lack of an available comprehensive comparative skeletoncollection (see 1.1).

3.2. Agamidae

The agamid lizard genera that live in southern China todayinclude species within Acanthosaura, Calotes, Draco, Japalura, Leio-lepis, Physignathus, Pseudocalotes, and possibly Mantheyus andPtyctolaemus (based on Zhao and Adler, 1993; Manthey, 2008, 2010;Nguyen et al., 2009). All agamids have an acrodont dentitionalthough most will have one or a few pleurodont teeth at theextreme mesial portion of the tooth row (Cooper et al., 1970). Theacrodont dentition typically will vary from being relatively small atthe symphysis to large at the distal end of the tooth row. The teethof some species can vary in the number of cusps from the mesial tothe distal portion of the tooth row (see Evans, 2008, and referenceswithin).

3.2.1. Sanhe CaveSpecimen V 19018.01 is a 4 mm fragment of a right maxilla

containing seven slightly-worn yet pointed acrodont teeth (Fig. 2Cand D; Table 1); the first which has both an anterocone and post-erocone. Based on the amount of wear and number of teeth, V19018.01 represents a small-to-minute taxon and not a hatchling orjuvenile of a larger form. The specimen is the anterior portion witha fragment of the facial process immediately posterior to the narialmargin. The facial process bends medially at about 45� from thetooth row, exhibiting a pronounced labial ridge (carina) just dorsaland parallel to the tooth row. The acute and medially-orientedfacial process illustrates that the lizard had a low, abbreviated,


Fig. 1. Map of China locating the Chongzuo caves (Table 1), Guangxi, containing Pleistocene lizard remains.

J.I. Mead et al. / Quaternary International xxx (2014) 1e6 3

and flattened nasal capsule such as found on Draco (extant today ingreater Southeast Asia), and small forms of Aphaniotis (PeninsularMalaysia, Indonesia), Ceratophora (Sri Lanka), Otocryptis (southernIndia, Sri Lanka), and Phoxophrys (Borneo) (Pethiyagoda andManamendra-Arachchi, 1998; Somaweera and Somaweera, 2009;Manthey, 2010; Grismer, 2011a,b), and is unlike the expressions ofthe facial region (maxilla and bones of the snout area) observed onAcanthosaura, Calotes, Japalura, Leiolepis, Physignathus, and Pseu-docalotes which live in southern China today. Of the genera thatappear to have themaxilla similar to V 19018.01, onlyDraco is foundin the region today (McGuire et al., 2007; Manthey, 2010). Directcomparisons were able to be made only with Draco, to which V19018.01 appeared near-identical. We accept that geographyshould not be the sole identifying character yet there is a generallack of availability of small agamid comparative skeleton speci-mens, thus we identify V 19018.01 as cf. Draco.

Specimen V 19018.2 is a fragment of a dentary containing threeheavily worn acrodont teeth. The specimen exhibits taphonomic


wear either via digestion or fluvial action. The specimen has theoverall size of the larger forms of agamid species, such as Acan-thosaura and Physignathus, but we feel that a satisfactory genericidentification cannot be made.

3.3. Gekkota

Gekkota encompasses the second most diverse clade of lizardsafter skinks (Scincidae). Currently Gekkota is considered tocomprise multiple clades/families of ‘geckos,’ including Pygopodi-dae, Carphodactylidae, and Diplodactylidae in Pygopodoidea, andGekkonidae, Phyllodactylidae, Sphaerodactylidae, and Eublephar-idae in Gekkonoidea (see discussion in Pianka and Vitt, 2003;Evans, 2008; Pyron et al., 2013; Daza et al., 2014). The Pygopodoi-dea are endemic to Australia, New Guinea, and the western Pacificislands and are not considered further here for our study. A typicalgekkonid dentary can resemble that of a few other Asian lizardssuch as dibamids (Dibamidae) and some scincids (Scincidae) in


Fig. 2. Lizard remains recovered from Chongzuo karst cave deposits along the ZuoRiver valley of the Guangxi, southern China. Gekko sp. from the early Late Pleistoceneof Zhiren Cave; A, right dentary (V 19020.1) and B, left dentary (V 19020.2). A rightmaxilla fragment identified as cf. Draco (V 19018.01); C, lateral view, D, medial view.Fragment of a right dentary of a skink (V 19018.4) identified as cf. Plestiodon; E, medialview; F, lateral view. Scale bars 2 mm.


having a closed and fused Meckelian groove. Often there is adistinct, wide dental gutter medial to the tooth bases; characters ofthe dentary and the teeth allow assignment to Gekkota (see Evans,2008).

Today there are numerous species of geckos living in easternand southeastern Asia. Zhao and Adler (1993) listed five genera(most within Gekkonidae) occurring in Guangxi area: Gehyra,Gekko, Goniurosaurus (Eublepharidae), Hemidactylus, and Hemi-phyllodactylus, and possibly Cyrtodactylus, Dixonius, and Ptycho-zoon; additional genera occur nearby in neighboring Vietnam,Thailand, Peninsular Malaysia, and greater Indonesia (Chan-Ardet al., 1999; Nguyen et al., 2009; Grismer, 2011a,b).

3.3.1. Zhiren CaveFive gekkonid dentaries were recovered including one complete

right (dentary row length 21.0 mm; V 19020.1; Fig. 2A) and onecomplete left (dentary row length 21.0 mm; V 19020.2; Fig. 2B),along with fragments of three others. Four of the specimens showexcellent preservation and all represent a large gekkonid species.Gekko is the largest of the geckos in Asia which has >10 speciesoccurring in China and many more in Vietnam, the PhilippineIslands, and elsewhere in Southeast Asia (at least 56 speciesdescribed; Grismer, 2011a,b). One of the largest species of Gekko isGekko gecko with a snout-vent length of 185 mm (Grismer, 2011a),and is known from the Guangxi region and elsewhere throughout


mainland Southeast Asia. Additional species outside of China occurwith long snout-vent lengths. The smaller Gekko adleri also isknown from the Guangxi region and has a snout-vent length<80 mm (Nguyen et al., 2013). The complete dentaries from ZhirenCave are near-identical in size to that of G. gecko with a snout-ventof 173 mm (dentary row length 22.5 mm). The dorso-ventral heightat the coronoid contact with the dentary is 4.5 mm on both den-taries from Zhiren Cave but that height on a G. gecko (svl¼ 173mm)is 7.0 mm. Although similar is overall size, the implication is thatthese fossils do not necessarily represent G. gecko, but clearly arelarger than all other gekkonids in the immediate region today. Thus,determining the species of Zhiren Cave Gekko is impractical at thistime without an adequate sample of skeletons of the various livingspecies; the fossil specimens (V 19020.1, V 19020.2) are identifiedas Gekko sp.

3.3.2. Queque CaveA fragment of a minute right dentary (3.5 mm long; V 19019.1;

Table 1) shows the closed Mecklian groove typical of gekkonids.Teeth are narrow, tall, with straight points for cusps. The identifi-cation is cf. Gekkonidae; further identification could not bedetermined.

3.3.3. Sanhe CaveV 19018.3 contains seven small (�5 mm length), highly frag-

mented dentaries and maxillae from a minute lizard taxon(Table 1). Three dentaries exhibit a pronounced dental gutter andclearly are a gekkonid. All elements contain delicate, parallel-sided,numerous teeth, all with pointed to blunt-pointed cusps. Due to thenumber of living genera in or potentially in southern China, onecannot satisfactorily identify any of these highly fragmented re-mains to genus. All these specimens are identified as Gekkonidae.

3.3.4. Baikong CaveThree dentaries and one maxilla (V 19016.1; Table 1) are heavily

fragmented by predatory chewing. All showaffinities with a speciesof minute gecko, Gekkonidae. Identifications could not be takenbelow Gekkonidae.

3.4. Scincidae

Skinks (Scincidae) comprise the largest and most species-abundant and morphologically diverse group of lizards in theworld today (Pianka and Vitt, 2003). Greer (1970) divided thefamily into four subfamilies: Scincinae, Acontinae, Lygosominae(includes the sphenomorphine skinks), and Feylininae (see updatediscussion in Hedges, 2014). As expected, China and especially thearea of Guangxi and nearby regions have an abundance of species ofmedium to minute skinks (Chan-Ard et al., 1999; Nguyen et al.,2009; Grismer, 2011a,b).

3.4.1. Queque CaveV 19019.2 is a fragment of a single minute right dentary (2.0 mm

long) containing an open Meckelian groove and diminutive,pointed teeth (Table 1). The identification can only be described cf.Sphenomorphini (due to the open Meckelian groove).

3.4.2. Sanhe CaveA fragment of a right dentary 4 mm long with five teeth was

recovered (V 19018.4; Table 1). There is no dental gutter, but teethare parallel-sided pillars with a squared-blunt cusp containing theapex in a sulcus oriented lingually. V 19018.4 shows all the affinitiesof belonging to large, robust form of Plestiodon (Fig. 2E and F;Scincinae) of which a few species exist in southern China andnearby regions today (Zhao and Adler, 1993; as Eumeces; Nguyen


J.I. Mead et al. / Quaternary International xxx (2014) 1e6 5

et al., 2009). Without an adequate skeleton collection of skinksfrom the region, the identification can only be stated as cf. Ples-tiodon, but there are likely other possibilities.

4. Discussion and conclusions

The caves in Chongzuo along the Zuo River valley of GuangxiZAR, southern China, produced a wealth of information aboutPleistocene-age faunal communities and environmental changesmainly through the detailed analyses of the recovered mammalremains. The micro-herpetofaunal remains received much lessattention until now. The squamates reported here from five cavesdating from approximately 2.0 Ma to 111 ka represent the firstlizard remains described in any detail from cave localities insouthern China. Most of the remains are highly fragmented andoften show breakage that can be assigned to the typical chewingaction by mammalian predators. This taphonomic aspect madeidentification difficult along with the lack of an adequate compre-hensive skeletal collection of lizard species living today in andadjacent to southern China. Most identifications made here areconsidered preliminary (cf.).

The recovery of agamids, skinks, a large species of Gekko, andsmaller forms of gekkonids is of interest in that the diversity oflizards for the caves is high given the extremely small sample sizefrom the five localities and the possible collection bias based onscreen-washing methodology. The implication is that with further,detailed sorting and delicate screenwashing, addition species couldbe recovered which ultimately should provide details as to how themodern-day lizard fauna developed.

The recovery of the minute maxilla showing affinities with thearboreal dragon cf.Draco (Fig. 2C and D) is intriguing.Draco is foundin the region of Sanhe Cave today, although there are at least 45species of this minute agamid living in Southeast Asia (McGuireet al., 2007). The description of cf. Draco reported here is the firstindication of the possible occurrence of this genus in the fossilrecord.

Even with an inadequate modern comparative collection avail-able for the project, some assumptions based on the identificationsimply that several interesting taxa lived in the region in the Early toLate Pleistocene (Table 1). No identifications represent taxa notfound in the region today; however, it is suspected that if and whenthe fossils can be identified to the species or species-group levelinteresting biogeographical information may well emerge. Thepreliminary identifications do show some level of diversity,including agamids, geckos, and skinks. Interestingly, nomembers ofVaranidae, Dibamidae, Anguidae, Shinisauridae, or Lacertidatewere noticed in any of the faunal remains recovered from the de-posits, yet representatives of these genera occur in southern Chinaand nearby regions today. Dentaries and maxillae of lizards withinthese families are distinct enough that they would have beennoticed had they been recovered. Our preliminary study here im-plies that they may not have been in the region during the Pleis-tocene, or alternatively, were not represented in the cave depositsamples we studied.

Guangxi and the adjacent territories in Vietnam and Laosobviously have a rich and diverse lizard fauna today. Cave depositswith established chronologies in these karst regions can help re-cord the diversity within the local squamate community throughtime. Although the fossil remains reported here appear to representmammalian prey remains (highly broken via chewing), caves thatcontain owl roost pellet deposits might have an additional advan-tage of containing specimens that are not as fragmented, therebypossibly permitting a greater number of identifications andincreasing the diversity of fossil taxa. Our project illustrates thatsquamates are readily recoverable from Pleistocene cave deposits


in Guangxi, thus in all likelihood equally-valuable herpetofaunasshould occur in the numerous caves found throughout southernChina and Southeast Asia. Increased attention to careful micro-mesh wet sieving should produce an elaborate squamate record.If one is to adequately reconstruct the evolving and changing her-petofaunal communities, then there is desperate need to create acomparative skeleton collection of all squamate species fromSouthern and Southeastern Asia.

Acknowledgements

We appreciate discussions with Larry Flynn (Peabody Museum),Jinyi Liu and Binghe Geng (Fossil Collections IVPP), Yusheng (Chris)Liu (National Science Foundation), Haowen Tong (IVPP), and YuanWang (Paleozoological Museum of China, IVPP) about the potentialof Chinese herpetofaunal deposits. We also thank Amanda Kauff-man for drawing our attention to the presence of lizard fossils inthese sediments. Initial support for the project was received in2006 from the American Philosophical Society for a FranklinResearch Grant to JI Mead. D Moscato received from the U.S. Na-tional Science Foundation an East Asia and Pacific Summer In-stitutes for U.S. Graduate Students award. This study was supportedby the Program of Chinese Academy of Sciences (KZZD-EW-03),National Natural Science Foundation of China (41072013 and41202017), and the Program of China Geological Survey(1212011220519). Additional funding was received from the Centerof Excellence in Paleontology and the Department of Geosciences,East Tennessee State University. We greatly appreciate the help anddiscussions of Sandra Swift and Kevin Chovanec, and Sandra forcreating the final illustrations. We appreciate the discussion andediting suggestions of Chris Bell, Chris Bae, and an anonymousreviewer.

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Pleistocene lizards (Squamata, Reptilia) from the karst caves in Chongzuo, Guangxi, southern China