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Original article A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae) § Révision des caméléonidés du Miocène inférieur de République tchèque, avec description d’une nouvelle espèce de Chamaeleo (Squamata, Chamaeleonidae) Andrej C ˇ ern ˇansky ´ Department of Geology and Paleontology, Faculty of Natural Sciences, Comenius, University in Bratislava, Mlynská dolina, 84215 Bratislava, Slovakia Received 10 November 2009; accepted 11 April 2010 Available online 6 August 2010 Abstract A revision of Chamaeleo caroliquarti Moody and Roc ˇek is presented. The comparisons of the holotypic left dentary with those of specimens subsequently assigned to C. caroliquarti and of the Recent species of Chamaeleo, Furcifer and Calumma is carried out. It is shown that the type dentaries of C. caroliquarti include two different morphotypes with the absence of unique features. Within the Recent chameleons, the exact determination of the individual species merely on the basis of the dentaries is impossible. The holotypic dentary of C. caroliquarti is basically identical with that of C. calyptratus. However, the same morphology of the dentary as present in C. caroliquarti is also present in other species of different genera such as Calumma globifer and Furcifer pardalis. The paratypic dentaries of C. caroliquarti have a different morphology to the holotype and are indistinguishable from that in the Recent C. chamaeleon. On the other hand, a new species of the genus Chamaeleo, C. andrusovi, is described on the basis of isolated cranial elements, which possess clear autapomorphic features. This material comes from the Lower Miocene (Ottnangian) zone MN 4 in the Dolnice locality of the Czech Republic, and it differs from Recent and fossil chameleons in the following combination of characters: (1) its typically developed strongly pustular ornamentation and its distribution on the external surfaces of the skull roofing bones; (2) the frontoparietal suture is digitiform with a well-developed, anteriorly directed mesial spine, and (3) the parietal bone narrows posteriorly at its midlength, it is not bowed dorsally and it does not contribute posteriorly to a dorsal sagittal crest. This new material expands our knowledge of the cranial anatomy of Lower Miocene chameleons. # 2010 Elsevier Masson SAS. All rights reserved. Keywords: Squamata; Chamaeleo andrusovi nov. sp.; Chamaeleo caroliquarti; Nomen dubium; Lower Miocene Résumé Dans le cadre de la révision de l’espèce Chamaeleo caroliquarti Moody et Roc ˇek, la comparaison de l’holotype dentaire gauche avec des spécimens attribués ultérieurement à C. caroliquarti ainsi qu’aux espèces actuelles de Chamaeleo, Furcifer et Calumma est réalisée. Nous montrons que les os dentaires de C. caroliquarti regroupent deux morphotypes différents, avec une absence de caractères uniques. Au sein des caméléons actuels, la détermination spécifique précise sur la base des os dentaires est impossible. L’holotype dentaire de C. caroliquarti est fondamentalement identique à celui de C. calyptratus ainsi qu’à celle d’autres espèces et d’autres genres, comme Calumma globifer et Furcifer pardalis. Les os dentaires paratypes de C. caroliquarti montrent des morphologies différentes de celle de l’holotype, et indistinguables de celle de l’actuel C. chamaeleon. D’autre part, une nouvelle espèce du genre Chamaeleo, C. andrusovi, est décrite à partir d’éléments crâniens isolés montrant des caractères autapomorphiques clairs. Le matériel provient du Miocène inférieur (Ottnangien, zone MN 4) de la localité de Dolnice, près de Cheb, en République tchèque. Cette nouvelle espèce diffère des autres espèces actuelles et fossiles de caméléons par la combinaison des caractères suivants : (1) une ornementation pustuleuse fortement développée typique, et sa distribution sur les surfaces externes des os du toit du crâne ; (2) une suture antérieure digitiforme avec une épine médiane bien développée et dirigée vers l’avant et (3) un rétrécissement bilatéral Geobios 43 (2010) 605613 § Corresponding editor: Gilles Escarguel. E-mail address: [email protected]. 0016-6995/$ see front matter # 2010 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2010.04.001

A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae)

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Page 1: A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae)

Original article

A revision of chamaeleonids from the Lower Miocene of the CzechRepublic with description of a new species of Chamaeleo

(Squamata, Chamaeleonidae)§

Révision des caméléonidés du Miocène inférieur de République tchèque, avec descriptiond’une nouvelle espèce de Chamaeleo (Squamata, Chamaeleonidae)

Andrej CernanskyDepartment of Geology and Paleontology, Faculty of Natural Sciences, Comenius, University in Bratislava, Mlynská dolina,

84215 Bratislava, Slovakia

Received 10 November 2009; accepted 11 April 2010

Available online 6 August 2010

Abstract

A revision of Chamaeleo caroliquarti Moody and Rocek is presented. The comparisons of the holotypic left dentary with those of specimenssubsequently assigned to C. caroliquarti and of the Recent species of Chamaeleo, Furcifer and Calumma is carried out. It is shown that the typedentaries of C. caroliquarti include two different morphotypes with the absence of unique features. Within the Recent chameleons, the exactdetermination of the individual species merely on the basis of the dentaries is impossible. The holotypic dentary of C. caroliquarti is basicallyidentical with that of C. calyptratus. However, the same morphology of the dentary as present in C. caroliquarti is also present in other species ofdifferent genera such as Calumma globifer and Furcifer pardalis. The paratypic dentaries of C. caroliquarti have a different morphology to theholotype and are indistinguishable from that in the Recent C. chamaeleon. On the other hand, a new species of the genus Chamaeleo, C. andrusovi,is described on the basis of isolated cranial elements, which possess clear autapomorphic features. This material comes from the Lower Miocene(Ottnangian) zone MN 4 in the Dolnice locality of the Czech Republic, and it differs from Recent and fossil chameleons in the followingcombination of characters: (1) its typically developed strongly pustular ornamentation and its distribution on the external surfaces of the skullroofing bones; (2) the frontoparietal suture is digitiform with a well-developed, anteriorly directed mesial spine, and (3) the parietal bone narrowsposteriorly at its midlength, it is not bowed dorsally and it does not contribute posteriorly to a dorsal sagittal crest. This new material expands ourknowledge of the cranial anatomy of Lower Miocene chameleons.# 2010 Elsevier Masson SAS. All rights reserved.

Keywords: Squamata; Chamaeleo andrusovi nov. sp.; Chamaeleo caroliquarti; Nomen dubium; Lower Miocene

Résumé

Dans le cadre de la révision de l’espèce Chamaeleo caroliquarti Moody et Rocek, la comparaison de l’holotype dentaire gauche avec desspécimens attribués ultérieurement à C. caroliquarti ainsi qu’aux espèces actuelles de Chamaeleo, Furcifer et Calumma est réalisée. Nousmontrons que les os dentaires de C. caroliquarti regroupent deux morphotypes différents, avec une absence de caractères uniques. Au sein descaméléons actuels, la détermination spécifique précise sur la base des os dentaires est impossible. L’holotype dentaire de C. caroliquarti estfondamentalement identique à celui de C. calyptratus ainsi qu’à celle d’autres espèces et d’autres genres, comme Calumma globifer et Furciferpardalis. Les os dentaires paratypes de C. caroliquarti montrent des morphologies différentes de celle de l’holotype, et indistinguables de celle del’actuel C. chamaeleon. D’autre part, une nouvelle espèce du genre Chamaeleo, C. andrusovi, est décrite à partir d’éléments crâniens isolésmontrant des caractères autapomorphiques clairs. Le matériel provient du Miocène inférieur (Ottnangien, zone MN 4) de la localité de Dolnice,près de Cheb, en République tchèque. Cette nouvelle espèce diffère des autres espèces actuelles et fossiles de caméléons par la combinaison descaractères suivants : (1) une ornementation pustuleuse fortement développée typique, et sa distribution sur les surfaces externes des os du toit ducrâne ; (2) une suture antérieure digitiforme avec une épine médiane bien développée et dirigée vers l’avant et (3) un rétrécissement bilatéral

Geobios 43 (2010) 605–613

§ Corresponding editor: Gilles Escarguel.E-mail address: [email protected].

0016-6995/$ – see front matter # 2010 Elsevier Masson SAS. All rights reserved.doi:10.1016/j.geobios.2010.04.001

Page 2: A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae)

postérieur du pariétal à mi-longueur, pariétal qui n’est pas arqué dorsalement et qui ne contribue pas postérieurement à une crête sagittale dorsale.Ce nouveau matériel élargi notre connaissance de l’anatomie crânienne des caméléons du Miocène inférieur.# 2010 Elsevier Masson SAS. Tous droits réservés.

Mots clés : Squamata ; Chamaeleo andrusovi nov. sp. ; Chamaeleo caroliquarti ; Nomen dubium ; Miocène inférieur

1. Introduction � the taxonomic revision of the species C. caroliquarti Moody

A. Cernansky / Geobios 43 (2010) 605–613606

The Chamaeleonidae is a bizarre family of lizards, whichincludes extant species from Africa, Madagascar, southwestAsia, the Mediterranean region of Europe and extinct speciesfrom the Miocene in the Czech Republic and Germany (Estes,1983) and in the Pleistocene of Spain. Five fossil species of thegenus Chamaeleo have been recovered from the Miocene ofEurope: C. caroliquarti Moody and Rocek, 1980 (MN 4, CzechRepublic and Germany), C. bavaricus Schleich, 1983 (MN 6,Germany), C. pfeili Schleich, 1984 (MN 4b, Germany), C.simplex Schleich, 1994 (MN 5, Germany) and C. sulcodentatusSchleich, 1994 (MN 5, Germany). From the Pleistocene ofSpain, the Recent species C. chamaeleon has been recorded(Talavera and Sanchiz, 1983).

Chameleons of the Czech Republic were first described byMoody and Rocek (1980) from the Lower Miocene (MN 4) inthe Dolnice locality. They described a new species ofchameleon, C. caroliquarti, based on several isolated dentariesand maxillae. Subsequently, Rocek (1984) attributed apostorbital from the same locality to this species. Chameleonsfrom a second Lower Miocene (MN 3) locality, Merkur inNorth Bohemia, comprise several maxillae and dentaries andthese were described by Fejfar and Schleich (1994) andVejvalka (1997, unpublished).

Moody and Rocek (1980) based their new species, C.caroliquarti, on a holotypic dentary and several paratypicmaxillae and several other fragmentary dentaries. The anteriorend of the Meckelian canal of the holotypic dentary turnsventrally and ends immediately anterior to the symphysis,which is oval in outline (Moody and Rocek, 1980: fig. 1, pl. 1,fig. 2). However, the paratypic dentaries are fragmentary, andthe anterior end of the Meckelian canal is straight penetratingthe lowermost portion of the symphysis, which is square inoutline (Moody and Rocek, 1980; this is particularly evident onthe relatively well-preserved dentary BSP 1937 II 19601: pl. 3,fig. 3). Hence, we have two different morphologies of theanterior portion of the dentary, and this requires revision of thecurrent understanding of the anatomy of the dentary ofchameleons. The new material from the Lower Miocenelocalities described below contributes to the solution of thisproblem.

This new chameleon material originates from bothpreviously known chameleon localities in the CzechRepublic (Dolnice and Merkur), and besides the maxillae,postorbital and dentaries previously described, it alsoincludes new cranial elements described here for the firsttime, thus expanding our knowledge of the cranial anatomyof Lower Miocene chameleons. Therefore, the aims of thispaper are:

and Rocek, 1980;� the description of new cranial material of chameleons from

the Lower Miocene localities in the Czech Republic.

2. Material and methods

This study is based mainly on fossils housed in theNational Museum at Prague, Czech Republic. Thesecomprise isolated elements collected by screen-washing orsurface prospecting from the Lower Miocene (Ottnangian),zone MN 4, of the Dolnice locality (Czech Republic) andon fossils housed in the Geological collection of theBilína opencast mine. This paper also deals with isolateddentaries collected by screen-washing or surface prospectingfrom the Lower Miocene (Eggenburgian), zone MN 3, ofthe Merkur-North locality (Czech Republic). All theChamaeleonidae skull-roofing bones can be assigned to asingle taxon, new species of Chamaeleo, on the basis of theunique external surface ornamentation. The material includesthe parietal (holotype), jugal, prefrontal, and postfrontalportion of the postorbitofrontal, as described in this paper.This same morphology is also present in the postorbitalportion of the postorbitofrontal (DP-FNSP 226), andincorrectly described by Rocek (1984) as belonging to thespecies C. caroliquarti. The new dentaries material isidentical to that of the paratypic dentaries of C. caroliquartidescribed by Moody and Rocek (1980). However, as can beseen below, they do not differ also from dentaries of theextant C. chamaeleon.

The following specimens of extant species of chameleonsof different ontogenetic stages deposited in the collections ofthe Department of Ecology, Comenius University in Bra-tislava, Faculty of Natural Sciences, have been used forcomparisons: C. calyptratus, DE 65, DE 74-77; C. Chamae-leon, DE 66; Furcifer pardalis, DE 80-81; Calumma globifer,DE 82-85. Comparisons based on the literature included recentdescriptions of extant species from Madagascar (Rieppel andCrumly, 1997) and North Africa (Schleich et al., 1996), andalso extinct species from Germany (Schleich, 1983, 1984,1994).

Abbreviations of repositories: Ah- number is theSGDB: geological collection of the Bílina opencast mineat Merkur-North, Czech Republic (Bílina city); DP-FNSP:Department of Paleontology, Charles University, Prague,Czech Republic; Pb: National Museum, Prague, CzechRepublic; DE: Collection of Department of Ecology,Comenius University, Bratislava, Slovakia; BSP: BayerischeStaatssammlung für Paläontologie und historische Geologiein Munich.

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Fig. 1. Comparison of the left dentaries in medial views: A. Fossil chamae-leonid Ah-772 SGDB with badly preserved anterior part. B. Ah-773 SGDB withwell preserved anterior part. C. Recent Chamaeleo chamaeleon DE-66. D.Recent Chamaeleo calyptratus DE–65.

A. Cernansky / Geobios 43 (2010) 605–613 607

3. Geological settings

Fossil chamaeleonids described in this paper come from twolocalities.

3.1. The Dolnice locality

Dolnice is located in the westernmost part of the Chebbasin near Cheb, at latitude/longitude of 508 05’ N/128 23’ E.The bones are derived from greenish calcareous marls andlimnic clay silts of the Lower Miocene (Ottnangian) zoneMN 4 and are interpreted as marginal, riparian facies (Fejfarand Kvacek, 1993). The fossil-bearing sediments are exposedat several sites in the roof of the local main brown coal seamof the basin. Besides chamaeleonids, this locality has alsoyielded rich materials of fossil lizards such as lacertids,cordylids and anguids (Klembara, 1979, 1981; Rocek,1984).

3.2. The Merkur-North locality

The locality is an opencast mine near Chomutov, which haslatitude/longitude of 508 25’ N/138 21’ E. The specimens arepreserved in grey calcareous marls at the base of the so-called‘‘Main Brown Coal Seam,’’ which are interpreted as reworkedvolcanic ash. This second Lower Miocene (MN 3) locality liesin the North West Bohemian rift. In addition to chamaeleonidsit yielded a rich material of various groups of amphibians andreptiles, e.g., frogs and lacertids (Vejvalka, 1997 unpublished;Cernansky and Joniak, 2009), choristoderans (Evans andKlembara, 2005), an anguiomorph lizard (Klembara, 2008) andsnakes (Ivanov, 2002). The sediments are also richlyfossiliferous with remains of limnic and terrestrial mollusks,plants, and mammals (Fejfar and Kvacek, 1993; Fejfar et al.,1997a, 1997b, 1998).

4. Systematic paleontology

In the following, a taxonomic revision of chameleoncurrently attributed to Chamaeleo caroliquarti (Moody andRocek, 1980) is presented. New materials of chameleons fromthe Lower Miocene of Czech Republic with clear diagnosticcharacters are described as a new species.

Order SQUAMATA Oppel, 1811.Family CHAMAELEONIDAE Gray, 1825.Genus Chamaeleo Linnaeus, 1758–1759.Chamaeleo caroliquarti Moody and Rocek, 1980 nomen

dubium.1980. C. caroliquarti – Moody and Rocek, p. 86, fig. 1B, pl.

1, figs. 1, 2; p. 88, pl. 2, figs. 1–4 , pl. 3, figs. 1, 2.1980. C. cf. caroliquarti – Moody and Rocek, p. 88, pl. 3,

figs. 3, 4, pl. 4, figs. 1–4.1994. C. cf. caroliquarti – Fejfar and Schleich, pp. 167–170,

pl. 1, figs. 1, 2.1997. cf. C. caroliquarti – Vejvalka, pp. 42–44, figs. 7, 8 , pl.

8, figs. 4–7.

Localities, horizons and material: (1) Merkur-North,Lower Miocene (Eggenburgian), zone MN 3: right dentaries(Ah-772 SGDB, 773 SGDB, 845 SGDB, 857 SGDB,859 SGDB, 860 SGDB, 861 SGDB), left dentaries (Ah-774 SGDB, 844 SGDB, 846 SGDB, 847 SGDB, 858 SGDB,862 SGDB, 863 SGDB); (2) Dolnice, Lower Miocene(Ottnangian), zone MN 4: left dentaries (Pb 02051, 02053),right dentary (Pb 02052).

Description:New material of dentaries: The new material described here

is identical with the paratypic dentaries of C. caroliquarti. Thedentary Ah-772 SGDB (Fig. 1(A)) is elongate and slightlyconvex laterally in the dorsal view and mediolaterallycompressed. In lateral view it heightens gradually posteriorly.The symphysial facet for the complementary dentary is squarein outline (Fig. 1(B)). An essentially straight subdental shelfsupports a single row of up to 10–12 typically tricuspid,acrodont teeth. A well-developed dental groove is situatedbelow the subdental shelf. The most prominent feature on thelingual side of the dentary is a deep groove for Meckel’scartilage (sulcus Meckeli; Fig. 1(A)), which is roofed by a well-developed supra-alveolar ridge and floored by a slightlythickened ventral edge (crista ventralis). For most of itsposterior length the Meckelian groove deepens and widens, butanteriorly, as best seen in Ah-773 SGDB (Fig. 1(B)). Itconstricts greatly as it extends beneath the symphysial facet, butremains narrowly separated from the ventral margin of the[()TD$FIG]

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A. Cernansky / Geobios 43 (2010) 605–613608

dentary by the ventral edge. The supra-alveolar ridge widensposteriorly; at about its mid-length and just below its ventralmargin a shallow notch marks the entrance of the anteriorlydirected anterior inferior alveolar canal. The posterior ventralcorner of the dentary is greatly elongated into a prominentangular process ( processus angularis), which is separated fromthe dorsal, distally incomplete coronoid process ( processuscoronoideus) and surangular process ( processus surangularis)by a deeply concave margin of the sinus surangularis. There isa series of rough-surfaced, antero-ventrally oriented, inter-dental grooves on the labial surface of the subdental shelf,which become most distinct at the midpoint of the dental series.The otherwise smooth labial surface of the dentary is piercedalong its upper margin by a longitudinal series of five foraminafor nerves and blood vessels ( foramina pro rami nervorumalveolarium inferiorum), whereas a shorter series of fourforamina occupies a more ventral position.

Dentition: All the teeth are acrodont, in contrast to thefamily Agamidae, where the anterior part of dentary displays apleurodont implantation (Moody, 1978; Moody and Rocek,1980). The first four posterior teeth of the dental series are farmore strongly developed than those of the remainder of theseries, which diminish in size anteriorly. In the posterior half ofthe dental series the teeth are tricuspid, with the central cuspbeing about twice the size of its fore and aft cusps, and with theformer cusp being slightly smaller than the latter. Anteriorly,the remaining portion of the series exhibits a gradualtransformation from bicuspid to monocuspid. The teeth arecompressed medio-laterally and the posterior edges of theirbases are slightly angled into a medial plane. The teeth exhibit aslight grooving of the crowns and are directed slightlyposteriorly. There is a slight, posterior swelling of the inter-dental gaps.

Remarks: The dentary of the fossil species C. bavaricus(Schleich, 1983) differs from the dentaries described in thispaper in the following features:

� the symphysial facet is triangular in shape;� Meckel’s groove is poorly developed;� the teeth are large and massive relative to the whole dentary;� the dentary is smaller.

The material of two Orleanium German species C. simplexand C. sulcodentatus, described by Schleich (1994), isfragmentary and makes comparison with other specimensextremely difficult. In contrast to the dentary material describedhere, the symphysial facet of C. simplex has a marginal bulgeand a small depression in its midpoint. The surface of the teethis smooth. The common feature is generally a similar squareshape of a symphysial facet and its base is also pierced byMeckel’s groove. C. sulcodentatus is characterized byfurrowed, high triangular teeth with bevelled mesio-distalmargins. The species C. pfeili, described by Schleich (1984;MN 4b), can be excluded from detailed comparison, since thisspecies is preserved only as fragments of maxilla.

Another extant species of chameleon, C. africanus, which isknown from North Africa and Greece (Böhme et al., 1998;

Kosuch et al., 1999), also exhibits similar dentary features.However, C. africanus differences are apparent:

� there are no narrow, antero-ventrally directed, labialinterdental grooves;� the dentary is not as slender.

According to a molecular based cladistic analysis presentedby Townsend and Larson (2002), C. calyptratus, C. africanusand C. chamaeleon represent a single clade in which C.africanus and C. Chamaeleo share a more recent commonancestor than either does with C. calyptratus.

Taxonomic status of C. caroliquarti: we have two differentmorphologies of the anterior portion of the dentary in the typematerial of the species C. caroliquarti, which are alsoobservable in the Recent species of chameleons. Thesemorphologies are not a matter of variability within a singlespecies. The holotypic dentary represents the first type. Theholotypic left dentary (DP-FNSP No. 101; Moody and Rocek,1980: fig. 1, pl. 1, fig. 2) served as the sole basis fordistinguishing C. caroliquarti from the typical Europeanspecies C. chamaeleon. The distinguishing features of theholotypic dentary include:

� the outline of the symphysial facet is slightly elliptical, ratherthan square;� the anterior-most portion of the Meckelian canal is curved

ventrally immediately in front of the symphysis, rather thancontinuing directly anteriorly;� the dentary is more slender than those in the Recent European

representative.

The present restudy of the holotypic dentary of C.caroliquarti shows that among the extant examined chameleonsspecies of Europe, Arabia, and Africa, the morphology of theholotypic dentary of C. caroliquarti (Moody and Rocek, 1980:fig. 1B, pl. 1, figs. 1, 2) is identical to that in the extant C.calyptratus (Fig. 1(D)). However, it is not possible to determineany of the Chamaeleo species, or in many cases, even the genus,based only on dentaries. The species Calumma globifer andFurcifer pardalis have almost identical dentary morphology tothat of C. calyptratus and C. caroliquarti.

The paratypic dentaries are very badly preserved, especiallywith respect to the observability of the continuance of Meckel’sgroove and the shape of the symphysis. In one case, the well-preserved paratypic dentary BSP 1937 II 19601 (Moody andRocek, 1980: pl. 3, figs. 3, 4) does not exhibit the features of theholotypic dentary DP-FNSP 101, and it represents a differenttype of morphology. Hence, these two types of dentarymorphology are characteristic for two different species, andthey do not occur in the same species as in the type material ofC. caroliquarti. Both morphologies of the anterior portions ofthe dentary are also present in Recent species of chameleonsand, therefore, are not unique. The paratypic dentaries BSP1937 II 19601 of C. cf. caroliquarti (Moody and Rocek, 1980:pl. 3, fig. 4) are basically identical to those of C. chamaeleon(Fig. 1(C)), particularly in:

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A. Cernansky / Geobios 43 (2010) 605–613 609

� the Meckelian canal continuing anteriorly directly above thesymphysis;� the outline of the symphysial facet being square.

The anterior end of the Meckelian canal is straight andpenetrates the lowermost portion of the symphysis, and thesymphysis is square in outline. For this reason, the exactdetermination of this fossil material without autapomorphicfeatures is impossible.

The new dentary specimens referred to here do not differ inany specific way from the paratypes described by Moody andRocek (1980) or those of the extant C. chamaeleon. Theposterior portion of the dentary Ah-772 SGDB (Fig. 1(A)) iswell preserved and the distinctive processus angularis and deepsinus surangularis separating it dorsally from the processuscoronoideus and processus surangularis are present andidentical to those in the Recent C. chamaeleon. Unfortunately,this portion of the dentary is not preserved in any of thespecimens described and figured by Moody and Rocek (1980)in their description of C. caroliquarti. The dentary DP-FNSPNo. 102 of C. caroliquarti from Dolnice (Moody and Rocek(1980: pl. 1, figs. 3, 4) and the dentaries BSP 1937 II 19602 and19603 of C. cf. caroliquarti from Bavaria (Eggenburg; Moodyand Rocek, 1980: pl. 4, figs. 1, 2, 3, 4) are badly preserved,especially in the region of the section of the Meckelian grooveand symphysial facet, hence their exact species determination istotally impossible.

The paratypic right maxilla of C. caroliquarti (DP-FNSPNo. 104), which belongs to an adult individual, exhibits anexternal surface sculpturing of the dorsal nasal processconsisting of irregular grooves, ridges and plain indistinctpustules (Fig. 2). This is in contrast to any of the recoveredfossil chamaeleonid skull roofing bones, which are describedbelow as a new species. In the recent material, the sculpturing[()TD$FIG]

Fig. 2. The right maxilla DP-FNSP No. 104, including sculptured nasal processand described by Moody and Rocek (1980) as Chamaeleo caroliquarti nov. sp.

of the dorsal nasal process of the maxilla is almost identical tothat on the prefrontal. The sculpturing between these two bonesis confluent. Without the whole skull, it is impossible toassociate the paratypic maxilla with other skull roofing bones.There are also no unambiguous associations with either type ofdentary and, therefore, no legitimate reason to place it togetherwith one type or the other. A similar situation applies to themaxillary fragment (Inv. No. 7408-3-1 University of Prague)described by Fejfar and Schleich (1994) from the localityMerkur-North as C. cf. caroliquarti, which lacks featuresnecessary for exact determination.

All the material of chameleons described by Moody andRocek (1980) as a new species is fragmentary and incomplete,and lacks the autapomorphic features necessary for an exactdetermination. In no case can it be determined to be a newspecies. For this reason, the taxon C. caroliquarti, withoutunique characters or the possibility to associate its type materialwith other cranial bones, is considered here as a speciesinquirenda and the associated specimen name is therefore anomen dubium.

Chamaeleo andrusovi nov. sp.(Figs. 3–6).1984. C. caroliquarti (part) - Rocek, p. 13, pl. 1, fig. 1.Etymology: Dimitrij Andrusov was a Slovak geologist of

Russian origin; his grandfather was the German archeologist H.Schliemann. Dimitrij Andrusov studied at the University in St.Petersburg and the Sorbonne in Paris. For a short time, he alsostudied and worked at the Charles University in Prague, afterwhich he moved to the Slovak Republic. He was the founderand first director of the Geological Institute of the SlovakAcademy of Sciences and the first professor of Geology atSlovak colleges. He had the most important influence on thedevelopment of Geological sciences in Slovakia.

Holotype: Pb 02056, nearly complete parietal but lackingthe posterior portion.

Paratypes: Right jugal (Pb 02057); left jugal (Pb 02058);left prefrontal (Pb 02059); postfrontal portion of the leftpostorbitofrontal (Pb 02050).

Type locality and horizon: Dolnice locality in the CzechRepublic, Lower Miocene (Ottnangian), zone MN 4.

Diagnosis: A small species of Chamaeleo distinguishablefrom other chamaeleonid species by:

� its typical strongly pustular ornamentation of the externalsurfaces of the skull roofing bones. Differs from adultChamaeleo calyptratus, Triceros hoehneli or Calummaglobifer in regard to the concentration, distribution andshape of the protuberances – mostly complicated pustularprotuberances are moderately spaced and rather evenlydistributed, covering the otherwise smooth external surface;� the digitiform mid-portion of the anterior parietal sutural

facet, including a small, anteriorly directed midsagittal spineor spike;� the posterior portion of the parietal, which narrows bilaterally

posteriorly and is not bowed dorsally, indicating the absenceof a posterior elevated midsagittal crest or casque. This last

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[()TD$FIG]

Fig. 3. Chamaeleo andrusovi nov. sp., holotype, Pb 02056. Parietal: A. Dorsal view; B. Ventral view.

[()TD$FIG]

Fig. 4. Chamaeleo andrusovi nov. sp., paratype, Pb 02057. Right jugal: A. External view; B. Internal view.

A. Cernansky / Geobios 43 (2010) 605–613610

feature is in contrast to all mainland African and Europeanrepresentatives of the genus Chamaeleo. A similar conditionis seen in some Malagasy chameleons, Brookesia and also insome African chameleons, e.g. Rampholeon, as well as insome species of the genus Calumma.

Description:Holotypic parietal: The single, unpaired holotypic parietal

Pb 02056 (Fig. 3), which roofed the braincase, lacks most of the

[()TD$FIG]

Fig. 5. Chamaeleo andrusovi nov. sp., paratype, Pb 02059. Left prefrontal inexternal view.

tapering posterior portion that gives it the typical chamaeleo-nid, triangular outline. Sutural facets for the frontals andpostorbitofrontals along the slightly convex anterior margin arepreserved and easily discernable, but not that of the squamosal.At about its mid-length, the parietal narrows posteriorly withslightly concave lateral margins which bordered the uppertemporal fossae (Fig. 3(A)). In contrast to most chamaeleonids,the parietal is not bowed dorsally, but rather it occupies ahorizontal plane and therefore did not contribute to a posterior

[()TD$FIG]

Fig. 6. Chamaeleo andrusovi nov. sp., paratype, Pb 02050. Postfrontal portionof the fused left postorbitofrontal in external view.

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A. Cernansky / Geobios 43 (2010) 605–613 611

elevated, midsagittal crest. The sutural margin along theanterior margin is digitiform at the midline area, with four ormore, small, short, narrow, rectangular spike-like projectionsthat are narrowly separated by smooth, saddle-like margins.There is no midline notch, but there is a slightly moredeveloped, anteriorly directed midsagittal spine or spike. Morelaterally, the sutural margin is irregularly wavy. A well-developed ornamentation of pustular protuberances that aremoderately spaced and rather evenly distributed covers theotherwise smooth external surface. The protuberances, how-ever, exhibit a gradual transformation toward the posteriormargin of the bone, becoming bigger, antero-posteriorlyelongated, and more crenulated. The internal surface of theparietal (Fig. 3(B)) is smooth, except for an anteriorly pointed,midsagittal ridge (crista medialis ossis parietalis) whichgradually enlarges as it extends from just posterior to theanterior margin to the incompletely preserved posterior margin.There is no parietal foramen.

Paratypic jugal: Two jugals, a right (Pb 02057; Fig. 4) and aleft (Pb 02058), are nearly complete and well preserved. Inlateral view, the jugal is crescent-shaped and can be divided intotwo processes, a suborbital process ( processus suborbitalis)and a postorbital process ( processus postorbitalis) which formthe postero-ventral and posterior margins of the orbit,respectively, and exhibit facets for the maxilla, postorbito-frontal, ectopterygoid, and squamosal. The postorbital processis more attenuated, tapering slightly to a sharp, acute angle,whereas the narrower suborbital process is slightly constrictedat its mid-length. At the junction of the two portions, a third,short, poorly defined process ( processus posteroventralis)extends postero-ventrally. A ventral facet of the suborbitalportion on the external surface of the jugal that contacted themedial surface of the maxilla is divided ( pars exterior and parsinterior areae maxillares ossis jugalis) by a blunt crest (carinainferior). This crest undoubtedly extended to the distal end ofthe suborbital process, but in both jugals the distal ends aremissing, making it possible to observe an internal canal fornerves and blood vessels. The maxillary facet ends ventrally ina sharp edge (carina zygomatica ossis jugalis), which continuesposteriorly and merges with the convex union of the twoprocesses. A low, but distinct, arched facet for the ectopterygoidis visible on the internal surface of the postorbital process,posterior to its thickened orbital margin. Extending the lengthof the external surface of the postorbital process in both jugalsis a double row of well-developed, pustular protuberances, withthose of one row alternating in serial position with those of theother row. The more posterior row contains seven protuber-ances and extends along the posterior edge of the process andends at the processus posteroventralis. The longer, moreanterior row of about eight protuberances, extends partly ontothe proximal portion of the suborbital process. Most of theprotuberances are lobate and crenulated, whereas some aresimple spike-like projections.

Paratypic prefrontal: The left prefrontal Pb 02059 (Fig. 5)lacks only a small ventral portion, which contacted the maxilla.It is narrowly triangular, gradually tapering to a sharply pointedapex as it curves slightly postero-dorsally with its posterior

margin forming the antero-dorsal portion of the orbit. The well-developed manubrium (manubrium ossis praefrontalis) is wellexposed in the posterolateral view, as is the sutural facet for thefrontal. A low but distinct supraorbital ridge (crista supraorbi-talis) extends the entire length of the manubrium. A smoothsurface of orbital lamina ( facies orbitalis) separating thesupraorbital ridge and the inferior margin (margo inferior) ofthe manubrium gradually widens ventrally. As in the other skullroofing bones, the external surface is ornamented with well-developed pustular protuberances. The protuberances thatborder the anterodorsal margin of the prefrontal are larger andmore crenulated and lobated than the smaller ones spread out onthe surface.

Paratypic postorbitofrontal: The postfrontal portion of thefused left postorbitofrontal Pb 02050 (Fig. 6), which formedpart of the lateral wall of the braincase and also contributed tothe skull roof, has an irregular pentagonal outline. Thepostorbital portion of the postorbitofrontal is not preservedhere. Although the sutural facet for the frontal is small, it isclearly discernable. The anterior orbital margin is verticallystraight, whereas the almost straight posterior margin, whilecontinuing to the postorbital, angles slightly anteroventrally.The entire element thins medio-laterally, and is best seen on theventral margin, from the thickened orbital margin to theposterior connection with the postorbital portion. The externalsurface exhibits an ornamentation of well-developed pustularprotuberances. These are most strongly developed along theorbital margin, where they are elongated dorso-ventrally andtypically deeply subdivided by juxtaposed fore and aft channelsinto three distinct ridges which extend distally to the summit ofthe protuberance, where some are separated by a shallow,concave emargination. The remainder of the external surfacepossesses approximately six greatly reduced protuberances,which are essentially aligned in vertical rows. They exhibit anirregular structure, being divided unevenly by a singlejuxtaposed pair of channels that may or may not divide thesummit of the protuberance. The interprotuberance surface ofthe postorbital becomes coarser posteriorly, but otherwise it issmooth and featureless.

Remarks: In a subsequent paper, Rocek (1984) assigned thepostorbital DP-FNSP 226 from the locality of Dolnice to C.caroliquarti, but no characteristics are given which wouldjustify this. The type of sculpturing of this postorbital isdifferent to that on the dorsal surface of the nasal process of theparatypic fragmentary maxilla, as originally described byMoody and Rocek (1980) (Fig. 2). The postorbital DP-FNSP226 exhibits strongly developed sculpturing of pustularprotuberances on the external surface of the skull roofingbones, as typical for C. andrusovi nov. sp. Therefore, the DP-FNSP 226 postorbital is considered here to belong to C.andrusovi nov. sp. Additionally, in adult chameleons, thepostorbital bone does not occur as an independent ossification.Therefore, the DP-FNSP 226 postorbital must represent onlythe postorbital portion of the compound postorbitofrontal. C.andrusovi nov. sp. cannot be compared to other fossil Europeanspecies because there are no homologous bones in otherspecies.

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5. Discussion and conclusions

The new material of skull roofing bones from the Dolnicelocality exhibits clear diagnostic characteristics, and shouldnow be described as a new species. The type material of C.caroliquarti is fragmentary, which makes comparisons withextant species extremely difficult. Revision shows that this typematerial of dentaries includes two different morphologies.Neither of these two types are unique and both are alsorepresented in Recent species. The exact determination of thetype material in the species C. caroliquarti is impossible, sinceit lacks autapomorphic features. Unfortunately, because suchdiagnostic characteristic states are absent, the exact determina-tion is impossible, as also is its association with the other skullroofing bones described herein. Therefore, this taxon is speciesinquirenda and the name C. caroliquarti is a nomen dubium(Stys, pers. comm.).

Only the fossil isolated cranial skull roofing bones from theLower Miocene of the Czech Republic have clearly definite andunique character of their features. This material has thereforebeen designated as the type material of the new species C.andrusovi. The parietal is designated as a holotype, because thecombined features of its shape and a mid-ventral ridge aretypical within the Chamaeleonidae (Rieppel and Crumly,1997), but this also has unique features.

The parietal of C. andrusovi nov. sp. shows the absence of aposterior dorsally orientated midsagittal crest or casque. As faras is known (Rieppel, 1987), in all mainland Africanrepresentatives of the genus Chamaeleo the parietal possessesa narrow, elevated midsaggital crest or casque on the narrowposterior portion of the parietal. Judging from the morphologyof the parietal in modern chameleons, the dorsal crest or casquein C. andrusovi nov. sp. was either very poorly developed orabsent. A similar condition is seen in some Malagasychameleons, such as Brookesia, and in some extant chame-leons in which the parietal narrows posteriorly, never forming atrue midsaggital crest (Rieppel et al., 1992). Considering theabove observations, the following conclusions can betentatively made concerning the appearance of the dorsalcrest or casque:

� it does not appear in this oldest known chameleon, where theparietal is preserved;� in as much as it is absent in the agamids, it can be assumed

that the last common ancestor of the group of chamaeleonidsplus agamids also most likely lacked a casque;� furthermore, a casque is also absent in early juveniles of

modern chameleons, appearing only later in ontogeny.

This means that the juveniles present the same morpho-logical feature as their adult ancestor. From this point of view,the feature of the absence of a casque could be regarded as aplesiomorphic feature within chameleons. These morpholo-gical conditions indicate that the development of the dorsalelevated parietal crest or casque in extant chameleons is aconsequence of a peramorphic heterochronic process. How-ever, this can only be confirmed or denied by new fossil

records. Partial confirmation is provided, however, by the otherknown relatively younger chameleon specimens belonging tothe 18 million year old C. intermedius from the MiddleMiocene of Kenya. These consist of only the posterior part ofparietals that supported the casque, but they closely match theparietal in the extant Calumma brevicornis from Madagascar,which possesses a weakly developed dorsal crest or casque(Rieppel et al., 1992). Measey et al. (2009) examined thefunction of this structure. The evolution of ecomorphs within aspecies may represent either unique evolutionary events, ormultiple convergent events in similar environments. Thetypical form is known to use casque size to communicatefighting ability between males. According to Measey et al.(2009), open habitat males had wider heads, biting harder thanclosed habitat males. Their data suggest honesty in signalingfor closed habitat ecomorphs, but for open habitat ecomorphscommunication is different, and this constitutes a findingcommensurate with the common framework for speciesradiations.

The material of fossil taxa, described by Schleich (1983,1984, 1994) from the Middle Miocene of Germany, consistsonly of isolated fragments of dentaries and maxillae. For thisreason, these taxa can be excluded from a detailed comparisonwith the skull roofing bones of the species C. andrusovi nov. sp.In spite of some clear diagnostic characteristics of the Germanmaterial, we should be more cautious in identifying fragmen-tary isolated material of dentaries and maxillae at the specieslevel, especially as regard the description of new species. Thegeographic distribution of Recent chameleons living in Europeand the Near East is different to that found in the Miocene. Thedistribution of the C. calyptratus group extends from AsirProvince, southwestern Saudi Arabia, to Aden, Yemen (Fritzand Shütte, 1987; Meerman and Boomsma, 1987). Thedistribution of the Recent C. chamaeleon includes aMediterranean region of Europe and North Africa (Pauloet al., 2002), and it was also recorded in the Pleistocene of Spain(Talavera and Sanchiz, 1983). In southern Greece, there is alsoa small population of C. africanus (Böhme et al., 1998; Kosuchet al., 1999). The discoveries of chameleons in the LowerMiocene of the Czech Republic and Germany indicate thesubtropical climatic conditions of this region this time.Remains of chameleons (Chamaeleonidae indet.) have alsobeen recorded in the Middle Miocene of North-West Germany(Mörs, 2002).

Acknowledgements

I am indebted to D. S. Berman (Carnegie Museum of NaturalHistory), P. Stys (commissioner ICZN London; CharlesUniversity in Prague) and S. Sumida (California StateUniversity, San Bernardino) for their helpful advices. Mythanks belong also to J. Klembara (Comenius University inBratislava) and M. Baladová (Charles University in Prague).For critically reading the manuscript and the text corrections Ithank J.C. Rage (Université de Paris) and A.M. Bauer(Villanova University in Pennsylvania). This project wassupported by the APVV Grant agency G. No. 0280-07.

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