identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
F46987B6191E4C0FFEEE26FEFA58F916.text	F46987B6191E4C0FFEEE26FEFA58F916.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ranoidea Gray 1825	<div><p>Ranoidea gen. et sp. indet.</p><p>(Fig. 1)</p><p>REFERRED MATERIAL. — One incomplete right humerus (MNHN.F.IBC1603; Fig. 1).</p><p>DESCRIPTION</p><p>This humerus was regarded as a possible Ranidae in the first publication of the In Becetèn fauna (Broin et al. 1974). MNHN.F.IBC1603 was later illustrated and shortly described in 1984 (Rage 1984: fig. 1A) and considered to be a “Ranoid”. The specimen is the distal half of a right humerus (Fig.1). Most of the distal head is preserved (Fig. 1). The diaphysis is straight and not curved ventrally (Fig. 1 A-F). The crista ventralis is well developed (Fig. 1D, F). Most of the preserved region of the crista ventralis is divided into two ridges, with an elongate groove between the two; only the distalmost portion forms a single crest (i.e., no crista paraventralis; Fig. 1A, D). The crista medialis is thin (Fig.1B,C, D, F). The fossa cubitalis ventralis is triangular and distally broader than its proximo-distal diameter (width greater than height; Fig. 1A, D). The fossa is shallow (although it seems deeper in Fig. 1A) but well delimited by a thin, distinct ridge (Fig. 1A, D). The eminentia capitata is well developed and not shifted laterally from the diaphysis axis (Fig.1A, D). The epicondylus ulnaris is well prominent, reaching distally the level of the eminentia capitata (Fig. 1A, B, D, E), while the epicondylus radialis appears to be confluent with the eminentia (Fig. 1A, B, D, E). The olecranon scar extends proximally, with a pointed and tapered proximal margin (Fig. 1B, E).</p><p>DISCUSSION</p><p>This humerus is similar to the neobatrachian morphology in having: 1) a well-developed eminentia capitata; and 2) asymmetrical epicondyles (Prasad &amp; Rage 2004; Rage et al. 2021). Furthermore, the crista ventralis is divided into two ridges in numerous neobatrachians (this may denote muscular insertion; Otero et al. 2014). Moreover, the presence of: 1) a straight diaphysis; 2) a shallow fossa cubitalis ventralis; 3) an eminentia capitata not shifted laterally;and 4) an epicondylus ulnaris more developed than the epicondylus radialis, is similar to ranoid morphology (Rage et al. 2013; Lapparent de Broin et al. 2020). In addition, a well-delimited fossa cubitalis ventralis, although rare (Rage 1984), is observed in several ranoid taxa from the Eocene of the Quercy Phosphorites (Rage 2016) and among extant ranoids (Worthy 2001). MNHN.F.IBC1603 is therefore similar to ranoids and can be attributed to the Ranoidea . The presence of a distinct crista medialis is often linked to sexual and/or ontogenetic dimorphism (Duellman &amp; Trueb 1994). This suggests that this humerus might belong to a male ranoid. A humerus from the Cenomanian of the Kem Kem (Morocco) with ranoid affinities has also been recently described (UCRC-PV104; see Lemierre &amp; Blackburn 2022: fig. 6A-C). Both humeri share: 1) a straight diaphysis; 2) a well-developed eminentia capitata; 3) asymmetrically developed epicondyles; 4) an eminentia capitata not shifted from the diaphysis axis; and 5) a shallow fossa cubitalis ventralis.However, MNHN.F.IBC1603 differs from UCRC-PV 104 in: 1) having a more developed crista ventralis divided into two ridges; 2) a well-delimited fossa cubitalis; 3) a more protruding epicondylus ulnaris; 4) an olecranon scar more extended proximally; and 5) in having a discrete crista medialis.Thus, we think that these humeri likely represent different taxa. Furthermore, MNHN.F.IBC1603 possesses more ranoid characters than UCRC-PV104.In conclusion, MNHN.F.IBC1603 is attributed to a new unidentified male ranoid, and thus represent the oldest occurrence of Ranoidea .</p></div>	https://treatment.plazi.org/id/F46987B6191E4C0FFEEE26FEFA58F916	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	Lemierre, Alfred;Folie, Annelise;Bailon, Salvador;Laurin, Michel	Lemierre, Alfred, Folie, Annelise, Bailon, Salvador, Laurin, Michel (2026): Anurans of In Becetèn (Republic of Niger): the most diverse site for amphibians in Mesozoic Africa. Geodiversitas 48 (4): 51-67, DOI: 10.5252/geodiversitas2026v48a4
F46987B6191E4C03FC152439FB11FDAB.text	F46987B6191E4C03FC152439FB11FDAB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Neobatrachia Reig 1958	<div><p>Neobatrachia gen. et sp. indet.</p><p>(Fig. 2 B-O)</p><p>REFERRED MATERIAL. — 22 fragmentary maxillae (MNHN.F. IBC1983, IBC1984 and IBC1985) and eight fragments of exocranium (MNHN.F.IBC1986) that include incomplete frontoparietal, nasals and squamosals.</p><p>DESCRIPTION</p><p>Maxilla</p><p>The maxilla is thick and heavily ossified (Fig. 2). Most fragments are ornamented by small closely spaced tubercles forming a honeycomb pattern (Fig. 2B, C). The ornamentation covers most of the preserved labial surface of maxillae, except for a narrow strip along the ventral margin of the bone (Fig. 2B). The pars facialis is incomplete but seems high on most of its length (Fig. 2B). The lamina horizontalis is prominent, rounded (in cross-section; Fig. 2D) and slightly project lingually (Fig. 2D, F). Although no teeth are preserved, the pars dentalis shows that the maxilla was toothed (12 teeth positions preserved on the largest maxilla fragment; Fig. 2 E-G). The pars dentalis is dorsoventrally shallow and extends posteriorly beyond the pterygoid process (Fig. 2F). The pterygoid process is well developed and projects lingually (Fig. 2F). Posterior to this process, the maxilla narrows dorsally (Fig. 2E). Unfortunately, the posterior end of the maxilla has not been preserved. Anterior to the pterygoid process, the lamina horizontalis dorsally bears a shallow groove for the palatoquadrate (Fig. 2C). The pars facialis is high near the anterior margin of the maxilla, and likely indicates the presence of an anterodorsal process (Fig. A, G) as in Calyptocephalella Strand, 1928 or Ceratophrys Wied-Neuwied, 1824 . The anterior tip of the maxilla shows a deep fossa for the articulation with the premaxilla (Fig. 2G).</p><p>Nasal</p><p>Two fragments (both included under MNHN.F.IBC1986) are slender, flat pieces of bone. The dorsal surface is covered by the same ornamentation as in the maxillae (Fig. 2H).</p><p>Frontoparietal</p><p>The dorsal surface bears ornamentation similar to the one recovered in maxillae and nasal fragments (Fig. 2I). The pars contacta is curved externally (Fig. 2J, K). The pars contacta delimits a dorsal flange. This flange is interpreted as the tectum supraorbitale (Fig. 3I, J).</p><p>Squamosal</p><p>Several fragments of ornamented bone are interpreted as fragments of the squamosal. The best-preserved bone (MNHN.F.IBC1986c; Fig. 2 L-M) represents the posterior process which bears a rounded posterior margin (Fig. 2L). It is missing its medial margin, which suggests that it might have extended further and that a ramus paroticus was present (Fig. 2M). The base of a posterolateralis process and of the lamella alaris are preserved (Fig. 2L, M). Other specimens preserve part of the lamella alaris (Fig. 2N, O). The external surface is ornamented and thickened internally, forming a triangular surface (Fig. 2O). This fragment likely represents the lamella alaris of the ramus zygomaticus. Unfortunately, there is no indication whether and how the squamosal articulates with the maxilla.</p><p>DISCUSSION</p><p>Ornamentation on dermal bones in anurans occurs in various anuran clades (Paluh et al. 2020). However, combination of osteological characters can be here used to exclude certain major clades. Latonia Meyer, 1843 ( Alytidae Fitzinger, 1843) is the only known alytid exhibiting dermal ornamentation (Roček 1994). However, its current stratigraphic range is limited only to Oligocene and Neogene of Europe and Northern Africa (Gardner &amp; Rage 2016; Syromyatnikova et al. 2019). In addition, our ornamented taxon can be differentiated from Latonia in having a rounded medial margin of the lamina horizontalis (pointed in Latonia). Pelobatids are known to exhibit dermal ornamentation (Roček 2013) that resembles the ornamentation of the In Becetèn specimens (tubercles closely spaced together). However, our specimens differ from all pelobatids in lacking an elongate palatine process of the maxilla (Roček 2013). Furthermore, no pelobatids are known within the fossil record of Gondwana (a “ Pelobatidae ” was mentioned in the Upper Cretaceous of Madagascar, but the attribution is considered erroneous; Asher &amp; Krause 1998; Gardner &amp; Rage 2016) (Gardner &amp; Rage 2016). The In Becetèn ornamented specimens can also be differentiated from Pipimorpha in: 1) having a well-developed squamosal (small and reduced squamosal in Pipimorpha); and 2) a rather thick maxilla (slender maxillae in Pipimorpha).Two anurans of indeterminate phylogenetic position from North America possess similar ornamentation: Scotiophryne Estes, 1969 and Theatonius Fox, 1976 . Scotiophryne is known from the Late Cretaceous and might also occur in Early Cretaceous and Paleocene (Roček 2013; Gardner &amp; DeMar 2013). Scotiophryne differs from our taxon in having a narrow horizontal lamina (Gardner 2008). Theatonius is known from Late Cretaceous deposits (Gardner &amp; DeMar 2013). Theatonius differs from our taxon in lacking teeth on its maxillae. Thus, our taxon can be excluded from Leiopelmatoidea, Alytoidea, Pipimorpha, and Pelobatoidea. The only major clade left is the Neobatrachia . This includes several extinct and extant hyperossified taxa.</p><p>Two neobatrachians are known in the Mesozoic of Africa, Beelzebufo ampinga Evans, Jones and Krause, 2008 from the Maastrichtian of Madagascar and Cretadhefdaa taouzensis Lemierre &amp; Blackburn, 2022 from the Cenomanian of Morocco. Beelzebufo ampinga, Cretadhefdaa taouzensis and our specimens share: 1) ornamentation extending on most of the lateral surface of the maxilla (Fig. 2B); and 2) the presence of a tectum supraorbitale (Fig. 2 I-J). The latter is also shared with Cretadhefdaa Lemierre &amp; Blackburn, 2022 . However, Beelzebufo does not possess a distinct processus frontalis (present in our taxon; Fig. 2F) and all known Beelzebufo specimens are far larger than our specimens. Our specimens can be differentiated from Cretadhefdaa in: 1) having an ornamentation made of small tubercles resembling a honeycomb pattern (deep pits and ridges in Cretadhefdaa); and 2) lacking ornamentation on the lateral surface of the pars dentalis (present in Cretadhefdaa; Lemierre &amp; Blackburn 2022: fig. 3E-G).</p><p>Another Mesozoic hyperossified neobatrachian is Hungarobatrachus szukacsi Szentesi &amp; Venczel, 2010 from the Santonian of Hungary. Our specimens are similar to Hungarobatrachus in having: 1) ornamentation extending on most of the lateral surface of the maxilla (Fig. 2B); 2) a well-developed pterygoid process of the maxilla (Fig. 2C, F); and 3) a tectum supraorbitale (Fig. 2 I-J). However, our specimens differ from H. szukacsi in: 1) having a well-developed lamina horizontalis (moderately developed in H. szukacsi); and 2) in having the maxillary tooth row extending posteriorly beyond the pterygoid process.</p><p>One last hyperossified neobatrachian from the Mesozoic is Baurubatrachus Báez &amp; Perí, 1989 from the Maastrichtian of Brazil. Baurubatrachus and our specimens differ in the ornamentation on their cranial remains (pits and ridges in Baurubatrachus; Báez &amp; Gómez 2018). However, both are similar in having: 1) the medial margin of the lamina horizontalis rounded (in cross-section, Fig. 2D); and 2) a crista dentalis ending slightly beyond the pterygoid process (Fig.2F).</p><p>In Africa, the next oldest hyperossified neobatrachian is the ranoid Rocekophryne ornata Rage et al., 2021 from the Early Eocene of Algeria. Our specimens can be differentiated from Rocekophryne in having: 1) a different ornamentation (deep pits and ridges in Rocekophryne; Rage et al. 2021); 2) ornamentation extending ventrally to cover the entire lateral surface of the pars dentalis; and 3) in having an anterodorsal process (Fig. 2A). However, our specimens and Rocekophryne are similar in having a rounded lamina horizontalis (Fig. 2H). Another extinct hyperossified ranoid is Thaumastosaurus De Stefano, 1903, of which at least three to five taxa are known (Lemierre et al. 2021; Georgalis et al. 2023). Our specimens differ from Thaumastosaurus in: 1) having an ornamentation made of small tubercles and pits (deep pits and ridges in Thaumastosaurus; Fig. 2B); and 2) having a well-developed pterygoid process (Fig. 2D, F).</p><p>Thus, the In Becetèn ornamented anuran can be mainly differentiated from all extinct hyperossified anurans with the following characters: 1) a dermal ornamentation made of small closely spaced tubercles forming a honeycomb pattern (shared with Theatonius and Scotiophryne); and 2) a well-developed pterygoid process (shared with Hungarobatrachus).</p><p>Known extant hyperossified and ornamented neobatrachians mainly occur among hyloids, except for three genera, Aubria Boulenger, 1917, Cornufer Tschudi, 1838 and Pyxicephalus Tschudi, 1838 . Aubria and Pyxicephalus (in the African endemic family Pyxicephalidae) both possess the same characteristics on their maxillae as Thaumastosaurus De Stefano, 1903, except for having a more developed pterygoid process (Sheil 1999). Thus, our specimens can be differentiated from all Pyxicephalidae ( Thaumastosaurus, Aubria and Pyxicephalus). Cornufer is a hyperossified ranoid (family Ceratobatrachidae) from Indonesia. Its maxilla bears no anterodorsal process, and its anterior region is low and mostly devoid of ornamentation. Furthermore, the lamina horizontalis projects lingually into a thin blade in Cornufer . Thus, our specimens likely differ from all known hyperossified ranoids.</p><p>All remaining ornamented taxa are found within the Hyloidea, mainly among Ceratophryidae Tschudi, 1838 and Calyptocephalellidae Reig, 1960 . Our taxon resembles the Ceratophryidae in (likely) having an anterodorsal process on its maxilla. Our taxon differs from all ceratophryids in having a lamina horizontalis slightly protruding from the lingual surface of the maxilla (flattened in Ceratophryidae).</p><p>Calyptocephalellidae and its type genus, Calyptocephalella, are known throughout the South American fossil record since the Late Cretaceous (Agnolin 2012; Muzzopappa 2019; Muzzopappa et al. 2020). Incomplete specimens attributed to this clade have all been referred to Calyptocephalella, an attribution considered equivocal (Nicoli et al. 2017). Hence, we will consider all materials referred to Calyptocephalella as Calyptocephalellidae in this comparison. Our specimens resemble calyptocephalellids in having: 1) ornamentation covering the lateral surface of the maxilla, except for a thin strip of bone ventrally along the margin of the crista dentalis (Fig. 2B); and 2) in having an anterodorsal process (Fig. 2A). However, our specimens differ from calyptocephalellids in lacking a distinct lamina horizontalis on most of the length of the maxilla (in calyptocephalellids, the lamina horizontalis is flattened lateromedially only at the level of the orbit; Muzzopappa &amp; Báez 2009: fig. 3.2).</p><p>To summarize, we can exclude our specimens from all non-neobatrachian anuran clades, but we did not recover known synapomorphies to firmly attribute these fossils to Neobatrachia . However, they seem to share several characters on the maxilla with neobatrachian clades, like the presence of an anterodorsal process. Thus, these ornamented specimens represent a new neobatrachian taxon, here informally called “ Neobatrachia gen. et sp. indet.”. We refrain from erecting a formal taxon in the absence of better-preserved materials. To assess its affinities, we included this unnamed neobatrachian in a morphological dataset composed of extant and extinct anurans with emphasis on neobatrachians.</p></div>	https://treatment.plazi.org/id/F46987B6191E4C03FC152439FB11FDAB	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	Lemierre, Alfred;Folie, Annelise;Bailon, Salvador;Laurin, Michel	Lemierre, Alfred, Folie, Annelise, Bailon, Salvador, Laurin, Michel (2026): Anurans of In Becetèn (Republic of Niger): the most diverse site for amphibians in Mesozoic Africa. Geodiversitas 48 (4): 51-67, DOI: 10.5252/geodiversitas2026v48a4
F46987B619124C02FC2D219BFAA2F810.text	F46987B619124C02FC2D219BFAA2F810.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Anura Dumeril 1805	<div><p>Anura gen. et sp. indet.</p><p>REFERRED MATERIAL. — Nine fragments of maxillae (MNHN.F.IBC1989a, IBC1989b, IBC1991a-IBC1991f, IBC2063) .</p><p>DESCRIPTION</p><p>Maxillae</p><p>Three maxilla morphotypes distinct from the unnamed neobatrachian can be identified within In Becetèn: maxillary morphotypes A, B and C.</p><p>The two fragments attributed to the maxillary morphotype A (MNHN.F.IBC1989a, IBC1989b) bear ornamentation made of pits and ridges on their labial surface (Fig. 3A, B). This ornamentation covers the whole pars facialis, and most of the pars dentalis, leaving only a thin strip of smooth bone along the margin of the crista dentalis (Fig. 3B). The crista dentalis is shallow. One fragment (MNHN.F.IBC1989a) bears a distinct, rounded lamina horizontalis lingually (Fig. 3C). On the second fragment (MNHN.F.IBC1989b), there is no lamina horizontalis (Fig. 3D). Morphotype A can be differentiated from the above unnamed neobatrachian in being ornamented by pits and ridges (instead of small tubercles).</p><p>Four fragments are assigned to the maxillary morphotype B (MNHN.F.IBC1991a, IBC1991b, IBC1991c, IBC1991d). Three of them lack ornamentation on their labial surface (Fig. 3E, F). The only exception is the largest fragment (MNHN.F.IBC1991a), where a faint patch of rugose ornamentation is present near the base of the frontal process (Fig. 3E). All fragments bear a crista dentalis, with poorly-preserved teeth (Fig. 3C, D, G, I). In lingual view, the rounded, prominent lamina horizontalis is distinct on all fragments (Fig. 3G). The recessus vaginiformis is shallow, but well delimited ventrally and anterodorsally by two crests (Fig. 3G). The anterodorsal crest (processus palatinus) extends dorsally onto the lingual surface of the frontal process. This morphotype can be differentiated from the maxillary morphotype A and the unnamed neobatrachian in: 1) lacking ornamentation on its labial surface; and 2) by a thin lamina horizontalis that protrudes lingually.</p><p>Two fragments are assigned to the maxillary morphotype C (MNHN.F.IBC1991e, IBC2063). Their labial surface is covered in small pits and ornamentation imparting rugosity (Fig. 3H). This ornamentation seems to extend onto the whole labial surface. The maxilla is toothed (Fig. 3I). The zygomaticomaxillaris process projects posterodorsally (Fig. 3H, I). The orbital margin strongly decreases in height posterior to the latter process (Fig. 3H). Lingually, the lamina horizontalis is indistinct from the lingual surface of the maxilla (Fig. 3I), and no pterygoid process is present. The presence of a zygomaticomaxillaris process indicates that squamosal and maxilla were articulated in the maxillary morphotype C. This morphotype differs therefore from maxillary morphotypes A and B and the unnamed neobatrachian in being ornamented over the whole labial surface of the maxilla. It is also differentiated from the unnamed neobatrachian in: 1) lacking a pterygoid process; and 2) lacking a distinct lamina horizontalis.</p><p>DISCUSSION AND ATTRIBUTION</p><p>All three maxillary morphotypes are distinct from the unnamed neobatrachian and appear to represent three other distinct taxa. It should be noted that the presence of teeth does not exclude an attribution to the four pipimorph taxa known (Lemierre et al. 2023, 2025). Although most pipids lack teeth (Trueb et al. 2000), xenopodines and extinct pipimorphs are known to have teeth (Henrici &amp; Báez 2001; Báez &amp; Púgener 2003). However, the ornamentation present in morphotypes A and C differs from that of Pachycentrata Báez &amp; Rage, 2004 and the unnamed pipimorph 2 (Báez &amp; Rage 1998; Lemierre et al. 2025), while Inbecetenanura Lemierre, Bailon, Folie &amp; Laurin, 2023 and the unnamed pipimorph 1 lack ornamentation. Hence, maxillary morphotypes A, B and C can be assigned neither to Pachycentrata nor to unnamed pipimorph 2. Furthermore, the presence of small patches of ornamentation in the maxillary morphotype B around the frontal process renders an attribution to Inbecetenanura unlikely, as such ornamentation would likely be present on the frontoparietal. Hence, the maxillary morphotype B could be attributed to the unnamed pipimorph 1 or to another non-pipid taxon in In Becetèn. Maxillary morphotypes A and C are assigned to indeterminate non-pipid anurans.</p><p>Thus, based on maxilla elements, between three and four non-pipid anuran taxa are present in In Becetèn.</p></div>	https://treatment.plazi.org/id/F46987B619124C02FC2D219BFAA2F810	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	Lemierre, Alfred;Folie, Annelise;Bailon, Salvador;Laurin, Michel	Lemierre, Alfred, Folie, Annelise, Bailon, Salvador, Laurin, Michel (2026): Anurans of In Becetèn (Republic of Niger): the most diverse site for amphibians in Mesozoic Africa. Geodiversitas 48 (4): 51-67, DOI: 10.5252/geodiversitas2026v48a4
