Anura

DESCRIPTION Maxillae Three maxilla morphotypes distinct from the unnamed neobatrachian can be identified within In Becetèn: maxillary morphotypes A, B and C. The two fragments attributed to the maxillary morphotype A (MNHN. F. IBC 1989 a, IBC 1989 b) bear ornamentation made of pits and ridges on their labial surface (Fig. 3 A, 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. 3 B). The crista dentalis is shallow. One fragment (MNHN. F. IBC 1989 a) bears a distinct, rounded lamina horizontalis lingually (Fig. 3 C). On the second fragment (MNHN. F. IBC 1989 b), there is no lamina horizontalis (Fig. 3 D). Morphotype A can be differentiated from the above unnamed neobatrachian in being ornamented by pits and ridges (instead of small tubercles). Four fragments are assigned to the maxillary morphotype B (MNHN. F. IBC 1991 a, IBC 1991 b, IBC 1991 c, IBC 1991 d). Three of them lack ornamentation on their labial surface (Fig. 3 E, F). The only exception is the largest fragment (MNHN. F. IBC 1991 a), where a faint patch of rugose ornamentation is present near the base of the frontal process (Fig. 3 E). All fragments bear a crista dentalis, with poorly-preserved teeth (Fig. 3 C, D, G, I). In lingual view, the rounded, prominent lamina horizontalis is distinct on all fragments (Fig. 3 G). The recessus vaginiformis is shallow, but well delimited ventrally and anterodorsally by two crests (Fig. 3 G). 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. Two fragments are assigned to the maxillary morphotype C (MNHN. F. IBC 1991 e, IBC 2063). Their labial surface is covered in small pits and ornamentation imparting rugosity (Fig. 3 H). This ornamentation seems to extend onto the whole labial surface. The maxilla is toothed (Fig. 3 I). The zygomaticomaxillaris process projects posterodorsally (Fig. 3 H, I). The orbital margin strongly decreases in height posterior to the latter process (Fig. 3 H). Lingually, the lamina horizontalis is indistinct from the lingual surface of the maxilla (Fig. 3 I), 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.

DISCUSSION AND ATTRIBUTION 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 & Báez 2001; Báez & Púgener 2003). However, the ornamentation present in morphotypes A and C differs from that of Pachycentrata Báez & Rage, 2004 and the unnamed pipimorph 2 (Báez & Rage 1998; Lemierre et al. 2025), while Inbecetenanura Lemierre, Bailon, Folie & 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. Thus, based on maxilla elements, between three and four non-pipid anuran taxa are present in In Becetèn.

REFERRED MATERIAL. — Nine fragments of maxillae (MNHN. F. IBC 1989 a, IBC 1989 b, IBC 1991 a-IBC 1991 f, IBC 2063).

REMARQUES Les Anura sont représentés par deux humérus gauches de Pelophylax sp. (Fig. 18). Les deux humérus sont attribués au genre Pelophylax Fitzinger, 1843 du fait de la combinaison d’une absence de crête paraventrale, d’une diaphyse droite et robuste avec le condyle situé dans le prolongement de l’axe diaphysaire et d’une crête mésiale courte et transverse (incurvée chez Rana Linnaeus, 1758; Bailon 1999). De plus la présence d’une crête mésiale uniquement chez MMSL- 00.002.0218 / 3 / 1 / A indique que c’est un humérus d’un individu mâle (MMSL- 00.002.0218 / 3 / 1 / B représentant un individu femelle).

Remarks. Numerous fossil elements can be assigned to anurans, but are either too poorly preserved or not taxonomically significant enough for a more refined identification.

Material. Hambach 6 C: four premaxillae (IPB-HaH 2268 / 2270, IPB-HaH 2299); 51 fragments of maxilla (IPB-HaH 2233 / 2265, IPB-HaH 2282 / 2298, IPB-HaH 2300); 17 trunk vertebrae (IPB-HaH 2089 / 2097, IPB-HaH 2144 / 2145, IPB-HaH 2157 / 2158, IPB-HaH 2181 / 2183, IPB-HaH 2192); seven sacral vertebrae (IPB-HaH 2087, IPB-HaH 2206 / 2208, IPB-HaH 2217 / 2218, IPB-HaH 2225); 12 urostyles (IPB-HaH 2088, IPB-HaH 2197 / 2198, IPB-HaH 2201 / 2205, IPB-HaH 2216, IPB-HaH 2226 / 2227, IPB-HaH 2306); three coracoids (IPB-HaH 2313 / 2314, IPB-HaH 2378); nine humeri (IPB-HaH 2063, IPB-HaH 2303 / 2304, IPB-HaH 2311 / 2312, IPB-HaH 2317 / 2318, IPB-HaH 2331); five radioulnae (IPB-HaH 2301 / 2302, IPB-HaH 2309, IPB-HaH 2330, IPB-HaH 2335); two ilia (IPB-HaH 2319 / 2320); one ischium (IPB-HaH 2322); one femur (IPB-HaH 2326); 14 tibiofibulae (IPB-HaH 2054, IPB-HaH 2062, IPB-HaH 2064 / 2068, IPB-HaH 2305, IPB-HaH 2310, IPB-HaH 2327, IPB-HaH 2332 / 2334, IPB-HaH 2400); four indeterminate elements (IPB-HaH 2308, IPB-HaH 2315, IPB-HaH 2328, IPB-HaH 2398). Hambach 11: 29 maxillae (IPB-HaR 2044 / 2070, IPB-HaR 2177 / 2178); one angular (IPB-HaR 2114); two atlases (IPB-HaR 2025, IPB-HaR 2035); five trunk vertebrae (IPB-HaR 2024; IPB-HaR 2027 / 2029; IPB-HaR 2037); two sacral vertebrae (IPB-HaR 2026, IPB-HaR 2036); three urostyles (IPB-HaR 2038 / 2040); two coracoids (IPB-HaR 2125 / 2126); two scapulae (IPB-HaR 2072 / 2073); 10 humeri (IPB-HaR 2089 / 2095, IPB-HaR 2108 / 2109, IPB-HaR 2153); 16 radioulnae (IPB-HaR 2074 / 2079, IPB-HaR 2107, IPB-HaR 2123 / 2124, IPB-HaR 2128, IPB-HaR 2155 / 2157, IPB-HaR 2180 / 2182); four ilia (IPB-HaR 2085 / 2086, IPB-HaR 2104 / 2105); one ischium (IPB-HaR 2106); three femurs (IPB-HaR 2133 / 2134, IPB-HaR 2141); 25 tibiofibulae (IPB-HaR 2080 / 2082, IPB-HaR 2118 / 2122, IPB-HaR 2135 / 2140, IPB-HaR 2158 / 2168); one indeterminate element (IPB-HaR 2169). Hambach 11 C: three maxillae (IPB-HaR 2422 / 2424); one ornamented bone fragment (IPB-HaR 2430); four tibiofibulae (IPB-HaR 2410 / 2413); one indeterminate element (IPB-HaR 2409).

DESCRIPTION Despite a swampy paleoenvironment that might be suitable for their occurrence and preservation, anurans are relatively scarce among the microvertebrate material from Angeac-Charente. In Angeac-Charente, anurans are represented by isolated, fragmentary bones, such as fused zeugopods (Fig. 10 A, B) and urostyles (Fig. 10 C, D) Among fused zeugopods, some are sufficiently well preserved to allow their identification. For instance, the general shape and length (even if the proximal part is missing) of specimen ANG M- 63 (Fig. 10 A, B), as well as several other features, allow its identification as a tibiofibula (Thomas 1996): presence of a tibiofibular groove, deeper on the anterior face, between the fused zeugopod, and presence of a tibial crest on the anterior face (Fig. 10 A). Among well-known Early Cretaceous anuran families are the Alytidae (= “ Discoglossidae ”), but within this family, the tibia and fibula remain unfused (Roček 2000), unlike the Angeac-Charente specimen. The specimen ANG M- 64 (Fig. 10 C, D) bears two broad and flattened articular facets anteriorly, and a small crest arises from its dorsal face. Four weakly marked ridges can be seen alongside this small crest, two on each side (Fig. 9 C). This morphology identifies it as a urostyle. To date, incompleteness of the material limits comparisons with other Mesozoic anurans.

REMARQUES Les tarses et les métapodes sont généralement des éléments peu discriminants, d’autant plus que le matériel de Ben Kérat est mal préservé, et ne peut pas être attribué à un genre d’anoure en particulier.

MATÉRIEL. — NR = 2; deux fragments de tarses ou de métapodes?

DESCRIPTION AND COMMENTS The fragment of maxilla bears teeth typical of most of the anurans (thus, among the current Asian anurans, Bufonidae members are edentulous), but it does not provide precise information. The sacral vertebra shows two posterior condyles (Figs 1 E; 2 F). The anterior face of the centrum is concave therefore this vertebra cannot be referred to the Alytidae or “ Ranidae ”. Its sacral expansions are broken off, but their bases demonstrate that they were not widely extended anteroposteriorly. It is not possible to determine whether they were cylindrical or moderately extended. Therefore, an assignment at a family level cannot be suggested. Similarly, it is not possible to state whether this vertebra belongs to the Anura indeterminata A from Sherullah 9 that is represented by procoelous sacral vertebrae.

LOCALITY AND AGE. — Hadji Rona, Sarobi basin, Afghanistan, late early Pliocene, late Ruscinian, MN 15.

MATERIAL EXAMINED. — One fragment of maxilla (AFG 1679), 1 incomplete sacral vertebra (AFG 1680).

LOCALITY AND AGE. — Pul-E Charkhi, Kabul basin, Afghanistan, earliest Pliocene, early Ruscinian, MN 13 / 14.

COMMENTS Generally, tibio-fibula provide poorly information for identification within anurans. The fragment from Pul-E Charkhi only shows that a frog was present. In any case, it forms the first evidence of frog occurence in this locality during the early Pliocene.

MATERIAL EXAMINED. — One fragment of tibio-fibula (AFG 1674).

DESCRIPTION AND COMMENTS Only the maxillae, vertebrae and ilia deserve comments. The other remains provide no information. The presence of teeth on the maxillae allows to rule out Bufonidae, where these bones are toothless. The vertebrae are only represented by their centra. Five centra of presacral vertebrae are amphicoelous, deeply biconcave, whereas seven are non-amphicoelous (it is not possible to determine their condition, procoelous or opisthocoelous). Vertebrae of Alytidae are opisthocoelous. In “ Ranidae ”, the last presacral vertebra is amphicoelous whereas the seven others are procoelous. Consequently, within the set of presacral vertebrae from Sherullah 9, the ratio amphicoelous / non-amphicoelous vertebrae is somewhat surprising. In a few anuran groups, all presacral vertebrae are amphicoelous: in the living Leiopelmatidae Mivart, 1869 (New Zealand) and Ascaphidae Fejérváry, 1923 (Western North America) the centra are clearly amphicoelous; in the Megophryidae Bonaparte, 1850 (southern Asia) and various Myobatrachidae Schlegel, 1850 (Australia) the intervertebral cartilages remain free in adults, therefore the vertebrae are amphicoelous but they are not deeply biconcave. Among extinct forms, Notobatrachus Reig, 1956 and likely Vieraella Reig, 1961 (both from the Jurassic of South America), as well as the Gobiatidae Roček & Nessov 1993 (Cretaceous of Central Asia) are amphicoelous (Báez & Basso 1996; Roček & Nessov 1993). But, comparisons between “ amphicoelous taxa ” and amphicoelous vertebrae from Sherullah 9 cannot be made because the latter specimens are known only by their centra. Based on the available specimens from Sherullah 9 a family assignment is not possible.

LOCALITY AND AGE. — Sherullah 9, Khordkabul basin, Afghanistan, late Miocene, late Vallesian-basal Turolian transition, MN 10 / 11.

Sacral vertebrae, as presacral ones, are represented by centra. The posterior face of all centra is bicondylar. But, in five sacral vertebrae the anterior face is convex (opisthocoelous) while it is concave (procoelous) in two specimens. The opisthocoelous sacral vertebrae may belong to Alytidae or “ Ranidae ”. But procoelous sacral vertebrae represent another family that cannot be identified. All ilia have a dorsal crest. This morphological feature is present in Discoglossinae (Alytidae), Pipidae Gray, 1825, “ Ranidae ”, Rhacophoridae and various Leptodactylidae Werner, 1896 and Hylidae Rafinesque, 1815. The morphology of the tuber superius and dorsal crest of the fossil leads to rule out Pipidae, Rhacophoridae, Leptodactylidae and Hylidae, but the poor preservation of these bones does not permit to refer them to either the Discoglossinae (Alytidae) or “ Ranidae ”.

MATERIAL EXAMINED. — Five fragments of toothed maxillae (AFG 1656), 1 fragment of angulosplenial (AFG 1657), 1 fragment of atlas (AFG 1658), 5 amphicoelous presacral vertebrae (AFG 1659), 7 non-amphicoelous presacral vertebrae (AFG 1660), 5 opisthocoelous sacral vertebrae (AFG 1661), 2 procoelous sacral vertebrae (AFG 1662), 5 fragments of urostyles (AFG 1663), 34 ilia (AFG 1664), 3 fragmentary humeri (AFG 1665), 3 fragments of radioulna (AFG 1666), 1 fragment of femur (AFG 1667).

Hylidae Dendropsophus nanus (Boulenger) Acanthocephala fam. gen. sp. Scinax nasicus (Cope) Centrorhynchus sp. Leptodactylidae Adenomera diptyx (Boettger) Centrorhynchus sp. Leptodactylus bufonius Boulenger Centrorhynchus sp. Leptodactylus chaquensis Cei Centrorhynchus sp. Leptodactylus latinasus Jiménez de la Espada Centrorhynchus sp. Pseudopaludicola boliviana Parker Centrorhynchus sp.

A well-preserved, procoelous, anuran vertebra, AMNH 8424, has a dorsoventrally compressed centrum (fig. 11 O, P). The neural arch is incomplete and tapers dorsally, and the transverse processes are short and robust. The length of the vertebra, the only one of its kind in the collection, is 1.6 mm.

Gonçalves et al. (2009) established high levels of genetic divergence within the Iberian Midwife Toad (Alytes cisternasii), but considered these to be within the range of typical intraspecific variation in amphibians. Carretero et al. (2009) disagreed with Zangari et al. ’ s (2006) decision to treat the Eastern Iberian Painted Frog as a subspecies, Discoglossus galganoi jeanneae. They found additional support for the species rank of these two taxa in Velo-Antón et al. (2008) and stated that, given the lack of more detailed studies allowing assessment of gene flow between both taxa in secondary contact areas, there is no reason to treat them as conspecific. This is in conflict with our fundamental appraisal that splitting a species can only be valid if the split is substantiated by scientific evidence, rather than considering taxa as species because of lack of reason to treat them as conspecific. As long as this is not the case, we promote conspecificity to be the rule. In fact, Velo-Antón et al. ’ s (2008) results are in agreement with Zangari et al. ’ s (2006) work. The fact that both studies found the same low level of nuclear differentiation with independent markers certainly calls for a reassessment of the validity of the specific status of jeanneae and reinforces our reluctance to treat is as a valid species. The comprehensive work of Frost et al. (2006) on amphibian systematics has provoked contrasting responses, including (among quite some others) a rather strong critique by Wiens (2007), which saw a subsequent rebuttal by Frost et al. (2008). Indirectly, Pauly et al. (2009) also criticised Frost et al. (2006), and also received a response (Frost et al. 2009). Overall, Speybroeck and Crochet’s (2007) treatment of the proposed changes seems to have been largely in correspondence to what other authors have concluded. An exception deserves, however, our renewed attention. While Speybroeck and Crochet (2007) proposed to attribute the European ‘ true toad’ species to 2 genera (Bufo and Epidalea), general consent in this case seems to be towards conserving Bufo as the genus for all, at least for the time being (Vences 2007; Bour et al. 2008; Lescure 2008). These authors argue that cases of natural hybridisation (e. g. a very recent record of hybridisation Bufo bufo x viridis by Duda 2008) should encourage rejection of a genus level split, as proposed by Dubois (1988) and applied to the case of Bufo for the first time by Dubois and Dinesh (2007). Concerning European species, Van Bocxlaer et al. (2009) provide some support for the generic arrangement proposed by Frost et al. (2006), which might very well make attribution of Green Toad (Bufo viridis (s. l. - see below )) to the genus Pseudepidalea and the Natterjack (Bufo calamita) to Epidalea a valid arrangement. Yet, with different relationships turning up from different studies (cf. also Pramuk et al. 2008) and many taxa still in need of investigation, it seems cautious not to draw any taxonomical conclusions just yet. Pending additional research, we therefore place all European species back in the single genus Bufo. Additionally, we note that according to Dubois and Bour (in press), the use of the name Pseudepidalea should be abandoned for that of the junior synonym Bufotes Rafinesque, 1815, while the name Epidalea Cope, 1864 remains available. We have previously been reluctant to accept Stöck et al. ’ s (2006) Bufo viridis (Green Toad) splits (Speybroeck and Crochet 2007). Stöck et al. (2008 a) described yet another new species from Sicily, Bufo siculus (Sicilian Green Toad). Despite Carretero et al. ’ s (2009) adoption of these new species, we still believe that mtDNA lineages alone cannot be used to substantiate new species, and that the level of divergence of the taxa, which are also supported by other characters (e. g. siculus which is also supported by nuclear and morphological data, albeit without comparing the taxon morphologically with its closest African relatives), is not high enough to be in itself evidence of specific status. To a certain degree at least, this seems to be corroborated by Van Bocxlaer et al. (2009): divergence between the Green Toad splits viridis and ‘ cf. variabilis ’ appears to be smaller than between the Common Toad (sub) species bufo and spinosus. We also treat the latter two taxa as conspecific. While we do not claim that the green toads of the Western Palearctic definitely belong to a single species, we maintain that the available information cannot (yet) support any species level split. As noted by Razzetti (2008), the correct name for the green toads of peninsular Italy, Corsica, Sardinia and northeastern Sicily is still controversial. Balletto et al. (2007), based on specimens from Venice, used Bufo lineatus Ninni, 1879 (type locality: surroundings of Venice - Frost, 2009) as the valid nomen for the clade of peninsular Italy, while Stöck et al. (2006, 2008 a) considered Bufo lineatus as a junior synonym of Bufo viridis, because they found specimens from Padua and Trieste that belong to the nominotypical lineage. Stöck et al. (2008 b) studied the phylogeography of the genus Hyla (tree frogs) around the Mediterranean. They identified three deeply divergent mitochondrial lineages in populations currently classified as Hyla arborea, each of them being supported by variation in one nuclear intron. In their mitochondrial tree (but not in their nuclear tree), treating H. sarda and H. intermedia as valid species could render H. arborea paraphyletic, because the three mitochondrial lineages identified within arborea are not necessarily each other's closest relatives. Since the specific status of intermedia is well supported by reproductive isolation in contact zones (Verardi et al. 2009), and since sarda displays distinct and well-known morphological and acoustic characters (Schneider 1974; Lanza 1983; Rosso et al. 2001, 2004; Castellano et al. 2002), we maintain them as valid species. As a consequence, the mitochondrial data provide strong evidence for recognising the Iberian taxon molleri and the eastern taxon orientalis (currently only known in Europe from the Black Sea Coast of Romania and European Turkey) as valid species as well. Nevertheless, the distributional limits of these two taxa remain unknown. There is no evidence of reproductive isolation in the continuous range of tree frogs in the Balkans, no known obvious morphological characters to separate them, and no obvious acoustic difference between molleri and arborea, nor orientalis and arborea (Schneider 1974, 2002). Accepting these two new European species would thus rest entirely on mtDNA data from a very small number of specimens (seven orientalis and only two molleri). Therefore, while a species level split is likely to be required, we prefer to wait for additional data, as specified, before recognising molleri and / or orientalis as valid species. Detailed study by Gvoždík et al. (2008) uncovered a complex pattern of geographical variation in morphology among populations of Hyla arborea and Hyla savignyi. The similarity among populations is not necessarily greater within species that between species. On the contrary, populations of different species inhabiting neighbouring regions are often more similar than populations of the same species inhabiting distant regions. Groups of populations defined by morphology do not correspond to the mitochondrial lineages defined by Stöck et al. (2008) either. In fact, Gvoždík et al. (2008) suggest that morphological variation of Hyla is more linked to climate variation than to evolutionary history. Stöck et al. ’ s (2008 b) unnamed clade of Hyla cf. intermedia from Switzerland corresponds with the northern clade of Canestrelli et al. (2007 a). Allozyme divergence between this northern clade and the southcentral clade of H. intermedia s. l. is typical of intraspecific level of divergence: Nei’s distance value of 0.07 according to Canestrelli et al. (2007 b), to be compared with Nei’s distance of 0.55 between arborea and intermedia (Verardi et al. 2009). Thus, in our opinion, the various clades within Hyla intermedia s. l. might well prove to constitute valid subspecies, but are unlikely to represent distinct species. In any case, we strongly advocate detailed analyses of contact zones prior to any formal proposal. A detailed phylogeographic analysis of the Pool Frog in Italy (Canestrelli and Nascetti 2008) supported the subspecific status of Pelophylax lessonae bergeri suggested by Crochet and Dubois (2004) and followed by Speybroeck and Crochet (2007). The same study confirmed that Sicilian pool frogs should also be recognised as a distinct subspecies (see also Santucci et al. 1996). Lymberakis et al. (2007) investigated Eastern Mediterranean water frog phylogeny by means of mitochondrial DNA. Their results reinforce the idea that Pelophylax kurtmuelleri (Greek Marsh Frog) should be treated as conspecific with central European populations of the P. ridibundus complex (Marsh Frog), as previously established with allozyme data (Beerli 1994). The precise status of these populations should be investigated in a range-wide analysis of the P. ridibundus complex. As long as mating call differences are the only support for specific treatment (Schneider et al. 1993), we suggest to no longer recognise kurtmuelleri as a valid species. Alleged contact zones between ridibundus and kurtmuelleri in Thrace (Schneider et al. 1993) seem to be in fact contact zones with Pelophylax bedriagae (Bedriaga’s Water Frog) rather than ridibundus, as Beerli (1994) identified Thracian water frogs unambiguously as Pelophylax bedriagae. This seems to have been confirmed by Lymberakis et al. ’ s (2007) results, which included a sample from Thrace (Dadia) attributed to P. bedriagae and closely related to Lesbos and Chios populations. However, their results also attributed a sample from a very nearby location, as well as other Thracian samples, to Pelophylax ridibundus. All these results suggest that bedriagae and “ European ridibundus ” form a contact zone in Thrace, where these two taxa are reproductively isolated (Schneider et al. 1993). This indicates that the two subclades B 5 and B 6 of Lymberakis et al. (2007) are valid biological species and thus supports the widely accepted species status of P. bedriagae and P. ridibundus (sensu lato). However, Lymberakis et al. (2007) did not include samples from the type locality of ridibundus (northern Caspian Sea area), so it remains to be determined if European populations of Marsh Frogs are conspecific with sensu stricto P. ridibundus or not. If not, the names Pelophylax ranaeformis (Laurenti, 1768) and Pelophylax fortis (Boulenger, 1884) might apply to the European Marsh Frog (Dubois & Ohler 1995 a, b). The former name relates (at least) to the populations of the Greek island Limnos (Dubois & Ohler 1995 b). Lymberakis et al. (2007) found Pelophylax cerigensis (Karpathos Water Frog) to be nested within their subclade B 5, corresponding to P. bedriagae. We note that these authors also attributed Rhodes populations to P. cerigensis, whereas the original description only considered this to be a possibility (Beerli et al. 1994). Indeed, based on biochemical data, Plötner (2005) placed Rhodes and Karpathos water frogs together, different from both bedriagae and ridibundus, and attributed populations from “ Karpathos and probably Rhodes ” to cerigensis. However, to our knowledge, no subsequent papers have provided definite evidence ascertaining the specific status of Rhodes water frogs. The results of Lymberakis et al. (2007) invalidate a P. cerigensis limited to Karpathos and Rhodes. Apart from the authors’ suggestion that P. c e r i g e n s i s could be treated as a junior synonym for P. bedriagae, alternative arrangements seem compatible with the available evidence: to restrict the name P. bedriagae to (at least some of) the more eastern populations (Syria and some surrounding areas, also Cyprus), whereas populations from Turkey, the eastern Aegean islands, including Karpathos and Rhodes could be attributed to P. caralitanus (Arıkan 1988), for which cerigensis (Beerli, Hotz, Tunner, Heppich and Uzzell 1994) would be a junior synonym. A second alternative could include splitting of the latter group, with the Karpathos populations being attributed to P. cerigensis and treating the Turkish, eastern Aegean and Rhodes populations as a different species. Under either of these alternative hypotheses, several other species would need to be recognised for Anatolian and Middle Eastern water frog populations. Our second alternative might result in retaining the validity of P. cerigensis (for Karpathos populations only). However, for the time being, we believe material from geographically intermediate populations is required to warrant these alternative arrangements, and therefore preliminarily consider P. cerigensis to represent a part of P. bedriagae rather than a separate species. The most recent available results about the contact zones between the western and central Anatolian lineages of water frogs (Akın et al. 2010) support the view that at least part of the genetic diversity within the bedriagae complex represents intraspecific variation. We thus suggest to recognise, for the time being, a single species of Middle East water frog, whose name should be either bedriagae or ranaeformis, depending on the identity of the water frogs of Limnos. The nomen cerigensis thus currently becomes a synonym of bedriagae at the species rank.

Bologna 1996), actual presence within considered area requires confirmation Podarcis Wagler, 1830 bocagei (Seoane, 1884) — Bocage’s Wall Lizard carbonelli Pérez-Mellado, 1981 — Carbonell’s Wall Lizard cretensis (Wettstein, 1952) — Cretan Wall Lizard erhardii (Bedriaga, 1876) — Erhard’s Wall Lizard filfolensis (Bedriaga, 1876) — Maltese Wall Lizard gaigeae (Werner, 1930) — Skyros Wall Lizard hispanicus (Steindachner, 1870) including s. s. morphotype, morphotype 1, and morphotype 2 — Iberian Wall Lizard

Family Salamandridae Goldfuss, 1820 (true salamanders and newts) Calotriton Gray, 1858

Iberolacerta Arribas, 1997 aranica (Arribas, 1993) — Aran Rock Lizard aurelioi (Arribas, 1994) — Aurelio’s Rock Lizard bonnali (Lantz, 1927) — Pyrenean Rock Lizard cyreni (Müller and Hellmich, 1937) — Cyren’s Rock Lizard galani Arribas, Carranza and Odierna, 2006 — Galan’s Rock Lizard horvathi (Méhely, 1904) — Horvath’s Rock Lizard martinezricai (Arribas, 1996) — Martinez-Rica’s or Peña de Francia Rock Lizard monticola (Boulenger, 1905) — (West-) Iberian Rock Lizard Lacerta Linnaeus, 1758 agilis Linnaeus, 1758 — Sand Lizard bilineata Daudin, 1802 — Western Green Lizard schreiberi Bedriaga, 1878 — Schreiber’s Green Lizard trilineata Bedriaga, 1886 — Balkan Green Lizard viridis (Laurenti, 1768) — Eastern Green Lizard Ophisops Ménétries, 1832 elegans Ménétries, 1832 — Snake-eyed Lacertid

Salamander strinatii (Aellen, 1958) — Strinati’s Cave Salamander supramontis (Lanza, Nascetti and Bullini, 1986) — Sopramonte Cave Salamander Family Proteidae Gray, 1825 (olms) Proteus Laurenti, 1768 anguinus Laurenti, 1768 — Olm Order Anura (frogs and toads) Family Alytidae Fitzinger, 1843 (painted frogs and midwife toads) Alytes Wagler, 1829 cisternasii Boscá, 1879 — Iberian Midwife Toad dickhilleni Arntzen and García-París, 1995 — Southern Midwife Toad muletensis (Sanchíz and Adrover, 1977) — Majorca Midwife Toad obstetricans (Laurenti, 1768) — Common Midwife Toad Discoglossus Otth, 1837 galganoi Capula, Nascetti, Lanza, Bullini and Crespo, 1985 — Iberian Painted Frog montalentii Lanza, Nascetti, Capula and Bullini, 1984 — Corsican Painted Frog pictus Otth, 1837 — Painted Frog sardus Tschudi in: Otth, 1837 — Tyrrhenian Painted Frog Family Bombinatoridae Gray, 1825 (fire-bellied toads) Bombina Oken, 1816 bombina (Linnaeus, 1761) — Fire-bellied Toad variegata (Linnaeus, 1758) — Yellow-bellied Toad Family Pelobatidae Bonaparte, 1850 (spadefoot toads) Pelobates Wagler, 1830 cultripes (Cuvier, 1829) — Western Spadefoot fuscus (Laurenti, 1768) — Common Spadefoot syriacus Boettger, 1889 — Eastern Spadefoot Family Pelodytidae Bonaparte, 1850 (parsley frogs) Pelodytes Bonaparte, 1838 ibericus Sánchez-Herráiz, Barbadillo, Machordom and Sanchiz, 2000 — Iberian Parsley Frog punctatus (Daudin, 1802) — Parsley Frog Family Bufonidae Gray, 1825 (true toads) Bufo Laurenti, 1768 bufo (Linnaeus, 1758) — Common Toad calamita (Laurenti, 1768) — Natterjack viridis (Laurenti, 1768) — Green Toad Family Hylidae Rafinesque, 1815 (tree frogs) Hyla Laurenti, 1768 arborea (Linnaeus, 1758) — Common Tree Frog intermedia Boulenger, 1882 — Italian Tree Frog meridionalis Boettger, 1874 — Stripeless Tree Frog sarda (de Betta, 1857) — Tyrrhenian Tree Frog Family Ranidae Rafinesque-Schmaltz, 1814 (true frogs) Pelophylax Fitzinger, 1843

arguta (Pallas, 1773) — Steppe Runner

jaculus (Linnaeus, 1758) — Sand Boa Family Psammophiidae Boie, 1827 (African sand snakes and Montpellier snakes) Malpolon Fitzinger, 1826

kl. grafi (Crochet, Dubois, Ohler and Tunner, 1995) — Graf’s Hybrid Frog lessonae (Camerano, 1882) — Pool Frog perezi (Seoane, 1885) — Iberian Water Frog ridibundus (Pallas, 1771) — Marsh Frog shqipericus (Hotz, Uzzell, Günther, Tunner and Heppich, 1987) — Albanian Pool Frog Rana Linnaeus, 1758 arvalis Nilsson, 1842 — Moor Frog dalmatina Fitzinger in Bonaparte, 1838 — Agile Frog graeca Boulenger, 1891 — Greek Stream Frog iberica Boulenger, 1879 — Iberian Stream Frog italica Dubois, 1987 — Italian Stream Frog latastei Boulenger, 1879 — Italian Agile Frog pyrenaica Serra — Cobo, 1993 — Pyrenean Stream Frog temporaria Linnaeus, 1758 — Grass Frog Class Reptilia (reptiles) Order Testudines or Chelonii (turtles, tortoises and terrapins) Family Cheloniidae Oppel, 1811 (sea turtles) Caretta Rafinesque-Schmaltz, 1814 caretta (Linnaeus, 1758) — Loggerhead (( Sea) Turtle) Family Dermochelyidae Fitzinger, 1843 (1825) (leatherbacks) Dermochelys de Blainville, 1816 coriacea (Vandelli, 1761) — Leatherback Family Testudinidae Batsch, 1788 (tortoises) Testudo Linnaeus, 1758 graeca Linnaeus, 1758 — Spur-thighed Tortoise hermanni Gmelin, 1789 — Hermann’s Tortoise marginata Schoepff, 1792 — Marginated Tortoise Family Geoemydidae Theobald, 1868 (Old World terrapins) Mauremys Gray, 1869 leprosa (Schweigger, 1812) — Spanish Terrapin rivulata (Valenciennes, 1833) — Balkan Terrapin Family Emydidae Rafinesque, 1815 (New World terrapins) Emys Duméril, 1805 orbicularis (Linnaeus, 1758) — European Pond Terrapin Order Squamata Suborder Sauria (lizards) Family Agamidae Spix, 1825 or Fitzinger, 1826 (agamas) Laudakia Gray, 1845 stellio (Linnaeus, 1758) — Starred Agama Family Chamaeleonidae Gray, 1825 or Rafinesque, 1815 (Chamaeleontidae) (chameleons) Chamaeleo Laurenti, 1768 africanus Laurenti, 1768 — African Chameleon chamaeleon (Linnaeus, 1758) — Mediterranean Chameleon Family Sphaerodactylidae Underwood, 1954 (least geckos) Euleptes Fitzinger, 1843 europaea (Gené, 1839) — European Leaf-toed Gecko Family Gekkonidae Oppel, 1811 or Gray, 1825 (true geckos) Hemidactylus Oken, 1817 turcicus (Linnaeus, 1758) — Turkish Gecko Mediodactylus Szczerbak and Golubev, 1977 kotschyi (Steindachner, 1870) — Kotschy’s Gecko Family Phyllodactylidae Gamble, Bauer, Greenbaum and Jackman, 2008 (leaf-toed geckos) Tarentola Gray, 1825 mauritanica (Linnaeus, 1758) — Moorish Gecko Family Lacertidae Batsch, 1788 (true lizards) Acanthodactylus Wiegmann, 1834 erythrurus (Schinz, 1833) — Spiny-footed Lizard Algyroides Bibron and Bory de Saint-Vincent, 1833 fitzingeri (Wiegmann, 1834) — Tyrrhenian Algyroides marchi Valverde, 1958 — Spanish Algyroides moreoticus Bibron and Bory de Saint-Vincent, 1833 — Greek Algyroides nigropunctatus (Duméril and Bibron, 1839) — Dalmatian Algyroides Archaeolacerta Mertens, 1921

liolepis (Boulenger, 1905) — Catalonian Wall Lizard melisellensis (Braun, 1877) — Dalmatian Wall Lizard milensis (Bedriaga, 1882) — Milos Wall Lizard muralis (Laurenti, 1768) — Common Wall Lizard peloponnesiacus (Bibron and Bory de Saint-Vincent, 1833) — Peloponnese Wall Lizard pityusensis (Boscá, 1883) — Ibiza Wall Lizard raffonei (Mertens, 1952) — Aeolian Wall Lizard siculus (Rafinesque-Schmaltz, 1810) — Italian Wall Lizard tauricus (Pallas, 1814) — Balkan Wall Lizard tiliguerta (Gmelin, 1789) — Tyrrhenian Wall Lizard vaucheri (Boulenger, 1905) — Vaucher’s Wall Lizard waglerianus Gistel, 1868 — Sicilian Wall Lizard Psammodromus Fitzinger, 1826 algirus (Linnaeus, 1758) — Large Psammodromus hispanicus Fitzinger, 1826 — Spanish Psammodromus Scelarcis Fitzinger, 1843 perspicillata (Duméril and Bibron, 1839) — Moroccan Rock Lizard Timon Tschudi, 1836 lepidus (Daudin, 1802) — Ocellated Lizard Zootoca Wagler, 1830 vivipara (Jacquin, 1787) or (Lichtenstein, 1823) — Viviparous Lizard Family Scincidae Oppel, 1811 or Gray, 1825 (skinks) Ablepharus Fitzinger in Eversmann, 1823 kitaibelii Bibron and Bory de Saint-Vincent, 1833 — Snake-eyed Skink Chalcides Laurenti, 1768 bedriagai (Boscá, 1880) — Bedriaga’s Skink chalcides (Linnaeus, 1758) — Italian Three-toed Skink ocellatus (Forskål, 1775) — Ocellated Skink striatus (Cuvier, 1829) — Iberian Three-toed Skink Ophiomorus Duméril and Bibron, 1839 punctatissimus (Bibron and Bory de Saint-Vincent, 1833) — Limbless Skink Family Anguidae Gray, 1825 (slow worms) Anguis Linnaeus, 1758 cephallonica Werner, 1894 — Peloponnese Slow Worm colchica (Nordmann, 1840) — Eastern Slow Worm fragilis Linnaeus, 1758 — Slow Worm graeca Bedriaga, 1881 — Greek Slow Worm Pseudopus Merrem, 1820 apodus (Pallas, 1775) — Glass Lizard Suborder Amphisbaenia (worm lizards)

lusitanica Bocage, 1864 — Golden-striped Salamander Euproctus Gené, 1839 montanus (Savi, 1838) — Corsican Brook Newt platycephalus (Gravenhorst, 1829) — Sardinian Brook Newt Ichthyosaura Sonnini and Latreille, 1801 alpestris (Laurenti, 1768) — Alpine Newt Lissotriton Bell, 1839 boscai (Lataste in Blanchard, 1879) — Bosca’s Newt helveticus (Razoumowsky, 1789) — Palmate Newt italicus (Peracca, 1898) — Italian Newt montandoni (Boulenger, 1880) — Montandon’s Newt vulgaris (Linnaeus, 1758) — Smooth Newt Lyciasalamandra Veith and Steinfartz, 2004 helverseni (Pieper, 1963) — Karpathos Salamander Pleurodeles Michahelles, 1830 waltl Michahelles, 1830 — Sharp — ribbed Newt Salamandra Garsault, 1764 atra (Laurenti, 1768) — Alpine Salamander corsica (Savi, 1838) — Corsican Fire Salamander lanzai (Nascetti, Andreone, Capula and Bullini, 1988) — Lanza’s (Alpine) Salamander salamandra (Linnaeus, 1758) — Fire Salamander Salamandrina Fitzinger, 1826 perspicillata (Savi, 1821) — Northern Spectacled Salamander terdigitata (Bonnaterre, 1789) — Southern Spectacled Salamander Triturus Rafinesque, 1815 arntzeni Litvinchuk, Borkin, Dzukić and Kalezić, 1999 — Arntzen’s Crested Newt carnifex (Laurenti, 1768) — Italian Crested Newt cristatus (Laurenti, 1768) — (Great or Northern) Crested Newt dobrogicus (Kiritzescu, 1903) — Danube Crested Newt? karelinii (Strauch, 1870) — Southern Crested Newt — presence in Europe depends on location of boundary with arntzeni macedonicus (Karaman, 1922) — Macedonian Crested Newt marmoratus (Latreille, 1800) — Marbled Newt pygmaeus (Wolterstorff, 1905) — Southern Marbled Newt Family Plethodontidae Gray, 1850 (lungless salamanders)

tessellata (Laurenti, 1768) — Dice Snake Family Colubridae Oppel, 1811 (colubrids) Coronella Laurenti, 1768 austriaca Laurenti, 1768 — Smooth Snake girondica (Daudin, 1803) — Southern Smooth Snake Dolichophis Gistel, 1868 caspius (Gmelin, 1789) — Caspian Whip Snake Eirenis Jan, 1863? modestus (Martin, 1838) — (Masked) Dwarf Snake — actual presence within the considered area requires confirmation Elaphe Fitzinger, 1833 quatuorlineata (Bonnaterre, 1790) — (Western) Four-lined Snake sauromates (Pallas, 1814) — Blotched Snake or Eastern Four-lined Snake Hemorrhois Boie, 1826 algirus (Jan, 1863) — Algerian Whip Snake hippocrepis (Linnaeus, 1758) — Horseshoe Whip Snake? nummifer (Reuss, 1834) — Coin-marked Snake — actual presence within considered area requires confirmation Hierophis Fitzinger in Bonaparte, 1834 gemonensis (Laurenti, 1768) — Balkan Whip Snake viridiflavus (Lacépède, 1789) — Western Whip Snake Macroprotodon Guichenot, 1850 brevis (Günther, 1862) — Western or Iberian False Smooth Snake cucullatus (Geoffroy Saint-Hilaire, 1809) — Eastern or African False Smooth Snake Platyceps Blyth, 1860 collaris (Müller, 1878) — Reddish Whip Snake najadum (Eichwald, 1831) — Dahl’s Whip Snake Rhinechis Michahelles in Wagler, 1833 scalaris (Schinz, 1822) — Ladder Snake Telescopus Wagler, 1830 fallax (Fleischmann, 1831) — Cat Snake Zamenis Wagler, 1830 lineatus (Camerano, 1891) — Italian Aesculapian Snake longissimus (Laurenti, 1768) — Aesculapian Snake situla (Linnaeus, 1758) — Leopard Snake Family Viperidae Oppel, 1811 (true vipers) Macrovipera Reuss, 1927 schweizeri (Werner, 1935) — Milos Viper Montivipera Nilson, Tuniyev, Andrén, Orlov, Joger and Herrmann, 1999 xanthina (Gray, 1849) — Ottoman Viper Vipera Garsault, 1764 ammodytes (Linnaeus, 1758) — Nose-horned Viper aspis (Linnaeus, 1758) — Asp Viper berus (Linnaeus, 1758) — Adder latastei (Boscá, 1878) — Lataste’s Viper seoanei (Lataste, 1879) — Seoane’s Viper ursinii (Bonaparte, 1835) — Meadow Viper