Bufotes, Rafinesque, 1815

Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin & Delfino, Massimo, 2024, New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans, Swiss Journal of Palaeontology (34) 143 (1), pp. 1-91 : 29-34

publication ID

https://doi.org/10.1186/s13358-024-00332-7

persistent identifier

https://treatment.plazi.org/id/E86287BF-FFF1-FFDB-BA71-A56EFEE6F884

treatment provided by

Felipe

scientific name

Bufotes
status

 

Bufotes View in CoL gr. viridis

Figure 25 View Fig

Material. Spilia 1: one femur ( UU SP1 1057); Spilia 3: two ilia ( UU SP3 669 and UU SP3 670); Spilia 4: two sacral vertebrae ( UU SP4 679 and UU SP4 680).

Description.

UU SP4 679 is the best-preserved sacral vertebra from Spilia ( Fig. 25a–e View Fig ). It has a single anterior cotyle, which is elliptical and mediolaterally elongated, and two posterior condyles, subelliptical as well but less elongated. Te walls of the neural arch are robustly built. Dorsally, the neural arch bears a sort of pseudozygosphenal structure anteriorly and a distinct neural crest. Te transverse processes are broken, but a laterally open fossa is present by their bases. A single prezygapophysis is preserved: it is wide and subcircular. UU SP4 680 is less preserved, but it shares the same morphology. Only the pseudozygosphenal structure is not evidently developed in this second specimen.

Te ilia from Spilia 3 are moderately small ( Fig. 25f, i View Fig ). Tey preserve a straight shaft devoid of dorsal crest and a small portion of the body. Te dorsal tubercle is better preserved in UU SP3 670 ( Fig. 25h, i View Fig ), displaying a rounded shape with small lobes dorsally. A deep preacetabular fossa is present, being more visible in UU SP3 669 ( Fig. 25f, g View Fig ).

Te femur (UU SP1 1057) from Spilia 1 is rather small and slender ( Fig. 25j View Fig ). It displays a well-developed and sharp crista femoris.

Remarks. Green toads ( Bufotes viridis complex) are present in the Spilia assemblage, identified through the criteria of Bailon (1999) and Ratnikov (2001): sacral vertebrae with an anterior cotyle, two posterior condyles, a distinct neural crest, and a laterally-open fossa close to each transverse process; ilia without dorsal crest, but provided with a multilobed dorsal tubercle and a deep preacetabular fossa; a femur with an undivided crista femoris. A pseudozygosphene is not commonly reported in green toad sacral vertebrae, but it was observed in at least few fossils from the Quaternary of southern Italy (Pirro Nord; M.D., pers. obs.) referred to this clade. Te taxonomic significance of this structure in these anurans may be worth further scrutiny. However, the structure observed in UU SP4 679 appears asymmetrical at close inspection, and so it may be interpreted as some sort of anomaly.

Bufonidae indet.

Figures 26 View Fig , 27 View Fig

Material. Chalicorrema: one sacral vertebra ( UU RA 501); Spilia 1: one humerus ( UU SP1 1041); Spilia 3: three trunk vertebrae ( UU SP3 661– UU SP3 663) and one humerus ( UU SP3 667); Spilia 4: one humerus ( UU SP4 681).

Description.

Te moderately large trunk vertebrae from Spilia 3 are robust ( Fig. 26a–e View Fig ). Te centrum is procoelous, well individualized and dorsoventrally compressed. Te neural arch has robust walls and a flat dorsal surface. A low carina neuralis is present. Te transverse processes are located posterior to the prezygapophyses.

UU RA 501 from Chalicorrema is the only bufonid element from that locality, representing a small fragment of a sacral vertebra ( Fig. 27 View Fig ). In spite of the small size, the vertebra was rather robustly built. Only the left half of the centrum and a small part of the left transverse process are preserved. Te centrum has an anterior cotyle. Posteriorly, there is an elliptical condyle. It can be assumed that another, similar condyle was present also on the missing right half of the centrum. Te two condyles were not coalescent. Te transverse process is too poorly preserved to give any significant morphological information. Te fragment has a total centrum length of 2.2 mm.

Te humeri from Spilia are medium sized ( Fig. 26f–h View Fig ). Tey have a straight shaft with no crista paraventralis. Te eminentia capitata is slightly shifted laterally. A moderately shallow fossa cubitalis ventralis is present. Te cristae lateralis and medialis are moderately developed. Te distal epiphysis is poorly ossified distally. Te olecranon scar is moderately wide.

Remarks. Tese fossils from Spilia share the diagnostic combination of features of bufonid anurans as reported by Bailon (1999). It is not possible, however, to identify them at lower taxonomic level, even though they could pertain to green toads as these are the only bufonids currently known from Spilia. Te straight humeral shaft, which is present in males of Bufo bufo (Linnaeus, 1758) and Bufotes viridis and absent in Epidalea calamita (Laurenti, 1768) and females of the preceding taxa according to Bailon (1999), may support this hypothesis.

Similarly, in spite of its very fragmentary status, the sacral vertebra UU RA 501 from Chalicorrema can still be assigned to a bufonid anuran, based on the overall robustness, the anterior cotyle, and the well-separated posterior condyles (Bailon, 1999). Further identification is not possible, but the size hints towards either a small species or a juvenile of a larger one.

Anura indet.

Figures 28 View Fig , 29 View Fig , 30 View Fig

Material. Chalicorrema: one trunk vertebra ( UU RA 508), one humerus ( UU RA 520), one radioulna ( UU RA 506), one femur ( UU RA 502), and two tibiofibulae ( UU R 503 and UU RA 507); Rema Marmara : eight bone fragments ( UU RMA 401– UU RMA 405); Spilia 1: one premaxilla ( UU SP1 1085), 14 maxillae ( UU SP1 1045, UU SP1 1051, UU SP1 1052, UU SP1 1055, UU SP1 1056, UU SP1 1060, UU SP1 1062– UU SP1 1064, UU SP1 1111, and UU SP1 1138), six angulars ( UU SP1 1106, UU SP1 1117, UU SP1 1139, and UU SP1 1149), one sculptured bone fragment ( UU SP1 1020), two atlantes ( UU SP1 1039 and UU SP1 1142), 10 trunk vertebrae ( UU SP1 1054, UU SP1 1061, and UU SP1 1151), seven fragments of vertebra ( UU SP1 1102, UU SP1 1104, UU SP1 1114, UU SP1 1116, UU SP1 1121, and UU SP1 1128), one sacral vertebra ( UU SP1 1040), two urostyles ( UU SP1 1137 and UU SP1 1148), one scapula ( UU SP1 1124), three coracoids ( UU SP1 1025, UU SP1 1118, and UU SP1 1153), 11 humeri ( UU SP1 1019, UU SP1 1059, UU SP1 1105, UU SP1 1131, UU SP1 1132, UU SP1 1140, UU SP1 1146, and UU SP1 1147), 16 radioulnae ( UU SP1 1050, UU SP1 1099, UU SP1 1103, UU SP1 1113, UU SP1 1122, UU SP1 1130, and UU SP1 1135), 13 ilia ( UU SP1 1029, UU SP1 1030, UU SP1 1032, UU SP1 1036, UU SP1 1037, UU SP1 1042, and UU SP1 1123), eight femora ( UU SP1 1058, UU SP1 1120, UU SP1 1129, UU SP1 1136, and UU SP1 1152), eight tibiofibulae ( UU SP1 1101, UU SP1 1107, UU SP1 1119, UU SP1 1125, and UU SP1 1150), nine phalanges ( UU SP1 1047– UU SP1 1049, UU SP1 1053, UU SP1 1112, UU SP1 1126, and UU SP1 1127), and 252 indeterminate elements ( UU SP1 1023, UU SP1 1084, UU SP1 1086, UU SP1 1091, and UU SP1 1095); Spilia 2b: one radioulna ( UU SP2b 503) and one indeterminate element ( UU SP2b 502); Spilia 2d: one maxilla ( UU SP2d 501) and one indeterminate element ( UU SP2d 502); Spilia 3: one maxilla ( UU SP3 671), seven trunk vertebrae ( UU SP3 616, UU SP3 656, UU SP3 666, UU SP3 684, and UU SP3 687), three fragments of vertebra ( UU SP3 682), one sacral vertebra ( UU SP3 619), five urostyles ( UU SP3 640, UU SP3 664, UU SP3 665, and UU SP3 686), one scapula ( UU SP3 657), two coracoids ( UU SP3 690), three humeri ( UU SP3 634, UU SP3 635, and UU SP3 689), 25 radioulnae ( UU SP3 620 and UU SP3 673), two ilia ( UU SP3 688), one ischium ( UU SP3 685), one tibiofibula ( UU SP3 683), and six indeterminate elements ( UU SP3 681 and UU SP3 691); Spilia 4: four premaxillae ( UU SP4 738), 22 maxillae ( UU SP4 512, UU SP4 689, UU SP4 690, UU SP4 696, and UU SP4 735), eight sculptured fragments ( UU SP4 739), four atlantes ( UU SP4 698 and UU SP4 731), 33 trunk vertebrae ( UU SP4 624 and UU SP4 724), 13 fragments of vertebra ( UU SP4 514), two sacral vertebrae ( UU SP4 697 and UU SP4 713), four urostyles ( UU SP4

522 and UU SP4 725), one scapula ( UU SP4 700), five coracoids ( UU SP4 677, UU SP4 678, and UU SP4 730), four humeri ( UU SP4 699 and UU SP4 729), 10 radioulnae ( UU SP4 614, UU SP4 694, and UU SP4 695), two ilia ( UU SP4 527 and UU SP4 726), three tibiofibulae ( UU SP4 727), two phalanges ( UU SP4 732 and UU SP4 733), and 95 indeterminate elements ( UU SP4 510 and UU SP4 511); Vevi: one premaxilla ( UU VE 548), one sacral vertebra ( UU VE 570), one humerus ( UU VE 569), three radioulnae ( UU VE 552), two tibiofibulae ( UU VE 539 and UU VE 571), one indeterminate autopodial element ( UU VE 572), one phalanx ( UU VE 551), and one distal phalanx ( UU VE 550).

Description and remarks.

Among the remains from Spilia belonging to indeterminate anurans ( Fig. 28 View Fig ), the largest and more robust likely pertain to either Latonia or a bufonid. A brief description of the different morphological features encountered in the phalanges from Spilia can be provided. On the whole, the phalanges from Spilia are medium-sized, elongated and rather slender ( Fig. 28c, d View Fig ). Tey narrow distinctly toward the distal tip. Te bulb at the tip is either smooth [eight specimens: UU SP1 1047, UU SP1 1048, UU SP1 1049, UU SP1 1126 ( Fig. 28d View Fig ), UU SP1 1127, and UU SP4 732] or rough [two specimens: UU SP1 1053 ( Fig. 28c View Fig ) and UU SP4 733]. Two lobes are slightly recognizable on the tip of UU SP1 1048, UU SP1 1126, and UU SP1 1127. Specimen UU SP1 1112 is too poorly preserved to recognize the shape of its tip. It is currently not clear, however, to what degree these morphologies have a taxonomical significance .

Similarly, the indeterminate anuran remains from Vevi

( Fig. 29 View Fig ) pertain perhaps to one of the taxa identified above from the same locality. Te distal phalanx (UU VE 550), in particular, is small and slender ( Fig. 29c View Fig ). It has an elongated triangular shape, strongly narrowing distally and ending with a globular tip with no evident lobes. Tis is similar to phalanxes of Latonia figured by Vasilyan (2020, fig. 6), but clear diagnostic features of anuran phalanxes even at supraspecific levels are not known at the moment. We thus refrain from referring this phalanx to the Vevi Latonia .

Te remains from Chalicorrema are even more fragmentary ( Fig. 30 View Fig ). Tese remains are too poorly preserved to give significant morphological information, allowing precise taxonomic allocation. Similarly, the even fewer remains from Rema Marmara can be assigned to indeterminate anurans, but they cannot be identified more precisely due to either poor preservation or scarce diagnostic value.

UU

University of Uppsala

Kingdom

Animalia

Phylum

Chordata

Class

Amphibia

Order

Anura

Family

Bufonidae

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