Biogeographical affinities of Late Cretaceous continental tetrapods of Europe: a review

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<ul><li><p>Biogeographical affinities of Late Cretaceous continental tetrapods of Europe:a review</p><p>XABIER PEREDA-SUBERBIOLA1</p><p>Key-words Late Cretaceous, Europe, Laurasia, Gondwana, Tetrapods, Diversity, Palaeobiogeography.</p><p>Abstract. The continental tetrapod assemblages from the Santonian-Maastrichtian of Europe consist of dinosaurs (the-ropods: Abelisauroidea, Alvarezsauridae, Dromaeosauridae, ?Oviraptorosauria, ?Troodontidae, and birds: Enantiorni-thes, basal Ornithurae; sauropods: Titanosauria; ankylosaurs: Nodosauridae; ornithopods: Hadrosauridae,Rhabdodontidae; and neoceratopsians), pterosaurs (Azhdarchidae), crocodyliforms (eusuchians: Alligatoroidea, Gavia-loidea, ?Hylaeochampsidae; sebecosuchian-like ziphosuchians; and, probably, basal neosuchians), choristoderes(?Champsosauridae), squamates (lacertilians: Iguanidae s.l., Paramacellodidae, Polyglyphanodontinae, Varanoidea; sna-kes: Madtsoiidae; possible amphisbaenians), turtles (cryptodires: Chelydroidea, Kallokibotionidae, Solemydidae; pleu-rodires: Bothremydidae, Dortokidae), lissamphibians (Albanerpetontidae; anurans: Discoglossidae, Palaeobatrachidae;caudates: Batrachosauroididae, Salamandridae), and mammals (multituberculates: Kogaionidae, ?Paracimexomysgroup; marsupials: Herpetotheriidae; eutherians: Zhelestidae).</p><p>The palaeobiogeographical affinities of the Late Cretaceous continental tetrapods of Europe are complex. Thefaunas are commonly considered to show a mixed pattern resulting from the addition of Asiamerican and Gondwananforms to European taxa. Albanerpetontids, both paramacellodid and polyglyphonodontine lizards, and hadrosaurid dino-saurs are taxa with Palaeolaurasian (or, in some case, even Neopangean) affinities. Other forms, such as paleobatrachidand batrachosauroidid lissamphibians, solemydid turtles, alligatoroid crocodyliforms, and nodosaurid dinosaurs can beconsidered as Euramerican taxa. Kallokibotionid and dortokid turtles, rhabdodontid dinosaurs and kogaionid mammalsare endemic to Europe. The Gondwanan taxa have been regarded as African immigrants that reached southern Europevia the Mediterranean Tethyan sill. Abelisaurid and titanosaurid dinosaurs, trematochampsid crocodyliforms,podocnemidid and bothremydid turtles, and boid and madtsoiid snakes constitute the basic pattern of the so-calledEurogondwanan fauna. However, the validity of some of these taxa is a disputed matter (Titanosauridae,Trematochampsidae), and the presence of other taxa in the Late Cretaceous of Europe is based on controversial data(Boidae, Podocnemididae). Only Abelisauroidea, Madtsoiidae and Bothremydidae (and, yet for confirming,Sebecosuchia) provide evidence of interchanges between Africa and Europe. At least abelisauroids might have reachedEurope before the Late Cretaceous. In conclusion, most of the continental tetrapod taxa from the latest Cretaceous ofEurope show affinities with those of northern Hemisphere landmasses. Latest Cretaceous trans-Tethyan dispersal eventsbetween Africa and Europe remain poorly documented.</p><p>Affinits palobiogographiques des ttrapodes continentaux du Crtac suprieur dEurope :une mise au point</p><p>Mots-cls. Crtac suprieur, Europe, Laurasie, Gondwana, Ttrapodes, Diversit, Palobiogographie.</p><p>Rsum. Les associations de ttrapodes continentaux du Santonien-Maastrichtien dEurope incluent des dinosaures (th-ropodes : Abelisauroidea, Alvarezsauridae, Dromaeosauridae, ?Oviraptorosauria, ?Troodontidae, et oiseaux: Enantiorni-thes, Ornithurae basaux ; sauropodes : Titanosauria ; ankylosaures : Nodosauridae; ornithopodes : Hadrosauridae,Rhabdodontidae ; et nocratopsiens), des ptrosaures (Azhdarchidae), des crocodyliformes (eusuchiens : Alligatoroidea,Gavialoidea, ?Hylaeochampsidae ; ziphosuchiens de type sbcosuchien, et probablement des nosuchiens basaux), deschoristodres (?Champsosauridae), des squamates (lacertiliens : Iguanidae s.l., Paramacellodidae, Polyglyphanodontinae,Varanoidea ; serpents : Madtsoiidae ; possibles amphisbnes), des tortues (cryptodires : Chelydroidea, Kallokibotionidae,Solemydidae ; pleurodires : Bothremydidae, Dortokidae), des lissamphibiens (Albanerpetontidae ; anoures : Discoglos-sidae, Palaeobatrachidae; urodles : Batrachosauroididae, Salamandridae), et des mammifres (multituberculs : Kogaio-nidae, ?groupe Paracimexomys ; marsupiaux : Herpetotheriidae ; euthriens : Zhelestidae).</p><p>Les affinits palobiogographiques des ttrapodes continentaux du Crtac suprieur dEurope sont complexes.Ces faunes sont gnralement interprtes comme ayant un cachet mixte rsultant de laddition de formes asiati-co-amricaines et gondwaniennes aux taxons europens. Les albanerpetontids, les lzards paramacellodids et poly-glyphanodontins, ainsi que les dinosaures hadrosaurids sont des taxons qui ont des affinits palolaurasiatiques (voiremme pangennes dans certains cas). Dautres formes, comme les lissamphibiens palobatrachids et batrachosauro-dids, les tortues solemydids, les crocodyliformes alligatorodes et les dinosaures nodosaurids sont considrs commetant des taxons euro-amricains. Les tortues kallokibotionids et dortokids, les dinosaures rhabdodontids et les mam-mifres kogaonids sont endmiques lEurope. Les formes gondwaniennes ont t considres comme des immi-</p><p>Bull. Soc. gol. Fr., 2009, t. 180, no 1, pp. 57-71</p><p>Bull. Soc. gol. Fr., 2009, no 1</p><p>1. Universidad del Pas Vasco/EHU, Facultad de Ciencia y Tecnologa, Departamento de Estratigrafa y Paleontologa, Apartado 644, 48080 Bilbao.xabier.pereda@ehu.esManuscrit dpos le 13 juillet 2007; accept aprs rvision le 30 juin 2008</p></li><li><p>grants qui atteignirent le Sud de lEurope travers le Seuil mditerranen. Les dinosaures ablisaurids ettitanosaurids, les crocodyliformes trmatochampsids, les tortues podocnmidids et bothrmydids, et les serpentsboids et madtsoiids constituent le modle de base de ce qui a t appel la faune euro-gondwanienne. Cependant, lavalidit de certains taxons est conteste (Titanosauridae, Trematochampsidae) et la prsence de quelques autres dans leCrtac suprieur dEurope est base sur des donnes controverses (Boidae, Podocnemididae). Il en rsulte que seule-ment les Abelisauroidea, Madtsoiidae et Bothremydidae (la prsence de Sebecosuchia reste confirmer) attestent deschanges entre lAfrique et lEurope. Au moins les ablisaurodes auraient pu atteindre lEurope avant le Crtac sup-rieur. En dfinitive, la plupart des taxons de ttrapodes continentaux du Crtac terminal dEurope montrent des affini-ts avec ceux des continents de lHmisphre nord. Les dispersions trans-tthysiennes entre lAfrique et lEurope aucours du Crtac terminal demeurent mal connues.</p><p>INTRODUCTION</p><p>Fossil remains of Late Cretaceous continental vertebrateshave been known in Europe since the first half of the nine-teenth century. The first discoveries were made in southernFrance between 1825 and 1846 [Buffetaut et al., 1993; Ta-quet, 2001]. Later, new finds mainly linked to the exploita-tion of natural resources in mines and quarries and thedevelopment of the network of roads and railways led tocollect abundant vertebrate fossils in France and othercountries: Austria, Belgium, the Netherlands, Portugal,Transylvania (now Romania), Spain, and Ukraine [Brink-mann, 1988; Buffetaut and Le Loeuff, 1991; Le Loeuff,1991; Weishampel et al., 1991; Pereda Suberbiola, 1999;Laurent, 2003; Dalla Vecchia, 2006]. In the last few deca-des, further discoveries have been made in old and new lo-calities, including those that have been found in Germany,Hungary, Italy, Slovenia and Sweden [Wellnhofer, 1994;Debeljak et al., 2002; Dal Sasso, 2003; si, 2006; Lindgrenet al., 2007]. Systematic excavations and the use of screen-washing and other modern techniques have considerablyimproved our knowledge of the European continental biotasand led to a better understanding of their composition andaffinities [see Rage, 2002 and references therein]. Most ofthe fossiliferous localities are Campanian-Maastrichtian inage, but significant Santonian sites have been recently dis-covered as well [e.g., si et al., 2003], enlarging our pictureof Late Cretaceous tetrapod assemblages of Europe.</p><p>Although some tetrapod fossils have been found in ma-rine rocks, most of the finds have been made in continental(i.e., fluviatile, lacustrine or deltaic) deposits, where precisedating is often problematic [Buffetaut and Le Loeuff,1991]. Marine-continental correlations are difficult, andthus many Late Cretaceous dinosaur localities of Europe arenot well-constrained stratigraphically. In a few cases,high-resolution stratigraphic correlation between dino-saur-bearing sites and marine deposits is likely [Lpez-Martnez et al., 2001]. However, the accurate age of a num-ber of tetrapod sites is still imprecise and some correlationproblems remain to the adjustment of the charophyte scaleto biochronological evidences [Laurent et al., 2002a].</p><p>Late Cretaceous continental tetrapods of Europe consistof lissamphibians, squamates, turtles, choristoderes, croco-dyliforms, dinosaurs (including birds), pterosaurs, andmammals. Le Loeuff [1991: Tables 1-2] compiled the conti-nental tetrapods from the Campanian-Maastrichtian of sou-thern Europe and recognized 16 families. Seven of them areretained here, the others are not taken into account becauseeither their status is questioned or they are doubtfully repre-sented by fossils (see below). Only continental tetrapods are</p><p>here considered. Taxa whose fossils have been collected inmarine or transitional deposits in association to those of ty-pical continental tetrapods are indicated, but fully marinetetrapods are dismissed. In addition to the Campa-nian-Maastrichtian, the Santonian tetrapod record of Europehas been taken into account. Moreover, the pterosaur, birdand choristodere families have been added to the list. In to-tal, about 40 families (some of them are incertae sedis or in-determinate) are listed in table I for the Santonian-Maastrichtian of Europe (about 20 Ma; [Gradstein et al.,2004]). In some cases (e.g., crocodyliforms and dinosaurs),a taxonomic level higher than the family has been adopted.</p><p>THE CONTINENTAL TETRAPOD RECORD FROMTHE SANTONIAN-MAASTRICHTIAN OF EUROPE</p><p>LissamphibiansThey are represented by the Albanerpetontidae, anurans(Discoglossidae, Palaeobatrachidae) and caudates (Batra-chosauroididae, Salamandridae) [Duffaud and Rage, 1999;Duffaud, 2000; Folie and Codrea, 2005]. The European re-cord has provided the oldest occurrences of palaeobatra-chids and salamandrids [Duffaud, 2000]. Albanerpetontidsfrom the Late Cretaceous consist of the genus Albanerpetonin southern France and Romania [Duffaud, 2000; Folie andCodrea, 2005; Venczel and Gardner, 2005]. Albanerpetonwas first included into the Prosirenidae (Caudata) [Estes,1981], and posteriorly transferred to a new family distinctfrom Caudata [Fox and Naylor, 1982]. Prosirenid fossils arecurrently unknown in Mesozoic deposits [Rocek, 2000]. Eu-ropean batrachosauroidids consist of aff. Batrachosauroidesfrom southern France [Duffaud, 1997]. Discoglossids havea relatively good record in Transylvania, with the presenceof Paralatonia and forms closely related to Eodiscoglossusand Paradiscoglossus [Codrea et al., 2002; Venczel and Csiki,2003; Folie and Codrea, 2005]. Palaeobatrachids seem tohave affinities with the genus Palaeobatrachus [Duffaud,2000]. Finally, Hatzegobatrachus is provisionally referred toAnura incertae sedis [Venczel and Csiki, 2003].</p><p>SquamatesThe squamate fossil record consists of lacertilians, snakesand possible amphisbaenians [Rage, 1996, 1999; Folie andCodrea, 2005]. Lacertilians include nonacrodontan igua-nians (Iguanidae sensu lato [Sig et al., 1997; Rage, 1999]),scincomorphans (Paramacellodidae, Polyglyphanodontinae)[Folie and Codrea, 2005; Makdi, 2006], and anguimorphs</p><p>58 PEREDA-SUBERBIOLA X.</p><p>Bull. Soc. gol. Fr., 2009, no 1</p></li><li><p>(Varanoidea indet.) [Laurent et al., 2002a]. The Iguanidae isretained as a metataxon because its monophyly is neithersupported nor rejected [Estes et al., 1988]. In Transylvania,the paramacellodids are represented by Becklesius and thepolyglyphanodontines by Bicuspidon [Folie and Codrea,2005]. The Teiidae sensu stricto has no demonstrablepre-Tertiary occurrence: teiid-like scincomorphans from theCretaceous of North America, Europe and Asia are now pla-ced into the Polyglyphanodontinae [sensu Estes, 1983] as</p><p>borioteiioid lizards [Nydam et al., 2007]. Snake fossils ofEurope have been referred to the Madtsoiidae [Rage, 1996,1999; Sig et al., 1997; Folie and Codrea, 2005]. The Ibe-rian record includes species of Madtsoia and Herensugea[Rage, 1996, 1999]. The Madtsoiidae are now excludedfrom the Boidae [McDowell, 1987]. The occurrence of scin-coids in Central Europe is likely [Smith et al., 2002; Folieet al., 2005]. On the contrary, the presence of both Aniliidaeand Boidae [see Rage, 1981; Astibia et al., 1990] is doubt-ful [Rage, 1999, pers. comm.]. Iberian remains have tentati-vely been referred to the Amphisbaenia [Astibia et al.,1990], but this assignment is uncertain; a referral to anguidlizards cannot be ruled out [Rage, 1999]. The relationshipsof the lacertilian Araeosaurus from the Campanian of Aus-tria are uncertain.</p><p>Turtles</p><p>Both cryptodire and pleurodire fossils are abundant in theLate Cretaceous continental sites of Europe [Broin, 1977;Lapparent de Broin, 2001]. They have been referred to thetaxa Kallokibotionidae, Solemydidae, Chelydroidea, Both-remydidae and Dortokidae [Antunes and Broin, 1988; Gaff-ney and Meylan, 1992; Lapparent de Broin and Murelaga,1996, 1999; Lapparent de Broin, 2001; Gaffney et al.,2006]. Kallokibotion from Transylvania [Nopcsa, 1923] hasbeen regarded as the sister group of Selmacryptodira [Gaff-ney and Meylan, 1992], although Joyce [2004] excluded itfrom Cryptodria and even from Testudines [see Joyce et al.,2004; Gaffney et al., 2006 for a discussion on turtle syste-matics]. The solemydid Solemys is known in several locali-ties of the Iberian Peninsula and southern France[Lapparent de Broin and Murelaga, 1996, 1999]. The rela-tionships of Solemydidae with Pleurosternidae are not wellestablished owing to the absence of skull remains associa-ted with shells in solemydids [Lapparent de Broin and Mu-relaga, 1999]. Solemys is regarded as a junior synonym ofTretosternon and included within Kallokibotionidae by Hi-rayama et al. [2000]. Chelydroids are only represented byan unnamed taxon from Provence [Lapparent de Broin,2001]. Among pleurodires, the bothremydids, which in-clude both freshwater and littoral forms, were the dominantturtles in southwestern Europe [Lapparent de Broin andMurelaga, 1999; Lapparent de Broin, 2001]. They consist ofthe genera Elochelys, Foxemys, Polysternon, Rosasia [Antu-nes and Broin, 1988; Lapparent de Broin and Murelaga,1996, 1999; Tong et al., 1998; Laurent et al., 2002b] and anew, unnamed taxon [Rabi and Botfalvai, 2006]. Finally, dor-tokids are represented by the genus Dortoka in the Iberian Pe-ninsula and southern France [Lapparent de Broin andMurelaga, 1996, 1999]. There is no evidence of podocnemi-dids reaching Europe befor...</p></li></ul>

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