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# Introduction to: Snakes from the Cenozoic of Europe –
towards a macroevolutionary and palaeobiogeographic
synthesis

Georgios L Georgalis, Hussam Zaher, Michel Laurin

► To cite this version:

Georgios L Georgalis, Hussam Zaher, Michel Laurin. Introduction to: Snakes from the Cenozoic of Europe – towards a macroevolutionary and palaeobiogeographic synthesis. Comptes Rendus. Palevol, 2025, 24 (3), pp.45-49. 10.5852/cr-palevol2025v24a3. hal-05344948

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Distributed under a Creative Commons Attribution 4.0 International License
Introduction to: Snakes from the Cenozoic
of Europe – towards a macroevolutionary
and palaeobiogeographic synthesis

Georgios L. GEORGALIS, Hussam ZAHER & Michel LAURIN

<figure><img src="image_3.png" /></figure>

SNAKES FROM THE CENOZOIC OF EUROPE
— TOWARDS A MACROEVOLUTIONARY AND PALAEOBIOGEOGRAPHIC SYNTHESIS

Edited by Georgios L. GEORGALIS, Hussam ZAHER & Michel LAURIN
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Micropaléontologie/Micropalaeontology

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Walter Joyce (Universität Freiburg, Switzerland)

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Paléomammalogie (petits mammifères sauf Euarchontoglires)/Palaeomammalogy (small mammals except for Euarchontoglires)

Robert Asher (Cambridge University, Cambridge)

Paléomammalogie (Euarchontoglires)/Palaeomammalogy (Euarchontoglires)

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Aurélien Mounier (CNRS/Muséum national d'Histoire naturelle, Paris)

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Nicolas Teyssandier (CNRS/Université de Toulouse, Toulouse)

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# Introduction to: Snakes from the Cenozoic of Europe – towards a macroevolutionary and palaeobiogeographic synthesis

## Georgios L. GEORGALIS

Institute of Systematics and Evolution of Animals,
Polish Academy of Sciences, Sławkowska 17, 31-016 Kraków (Poland)
[georgalis@isez.pan.krakow.pl](mailto:georgalis@isez.pan.krakow.pl) (corresponding author)

## Hussam ZAHER

Museu de Zoologia, Universidade de São Paulo, Avenida Nazaré 481,
Ipiranga 04263-000, São Paulo, SP (Brazil)

## Michel LAURIN

Centre de Recherches en paléontologie – Paris (CR2P), Centre national de la Recherche scientifique (CNRS), Muséum national d'Histoire naturelle (MNHN), Sorbonne Université, Département Origines & Évolution, CP 38, 57 rue Cuvier, F-75231 Paris cedex 05 (France)

Submitted on 20 December 2024 | Accepted on 4 February 2025 | Published on 19 February 2025

urn:lsid:zoobank.org:pub:0AB7A60B-74D4-4B18-AFA0-2D68341C8960

Georgalis G. L., Zaher H. & Laurin M. 2025. — Introduction to: Snakes from the Cenozoic of Europe – towards a macroevolutionary and palaeobiogeographic synthesis, in Georgalis G. L., Zaher H. & Laurin M. (eds), Snakes from the Cenozoic of Europe – towards a macroevolutionary and palaeobiogeographic synthesis. *Comptes Rendus Palevol* 24 (3): 45-49. https://doi.org/10.5852/cr-palevol2025v24a3

Snakes have long been important components of most European ecosystems. This is not readily obvious to herpetologists given that the modest extant taxonomic richness represents simply a glimpse of their astonishing abundance in Europe during much of the Cenozoic. Indeed, the fossil record documents a high taxonomic diversity of snakes, which included a plethora of distinct shapes, sizes, feeding and reproductive adaptations, ecological strategies, and locomotion (Rage 1984; Ivanov 2022; Smith & Georgalis 2022). Interestingly (as of February 2025), as many as 162 extinct species have been established from the Cenozoic of Europe; these 162 species have been assigned to 92 different genera, among which 56 genera are extinct and established upon European fossil material, 10 genera are extinct and have originally established upon fossil material from other continents, and 26 represent extant genera (GLG, unpublished data). This high diversity included both early representatives of extant clades that now inhabit the continent, such as erycids, colubrids, natricids, and viperids (Szyndlar 1984, 1991a, b; Ivanov 2022), giant forms of extant clades of the continent (Georgalis *et al.* 2016), as

well as taxa that are now absent from Europe, such as pythons and elapids (Rage 1984; Szyndlar 1991a, b; Szyndlar & Rage 2003; Zaher & Smith 2020; Ivanov 2022), but also a parade of unique, bizarre, and now extinct clades, such as palaeophiids, russellophiids, and anomalophiids (Rage 1983, 1984; Georgalis *et al.* 2020; Smith & Georgalis 2022; Fig. 1).

Fossil snake remains from the European Cenozoic consist primarily of isolated vertebrae (Rage 1984; Szyndlar 1984, 1991a, b; Venczel 2000; Szyndlar & Rage 2003; Ivanov 2022; Smith & Georgalis 2022; Fig. 1). Articulated vertebrae and cranial remains are less common (e.g. Owen 1850; Szunyoghy 1932; Hoffstetter 1939; Szyndlar 1984; Ivanov 1999; Venczel 2000; Szyndlar & Rage 2003; Georgalis *et al.* 2021; Georgalis & Scheyer 2022; Čerňanský 2024; Fig. 1), while complete skeletons are exceptionally rare and typically limited to a few Fossillagerstätte localities (Meyer 1845; Mas salongo 1859; Janensch 1906; Kuhn 1939; Szyndlar 1992; Szyndlar & Böhme 1996; Baszio 2004; Smith & Scanferla 2016, 2021, 2022; Scanferla & Smith 2020; Zaher & Smith 2020; Georgalis *et al.* 2021; Seghetti *et al.* 2022; Palci *et al.*
<figure><img src="image_4.png" /><figcaption>FIG. 1. — Selected type specimens of European Cenozoic snakes: A, paratype skeleton (HLMD-Be 165) of *Messelopython freyi* Zaher & Smith, 2020, from the early to middle Eocene of Messel, Germany; B, holotype skeleton (SMF-ME 1828 A-B) of *Messelophis variatus* Baszio, 2004, from the early to middle Eocene of Messel, Germany; C, syntype trunk vertebra (NHMUK PV OR 33209) of *Palaeophis typhaeus* Owen, 1850, in right lateral view, from the early Eocene of Bracklesham, England; D, holotype trunk vertebra (ISEZ FR/RKI-10000) of *Natrix longivertebrata* Szyndlar, 1984, in right lateral view, from the</figcaption></figure>
2024; Fig. 1). Remarkably, even mummified portions of snake
bodies have been recovered in exceptional cases (Filhol 1877;
Rochebrune 1884; Georgalis *et al.* 2021; Fig. 1)! Notably, the
beautifully preserved skeletons from the Fossillagerstätte locali-
ties of Messel and Geiseltal in Germany have provided and
continue to provide critical insight into the cranial anatomy
and intracolumnar variation of multiple extinct snake taxa
(Kuhn 1939; Baszio 2004; Scanferla & Smith 2020; Zaher &
Smith 2020; Smith & Scanferla 2021, 2022; Georgalis *et al.*
2021; Palci *et al.* 2024; Fig. 1). Additionally, these fossils offer
valuable information on palaeoecology and trophic interac-
tions (e.g. Smith & Scanferla 2016).

The European Cenozoic fossil record of snakes dates back to
the Paleocene and encompasses a wide geographic distribution.
As such, it provides valuable insights into major climatic events,
dispersals, faunal turnovers, and extinctions that shaped the
European Cenozoic faunas, ultimately leading to the emergence
of extant taxa. Such prominent events that affected European
snake faunas include the high temperatures observed during the
Paleocene-Eocene Thermal Maximum (PETM) and the Early
Eocene Climatic Optimum (EECO), the cooling that triggered
the "Grande Coupure" at the Eocene-Oligocene boundary, or
the distinct palaeogeographic settings that arose during the
formation of the "Gomphotherium Landbridge" during the
Early Miocene, and the Messinian Salinity Crisis during the
Miocene-Pliocene boundary (Szyndlar & Rage 2003; Rage
2013; Zaher et al. 2019; Georgalis et al. 2021; Ivanov 2022;
Georgalis & Szyndlar 2022; Smith & Georgalis 2022). How-
ever, significant gaps remain from both a stratigraphic and a
geographic point of view. Indeed, stratigraphically speaking,
some geological epochs, such as the Paleocene and the Oli-
gocene, are underrepresented, compared to other epochs, in
the European snake fossil record (Smith & Georgalis 2022).
Most notably from a geographic point of view, the Eastern
portions of the continent remain notably less sampled for
fossil snakes than Western and Central Europe (e.g. Ivanov
2022; Smith & Georgalis 2022).

European snake fossils have been known for over two centu-
ries, with the earliest discoveries dating back to the early 19th
century (Karg 1805). Throughout that century, notable fossil
snake discoveries continued to capture the interest of European
palaeontologists, including remarkable or enigmatic forms,
such as the giant aquatic genus *Palaeophis* Owen, 1841, from
England, the giant viper *Laophis* Owen, 1857, and *Python*
*euboicus* Römer, 1870 – representing the first record of the

extant genus *Python* Daudin, 1803, in Europe – both from
Greece, as well as the constrictor *Palaeopython* Rochebrune,
1880, from France (Owen 1841, 1850, 1857; Meyer 1845;
Römer 1870; Filhol 1877; Rochebrune 1880, 1884). The 20th
century witnessed continued advancements in fossil snake
discoveries in Europe, driven particularly by the comprehen-
sive and meticulous work of Robert Hoffstetter, Jean-Claude
Rage, and Zbigniew Szyndlar, who significantly enhanced our
understanding of taxonomic diversity, cranial and vertebral
diagnostic features, and intracolumnar variation in snakes
(e.g. Hoffstetter 1939; Hoffstetter & Rage 1972; Rage 1974,
1983, 1984; Szyndlar 1984, 1991a, 1991b). In recent decades,
our understanding of the taxonomic diversity and anatomy of
Cenozoic fossil snakes from Europe has accelerated significantly,
aided by the use of recent technologies such as micro-com-
puted tomography (µCT) scanning (Bell et al. 2021) and by
an improved knowledge of the skeletal anatomy of extant taxa
(Zaher et al. 2019; Szyndlar & Georgalis 2023). Despite these
advances, significant gaps remain, as many taxa have yet to
be adequately described and the phylogenetic relationships
of several groups are still poorly understood, hindering a full
understanding of snake evolution in the European Cenozoic.

With this new volume in *Comptes Rendus Palevol*, we aim
to address significant gaps in our knowledge of the snake
fossil record during the European Cenozoic. The forthcoming
contributions in this volume will encompass a range of
topics, including:

- descriptions of new taxa and revisions of poorly documented taxa;

- identifications of novel anatomical features in the skull and vertebrae of fossil snakes;

- studies of articulated skeletons from Fossillagerstätte localities, enhanced by the application of micro-computed tomography (µCT) scanning;

- documentation of new fossil snake occurrences from
  underexplored regions of Europe;

- revised taxonomic frameworks for enigmatic and previously ambiguous extinct snake groups;

- studies of palaeobiogeography, dispersal events, extinction and survivorship dynamics of snakes across the Cenozoic of Europe, using sophisticated analytical methods.

We anticipate that this volume will provide fresh perspec-
tives on the evolution of Cenozoic snakes in Europe, while
also serving as a trigger for future research and new discoveries
about these fascinating and charismatic reptiles!

Late Pliocene (MN 16), of Rěbielice Królewskie IA, Poland; **E**, holotype trunk vertebra (MNHN.F.VCO29) of *Python europaeus* Szyndlar & Rage, 2003, in dorsal view, from the Early-Middle Miocene (MN 4/5) of Vieux-Collonges, France; **F**, syntype trunk vertebra (MNHN.F.LGA53) of "Typhlops" *grivensis* Hoffstetter, 1946, in dorsal view, from the Middle Miocene (MN 7/8) of La Grive, France; **G**, holotype trunk vertebra (MNHN.F.CB1603) of *Russellophis tenuis* Rage, 1975, in ventral view, from the early Eocene (MP 8/9) of Condé-en-Brie, France; **H**, right maxilla (GMH Ce I-5826-1926) part of the lectotype of *Eoconstrictor spinifer* (Barnes, 1927), in labial view, from the late early to early middle Eocene of Geiseltal, Germany; **I**, holotype trunk vertebra (PIMUZ A/III 634) of *Palaeopython helveticus* Georgalis & Scheyer, 2019, in anterior view, from the late middle to late Eocene of Dielsdorf, Switzerland; **J**, paralectotype mummified trunk portion (MNHN.F.QU16329) of *Rageophis lafonti* (Filhol, 1877) from the Eocene of the Phosphorites du Quercy, France (this specimen serves also as a paralectotype of *Tachyophis nitidus* Rochebrune, 1884); **K**, holotype posterior caudal vertebra (MNHN.F.COD1) of *Bransateryx vireti* Hoffstetter & Rage, 1972, in anterior view, from the late Oligocene (MP 30) of Coderet, France; **L**, paralectotype left dentary (MNHN.F.QU16336) of *Palaelaphis antiquus* Rochebrune, 1884, in medial view, from the Eocene or Oligocene of the Phosphorites du Quercy, France. Collections: A, Hessisches Landesmuseum Darmstadt; B, Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, C, Natural History Museum, London, D, Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow; E-G, J-L, Muséum national d'Histoire naturelle, Paris; H, Geiseltalmuseum of Martin-Luther Universität Halle-Wittenberg, Halle; I, Palaeontological Institute of the University of Zurich. Scale bars: A, 50 mm; B, 40 mm; C, J, 10 mm; D, F, 1 mm; E, 3 mm; G, K, 2 mm; H, I, L, 5 mm. Photo credits: A, HZ; B, Anika Vogel; C-L, GLG.
## Acknowledgements

For access to specimens under their care, we thank Nour-Eddine Jalil (MNHN), Marc Jones and Mike Day (NHMUK), Krister Smith (SMF-ME), Oliver Wings and Roberto Rozzi (GMH), Christian Klug (PIMUZ), and Torsten Wappler (HLMD). GLG acknowledges funding from the research project no. 2023/49/B/ST10/02631 financed by the National Science Center of Poland (Narodowe Centrum Nauki), as well as travel support from SYNTHESYS FR-TAF_Call4_035 (MNHN) and the 2023 Richard Owen Fund of the Palaeontographical Society.

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Submitted on 20 December 2024;
accepted on 4 February 2025;
published on 19 February 2025.