Review of Mesozoic and Cenozoic Sea Turtles from the Former Ussr

Home , Adocus, Eosphargis, Glarichelys, Ordosemys, Psephophorus, Sinemys

Russian Journal of Herpetology Vol. 9, No. 2, 2002, pp. 137 – 154

REVIEW OF MESOZOIC AND CENOZOIC SEA TURTLES FROM THE FORMER USSR

Alexander O. Averianov1

Submitted December 17, 2001.

A list of 51 localities of fossil sea turtles from the Middle Jurassic, Early and Late Cretaceous, Paleocene, Eocene and Miocene of Russia, Uzbekistan, Kazakhstan, Tajikistan, and Ukraine is provided. For four localities in Tajikistan, Kazakhstan, and Ukraine presence of sea turtles is not confirmed by current data. Two new taxa, Dollochelys rogovichi sp.nov. (Cheloniidae, Eocene, Ukraine) and Turgaiscapha kushmurunica gen. et sp.nov. (Chelonioidea indet., cf. Dermochelyidae, Late Cretaceous, Kazakhstan), are described.

Key words: Cheloniidae, Chelonioidea, Dermochelyidae, Protostegidae, sea turtles, Thalassemydidae.

INTRODUCTION St. Petersburg, Russia; EIMIZ, Natural History Mu- seum named after Zardabi, Baku, Azerbaijan; IPG, Turtles entered the open sea several times since Institute of Paleobiology, Georgian Academy of Sci- the beginning of their geologic history. Pronounced ences, Tbilisi, Georgia; IZK, Institute of Zoology, adaptation to this new environment (fore limbs modi- Kazakhstan Academy of Science, Alma-Ata, Ka- fied to swimming, shell lightened by costoperipheral zakhstan; IZU, Institute of Zoology, Ukraine Acad- and plastral fontanels) are found in Late Jurassic Eu- emy of Science, Kiev, Ukraine; PIN, Paleontological ropean Eurysternidae and Early Cretaceous to Recent Institute, Russian Academy of Sciences, Moscow, globally distributed Chelonioidea (monophyletic Russia; ZISP PH (or ZISP PHT), Paleoherpetologi- group, sensu Gaffney and Meylan, 1988). This re- cal collection, Zoological Institute, Russian Acad- view includes all fossil sea turtles localities for the emy of Sciences, St. Petersburg, Russia. territory of the USSR, as known previously, as well as reported here for the first time. Majority of these localities confined to the areas of the Cretaceous- REVIEW OF THE MESOZOIC Paleogene Tethys Ocean, Cretaceous Turgai Straight, AND CENOZOIC SEA TURTLES epicontinental sea of the Russian Platform, and Neo- LOCALITIES gene Paratethys basins (Fig. 1; Table 1). In the list of localities references to all descriptions and illustra- Jurassic tions of the material are given. This review does not include some unpublished materials, currently stud- 1. Peski, Moscow Region, Russia. ied by Prof. V. B. Sukhanov, by Dr. A. A. Yarkov, Geology. Black clays of Moskva Reka Forma- and by the author. Yaxartemys longicauda Riabinin, tion, Meshera Horizon, filling fissures in the Carbon- 1948, known from the juvenile specimens only from iferous limestone. Late Jurassic of Kazakhstan, originally referred to Age. Middle Jurassic, Bajocian-Bathonian. Thalassemydidae (Riabinin, 1948), possibly belongs Material and references. “Bones of skeleton to the Xinjiangchelyidae, a group of Asiatic fresh- and shell fragments of a turtle” (Krupina, 1995:129). water and estuarine waters turtles (Kaznyshkin et al., New genus and species of “Triassic generation” tur- 1990), which are not considered here. tles (Novikov et al. 1998:58). A new genus of Cryp- Collection abbreviations. CCMGE, Cherny- todira “more archaic than the Early Jurassic Kayenta- shev’s Central Museum of Geological Exploration, chelys” (Alifanov and Sennikov, 2001:75). This is an archaic sea turtle, not closely related to other sea tur- 1 Zoological Institute of the Russian Academy of Sciences, Uni- tles, recently described as Heckerochelys romani Su- versitetskaya nab. 1, St. Petersburg 199034, Russia. khanov, 2001 and referred to a new family Heckero-

Fig. 1. Localities of the fossil sea turtles for the territory of the USSR. Localities: 1, Peski; 2, Motnya 1; 3, Khodzhakul’ 1; 4, Gubkin; 5, Sheikhdzheili 2; 6, Khodzhakul’sai; 7, Itemir; 8, Chukhonastovka; 9, Bagaevka; 10, Saratov; 11, Nogaity; 12, Polunino; 13, Alymtau; 14, Kyrkkuduk 2; 15, Malaya Serdoba; 16, Rychkovo; 17, Zhuravlevskii; 18, Kushmurun; 19, Malaya Ivanovka; 20, Karpovka; 21, Bakh- chisarai; 22, Vostochnyi; 23, Kol’tso; 24, Darbaza 1 and 2; 25, Dzhylga 1a; 26, Dzhylga 1b and 2; 27, Andarak 2; 28, Getkilnin; 29, Dzhe- roi 1; 30, Dzheroi 2; 31, Khanabad; 32, Maisk; 33, Kentau; 35, Bakhmutovka; 36, Krasnorechenskoe; 37, Kiev; 38, Pirogovo; 39, Koktur- nak; 40, Prolom; 43, Otradnaya; 44, Abadzekhskaya; 45, Tchelkar-Teniz; 47, Perekischkyul’; 48, Agyspe; 50, Belomechetskaya; 51, Dzaudzhikau. The remaining localities cannot be located with confidence. chelyidae (Sukhanov, 2001). Material in the PIN 3. Khodzhakul’ 1, Karakalpakistan, Uzbekistan. collection. Geology. Lower or middle part of the Khodzhakul’ Formation. Early Cretaceous Age. Early Cretaceous, late Albian (possibly, Late Cretaceous, early Cenomanian). 2. Motnya 1, Buryatiya, Russia. Geology. Lower part of Khilok Formation. Material and references. Basisphenoid, CCMGE 2/11478, the holotype of Oxemys gutta Age. Early Cretaceous, Berriasian?. Nessov, 1977 (Nessov, 1977b:47, Figs. V and G; Material and references. Two bridge peripherals (ZISP PHT B59 – 11) and part of the plastron 1987b, pl. 2, Fig. 6). “Oxemys gutta (Chelonioidea) (?hypoplastron; ZISP PHT B59-12) of “?Chelonio- (Nessov, 1977a:156). “Some Chelonioidea (genus idea fam., gen. et sp. indet.” (Nessov and Khosatzky, Oxemys) (Nessov and Khosatzky, 1977:158). “Oxe- 1981:77, pl. 4, Figs. 11 and 12). “Some poorly deter- mys gutta Ness. (Chelonioidea incertae sedis)” (Ne- mined forms (Chelonioidea?), small and middle- ssov, 1984:216). “Oxemys gutta Ness.” (Nessov, sized” (Nessov, 1984:216). “Chelonioidea (?) indet.” 1988b:96). “Turtles Chelonioidea...: Oxemys gutta” (Nessov, 1997:121, pl. 28, Figs. 13 and 14). (Nessov, 1997:135, pl. 27, Fig. 1). Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 139

Comments. The basisphenoid of O. gutta is TABLE 1. Taxa of the Sea Turtles Found in the Mesozoic and Ce- clearly of chelonioid type. The rostrum basisphenoi- nozoic of the Former USSR (Numbers of Localities Correspond to Those in the Text and Fig. 1). dale much reduced. The sella turcica obliterated. The Localities (* indicates foramina anterior canalis carotici interni lie close to- Taxon gether (separated by the 1.5 mm distance). The dor- a type locality) sum sellae high, not overhanging the sella turcica, Jurassic and separated from the latter by a bone space with a Heckerochelys romani 1* saggittal crest. The systematic position of this turtle Cretaceous remains enigmatic. Among thousands of turtle shell Oxemys gutta 3* fragments from the Khodzhakul’ locality none could cf. Oxemys sp. 5, 6 be attributed to a chelonioid. The turtle assemblage is Parathalassemys cava 7* dominated by macrobaenid-sinemydid (Kirgizemys, Chelonioidea indet. 2, 10, 12 14, 16, 17 Tienfucheloides) and trionychoid (Adocus, Fergane- Cheloniidae indet. 14 mys, Kizylkumemys, Trionychidae indet.) forms, all Dermochelyidae indet. 4 of which have quite distinct structure of basisphenoid Turgaiscapha kushmurunica 18* Protostegidae indet. 9, 11, 15 (known for trionychoids mentioned, for macrobae- Teguliscapha rossica 4* nids Hangaiemys, Ordosemys, Dracochelys, and si- Teguliscapha(?) sp. 8 nemydid Sinemys). Cenozoic Chelonioidea indet. 21–23, 28, 29, 33–36, 38, 39, 4. Gubkin and Staryi Oskol, 41, 42, 49, 50 Belgorod Region, Russia. Cheloniidae indet. 27, 30 32, 40, 43, 44, 48, 51 Geology. Phosphorite conglomerate within yel- Osteopyginae indet. 20 low and gray sands of the Sekmenov Formation, out- Tasbacka aldabergeni 24, 25*, 26 cropped in the ore quarries Lebedinskyi and Stoilen- cf. Tasbacka sp. 19 Dollochelys rogovichi 37* skii near the cities Gubkin and Staryi Oskol respec- Glarichelys gwinneri 45–47 tively. Age. Early – Late Cretaceous, late Albian — early Cenomanian. 1980 – 1994, by A. O. Averianov and his colleagues Material and references. “Sea turtles in 1991 – 2000. Chelosp[h]arginae” (Nessov, 1985:148). “Plastron, carapace and skull elements, dentary (with moder- Late Cretaceous ately elongated symphysis, without saggittal crest)... of small-sized, primitive, ... oldest known Protostegi- 5. Sheikhdzheili 2, Karakalpakistan, Uzbekistan. dae (Chelospharginae)” (Nessov, 1987a:77). “Chelo- Geology. Upper part of the Khodzhakul’ Forma- spharginae (Teguliscapha)” (Glickman et al., 1987: tion. 258). “Sea turtles of the genus Teguliscapha Nessov, Age. Late Cretaceous, early Cenomanian. 1987 [sic]... relatively advanced in lower jaw struc- Material and references. “Testudines indet.: cf. ture... but retained primitive humerus of thalassoid Oxemys sp.” (Nessov, 1997:139). type” (Nessov, 1987b:89, Fig. 4). “Chelosphargi- 6. Khodzhakul’sai, Karakalpakistan, Uzbekistan. nae... with narrow long shell, thalassoid humerus... Geology. Upper part of the Khodzhakul’ Forma- referred to a new genus Teguliscapha Nessov, tion. gen.nov.” (Nessov et al., 1988:130, Figs. 1(17), 2). Age. Late Cretaceous, early Cenomanian. “Turtles Protostegidae and other [sea turtles]” (Ne- Material and references. “Testudines indet.: cf. ssov, 1997:126). The frontal, parietal, opisthotic, Oxemys sp.” (Nessov, 1997:140). some other cranial fragments, dentary, a number of shell fragments of medium-sized Teguliscapha rossi- 7. Itemir, Bukhara Viloyat, Uzbekistan. ca Nessov, 1988 (Protostegidae, Chelospharginae) Geology. Kulbecke Formation. and much rarer shell fragments of a gigantic archaic Age. Early(?)-Late Cretaceous, late Albian? Dermochelyidae indet. The material (CCMGE and – early Cenomanian. ZISP PH collections) will be fully described by Ave- Material and references. “Thalassemydidae rianov and Danilov (in preparation). Material col- (?)” (Nessov, 1984:216). Imprints of carapace and lected by L. A. Nessov and his colleagues in plastron fragments, CCMGE 1/12086, the holotype 140 Alexander O. Averianov

Age. Late Cretaceous, Cenomanian. a Material and references. “Vertebra of Chelo- spharginae indet.” (Arkhangelskii, 1999:142, Fig. 1B). 10. Saratov (= Lysaya gora, = Zavokzal’noe ushchel’e), Saratov Region, Russia. b Geology. White or yellow quartz-glauconite sands. Age. Late Cretaceous, late Cenomanian. Material and references. “Sea forms [of turtles] (Chelonioidea)” (Nessov and Khosatzky, 1977:158; Chkhikvadze, 1983:29). “Carapace plate of the turtle from the family Protostegidae” (Arkhangelskii, 1998:12). “?Peripheral plate of Chelospharginae indet.” (Arkhangelskii, 1999:142, Fig. 1A). Two c fragmented right dentaries, from an immature and an adult specimens (ZISP PH 1 – 2/39, Fig. 2), apparently belonging to one species, and a possibly femur fragment. Material from the ZISP collection (Lysaya gora locality) collected by L. S. Glickman in 1949. Comments. On the adult dentary specimen posterior part of the symphysis is preserved. Symphysis d seems to be short, with some emargination to its midline on the dorsal surface, possible indicating presence of a saggittal crest. The triturating surface is narrow, flat or slightly convex. The cutting edge of the dentary is not sharp. The ventral (chin) shelf at the symphysis is wider than the dorsal one. The sulcus Fig. 2. Fragmented right dentaries of Chelonioidea indet., cartilaginis meckeli is relatively narrow. Generally ZISP PH 1/39, immature specimen in internal view (a) and these dentaries are similar with those in Chelosphar- / ZISP PH 2 39, adult specimen in internal (b), dorsal (c), and lat- gis advena (Hay, 1908) (Zangerl, 1953a: Fig. 21C, eral (d) views. Locality 10, Saratov (Lysaya gora); Cenomanian. Scale bar is 1 cm. D), Toxochelys latiremis Cope, 1873 and T. moore- villensis Zangerl, 1953 (Zangerl, 1953b: pls. 12, 13, 14) from the Late Cretaceous of North America. This of Parathalassemys cava Nessov, 1984 (Thalassemy- turtle is determined here as Chelonioidea indet. From didae) (Nessov and Krasovskaya, 1984:22, Fig. 4(9 the chelosphargine Teguliscapha rossica from the – 12)). “Testudines inc. sedis: Parathalassemys cava, Albian-Cenomanian of Belgorod Region (see item 4) turtles show similarities with Plesiochelyidae, previ- it differs by evidently shorter symphysis and possible ously were referred to Thalassemydidae” (Nessov, presence of a saggittal crest. 1997:137, pl. 28, Figs.4–7,11,pl.35,Fig. 3). 11. Nogaity, Aktyubinsk Region, Kazakhstan. Geology. Nogaity Formation. 8. Chukhonastovka, Volgograd Region, Russia. Age. Late Cretaceous, late Cenomanian. Age. Late Cretaceous, Cenomanian. Material and references. Centrum of the trunk Material and references. The costal and periph- vertebra of a large sized Protostegidae? indet., ZISP eral (ZISP PH 1/22 and 1/22) of an immature speci- PH 1/33 (Fig. 3). Material collected by V. I. Zhelez- mens of Teguliscapha(?) sp. (Protostegidae, Chelo- ko in 1970. spharginae) (Averianov and Yarkov, 2000:162, Fig. 1). Material collected by A. A. Yarkov in 1988. 12. Polunino 2, Ol’khov District, Volgograd Region, Russia. 9. Bagaevka, Saratov Region, Russia. Geology. Quartz-glauconite sands and sand- Geology. Material reworked from the Cenomani- stone. For locality description see Pervushov et al. an Melovatka Svita into the Turonian basal phos- (1999). phorite horizon. Age. Upper Cretaceous (Campanian). Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 141 a b a b

c c

Fig. 4. Neural of Chelonioidea indet., ZISP PH 1/34, in dorsal Fig. 3. Centrum of trunk vertebra of Protostegidae? indet., (a), ventral (b), and anterior (c) views. Locality 16, Rychkovo; ZISP PH 1/33, in dorsal (a), ventral (b), and anterior(?) (c) views. early Campanian. Scale bar is 1 cm. Locality 11, Nogaity, Nogaity Formation; late Cenomanian. Scale bar is 1 cm.

Material and references. “Lophochelyinae Material and references. The metapodial or indet. [and] Chelonioidea indet.” (Averianov and Ne- phalanx, right dentary fragment, and costal fragment ssov, 1995:68). “Turtles Toxochelyidae: Lophoche- of the Chelonioidea indet. (ZISP PH 3 – 5/22; Ave- lyinae indet.” (Nessov, 1997:113). rianov and Yarkov, 2000:162, Fig. 2). Material collected by A. A. Yarkov in 1984 – 1985. 15. Malaya Serdoba 1, Penza Region, Russia. 13. Alymtau, Chimkent Region, Kazakhstan. Geology. Phosphorite horizon (layer 2). Geology. Syuksyuk or Darbaza Formation. Age. Late Cretaceous, early Campanian. Age. Late Cretaceous, late Santonian Material and references. “Remains of a turtle – Campanian. from the family Protostegidae” (Arkhangelskii, Material and references. Posterior peripheral, 1998:13). “A propodium fragment of Chelosphargi- ZISP PHT T78-1. “Very large turtle from the super- nae indet.” (Arkhangelskii, 1999:143, Fig. 1V). family Chelonioidea” (Nessov, 1981:71). “Very large Chelonioidea (Desmatochelyidae?)” (Nessov, 16. Rychkovo, Volgograd Region, Russia. 1984:220). “Peripheral plate of Desmatochelyidae? ” Age. Late Cretaceous, early Campanian. (Nessov and Udovichenko, 1986: pl. 1, Fig. 13). Material and references. “Neural plate of a “Very large Desmatochelyidae(?)” (Nessov, 1987a: small-sized turtle from Lophochelyinae (Toxochely- 78). “Large-sized sea Desmatochelyidae” (Glickman idae)” (Nessov and Yarkov, 1993:45). “Sea turtles” et al., 1987:258). “Very large turtles from Chelonio- (Nessov, 1997:128). Material collected by L. A. Ne- idea (Desmatochelyidae?)” (Nessov, 1997:107). Ma- ssov and A. A. Yarkov in 1991. terial collected by L. A. Nessov in 1978. Comments. The neural (ZISP PH 1/34; Fig. 4) is 14. Kyrkkuduk 2, Chimkent Region, Kazakhstan. relatively narrow (length 21.3 mm, maximum width Geology. Gray sands of the lower part of Darba- 12.1; possible not laterally complete), keeled, with za Formation. smooth surface and without a horn sulcus. Referred Age. Late Cretaceous, early Campanian. here to as Chelonioidea indet. 142 Alexander O. Averianov

17. Zhuravlevskii, Kustanai Region, Kazakhstan. Material collected by N. I. Udovichenko in 1972 – Geology. Sandy clays with phosphorites in the 1979. lower part of Zhuravlevskii Formation. 23. Kol’tso, Osh Region, Kirghizia. Age. Late Cretaceous, late Campanian. Geology. White quartzite sands (“Sulyukta Material and references. “Remains of... sea tur- sands”) within the Bukhara beds. tles” (Levina et al., 1990:44). “Turtles Chelonioidea Age. Late Paleocene, Thanetian. indet.” (Nessov, 1997:112). Material and references. A few very incom- 18. Kushmurun, Kustanai Region, Kazakhstan. plete shell fragments of cheloniid(?) turtles. Material Geology. Gray gravelites and sands with phos- collected by A. O. Averianov in 1988. phorites in the lower part of Eginsai Formation. 24. Darbaza 1 and 2, Chimkent Region, Age. Late Cretaceous, late Campanian – ?early Kazakhstan. Maastrichtian. Geology. White and yellow quartz sands. Material and references. “Protostegidae” (Pri- Age. Late Paleocene, Thanetian. zemlin, 1993:34, 44). “Turtles Protostegidae (not Material and references. Rare and fragmented large sized)” (Nessov, 1997:114). Most materials in shell fragments of Tasbacka aldabergeni. Material ZISP collection collected by L. A. Nessov in 1992. collected by A. O. Averianov in 1989. Comments. Shell fragments of a cf. dermochelyid Turgaiscapha kushmurunica gen. et sp.nov., de- 25. Dzhylga 1a, Chimkent Region, Kazakhstan. scribed below. Geology. White quartz sands. Age. Latest Paleocene or, less probably, earliest Paleocene Eocene, Thanetian – ?Ypresian. Material and references. More than 1500 shell, 19. Malaya Ivanovka, Volgograd Region, Russia. cranial and postcranial elements, including one skull Geology. Yellowish sands with gravel. and one nearly complete carapace, of Tasbacka alda- Age. Late Paleocene, Thanetian. bergeni Nessov, 1987 (Cheloniidae, Eocheloniinae), Material and references. “Sea turtles, similar collections CCMGE 12175 and ZISP PH (Nessov, with Tasbacka aldabergeni Ness., but twice larger” 1986a:180, Figs. 1, 2; 1986b: Fig. 6; 1987a:81, (Nessov and Yarkov, 1989:80). Figs.1–3,pls. 7, 8; 1988b:98; Chkhikvadze, 1990: 8, 24; 1999:258; Averianov et al., 1993:81). Material 20. Karpovka, Volgograd Region, Russia. collected by L. A. Nessov and his colleagues in Geology. Green-gray quartz-glauconite sands 1977 – 1992, by A. O. Averianov in 1989. with phosphorites within Saratov beds. Age. Late Paleocene, Thanetian. Eocene Material and references. “Shell fragments of sea turtles (cheloniids)” (Efimov and Yarkov, 1993: 26. Dzhylga 1b and Dzhylga 2, Chimkent Region, 88). The fragmented right maxilla and premaxilla Kazakhstan. (ZISP PH 6/22) and some shell fragments (ZISP PH Geology. White quartz sands, level 2–3m 7-9/22) of Osteopyginae indet. (Cheloniidae) (Ave- stratigraphically higher than the Dzhylga 1a locality. rianov and Yarkov, 2000:164, Figs. 3a – e, 4). Mate- Age. Early Eocene, Ypresian. rial collected by A. A. Yarkov in 1990. Material and references. Cranial and shell fragments of Tasbacka aldabergeni (Fig. 5). Material 21. Bakhchisarai, Crimea, Ukraine. collected by A. O. Averianov in 1989 and 1996. Age. Late Paleocene, Thanetian. Material and references. “Bones of [sea] tur- 27. Andarak 2, Osh Region, Kirghizia. tles” (Nessov, 1987a:77; Chkhikvadze, 1990:5). Ma- Geology. Conglomerate with oyster shells within terial collected by N. I. Udovichenko. yellow sands of lower Alay beds. Age. Early Eocene, late Ypresian. 22. Vostochnyi, Osh Region, Kirghizia. Material and references. “Remains of sea tur- Geology. White quartz sands (layer 3; “Sulyukta tles” (Chkhikvadze, 1983:29). “Middle sized sea tur- sands”) within the Bukhara beds. tle Cheloniidae” (Chkhikvadze, 1984b:75). “Chelo- Age. Late Paleocene, Thanetian. niidae gen. indet.” (Chkhikvadze, 1990:6). The prox- Material and references. “Fragment of the free imal part of humerus (ZISP PH 3/3) and rare shell rib end of [sea] turtle” (Udovichenko, 1984:134). fragments of Cheloniinae gen. et sp. indet. (Danilov Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 143

a b Fig. 6. Neural of Cheloniidae indet., ZISP PH 1/35, in dorsal (a) and ventral (b) views. Locality 31, Khanabad, upper Turkestan Fig. 5. Nuchal of Tasbacka aldabergeni, ZISP PH 1/38, in dorsal beds; middle Eocene. Scale bar is 1 cm. (a) and ventral (b) views. Locality 26, Dzhylga 1b; early Eocene. Scale bar is 1 cm. Material and references. “Small-sized sea turtles” (Nessov, 1986c:37; 1987a:77; Chkhikvadze, and Averianov, 1997:43, Fig. 3). Material collected 1990:7). “Sea turtles from archaic Eochelyinae (1.5 by N. S. Shevyreva and V. Yu. Reshetov in 1975, by times larger than the Paleocene Tasbacka aldaberge- A. O. Averianov in 1988 – 1995. ni Ness.) and cf. Syllomiinae” (Nessov, 1988a:122). “Sea turtles of two genera” (Nessov, 1992a:35). “Eo- 28. Getkilnin, Kamchatka Region, Russia. chelyinae and syllomiinae(?) sea turtles” (Nessov, Geology. Sandstone of Kamchikvayam (= 1992b:474). “Two forms of sea turtles” (Panteleev “Kamchikskaya”) Formation. and Nessov, 1993:97). Material collected by L. A. Age. Early-middle Eocene. Nessov and A. O. Averianov in 1987, by A. O. Ave- Material and references. “Costal plate... of a ju- rianov and A. V. Panteleev in 1989, by D. Ward, venile specimen... of a sea turtle similar with or refer- C. King, and A. O. Averianov in 1997. able to Puppigerus kemperi [sic, l.c. for “camperi”]” Comments. The material is fully described by (Chkhikvadze, 1984a:46). “Costal plate of sea turtle, Averianov (in preparation). similar with Puppigerus Cope” (Nessov, 1987a:77). “Puppigerus sp.” (Chkhikvadze, 1990:7). “Costal 31. Khanabad, Khodzhent Region, Tajikistan. plate of a relatively small sea turtle, apparently Pup- Geology. Clays of upper Turkestan beds. pigerus sp. (Chkhikvadze, 1999:259). Material col- Age. Middle Eocene, Lutetian-Bartonian. lected by M. B. Efimov in 1979. Material and references. “Bones of small- Comments. The generic attribution could not be sized... sea turtles” (Nessov, 1987a:76; Chkhikvadze, made upon such a material. The material should be 1990:9). One complete neural, ZISP PH 1/35 (Fig. 6) referred to as Chelonioidea or Cheloniidae(?) indet. and a few very incomplete shell fragments. Material 29. Dzheroi 1, Bukhara Viloyat, Uzbekistan. collected by N. I. Udovichenko in 1981 and 1982. Geology. Phosphorite bed. Comments. The neural is a relatively thin plate Age. Middle Eocene, late Lutetian. (maximum thickness 6.5 mm, at the posterior end), Material and references. “Shell bones of... sea with deeply incised anterior margin. It differs from turtles” (Nessov et al., 1987:74). Material collected those in Tasbacka aldabergeni and Puppigerus cam- by L. A. Nessov in 1985. peri by considerably narrower posterior margin and referred here to as Cheloniidae indet. 30. Dzheroi 2, Bukhara Viloyat, Uzbekistan. Geology. Gray and yellow sands with phos- 32. Maisk, Tashkent Viloyat, Uzbekistan. phorites. Geology. Yellow sands. Age. Middle Eocene, late Lutetian. Age. Middle Eocene, Lutetian-Bartonian. 144 Alexander O. Averianov

chelyidae?). Material collected by N. I. Udovichenko in 1988. Comments. One even costal (ZISP PH 1/20; Fig. 7) is a thin and mediolaterally short plate with reduced vertebral scute, shallow scute sulci and sculpture of irregular ridges radiating from the intercostal sulcus. 35. Bakhmutovka, Lugansk Region, Ukraine. Geology. Yellowish sand and sandstone of Bu- chak(?) Formation, below phosphate horizon at the bottom of Kiev Formation. Age. Middle Eocene, Lutetian-Bartonian. Material and references. Shell fragments of Chelonioidea? indet. with sculptured surface. Mate- rial collected by N. I. Udovichenko in 1990. Fig. 7. Costal of Chelonioidea indet. (Dermochelyidae?), ZISP PH 1/20, in dorsal view. Unknown locality (34) in Lugansk 36. Krasnorechenskoe, Lugansk Region, Ukraine. Region; middle Eocene. Scale bar is 1 cm. Geology. Light colored sand of Buchak(?) For- mation, below phosphate horizon at the bottom of Kiev Formation. Material and references. “Cheloniidae” (Udo- Age. Middle Eocene, Lutetian-Bartonian. vichenko and Nessov, 1982:35). “Small-sized sea Material and references. Shell fragments of turtles, similar to those from the Dzhylga assem- Chelonioidea? indet. with sculptured surface. Collec- blage, but perhaps representing a distinct species” tion ZISP PH 19. Material collected by N. I. Udovi- (Nessov, 1987a:77; Chkhikvadze, 1990:8). Material chenko in 1989. collected by L. A. Nessov and N. I. Udovichenko in 1982. 37. Kiev, Kiev Region, Ukraine. Geology. Blue carbonate clays (= “Kiev marl,” = Comments. The frontal and parietal (collection “Spondylus clay”) within the Kiev Formation. CCMGE 12087) of this turtle were briefly described Age. Middle Eocene, Bartonian. and the last bone was figured (Udovichenko and Material and references. Some bones of sea tur- Nessov, 1987:170, Fig. 15). The parietal differs from tles were figured by A. S. Rogovich (1870: pl. 10, those of T. aldabergeni (Nessov, 1987a: Fig. 1) and Figs. 47, 48, 51, and possibly others), but misinter- Puppigerus camperi (Gray, 1831) (Moody, 1974, preted by him as bones of a fish “Hypsodon kiovien- pl. 3) by separated frontal and upper occipital scutes sis Rog.” “Small sea turtle” (Khosatzky, 1950:24). and very narrow upper occipital scute. It may belong “Specimen of an Eocene sea turtle” (Tarashchuk, to a distinct species of Tasbacka or related turtle, but 1971:56). “Cheloniidae” (Dubrovo and Kapelist, the material at hands (except cranial bones there are 1979:10). Shell fragments and phalanges of Puppige- two incomplete costals and undeterminably shell rus sp., collection IZU, without numbers, from Vysh- fragments) is insufficient for more complete determi- gorod in the vicinity of Kiev (Chkhikvadze, 1983:30, nation. Figs. 11 – 13; 1990:5; 1999:259). “Puppigerus” sp.” 33. Kentau, Chimkent Region, Kazakhstan. (Nessov, 1987a:82). “Sea turtles new to science” Age. Middle Eocene, Lutetian-Bartonian. (Averianov et al., 1990:4). Apparently, this locality Material and references. “Sea turtles” (Nessov, was in mind of I. G. Pidoplichko when he wrote “in 1992a:36). Material collected by L. A. Nessov in the Eocene deposits of Northern Ukraine remains of 1990. sea turtles were found” (Pidoplichko, 1961:90). The premaxilla, fragmented dentaries, ceratobranchiale I, 34. Unknown locality, Lugansk Region, Ukraine. and fourth(?) costal of Dollochelys rogovichi, sp. Geology. Below the marl horizon of Kiev Forma- nov., described below, in the ZISP PH collection. tion. Material from the ZISP PH collection collected by Age. Middle Eocene, Lutetian-Bartonian. A. S. Rogovich in 1872 – 1874. Material and references. Shell fragments of 38. Pirogovo, Kiev Region, Ukraine. Chelonioidea indet. with sculptured surface (Dermo- Geology. Upper part of Kiev Formation. Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 145

Age. Middle(?) Eocene, Bartonian(?). Oligocene Material and references. “Bones of small sized 44. Abadzekhskaya, Krasnodar Region, Russia. sea turtles” (Nessov, 1987a:76; Chkhikvadze, 1990: Geology. Gray clays of Khadum beds of the 5). “Shell fragment of sea(?) turtle [collection Maikop Formation. CCMGE 12087]” (Udovichenko and Nessov, 1987: 171). Material collected by N. I. Udovichenko in Age. Eocene? or Early Oligocene. 1971 – 1972. Material and references. “Sea turtle” (Chkhi- kvadze, 1983:30; 1999:259). Possibly “Eochely- 39. Kokturnak, Kzyl-Orda Region, Kazakhstan. [i]nae — relatively small sea turtles of Puppigerus Geology. Sands on the Kokturnak Peninsula, for- type” (Chkhikvadze, 1987:85). “Remains... of a sea mer northern coast of the Aral sea. turtle” (Chkhikvadze, 1990:5; 1999:259; Nessov, Age. Middle Eocene, (?)Bartonian. 1992a:39). Material collected by A. A. Svichenskaya Material and references. “Remains of in 1972. [Chelonioidea]” (Khosatzky, 1975:439; Chkhikvad- 45. Tchelkar-Teniz, Turgai Region, Kazakhstan. ze, 1983:29; 1990:8). “Remains of sea turtles” (Asla- Age. Middle Oligocene. nova et al., 1979:17). Material collected by L. S. Material and references. “Chelonia sp.” Glickman in 1960. (Chkhikvadze, 1969:145). “Remains of sea turtles” 40. Prolom (= Belogorsk), Crimea, Ukraine. (Chkhikvadze, 1983:29). Referred, with some reser- Geology. Phosphatic marl layer within thick vation, to Glarichelys gwinneri (Wegner, 1911) (As- sequence of nummulitic limestone of Bodrak(?) lanova et al., 1979:18; Chkhikvadze, 1983:31). Horizon. 46. Ashchiktaipak, Ural’sk Region, Kazakhstan. Age. Middle – ?Late Eocene, Bartonian – ?Pri- Geology. Solenaya (= “Solenovskii”) horizon. abonian. Age. Middle Oligocene. Material and references. “Bones of [sea] tur- Material and references. Hyoplastron of Glari- tles” (Nessov, 1987a:77; Chkhikvadze, 1990:5). “Re- chelys gwinneri (Wegner, 1911), IPG 6-13-1 (Asla- mains of... sea turtles” (Nessov, 1992:38). Shell frag- nova et al., 1979:18, Fig. 3; Chkhikvadze, 1990:7, ments of Cheloniidae indet. Collection ZISP PH 18. 25). “Sea turtles, possibly close or attributing to Gla- Material collected by N. I. Udovichenko in different richelis gwinneri” (Chkhikvadze, 1999:262). Mate- years, including 1994. rial collected by V. G. Pronin. 41. Unknown locality in the Crimea, Ukraine. 47. Perekishkyul’, Baku Region, Azerbaijan. Age. Eocene. Geology. Gray and brown clays in the upper part Material and references. “There was informa- (= Miatly-Mutsidakal horizon) of the Maikop For- tion about a specimen of sea turtle, nowadays, per- mation. haps, lost, from the Eocene of Crimea” (Khosatzky, Age. Middle – late? Oligocene. 1950:24). The reliability of this report cannot be Material and references. “Sea turtle from the tested now. family Cheloniidae” (Aslanova, 1965:100). “Islami- 42. Unknown locality in Ukraine. chelys azerbajdzanica Asl., gen. et sp.nov.” [the Age. Eocene. name is not available according to the ICZN Article Material and references. “Remains of sea tur- 7b, but not a nomen nudum, contra Chkhikvadze tles” (Khosatzky, 1975:440). (1977:226) and Aslanova et al.(1979:18)] (Aslanova, 1969:9). “Remains of sea turtles” (Khosatzky, 43. Otradnaya, Krasnodar Region, Russia. 1975:440; Nessov, 1992a:39). “Glarichelys gwinneri Geology. Chalk. (Wegner, 1911)” (Chkhikvadze, 1987:85). “Remains Age. Eocene?. of large sea turtles Glarichelys gwinneri (Chkhikvad- Material and references. “?Chelonia sp.” ze, 1999:262). Complete skull (EIMIZ 62/1), consid- (Chkhikvadze, 1977:226). “Nearly complete cara- erable portions of the shells and postcranial elements pace of Cheloniidae gen. et sp. indet.” (Chkhikvadze, of Glarichelys gwinneri (Wegner, 1911), collection 1983:39; Chkhikvadze, 1990:5), collection IPG, EIMIZ 62 (Aslanova et al., 1979:18, Figs. 1 and 2; mostly immersed in the rock. Possibly “Eochely- Chkhikvadze, 1983:31, Figs. 14 – 16; 1990:25; [i]nae — relatively small sea turtles of Puppigerus Chkhikvadze, 1990:5). Material collected by S. M. type” (Chkhikvadze, 1987:85). Aslanova in 1959 – 1963. 146 Alexander O. Averianov

48. Agyspe [= Akespe], Kzyl-Orda Region, va et al., 1979:17). “Chelonia caucasica Riabinin, Kazakhstan. 1929, which is a juvenile specimen of a sea turtle” Age. Late Oligocene. (Chkhikvadze, 1987:85). Nearly complete, but badly Material and references. Costal, two connected preserved skeleton of juvenile specimen of “Chelo- peripherals, and coracoid of Chelonia aralis Kho- nia” caucasica Riabinin, 1929” (Chkhikvadze, 1983: satzky, 1945, PIN, without numbers [collection 417?] 36). “Imprint of a juvenile sea turtle ... Chelonia cau- (Khosatzky, 1975:54, Figs. 1 and 2). “Remains of casica” (Chkhikvadze, 1999:257). Generic attribu- sea turtles” (Khosatzky, 1950:24). “Chelonia aralis tion is questionable (Sukhanov, 1964:422; Chkhi- Khos.” (Kuznetsov, 1963:10). “Remains of relatively kvadze, 1983:38). Material collected by V. P. Smir- large sea turtle Chelonia aralis Khos.” (Khosatzky, nov. 1975:440). “Chelonia” aralis Khosatzky, 1945” (Chkhikvadze, 1990:5, 25). List of Localities for Which Reports of Sea Comments. Generic attribution of this turtle is Turtles were not Confirmed by Further Data doubtful (Sukhanov, 1964:422), it may belongs to 1. Kansai, Khodzhent Region, Tajikistan. Carettinae or Chelydridae (Aslanova et al., 1979:17; Geology. Red sands of Yalovach Formation. Chkhikvadze, 1983:29; Chkhikvadze, 1999:262). Age. Late Cretaceous, early Santonian. Material collected by Yu. A. Orlov and Ya. M. Eglon Material, references and comments. “Obvi- in 1938. ously sea turtles” (Verzilin, 1966:21), or “typical sea turtles” (Verzilin, 1967:55) are absent from this lo- Uncertain Paleogene cality (Nessov, 1997:131) (these determinations have 49. Unknown locality in Middle Povolzhye, Russia. been probably based on Anatolemys remains). Age. Paleogene 2. Gusinyi Perelet, Pavlodar Region, Kazakhstan. Material and references. “Remains of sea tur- Geology. Pavlodar Formation. tles” (Khosatzky, 1975:440; Chkhikvadze, 1983:29). Age. Late Miocene, middle Turolian. Miocene Material and references. Fragment of costal plate (IZK R-654) of “one form of sea turtles of the 50. Belomechetskaya, Stavropol’ Region, Russia. order [sic] Cheloniidea [sic] (apparently, family Che- Geology. Deposits with marine mollusks Pectun- loniidae)” (Kuznetsov, 1958:69). Material collected culus(?), below the level with the Middle Miocene by V. V. Lavrov in 1954. (MN 6) Platybelodon mammal fauna. Comments. The costal is referred to Chelydrop- Age. Oligocene? or Miocene. sis kusnetzovi Chkhikvadze, 1985 (Chelydridae) (As- Material and references. “Fragment of costal or lanova et al., 1979:17; Gaiduchenko and Chkhikvad- plastron of Cheloniidae? ” (Alekperov, 1978:184, ze, 1985:117). Fig. 52b, g). Material collected by N. K. Vereschagin in 1950. 3. Odessa [= Novoelizavetovka], Comments. Determination by L. I. Khosatzky Odessa Region, Ukraine. influenced heavily by finding of this fossil in marine Geology. Pontian limestone. strata. The figures published are not adequate. Attri- Age. Late Miocene, late Turolian. bution of this fragment to a marine turtle remains to Material and references. “Cheloniidae” (Tara- be demonstrated. shchuk, 1971:57). “Chelonia sp.” (Dubrovo and Ka- 51. Dzaudzhikau (= Chernaya Rechka), pelist, 1979:55). Comments. This material is now referred to Che- Northern Osetia, Russia. lydropsis nopcsai (Szalai, 1934) (= Macrocephalo- Geology. Gray shales. chelys pontica Tarashchuk, 1960) (Chelydridae) (As- Age. Early Miocene, Burdigalian. lanova et al., 1979:17). Material and references. The imprint of the juvenile turtle skeleton, the holotype of Chelonia cau- 4. Kuchurgan, Odessa Region, Ukraine. casica Riabinin, 1929 (Riabinin, 1929:194, pl. 1). Geology. Kuchurgan gravel. “Remains of sea turtles” (Khosatzky, 1950:24). “Re- Age. Late Pliocene, early Ruscinian. mains of a distinct species of sea turtles Chelonia Material and references. “Sea turtles of the caucasica Riab.” (Khosatzky, 1975:440). “Chelonia family Cheloniidae” (Tarashchuk, 1971:57). caucasica” (Chkhikvadze, 1977:226). “Juvenile Comments. This material is referred to Chely- specimen of sea turtle Chelonia caucasica” (Aslano- dropsis nopcsai (Szalai, 1934) [= Macrocephaloche- Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 147 lys pontica Tarashchuk, 1960] (Chelydridae) (Asla- Composition. Four species: D. atlantica (Zan- nova et al., 1979:17). gerl, 1953), D. coatesi Weems, 1988, D. casieri, and D. rogovichi sp.nov. Distribution. Maastrichtian – late Paleocene of SYSTEMATIC PALEONTOLOGY eastern North America; early-middle Eocene and middle Oligocene of Europe. Superfamily Chelonioidea Baur, 1893 Dollochelys rogovichi sp.nov. Family Cheloniidae Gray, 1825 (Figs.8–10) Dollochelys Zangerl, 1971, new assignment Hypsodon kioviensis Rog. [partim]: Rogovich, 1870: pl. 10, Figs. 47, 48, and 51. Type species. D. casieri Zangerl, 1971. Anthracotherium alsaticum Cuv. [partim]: Rogo- Diagnosis [after Zangerl (1971) and Parris et al. vich, 1875a:36; Rogovich, 1875b: 46. (1986), emended]. Medium to large-sized sea turtles. Cheloniidae [indet.]: Dubrovo and Kapelist, Skull with primary palate. Dentary of very pro- 1979:10. nounced cutting device. Carapace elongated, width Puppigerus sp.: Chkhikvadze, 1983:30, Figs. 11 less than 90% of its length. Maximum width of cara- – 13; Chkhikvadze, 1990:5; Chkhikvadze, 1999:259. pace disc less than 60% of total carapace length. Cos- “Puppigerus” sp.: Nessov, 1987a:82. tonuchal fontanels absent. Costoperipheral fontanels Holotype. ZISP PH 1/36, right fragmented den- large. Antero-lateral peripherals very narrow. Pygal tary of an adult specimen (Fig. 8j – k). Kiev, Kiev as long or longer than wide. Plastral index relatively Formation; middle Eocene, Bartonian. low (about 37 – 47). Material. Holotype and ZISP PH 2/36, prema- Remarks. Originally Dollochelys was referred to xillae with anterior portion of left maxilla, ZISP PH “Toxochelyinae,” which is now considered as a para- 3, 4/36, left and right fragmented dentaries, ZISP PH phyletic group relative to Euchelonioidea sensu Gaff- 5/36, right first ceratobranchial arch, all possible ney and Meylan (1988). This influenced reconstruc- from one juvenile specimen (Fig. 8a – g, l), ZISP PH tion of the anterior carapace end, unknown for both 6/36, left fragmented dentary (possibly one specimen D. casieri and D. atlantica at that time, as having with the holotype) and ZISP PH 7/36, right fourth(?) costonuchal fontanels (Zangerl, 1971: pl. 2, Fig. 2), costal from adult specimens (Figs. 8h and 10); IZU, which was reproduced without reservation in M³y- without numbers, fragmented nuchal, middle cara- narski’s “Encyclopedia of paleoherpetology, Tes- pace fragment (parts of neurals 3 – 5, right and left tudines” (1976: Fig. 40). Actually, even juvenile costals 3 – 4), and left hypoplastron fragment, all specimens of D. atlantica (Parris et al., 1986:99) and from one juvenile specimen (Fig. 9). All specimens D. rogovichi sp.nov. (Chkhikvadze, 1983:31) lack from the same or close localities. costonuchal fontanels. This leaves no longer basis for Differential diagnosis. Differs from D. casieri attribution of Dollochelys to “Toxochelyinae”; it is by somewhat wider vertebral scutes and larger costo- best placed in the Cheloniidae, being together with peripheral fontaneles, from D. atlantica by larger Eochelone Dollo, 1903 the most archaic members of central fontanel on plastron and possibly by a consid- the group (primary or incipient secondary bone pal- erably space between hypoplastra, from D. coatesi by ate, relatively low plastral index). considerably wider vertebral scutes and thicker shell bones. The dentary fragment referred to Toxochelys at- Description. Large turtles, with the carapace lantica by Zangerl (1953b: pl. 15, Figs. 1, 2) is of length estimated as about 1 m for adults. The pre- crushing device characteristic for toxochelyids and maxillae fused. Its labial ridge is exceedingly sharp. apparently does not belong to Dollochelys, having The foramina praepalatina lie close to the anterior cutting jaws. end of the bone. Generally the premaxillae are similar The materials from the middle Oligocene of Ger- with those in Toxochelys latiremis Cope, 1873 (Gaff- many attributed to D. casieri (Karl, 1991) are similar ney, 1979: Fig. 197) and Tasbacka aldabergeni (Ne- to D. rogovichi sp.nov. by relatively narrow (medio- ssov, 1987a: Fig. 1) and relatively lower than pre- laterally short) costals, large costoperipheral fonta- maxillae in Eochelone brabantica Dollo, 1903 (Gaff- nels, but differs from the latter species and D. casieri ney, 1979: Fig. 209). The cutting edge of the dentary in the shape of dentary. This turtle may belongs to a is exceedingly sharp, nearly straight on the preserved new species of Dollochelys. fragments. The triturating surface is narrow and 148 Alexander O. Averianov

a b c e

l g

j h

k a–h l

Fig. 8. Cranial elements of Dollochelys rogovichi sp.nov., locality 37, Kiev, Kiev Formation; middle Eocene: a – c, ZISP PH 2/36, fused premaxillae with anterior portion of left maxilla in anterior (a), dorsal (b), and ventral (c) views; d, ZISP PH 3/36, left dentary of a juvenile specimen in internal view; e – g, ZISP PH 4/36, right dentary of a juvenile specimen in internal (e), dorsal ( f ), and lateral (g) views; h, ZISP PH 6/36, left dentary of an adult specimen in internal view; i – k, ZISP PH 1/36, holotype, right dentary of an adult specimen in internal (i), dorsal ( j ), and lateral (k) views; l, ZISP PH 5/36, right ceratobranchiale I in ventral view. Scale bars are 1 cm. short, concave. The ventral (chin) shelf at the sym- Fig. 2A). On the carapace costoperipheral fontanels physis is longer than the dorsal shelf. The adult den- are well developed. The nuchal is broad and short, taries twice larger than the juvenile, with a thin, with the rugose surface for the seventh cervical neu- weakly developed dorsal shelf and wider and less ral attachment on its ventral surface near the medial concave triturating surface. The sulcus cartilaginis border. The costonuchal fontanels absent. The neu- meckeli is relatively larger and goes closer to the rals elongated, six-sided, with short antero-lateral symphysis in juvenile dentaries. The first ceratobran- edges. The costals relatively short, which possibly in- chial arch (first branchial horn) is incomplete, it is dicate that carapace (or carapace disc only?) was bent considerably about at midline, as in Chelosphar- more narrow compared with other species of Dollo- gis advena (Zangerl, 1953a: Fig. 21E). Its distal chelys. The vertebral scutes wide, six-sided, occupy- portion is relatively longer than in Tasbacka aldaber- ing about 50% or more of the costal length. There is a geni (Nessov, 1987a: pl. 7, Fig. 5), but relatively weakly defined abdomino-femoral sulcus on the hy- shorter than in Toxochelys latiremis (Nicholls, 1988: poplastron. The central plastral fontanel is large, ap- Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 149 proximately 5 cm in wide in the juvenile specimen. a The hypoplastra probably did not contact, their medial projections quite a broad. The spine projection of the hypoplastron for fixing of the xiphiplastron is well developed. Distribution. Pelagic of the middle Eocene epicontinental sea on eastern Europe. Etymology. In memory of A. S. Rogovich.

Order Chelonioidea Family cf. Dermochelyidae Gray, 1825 Turgaiscapha gen.nov. Type species. Turgaiscapha kushmurunica sp.nov. Late Cretaceous, northern Kazakhstan. Differential diagnosis. Differs from Corsoche- lys Zangerl, 1960 by suture contact between pygal and second suprapygal, carapace sculptured, from Mesodermochelys Hirayama and Chitoku, 1996 by scute sulci on costals more developed, suprapygal much narrower [inferred from the pygal-suprapygal b suture], posterior notch on pygal, from Eosphargis Lydekker, 1889, Psephophorus v. Meyer, 1847, and Dermochelys Blainville, 1816 by scute sulci present, costals not reduced. Remarks. Turgaiscapha gen.nov. is similar with Dermochelyidae by scute sulci of shell rudimentary and similarity of the shell fragments preserved with those in primitive Cretaceous dermochelyids Corso- chelys haliniches Zangerl, 1960 and Mesodermoche- lys undulatus Hirayama and Chitoku, 1996. If Tur- gaiscapha gen.nov. belongs to Dermochelyidae, it Fig. 9. Hypothetical restoration of the shell of Dollochelys rogovichi sp.nov. juvenile specimen (IZU material), based on D. atlan- would be the most archaic dermochelyid known (size tica carapace [a, after Parris et al. (1986: Fig. 2)] and D. casieri small, shell ossification well developed, scute sulci plastron [b, after Zangerl (1971: pl. 3, Fig. A)]. Shaded area shows rudimentary to a lesser extent than in any other der- known fragments. Scale bar is 10 cm. mochelyid). The rarity of the Mesozoic Dermochely- idae, known so far from USA (two records) and Ja- pan and now probably from Kazakhstan may be ex- plained predominantly by their extremely pelagic mode of life (Hirayama, 1997; Parham and Stidham, 1999). Nessov (1997:144) indicated a toxochelyid cf. Porthochelys from the Kushmurun assemblage and noted, that remains of this turtle were possibly referred to as “Dermatemydidae” by Prizemlin (1993: 34, 44). I do not know upon which material this determination was based. In the materials from Kushmu- run available to me there are no elements which could be referred to a “toxochelyid” turtle. Probably this determination was based (partly?) on costal fragment, referred here to juvenile specimen of Turgai- Fig. 10. Right fourth(?) costal of Dollochelys rogovichi sp.nov., scapha kushmurunica gen. et sp.nov. adult specimen, ZISP PH 7/36, in dorsal view. Locality 37, Kiev, Composition. Type species only. Kiev Formation; middle Eocene. Scale bar is 1 cm. 150 Alexander O. Averianov

a b j

c k d

e f

g i h l

Fig. 11. Shell elements of Turgaiscapha kushmurunica, gen. et sp.nov., locality 18, Kushmurun, lower part of Eginsai Formation; Late Cretaceous, late Campanian – ?early Maastrichtian: a – h, ZISP PH 2 – 5/37, fragmented costals in ventral (a, d, f, h) and dorsal (b, c, e, g) views; i, ZISP PH 6/37, left(?) bridge peripheral in dorsal(?) view; j, ZISP PH 1/37, holotype, pygal in dorsal view; k, l, ZISP PH 7/37, proximal fragment of left xiphiplastron in ventral (k) and dorsal (l) views. Scale bar is 1 cm.

Distribution. As for the type species. Description. An anterior (second?) neural plate Etymology. From Turgai Straight and Greek is hexagonal, with deeply concave anterior edge and “scapha,” a boat. convex posterior edge. The posterolateral edge is twice longer than the anterolateral edge; both are Turgaiscapha kushmurunica sp.nov. meeting at an angle of ~145°. The neural is a flat Figs. 11, 12 plate, indicating a rather flat carapace. The dorsal Protostegidae [indet.]: Prizemlin, 1993:34, 44; surface is sculptured by reticulate ridges in the Nessov, 1997:114. centrum and posteriorly (accentuated by deep pits in Holotype. ZISP PH 1/37, pygal (Fig. 11j). the central part of the plate), and by radiating ridges Kushmurun, northern Kazakhstan. Lower part of in anterior part. There are no traces of horn sulci. The Eginsai Formation. Late Cretaceous, late Campanian maximum length of the plate is 88 mm, maximum – ?early Maastrichtian. width is 57.5 mm. Material. Holotype and ZISP PH 2-5/37, frag- Due to the weak development of horn sulci, the mented costals of different size (Fig. 11a–h), ZISP position of the known costals within the carapace PH 6/37, left(?) bridge peripheral (Fig. 11i), ZISP cannot be determined. The rib head is distinctly more PH 7/37, proximal fragment of left xiphiplastron anteroposteriorly compressed in smaller costals (Fig. 11k, l). All specimens from the same locality. (ZISP PH 4, 5/37), compared with larger costals Tentatively assigned specimen. ZISP PH 8/37, (ZISP PH 2, 3/37). On the ZISP PH 3/37 specimen anterior neural (Fig. 12). the dorsal surface is sculptured by a net of irregular Diagnosis. As for the genus. notches. On other costals the dorsal sculpture is pres- Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 151 ent by very fine pits and furrows, radiating from the horn sulcus. The medial edge in ZISP PH 3/37 specimen is almost straight, indicating possibly absence of a neural plate medial to it. If so, this costal may belong to the posterior portion of the carapace. The bridge peripheral is 94 mm long, with medial margin broken off (the rib pit is not preserved). The intermarginal sulcus is weak, but visible on both dorsal and ventral surfaces. The lateral margin is slightly incised at the intermarginal sulcus. The sculpture is fine striations radiating from the intermarginal sulcus. The pygal is 67 mm long and 84 mm wide, of tra- a b pezoid shape. It is most thick at the suprapygal-pygal Fig. 12. An anterior (second?) neural tentatively assigned to Tur- suture and thinning and laterally widening towards gaiscapha kushmurunica, gen. et sp.nov., ZISP PH 8/37 in dorsal the caudal end. The cranial margin is convex dorsal- (a) and ventral (b) views. Locality 18, Kushmurun, lower part of Eginsai Formation; Late Cretaceous, late Campanian – ?early Ma- ly. There are three distinct facets on the cranial mar- astrichtian. Scale bar is 1 cm. gin, all appears to represent sutural contact with the suprapygal. There is a small posterior notch and short posterior sulcus along the midline of the dorsal and ventral surfaces (somewhat longer on the ventral sur- REFERENCES face). On the dorsal surface there are very fine striations directed towards the pygal-peripherals suturae. Alekperov A. M. (1978), Amphibians and Reptiles of The proximal portion of the xiphiplastron is nar- Azerbaijan, Elm, Baku [in Russian]. Alifanov V. R. and Sennikov A. G. (2001), “About dis- row, tapering and laterally bending proximally. On covery of dinosaur remains in Moscow region,” Dokl. the dorsal (visceral) surface there is a deep cleft for RAN, 376, 73 – 75 [in Russian]. the spine projection of the hypoplastron. On the ven- Arkhangelskii M. S. (1998), Sea Reptiles of the Mesozoic tral (outer) surface, close to the proximal end, there is of Saratov Volga Region, Its Stratigraphic and Bio- a straight and weak femoroanal sulcus. Both dorsal nomic Significance. Author’s Abstract of Candidate’s and ventral surfaces covered by fine longitudinal Thesis, Saratov [in Russian]. striations. Arkhangelskii M. S. (1999), “On the findings of turtle remains in the Upper Cretaceous of Lower Volga region,” Remarks. ZISP PH 8/37 is assigned to Turgai- Trudy NII Geol. Saratovsk. Gos. Univ. Nov. Ser., 1, scapha kushmurunica, gen. et sp.nov. because of the 142 – 144 [in Russian]. same provenance and similarity in the sculpture with Aslanova S. M. (1965), “Some data about Perekischkul’ some of the costal plates, referred to that taxon. In Maikop fauna of vertebrates,” Uch. Zap. Azerb. Gos. Mesodermochelys, Corsochelys, and Eosphargis Univ., 2, 99 – 101 [in Russian]. neurals are rectangular, rather than hexagonal (Hira- Aslanova S. M. (1969), Perekischkul’ Fauna of Higher Vertebrates from Maikop Svita. Author’s Abstract of yama, 1996:610). Hexagonal neurals, if really belong Candidate’s Thesis, Baku [in Russian]. to Turgaiscapha gen.nov., could be another primitive Aslanova S. M., Pronin V. G. and Chkhikvadze V. M. character of this taxon. (1979), “Remains of sea turtles from the Oligocene de- Distribution. Pelagic of the Late Cretaceous posits of Aphsheron and northern Ustyurt,” Vest. Zool., (Campanian – ?Maastrichtian) epicontinental sea on 6, 17 – 22 [in Russian]. Averianov A. O. and Nessov L. A. (1995), “A new Creta- Central Asia (Kazakhstan). ceous mammal from the Campanian of Kazakhstan,” Etymology. From Kushmurun locality. Neues Jahrb. Geol. Palaontol. Monatsch., 2,65–74. Averianov A. O., Nessov L. A., and Udovichenko N. I. Acknowledgments. The author is grateful to I. G. Da- (1993), “Late Paleocene complex of bony fishes and nilov and J. Parham for reading the manuscript and provid- other vertebrates from locality Dzhylga in southern Ka- ing useful comments, to I. G. Danilov also for help with zakhstan,” Mater. Ist. Fauny Flory Kazakhstana, 12, preparation of Figs. 11 and 12. 79 – 91 [in Russian]. 152 Alexander O. Averianov

Averianov A. O. and Yarkov A. A. (2000), “Some turtles gene deposits of Pavlodar Priirtyshye,” Geol. Geofiz., remains from the Cretaceous and Paleogene of Volgo- 1, 116 – 118 [in Russian]. grad Region, Russia,” Russ. J. Herpetol., 7(2), 161 – Glickman L. S., Mertinene R. A., Nessov L. A., Rozhde- 166. stvenskii A. K., Khosatzky L. I., and Yakovlev V. N. Averianov A. O., Potapova O. R., and Nessov L. A. (1987), “Vertebrates,” in: M. M. Moskvin (ed.), Strati- (1990), “On original native bone finds of ancient grafiya SSSR. Melovaya sistema (Stratigraphy of the birds,” Trudy Zool. Inst. AN SSSR, 210, 3–9 [in USSR. The Cretaceous System). Semivol. 2, Nedra, Russian]. Moscow, pp. 255 – 262 [in Russian]. Chkhikvadze V. M. (1969), “New turtles from the Paleo- Hirayama R. (1997), “Distribution and diversity of Creta- gene deposits of Zaisan Depression and Central Ka- ceous chelonioids,” in: J. M. Callaway and E. L. Nicholls zakhstan,” Byull. Mosk. Obshch. Ispyt. Prirody. Otdel (eds.), Ancient Marine Reptiles, Acad. Press, San Geol., 44(6), 145 – 146 [in Russian]. Diego, pp. 225 – 241. Chkhikvadze V. M. (1977), “Review of knowledge on Efimov M. B. and Yarkov A. A. (1993), “A Paleocene fossil turtles of the Northern Caucasus,” in: I. S. Darev- crocodile from the lower reaches of the Volga basin,” skii (ed.), Voprosy Gerpetologii (Questions of Herpe- Paleontol. Zh., 2, 87 – 91 [in Russian]. tology). Vol. 4, Nauka, Leningrad, pp. 226 – 227 [in Karl H.-V. (1991), “Die toxochelyiden Seeschildkröten Russian]. (Chelonioidea, Toxochelyidae) von Sachsen,” Mauri- Chkhikvadze V. M. (1983), Fossil Turtles of Caucasus tiana, 13, 233 – 245. and Northern Black Seaside Region, Tbilisi [in Kaznyshkin M. N., Nalbandyan L. A., and Nessov L. A. Russian]. (1990), “Turtles of middle and late Jurassic of Fergana Chkhikvadze V. M. (1984a), “First record of a fossil turtle (Kirghiz SSR),” Ezhegod. Vsesoyuz. Paleontol. on Kamchatka,” in: Vyd i Ego Produktivnost’ v Areale Obshch., 33, 185 – 204 [in Russian]. (Species and Its Productivity in Its Areal), Sverdlovsk, Khosatzky L. I. (1945), “Finding of sea turtle remains in pp. 46 – 47 [in Russian]. the Oligocene deposits of Priaral’ye,” Dokl. AN SSSR, Chkhikvadze (Tchikvadze) V. M. (1984b), “A new spe- 49, 53 – 55 [in Russian]. cies of land tortoise from the middle Eocene of Fer- Khosatzky L. I. (1950), “Paleontological and stratigraphi- gana,” Paleontol. Sb., 21, 74 – 78 [in Russian]. cal significance of the fossil turtles,” Vopr. Paleontol., Chkhikvadze V. M. (1987), “Recent notion on fossil am- 1, 20 – 31 [in Russian]. phibians and reptiles from the territory of Northern Khosatzky L. I. (1975), “Amphibians and reptiles,” in: Caucasus,” in: Problemy Regional’noi Fauny i Ekologii V. A. Grossheim and I. A. Korobkov (eds.), Stratigrafia Zhivotnykh (Problems of Regional Fauna and Ecology SSSR. Paleogenovaya Sistema (Stratigraphy of the of Animals), Stavropol, pp. 84 – 90 [in Russian]. USSR. The Paleogene System). Semivol. 2, Nedra, Chkhikvadze V. M. (1990), Paleogene Turtles of the Moscow, pp. 439 – 441 [in Russian]. USSR, Tbilisi [in Russian]. Krupina N. I. (1995), “The first finding of Jurassic cerato- Chkhikvadze V. M. (1999), “The history of the develop- dontids (Dipnoi) from the Moscow Region,” Paleontol. ment of the Paleogene herpetofauna of the former Zh., 1, 129 – 131 [in Russian]. Soviet Union territory,” in: Problemy Paleobiologii Kuznetsov V. V. (1958), “Sea turtle from Neogene of the (Problems of Paleobiology). Vol. 1, Tbilisi, pp. 256 Pavlodar Priirtyshye,” Mater. Ist. Fauny Flory Kazakh- – 279 [in Russian]. stana, 2, 69 – 71 [in Russian]. Danilov I. G. and Averianov A. O. (1997), “New data on Kuznetsov V. V. (1963), Turtles of Paleogene and Neo- the turtles from the early Eocene of Kirghizia,” Russ. J. gene of Turgai and northern Priaral’ye. Author’s Ab- Herpetol., 4(1), 40 – 45. stract of Candidate’s Thesis, Alma-Ata [in Russian]. Dubrovo I. A. and Kapelist K. V. (1979), Catalogue of Levina A. P., Naidin D. P., Ponomarenko Z. K. and the Tertiary Vertebrate Localities of the Ukrainian SSR, Paskar’ Z. S. (1990), “Outcrops along the Ayat River,” Moscow [in Russian]. in: G. N. Papulov, V. I. Zhelezko, and A. P. Levi- Gaffney E. S. (1979), “Comparative cranial morphology na (eds.), Verkhnemelovye Otlozheniya Yuzhnogo Za- of Recent and fossil turtles,” Bull. Am. Mus. Nat. Hist., ural’ya (Raion Verkhnego Pritobol’ya)[Upper Creta- 164, 65 – 376. ceous Deposits of the South Transurals (Region of Up- Gaffney E. S. and Meylan P. A. (1988), “A phylogeny of per Priob’e)], Sverdlovsk, pp. 26 – 46 [in Russian]. turtles,” in: Benton M. J. (ed.), The Phylogeny and Mùynarski M. (1976), Encyclopedia of Paleoherpetology. Classification of the Tetrapods. Vol. 1. Amphibians, Testudines, Stuttgart – New York. Reptiles, Birds, Clarendon Press, Oxford, pp. 157 – Moody R. T. J. (1974), “The taxonomy and morphology 219. of Puppigerus camperi (Gray), an Eocene sea-turtle Gaiduchenko L. L. and Chkhikvadze V. M. (1985), from northern Europe,” Bull. Brit. Mus. (Nat. Hist.). “First discovery of cayman turtle remains in the Neo- Geol., 25, 153 – 186. Review of Mesozoic and Cenozoic Sea Turtles from the Former USSR 153

Nessov L. A. (1977a), “Turtles and some other reptiles Nessov L. A. (1992a), “Review of localities and remains from the Cretaceous of Karakalpakia,” in: I. S. Darev- of Mesozoic and Paleogene birds of the USSR and the skii (ed.), Voprosy Gerpetologii (Questions of Herpe- description of new findings,” Russ. Ornitol. Zh., 1(1), tology). Vol. 4, Nauka, Leningrad, pp. 155 – 156 [in 7 – 50 [in Russian]. Russian]. Nessov L. A. (1992b), “Mesozoic and Paleogene birds of Nessov L. A. (1977b), “On some cranial features of two the USSR and their paleoenvironments,” in: K. E. Camp- late Cretaceous turtles,” Vestn. LGU. Ser. Biol., 21, bell, Jr. (ed.), Papers in Avian Paleontology Honoring 45 – 48 [in Russian]. Pierce Brodkorb, Los Angeles, pp. 465 – 478. Nessov L. A. (1981), “On a turtle of the family Dermate- Nessov L. A. (1997), Cretaceous Nonmarine Vertebrates mydidae from the Cretaceous of Amur River basin and of Northern Eurasia, St. Petersburg [in Russian]. some rare records of remains of fossil turtles of Asia,” Nessov L. A. and Khosatzky L. I. (1977), “Turtles of Me- in: L. Ya. Borkin (ed.), Gerpetologicheskie Issledova- sozoic of the USSR,” in: I. S. Darevskii (ed.), Voprosy niya v Sibiri i na Dal’nem Vostoke (Herpetological Gerpetologii (Questions of Herpetology). Vol. 4, Invetigations in Siberia and Far East), Zool. Inst. AN Nauka, Leningrad, pp. 157 – 159 [in Russian]. SSSR, Leningrad, pp. 69 – 73 [in Russian]. Nessov L. A. and Khosatzky L. I. (1981), “Early Creta- Nessov L. A. (1984), “Data on late Mesozoic turtles from ceous turtles of the Baical Lake region,” in: L. Ya. Bor- the USSR,” Stvdia Geol. Salmantic., 2(1) (Stvdia Pa- kin (ed.), Gerpetologicheskie Issledovaniya v Sibiri i na laeochelonol., 1), 215 – 223. Dal’nem Vostoke (Herpetological Investigations in Si- Nessov L. A. (1985), “New data on vertebrates of Late beria and the Far East), Zool. Inst. AN SSSR, Lenin- Mesozoic of the USSR,” in: I. S. Darevskii (ed.), Vo- grad, pp. 74 – 78 [in Russian]. prosy Gerpetologii (Questions of Herpetology). Vol. 6, Nessov L. A. and Krasovskaya T. B. (1984), “Changes in Zool. Inst. AN SSSR, Leningrad, pp. 148 – 149 [in the composition of Cretaceous turtle assemblages in Russian]. Middle Asia,” Vestn. LGU. Ser. Biol., 3, 15–25[in Russian]. Nessov L. A. (1986a), “On the levels of organization and phylogenetic branches in the evolution of sea turtles,” Nessov L. A., Mertinene R. A., Golovneva L. B., et al. in: E. I. Vorob’eva and N. S. Lebedkina (eds.), Mor- (1988), “New findings of ancient organisms in Belgo- phology and Evolution of Animals, Nauka, Moscow, rod and Kursk Regions,” in: V. S. Ipatov, L. A. Kiri- pp. 179 – 186 [in Russian]. kova, and V. I. Vasilevich (eds.), Kompleksnye Issledo- vaniya Biogeotsenozov Lesostepnykh Dubrav (Complex Nessov L. A. (1986b), “Some late Mesozoic and Paleo- Investigations of Forest-Steppe Oak-Trees Biogeo- cene turtles of Soviet Middle Asia,” Stvdia Geol. Sal- cenoses), Leningrad, pp. 124 – 131 [in Russian]. mantic., 2(1) (Stvdia Palaeochelonol., 2), 7 – 22. Nessov L. A. and Udovichenko N. I. (1986), “New find- Nessov L. A. (1986c), “The first finding of late Cretaceous ings of remains of Cretaceous and Paleogene verte- bird Ichthyornis in Old World and some other bird brates of Middle Asia,” Vopr. Paleontol., 9, 129 – 136 bones from Cretaceous and Paleogene of Soviet Middle [in Russian]. Asia,” Trudy Zool. Inst. AN SSSR, 147, 31–38[in Nessov L. A., Shabanina N. V., and Udovichenko N. I. Russian]. (1987), “New localities of Vertebrate remains in the Nessov L. A. (1987a), “About sea turtles of the Paleogene middle Eocene phosphorites of Central Kzylkum and of southern Kazakhstan and phylogenetic relations of conditions of the granular phosphorites formation,” Toxochelyidae and Cheloniidae,” Paleontol. Zh., 4, Uzb. Geol. Zh., 4, 72 – 77 [in Russian]. 76 – 87 [in Russian]. Nessov L. A. and Yarkov A. A. (1989), “New Cretace- Nessov L. A. (1987b), “On some Mesozoic turtles of the ous-Paleogene birds of the USSR and some remarks on Soviet Union, Mongolia and China, with comments on the origin and evolution of the class Aves,” Trudy Zool. systematics,” Stvdia Geol. Salmantic., 2(4) (Stvdia Pa- Inst. RAN, 197, 78 – 97 [in Russian]. laeochelonol., 2), 87 – 102. Nessov L. A. and Yarkov A. A. (1993), “Hesperornithi- Nessov L. A. (1988a), “New Cretaceous and Paleogene forms in Russia,” Russ. Ornitol. Zh., 2(1), 37 – 54 [in birds of Soviet Middle Asia and Kazachstan and envi- Russian]. ronments,” Trudy Zool. Inst. AN SSSR, 182, 116 – 123 Nicholls E. L. (1988), “New material of Toxochelys latire- [in Russian]. mis Cope, and a revision of the genus Toxochelys (Tes- Nessov L. A. (1988b), “Late Mesozoic and Paleocene ver- tudines, Chelonioidea),” J. Vert. Paleontol., 8, 181 – tebrate assemblages of central Asia,” in: T. N. Bogda- 187. nova and M. V. Oshurkova (eds.), Stanovlenie i Evolu- Novikov I. V., Lebedev O. A., and Alifanov V. R. (1998), tsiya Kontinental’nykh Biot (Formation and Evolution “New Mesozoic vertebrate fossil sites of Russia,” in: of the Continental Biotas). Proc. of the 31 meeting All- J. W. M. Jagt, P. H. Lambers, E. W. A. Mulder, and Union Paleontol. Soc., Nauka, Leningrad, pp. 93 – 101 A. S. Schulp (eds.), Third Eur. Workshop on Vertebrate [in Russian]. Palaeontol. Maastricht, May6–9, 1998,p.58. 154 Alexander O. Averianov

Panteleev A. V. and Nessov L. A. (1993), “A small tubu- N. N. Iordansky, S. L. Kuzmin, and V. F. Orlova (eds.), nare (Aves: Procellariiformes) from the Eocene of Mid- Voprosy Gerpetologii (Questions of Herpetology), Pu- dle Asia,” Trudy Zool. Inst. RAN, 252, 95 – 103 [in shchino – Moscow, pp. 282 – 284 [in Russian]. Russian]. Tarashchuk V. I. (1971), “Turtles of Neogene and Anthro- Parham J. F. and Stidham T. A. (1999), “Late Creta- pogene deposits in the Ukraine. Communication 1. ceous sea turtles from the Chico Formation of Califor- Family of large-headed turtles (Platysternidae),” Vest. nia,” PaleoBios, 19(3),1–7. Zool., 2, 56 – 62 [in Russian]. Parris D. C., DeTample C., and Benton R. C. (1986), Udovichenko N. I. (1984), “About age of the lower part of “Osteological notes on the fossil turtle ?Dollochelys at- Paleogene deposits in South-Western Fergana by paleo- lantica (Zangerl),” The Mosasaur, 3, 97 – 108. ichthyological data,” in: Paleontologiya i Stratigrafiya Pidoplichko I. G. (1961), “Study of ancient vertebrates of Fanerozoya Ukrainy (Paleontology and Stratigraphy Ukraine for 40 years,” in: A. I. Tolmachev (ed.), Sorok of the Phanerozoic Ukraine), Kiev, pp. 133 – 136 [in Let Sovetskoi Paleontologii (Forty Years of the Soviet Russian]. Paleontology). Proc. of the 4th meeting All-Union Pa- Udovichenko N. I. and Nessov L. A. (1982), “Paleogene leontol. Soc., Gosgeoltekhizdat, Moscow, pp. 84 – 91 sharks and ray assemblage of the Tashkent region and [in Russian]. its temporaries equivalents in Ukraine,” in: Novye Dan- Prizemlin B. V. (1993), “Late Cretaceous complex of rep- nye po Stratigrafii Kainozoya Ukrainy I Nekotorykh tiles from ‘Priozernyi’ quarry (northern Kazakhstan),” Drugikh Territorii (k V sessii Ukrainskogo Paleontolo- Mater. Ist. Fauny Flory Kazakhstana, 12, 30–45[in gicheskogo Obschestva [New Data on Stratigraphy of Russian]. the Cenosoic of Ukraine and Some Other Territories (to Riabinin A. N. (1929), “Sea turtle from the Paleogene of the Fifth Session of the Ukrainian Paleontological So- the Vladikavkaz vicinity,” Zap. Ross. Mineral. Obshch. ciety)], Kiev, pp. 35 – 36 [in Russian]. Ser. 2, 58, 193 – 199 [in Russian]. Udovichenko N. I. and Nessov L. A. (1987), “Compari- Riabinin A. N. (1948), “Turtle from the Jurassic of Kara- son of chondrichthyan and other vertebrates assem- tau,” Trudy Paleontol. Inst., 15, 94 – 98 [in Russian]. blages from the Paleogene of Tashkent Chuli (Maisk) Rogovich A. S. (1870), “Fossil fishes of the Kiev Tertiary and Ukraine (Pirogovo),” in: O. S. Vyalov (ed.), Bio- basin and adjacent formations,” in: G. E. Tshurovskii stratigrafiya, Paleontologiya Osadochnogo Chekhla (ed.), Trans. of the Seond Congr. Of Russ. Naturalists Ukrainy (Biostratigraphy and Paleontology of the onthe Sect. of Mineralogy, Geology, and Paleontology, Depositional Cover of Ukraine), Naukova dumka, Moscow, pp. 19 – 54 [in Russian]. Kiev, pp. 167 – 174 [in Russian]. Rogovich A. S. (1875a), “Note on the localities of fossil Verzilin N. N. (1966), “About possibility to use some pa- mammal animal bones in the southwestern Russia,” leogeographic objective laws for investigation of the Zap. Kievsk. Obsch. Estestvoispyt., 4, 33–45 [in general conditions of continental deposits formation,” Russian]. Vestn. LGU. Ser. Geol. Geogr., 12, 16 – 22 [in Russian]. Rogovich A. S. (1875b), “Investigation of the brown coal Verzilin N. N. (1967), “Cretaceous deposits of the south of formation in the Kiev Province,” Zap. Kievsk. Obsch. Fergana Depression and their oil-and-gas content,” Estestvoispyt., 4, 46 – 49 [in Russian]. Trudy Leningr. Obshch. Estestvoispyt., 71, 1 – 138 [in Sukhanov V. B. (1964), “Subclass Testudinata. Testudi- Russian]. nates,” in: A. A. Rozhdestvenskii and L. P. Tatarinov Zangerl R. (1953a), “The vertebrate fauna of the Selma (eds.), Osnovy Paleontologii. Zemnovodnye, Presmyka- Formation of Alabama. Part 3. The turtles of the family yushchiesya i Ptitsy (Fundamentals of Paleontology. Protostegidae,” Fieldiana. Geol. Mem., 3, 61 – 133. Amphibians, Reptiles, Birds), Nauka, Moscow, pp. 354 Zangerl R. (1953b), “The vertebrate fauna of the Selma – 438 [in Russian]. Formation of Alabama. Part 4. The turtles of the family Sukhanov V. B. (2001), “An archaic turtle from the Toxochelyidae,” Fieldiana. Geol. Mem., 3, 135 – 277. Middle Jurassic of Moskovskaya Province and its posi- Zangerl R. (1971), “Two toxochelyid sea turtles from the tion in basal radiation of the order Testudines,” in: Landenian sands of Erquelinnes (Hainaut) of Belgium,” N. B. Ananjeva, I. S. Darevsky, E. A. Dunayev, Mem. Inst. Roy. Sci. Nat. Belg., 169,1–32.