Osteology of Russian Toads and Frogs for Paleontological Researches
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Acta zoologica cracoviensia, 44(1): 1-23, Krakow, 30 March, 2001 Osteology of Russian toads and frogs for paleontological researches V RATNIKO . Yu v Viatchesla Received: 24 May, 2000 Accepted for publication: 28 Dec., 2000 RATNIKOV V. Yu. 2001. Osteology of Russian toads and frogs for paleontological re- searches. Acta zoologica cracoviensia, 44(1): 1-23. Abstract. Osteological diagnosis of modern genera Bufo and Rana as well as species Bufo bufo, B. gargarizans, B. verrucosissimus, B. viridis, B. raddei, B. calamita, Rana tempo- raria, R. chensinensis, R. arvalis, R. asiatica, R. amurensis, R. ridibunda, R. lessonae and R. nigromaculata are given. The genera investigated are distinguished very easily, be- cause practically all the skeletal elements, with the exception of distal elements of the limbs, have their specific generic features. Bufonids have 9-11 bones used for species identification. The most important of them are ilium, frontoparietale, maxillare, and parasphenoideum. Ranids have 6-7 such bones, the most important of which are ilium, frontoparietale, and scapula. Russia , species n Moder Rana, Bufo, , Osteology : words y Ke Viatcheslav Yu. RATNIKOV, Geological Faculty, Voronezh State University, University Sq. 1, Voronezh, 394693 Russia. E-mail: [email protected] I. INTRODUCTION The geological researches of anhtropogenic deposits of the East-European platform conducted recently, have shown, that amphibian and reptile remains in them occur not so rarely as had been previously supposed. As a part of fossil fauna, they bear information concerning the geological past and can be used for the reconstruction of paleogeographical conditions and the determination of the geological age of bone-bearing deposits (RATNIKOV, 1995, 1996a). Most of the remains in quater- nary sediments belong to modern species, requiring their careful study for comparison with fossil materials. Paleontologists should also take into consideration the change of systematic status of some forms in connection with the new ideas of the neoherpetologists and finding osteological distinctions in newly described species. In the present paper the author brings criteria of identification of skeleton elements of some modern representatives of genera Bufo and Rana, which are often found in anthropogene deposits of Russia. He found these criteria when comparing the fossil remains with samples of osteological col- lection, which contains (sensu BORKIN, DAREVSKY, 1987) (the digits denote the number of speci- mens): Bufo bufo (LINNAEUS) -21, Bufo verrucosissimus PALLAS - 8, V. Yu. RATNIKOV 2 , 9 - R CANTO gargarizans Bufo Bufo viridis LAURENTI - 14, Bufo raddei STRAUCH - 12, , 3 - I LAURENT calamita Bufo Rana temporaria LINNAEUS - 10, , —11 D DAVI chensinensis Rana Rana arvalis NlLSSON - 37, , 6 - A BEDRIAG asiatica Rana , 6 - R BOULENGE amurensis Rana , 19 - S PALLA ridibunda Rana Rana lessonae CAMERANO - 8, Rana nigromaculata HALLO WELL - 6. g describin s article n i d mentione y rarel y ver e ar s bone g fro d an d toa f o s attribute c generi e Th fossil finds. Some information is contained in the works of BOHME, 1977; HOLMAN, 1998; RAGE, 1974, but it is not complete. Therefore the present author gives a criterion of the generic identifica- tion he used, and where possible avoids mention data from those papers. He used the terminology of the above-mentioned authors and given by SANCHIZ (1998). Acknowledgements. The author thanks Prof. Z. ROCEK for examination of e th g translatin n i p hel r he r fo A L.TlTOV o t l grateful o als s i e h d an s remark e valuabl d an r pape s thi work in to English. This study was possible thanks to L. BORKTN, A. MASALYKIN, and V. ORLOVA . collection l osteologica e th e complet o t r autho e th d helpe o wh II. FAMILY BUFONIDAE Genus Bufo LAURENTI, 1768 Condylus occipitalis is wide, with parallel lateral margins; the borders of fossa condyloidea practically coincide with those of foramen jugulare (Fig. 1 A, B). Small and large openings of the - Na . dimentions r simila e hav d an r othe h eac o t e clos y ver e ar l cana r semicircula r anterio t shor y ver sale are of subtriangular form or with a small S-shaped bend (Fig. 2 C-F). Pars lateralis ossis squa- n tha r wide s i i parasphenoide s Corpu . C-H) 4 . (Fig s zygomaticu s ramu o t e angl e acut n a t a s i mosi lateral processes, the length of the bone being less than the distance between the ends of lateral pro- cesses (Fig. 3 A-F). Two blades of the anterior pterygoid branch coincide in width along the greater s i e Gonial . A) 4 . (Fig e blad l ventra e th n o n projectio d angle-shape n a s i e ther d an , length s it f o t par comparatively high, with low processus coronoideus considerably deviated laterally; dorsal margin s i e bon e th f o t par r posterio ; process d coronoi a r unde s end h whic , crest a s a s proceed e bon e th f o abruptly curved in S manner, the lateral margin forms a visible obtuse angle; the ventral part of the bone under the coronoid process is considerably compressed but does not form a crest; channel for o tw n betwee e spac e som s i e Ther . O) 4 . (Fig e bon e th f o t par e middl e th n i s i e cartilag i Mekkel cotyles of atlas; neural arch in the upper part is very thick (Fig. 5 A-B). Vertebrae with large, often compressed bodies; the neural arch is massive in the upper part at breast vertebras and is flattened dorsally at abdominal ones, being almost in the same plane with diapophises; canalis neuralis of ab- dominal vertebrae is narrow especially at presacral vertebra (Fig. 5 D, E, H, I). Joint pits of urostyl - ven ; view r anterio n i e triangl l equilatera n a f o m for g havin , low s i l channe l neura ; separated t no e ar - widen e considerabl h wit y usuall , medially s widen Clavicula . E) 6 . (Fig x conve s i e surfac e bon l tra ing on the lateral end (Fig. 6 G-J). Cleithrum is of V-shape; the posterior margin of lamina marginalis cleithri, approaching margo scapularis, turns in caudal direction, forming subtriangular expansion of l.marginalis (Fig. 6 L). Pars acromialis and pars glenoidalis of scapula do not cover s frog d an s toad n Russia f o y Osteolog 4mm - prooti - E D- temporaria; Rana - C viridis, Bufo - B bufo, Bufo - A : view r posterio , exoocipital - C A- . bones l Crania . 1 . Fig cum, dorsal and ventral views: D - Rana temporaria, E - Rana ridibunda; F-L - maxillare, F-H, K, L - outer view, I-J - in- Bufo — K raddei, Bufo - } viridis, Bufo - I verrucasissimus, Bufo - H gargarizans, Bufo - G bufo, -F Bufo : view r ne - N bufo, Bufo - M : views ) anterior d an - P , M , (J l ventra , dorsal , sphenethmoideum - S M- arvalis; Rana - L calamita, temporaria. Rana - S calamita, Bufo - R raddei, Bufo - Q viridis, Bufo - P verrucosissimus, Bufo - O gargarizans, Bufo V. Yu. RATNIKOV , nasale - G C- ; view l dorsa temporaria, Rana - B ; view r anterio viridis, Bufo - A : praemaxillare - B A- . bones l Crania . 2 . Fig dorsal view Bufo- C : bufo, Bufo- D viridis, E—Bufo raddei, —F Bufo calamita, Rana- G temporaria; H-U frontoparie- - tale, dorsal view: H - Bufo bufo (fused with prooticum and exoccipitale), 1 - Bufo gargarizans (fused with prooticum), J - Bufo verrucosissimus, K - Bufo viridis (fused with prooticum), L - Bufo raddei (fused with prooticum and exoccipitale), M - Bufo calamita (fused with prooticum and exoccipitale), N - Rana temporaria, O - Rana chensinensis, P - Rana ar- valis, Q - Rana asiatica, R - Rana amurensis, S - Rana ridibunda, T — Rana lessonae, U - Rana nigromaculata. Osteology of Russian toads and frogs - E viridis, Bufo - D verrucosissimus, Bufo - C gargarizans, Bufo - B bufo, Bufo - A . view l ventra , Parasphenoideum . 3 . Fig Bufo calamita, Bufo- F raddei, Rana- G temporaria, Rana- H chensinensis, Rana- I arvalis, —J .Sara asiatica,K San- N anigromaculata. San- M alessonae, ffa«- L aridibunda, Xa«aamurensis, V. Yu. RATNIKOV - C : view r oute , squamosum - P C- arvalis; Rana — B bufo, Bufo - A : view r oute , pterygoideum - A-B . bones l Crania . 4 . Fig Bufo bufo, D - Bufo gargarizans, E — Bufo verrucosissimus, F — Bufo viridis, G - Bufo raddei, H — Bufo calamita, I — ridibunda, ^- f ana N amurensis, a Ran M— asiatica, Rana - L chensinensis, Rana - K temporaria, Rana - J arvalis, Rana arvalis. -R Rana bufo, Bufo - Q : views l interna d an l dorsa , goniale - R Q- nigromaculata; Rana - P lessonae, Rana - O Osteology of Russian toads and frogs - G D- temporaria; Rana - C viridis, Bufo - B bufo, Bufo - A : views r posterio d an r anterio , atlas - C A- . Vertebrae . 5 . Fig - D : views r anterio d an l ventra , dorsal - G , views l ventra d an l dorsa - F , views r anterio d an l dorsa - E D- , vertebra t breas Bufo bufo, E—Bufo viridis, F - Rana temporaria, G — Rana ridibunda; H-L — abdominal vertebra, H-I - dorsal and anterior -J Rana viridis, Bufo - I bufo, Bufo - H : views l ventra d an l dorsa - L K- , views r anterio d an l ventra , dorsal - J , views arvalis. Rana - L temporaria, Rana - K ridibunda, V. Yu. RATNIKOV M 4mm N Fig. 6. A-D - sacrum, A-B - dorsal and anterior views, C-D - dorsal view: A- Bufo bufo, B - Bufo viridis, C - Rana tempo- raria, D - Rana ridibunda; E-F - urostyl, anterior and lateral views: E - Bufo bufo, F - Rana temporaria; G-K - clavicula, internal view: G - Bufo gargarizans, H - Bufo viridis, I - Bufo raddei, J - Bufo calamita, K — Rana arvalis; L-M - clei- Rana - P O- gargarizans, Bufo - N : view r inne , coracoideum - P N- arvalis; Rana - M viridis, Bufo - L : view r inne , thrum temporaria: O - male, P - female.