taxonID	type	description	language	source
03B787BFFFACFFE2FF2FFB6AFBF0F815.taxon	description	(Fig. 2)	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFACFFE2FF2FFB6AFBF0F815.taxon	materials_examined	MATERIAL. — Suceag 1: 11 trunk vertebrae (UBB V 1042 / 1 - 2; UBB V 1043 / 1 - 9).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFACFFE2FF2FFB6AFBF0F815.taxon	description	DESCRIPTION UBB V 1042 / 1, is the largest available specimen representing a middle trunk vertebra, lacking its prezygapophyseal areas and the right posterolateral side of its neural arch. In dorsal view, the vertebra appears slightly longer than wide featuring a relatively deep interzygapophyseal constriction, whereas the posterior notch of the neural arch is also deep, bordered by the nearly straight posteromedian margins of the neural arch (Fig. 2 A). The neural spine, representing about one third of the length of the neural arch, is extremely low and widened, and without an anterior or posterior overhang; tiny tubercles are present on its dorsal anterior limit. The anterior margin of the zygosphene is slightly convex, with two protruding lateral lobes (the left one is damaged). In ventral view, the centrum is longer than wide (centrum length (CL) ≈ 3.85 mm; centrum width (CW) ≈ 3.26 mm; CL / CW ≈ 1.18), whereas the haemal keel is prominent, somewhat broadened anteriorly and posteriorly, but tapers near the condylar neck; on the right side a distinct subcotylar tubercle is preserved. The subcentral grooves are indistinct, whereas the subcentral foramina are present; on the right side a distinctly large subcentral foramen is preserved. Only the right paradiapophysis is preserved; it strongly projects laterally and features a well-developed tubercle on the anterior margin of the prezygapophyseal buttress (Fig. 2 B). The surface of the left postzygapophysis preserves numerous lines of arrested growths (LAGs) (for their interpretation, see Venczel et al. 2015), of which six are more discernible, considered as indication of intermittent seasonal growths (Venczel et al. 2015; Venczel 2023). In lateral view, the haemal keel is prominent and its ventral margin is relatively straight, whereas the subcentral ridges are weakly defined (Fig. 2 C). In anterior view, the roof of the zygosphene is straight with the lateral lobes tilting upward; the cotylar rim is damaged (Fig. 2 D). In posterior view, the neural arch is depressed, slightly convex dorsally and preserves on the left dorsal side of the postzygapophysis a low bony ridge extending anteroposteriorly. No parazygantral foramen is preserved (Fig. 2 E). The remaining trunk vertebrae are smaller and fragmentary, exhibiting some morphological variation compared to the UBB V 1042 / 1 specimen. The neural spine is consistently low and strongly broadened and with either parallel lateral margins (Fig. 2 F), or distinct posterior widening (Fig. 2 G). Some specimens feature a well-developed median lobe on the zygosphene (Fig. 2 F, G). In the UBB V 1042 / 2 specimen, the paradiapophyses are well-preserved and somewhat differentiated into diapophyseal and parapophyseal portions. The portions of prezygapophyseal buttresses between the paradiapophyses and prezygapophyseal processes, as seen in Falseryx neervelpensis (see Szyndlar et al. 2008), are developed into prominent, anteriorly facing tubercles (Fig. 2 H). In anterior view, the outline of the cotyle appears circular, without paracotylar foramina (Fig. 2 I). In UBB V 1043 / 1, representing an anterior trunk vertebra (Fig. 2 J, K), the neural arch appears more vaulted and less elongated, whereas the haemal keel is more prominent and extends posteriorly beyond the condylar neck.	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFACFFE2FF2FFB6AFBF0F815.taxon	discussion	REMARKS The size and morphology of the above-described vertebrae from Suceag 1 are closely resemble those of Falseryx neervelpensis known from the early Oligocene (MP 21) locality of Boutersem TGV, Belgium. In particular, the prominent tubercles protruding anteriorly on the prezygapophyseal buttresses are highly similar to those observed in this species (Szyndlar et al. 2008: figs 1, 2). The vertebrae of the type species of the genus (i. e., Falseryx petersbuchi Szyndlar & Rage, 2003), known from the Early Miocene (MN 4) localities of Petersbuch 2, Germany and Dolnice, Czech Republic (Szyndlar & Rage 2003), also bear resemblance to the specimens from Suceag 1. However, they lack the tubercles developed on the prezygapophyseal buttresses, which are considered diagnostic of F. neervelpensis (see Szyndlar et al. 2008).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFACFFE2FE90FDA8FCBBFB57.taxon	discussion	REMARKS The genus Falseryx Szyndlar & Rage, 2003, was originally established as a member of Boinae Gray, 1825 or Tropidophiinae Brongersma, 1951 (Szyndlar & Rage 2003), ‘ Tropidophiidae’ (Szyndlar et al. 2008), Tropidophiidae Brongersma, 1951 (Ivanov 2022), and finally classified as Alethinophidia incertae sedis (Smith & Georgalis 2022). The uncertainties regarding the phylogenetic position of Falseryx stem primarily from the fact that it is known exclusively from vertebrae. There is a single published vertebral synapomorphy of tropidophiids: the presence of an anteroventral straight corner of the hypapophyses of trunk vertebrae (Smith & Georgalis 2022; Szyndlar & Georgalis 2023; Zaher et al. 2024); notably, this feature is absent in Falseryx, which has a haemal keel instead of a hypapophysis in its mid- and posterior trunk vertebrae, hinting further that this genus does not belong to tropidophiids.	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFAFFFE1FE80F9EDFB37F815.taxon	description	(sensu Burbrink et al. 2020)	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFAFFFE1FE80F9EDFB37F815.taxon	discussion	REMARKS Ungaliophiidae were previously classified as a subfamily within Tropidophiidae (originally Ungaliopheinae of McDowell 1987). However, based on molecular data, ungaliophiids are considered to be much more distantly related to tropidophiids and instead represent distinct booids. This taxonomic classification is usually regarded as a subfamily within Charinaidae Gray, 1849 (Ungaliophiinae of Pyron et al. 2014; Georgalis & Smith 2020), or as a distinct family of Booidea, the Ungaliophiidae (Burbrink et al. 2020; Szyndlar & Georgalis 2023; Zaher et al. 2023). Parsimony analyses have recovered the latter as the sister clade of Charinaidae (Burbrink et al. 2020) (for further details see also Szyndlar & Georgalis 2023). Messelophis Baszio, 2004, is a genus of fossil dwarf boas from the middle Eocene of Messel locality, Germany, described firstly by Baszio (2004), based mainly on the morphology of its vertebrae. Another species of the genus, Messelophis ermannorum Schaal & Baszio, 2004, also from Messel, was later reassigned to a new genus by Scanferla et al. (2016): Rieppelophis Scanferla, Smith & Schaal, 2016. The latter genus appears in parsimony analyses as the sister taxon of Messelophis, while Messelophis + Rieppelophis has been recovered as the sister taxon of Exiliboa + Ungaliophis (Scanferla & Smith 2020 a, b).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFAEFFEEFF63FA0DFB0AFCA9.taxon	description	(Figs 3 A-K; 4 A-I)	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFAEFFEEFF63FA0DFB0AFCA9.taxon	materials_examined	MATERIAL. — CetăŢuia Hill: one anterior trunk vertebra (UBB V 1045), four trunk vertebrae (UBB V 1046 / 1 - 4); Suceag 1: eight trunk vertebrae (UBB V 1047 / 1 - 8), one caudal vertebra (UBB V 1044).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFAEFFEEFF63FA0DFB0AFCA9.taxon	description	DESCRIPTION The specimen UBB V 1045 represents a partial anterior trunk vertebra with the roof of its neural arch broken off (Fig. 3 A, B). The centrum is slightly longer than wide and features a salient and posteroventrally projecting hypapophysis. Two small subcentral foramina are present and the paradiapophyses appear robust and undifferentiated, however their surfaces are strongly eroded. One of the best preserved mid-trunk vertebrae from the CetăŢuia Hill locality is represented by UBB V 1046 / 1 (Fig. 3 C-F); however, in the specimen the right prezygapophysis and the left postzygapophysis are completely broken off, whereas the lateral margin of the left prezygapophysis, the ventral margin of the cotylar lip and the posterior tip of the condyle are also missing. The estimated centrum length of the UBB V 1046 / 1 specimen approaches 2.42 mm, whereas its centrum width approaches 2.39 mm; in the UBB V 1047 / 1 specimen from Suceag 1 locality (Fig. 4 A-C) the centrum length reaches 2.64 mm, whereas its centrum width equals 2.46 mm. In dorsal view, the most striking feature is that the neural spine is extremely short and point-like, positioned on the upraised dorsoposterior border of the neural arch, whereas the area in front of the neural spine is flat (Figs 3 C, G, H; 4 A, D, E); however, in all the specimens the distal tip of the neural spine is broken off (Figs 3 G, H; 4 H). The interzygapophyseal constriction is moderate and a well-defined posterior notch is present on the neural arch. The anterior margin of the zygosphene is provided with two lateral lobes and with a less developed median lobe (Figs 3 C; 4 A). In ventral view, the haemal keel is well-defined, more salient in the specimens from CetăŢuia Hill (Figs 3 D, I; 4 F), whereas in those from Suceag 1, it is somewhat widened and flattened (Fig. 4 B). The subcentral foramina are present and of variable size, the subcentral grooves are weakly defined, whereas the paradiapophyses project strongly laterally; the prezygapophyseal process is vestigial (Fig. 3 I, J). In lateral view, the vertebrae appear moderately flattened with the neural arch elevated near the neural spine (Fig. 4 C, D). The paradiapophyses are undifferentiated and robustly built, as seen in specimen UBB V 1046 / 3 (Fig. 3 J). In anterior view, the neural arch is slightly flattened, the neural spine is relatively low, whereas the cotyle is circular and the paracotylar foramina are lacking; the zygosphenal roof is almost horizontal (Fig. 3 E, H). In posterior view, the neural arch appears slightly convex, the condyle is rounded, whereas the parazygantral foramina are lacking (Fig. 3 F, K). A single caudal vertebra (UBB V 1044), representing a small sized individual (however, about five lines of the arrested growths are observed on its prezygapophysis indicating that it was an adult snake), possesses a low neural spine extending from the base of the zygosphene to the posterior notch of the neural arch (Fig. 4 G). The centrum is longer than wide, whereas the neural arch is depressed possessing a weak interzygapophyseal constriction (Fig. 4 H); below the right prezygapophysis, a tiny prezygapophyseal accessory process is preserved, and the prezygapophysis itself is elongated. In ventral view, the pleurapophyses are broken off; however, the remnants of the paired haemapophyses are partially preserved (Fig. 4 H, I).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFAEFFEEFF63FA0DFB0AFCA9.taxon	discussion	REMARKS Some of the phenotypical features of the vertebrae described above closely resemble those of Messelophis variatus (e. g. presence of an extremely short “ point-like ” neural spine, developed near the posterior border of the neural arch and the paracotylar foramina absent). Similar to the specimens of our study, the vertebrae of M. variatus are elongated, with extremely short neural spines developed at the posterior border of the neural arch, the paracotylar foramina are absent and the prezygapophyseal accessory processes are weakly developed (Baszio 2004; Schaal & Baszio 2004; Scanferla et al. 2016: fig. 1 B). Moreover, in the holotype of M. variatus (SMF-ME 1828 A-B), exposing a short portion of the ventral side of its vertebral column (MV pers. obs.), and in the recently referred specimen SMF-ME 513 a from the early middle Eocene of Messel, Germany, exposing the ventral side of its vertebral column (Scanferla & Smith 2020 b: fig. 1 B), the trunk vertebrae possess prominent haemal keels of variable shape (i. e., of gladiate or spatulate-shape, sensu Auffenberg 1963). The caudal vertebra UBB V 1044, similarly to M. variatus, possesses a low neural spine and provided with paired haemapophyses (Scanferla et al. 2016: fig. 18 A); however, in M. variatus the haemapophyses appear laminar, whereas in UBB V 1044 the remnants of the latter structure comparatively are less elongated anteroposteriorly. A number of comparable attributes of the studied specimens should be noted also in the fossil genus Dunnophis Hecht in McGrew et al., 1959, which was originally designated based on isolated vertebrae as an incertae sedis snake from the late early Eocene of North America (Hecht 1959), and recorded later from the Eocene of Western Europe (e. g. Rage 1973, 1974, 1984, 2006; Rage & Ford 1980; Rage & Augé 2003, 2010; see Smith & Georgalis 2022), and even from the late Palaeocene of Morocco (Augé & Rage 2006). Among the peculiar features of Dunnophis, as noted by the above authors, are the following: the centrum is depressed, narrow and longer than wide; the extremely short neural spine is developed near the margin of the deep posterior notch of the neural arch; the prezygapophyseal processes are absent or vestigial; and the caudal vertebrae, rarely present in the fossil material, likely possessed haemapophyses, as seen in the North American type species of the genus, Dunnophis microechinis Hecht in McGrew et al., 1959 (see Hecht 1959: plate 56: 7 - 10). Resemblance between Dunnophis and Ungaliophis Müller, 1880 was already noted by Bogert (1968), while similarity between Dunnophis and Messelophis was mentioned by Baszio (2004). Rage & Augé (2010) suggested that Messelophis might be considered a junior synonym of Dunnophis. On the other hand, the specimens from the early Oligocene of Romania described herein, share with the two recent genera of ungaliophiids (i. e., Exiliboa Bogert, 1968, and Ungaliophis) a number of phenotypic features, as follows: lightly built vertebrae with their centrum longer than wide, the presence of a prominent haemal keel in the trunk vertebrae (according to Szyndlar & Georgalis, 2023, the haemal keel is less developed in Ungaliophis), the absence of paracotylar foramina, and weakly developed or vestigial prezygapophyseal accessory processes. Some of these characters (e. g. elongation of the trunk vertebrae and presence of haemal keel in the caudal vertebrae) have been considered synapomorphies of Ungaliophiidae within the clade of Constrictores (Smith 2013).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFA0FFEEFC66FC08FAA9F954.taxon	description	(Fig. 4 J-N)	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFA0FFEEFC66FC08FAA9F954.taxon	materials_examined	MATERIAL. — Treznea: one posterior caudal vertebra (UBB V 1048).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFA0FFEEFC66FC08FAA9F954.taxon	description	DESCRIPTION The only available caudal vertebra (UBB V 1048) is known from the Treznea locality (Fig. 4 J-N). It appears well-preserved; however, its surface displays some signs of erosion. The neural arch is strongly depressed and the neural spine is reduced to a low keel. In ventral and lateral views, the centrum possesses a prominent haemal keel, connected to a rounded condyle. The haemal keel is somewhat enlarged posteriorly and displays on the posteroventral side two small knobs (Fig. 4 K).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
03B787BFFFA0FFEEFC66FC08FAA9F954.taxon	discussion	REMARKS The main feature of the single available specimen (UBB V 1048) is the presence of a well-developed haemal keel on the subcentral surface, reminiscent of Rieppelophis ermannorum (Schaal & Baszio, 2004), in which the haemapophyses in the caudal vertebrae are replaced by paired knobs (Scanferla et al. 2016: fig. 18 B). Additionally, it shares similarities with recent ungaliophiids (Exiliboa and Ungaliophis), which possess a distinct haemal keel in the caudal vertebrae, instead of paired haemapophyses (Szyndlar & Georgalis 2023).	en	Venczel, Márton, Codrea, Vlad A., Solomon, Alexandru A., Fărcaș, Cristina, Bordeianu, Marian (2025): Late Eocene-early Oligocene snakes from the Transylvanian Basin (Romania). Comptes Rendus Palevol 24 (13): 229-240, DOI: 10.5852/cr-palevol2025v24a13, URL: https://doi.org/10.5852/cr-palevol2025v24a13
