identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
05CA534F9A455C7D8499D2F6B547A667.text	05CA534F9A455C7D8499D2F6B547A667.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Caenophidia Hoffstetter 1939	<div><p>Caenophidia Hoffstetter, 1939</p><p>Colubriformes Günther, 1864 (sensu Zaher et al., 2009)</p><p>Remarks.</p><p>Most of our snake vertebrae describe here belong to the group of Colubriformes. Within Colubriformes, hypapophyses are typically present in mid- and posterior trunk vertebrae of several subgroups, among others, natricids, elapids, and viperids (Georgalis and Scheyer 2022). The absence of hypapophyses in mid- and posterior trunk vertebrae has long been considered characteristic of Colubridae (e. g., Boulenger 1896; Bogert 1940; Underwood 1967; Bourgeois 1968; Dowling and Duellman 1978) or of the paraphyletic group traditionally referred to as “ Colubrinae ” (Rage 1984; Szyndlar 1987, 1991 a, 2012; Georgalis et al. 2018). However, recent studies have shown that more distantly related groups, such as dipsadids and psammophiids, may also lack hypapophyses in mid- and posterior trunk vertebrae (Georgalis et al. 2019, 2024; Zaher et al. 2019; Georgalis and Scheyer 2022; Georgalis and Szyndlar 2022). It is important to note that both of these non-colubrid groups are native to the New World and are absent from Taiwan.</p></div>	https://treatment.plazi.org/id/05CA534F9A455C7D8499D2F6B547A667	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	Lin, Chien-Hsiang;Lin, Si-Min;Chien, Chi-Wei;Lin, Te-En;Nazir, Haroon;Singh, Ningthoujam Premjit	Lin, Chien-Hsiang, Lin, Si-Min, Chien, Chi-Wei, Lin, Te-En, Nazir, Haroon, Singh, Ningthoujam Premjit (2025): First fossil frog and snake assemblage from southern Taiwan: a window into Pleistocene herpetofauna and palaeoenvironments in subtropical East Asia. Fossil Record 28 (2): 293-308, DOI: 10.3897/fr.28.164568
28742CDC146C59DCBDE782F8B1E5E083.text	28742CDC146C59DCBDE782F8B1E5E083.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Duttaphrynus melanostictus (Schneider 1799)	<div><p>Duttaphrynus melanostictus (Schneider, 1799)</p><p>Fig. 2 A</p><p>Material.</p><p>Trunk vertebra (ASIZF 0101045).</p><p>Description.</p><p>The vertebra is large, wider than long, with a centrum that is procoelous and slightly dorsoventrally compressed. The neural canal is larger than both cotyle and condyle. The walls of the neural arch are robustly built, bearing a low but distinct carina neuralis. The prezygapophyses and postzygapophyses are dorsally elevated and extend laterally. A single transverse process is preserved and extends distally.</p><p>Remarks.</p><p>The substantial size of the vertebra (&gt; 4.5 mm in length), its proportions, and robust construction suggest that it belongs to the Bufonidae . Given its dimension and general morphology, the vertebra most likely represents a mid-trunk element, probably the 3 rd or 4 th presacral vertebra. Of the more than 30 native anuran species in Taiwan, only two bufonids, Bufo bankorensis (Barbour, 1908) and Duttaphrynus melanostictus, attain a comparable size. Trunk vertebrae of both species are morphologically similar; however, the centrum of B. bankorensis is more elevated in lateral view (Fig. 2 C 5), whereas that of D. melanostictus is flatter and more dorsoventrally compressed.</p></div>	https://treatment.plazi.org/id/28742CDC146C59DCBDE782F8B1E5E083	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	Lin, Chien-Hsiang;Lin, Si-Min;Chien, Chi-Wei;Lin, Te-En;Nazir, Haroon;Singh, Ningthoujam Premjit	Lin, Chien-Hsiang, Lin, Si-Min, Chien, Chi-Wei, Lin, Te-En, Nazir, Haroon, Singh, Ningthoujam Premjit (2025): First fossil frog and snake assemblage from southern Taiwan: a window into Pleistocene herpetofauna and palaeoenvironments in subtropical East Asia. Fossil Record 28 (2): 293-308, DOI: 10.3897/fr.28.164568
5FDAFE8240885374A3209187926FC481.text	5FDAFE8240885374A3209187926FC481.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lycodon rufozonatum Cantor 1842	<div><p>Lycodon rufozonatum Cantor, 1842</p><p>Fig. 3 A, B</p><p>Material.</p><p>Two trunk vertebrae (ASIZF 0101048 –1049).</p><p>Description.</p><p>In anterior view, the cotyle is rounded in outline. Paracotylar foramina are present on either side of the cotyle. The prezygapophyses are well-developed, and the prezygapophyseal accessory processes are laterally extended. The zygosphene roof is thin and dorsally convex. The zygosphenal articular facets of the zygosphene are inclined dorsally. The neural canal is tunnel-like, with a width approximately equal to that of the cotyle.</p><p>In posterior view, the condyle is circular in shape and proportionally similar in height and width. The neural arch is extremely depressed to depressed (vaulting ratio = 0.19–0.23). The zygantral area is deep and contains paired endozygantral foramina.</p><p>In ventral view, the centrum is triangular and relatively short. Small paired subcentral foramina are visible flanking a well-developed, oblanceolate haemal keel that extends nearly to the condyle. Subcentral grooves are deep and long. Postzygapophyseal articular facets are oval. Synapophyses are clearly divided into diapophyses and parapophyses, with the absence of parapophyseal processes.</p><p>In dorsal view, the vertebrae are slightly wider than long. The anterior margin of the zygosphene is straight with two blunt lobes on the lateral sides. The posterior median notch is distinct and deep. Prezygapophyseal articular facets are oval.</p><p>In lateral view, both interzygapophyseal and subcentral ridges are straight. The neural spine is high, with the posterior end protruding above its base and the anterior end overhanging. Lateral foramina are present.</p><p>Remarks.</p><p>The two trunk vertebrae differ from those of Ptyas by lacking epizygapophyseal spines (Ikeda 2007; Nakamura et al. 2013). Additionally, several features, including the straight zygosphenal lip with two small blunt lobes on the lateral sides, exclude their allocation to Coronella, which possesses a crenate zygosphenal lip, and a wide haemal keel (Ivanov et al. 2018). The fossils can also be distinguished from Texasophis and Telescopus by their relatively large size and distinct subcentral ridges (Szyndlar 1987, 2005; Čerňanský et al. 2017).</p><p>The fossil vertebrae also closely resemble those of Elaphe and Lycodon, sharing characters including undeveloped parapophyseal process, well-developedprezygapophyseal accessory processes, and oval zygosphenal articular facets (Chen 2020). However, in Elaphe, the trunk vertebrae generally exhibit less prominent subcentral ridges and indistinctly developed subcentral grooves (Ratnikov 2004; Ivanov et al. 2018), which is not identical to the prominent subcentral ridges and deep subcentral grooves in the fossils, although Ratnikov (2022) noted that Elaphe trunk vertebrae show progressive morphological changes along the column, such as deepening of subcentral grooves, more distinct subcentral ridges, reduced haemal keel width, and increased neural spine height posteriorly.</p><p>Two extant species of Lycodon, Lycodon ruhstrati (Fischer, 1886) and Lycodon rufozonatus (Cantor, 1842), are found in Taiwan. The vertebrae of L. ruhstrati possess flat and very low neural spines, whereas in L. rufozonatus, these are substantially high as in our fossils (Fig. 3 C, D).</p></div>	https://treatment.plazi.org/id/5FDAFE8240885374A3209187926FC481	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	Lin, Chien-Hsiang;Lin, Si-Min;Chien, Chi-Wei;Lin, Te-En;Nazir, Haroon;Singh, Ningthoujam Premjit	Lin, Chien-Hsiang, Lin, Si-Min, Chien, Chi-Wei, Lin, Te-En, Nazir, Haroon, Singh, Ningthoujam Premjit (2025): First fossil frog and snake assemblage from southern Taiwan: a window into Pleistocene herpetofauna and palaeoenvironments in subtropical East Asia. Fossil Record 28 (2): 293-308, DOI: 10.3897/fr.28.164568
91B799CA08E55172A2B26FE22A442BC1.text	91B799CA08E55172A2B26FE22A442BC1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ptyas mucosa (Linnaeus 1758)	<div><p>Ptyas mucosa (Linnaeus, 1758)</p><p>Fig. 4 A</p><p>Material.</p><p>One trunk vertebra (ASIZF 0101050).</p><p>Description.</p><p>In anterior view, the cotyle appears rounded and flanked by paracotylar foramina. The neural canal is large and circular. The prezygapophyses are well-developed, with the ending portions of their accessory processes prominently directed forward and slightly ventrally inclined. The zygosphene roof is straight, and its articular facets are dorsally tilted.</p><p>In posterior view, the condyle is rounded and the neural arch is depressed (vaulting ratio = 0.24), with paired endozygantral foramina visible. In ventral view, the centrum is elongated and triangular in shape. Subcentral foramina are present, and both subcentral grooves and ridges are clearly defined. The haemal keel is well-developed and oblanceolate. Postzygapophyseal articular facets are oval. Epizygapophyseal spines are developed and directed posterolaterally. Diapophyses and parapophyses are clearly visible, though the parapophyseal processes are absent.</p><p>In dorsal view, the vertebra is slightly laterally compressed. The anterior margin of the zygosphene is straight with two small, blunt lateral lobes. The posterior median notch is markedly deep. Prezygapophyseal articular facets are oval, and the prezygapophyseal accessory processes are well-developed, pointed, and directed anterolaterally. In lateral view, the interzygapophyseal and subcentral ridges are straight. The neural spine is damaged. The lateral foramina are observed.</p><p>Remarks.</p><p>The fossil vertebra can confidently be attributed to colubrids or “ colubrines ”, further supported by several combined features: a well-developed haemalkeel (instead of a hypapophysis), the presence of paracotylar foramina, a pronounced division between the para- and diapophyses, and a moderately high neural spine. Moreover, its overall vertebral morphology closely matches that of Ptyas, which share characteristics such as epizygapophyseal spines, pointed prezygapophyseal accessory processes, and distinct subcentral grooves and ridges (see Ikeda 2007; Shi et al. 2023).</p><p>The strongly anteriorly directed prezygapophyseal accessory processes, however, are a distinctive feature seen only in the two extant Ptyas species from Taiwan, Ptyas major (Günther, 1858) and Ptyas mucosa (Linnaeus, 1758) (see Chen 2020). Among these, P. major is a medium-sized species, while P. mucosa notably larger, and the vertebrae of P. major are consistently smaller than those of Ptyas mucosa, as seen in our comparative specimens (Fig. 4 B, C).</p></div>	https://treatment.plazi.org/id/91B799CA08E55172A2B26FE22A442BC1	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	Lin, Chien-Hsiang;Lin, Si-Min;Chien, Chi-Wei;Lin, Te-En;Nazir, Haroon;Singh, Ningthoujam Premjit	Lin, Chien-Hsiang, Lin, Si-Min, Chien, Chi-Wei, Lin, Te-En, Nazir, Haroon, Singh, Ningthoujam Premjit (2025): First fossil frog and snake assemblage from southern Taiwan: a window into Pleistocene herpetofauna and palaeoenvironments in subtropical East Asia. Fossil Record 28 (2): 293-308, DOI: 10.3897/fr.28.164568
B8C4963B851E5B2BABD891D2430B5CA7.text	B8C4963B851E5B2BABD891D2430B5CA7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ptyas mucosa (Linnaeus 1758)	<div><p>Ptyas cf. mucosa (Linnaeus, 1758)</p><p>Fig. 5</p><p>Material.</p><p>Two trunk vertebrae (ASIZF 0101051 –1052).</p><p>Description.</p><p>In anterior view, the cotyle is rounded. In specimen ASIZF 0101052, paracotylar foramina are present, located on the lateral margins of the cotyle, whereas in specimen ASIZF 0101051, these foramina are not clearly visible. The prezygapophyses are well-developed, but the prezygapophyseal accessory processes are broken laterally. The zygosphene roof is dorsally convex and wider than the cotyle, with articular facets inclined dorsally. The neural canal is tunnel-like.</p><p>In posterior view, the condyle is circular, with height similar to width. The neural arch is depressed (vaulting ratio = 0.22–0.25). The zygantral area is deep and the paired endozygantral foramina are present.</p><p>In ventral view, the centrum is distinctly longer than wide and triangular in outline. A well-developed, oblanceolate haemal keel is present instead of a hypapophysis, almost reaching the condyle. Subcentral foramina are not clearly visible. The subcentral grooves are deep and long. Postzygapophyseal articular facets are oval. The paradiapophyses comprise a laterallysalient diapophysis and an almost flat parapophysis.</p><p>In dorsal view, the anterior edge of the zygosphene is straight with two small pointed lobes on the lateral sides. The posterior median notch is distinctly deep. Prezygapophyseal articular facets are oval.</p><p>In lateral view, the height of the neural spine is indeterminate in specimen ASIZF 0101051 due to poor preservation, whereas it is very high in ASIZF 0101052. Both interzygapophyseal and subcentral ridges are distinct. Lateral foramina are present.</p><p>Remarks.</p><p>Although the prezygapophyseal accessory processes are only partially preserved, the remaining portions appear to be anteriorly directed. Combined with the large overall size of the vertebrae and the presence of distinct subcentral ridges with deep grooves, these features most closely match Ptyas mucosa, as described above. However, the poor preservation of both specimens — particularly the absence of epizygapophyseal spines, which may have been worn away — precludes a definitive assignment.</p></div>	https://treatment.plazi.org/id/B8C4963B851E5B2BABD891D2430B5CA7	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	Lin, Chien-Hsiang;Lin, Si-Min;Chien, Chi-Wei;Lin, Te-En;Nazir, Haroon;Singh, Ningthoujam Premjit	Lin, Chien-Hsiang, Lin, Si-Min, Chien, Chi-Wei, Lin, Te-En, Nazir, Haroon, Singh, Ningthoujam Premjit (2025): First fossil frog and snake assemblage from southern Taiwan: a window into Pleistocene herpetofauna and palaeoenvironments in subtropical East Asia. Fossil Record 28 (2): 293-308, DOI: 10.3897/fr.28.164568
279252DD2CC1576310B925A70ED38170.text	279252DD2CC1576310B925A70ED38170.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Undetermined undetermined Oppel 1811	<div><p>Colubridae indet.</p><p>Fig. 6 A, B</p><p>Material. Two caudal vertebrae (ASIZF 0101046 –1047).</p><p>Description. In anterior view, the cotyle is rounded. Paracotylar foramina are present, located on the lateral margins of the cotyle. The prezygapophyseal accessory processes and pleurapophyses are developed and pointed in specimen ASIZF 0101046, but they are broken laterally in specimen ASIZF 0101047. The zygosphene roof is dorsally convex and about the same width as the cotyle, with articular facets inclined dorsally. The neural canal is tunnel-like.</p><p>In posterior view, the condyle is circular, with height similar to width. The neural arch is depressed (vaulting ratio = 0.23). The zygantral area is deep and the paired endozygantral foramina are present. In ventral view, the centrum is distinctly longer than wide. Well-developed haemapophyses are partially present in specimen ASIZF 0101046, almost reaching the condyle. Subcentral foramina are visible. Postzygapophyseal articular facets are oval in ASIZF 0101046.</p><p>In dorsal view, the posterior median notch is distinctly deep in specimen ASIZF 0101046. Prezygapophyseal articular facets are oval. In lateral view, the height of the neural spine is indeterminate in ASIZF 0101047 due to poor preservation, whereas it is high in ASIZF 0101046. Both interzygapophyseal and subcentral ridges, and lateral foramina are distinct in ASIZF 0101046.</p><p>Remarks. The caudal vertebrae closely resemble those of Elaphe as described by Ratnikov (2022), particularly in terms of their elongated morphology from anterior to posterior, and well-developed haemapophyses and pleurapophyses. However, colubrid caudal vertebrae are often not suitable for genus-level determination.</p></div>	https://treatment.plazi.org/id/279252DD2CC1576310B925A70ED38170	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	Lin, Chien-Hsiang;Lin, Si-Min;Chien, Chi-Wei;Lin, Te-En;Nazir, Haroon;Singh, Ningthoujam Premjit	Lin, Chien-Hsiang, Lin, Si-Min, Chien, Chi-Wei, Lin, Te-En, Nazir, Haroon, Singh, Ningthoujam Premjit (2025): First fossil frog and snake assemblage from southern Taiwan: a window into Pleistocene herpetofauna and palaeoenvironments in subtropical East Asia. Fossil Record 28 (2): 293-308, DOI: 10.3897/fr.28.164568
