89e8144ea1d903ef4e294d947c426fc8 comptes-rendus-palevol2026v25a7.pdf 770a8813af51870b4ee0c67ae3d50208941739d0 comptes-rendus-palevol2026v25a7.pdf 7e69361dc2e23a823a18a3e331da74e63165918f1fa25e49f5acc9bf1b0e9dcb comptes-rendus-palevol2026v25a7.pdf Title: First fossil remains of the extant colubrid Zamenis situla (Linnaeus, 1758) (Serpentes) from the island of Crete, Greece Subject: English abstract (résumé anglais) We here provide the first description of fossil remains of the colubrid snake cf. Zamenis situla from Crete, Greece, originating from the Late Pleistocene of the Rethymnon fissure. Although Zamenis situla (Linnaeus, 1758) is a widespread snake species in Greece, including Crete, its fossil record in the area has thus far been restricted to Middle Pleistocene material from Tourkobounia 2, near Athens, which also represents the sole confirmed fossil occurrence of the species. We provide a comparison of the fossil specimens to other extant Cretan snakes, and highlight key morphological features that distinguish it from other colubrids. This fossil material provides further support to recent molecular studies suggesting long-term presence of Z. situla on the island. However, due to the scarcity of the fossil record, it remains unclear whether Z. situla inhabited Crete continuously since the Late Pleistocene or was instead reintroduced to it later on. Finally, we investigate skeletal material of Z. situla, providing anatomical descriptions of key cranial and vertebral structures, which could facilitate in future fossil identifications and comparisons among colubrid snakes. French abstract (résumé français) Nous fournissons ici la première description de restes fossiles du serpent colubridé cf. Zamenis situla en provenance de Crète, en Grèce, datant du Pléistocène tardif et issus de la fissure de Réthymnon. Bien que Zamenis situla (Linnaeus, 1758) soit une espèce de serpent largement répandue en Grèce, y compris en Crète, son registre fossile dans la région était jusqu’à présent limité au Pléistocène moyen de Tourkobounia 2, près d’Athènes, qui constitue également la seule occurrence fossile confirmée de l’espèce. Nous comparons les spécimens fossiles à d’autres serpents crétois actuels et mettons en évidence les caractéristiques morphologiques clés qui le distinguent des autres colubridés. Ce matériel fossile apporte un soutien supplémentaire aux études moléculaires récentes suggérant une présence à long terme de Z. situla sur l’île. Cependant, en raison de la rareté du registre fossile, il reste incertain si Z. situla a habité la Crète de manière continue depuis le Pléistocène tardif ou s’il y a été réintroduit ultérieurement. Enfin, nous examinons le matériel squelettique de Z. situla, en fournissant des ­descriptions anatomiques des structures clés du crâne et des vertèbres, ce qui pourrait faciliter les futures identifications fossiles et les comparaisons entre les colubridés. Author: Olaf LIZAK Creator: Adobe InDesign 20.5 (Windows) Producer: Adobe PDF Library 17.0 CreationDate: Wed Apr 22 14:55:07 2026 CEST ModDate: Wed Apr 22 14:59:34 2026 CEST Custom Metadata: no Metadata Stream: yes Tagged: yes UserProperties: no Suspects: no Form: none JavaScript: no Pages: 14 Encrypted: no Page size: 595.276 x 841.89 pts (A4) Page rot: 0 File size: 7814191 bytes Optimized: no PDF version: 1.5 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- AULWKS+Wingdings-Regular CID TrueType Identity-H yes yes yes 1485 0 NQUWKS+ITCStoneSans-Semibold Type 1C Custom yes yes yes 1488 0 VLFTDX+HelveticaNeue CID TrueType Identity-H yes yes yes 106 0 ANUCVN+ITCStoneSans-SemiboldItalic Type 1C WinAnsi yes yes yes 105 0 NQUWKS+ITCStoneSans-Semibold Type 1C WinAnsi yes yes yes 104 0 FNDBND+HelveticaNeue TrueType WinAnsi yes yes yes 100 0 FNDBND+HelveticaNeue-Italic TrueType WinAnsi yes yes yes 96 0 PKGJXJ+HelveticaNeue-SC750 TrueType WinAnsi yes yes yes 92 0 PKGJXJ+HelveticaNeue-Italic-SC750 TrueType WinAnsi yes yes yes 91 0 ZMERHP+HelveticaNeue-BoldItalic TrueType WinAnsi yes yes yes 87 0 ZMERHP+HelveticaNeue-Bold TrueType WinAnsi yes yes yes 83 0 ZMERHP+AGaramondPro-Regular Type 1C Custom yes yes yes 125 0 JMBPRV+AGaramondPro-Semibold Type 1C WinAnsi yes yes yes 121 0 NIVIVN+AGaramondPro-Italic Type 1C Custom yes yes yes 116 0 MHFSHP+MyriadPro-Regular Type 1C Custom yes yes yes 115 0 ZMERHP+AGaramondPro-Regular Type 1C WinAnsi yes yes yes 135 0 NIVIVN+AGaramondPro-Italic Type 1C WinAnsi yes yes yes 133 0 FNDBND+HelveticaNeue TrueType WinAnsi yes yes yes 131 0 MHFSHP+AGaramond-Regular-SC750 Type 1C WinAnsi yes yes yes 143 0 QIELLH+AGaramondPro-SemiboldItalic Type 1C WinAnsi yes yes yes 142 0 ZMERHP+AGaramondPro-Regular Type 1C WinAnsi yes yes yes 140 0 NIVIVN+AGaramondPro-Italic Type 1C WinAnsi yes yes yes 138 0 FNDBND+HelveticaNeue TrueType WinAnsi yes yes yes 148 0 QIELLH+HelveticaNeue Type 1C WinAnsi yes yes no 149 0 QIELLH+HelveticaNeue Type 1C WinAnsi yes yes no 152 0 QIELLH+HelveticaNeue Type 1C WinAnsi yes yes no 156 0 QIELLH+HelveticaNeue Type 1C WinAnsi yes yes no 157 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2026-06-04 03:57:42 CEST RepresentationInformation: comptes-rendus-palevol2026v25a7.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-06-03 08:35:23 CEST Size: 7814191 Format: PDF Version: 1.5 Status: Well-Formed, but not valid SignatureMatches: PDF-hul ErrorMessage: Annotation object is not a dictionary ID: PDF-HUL-114 Offset: 5089387 MIMEtype: application/pdf PDFMetadata: Objects: 1494 FreeObjects: 1241 IncrementalUpdates: 2 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: false NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseOutlines Language: fr-FR Outlines: Item: Title: Cover Item: Title: Title Item: Title: ABSTRACT Item: Title: RÉSUMÉ Item: Title: INTRODUCTION Children: Item: Title: Institutional abbreviations Item: Title: SYSTEMATIC PALAEONTOLOGY Children: Item: Title: SERPENTES Linnaeus, 1758 Children: Item: Title: ALETHINOPHIDIA Nopcsa, 1923 Children: Item: Title: CAENOPHIDIA Hoffstetter, 1939 Children: Item: Title: COLUBROIDES Zaher, Grazziotin, Cadle, Murphy, Moura-Leite & Bonatto, 2009 Children: Item: Title: COLUBRIFORMES Schlegel, 1837 (sensu Zaher et al., 2009) Children: Item: Title: COLUBROIDEA Oppel, 1811 (sensu Zaher et al., 2009) Children: Item: Title: COLUBRIDAE Oppel, 1811 (sensu Zaher et al., 2009) Children: Item: Title: Zamenis Wagler, 1830 Children: Item: Title: Zamenis situla (Linnaeus, 1758) Children: Item: Title: cf. Zamenis situla (Figs 1; 2) Children: Item: Title: Material examined Item: Title: Description Item: Title: SKELETAL ANATOMY OF ZAMENIS SITULA Item: Title: DISCUSSION Item: Title: Acknowledgements Item: Title: REFERENCES Info: Author: Olaf LIZAK Subject: English abstract (résumé anglais) We here provide the first description of fossil remains of the colubrid snake cf. Zamenis s椀琀甀氀愀 昀爀漀洀 䌀爀攀琀攀Ⰰ 䜀爀攀攀挀攀Ⰰ 漀爀椀最椀渀愀琀椀渀最 昀爀漀洀 琀栀攀 䰀愀琀攀 倀氀攀椀猀琀漀挀攀渀攀 漀昀 琀栀攀 刀攀琀栀礀洀渀漀渀 昀椀猀猀甀爀攀⸀ 䄀氀琀栀漀甀最栀 娀愀洀攀渀椀猀 猀椀琀甀氀愀 䰀椀渀渀愀攀甀猀Ⰰ ㄀㜀㔀㠀 椀猀 愀 眀椀搀攀猀瀀爀攀愀搀 猀渀愀欀攀 猀瀀攀挀椀攀猀 椀渀 䜀爀攀攀挀攀Ⰰ 椀渀挀氀甀搀椀渀最 䌀爀攀琀攀Ⰰ 椀琀猀 昀漀猀猀椀氀 爀攀挀漀爀搀 椀渀 琀栀攀 愀爀攀愀 栀愀猀 琀栀甀猀 昀愀爀 戀攀攀渀 爀攀猀琀爀椀挀琀攀搀 琀漀 䴀椀搀搀氀攀 倀氀攀椀猀琀漀挀攀渀攀 洀愀琀攀爀椀愀氀 昀爀漀洀 吀漀甀爀欀漀戀漀甀渀椀愀 ㈀Ⰰ 渀攀愀爀 䄀琀栀攀渀猀Ⰰ 眀栀椀挀栀 愀氀猀漀 爀攀瀀爀攀猀攀渀琀猀 琀栀攀 猀漀氀攀 挀漀渀昀椀爀洀攀搀 昀漀猀猀椀氀 漀挀挀甀爀爀攀渀挀攀 漀昀 琀栀攀 猀瀀cies. We provide a comparison of the fossil specimens to other extant Cretan snakes, and highlight key morphological features t栀愀琀 搀椀猀琀椀渀最甀椀猀栀 椀琀 昀爀漀洀 漀琀栀攀爀 挀漀氀甀戀爀椀搀猀⸀ 吀栀椀猀 昀漀猀猀椀氀 洀愀琀攀爀椀愀氀 瀀爀漀瘀椀搀攀猀 昀甀爀琀栀攀爀 猀甀瀀瀀漀爀琀 琀漀 爀攀挀攀渀琀 洀漀氀攀挀甀氀愀爀 猀琀甀搀椀攀猀 猀甀最最攀猀琀椀渀最 氀ng-term presence of Z. situla on the island. However, due to the scarcity of the fossil record, it remains unclear whether Z. s椀琀甀氀愀 椀渀栀愀戀椀琀攀搀 䌀爀攀琀攀 挀漀渀琀椀渀甀漀甀猀氀礀 猀椀渀挀攀 琀栀攀 䰀愀琀攀 倀氀攀椀猀琀漀挀攀渀攀 漀爀 眀愀猀 椀渀猀琀攀愀搀 爀攀椀渀琀爀漀搀甀挀攀搀 琀漀 椀琀 氀愀琀攀爀 漀渀⸀ 䘀椀渀愀氀氀礀Ⰰ 眀攀 椀渀瘀攀猀琀椀最愀e skeletal material of Z. situla, providing anatomical descriptions of key cranial and vertebral structures, which could facili琀愀琀攀 椀渀 昀甀琀甀爀攀 昀漀猀猀椀氀 椀搀攀渀琀椀昀椀挀愀琀椀漀渀猀 愀渀搀 挀漀洀瀀愀爀椀猀漀渀猀 愀洀漀渀最 挀漀氀甀戀爀椀搀 猀渀愀欀攀猀⸀䘀爀攀渀挀栀 愀戀猀琀爀愀挀琀 爀猀甀洀 昀爀愀渀愀椀猀一漀甀猀 昀漀甀爀渀椀猀猀漀渀猀 椀挀椀 氀愀 瀀爀攀洀椀爀攀 搀攀猀挀爀椀瀀琀椀漀渀 搀攀 爀攀猀琀攀猀 昀漀猀猀椀氀攀猀 搀甀 猀攀爀瀀攀渀琀 挀漀氀甀戀爀椀搀 挀昀⸀ 娀愀洀攀渀椀猀 猀椀琀甀氀愀 攀渀 瀀爀漀瘀攀渀愀渀挀攀 搀攀 䌀爀琀攀Ⰰ 攀渀 䜀爀挀攀Ⰰ 搀愀ant du Pléistocène tardif et issus de la fissure de Réthymnon. Bien que Zamenis situla (Linnaeus, 1758) soit une espèce de ser瀀攀渀琀 氀愀爀最攀洀攀渀琀 爀瀀愀渀搀甀攀 攀渀 䜀爀挀攀Ⰰ 礀 挀漀洀瀀爀椀猀 攀渀 䌀爀琀攀Ⰰ 猀漀渀 爀攀最椀猀琀爀攀 昀漀猀猀椀氀攀 搀愀渀猀 氀愀 爀最椀漀渀 琀愀椀琀 樀甀猀焀甠ᤀ 瀀爀猀攀渀琀 氀椀洀椀琀 愀甀 倀氀椀猀ocène moyen de Tourkobounia 2, près d’Athènes, qui constitue également la seule occurrence fossile confirmée de l’espèce. Nous 挀漀洀瀀愀爀漀渀猀 氀攀猀 猀瀀挀椀洀攀渀猀 昀漀猀猀椀氀攀猀  搠ᤀ愀甀琀爀攀猀 猀攀爀瀀攀渀琀猀 挀爀琀漀椀猀 愀挀琀甀攀氀猀 攀琀 洀攀琀琀漀渀猀 攀渀 瘀椀搀攀渀挀攀 氀攀猀 挀愀爀愀挀琀爀椀猀琀椀焀甀攀猀 洀漀爀瀀栀漀氀漀最椀焀甀攀猀clés qui le distinguent des autres colubridés. Ce matériel fossile apporte un soutien supplémentaire aux études moléculaires ré挀攀渀琀攀猀 猀甀最最爀愀渀琀 甀渀攀 瀀爀猀攀渀挀攀  氀漀渀最 琀攀爀洀攀 搀攀 娀⸀ 猀椀琀甀氀愀 猀甀爀 氠ᤀ氀攀⸀ 䌀攀瀀攀渀搀愀渀琀Ⰰ 攀渀 爀愀椀猀漀渀 搀攀 氀愀 爀愀爀攀琀 搀甀 爀攀最椀猀琀爀攀 昀漀猀猀椀氀攀Ⰰ 椀氀 爀攀te incertain si Z. situla a habité la Crète de manière continue depuis le Pléistocène tardif ou s’il y a été réintroduit ultéri攀甀爀攀洀攀渀琀⸀ 䔀渀昀椀渀Ⰰ 渀漀甀猀 攀砀愀洀椀渀漀渀猀 氀攀 洀愀琀爀椀攀氀 猀焀甀攀氀攀琀琀椀焀甀攀 搀攀 娀⸀ 猀椀琀甀氀愀Ⰰ 攀渀 昀漀甀爀渀椀猀猀愀渀琀 搀攀猀 관搀攀猀挀爀椀瀀琀椀漀渀猀 愀渀愀琀漀洀椀焀甀攀猀 搀攀猀 猀琀爀甀挀琀甀es clés du crâne et des vertèbres, ce qui pourrait faciliter les futures identifications fossiles et les comparaisons entre les 挀漀氀甀戀爀椀搀猀⸩⽔楴汥⡆楲獴⁦潳獩氠牥浡楮猠潦⁴桥⁥硴慮琠捯汵扲楤⁚慭敮楳⁳楴畬愠楮湡敵猬‱㜵㠩
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application/x-indesign to application/pdf Adobe InDesign 20.5 (Windows) / 2026-04-22T14:55:07+02:00 application/pdf Olaf LIZAK Zbigniew SZYNDLAR† Massimo DELFINO Georgios L. GEORGALIS Published under CC-BY 4.0 Licence / Publications scientifiques du Muséum national d’Histoire naturelle, Académie des sciences, Paris, 2026 — Comptes Rendus Palevol — http://sciencepress.mnhn.fr English abstract (résumé anglais) We here provide the first description of fossil remains of the colubrid snake cf. Zamenis situla from Crete, Greece, originating from the Late Pleistocene of the Rethymnon fissure. Although Zamenis situla (Linnaeus, 1758) is a widespread snake species in Greece, including Crete, its fossil record in the area has thus far been restricted to Middle Pleistocene material from Tourkobounia 2, near Athens, which also represents the sole confirmed fossil occurrence of the species. We provide a comparison of the fossil specimens to other extant Cretan snakes, and highlight key morphological features that distinguish it from other colubrids. This fossil material provides further support to recent molecular studies suggesting long-term presence of Z. situla on the island. However, due to the scarcity of the fossil record, it remains unclear whether Z. situla inhabited Crete continuously since the Late Pleistocene or was instead reintroduced to it later on. Finally, we investigate skeletal material of Z. situla, providing anatomical descriptions of key cranial and vertebral structures, which could facilitate in future fossil identifications and comparisons among colubrid snakes. French abstract (résumé français) Nous fournissons ici la première description de restes fossiles du serpent colubridé cf. Zamenis situla en provenance de Crète, en Grèce, datant du Pléistocène tardif et issus de la fissure de Réthymnon. Bien que Zamenis situla (Linnaeus, 1758) soit une espèce de serpent largement répandue en Grèce, y compris en Crète, son registre fossile dans la région était jusqu’à présent limité au Pléistocène moyen de Tourkobounia 2, près d’Athènes, qui constitue également la seule occurrence fossile confirmée de l’espèce. Nous comparons les spécimens fossiles à d’autres serpents crétois actuels et mettons en évidence les caractéristiques morphologiques clés qui le distinguent des autres colubridés. Ce matériel fossile apporte un soutien supplémentaire aux études moléculaires récentes suggérant une présence à long terme de Z. situla sur l’île. Cependant, en raison de la rareté du registre fossile, il reste incertain si Z. situla a habité la Crète de manière continue depuis le Pléistocène tardif ou s’il y a été réintroduit ultérieurement. Enfin, nous examinons le matériel squelettique de Z. situla, en fournissant des ­descriptions anatomiques des structures clés du crâne et des vertèbres, ce qui pourrait faciliter les futures identifications fossiles et les comparaisons entre les colubridés. First fossil remains of the extant colubrid Zamenis situla (Linnaeus, 1758) (Serpentes) from the island of Crete, Greece Serpentes Colubridae Crete Pleistocene skeletal anatomy Serpentes Colubridae Crète Pléistocène anatomie du squelette http://cr-palevol.fr/25/7 False Adobe PDF Library 17.0 False Muséum national d'Histoire naturelle, Publications scientifiques, Palevol Checksum: cb1e92b5 Type: CRC32 Checksum: 89e8144ea1d903ef4e294d947c426fc8 Type: MD5 Checksum: 770a8813af51870b4ee0c67ae3d50208941739d0 Type: SHA-1 Checksum: 7e69361dc2e23a823a18a3e331da74e63165918f1fa25e49f5acc9bf1b0e9dcb Type: SHA-256