identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03C687BCFF80FFE07E4C4727366BB143.text	03C687BCFF80FFE07E4C4727366BB143.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cheilophis Gilmore 1938	<div><p>Genus Cheilophis Gilmore, 1938</p><p>TYPE SPECIES. — Cheilophis huerfanoensis Gilmore, 1938 .</p><p>EMENDED DIAGNOSIS. — Cheilophis can be differentiated from all other snakes by its unique combination of features: wide haemal keel (on the posterior mid- and posterior trunk vertebrae) that bears a constriction at around its mid-length and becomes wider and blunt wedged-shaped at its posterior termination; distinct “notches” of the haemal keel in ventral view; ventral expansion of the haemal keel in lateral view; thick neural spine that is mostly confined at the posterior portion of the neural arch; relatively thin zygosphene with a distinct median lamina (“lip”) and prominent lateral lobes; moderately vaulted neural arch; deep posterior median notch of the neural arch; deep interzygapophyseal constriction; almost circular cotyle and condyle; widely extended parapophyseal process of the paradiapophyses; almost horizontal to only slightly dorsally inclined prezygapophyses; and large prezygapophyseal and postzygapophyseal articular facets. In addition to these vertebral characters, Rage (1984b) also indicated the following cranial features as diagnostic for the genus: palatal process of the maxilla directed backwards; surangular lamina of the compound bone higher than the prearticular; coronoid present.</p></div>	https://treatment.plazi.org/id/03C687BCFF80FFE07E4C4727366BB143	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Mennecart, Bastien	Georgalis, Georgios L., Mennecart, Bastien (2025): Another wanderer from the Paleocene-Eocene Thermal Maximum? A new species of the North American snake genus Cheilophis Gilmore, 1938 from the early Eocene of France. Comptes Rendus Palevol 24 (16): 317-331, DOI: 10.5852/cr-palevol2025v24a16, URL: https://doi.org/10.5852/cr-palevol2025v24a16
03C687BCFF80FFEE7C88454637BCB354.text	03C687BCFF80FFEE7C88454637BCB354.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cheilophis periplanetes Georgalis & Mennecart 2025	<div><p>Cheilophis periplanetes n. sp.</p><p>(Figs 2-7)</p><p>urn:lsid:zoobank.org:act: 97488C93-5A21-4B5F-8DB2-52964DD335F5</p><p>TYPE MATERIAL. — Holotype. France • 1 specimen (a trunk vertebra); Cuis; Formation of Unios and Teredinids Sands; 49.86 ± 0.455 Ma; MP 10, late Ypresian (early Eocene); NMB T.S.1100 (Figs 2; 3). Paratyp e. France • 1 specimen (a trunk vertebra); Cuis; Formation of Unios and Teredinids Sands; 49.86 ± 0.455 Ma; MP 10, late Ypresian (early Eocene); NMB T.S.1101 (Figs 4; 5).</p><p>REFERRED SPECIMENS. — Five trunk vertebrae (NMBT.S.1063, NMB T.S.1067, NMB T.S.1071, NMB T.S.1084, and NMB T.S.1092). Tentatively also, an anterior trunk vertebra (NMB T.S.1065) and two caudal vertebrae (NMB T.S.1048 and NMB T.S.1066).</p><p>TYPE LOCALITY AND AGE. — All specimens originate from the locality of Cuis; MP 10 (around 49.86 Ma ± 0.455), early Eocene.</p><p>ETYMOLOGY. — The new species epithet is from the Greek word “periplanetes ” (“πΕΡΙπλανητής”), which means “wanderer”, reflecting the potential wandering that occurred during the dispersals among North America and Europe.</p><p>D</p><p>DIAGNOSIS. — Cheilophis periplanetes n. sp. can be referred to the genus Cheilophis on the basis of the full list of diagnostic vertebral features of the genus (see above). Cheilophis periplanetes n. sp. can be differentiated from the type and only other known species of Cheilophis, i.e., Cheilophis huerfanoensis, by its much smaller size (centrum length [CL] almost half in size), shorter centrum, the absence of prezygapophyseal accessory processes, the more massive prezygapophyseal and postzygapophyseal articular facets, the shorter neural spine, the rare presence of paracotylar foramina, and the more dorsally expanded lateral lobes of the zygosphene.</p><p>DESCRIPTION</p><p>Holotype</p><p>The holotype vertebra (NMB T.S.1100) is incomplete, missing most of the left postzygapophysis and part of the right prezygapophysis, while the paradiapophyses are eroded (Figs 2; 3). The vertebra has a centrum length (CL) of 3 mm and a neural arch width (NAW) of 3.2 mm. In anterior view (Figs 2A; 3A), the zygosphene is moderately thin and bears a distinct median lamina, resembling a lip. There are two distinct lateral lobes on the zygosphene that protrude much dorsally, like horns. The neural canal is semicircular (i.e., like an arch, rounded above and flat below). The prezygapophyses are almost horizontal, being only very slightly dorsally extended. The cotyle is large and almost circular. There are no paracotylar foramina. In posterior view (Figs 2B; 3E), the neural arch is moderately vaulted, with a vaulting ratio (sensu Georgalis et al. 2021b) equal to 0.47. The zygantral roof is relatively thick. The condyle is large and circular. In dorsal view (Figs 2C; 3D), the neural spine is very thick and is mostly confined to the posterior half of the neural arch. The zygosphene is trilobed, with the two lateral lobes being more prominent than the central one. The prezygapophyses project anterolaterally. The prezygapophyseal articular facets are large and oval. The interzygapophyseal constriction is deep (unlike other Eocene European constrictors, such as Eoconstrictor Scanferla &amp; Smith, 2020, and Palaeopython Rochebrune, 1880). The posterior median notch of the neural arch is deep. In ventral view (Figs 2D; 3I), the centrum is wider than long (ratio CL/ NAW equal to 0.94). The haemal keel runs along most of the midline of the centrum; it commences anteriorly almost at the ventral level of the cotyle and terminates posteriorly at the level of the condyle. The width of the haemal keel is not consistent throughout its length; instead, there is a prominent constriction at the anterior half of the centrum and it becomes gradually wider and more prominent towards its posterior termination. Notably, there are distinct “notches” around its posterior portion. The subcentral grooves are deep. Subcentral foramina are present. The paradiapophyses are much eroded and their exact shape and size cannot be precisely deciphered, but it seems that they were not divided (or were only slightly divided) into distinct diapophyses and parapophyses. There are no prezygapophyseal accessory processes. The postzygapophyseal articular facets are massive and broad. In lateral view (Figs 2E, F;3G), the neural spine is very short; it commences far behind the level of the zygosphene and gradually increases in height, being developed mostly at the posterior half of the neural arch. The zygosphenal facets are broad. The subcentral ridges are almost straight. The haemal keel protrudes ventrally at the level of the posterior half of the centrum, with its height gradually developed and culminating around its posteriormost edge. Judging from the width of the haemal keel, the vaulting of the neural arch, and the height of the neural spine, we assume that the vertebra originates somewhere from the mid- or posterior mid- trunk portion of the vertebral column.</p><p>Paratype</p><p>The paratype vertebra (NMB T.S.1101) is incomplete, missing both prezygapophyses and the right postzygapophysis (Figs 4; 5). The vertebra is small, with a centrum length (CL) equal to 2.8 mm and a neural arch width (NAW) equal to 3 mm. The paratype vertebra NMB T.S.1101 is bears an overall resemblance to the holotype, exhibiting typical features of the species, such as the thin zygosphene with the distinct “lip”, the prominent lateral lobes of the zygosphene, the dorsoventrally short neural spine that is mostly confined to the</p><p>E posterior portion of the neural arch, and the wide haemal keel with distinct “notches”. Nevertheless, there are some striking features observed in the paratype. Most prominently, the paratype vertebra exhibits the thickest neural spine known for the species (Figs 4B; 5E). Interestingly, the paratype vertebra NMB T.S.1101 demonstrates the widely extended parapophyseal process of the paradiapophyses (Fig. 5A, B), a feature that was highlighted as diagnostic for Cheilophis by Rage (1984a). This structure is otherwise not preserved in any other specimen of the new taxon (including the holotype [though a hint about such extended parapophyseal process might be observed on the right side of this specimen]) and thus, its observation in NMB T.S.1101 is of particular importance, as it further reinforces a referal of the new taxon from Cuis to the North American Cheilophis . The neural arch of NMB T.S.1101 is exceedingly depressed (Figs 4F; 5J), with a vaulting ratio (sensu Georgalis et al. 2021b) reaching only 0.16. The haemal keel is wide, and, similarly to the holotype, there are distinct “notches” around its posterior portion (Figs 4E; 5G, H). What is further notable for NMB T.S.1101 is the presence of a small paracotylar foramen on the right of the cotyle (Figs 4A; 5A, B, D) – our observation of the μ CT files of this specimen confirms that this foramen goes relatively deeply into the vertebra. We attribute all these differences to intracolumnar variation. Judging from these features, we consider that the paratype vertebra NMB T.S.1101 originates from the posterior mid-trunk portion of the column (more posterior than the holotype).</p><p>Referred specimens – intracolumnar variation</p><p>All referred specimens are much incomplete but they nevertheless preserve certain anatomical structures that allow a taxonomic attribution to Cheilophis periplanetes n. sp. Especially NMB T.S.1071, NMB T.S.1084, NMB NMB T.S.1092, and NMB T.S.1063 are all rather fragmentary and/or eroded. Most available trunk vertebrae share with the</p><p>holotype an overall similar morphology, including a wider than long centrum, a thin zygosphene with a median lamina (“lip”) and prominent lateral lobes that protrude dorsally, a relatively short neural spine that is mostly confined at the posterior portion of the neural arch, a moderately vaulted neural arch, absence of prezygapophyseal accessory processes, and large prezygapophyseal articular facets (Fig. 6). Nevertheless, some differences are apparent among the few available specimens, dealing with the shape and width of the haemal keel, the size of the prezygapophyseal and postzygapophyseal articular facets, and the height, anteroposterior length, and thickness of the neural spine. All these differences can most probably be attributed to intracolumnar variation (see e.g. Smith 2013; Szyndlar &amp; Georgalis 2023) though some other kind of variation, such as ontogenetic (e.g. Georgalis &amp; Scheyer 2019) cannot be excluded.</p><p>As is common in snakes, anterior trunk vertebrae are characterized by the presence of a hypapophysis instead of haemal keel, a more vaulted neural arch, and a higher and posteriorly inclined neural spine. The only anterior trunk vertebra from our sample is NMB T.S.1065 (Fig. 6 A-F), though still a referral of this specimen to Cheilophis periplanetes n. sp. should be considered as tentative. NMB T.S.1065 possesses hypapophysis (instead of haemal keel), which is short and posteroventrally directed, as well as a posteriorly inclined neural spine, which is considerably higher than in other vertebrae. Other differences include the narrower centrum and the (slightly) more vaulted neural arch. Nevertheless, other typical features of the new species are still present in the specimen NMB T.S.1065, such as the thin zygosphene with a median lamina (“lip”) and two prominent lateral lobes, and the massive postzygapophyseal articular facets.</p><p>NMB T.S.1067could be a posterior trunk vertebra (Fig.6 G-L), judging from the wide haemal keel, or even an anterior cloacal vertebra, judging from the high neural arch. However, features such as the not-so-short neural spine should make an exact intracolumnar assignment uncertain. Rage (1984b) had already highlighted that unlike most snakes, posterior trunk vertebrae of Cheilophis huerfanoensis did not possess short neural spines compared to preceding vertebrae, and therefore this observation may apply also to the new European species. Nevertheless, certain diagnostic features of the new species are apparent in this specimen, such as the zygosphene with the “lip” and the distinct lateral lobes, as well as the massive prezygapophyseal and postzygapophyseal articular facets.</p><p>NMB T.S.1048 and NMB T.S.1066 are the only caudal vertebrae (Fig. 7) that are, at least tentatively, referred to Cheilophis periplanetes n. sp. They are characterized by the presence of haemapophyses and pleurapophyses. These specimens too possess a thick neural spine that is confined to the posterior portion of the neural arch. Judging from the elongation of their centrum and the shape of the haemapophyses, NMB T.S.1048 originates from the anterior caudal region (Fig.7 A-E), while NMB T.S.1066 represents a posterior caudal vertebra (Fig. 7 F-J). Interestingly, in the anterior caudal vertebra NMB T.S.1048, the haemapophyses are short and their base is situated onto a distinct keel in the centrum (Fig. 7A, B, D, E).</p></div>	https://treatment.plazi.org/id/03C687BCFF80FFEE7C88454637BCB354	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Mennecart, Bastien	Georgalis, Georgios L., Mennecart, Bastien (2025): Another wanderer from the Paleocene-Eocene Thermal Maximum? A new species of the North American snake genus Cheilophis Gilmore, 1938 from the early Eocene of France. Comptes Rendus Palevol 24 (16): 317-331, DOI: 10.5852/cr-palevol2025v24a16, URL: https://doi.org/10.5852/cr-palevol2025v24a16
