taxonID	type	description	language	source
03F52665D050FF8FFC609E34FB8C74CD.taxon	description	Figures 3 – 4	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D050FF8FFC609E34FB8C74CD.taxon	materials_examined	Material. Hambach 6 C: three atlases (IPB-HaH 2117, IPB-HaH 2175, IPB-HaH 2164); one trunk vertebra (IPB-HaH 2162). Hambach 11: two dentaries (IPB-HaR 2043, IPB-HaR 2071); one anterior trunk vertebra (IPB-HaR 2183); one trunk vertebra (IPB-HaR 2008). Hambach 11 C: five trunk vertebrae (IPB-HaR 2404 / 2408).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D050FF8FFC609E34FB8C74CD.taxon	description	Description. IPB-HaR 2071 (Figure 3) represents part of the posterior end of a very large dentary. It is very fragmentary. IPB-HaR 2043 is also very large and robust. It represents a portion originally located somewhere in the middle of the dentary. The pars dentalis of the dentary is composed by a very high dental lamina and a very low subdental lamina. On the medial side, nine tooth positions are visible, hosting the poorly-preserved remains of pleurodont, very high, rather narrow and closely spaced teeth provided with thick walls. Ventral to the pars dentalis, there is a robust shelf, which is not strongly developed medially. The preservation of this shelf is rather poor, but a narrow and shallow groove is visible ventrally. The groove moves towards the medial surface near the anterior end of the fragment. The lateral surface is smooth. In lateral / medial view, the ventral margin is somehow concave, suggesting ventral development of the posterior part of the dentary. IPB-HaH 2164 is a very small atlas, but the other ones are larger. The length of the largest atlas, IPB-HaH 2175 (Figure 4 F-I), is 6 mm. All atlases miss almost completely the neural arch, preserving only the centrum. Anteriorly, the occipital joints are wide and mediolaterally elongated, with a suboval / subelliptical shape (i. e., slightly dorsoventrally compressed) in anterior view. They are very shallowly concave and do not coalesce in the middle, being separated by a thin and long processus odontoideus. The processus is slightly slenderer in IPB-HaH 2175 than in the other two specimens. It has a flat dorsal surface and an anteroventrally-directed and strip-like articular surface that is not separated into two distinct lateral areas. There is no postodontoideus foramen at the base of the process. The posterior end of the centrum is represented by a posterior cotyle with a notochordal pit in the middle. The cotyle is circular in IPB-HaH 2175 (Figure 4 I) and more mediolaterally compressed in IPB-HaH 2117 (Figure 4 D). In both specimens, it is larger than the processus odontoideus. This feature cannot be evaluated in IPB-HaH 2164 because the area is damaged. In IPB-HaH 2715, the ventral surface of the centrum shows a concave area with a number of large foramina (Figure 4 G). On the other hand, IPB-HaH 2117 bears a very deep fossa in the middle of the centrum, which is flanked by two smaller symmetrical foramina by the sides and by other even smaller foramina posteriorly (Figure 4 B). The ventral surface of IPB-HaH 2164 displays a depressed area by each side of the centrum. Foramina also cover the lateral surface of each processus lateralis in all specimens. Both IPB-HaR 2008 (Figure 4 O-T) and IPB-HaR 2408 (Figure 4 U-Z) are amphicoelous and large sized (the centrum length reaches 6.5 mm and 9.5 mm, respectively). A notochordal pit is present in the middle of the large and hourglassshaped centrum. The cotyles are circular in both anterior and posterior views. The ventral surface of the centrum bears two high and sharp longitudinal basapophyses, which run parallel along the entire length of the vertebra. Only in IPB-HaR 2408, the basapophyses contact in the middle at about one third of the centrum length (Figure 4 V). Between the basapophyses, various central foramina are present. At least three large ones are visible on IPB-HaR 2008 (Figure 4 P), surrounded by other smaller ones, whereas only a number of small ones are visible on IPB-HaR 2408 (Figure 4 V). The neural arch is robust and dorsally flattened, being better preserved in IPB-HaR 2408. A neurapophysis is present. It is low anteriorly, but rises to a moderate degree posteriorly. A robust spine is present at the posterior end of the arch, strongly projecting posterodorsally beyond the postzygapophyses. The posterior end of the spine is truncated. A wide and deep, U-shaped anterior notch on the neural arch is visible on IPB-HaR 2408 (Figure 4 U), allowing the anterior cotyle to be visible in dorsal view. The deepest part of the notch reaches the posterior margin of the prezygapophyses. The zygapophyses are more or less horizontal in both anterior and posterior view. The zygapophyseal facets are suboval. In posterior view, two shallow depressions are visible on the posteroventral surface of the neural arch, flanking a low longitudinal ridge running along the ventral side of the neural spine. The transverse processes are moderately developed and posterolaterally directed. Ventrally, they are connected to the centrum by moderately- (IPB-HaR 2008) or well-developed (IPB-HaR 2408) anterior ventral crests (anterior alar process in Vasilyan and Yanenko, 2020) and little-developed posterior ventral crests. On the other hand, the zygapophyseal crests are not developed. The other vertebrae only preserve isolated centra or fragments of centra, some of them being much smaller than the two previously described (IPB-HaR 2406 is about 4 mm long). Nevertheless, they share the same morphology. IPB-HaR 2183 (Figure 4 J-N) is the only trunk vertebra displaying some differences from the other ones. This large vertebra has a very massive, almost 6 mm long centrum, which is amphicoelous, hour-glass-shaped and notochordal. The overall aspect is relatively shorter compared to the trunk vertebra IPB-HaR 2008, coming from the same level. The anterior cotyle is moderately mediolaterally compressed, appearing subelliptical in anterior view. The posterior cotyle, on the other hand, is subcircular in posterior view. The ventral surface of the centrum bears a narrow keel and no basapophyses. A number of small foramina are present by the ventrolateral sides of the centrum, in place of real subcentral foramina. Most of the neural arch is missing, preserving only the right lateral wall and the base of the left one. The transverse processes are long and moderately robust; they are directed posterolaterally. The anterior ventral crest is very low, whereas the posterior one is slightly more developed. Zygapophyseal crests are not developed. The base of the right prezygapophysis is also preserved, but not the zygapophysis itself. The shortness of this vertebra, together with its ventral keel, identify it as an anterior trunk vertebra (Estes et al., 1967).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D050FF8FFC609E34FB8C74CD.taxon	discussion	Remarks. The diagnostic features recently reported by Vasilyan and Yanenko (2020) allow a rather straightforward identification for these remains as a batrachosauroidid salamander. The dentaries of these urodeles have thick-walled teeth and a ventrally-projecting posterior part. Trunk vertebrae are amphicoelous, with subcircular cotyles, basapophyses, and a posterodorsally-projecting neural spine. The atlases are provided with both anterior and posterior cotyles with a rounded outline, among which the former are large and concave. The well-developed paired ventral basapophyses, as well as maybe the flat neural arch and developed anterior ventral crest, suggest that the batrachosauroidid from Hambach is a member of the European genus Palaeoproteus (Vasilyan and Yanenko, 2020). In particular, the following combination of features observed in the studied material is diagnostic of P. miocenicus following Vasilyan and Yanenko (2020): overall large size; poorly-developed subdental shelf of dentary; vertebrae with a long neural spine; weakly-concave and slightly dorsoventrally-compressed anterior cotyles of the atlas; external surface of the atlas pierced by foramina of different sizes; strongly pronounced, lip-shaped odontoid process of the atlas (Vasilyan and Yanenko, 2020: p. 8, stated that “ In P. miocenicus, the odontoid process is very large in comparison to all known batrachosauroidids ”). However, the atlases from Hambach 6 C show no postodontoid foramen, in contrast with the type and referred material of P. miocenicus (Vasilyan and Yanenko, 2020). All four P. miocenicus atlases share the presence of this foramen, whereas it is absent in Palaeoproteus klatti Herre, 1935, and Palaeoproteus gallicus Estes et al., 1967. Despite this difference, we refrain to name a new species here and attribute the fossils from Hambach 6 C to Palaeoproteus cf. miocenicus, due to the overall similarity between bones of P. miocenicus and the German taxon as well as the low sample of atlases that hinders a complete understanding of the real variation of this feature in these animals. Material coming from the youngest level in Hambach also shares the same attribution, given that atlases from this level are currently unknown, and so it is not possible to verify the presence or absence of a postodontoid foramen.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D053FF80FC199AE1FE0E7493.taxon	description	Figure 5	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D053FF80FC199AE1FE0E7493.taxon	materials_examined	Material. Hambach 6 C: one humerus (IPB-HaH 2390).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D053FF80FC199AE1FE0E7493.taxon	description	Description. This humerus is rather large and robust. It misses only most of the distal epiphysis. The length of the preserved portion is about 16 mm. The bone expands strongly towards the epiphyses. Based on the preserved portion, it appears rather short. The proximal epiphysis is poorly ossified. It has an asymmetrical shape, more expanded toward the ventral side. The posterior surface displays a distinct fossa.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D053FF80FC199AE1FE0E7493.taxon	discussion	Remarks. The size of this humerus suggests that it belongs to one of the two largest caudate taxa recognised in Hambach (i. e., either cryptobranchids or batrachosauroidids). The bone appears rather short in general appearance, which hints against Chelotriton, cryptobranchids, and possibly also proteids. Its shortness rather recalls the shortened limbs of P. klatti (see Vasilyan and Yanenko, 2020). Furthermore, the general morphology of the bone differs from the fragment of humerus attributed to Mioproteus gardneri Venczel and Codrea, 2018, whereas the poorly ossified proximal end hints against Chelotriton. Therefore, IPB-HaH 2390 is here attributed with doubt to the batrachosauroidid taxon found in the Hambach assemblage, Palaeoproteus cf. miocenicus.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D05CFF80FED19B23FE5F7218.taxon	materials_examined	Material. Hambach 6 C: five atlases (IPB-HaH 2119 / 2121, IPB-HaH 2150, IPB-HaH 2165).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D05CFF80FED19B23FE5F7218.taxon	discussion	Remarks. These atlases were recently described as types of a new proteid taxon, E. grogu. The reader is referred to Macaluso et al. (2022 b) for further information.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D05CFF81FED59CB4FDD47309.taxon	description	Figure 6	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D05CFF81FED59CB4FDD47309.taxon	materials_examined	Material. Hambach 11: nine trunk vertebrae (IPB-HaR 2000 / 2007, IPB-HaR 2013). Hambach 11 C: six trunk vertebrae (IPB-HaR 2400 / 2403, IPB-HaR 2427 / 2428).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D05CFF81FED59CB4FDD47309.taxon	description	Description. These vertebrae (Figure 6) reach a large size and are robustly ossified. The largest one has a centrum length of about 6.5 mm. They display a range of different degree of preservational statuses, from very fragmentary to well preserved. The centrum is amphicoelous and pierced by a wide notochordal canal. Ventrally, the centrum bears a robust and high keel, which is more or less narrow (from sharp to slightly thicker). By the anterior and posterior ends, the keel expands into triangular and flat ventral surfaces. The keel is flanked by small subcentral foramina, sometimes more than one per side. Posteriorly, poorly-developed basapophyses are present. The neural arch is low, without a significant rising posteriorly, and dorsally flat. A neurapophysis is present, being either rather low or slightly higher. It runs almost entirely along the dorsal surface of the arch. The anterior margin of the arch appears straight in dorsal view, even though it is never completely preserved. It is located roughly at midlength of the prezygapophyses. The posterior margin is rather straight (but very slightly wavy, even though with no median notch) and ends well before the end of the postzygapophyses. By each side of the neurapophysis, the posterior end of the dorsal surface of the neural arch carries a well-developed spine, which projects well beyond the posterior margin, but not beyond the postzygapophyses. The spines are well spaced in dorsal view, not strongly extended anteriorly (they do not reach the level of the transverse processes) and not connected to the neurapophysis medially. The zygapophyses are suboval and almost horizontal. The ventral lamina has a subtrapezoidal shape, originated by very well-developed anterior ventral crests and less-developed posterior ones. The zygapophyseal crests are also well developed, but not as much as the anterior ventral crests. The posterior zygapophyseal crests are distinctly convex dorsally in lateral view. Due to the strong development of both the ventral lamina and the zygapophyseal crests, the vertebrae assume a wide appearance in both ventral and dorsal views. The transverse processes are reduced to a small single structure, which is slender and posterolaterally directed. Individualized para- and diapophyses are not clearly discernible. A more-or-less large foramen is visible anterior to the base of the transverse process.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D05CFF81FED59CB4FDD47309.taxon	discussion	Remarks. These vertebrae are referred to Mioproteus because of the following combination of features (Estes and Darevsky, 1977): robust and wellossified aspect; well-spaced posterior spines; wide appearance in dorsal and ventral views (i. e., wide neural arch, wide ventral lamina); presence of basapophyses. Three species of Mioproteus are currently known: Mioproteus caucasicus Estes and Darevsky, 1977, M. gardneri, and M. wezei. The vertebrae from Hambach are concordant in size with both M. caucasicus and M. wezei, whereas they are larger than M. gardneri. Regarding their morphology, they differ from M. gardneri in the more developed zygapophyseal crests, the taller neurapophysis, and the more laterally-trending prezygapophyses (Venczel and Codrea, 2018). Most characters reported to be diagnostic for M. wezei are variable, but at least the consistent absence of a connection between the posterior spines and the neurapophysis seems to be sufficiently reliable (Syromyatnikova et al., 2021). This condition is shown by the Hambach material, whereas M. caucasicus either has posterior spine that connect or not to the neurapophysis medially. Strongly dorsally-convex posterior zygapophyseal crests may also be typical for M. wezei (Młynarski et al., 1984; Bailon, 1995; even though Syromyatnikova et al., 2021, mentioned some variation of this feature in Mioproteus vertebrae). Considering this, we here tentatively attribute the Hambach Mioproteus vertebrae to M. wezei, even though pointing out uncertainty over this identification pending a clarification of the diagnostic features within Neogene Mioproteus.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D043FF92FCEB98DBFD3E70B3.taxon	description	Figures 16 – 17	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D043FF92FCEB98DBFD3E70B3.taxon	materials_examined	Material. Hambach 11: one sphenethmoid (IPB-HaR 2021); 11 angulars (IPB-HaR 2110 / 2113, IPB-HaR 2129 / 2132, IPB-HaR 2144 / 2146); one trunk vertebra (IPB-HaR 2030); five humeri (IPB-HaR 2148 / 2152); five ilia (IPB-HaR 2099 / 2102, IPB-HaR 2147). Hambach 11 C: one maxilla (IPB-HaR 2429); one angular (IPB-HaR 2418); two humeri (IPB-HaR 2419 / 2420).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D043FF92FCEB98DBFD3E70B3.taxon	description	Description. IPB-HaR 2429 (Figure 16 A-C) is a fragment of maxilla, measuring about 5.5 mm in length and preserving only the area of the processus palatinus. The bone is robustly built. On the medial side, part of the tooth row is preserved, even though in bad conditions. Four wide tooth positions are preserved, one of them still hosting the base of a tooth. The tooth positions are separated by knob-like structures. The lamina horizontalis is mostly broken, but it clearly extended medially with a toothless portion. No clear ridge is visible ventrally separating the toothed and toothless portion of the lamina. On its dorsal side, a deep recessus vaginiformis is present. The processus palatinus is represented in this specimen by a low, subtriangular structure with a truncated dorsal tip. In dorsal view, it is shifted medially, thus originating a concavity on the lateral surface of the maxilla. Both the anterior and posterior margins of the process are distinctly irregular, and its lateral surface displays few foramina. IPB-HaR 2021 (Figure 16 D-G) is a fragmentary sphenethmoid. It is anteroposteriorly elongated and large-sized. The lateral margins of the bone are rather eroded, but distinctly developed laminae supraorbitalis and trabecula seem not to be present. In dorsal view, a long and U-shaped fenestra frontoparietalis is recognizable, even if the left portion of the bone is lacking. Anteriorly to the fenestra, the dorsal surface of the bone is smooth. The contact surface with the parasphenoid is visible on the ventral surface: it is delimited laterally by two low ridges and widens anteriorly. Angulars are robust and large-sized. They have a rather deep sulcus cartilagine Meckeli and an anteroposteriorly elongate, stocky and dorsoventrally compressed processus coronoideus. The dorsal surface of the latter is strongly irregular, with pits and / or ridges, and shows a certain degree of individual variation (Figure 17). A few angulars, such as e. g., IPB-HaR 2111 (Figure 17 A) and IPB-HaR 2418, bear a small tubercle on the lateral side, in correspondence with the anterior end of the processus coronoideus. The extremitas spatulata is short and broad. IPB-HaR 2030 is a fragmentary trunk vertebra provided with a very dorsoventrally compressed centrum. The latter is procoelous, wide, and shows numerous small pits on the ventral surface. Humeri (Figure 16 H-M) are very large-sized and lack a fossa cubitalis ventralis. The eminentia capitata and the epicondyles are distally eroded and were probably partly cartilaginous in the living animal. The epicondylus ulnaris is only slightly larger than the epicondylus radialis. The olecranon scar is depressed, but the articular surface with the olecranon of the radioulna is small and poorly developed. All specimens but IPB-HaR 2420 preserve only the distal epiphysis and the distal part of the diaphysis. IPB-HaR 2420 is more preserved, but the proximal end of the bone is still missing. A hint of a robust crista ventralis is recognizable on the ventral surface of the humeri. This is particularly evident in IPB-HaR 2420, which also express the base of a crista paraventralis. Cristae medialis and lateralis are not developed. Ilia (Figure 16 N-Q) show a well-developed and elongated dorsal tubercle, but no dorsal crest. The tubercle bends in lateral direction. The large acetabular fossa has a prominent anteroventral rim. The ventral acetabular expansion is not developed, whereas the dorsal one is moderately developed. A supraacetabular fossa is visible dorsally to the acetabulum. A wide and deep interiliac groove is visible on the medial surface of the body of the bone.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D043FF92FCEB98DBFD3E70B3.taxon	discussion	Remarks. Clear diagnostic features of palaeobatrachid anurans in the above-described material are the following (Wuttke et al., 2012; Roček, 2013; Roček et al., 2021): knob-like structures separating tooth positions in the maxilla; elongated sphenethmoid, provided with frontoparietal fenestra longer than half the total length of the bone and with two parallel ridges delimiting the articulation area for the parasphenoid on the ventral surface; coronoid process of the angular either smooth or bearing muscle scars on dorsal surface; vertebral centrum strongly dorsoventrally compressed and with numerous pits on the ventral surface; humerus devoid of fossa cubitalis ventralis (different from Eocene palaeobatrachids, though) and provided with a comparatively small eminentia capitata that is located on or near the long axis of bone, as well as similar-sized epicondyles; ilium with massive dorsal acetabular expansion; large acetabular fossa extending anteroventrally beyond the margin of the ilial body, thus concealing the ventral expansion; dorsal tubercle protruding only slightly in dorsal direction, but more prominent laterally and bearing muscle scars on the lateral surface; distinct horizontal depression on the inner surface of the iliac shaft. Specific attribution to P. eurydices appears also justified, due to the following combination of features (Villa et al., 2016): the interorbital section of the processus cultriformis of the parasphenoid was narrow, but the process becomes wider towards the anterior (as suggested by the divergent longitudinal ridges on the ventral side of sphenethmoid, which delimit laterally the area of attachment of the parasphenoid); the sphenethmoid lacks articular facets for the nasals; the dorsal surface of the sphenethmoid shows no median ridge; the processus coronoideus of the angular extends parallel to most of the extremitas spatulata; the extremitas spatulata is short and broad; the fossa cubitalis ventralis of humerus is absent. Despite its very fragmentary status, the maxilla IPB-HaR 2429 is also very similar in morphology to maxillae of P. eurydices. In particular, it could be somehow representative of a sort of “ intermediate ” stage between the supposed juvenile maxilla reported by Villa et al. (2016) and the adult ones. Similar to the purported juvenile specimen, IPB-HaR 2429 has a subtriangular processus palatinus, which gives to the bone a concave lateral appearance in dorsal view due to a slight medial shifting. However, the size of the specimen, as well as the number of teeth in the processus palatinus area and the absence of a ridge separating the toothed and toothless portions of the ventral surface of the lamina horizontalis, seem to be more comparable with the adult holotype maxilla of P. eurydices.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04EFF93FEF79EE2FCD174ED.taxon	description	Figure 18	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04EFF93FEF79EE2FCD174ED.taxon	materials_examined	Material. Hambach 6 C: one maxilla (IPB-HaH 2279).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04EFF93FEF79EE2FCD174ED.taxon	description	Description. IPB-HaH 2279 is a fragmentary and robust maxilla, lacking the anterior and posterior portions. The lamina horizontalis is robust and moderately developed in medial direction. On the ventral surface of the lamina, a low longitudinal ridge separates a toothless medial half from the crista dentalis. Teeth were pleurodont, but no one is preserved; they were separated by small processes, which are now eroded. Nine tooth positions are recognizable. In the middle of the dorsal surface of the lamina, a deep and circular recess is visible. By the recess, the lateral wall of the maxilla bends medially. The processus palatinus should be located by this bending, but it is not clearly recognizable in this specimen; this could be either due to a real absence of the processus or, more likely, to preservational reasons. The lateral surface of the bone is smooth.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04EFF93FEF79EE2FCD174ED.taxon	discussion	Remarks. This specimen from Hambach 6 C is rather similar to the maxillae of P. eurydices in (Villa et al., 2016): overall robustness; presence of the circular recess (recessus vaginiformis for the processus maxillaris anterior, which is the anterior termination of the subocular bar; Roček, 2003: 1941) on the lamina horizontalis; presence of processes separating the teeth (even though it is not possible to say if they were knob-like in origin). On the ventral surface of the lamina horizontalis, the low ridge delimitating medially the toothed area recalls the maxilla RGM 632039, possibly referred to a young P. eurydices by Villa et al. (2016). However, the Hambach 6 C specimen has more teeth than the latter. IPB-HaH 2279 may thus represent a palaeobatrachid, maybe even related to P. eurydices, but in absence of further, better preserved and more taxonomically significant, material from the Miocene level of Hambach, it appears more confident to propose only a cautious identification.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04FFF93FE9D9B21FC24716D.taxon	description	Figure 19 A-J	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04FFF93FE9D9B21FC24716D.taxon	materials_examined	Material. Hambach 11: one sacral vertebra (IPB-HaR 2179); one humerus (IPB-HaR 2154); two ilia (IPB-HaR 2084, IPB-HaR 2103).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04FFF93FE9D9B21FC24716D.taxon	description	Description. IPB-HaR 2179 (Figure 19 A-D) is a moderately small sacral vertebra and has an amphicoelous and cylindrical centrum. The neural canal is circular and the dorsal surface of the neural arch is smooth. Both the prezygapophyses and the transverse processes are broken off. The latter are anteroposteriorly extended. IPB-HaR 2154 (Figure 19 E-F) is a poorly preserved humerus provided with a curved diaphysis (though it is broken and misses the proximal portion) and an eminentia capitata that is shifted laterally compared to the main axis of the bone. A deep fossa cubitalis ventralis is present; it opens on the lateral side. There are no cristae medialis and lateralis. Ilia (Figure 19 G-J) lack a dorsal tubercle and a dorsal crest. They have an acetabular fossa provided with a strong anteroventral rim. The dorsal acetabular expansion is rather short. No supraacetabular fossa, preacetabular fossa, interiliac groove, or interiliac tubercle are visible. The spiral groove is not distinct. The posterior end of the bone is slightly eroded in both specimens, but very light striae are visible on their posteromedial surface.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04FFF93FE9D9B21FC24716D.taxon	discussion	Remarks. Few elements from Hambach 11 are attributed to Pelobatidae because of the following combination of features (Bailon, 1999; Roček, 2013): curved diaphysis of the humerus; laterallyshifted eminentia capitata; deep fossa cubitalis ventralis, which is open laterally; ilia with no dorsal crest and no dorsal tubercle; no preacetabular and supracetabular fossae; striae on the medial side of the ilial body. Within pelobatids, the absence of a deep spiral (or oblique) groove is used to distinguish Eopelobates from Pelobates Wagler, 1830 (Böhme, 2010; Syromyatnikova, 2019), and thus these fossils are here assigned to the former genus. However, this identification is only considered tentative here, because not all authors deem isolated postcranial elements sufficient for genus level discrimination (Rage and Roček, 2003). Known Eopelobates species are all based on articulated material (Roček et al., 2014), making comparison with the disarticulated specimens from Hambach difficult. Furthermore, most of the diagnostic features of the species are on cranial elements. Nevertheless, at least Eopelobates deani Roček et al., 2014, and Eopelobates grandis Zweifel, 1956, seem to differ from the Hambach ilia in the low dorsal tubercle and the low dorsal crest in the anterior portion of the shaft respectively. The sacral vertebra IPB-HaR 2179 is also tentatively attributed to cf. Eopelobates sp. because of the cylindrical centrum and the extended transverse processes, as well as the presence of the spinal foramina. In pelobatids, sacral vertebrae not fused to the urostyle are present in both Eopelobates and Pelobates (Bailon, 1999; Roček et al., 2014; Syromyatnikova, 2017). However, vertebrae of pelobatids are usually procoelous. Amphicoelous vertebrae, followed by a cartilaginous disk, are known only in E. grandis (even though its referral to Eopelobates is called into question by some authors also because of this feature; Roček et al., 2014). The holotype and only known specimen of E. grandis does not preserve the sacral centrum or the anterior part of the urostyle, and so an amphicoelous condition of the former cannot be evaluated. Amphicoelous sacral centra are shown by Ascaphus Stejneger, 1899, and some extinct Mesozoic frogs (Reilly and Jorgensen, 2011), but accounting for the absence of any existing evidence supporting the possible presence of the North American ascaphids in Europe at any moment in time as well as the highly unlikely circumstance of a survival of an early-branching Mesozoic anuran lineage in the Pliocene of the continent, we here consider more probable that IPB-HaR 2179 could represent a pelobatid with either a peculiar vertebral morphology or an anomalous condition due to ontogenetic or pathologic circumstances.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D049FF95FE279D2CFC18746E.taxon	description	Figure 19 P-U	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D049FF95FE279D2CFC18746E.taxon	materials_examined	Material. Hambach 6 C: one trunk vertebra (IPB-HaH 2220); one humerus (IPB-HaH 2399).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D049FF95FE279D2CFC18746E.taxon	discussion	Discussion. IPB-HaH 2220 (Figure 19 P-S) is a moderately large-sized and procoelous trunk vertebra. It has a cylindrical centrum and a long neural arch, with a low carina neuralis and a well-developed posterior point. Transverse processes are broken off, but they were located under the prezygapophyses. The humerus (Figure 19 T-U) is small, but preserves part of the diaphysis and part of the epiphysis. The eminentia capitata is shifted laterally. A shallow fossa cubitalis ventralis, which is open on the lateral side, is present. The epicondyles are missing. The cristae medialis and lateralis are not developed. On the dorsal side, the visible part of the olecranon scar is poorly marked. Remarks. Following Bailon (1999), these two fossils can be assigned to pelobatids. This attribution is based on: vertebra procoelous, with a long neural arch, transverse processes located ventral to the prezygapophyses, a cylindrical centrum, and a well-developed posterior point; humerus with a laterally-open fossa cubitalis ventralis and maybe also lateral shift of the eminentia capitata. The absence of more taxonomically significant elements in the Miocene level of Hambach, however, hinders a discrimination of either Eopelobates or Pelobates.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04AFF96FC199902FC0B75F8.taxon	description	Figure 21	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04AFF96FC199902FC0B75F8.taxon	materials_examined	Material. Hambach 11: one sacral vertebra (IPB-HaR 2020).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04AFF96FC199902FC0B75F8.taxon	description	Description. IPB-HaR 2020 is a medium-sized sacral vertebra provided with an anterior cotyle and two posterior condyles. The neural canal is subelliptical. A low carina neuralis is present on the dorsal surface of the neural arch, whereas there are no fossettes. Transverse processes are broken and so it is not easy to evaluate their complete anteroposterior extension. Nevertheless, the preserved bases suggest that they were not cylindrical and also not strongly expanded.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D04AFF96FC199902FC0B75F8.taxon	discussion	Remarks. This sacral vertebra is here attributed to a representative of the common toad species group due to (Bailon, 1999): presence of an anterior cotyle; transverse processes not strongly extended; presence of a carina neuralis; and absence of fossettes on the dorsal surface of the neural arch.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D074FFA8FE309AC3FA7F74B3.taxon	description	Figure 22 G-N	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D074FFA8FE309AC3FA7F74B3.taxon	materials_examined	Material. Hambach 6 C: nine sacral vertebrae (IPB-HaH 2193 / 2194, IPB-HaH 2209 / 2215); one scapula (IPB-HaH 2323). Hambach 11: one trunk vertebra (IPB-HaR 2184); three sacral vertebrae (IPB-HaR 2018 / 2019; IPB-HaR 2031); one ilium (IPB-HaR 2087).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D074FFA8FE309AC3FA7F74B3.taxon	description	Description. IPB-HaR 2184 is a small and amphicoelous trunk vertebra, with a short neural arch and laterally directed transverse processes. Sacral vertebrae (Figure 22 G-J) are small and provided with one anterior and two posterior condyles. They have cylindrical transverse processes. IPB-HaH 2323 (Figure 22 K-L) is an elongated scapula. It has no ridges on the anterior margin, but it is provided with a ridge on the inner surface. The cavitas glenoidalis opens in posterior direction and is partially hidden by the pars acromialis in ventral view. The ilium (Figure 22 M-N) is small and poorly preserved, missing completely the shaft. It has a large and subcircular acetabular fossa. Dorsal to the latter, a deep supraacetabular fossa is present. The dorsal tubercle is also present, even though mostly broken away. Nevertheless, it appears moderately distinct. Anteriorly, it is in continuation with a dorsal crest. Both the dorsal and ventral acetabular expansions are broken, but the former appears well developed based on what is preserved. Medially, there is no interiliac groove or tubercle. The ilioischiatic juncture seems rather low and large, but it is not completely preserved.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D074FFA8FE309AC3FA7F74B3.taxon	discussion	Remarks. All these specimens present a combination of features supporting attribution to indeterminate ranids according to the criteria presented by Bailon (1999). For the trunk vertebra, this combination includes amphicoely, the short neural arch, and the laterally directed processes; this further suggests that it represents the eighth vertebra in the column (Bailon, 1999). In the sacral vertebra, the diagnostic combination consists of the anterior condyle, the two posterior condyles, and the cylindrical processes. For the scapula, significant are the elongation, the absence of crista anterior, the cavitas glenoidalis hidden in ventral view, as well as the presence of an inner ridge on scapula. The attribution of the ilium is supported by the presence of the dorsal crest and the absence of interiliac tubercles and grooves. The low and large juncture could hint at green frogs for the ilium (Gleed-Owen, 1998), but it is not clear how much this apparent morphology may be influenced by the preservation.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D074FFA9FB8A9AE3FDCE7071.taxon	materials_examined	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).	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
03F52665D074FFA9FB8A9AE3FDCE7071.taxon	discussion	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.	en	Villa, Andrea, Macaluso, Loredana, Mörs, Thomas (2024): Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe. Palaeontologia Electronica (a 3) 27 (1): 1-56, DOI: 10.26879/1323, URL: http://dx.doi.org/10.26879/1323
