M. Delfino1, J.C. Rage2 & L. Rook1 1 Earth Sciences Department, University of Florence 2 Laboratoire de Paléontologie, Paris

Tertiary mammal turnover phenomena: what happened to the herpetofauna?

Delfino, M., Rage, J.C. & Rook, L., 2003 - Tertiary mammal turnover phenomena: what happened to the herpetofauna? - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 153- 161 [ISSN 0923-9308] Published 1 December 2003

Since the and Mammal Units are based on mammal turnovers that are the expression of more general changes in biodiversity, we question whether other vertebrates might contribute to define the biodiversity dynamics and consequently the limits of the stratigraphic units. Theoretically, due to their high sensitivity to environmental factors, and reptiles could act as precise signals of changes in biodiversity. How much did the European herpetofauna evolve during the Cenozoic? Did it experience any turnover phenomenon? Is it possible to find any synchrony in mammalian and herpetofaunal turnover? Focusing on some major mammal turnovers (i.e. those limiting different epochs) it appears evident that also the herpetofauna experienced some more or less abrupt changes that are sometimes perceivable at suprabasinial scale but are often detected in relatively restricted areas.

Correspondence: M. Delfino and L. Rook: Earth Sciences Department and Museum of Natural History (Section of Geology and Palaeontology), University of Florence, Via G. La Pira, 4, I- 50121 Florence, Italy, e-mail: [email protected]; [email protected]; J.C. Rage: Laboratoire de Paleontologie, UMR CNRS 8569, Muséum National d'Histoire Naturelle, 8 rue Buffon, F-75005 Paris, France, e-mail: [email protected]

Keywords: Tertiary, Europe, Amphibians, Reptiles, Mammal Turnover

INTRODUCTION suggested the application of term ‘biochron’ Vertebrate paleontologists have a long and for units of geologic time that are based on rich tradition of placing vertebrate evolution paleontologic data without reference to litho- into stratigraphic and biochronologic con- stratigraphy or rock units. On this conceptual texts. However, the term biochronology was basis, specific mammal biochronologic scales rarely used prior to the 1970s, when the have been constructed for North and South application of radiometric dating became America and Europe extensively, but also for widespread in geology, and the distinction Asia, Africa and Australia (cf. Lindsay 2001). was made between radiochronology and bio- For the Paleogene and Neogene of Europe, chronology as different aspects of geochrono- mammal paleontologists have developed two logy (cf. Berggren & Van Couvering 1974). unit systems, MP and MN respectively, dis- Based on the original definition of a ‘bioch- cussed among others, by Schmidt-Kittler et ron’ being ‘a time unit whose measure is the al . (1987), Vianey-Liaud (1997), Mein (1975, endurance of organic character’ (Williams 1989) and De Bruijn et al. (1992). Although 1901), Berggren & Van Couvering (1974) it has been stressed that units lack boundaries

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by definition (De Bruijn et al. 1992, Sigé & faunas and mammal faunas can be documen- Legendre 1997), an implicit idea of bounda- ted, then herpetofaunas should provide consi- ries cannot be denied (Van der Made 1997) so derable new information from both taxono- that Agustí (1999) recently proposed "to re- mic and a palaeoecologic points of view. This open the discussion on the MN unit system would improve the resolution of the ‘bounda- and to re-define and correlate them in terms ries’ between various units based on mammals. of faunal dispersals and extinctions (FAD and Theoretically, a broad analysis of the chan- LAD)". The ‘borders’ of these units (in other ges in the composition of European herpeto- words, the reference localities of each unit) faunas is biased by some main aspects: can therefore be defined with the aid of mar- țthe physiographic heterogeneity of different ked changes in the composition of the European regions through geologic ages; Tertiary mammal assemblages, changes that țthe presence of herpetological biogeogra- are relatively well known (for West Europe: phic units in pre-Pleistocene Europe Agustí et al. 2001) and frequently described (Barbadillo et al. 1997); and analysed in terms of dispersal events, țthe geographical and chronological non-uni turnover, provinciality, similarity and diversi- formity of the fossil record (e.g. Paleogene ty (cf. Bernor et al. 1996, Van der Made amphibians and squamates are scantily 1997). Fortelius et al. (1996a,b) recently syn- represented in Italy and Central Europe); thesised the theoretical and practical aspects țthe different status of the paleoherpetologi- of the study of these changes creating solid cal studies in different countries, an aspect bases for well-grounded future analyses. directly related to the local research tradi- If one considers that a mammal turnover is tions (among others, French, German, simply a signal of complex biodiversity dyna- Polish and Spanish paleoherpetofaunas are mics (e.g. addition of origination and extinc- particularly well known). tion of populations, taxa and communities; Taking into consideration the disparities lis- McKinney 1998) that results from changes in ted above, and limiting the analysis to family terrestrial paleoenvironments, it is worth and genus level, in order to avoid the doubt- noting that the rest of the terrestrial vertebra- ful nature of several specific allocations, te fauna has been taken into little account in some general issues can be outlined. For the the definition of these dynamics, perhaps also purposes of the present study crocodylians because it is usually not studied for strati- have not been considered. graphic purposes. It has been long said that amphibians and reptiles have little or no stra- THE MAIN TURNOVERS IN EURO- tigraphic significance. More in particular, as PE far as amphibians and reptiles are concerned, In the past decades, the researches dealing the terms ‘turnover’ and ‘biostratigraphy’ are with the fossil faunas of and reptile seldomly found in the European paleoherpe- from the European Cenozoic have produced a tological literature (interesting exceptions are huge amount of data. For example, Italy alone, represented by Vigne et al. 1997, and Gleed- whose Cenozoic fossil record has not been as Owen 1999). This is perhaps related to the extensively studied as those of other countries, widely accepted opinion that amphibians and offers at present more than 800 taxon/locality reptiles evolved gradually and that in the late data combinations (Delfino 2002). Although Quaternary, contrary to the mammals, they the European record has never been object of show little or no change (Holman 1995, a detailed wide scale analysis in terms of stra- 1998). In any case, this assumption does not tigraphical and geographical distribution, necessarily indicate stability or gradual evo- some major changes in the composition of the lution throughout the Cenozoic. Assuming European Tertiary herpetofauna are easily that synchronous changes between herpeto- detectable in correspondence with the limits

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between different epochs. eastern parts of the continent. Palaeocene/Eocene boundary /Miocene boundary Pleurosternid turtles, which were present in The transition between the Oligocene and Europe during the late Palaeocene, did not Miocene corresponds to a ‘modernisation’ of survive into the Eocene. Moreover, dicamp- the European herpetofauna but the change todontid salamanders temporarily disappeared was not an abrupt turnover. At family level, from the continent. On the other hand, the only cordylid lizards and tropidophiid snakes earliest Eocene (MP7) is marked by immigra- disappeared from Europe at the end of the tions, newcomers significantly increased the Oligocene (MP30). On the other hand, only European herpetofauna: pelobatid ; one family (viperid snakes) appeared in the podocnemidid and geoemydine turtles; iguan- earliest Miocene (MN1). However, in id, agamid, gekkonid, varanid, helodermatid Western Europe, a lineage of lacertid lizards and ascertained lacertid lizards; scolecophi- with ‘molariform’ teeth markedly developed dian, tropidophiid, and russellophiid snakes during the Oligocene. At the same time the (Rage & Auge 1993, Rage 1997, Duffaud number of erycine boids (fossorial snakes) 2000). As a result, the Palaeocene/Eocene strongly increased. Consequently, the limit represents an important turnover in squamate fauna at the end of the Oligocene European herpetofaunas. was highly peculiar, with very high percenta- ges of unusual lizards and fossorial snakes. Eocene/Oligocene boundary This squamate fauna suddenly disappeared at The limit between the Eocene and Oligocene the Oligocene/Miocene boundary (Rage marks a prominent turnover, at least in 1987). Given that "there is not a clear crite- Western Europe. Some changes can be detec- rion for separating the uppermost Oligocene ted at the family level, although squamates terrestrial levels from those of the lowermost show considerable modification at the species Miocene, the use of particular segments of level. By the end of the Eocene, russellophiid some evolutionary lineages excepted" (Agustí snakes became definitively extinct and lepto- 1999; see Agustí et al. 2001 for a detailed dactylid frogs, podocnemid turtles, and igu- discussion), the presence/absence of the anid lizards disappeared from the continent above mentioned herpetofauna should be (Lapparent de Broin 2001, Rage & Auge investigated in detail since they might be use- 1993). Pelodytid frogs temporarily withdrew ful markers. from Europe. At the species level, amphi- bians and non-squamate reptiles were appa- Other turnovers rently not prominently affected (Duffaud Apart from the three above-mentioned main 2000) but at least three-fourths of the turnovers, other changes have been identified, squamate species became extinct at the end but have not been studied in a comprehensive of the Eocene (Rage 1984). At the beginning way yet: MP22 during the Oligocene, of the Oligocene (MP21), a few survivors and early/middle Miocene transition (for snakes very rare immigrants formed a depauperated see Ivanov 2001), and perhaps MP15/MP16 herpetofauna. During the Early Oligocene in the late Eocene. Other minor changes (after MP21), a combination of Asian immi- might also be significant from a stratigraphic grants and autochthonously evolved species point of view, mainly in the Eocene and increased herpetofaunal diversity. Prior to the Oligocene. Eocene/Oligocene limit, the European herpe- tofauna had an Euro-American aspect; after The problem of the late Neogene this event, it had an Eurasian aspect. This No significant turnover can be detected event is well documented in Western Europe among the European herpetofauna at the but it is still poorly known in central and Miocene/ boundary although global

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vegetation and faunal changes have been thermic members of which are highly sensiti- reported (Cerling et al. 1997, and reference ve to temperature and humidity. A general therein quoted). Modifications in the herpeto- discussion of faunal changes in Europe is dis- logical communities are apparently scattered cussed in Azzaroli et al. (1988) and evidences along the transition. As already noticed by have been reported from Spain, France, Italy Barbadillo et al. (1997) and Rage (1997), the (Alberdi et al. 1997, Gliozzi et al. 1997) and late Neogene is apparently less affected by Eastern European countries (summarised by marked faunal changes in the sense that, at Vislobokova et al. 1993). least at family level, there are only few Although attractive, this hypothesis is per- entries and, on the whole, it seems to be haps biased by the fact that the presence in dominated by a progressive decrease in diver- the middle Pliocene (MN16) of one of the sity. The late Neogene herpetofauna consisted more informative Pliocene herpetofaunas of a mixture of taxa that were mainly relics (Balaruc II; Bailon 1989, 1991) is not balan- of the thermophilous Miocene fauna, or new ced in the late Pliocene (MN17) by any immigrants from Asia or North Africa equally representative fauna. Moreover, the (Bailon 1991): curiously, amphibians show recent split of the Pliocene into three parts no direct exchange with Africa (Sanchiz (Rio et al. 1994) is potentially misleading 1997) but only with Asia, whereas squamates since some of those sites previously conside- immigrate from both (Rage & Auge 1993). red to be late Pliocene, could be middle Pliocene in age: a critical review of the chro- All the families (as well as the most of the nological allocation of several sites is there- genera and several species) of the modern fore required. European herpetofauna possess a fossil It is worth mentioning that at the same record that goes back at least to the Pliocene time, several reptile taxa that are now restric- when Europe was still inhabited by some ted to the Balkans and few Aegean Islands families and genera that are now definitely were still present in West Europe (Bailon extinct or that we can consider exotic. With 1991): Agamidae, Scolecophidia, Boidae, the beginning of the climatic deterioration, large Viperidae (the ‘Oriental vipers’). we can perceive a whole string of extinctions Starting from the late Pliocene their ranges and range re-arrangements. The timing of markedly shrunk leading to a further decrease these changes is hard to define in detail since in diversity mainly in West and Central the late Pliocene (MN17) is represented by a Europe. This N-S and W-E contraction of the limited number of sites. Assuming that the amphibian and reptilian ranges continued late Pliocene record is sufficiently informati- during the Quaternary when Europe lost the ve, a sharp drop in species diversity can be Hynobiidae (Venczel 1999) and when, at the perhaps identified at the middle-late Pliocene same time, the family Palaeobatrachidae transition: after MN 16, cryptobranchid sala- (Sanchiz 1998) and the genus Latonia defini- manders, varanid lizards, aniliid and elaphid tely disappeared (Delfino 2002). snakes and perhaps chelydrid turtles are not It should be noted that amphibians and rep- found in Europe (Broin 1977, Rage 1997). tiles, although traditionally placed together in This change could have been triggered by the the herpetofauna since they are the only ecto- earliest glacial-interglacial cycles and the thermic terrestrial vertebrates, display some severe coolings that Europe faced since 2,6 differences in terms of extinction and survi- million years (Bertini 1995, Suc et al. 1995), val patterns. The reptilian extinction pattern coolings that certainly disrupted the faunal follows not only a N-S trend but also a mar- assemblages (for Italian mammals see: ked W-E one. The amphibian extinction pat- Gliozzi et al. 1997) and particularly affected tern appears to be not so directional (e.g. the herpetofaunal communities, most ecto- palaeobatrachids survived only in Central

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Figure 1 Simplified distribution of selected herpetofaunal taxa of stratigraphic interest (crocodylians are not considered) showing the ‘time-slices’ in which significant turnover occured (see text for details). Appearances and extinctions are plotted on the figure only when they correspond to a turnover.When needed, dotted lines have been added to the black squares in order to avoid the false impression of ‘flash permanence’ of taxa; therefore, they do not represent the real distribution of the taxa. ° = taxa extinct in Europe; * = taxa extinct globally.

Europe before extinction) as they are highly of the fossil record suggests a withdrawal of sensitive to the environmental moisture that the ranges from W to E, withdrawal that has generally shows a more complex distribution not been completed for the taxa that still per- than temperature. As further evidence, we can sist in South Eastern Europe. Therefore, the consider that among those taxa (reptiles only) above-mentioned -often gradual- changes of that deeply restricted their range during the geographic ranges hamper the use of amphi- considered time span and that are still present bians and reptiles in the analysis of the late in Europe, many are widespread in the Neogene stratigraphy. Balkans or in the Aegean Islands only. This apparently contrasts with the fact that Iberia CONCLUSIONS is warmer than the Balkans at present. The comparison of the paleoherpetological Concerning this topic, Cheylan (1995) noti- record of the European Tertiary with that of ced that the centre of dispersion for most of the mammals is biased by several factors, the the European taxa was Asia and the speed of main one being the absence of data from the Plio/Pleistocene range shrinkage has been various areas and periods from which fossil higher in those regions (i.e. Iberia and amphibians and reptiles have not yet been France) that are the farthest from the centre studied, although it may be presumed that of dispersion. As a consequence, the analysis their remains have already been collected

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along with those of mammals. However, a in terms of time and space, a detailed analy- general analysis of the available record from sis would benefit from the development of a a chronological point of view allows us to broad taxon/locality data-set and an accuracy observe that the herpetofauna evolved, at of investigation similar to those that are cur- least prior to the late Neogene, through a rently performed by programs such as NOW series of turnovers influenced by the same (Neogene of the Old World; http://www.hels- external forces that, being responsible for inki.fi/science/now/) and EEDEN more general biodiversity dynamics, shaped (Environments and Ecosystem Dynamics of the terrestrial palaeoenvironments. the Eurasian Neogene; http://www.esf.org/life/ Several turnover episodes, corresponding to lp/ EEDEN/eedena.htm). some of the limits between different epochs, can be evidenced (Fig. 1): Paleocene/Eocene, ACKNOWLEDGEMENTS Eocene/Oligocene and Oligocene/Miocene We thank R. Bernor, M. Fortelius, F. de boundaries (but also MP 22 during the Lapparent de Broin, P. Mazza and D. Torre Oligocene, the early/middle Miocene transi- for providing useful information and profita- tion, perhaps MP15/MP16 in the late Eocene, ble discussion. This work has been developed and others). The pattern of evolution of the within the frame of researches supported by Tertiary European herpetofauna follows some MURST 1999 COFIN grant ‘Paleogeografia of the guidelines already identified for mam- del Mediterraneo centrale dal Miocene supe- mals. The turnover of herpetological faunas is riore al Pleistocene medio’ (scientific respon- mainly connected to East-West dispersal sible D. Torre). events as underlined for mammals by Werdelin & Fortelius (1997). REFERENCES Due to the fact that the geographic and Agustí, J., 1999 - A critical re-evaluation of the Miocene chronological distribution of the paleoherpe- mammal units in Western Europe: dispersal events tofaunistic data are far from being uniform at and problems of correlation - in: Agustí, J., Rook, L. the moment, a perusal of aspects such as pro- & Andrews, P. (eds.) - Hominoid evolution of vinciality and diachrony cannot be easily Neogene terrestrial ecosystems in Europe - pp. 84- accomplished. Nonetheless, faunal changes 112, Cambridge University Press, Cambridge that are markedly perceivable inside one Agustí, J., Cabrera, L., Garcés, M., Krijgsman, W., Oms, single region (French Late Oligocene; Rage O. & Parés J.M., 2001 - A calibrated mammal scale 1987) are, at least partly, the product of broad for the Neogene of Western Europe State of the art - renewals that left evidences even at supraba- Earth-Science Reviews 52: 247-260 sinial level. For example, the extinction of Alberdi, M.T., Azanza, B., Cerdeño, E. & Prado, J.L., lizards with ‘molariform’ teeth at the end of 1997 - Similarity relationships betweens mammal the Oligocene is perhaps a phenomenon res- faunas and biochronology from latest Miocene to tricted to one region (southern France) but, Pleistocene in the westernen mediterranean area - just after the extinction of these lizards, the Eclogae Geologicae Helvetiae 90: 115-132 genus Vipera certainly appeared across most Azzaroli, A., De Giuli, C., Ficcarelli, G. & Torre, D., of Europe. 1988 - Late Pliocene to early Mid-Pleistocene Although it is unlikely that the biochrono- Mammals in Eurasia: Faunal Succession and logic use of the European Cenozoic herpeto- Dispersal events - Palaeogeography,Palaeo- fauna will ever attain the resolution of mam- climatology, Palaeoecology 66: 77-100 mals, it may be possible to foresee a role of Bailon, S., 1989 - Les Amphibiens et les Reptiles du these vertebrates in augmenting mammals for Pliocène Supérieur de Balaruc II (Hérault, France) - biochronologic and, even more so, paleoeco- Palaeovertebrata 19 (1): 7-28 logic characterisations. Because of the high Bailon, S., 1991 - Amphibiens et reptiles du Pliocène et number of taxa involved and the wide range du Quaternaire de France et d’Espagne: mise en place

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et évolution des faunes - Ph.D. Thesis, Université de Delfino, M., 2002 - Erpetofaune italiane del Neogene e Paris VII, 2 volumes del Quaternario - Ph.D. dissertation, Modena & Barbadillo, L.J., Garcia-Paris, M. & Sanchiz, B., 1997 - Reggio Emilia University Origenes y relaciones evolutivas de la herpetofauna Duffaud, S., 2000 - Les faunes d’amphibiens du Crétacé Iberica - in: Pleguezuelos, J.M. (ed.) - Distribucion y supérieur à l’Oligocène inférieur en Europe: paléo- biogeografia de los anfibios y reptiles en España y biodiversité, évolution, mise en place - Thesis, Portugal. Monografías de Herpetología, Volumen 3 - Muséum national d’Histoire naturelle, 2 volumes pp. 47-100, Universidad de Granada & Asociación Fortelius, M., Andrews, P., Bernor, R.L., Viranta, S. & Herpetológica Española, Granada Werdelin, L., 1996 - Preliminary analysis of taxono- Berggren, W.A. & Van Couvering, J.A., 1974 - The late mic diversity, turnover and provinciality in a sub Neogene, biostratigraphy, geochronology, and paleo sample of large land mammals from the later climatology of the last 15 million years in marine and Miocene of western Eurasia - Acta Zoologica continental sequences - Palaeogeography, Cracoviensia 39: 167-178 Palaeoclimatology, Palaeoecology 16 (1-2): 1- 216 Fortelius, M., Werdelin, L., Andrews, P., Bernor, R.L., Bernor, R.L., Fahlbusch, V., Andrews, P., de Bruijn, H., Gentry, A., Humphrey, L., Mittmann, H.-W. & Fortelius, M., Rögl, F., Steininger, F.F. & Werdelin, Viranta, S., 1996 - Provinciality, diversity, turnover L., 1996 - The evolution of western Eurasian and paleoecology in land mammal faunas of the later Neogene Mammal faunas: a chronologic, systematic, Miocene of western Eurasia - in: Bernor, R., Fahl- biogeographic and paleoenvironmental synthesis - in: busch, V. & Mittmann, W.-H. (eds.) - The Evolution Bernor, R., Fahlbusch, V., & Mittmann, W.-H. (eds.) - of Western Eurasian Neogene Mammal Faunas - pp. The Evolution of Western Eurasian Neogene 414-448, Columbia University Press, New York Mammal Faunas - pp. 449-470, Columbia University Gleed-Owen, C.P., 1999 - The palaeoclimatic and bio- Press, New York stratigraphic significance of herpetofaunal remains Bertini, A., 1995 - La végétation et le climat en Italie from the British Quaternary - in: Andrews, P. & centro-septentrinale entre 7 et 1.6 millions d’années Banham, P. (eds.) - Late Cenozoic Environments and (du Messinien au Pléistocéne inferieur) - in: La Hominid Evolution: a tribute to Bill Bishop - pp. Mediterranée: variabilités climatiques, environnement 201-215, Geological Society, London et biodiversité - pp. 91-97, Actes du Colloque Gliozzi, E., Abbazzi, L., Argenti, P., Azzaroli, A., Caloi, scientifique, 6-7 Avril 1995, Montpellier L., Capasso Barbato, L., Di Stefano, G., Esu, D., Cerling, T.E., Harris, J.M., Macfadden, B.J., Leakey, Ficcarelli, G., Girotti, O., Kotsakis, T., Masini, F., M.G., Quade, J., Elsenmann, V. & Ehleringer, J.R., Mazza, P., Mezzabotta, C., Palombo, M.R., Petronio, 1997 - Global vegetation change through the C., Rook, L., Sala, B., Sardella, R., Zanalda, E. & Miocene/Pliocene boundary - Nature 389: 153-158 Torre, D. 1997 - Biochronology of selected mammals, Cheylan, M., 1995 - Les reptiles du Paléarctique occi molluscs and ostracods from the Middle Pliocene to dental. Diversité et conservation - These Ecole the Late Pleistocene in Italy. The state of the art - Pratique des Hautes Etudes, Montpellier Rivista Italiana di Paleontologia e Stratigrafia 103 De Broin, F., 1977 - Contribution à l’étude des (3): 369-388 Chéloniens. Chéloniens continentaux du Crétacé et Holman, J.A., 1995 - Pleistocene Amphibians and du Tertiaire de France - Mémoires du Muséum Reptiles in North America - Oxford Monographs on national d'Histoire naturelle, C 38: i-ix, 1-366 Geology and Geophysics No. 32. Oxford University De Bruijn, H., Daams, R., Daxner-Hock, G., Fahlbush, Press, New York V., Ginsburg, L., Mein, P. & Morales J., 1992 - Holman, J.A., 1998 - Pleistocene Amphibians and Report of the RCMNS working group on fossil Reptiles in Britain and Europe - Oxford Monographs mammals, Reisenburg 1990 - Newsletter on on Geology and Geophysics No. 38. Oxford Univer- Stratigraphy 26 (2/3): 65-118 sity Press, New York De Lapparent de Broin, F., 2001 - The European turtle Ivanov, M., 2001 - Changes in the composition of the fauna from the to the present - Dumerilia 4 European snake fauna during the Early Miocene and (3): 155-217 the Early/Middle Miocene transition - Paläonto-

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logische Zeitschrift 74 (4): 563-573. 185-206, University Complutense, Madrid Lindsay, E., 2001 - Correlation of mammalian biochro- Sanchiz, B., 1998. Salientia - in: Wellnhofer, P. (ed.) - nology with the Geomagnetic Polarity Time Scale - Handbuch der Paläoherpetologie, part 4 - Gustav in: Rook, L. & Torre, D. (eds.) - Neogene and Fischer Verlag, Stuttgart - New York Quaternary continental stratigraphy and mammal Schmidt-Kittler, N. (ed.), 1987 - International symposium evolution. Papers in honour of Augusto Azzaroli’s on Mammalian Biostratigraphy and Paleoecology of outstanding contribution in Geology and paleontology the European Paleogene - Münchner geowissen - Bollettino della Società Geologica Italiana 40 (2): schaftliche Abhandlungen, A: 10 225-233 Sigé, B., & Legendre, S., 1997 - Un outil de la strati McKinney, M.L., 1998 - Biodiversity dynamics: niche graphie du Tertiaire continental: l'échelle de niveaux- preemption and saturation in diversity equilibria - in: repères de mammifères. Principes et spécificité; McKinney, M.L. & Drake, J.A. (eds.) - Biodiversity interêt relié des faunes karstiques - in: Aguilar, J.P., dynamics. Turnover of populations, taxa and commu Legendre S. & Michaux, J. (eds.) - Actes du Congrès nities - pp. 1-16, Columbia University Press, New BiochroM’97. Mémoires et Traveaux de l’Institut de York Montpellier de l'École Pratique des Hautes Études, Mein, P., 1975 - Report on activity RCMNS-working Institute Montpellier 21: 47-54 groups (1971-1975) - pp. 78-81, Bratislava Suc, J.-P., Bertini, A., Combourieu-Nebout, N., Diniz, F., Mein, P., 1989 - European mammal correlations - in: Leroy, S., Russo-Ermolli E., Zheng Z., Bessais E. & Lindsay, E.H., Fahlbush, V. & Mein, P. (eds.) - Ferrier J., 1995 - Structure of West Mediterranean European Neogene Mammal Chronology - pp. 73-90, vegetation and climate since 5.3 ma - Acta Zoologica Plenum Press, New York Cracoviensia 38 (1): 3-16 Rage, J.C., 1984 - La "Grande Coupure" éocène/oligocè Van Der Made, J., 1997 - Intercontinental dispersal ne et les herpétofaunes (Amphibiens et Reptiles): events, eustatic sea level and early and middle problèmes du synchronisme des événements paléo Miocene stratigraphy - in: Aguilar, J.P., Legendre S., biogéographiques - Bulletin de la Société géologique & Michaux, J. (eds.) - Actes du Congrès Biochro de France 26 (6): 1251-1257 M’97. Mémoires et Traveaux de l’Institut de Rage, J.C., 1987 - Extinctions chez les Squamates Montpellier de l'École Pratique des Hautes Études, (Reptiles) à la fin de l'Oligocène en France. Institute Montpellier 21: 75-81 Adaptations et modifications de l'environnement - Venczel, M., 1999 - Land salamanders of the family Mémoires de la Société géologique de France, N.S. Hynobiidae from the Neogene and Quaternary of 150: 145-150 Europe - Amphibia-Reptilia 20: 401-412 Rage, J.C., 1997 - Palaeobiological and palaeogeographi Vianey-Liaud, M. (ed.), 1997 - Biochronologie du cal background of the European Herpetofauna - in: Paléogène - in: Aguilar, J.P., Legendre S. & Michaux, Gasc, J.-P. et al. (eds.) - Atlas of Amphibians and J. (eds.) - Actes du Congrès BiochroM’97. Mémoires Reptiles in Europe - pp. 23-29. Societas Europaea et Traveaux de l’Institut de Montpellier de l'École Herpetologica and Muséum national d'Histoire Pratique des Hautes Études, Institut Montpellier 21: naturelle (IEGB/SPN), Paris 770-794 Rage, J.C. & Auge, M., 1993 - Squamates from the Vigne, J.-D., Bailon, S. & Cuisin J., 1997 - Biostrat- Cainozoic of the western part of Europe. A review - igraphy of amphibians, reptiles, birds and mammals Revue de Paléobiologie, Special Volume. 7: 199-216 in Corsica and the role of man in the Holocene faunal Rio, D., Sprovieri, R. & Di Stefano, E., 1994 - The turnover - Anthropozoologica 25-26: 587-604 Gelasian stage: a proposal of a new chronostratigra- Vislobokova, I.A., Erbaeva, M.A. & Sotnikova, M.V., phic unit of the Pliocene series - Rivista Italiana di 1993 - The Early Villafranchian stage in the develop Paleontologia e Stratigrafia 100 (1): 103-124 ment of the mammalian fauna of Northern Eurasia - Sanchiz, B., 1997 - Factores causales de biodiversidad: Stratigraphy and Geological Correlation 1 (5): 555- especiatión, extinctión y migración en la formación 564 de la batracofauna europea - in: Aguirre, E. et al. Werdelin, L. & Fortelius, M., 1997 - Biogeographic (eds.) - Registros fósiles e historia de la Tierra - pp. characterization of MN unit reference localities - in:

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DELFINO et al.:Tertiary herpetofauna

Aguilar, J.P., Legendre S. & Michaux, J. (eds.) - Actes du Congrès BiochroM’97. Mémoires et Williams, H.S., 1901 - Discrimination of time value in Traveaux de l’Institut de Montpellier de l'École geology - Journal of Geology, 9: 570-585 Pratique des Hautes Études, Institut Montpellier 21: 67-73 Received 19 May 2001 Accepted 04 November 2002

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