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Late Cenozoic catfishes of Southeastern Europe with inference to their taxonomy and palaeogeography

Oleksandr Kovalchuk and Carl J. Ferraris

Plain Language Abstract

Catfish fossils (Siluriformes) from late Miocene through Pleistocene strata of Southeastern Europe are described in our paper. The material under study is represented by disarticulated bones from 26 localities in the territory of Southern Ukraine and the Russian Federation. Catfish fossils in that region are represented by members of two genera, Heterobranchus and Silurus, of two families (Clariidae and Siluridae, respectively), and at least five species, including one as new for science (Silurus spinosus n. sp.). These discoveries shed new light upon the evolutionary history of these fishes and establish some characteristics of their distribution during the late Cenozoic.

Resumen en Español

Peces gato del Cenozoico tardío de Europa sudoriental con inferencias respecto a su taxonomía y paleogeografía

Describimos fósiles de peces gato (Siluriformes) del Mioceno tardío hasta los estratos del Pleistoceno del sudeste de Europa. El material en estudio está representado por huesos desarticulados de 26 localidades en el territorio del sur de Ucrania y la Federación Rusa. Los fósiles de pez gato en esta región están representados por dos géneros, Heterobranchus y Silurus, de dos familias (Clariidae y Siluridae, respectivamente) y cinco especies, una de ellas nueva para la ciencia (Silurus spinosus n. sp.) y dos nombradas con nomenclatura abierta (Silurus cf. S. glanis, Silurus sp.). La presencia de †Heterobranchus austriacus en el Sarmatiense tardío del sur de Ucrania puede explicarse por la expansión de esta especie de África a Europa durante el Mioceno tardío (los primeros restos fósiles de este taxón sólo se documentaron en Europa central y occidental). La distribución geográfica de Silurus soldatovi fue más amplia en el pasado, como queda evidenciado por los numerosos hallazgos de sus huesos en los depósitos del Mioceno y Plioceno tardío de Europa sudoriental. El moderno Silurus glanis apareció por primera vez en la etapa final del Mioceno tardío (Pontiense) y probablemente sustituyó a S. soldatovi debido a su mayor plasticidad ecológica (especialmente su tolerancia a la temperatura - esta especie puede desovar en aguas más frías). Se ha observado una tendencia evolutiva interesante en el cambio de tamaño, forma y orientación de las denticulaciones en el eje de la espina de la aleta pectoral en diferentes especies del género Silurus. Esta tendencia puede explicarse en términos de la oligomerización de estructuras homólogas, pero esta cuestión merece un estudio especial y no se aborda aquí.

Palabras clave: Clariidae; Siluridae; nueva especie; Mioceno tardío; Plioceno; Pleistoceno

Traducción: Enrique Peñalver (Sociedad Española de Paleontología)

Résumé en Français

Des poissons-chats du Cénozoïque récent du sud-est de l'Europe : implications taxinomiques et paléogéographiques

Nous décrivons du matériel de poissons-chats (Siluriformes) fossiles provenant de couches du sud-est de l'Europe et datant du Miocène récent au Pléistocène. Le matériel étudié est représenté par des os désarticulés de 26 localités du territoire d'Ukraine du Sud et de la fédération de Russie. Les fossiles de poissons-chats de cette région sont représentés par des membres appartenant à deux familles (Clariidae et Siluridae), deux genres (respectivement Heterobranchus et Silurus), et cinq espèces, dont une nouvelle espèce (Silurus spinosus n. sp.) et deux espèces décrites en nomenclature ouverte (Silurus cf. S. glanis, Silurus sp.). La présence de †Heterobranchus austriacus dans le Sarmatien récent de l'Ukraine du Sud peut s'expliquer par l'expansion de la répartition géographique de cette espèce d'Afrique en Europe pendant le Miocène récent (des restes plus anciens de ce taxon ont été documentés uniquement en Europe centrale et occidentale). La répartition géographique de Silurus soldatovi était plus étendue dans le passé comme l'indiquent de nombreuses découvertes de ses os dans des dépôts du Miocène récent et du Pliocène dans le sud de l'Europe. L'espèce actuelle Silurus glanis est apparue pendant le stade final du Miocène récent (Pontien) et a probablement supplanté S. soldatovi en raison de sa plus grande plasticité écologique (et en particulier sa tolérance des variations de température, cette espèce pouvant frayer dans des eaux plus froides). Une tendance évolutive intéressante a été observée et concerne le changement de taille, de forme et d'orientation des denticules du corps de l'épine de la nageoire pectorale dans différentes espèces du genre Silurus. Une telle tendance pourrait s'expliquer par une oligomérisation des structures homologues, mais cette question demanderait se propre étude et n'est donc pas abordée ici.

Mots-clés : Clariidae ; Siluridae ; nouvelle espèce ; Miocène récent ; Pliocène ; Pléistocène

Translator: Antoine Souron

Deutsche Zusammenfassung

In progress

Translator: Eva Gebauer

Arabic

Translator: Ashraf M.T. Elewa

 

 

TABLE 1. Stratigraphic distribution of investigated localities with catfish remains.

Epoch

Stage

Age, Ma

No. (Figure 1)

Investigated localities

Heterobranchus austriacus

Silurus spinosus n. sp.

Silurus soldatovi

Silurus glanis

Silurus cf. S. glanis

Silurus sp.

Pleistocene Middle 0.77 26 Semibalka 1            
Early 2.58 21 Kotlovina 3            
Pliocene Piacenzian 10 Cherevichne 2            
  24 Obukhovka 2            
3.60 23 Shirokino            
Zanclean 21 Kotlovina 1            
  20 Kamenskoe            
  25 Novokyivka            
  19 Novopetrovka            
5.33 18 Trudomirovka            
Late Miocene Pontain 22 Shkodova Gora            
  17 Nerubaiske            
  16 Vinogradovka 1            
7.25 15 Pontian Lectostratotype            
Maeotian 14 Novoukrainka 1            
  13 Egorovka 1            
  13 Egorovka 2            
  12 Verkhnya Krynitsya 2            
  11 Vasylivka 1            
9.88 10 Cherevichne 3            
Late Sarmatian s.l. 9 Novoelizavetovka 2            
  8 Kubanka 2            
  7 Lysa Gora 2            
  6 Otradovo            
  5 Palievo            
  4 Frunzovka 2            
  3 Lobkovo            
  2 Mykhailivka 2            
11.0 1 Popovo 3            

 

 

TABLE 2. ­­­Anatomical distribution of investigated late Cenozoic catfish remains.

Skeletal element

Heterobranchus
austriacus

Silurus
spinosus
n. sp.

Silurus
soldatovi

Silurus
glanis

Silurus cf.
S. glanis

Silurus sp.

Total

Parasphenoid - 1 - 3 1 - 5
Praemaxilla - 6 - - - 11 17
Quadrate - 6 5 1 - - 12
Articular - 2 31 1 - - 34
Dentary - 40 35 2 - 60 137
Isolated jaw teeth - - - 2 - 12 14
Cleithrum - 10 74 2 - 6 96
Pectoral spine 102 154 256 19 4 69 604
Weberian centrum - - 27 - - - 27
Centrum - - 20 2 9 1354 1385
Total 102 219 448 32 14 1512 2327
 

kovalchukKovalchuk, Oleksandr. Department of Paleontology, National Museum of Natural History, National Academy of Sciences of Ukraine, 15 Bogdan Khmelnitsky str., Kyiv 01601 Ukraine. This email address is being protected from spambots. You need JavaScript enabled to view it.

Dr Oleksandr Kovalchuk, of Kyiv, Ukraine, is a researcher and curator of Cenozoic fish paleontology in the Department of Palaeontology, National Museum of Natural History of the National Academy of Sciences of Ukraine, specialist in the field of palaeoichthyology. Scientific interests are related to morphology, taxonomy, historical faunistics and palaeoecology of Neogene and Anthropogene freshwater fishes of Southeastern Europe.

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ferrarisFerraris, Carl J. National Museum of Natural History, Smithsonian Institution, Washington; 545 NE Floral Place, Portland, Oregon 97232 U.S.A. This email address is being protected from spambots. You need JavaScript enabled to view it.

Dr Carl Ferraris, of Portland, Oregon, U.S.A., has been involved for more than 25 years in fish research, with an emphasis on revisionary studies of catfishes and the nomenclature of fishes. Among the publications authored are the Checklist of Catfishes (in Zootaxa), and the Species of Fishes (in the Catalog of Fishes), as well as papers redescribing or newly describing more than 100 species of catfishes. He was an editor of the Checklist of the Freshwater Fishes of South and Central America, and Checklist of the Freshwater Fishes of the Guiana Shield, and was a Co-Principal Investigator of the recently concluded Planetary Biodiversity Inventory (PBI), Phase I: All Catfish (Siluriformes) Species Inventory (2003-2009), funded by the U.S. National Science Foundation.

 

APPENDIX

Comparative material examined. All specimens are dried skeletons and c&s (cleared and stained) bones. Each number represents an individual specimen.

Silurus glanis: NMNHU-P 1/113, 114, 115, 117, 118, 119, 120, 121, 122, 123, 124, 125, 216, 218 (dried skeletons); NMNHU-P 1/126 (c&s).

Silurus soldatovi: PIN (comparative collection) - quadrate, articular, dentary, hyomandibular, epihyal, ceratohyal, cleithrum, pectoral spine, Weberian centrum (c&s).

Silurus asotus: PIN (comparative collection) - pectoral spine (c&s).

Additional data for comparison were obtained from literature:

Heterobranchus austriacus: pectoral spine (see Thenius, 1952; Weinfurter, 1954; Böhme, 2002).

Parasilurus altus: complete skeleton (see Sytchevskaya, 1989).

Silurus joergi: complete skeleton, isolated bones (see Gaudant, 2015).

Silurus cf. S. soldatovi: isolated bones (see Sytchevskaya, 1989).

Silurus aristotelis: skull, shoulder girdle, pectoral spine (see Kobayakawa, 1989).

Silurus biwaensis: pectoral spine (see Kobayakawa, 1989); cleithrum, praemaxilla, pectoral spine, Weberian centrum (see Kobayakawa and Okuyama, 1994).

Silurus cochinchinensis: skull, shoulder girdle, pectoral spine (see Kobayakawa, 1989).

Silurus glanis: pectoral spine (see Kobayakawa, 1989); skull (see Ünlü et al., 2012).

Silurus grahami: skull, pectoral spine (see Kobayakawa, 1989).

Silurus microdorsalis: skull, shoulder girdle, pectoral spine (see Kobayakawa, 1989).

Silurus triostegus: skull, pectoral spine (see Ünlü et al., 2012).

 

FIGURE 1. Investigated localities with late Miocene, Pliocene and Pleistocene catfish remains. 1, Popovo. 2, Mykhailivka. 3, Lobkovo. 4, Frunzovka. 5, Palievo. 6 , Otradovo. 7, Lysa Gora. 8 , Kubanka. 9, Novoelizavetovka. 10, Cherevichne. 11, Vasylivka. 12, Verkhnya Krynitsya. 13, Egorovka. 14, Novoukrainka. 15, Pontian lectostratotype. 16, Vinogradovka. 17, Nerubaiske. 18, Trudomirovka. 19, Novopetrovka. 20, Kamenskoe. 21, Kotlovina. 22 , Shkodova Gora. 23, Shirokino. 24, Obukhovka. 25, Novokyivka. 26, Semibalka.

 figure1

FIGURE 2. Isolated bones assigned to † Heterobranchus austriacus Thenius, 1952. 1-3, left pectoral-fin spine (NMNHU-P 33/570, Mykhailivka 2): dorsal view ( 1), ventral view ( 2), proximal articulation surface ( 3). 4-5, right pectoral-fin spine (NMNHU-P 41/3732, Kubanka 2): dorsal view ( 4), proximal articulation surface ( 5). 6-7, right pectoral-fin spine (NMNHU-P 41/2891, Frunzovka 2): dorsal view ( 6), proximal articulation surface ( 7). 8 , right pectoral-fin spine with wrong accrete fracture (NMNHU-P 33/574, Mykhailivka 2), lateral view. 9-10, right pectoral-fin spine (NMNHU-P 41/2908, Frunzovka 2): lateral view ( 9), proximal articulation surface ( 10). Scale bars equal 5 mm (Figure 2.1-2, 2.4, 2.6, 2.8-9) and 2 mm (Figure 2.3, 2.5, 2.7, 2.10).

 figure2

FIGURE 3. Isolated bones assigned to † Silurus spinosus n. sp. 1-5 , holotype, left pectoral-fin spine (NMNHU-P 29/1747, Popovo 3): dorsal view ( 1), ventral view ( 2), medial view ( 3), lateral view ( 4), proximal articulation surface ( 5). 6-7, distal fragments of pectoral-fin spines (NMNHU-P 29/4034-4035, Lobkovo). 8, medial part of parasphenoid (NMNHU-P 29/1756, Popovo 3), dorsal view. 9, right dentary (NMNHU-P 29/4002, Lobkovo), medial view. 10, symphysal fragment of cleithrum (NMNHU-P 29/1814, Popovo 3), lateral view. 11 , articular (NMNHU-P 29/1852, Popovo 3), lateral view. 12 , praemaxilla (NMNHU-P 29/1906, Popovo 3), ventral view. 13 , articular facet of quadrate (NMNHU-P 29/1911, Popovo 3), anterior view. Scale bars equal 10 mm.

 figure3

FIGURE 4. Isolated bones assigned to Silurus soldatovi Nikolsky and Soin, 1948. 1, left pectoral-fin spine (NMNHU-P 41/2894, Frunzovka 2), medial view. 2-4, right pectoral-fin spine (NMNHU-P 41/2454, Pontian lectostratotype): lateral view ( 2), medial view ( 3), proximal articulation surface ( 4). 5, left pectoral-fin spine (NMNHU-P 37/736, Novopetrovka), dorsal view. 6, left pectoral-fin spine (NMNHU-P 41/4432, Egorovka 2), dorsal view. 7, left pectoral-fin spine with wrong accrete fracture (NMNHU-P 41/4457, Egorovka 2), medial view. 8-9, right pectoral-fin spine (NMNHU-P 42/213, Kamenskoe): dorsal view ( 8), medial view ( 9). 10, distal part of pectoral-fin spine (NMNHU-P 42/254, Kamenskoe), ventral view. 11, distal part of pectoral-fin spine (NMNHU-P 53/4508, Novokyivka), dorsal view. 12-14, left pectoral-fin spine (NMNHU-P 53/4507, Novokyivka): dorsal view ( 12), lateral view ( 13), proximal articulation surface ( 14). 15, symphysal part of cleithrum (NMNHU-P 29/1223, Vasylivka 1), lateral view. 16, fragment of cleithrum (NMNHU-P 29/664, Verkhnya Krynitsya 2), lateral view. 17, symphysal part of cleithrum (NMNHU-P 41/4458, Egorovka 2), lateral view. 18, articular facet of quadrate (NMNHU-P 42/784, Kamenskoe), anterior view. 19, fragment of left dentary (NMNHU-P 29/1238, Vasylivka 1), lateral view. 20, fragment of right dentary (NMNHU-P 42/885, Kamenskoe), lateral view. 21, right cleithrum (NMNHU-P 53/4614, Novokyivka), lateral view. 22, left dentary (NMNHU-P 53/4610, Novokyivka), lateral view. 23, right articular (NMNHU-P 29/1237, Vasylivka 1), medial view. 24, right articular (NMNHU-P 53/4593, Novokyivka), medial view. 25-26, Weberian centrum (NMNHU-P 42/207, Kamenskoe), lateral view. 27, Weberian centrum (NMNHU-P 53/4580, Novokyivka), lateral view. 28, abdominal centrum (NMNHU-P 42/1249, Kamenskoe), anterior view. Scale bars equal 10 mm (Figure 4.1-5, 4.8-16, 4.18, 4.20-26) and 5 mm (Figure 4.6-7, 4.17, 4.19, 4.27-28).

figure4 

FIGURE 5. Isolated bones assigned to Silurus glanis Linnaeus, 1758. 1-4, right pectoral-fin spines (NMNHU-P 41/609, 621, Shkodova Gora): dorsal view ( 1), lateral view ( 2), proximal articulation surface ( 3), fragment of the shaft with denticulations ( 4). 5, isolated jaw tooth (NMNHU-P 41/2530, Kotlovina 1), lateral view. 6, fragment of left dentary (NMNHU-P 41/52, Shkodova Gora): dorsal view. 7-8, right articular (NMNHU-P 41/626, Shkodova Gora): lateral view ( 7dorsal view ( 8). 9-10, parasphenoid with basioccipital bone (NMNHU-P 41/717, Shkodova Gora): ventral view ( 9), dorsal view ( 10). 11, fragment of quadrate (NMNHU-P 41/623, Shkodova Gora), anterior view on the articular facet. 12, fragment of cleithrum (NMNHU-P 41/615, Shkodova Gora), lateral view. Scale bars equal 20 mm (Figure 5.9-10, 5.12), 10 mm (Figure 5.1-4, 5.6-8, 5.11) and 5 mm (Figure 5.5).

figure5 

FIGURE 6. Isolated bones assigned to Silurus cf. S. glanis Linnaeus, 1758. 1, centrum (NMNHU-P 41/2731, Vinogradovka 1). 2, centrum (NMNHU-P 53/4698, Nerubaiske). 3, centrum (NMNHU-P 41/1165, Shkodova Gora). 4, centrum (NMNHU-P 29/3973, Shirokino). 5, centrum (NMNHU-P 37/797, Trudomirovka). 6, centrum (NMNHU-P 45/4052, Cherevichne 2). All centra (Figure 6.1, 6.3-6 - abdominal, Figure 6.2 - caudal) are presented in anterior view. 7, fragment of parasphenoid (NMNHU-P 41/680, Shkodova Gora). Scale bars equal 10 mm.