taxonID	type	description	language	source
039987DFFFE9C7577D23C6FBFC03F872.taxon	description	(Text-fig. 2: 1 – 6)	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFE9C7577D23C6FBFC03F872.taxon	description	In the Oligocene, alytids occur from MP 21 to MP 25 and in MP 28. Alytids from the Oligocene were all referred to as Discoglossus cf. giganteus by de Bonis et al. (1973: table 2 – 5). D. giganteus WETTSTEIN- WESTERHEIMB, 1955 is now included in the genus Latonia, as L. gigantea (Roček 1994, Rage and Hossini 2000). Before Roček’s revision of Latonia (Roček 1994), various alytids from the Cainozoic were erroneously assigned to D. giganteus. However, there is likely more than one alytid taxon in the Oligocene of the Phosphorites. The few alytids from the early Oligocene (MP 21 and MP 22) appear to be about twice the size of the younger specimens (Text-fig. 2: 2, 3) and, based on urostyles, their size is similar to that of the only specimen known from the Eocene. This difference in size may correspond to a taxonomic distinction, but this cannot be demonstrated on the basis of the available material. In addition, in MP 23, large and smaller alytids coexisted. Duffaud (2000) suggested that Latonia vertaizoni FRIANT, 1944 might be present in the Oligocene of the Phosphorites. The latter taxon is known by a single specimen from Vertaizon, a locality relatively close to the Quercy and whose age is regarded as late Oligocene, although an early Miocene age cannot be rejected (Gaudant 1993). Unfortunately, comparisons are almost impossible because the holotype of L. vertaizoni is an articulated skeleton whose bones are poorly preserved (Roček 1994: fig. 19). The Alytidae from the Phosphorites cannot be identified to genus level. However, various features observable on the available bones demonstrate that these alytids belong to the ‘ Discoglossus group’ (presence of a dorsal crest on the ilium, tuber superius formed by a thickening of the posterior part of the latter crest, vertebral centrum cylindrical, neural spine projecting posteriorly beyond the level of the postzygapophyses, distal part of the humerus relatively expanded transversely; Text-fig. 2: 4 – 6).	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFEAC7567FFDC5B1FB31FF2E.taxon	description	(Text-fig. 2: 7, 8)	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFEAC7567FFDC5B1FB31FF2E.taxon	description	Duffaud (2000) mentioned Eopelobates aff. bayeri from the late Eocene (MP 19) and the earliest Oligocene (MP 21) of the Phosphorites. Rage (2006) reported those fossils as Eopelobates aff. anthracinus because Sanchiz (1998) regarded E. bayeri ŠPINAR, 1952 as a junior synonym of E. anthracinus PARKER, 1929; however, this synonymy was not recognized by Roček et al. (2014). However, the available material does not permit such precise identification. At present, it is only possible to state that Pelobatidae are present in the Phosphorites from MP 16, late middle Eocene (Rage 1988) to MP 23, early / middle Oligocene (Duffaud 2000). In younger levels of the Phosphorites, anurans become rare and, despite the report of a pelobatid as cf. Pelobates from the late Oligocene (MP 28) (Crochet 1972), the presence of representatives of the family after MP 23 has not been confirmed.	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFEBC7517FD4C198FC57FB6F.taxon	description	(Text-fig. 3) In the Phosphorites, the taxon is represented by vertebrae, ilia, humeri and one fused tibiale-fibulare (Text-fig. 3). Vertebrae show a combination of features that is characteristic of Pelodytes (Text-fig. 3: 1, 2). They are proceolous, lightly built and the centrum, cotyle and condyle all are markedly depressed dorsoventrally. In ventral aspect, the ventral parts of the lateral walls typically extend laterally on either side of the centrum. The neural arch is of the imbricate type, i. e. it is anteroposteriorly long. The posterior projection of the neural spine is relatively weak. In posterior presacral vertebrae, the bases of the transverse processes originate approximately below the prezygapophyses and they project markedly anterolaterally. One incomplete sacral vertebra may belong to this taxon; however, it is regarded as belonging to an indeterminate anuran because it bears postzygapophyses (see below: ‘ Problematic taxa’). The ilia lack both a dorsal crest and a tuber superius, as do those of pelobatids (Text-fig. 3: 4). However, their shaft is markedly curved in lateral view and the dorsal border of their pars ascendens (ischiatic process) is slightly curved dorsally, which distinguishes them from pelobatid ilia. Humeri are very similar to those of Pelodytes (Text-fig. 3: 3). They have a slender and almost straight diaphysis that bears ventral and paraventral crests. The articular ball is well defined, comparatively small and scarcely shifted laterally. The distal part of the lateral border of the bone (crista lateralis not taken into account), close to the articular ball, forms a gentle curve that is slightly convex laterally; this part of the humerus is generally concave in anurans. A fused tibiale-fibulare (Text-fig. 3: 5) belonging to a pelodytid was recovered among the specimens of anurans from the late Eocene of Malpérié (MP 17) by Duffaud (2000). In almost all anurans, the tibiale and fibulare are elongate and fused only at their extremities. However, in pelodytids, centrolenids (an extant Central and South American family) and Tephrodytes HENRICI, 1994 (but not in rhinophrynids, contra Trueb 1973), they are fused throughout their length and form a single element. This element thus resembles the anuran tibiofibula; however, it is shorter and its extremities are more expanded. In addition the proximal ends of the tibiale and fibulare are approximately circular, whereas the distal extremities are clearly flattened. Traces of the fusion remain as grooves, except in the central portion of the bone. This tibiale-fibulare from Malpérié clearly displays this morphology, which is characteristic of Pelodytes (Sanchiz 1978). It differs from that of Miopelodytes TAYLOR, 1941 in being more slender; more specifically, its extremities are less expanded. The fused tibiale-fibulare of centrolenids is markedly more elongate and slender (Guayasamin et al. 2009). The fused tibiale-fibulare from Malpérié cannot be distinguished from the tibiale-fibulare of extant Pelodytes and it also closely resembles that of Tephrodytes (Oligocene-Miocene transition of the USA; Henrici 1994). Tephrodytes was first referred to the Pelodytidae (Henrici 1994) but Henrici et al. (2013) placed it among pelobatids s. l. However, the morphology of various bones suggests that Tephrodytes may be more closely related to the Pelodytidae. In conclusion, the specimens from the Phosphorites allocated to Pelodytidae are quite similar to those of extant Pelodytes. Henrici et al. (2013: 304) noted that reliable referral of the remains from the Eocene of Europe to pelodytids (as cf. Pelodytes) would require finding of a fused tibiale-fibulare in the same localities. The recovery of such a tibiale-fibulare confirms that a fossil taxon closely related to Pelodytes, if not Pelodytes, is present in the late middle and late Eocene (from MP 16 to MP 19) of western Europe, more specifically in the Phosphorites.	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFECC7517FF5C698FA92F853.taxon	description	The earliest ranoid is represented by a few isolated bones from the Cenomanian of Sudan (Báez and Werner 1996); unfortunately, these specimens remain undescribed. Stratigraphically, the next possible ranoid occurs in the Coniacian- Santonian of In Beceten, Niger (de Broin et al. 1974, Rage 1984 b, in progress). Ranoids may also be present in the Paleocene of Cernay, France (Estes et al. 1967, Rage 1984 b). Ranoids become more frequent, although still not numerous, in the Eocene and they are present in the Phosphorites. Two ranoid groups from the Phosphorites are considered separately below: the genus Thaumastosaurus and unidentified taxa. Family indeterminate	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFECC7537FDFC5E2FEBFFE2E.taxon	description	In the 1970 s, new excavations in the Phosphorites yielded several specimens belonging to Thaumastosaurus. Surprisingly, as with de Stefano’s lost specimen, they were all represented by the posterior part of a braincase plus otic capsules and fused posterior parts of the frontoparietals. Fossilization of this part of the skull of Thaumastosaurus, which is marked by hyperossification, clearly is favored. Based on this new material, Crochet et al. (1981: tab. 2 - 1) reported the taxon (as Enigmatosaurus bottii) from six (plus perhaps another two) localities of late Eocene age in the Phosphorites. The species then was regarded as a possible Leptodactylidae, an assignment that was not discussed in the article. The tentative referral to the ‘ leptodactylids’, an assemblage now regarded paraphyletic, was briefly discussed by Rage (1981). The assignment was based on the similarity of the skull with that of ‘ ceratophryines’ (then included in the ‘ leptodactylids’) and on the presence, in the same localities as the skull bones, of fragments of bony plates bearing a pustular ornamentation similar to that of the dorsal shield of ‘ ceratophryines’. Thaumastosaurus was therefore regarded as an anuran with South American affinities (‘ ceratophryines’ being restricted to South America). Thereafter, such biogeographical affinities were supported by Roček and Lamaud (1995), Rage and Roček (2007), Evans et al. (2008, 2014) and Agnolin (2012). Roček and Lamaud (1995) provided the first detailed description of Thaumastosaurus bottii, based on various skull bones from La Bouffie, a late Eocene (MP 17) locality in the Phosphorites. They also showed that the one-letter difference between Thaumastosaurus DE STEFANO, 1903 and Thaumatosaurus VON MEYER, 1841 (or the misspelled Thaumattosaurus) prevents homonymy, and that the name Enigmatosaurus NOPCSA, 1908 is a junior synonym of Thaumastosaurus DE STEFANO, 1903. By the early 2000 s, the skull MNHN. F. QU 17736, formerly described by Piveteau (1927) and temporarily lost, was found in the collections. Rage and Roček (2007) described it, showed that it belonged to Thaumastosaurus and demonstrated that it represents a second species, T. gezei. At that time, the latter species was known only by two specimens from the old collections: the skull (i. e. the holotype) and a squamosal. Laloy et al. (2013), using tomography, studied the so-called ‘ mummy’ of Rana plicata FILHOL, 1876 (i. e., Rana cadurcorum MARTÍN, ALONSO- ZARAZAGA et SANCHIZ, 2012). They also scanned the ‘ mummy’ of a forelimb, which might have broken off the main ‘ mummy’ according to Filhol (1877). It was not possible to confirm whether the forelimb belongs to the ‘ mummy’. The tomographic study revealed that the skull of the ‘ mummy’ is identical to that of Thaumastosaurus gezei and, on that basis, the ‘ mummy’ was referred to the latter species. Thanks to a large part of the post-cranial skeleton being preserved in the ‘ mummy’, much of that region is now reliably known for T. gezei (e. g., Text-fig. 4: 1, 2). A recent phylogenetic analysis including post-cranial characters demonstrated that Thaumastosaurus does not belong to a South American clade, but that it belongs instead to the ranoid assemblage; more specifically, it appears to be related to pyxicephalids, an endemic African group (Laloy et al. 2013). This is a good example of how an over-reliance on cranial features related to hyperossification, which can be convergent among unrelated groups of anurans, may adversely affect phylogenetic analyses (Báez and Gómez 2014, Evans et al. 2014). New knowledge about part of the post-cranial skeleton now can be used to identify isolated post-cranial bones in the material collected during recent excavations. Unfortunately, post-cranial bones are known only in T. gezei, thus differences with post-cranial elements of T. bottii remain unknown. Therefore, in this paper, all disarticulated post-cranial bones from the Phosphorites similar to those of T. gezei are referred to as Thaumastosaurus sp. Such is the case, for instance, of scapulae (characterized by a marked dorsoventral elongation; Text-fig. 4: 3) and one humerus (characterized by an articular ball that rather weakly protrudes and is slightly shifted laterally, and by a moderately developed lateral epicondyle; Text-fig. 4: 4). This is also the case for various cranial bones that cannot be allocated at species level (Rage and Roček 2007). It is worth noting that, although a part of the ilium is preserved in the ‘ mummy’ of T. gezei, it is not possible to rank this bone among the post-cranial elements of Thaumastosaurus whose morphology is known. Ilia are among the most frequently recovered anuran bones and among the most useful ones for purposes of identification (Roček et al. 2013, Gómez and Turazzini 2016). Unfortunately, in T. gezei only the anterior extremity of one ilium is known, and it shows only that the shaft bears a medially inclined dorsal crest (Laloy et al. 2013). Unfortunately, this feature is not sufficient for identification within ranoids; consequently, no isolated ilium from the Phosphorites or elsewhere may be confidently assigned to Thaumastosaurus. For the time being, in the Phosphorites we must distinguish between Thaumastosaurus bottii (the type species), T. gezei and Thaumastosaurus sp. Outside of the Phosphorites, two species were assigned to Thaumastosaurus: T. wardi and T. sulcatus, described by Holman and Harrison (2002) and Holman and Harrison (2003), respectively. Both non-Phosphorites species were recovered from the late Eocene (MP 17) of southern England.	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFEEC7537CDCC148FD46FA81.taxon	description	(Text-fig. 4: 1, 2)	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
039987DFFFEEC7527D57C7A6FD94F9A2.taxon	description	(Text-fig. 4: 3, 4) Among specimens referable to Thaumastosaurus sp. are a humerus, a 8 th presacral and a sacral vertebrae that were all regarded as belonging to a large Ranidae RAFINESQUE, 1814 by Rage (1984 b: fig. 2 A, C, D). During the recent excavations, specimens referable to Thaumastosaurus sp. were recovered in localities ranging from MP 16 (localities of Le Bretou and Lavergne; late middle Eocene) to MP 19, perhaps MP 20 (latest Eocene); the youngest specimen is a fragment of maxilla from Tabarly (MP 20) that is tentatively assigned to the genus. Indeterminate ranoids (Text-fig. 4: 5 – 9) Indeterminate ranoids from the Phosphorites include part of the ‘ Ranidae’ reported by Rage (1984 b). These ‘ Ranidae’ are all represented by isolated bones. At that time, the concept of Ranidae was broader than it is today. The available bones closely resemble those of Recent European ranids and, therefore, they were assigned to the Ranidae (Rage 1984 b). Sanchiz (1998) even suggested they probably represent the genus Rana LINNAEUS, 1758 (Pelophylax FITZINGER, 1843 included in Sanchiz’s concept of a broader Rana). However, it does not seem possible to demonstrate that these remains all belong to the Ranidae sensu stricto, as that family is now defined by Frost et al. (2006). In addition, as stated above, some bones (but not all) allocated to the ‘ large form’ by Rage (1984 b) actually belong to Thaumastosaurus sp. In addition to Thaumastosaurus, at least three taxa of ranoids are present in the Phosphorites. This number of taxa is based on humeri. Indeed, three clearly distinct sizes are recognized. These specimens may be regarded as humeri of adult individuals (Text-fig. 4: 5 – 7), on the basis that for each morph the articular ball is entirely preserved and well shaped. The small-sized ranoid (Text-fig. 4: 5) apparently occurs only in Eocene localities, from MP 16 (Lavergne) to MP 18 (Sainte-Néboule). In addition to humeri, at least one ilium, one scapula and some vertebrae may be referred to this small ranoid (Rage 1984 b). The size and the morphology of the bones are homogenous, which suggests that this small form likely represents a single taxon. The mid-sized ranoid is present in the Eocene (Textfig. 4: 6) and Oligocene. Bones are rare and scattered in various localities. It is not possible to determine whether only one taxon is represented. The size of the large form is similar to that of Thaumastosaurus. Humeri are distinguished from those of Thaumastosaurus by their more projecting articular ball, more developed lateral epicondyle and less slender diaphysis (Text-fig. 4: 7). A coracoid (Text-fig. 4: 9) from Escamps (MP 19) may be referred to this large taxon. It differs from that of Thaumastosaurus (Text-fig. 4: 2) in having a neck (corpus coracoidis, Špinar 1972) with more parallel borders and a more flaring pars epicoracoidalis. This large ranoid is reliably known from MP 16 to MP 19. It should be noted that Rage (1984 b: fig. 2) reported a large ranoid from Lavergne (MP 16). However, as stated above, it is now possible to refer the humerus and vertebrae of this ranoid to Thaumastosaurus. The ilium of Thaumastosaurus being unknown, the ilium illustrated by Rage (1984 b: fig. 2 B) cannot be assigned within ranoids. Among ranoid bones from the Phosphorites figured by Rage (1984 b), only the coracoid appears to belong to the large, indetermined ranoid. As far as ranoids from the Eocene of the Phosphorites are concerned, the case of Rhacophoridae must be briefly addressed. This family was reported by Sanchiz (1998) from the Eocene of Escamps (MP 19). However, Sanchiz did not provide descriptions and he did not indicate on what bone (s) he based this identification. In view of the difficulty of identifying taxa within ranoids on the basis of isolated bones, neither the presence nor absence of rhacophorids may be confirmed. Today, the family occurs in sub-Saharan Africa and in southern and southeastern Asia. Fossils are known only from the Quaternary of Japan and perhaps from the Pliocene of eastern Europe (Sanchiz 1998). Ranoids from the Oligocene of the Phosphorites are less frequent than in the Eocene. The few available bones do not permit reliable comparisons with specimens from the Eocene. Ranoid bones are unknown at Phosphorites from the basal Oligocene (MP 21) and apparently from MP 24 to MP 27. A report from the late Oligocene of Pech-du-Fraysse (MP 28; Crochet 1972) is not confirmed. This scarcity of material likely reflects increasing aridity during the Oligocene. Problematic taxa	en	Rage, Jean-Claude (2016): Frogs (Amphibia, Anura) From The Eocene And Oligocene Of The Phosphorites Du Quercy (France). An Overview. Fossil Imprint 72 (1 - 2): 53-66, DOI: 10.14446/FI.2016.53
