Mesozoic marine reptiles of Russia and other former Soviet republics
GLENN W. S'I'OKKS, hf;\XIM S. ;\RKII;\N<;EL'SKII, .AN11 VL:\I>IMIR hl. EFIhfO\'
Introduction liptian). The phosphorites typically produce only fragmentary and disarticulated remains, but complete Marine reptile remains have often been fbund in the skeletons are common in the clays, shales, and marls of extensive Mesozoic epicontinental marine units of the the Volga. There is good potential for discoveries in former Soviet Union, and in particular, the Russian other quarries, but these are currently unmonitored. Platform of the European part of Russia. These fossils Llistorical localities of the Moscow Region generally include relatively common plesiosaurs, ichthyosaurs no longer exist, as these quarries have been al~andoned and mosasaurs, and rarer crocodilians and turtles. The over time. The best potential for immediate results Moscow Basin and the Ul'yanovsk, Samara, and therefore lies with the excellent cliff-face exposures Saratov regions of the Volga River Basin have lxen along the Volga. particularly productive. These areas not only contain Numerous Russian reports refer to localities in large exposures of UpperJurassic and Cretaceous sed- 'Povolzh'e' or 'Zavolzh'e'. These are Russian conven- iments, but have proved relatively accessible to tions for the Volga River Basin, Povolzh'e indicating Russian workers over the years. Sadly, there has l~een the right bank of the river (Ul'yanovsk and Saratov little in-depth sti~dyand analysis of marine reptile shore; west bank), and Zavolzh'e the left (Samara fossils in the former Soviet Union in recent years, shore; east bank). In the central Volga Basin near although a great many historical works have been Ul'yanovsk, marine reptile remains have been found in devoted to them. In this article, we review the current all ammonite zones from the Callovian to the .l\lbian. state of knowledge of these interesting fossils and The greatest concentrations, however, occur in the provide a lmis for informed future study. Middle Volgian Dovsop1~zuitc.spnndcvi, Epiziivgntites niki- The quality of most Russian marine reptile lholo- tint, and Upper Volgian Cvnsprditcs subditrs ammonite types is poor and most specimens have been recovered zones and in the Hauterivian Spertonicev~zsvevsicolov as float fiorn the Volga and Moscow River Basins, or as Zone. Uniforrn formational or 'svita' (suite) names chance occurrences in quarries and oil shale mines. have not been established generally for the Russian Almost no deliberate excavations have been under- Platform marine rocks, and the use of local ammonite taken. Currently, the most productive Russian local- zones is preferred. The Russian Volgian essentially ities for plesiosaurs and ichthyosaurs are the equals the 'rithonian. phosphorite quarries near Voskresensk in the Moscow Lower Volga sediments producing ichthyosaurs, Region, quarries near the village of Sundokovo in plesiosaurs, mosasaurs, and turtles in the Saratov and Tataria, the vicinity of Kashpir in the Samara Region lJenza Regions (Povolzh'e) are Senonian in age, typi- (UpperJurassic to Aptian), the 30 km of Volga River cally Campanian and Maastrichtian. 'l'he precincts of shoreline from the village of Kriushi to Mordovo the Serdoha River near the village of Malaya Serdoba, (Barremian to illhian), and the 24 km stretch of the Penza, have been particularly rich in fragmentary Volga from Ul'yanovsk to Undory (Kimmerid~'nail to remains of Campano-Maastrichtian age (pers. obs., G.W. STORRS t~al.
M.S.A.). Jurassic and Cretaceous marine deposits con- (1 947), Meniler (1948), Novozhilov (1948a, b, 1964), taining fossil reptile remains are also found in some of and Ochev (197ha, 1977), for example, have written on the central Asian nations that were formerly republics Russian plesiosaur remains more recently. Little study of the Soviet Union, but these areas have been less has been attempted in the past 20 years. Older workers well studied. Kazakhstan, Uzbekistan, and Azerbaijan (e.g. Bogolyubov, 191 1; Menner, 1948) sometimes are notable examples (Bazhanov, 1958; relied upon disassociated collections of bones for their Rozhdestvenskii, 1973; Nesov and Krasovskaya, 1984; type series. Where possible, holotypes in these cases Glikman etal., 1987). have been selected on the basis of page priority. Bogolyubov (1911) named numerous Russian ple- siosaur species all of which, however, are indetermi- Repository abbreviation^ nate and must be considered nomina dubia (l'able 11.1), KGU, Geology and Mineralogy Museum, Kazan State even though some of these were retained by Universit): Kazan'; MGRI, Moscow Geological Pravoslavlev (1915) (see Welles, 1962). Indeed, virtu- Prospecting Institute, Vernadskii State Geological ally all of the 'species' unique to Russia are nomina Museum, Moscow; PIN, Paleontological Institute, dubia, and not a single complete skeleton has been Russian Academy of Sciences, Moscow; PMK, described. Most are based upon isolated vertebrae and Pugachev Regional Museum, Pugachev, Saratov teeth and as such are non-diagnostic below the subor- region; POKM, Penza Regional Local History dinal, or perhaps the familial level. Most, if not all, of Museum, Penza, Penza Region; SGU, Paleontology Bogolyubov's (191 1) holotypes, from the old museum Museum of the Department of Historical Geology of the Geological Cabinet of Moscow University, are and Paleontology, Saratov State University, Saratov; now housed in MGRI. Some of these types have been 'Simbirtsit', Paleontological collection of the identified (Table 11.1), but through neglect while 'Simbirtsit' Industrial Works (a free enterprise under communist authority, parts of the collection are company), All-Russian Cultural Fund, Undory, inaccessible. It is believed that a planned renovation Ul'yanovsk Region; TsGM, Central Geological project, now underway, will uncover the remaining Museum, St. Petersburg; UPM, Undory specimens. Kipriyanov's (1 882, 1883) material is pre- Palaeontological Musuem, Undory, Ul'yanovsk sumably in the museum of the Academy of Sciences, region; ZIN, Zoological Institute, Russian Academy of St. Petersburg. The whereabouts of other collections, Sciences, St. Petersburg. such as those of Eichwald and Fischer von Waldheim, are unknown at present. Ostensibly, the genera Cimoliasaurus, Colymbosaurus, Systematic survey Cryptoclidus, Elasmosaurus, Eretmosaurus, Georgiasaurus, DIAPSIDA Osborn, 1903 Leutkesaurus, Liopleurodon, Muraenosaurus, Neoplio- SAUROPTERYGIA Owen, 1860 saurus, Peloneu.rte.r, Plesiosaurus, Pliosaurus, Polycotylus, PLESIOSAURIA de Blainville, 1835 Polyptychodon, 'Rhino.raurus: Scanisaurus, Simolestes, Comments. Over two dozen plesiosaur species have Spondylosaurus, Strongylokrotaphus, and Thaumatosaurus been named, and numerous additional taxa recog- are present in Russian rocks (Welles, 1962; Persson, nized, from Russian sediments by Russian and 1963; Novozhilov, 1964). However, most generic German workers. Most of this effort was undertaken in identifications and assignments to previously known the pre-revolutionary years of the nineteenth and Western taxa have been based upon stratigraphical, early twentieth centuries (e.g. Fischer von Waldheim, rather than morphological information, with an his- 1845, 1846; Eichwald, 1865-1868; Kipriyanov, 1883; torical readiness to name new species based upon geo- Ryabinin, 1909, 1915; Bogolyubov, 191 1; Pravoslavlev, graphic occurrence. Few of these identifications can 1915, 1916). Zhuravlev (1941, 1913), Rozhdestvenskii be considered reliable. Indeed, most Western species Table 1 1.1. Compilation of plesiosauvian taxa based upon mateiial fi-om the fivmev Soviet CJnion.Holotypes indicated by vepositovy abbveviation ovspecime?~numbev, where known
Taxon Holotype Material Locality Horizon Status
Colymbosauvus sklerodivus Bogolyubov, 19 11 MGRI fragmentary skeleton Moscow Region Volgian Plesiosauria indet. Cryptoclidus simbivskensis Bogolyubov, 1909 hlGRI vertebrae/limb frags. Ul'yanovsk Region Callovian-Oxfordian Plesiosauria indet. Elastnosaurus amalitskii Pravoslavlev, 19 16 - vertebral series Don Region Turonian Elasmosauridae indet. Elasmosaurus antiquus Dubeikovskii & Ochev, SGU 104a/17,18, 19 cervical centra Kama River Basin Hauterivian Elasmosauridae 1967 indet. Elasmosauvus kuvskennsis Bogolyubov, 191 1 MGRI [Kipr., 18821 med. cervical centrum Kursk Region Cenomanian Plesiosauria indet. [see Welles, 19621 Elasmosaurus ovskensis Bogolyubov, 19 11 lZIGRI cervical centra Orenburg Region Senonian Elasmosauridae indet. Elasmosaarus sevdobensis Bogolyubov, 19 11 hIGRI ant. cervical centrum Penza Region Cainpanian Elasinosauridae indet. ?Elasmosauvussachalinensis Ryabinin, 19 15 - phalanx Sakhalin Island Lower Senonian Plesiosauria indet. Eretmosauvus vzasnickii hfenner, 1918 hfGR1 VI 6111 cervical centrum Vilyui River, Siberia IvIiddleJurassic Plesiosauria indet. ?Evetmosauvus,jakowlewihfenner, 1918 MGRIVI61/15 caudal centrum Vilyui River, Siberia hliddle Jurassic Plesiosauria indet. Georgiasauvus (Geovgia)penzensis (Ochev, POKlLI No. 11658 partial skull/skel. Penza Region Santonian presumed valid 1976a) Leutkesauvus [no sp.] Kipriyanov, 1883 ZIN? teeth and vertebrae Kursk Region Cenomanian ? Polypvchodon sp. Muraenosauvus elasmosauvoides Bogolyubov, MGRI cervical centrum hfoscow Region Volgian ?Elasmosauridae 1911 indet. .I?uraetzosauvus kamerzsis Dubeikovskii & SGU 104al16 [lost] cervical vertebrae Kaina River Basin Volgian ?Elasmosauridae Ochev, 1967 indet. lMurnenosauvuspuvbecki Bogolyubov, 191 1 hfGRI centrum $loscow Region Volgian Plesiosauria indet. RTeopliosauvus [no sp.] Sintzov, 1899 - vertebraelhumeri Penza Region Senonian ?Polycotylidae indet. Plesiosaurus helmevserzi Kipriyanov, 1882 ZIN cervical centrum Penza Region Campanian Elasmosauridae (emend. Bog. '1 1) indet. [see Persson, 19591 propodial fragment Criinea Neocomian Ichthyosauria indet. [see Rvabinin, 1946bl Pliosaurusgiganteus Trautschold, 1860 - tooth bloscow Region Oxfordian L iopleuvodotz ? fivox P1ro.inuru.r vossicus Novozhilov, 1918a PIN 301 partial skull/skel Chuvashia Volgian Liopleuvodon vossicus [see Halstead, 19711
I Mesozoic marine reptiles
of 'Plesiosaurus' suffer from the same problem (Storrs, the remainder beiilg saved only hy chance. Halstead 1996) while Cimoliasarirus is apparently a nondiagnos- (1971) described a number of additional hones as able plesiosaur (?elasmosaur) and 'waste-basket taxon' belonging to the holotype, but inadequate collectioil (Williston, 1903). Fischer von Waldheirn's (1847) management practices have seemingly allowed confu- Rbinos~iurr~s,described as a plesiosaur, was based upon sion of at least some of the hones of this specimen a Permian anthracosaur skull, the origin of which was with those of 'Stro~~~lokrota~)buiirgi.r~?z~i.r (discussed perhaps misinterpreted through confusion within the below). Ochev (pers. cornin., 199.5) insists that only the containing collection, and is no\\, known as skull and pectrum of 'P.' rossicus were recovered. Rhinosauriscus (Kabanov, I 959; Rozhdestvenskii, 1973). Novozhilov (1948a, b) described only the skull, and Rarer plesiosaur remains are known from some of the later (Novozhilov, 1964) figured the pectrum. l'he other former Soviet republics hut most, if not all, of short dorsal blade and anteroventral rarnus of the these are also generically indeterminate. scapula suggest that the animal was immature. Few species of Russian plesiosaur are known from Halstead (1 9 71 ) considered 'P.' ro.rsicus to represent material of adequate quality to justify their retention Liopleurodo?z largely on the basis of its short mandibn- ('Table 11.1). Only three of these potentially distinct lar symphysis. It has, however, trihedral tooth cross- species are unique to Russia. The three include the sections unlike the Oxfordian Lioplrurodo?~firox, but best Uussiail specimens and are known from partial similar to Pliosaurus, a genus with a significantly longer skulls, although they have been described only in a syrnphysis. 'l'arlo's (1 959) Kimmeridgian genus with preliminary fashion, and a comprehensive review of trihedrally sectioned teeth and a short symphysis, each is required. Stretosaurus, was based upon incorrectly identihed material and is hence invalid, much of the referred PLI OSiZU UI DL%ESeeley, 1874a material probably belonging to Liopleurodo??(flalstead, Liopleurodon Sauvage, 1 873 1989). Pending new reviews of P1ios~iuru.r and Diagnosis. Large pliosaur distinguished from other I,iopleurodo?z, Halstead's interpretation of 1,. rossicus is forms by relatively short and straight-sided inandibu- accepted here. Points worth noting are that the bar lar symphysis hearing 5-7 pairs of teeth; dorsal aspect between the naris and orbit is not as narrow as is sug- of symphysis tapers anteriorly to blunt V. This diag- gested in the drawings in Novozhilov (194Ha, 1964) nosis conforins to the concept of I,ioplezirodo?2in Tarlo and the naris itself is much smaller than shown. (1960), although the genus is founded upon a single I;urthermore, contrary to Novozhilov (I 948a), there is tooth (Sauvage, 1873) that may prove problematic. no nasal bone or lacrimal. IIalstead (1971) also equated the enormous rostrum Liopkurodo?~vossicus (Novozhi lov, 1948a) of 'Plios~iuru.rcf.P gr~i?zdii(PIN 2440/1), described hy See Figure 1 1.1. Uozhdestvenskii (1047) fiom the left L'olga hank Holo{ype and locality. PIN 304, most of a skull and a (Zavolzh'e), Ozink Mine, Saratov Province, D.pa?ideri partial pectrum (scapulae and coracoid), with perhaps Zone, with L. vo.rsicu.r. This specimen too, was discov- a few other elements associated; Buinsk Mine oil ered complete, but only the rostrum, the proximal end shales, Ibresi District, right bank of Volga (Povolzh'e), of a humerus, a phalanx, and some rib frag~nentswere Autonomous Republic of Chuvashia, Russia. saved from the mining operations (Rozhdestvenskii, Horizon. Middle Volgian (Dorsopla?zite.rpandcrtZone). 1947). The complete hind limb noted by Halstead Comments. Originally described as 'Pliosaurus' ros.ricus (1971) undoubtedly belongs to 'Stro?zgylokrot~ipbus' Novozhilov, 1948a, this is a large, short-necked animal (PIN 426), as figured hy Zhuravlev (1943) and typical of the Pliosauridae. lMost of what was once a Novozhilov (1964). 'l'he ma~ldihular symphysis of complete skeleton was destroyed in 1938 during the PIN 244011 contains approxin~atelysix sharply trihe- process of mining the oil shales in which it was found, dral teeth per ralnus. G.W. STORRS etal.
Figure 11.1. Liopleurodon (Pliosaurus) rossicus(Novozhilov, 1948a), PIN 30411, from the middle Volgian oil shales of Buinsk Mine (DorsoplanitespanderiZone), Ibresi District, Chuvashia. Skull and mandible in dorsal (A), right lateral (B), and left lateral (C) views. Scale bars, 100 mm. Compare with Novozhilov (1964).
Pliosaurus Owen, 1842 Holotype and locality. PIN 426, a large partial skull, a Diagnosis. Very large pliosaurid up to approximately partial vertebral column, and an articulated hind limb; 10 m, distinguished by long mandibular symphysis oil shales of Savel'evsk Mine No. 1, near Gorny, about and trihedrally sectioned tooth crowns; approximately 35 km southwest of Pugachev, eastern Saratov 11 pairs of caniniform teeth within symphysis; labial Province, Russia. edges of mandibular symphysis approximately par- Horzzon. Savel'evsk oil shales, Volgian. allel. Comments. Most commonly known in Russian palae- ontological circles as 'Strongylokrotapbus' irgisensis, this Pliosaurus irgisensis (Novozhilov, 1948a) fossil was found during mining operations in 1933 See Figure 11.2. (Zhuravlev, 1941,1943; Novozhilov, 1948a, 1964). The Mesozoic marine reptiles
Figure 11.2. Pliosaurus (Stmngylokrotaphus) irgisensis (Novozhilov, 1948a), PIN 426, from the Volgian oil shales of Savel'evsk Mine No. 1, near Gornyi, Pugachev District, Saratov Region. A, Skull roof, occiput, and right mandibular ramus in dorsal aspect (posterior at top). B, C, Anterior cervical vertebra in anterior and left lateral views, respectively. D, Right hind limb. Scale bars, 100 mm. Compare with Novozhilov (1964).
skeleton, as formerly exhibited in both the Pugachev Novozhilov (1964), there appears little to differentiate Museum and PIN, was heavily restored and is less well the species from Pliosaurus, and his creation of a preserved than appears in previously published unique genus (Stongylokrotaphus) appears unjustified. photographs (Zhuravlev, 1941, 1943). All elements are The cervical centra of Pliosaurus irgisensir are large now greatly affected by pyrite decay and the skull and and blocky, but very short, as in Pliosauws, and their mandible, in particular, are nearing destruction unless ribs are distinctly double-headed as is typical for plio- emergency conservation measures are soon employed, saurids. Novozhilov's (1964) placement of this species an unlikely scenario under present constraints. in the Polycotylidae (= 'Trinacromeriidae') is entirely Tarlo (1960) equated this taxon with Pliosaurus, inappropriate. The figures in Novozhilov (1948a, noting that Novozhilov's (1948a) original assignment 1964) are misleading; the ovate 'fenestrae' seen lateral of this very large specimen to the relatively small to the posterior ends of the parietal crest represent an Peloneustes was seemingly only on the basis of its elon- oblique dorsal view through the distorted posttempo- gated rostrum (of which only the extreme anterior tip, ral fenestrae. The posterior interpterygoid vacuities found in place, is preserved), and a misunderstanding are not visible dorsally. The apparent 'foramina' in the of the characters of Pliosaurus. Indeed, contra occiput of Novozhilov's (1964) figure represent the G.W. STORRS etal.
Figure 11.3. Georgzasauru~(Geo7gia)penzensi.r (Ochev, 1976a), POKM 11658, from a building stone quarry in Santonian rocks at Zatolokino, Bekovo District, Penza Region. Natural mould of braincase, dorsal surface of palate, and rostrum. Total length of skull approximately 800 mm. Compare with Ochev (1976a).
splayed halves of the crushed foramen magnum with POLYCOTYLIDAE Williston, 1908 the exoccipitals forced laterally. There is no lacrimal Georgiasaurus Ochev, 1977 preserved. Indeed, if once present, the orbits and ante- See Figure 11.3. rior rami of the squamosals are now lost. Neither is Diagnosis. Relatively large polycotylid pliosaur distin- Novozhilov7s (1964) figure of the right hind limb guished by its size (+5 m) and relatively elongate entirely accurate, as comparison with Figure 11.2 rostrum; details of palate and braincase noted by indicates. Ochev (1976a) may also be significant. The specimen has been cited as one of the few known pliosaurs with gut contents, i.e. associated Georgiasauruspenzensis (Ochev, 19 76a) cephalopod hooklets (Zhuravlev, 1943; Gekker and Holotype and locality. POKM 11658, the natural moulds Gekker, 1955). Zhuravlev (1943) also notes the associ- of a skull and partial skeleton from a building stone ation of an entire fish, shark teeth, and sand or gravel. quarry in the Penza Region, Russia. Surely the shark teeth are the result of scavenging of Horizon. Upper Cretaceous (Santonian). the carcass by these animals, but the current state of Comments. Georgiasaurus Ochev, 1977 replaces the pre- conservation of this fossil reptile prevents any addi- occupied Georgia Ochev, 1976a. The snout of tional comment. Apparently, none of the associated Georgiasauruspenzensis is narrow and elongate and the remains has been preserved in the collections. cervical ribs are single-headed as in other polycotyl- hlesozoic marine reptiles ids. Ochev (1976a) estimates the total length of the Basin, and Ples in the Kostrolna Region (Bogolyubov, animal in life at t5m, relatively large fix most poly- 1911; Dubeikovskii and Ochel., 1067). Volgian to cotylids. Although the holotype material is poorly Valanginian records of 'LMuvcretzo.muvtr.r' come from preserved, this taxon is provisionally retained on the hloscow and Orenburg, but these identifications hare basis ofthe adequate description by Ochev (1 976a). not been confirmed. Pliosaurids, of the sort common in the English Upper Jurassic and represented in Kussia by I. io~~~lc'urodotlrossicus and I'lios~~urusirgi.ret~.ri.r, are known The primary contribution of Iiussian plesiosaur fro~nseveral localities in addition to those listed above. material is the supplerrlentrlry information that it pro- 'Plios~~uru.r',perhaps representing hoth of the above vides on the stratigraphic rlnd geographic occurrence taxa, has been reported from the Moscow Basin of the group, and particularly for suprageneric-level Oxfordian, Callovian, Kirnrneridgian, and Lrolgian raxa. The earliest sauropterygian material from Iiussia ('l'rautscl~old, 1860; Ryahinin, 1000; B(>g(>ly~l)ov, was reported by l,azurkin and Ocher (1068) from the 191 I), the liyazan' Region Callovian, and the Volgian Ladinian of the left bank of the Taas-Krest River, of the Samara, Ural'sk, western Kazakhstan, and Lena liiver Basin, near the Laptev Sea, northern Chuvasliia regions (Kozlidestvenskii, 1947; Russia. This fossil, an isolated dorsal centrum (SGU Novozhilov, 1948a; Bazhanov, 1958). So-called 104all5, now in PIN), ~rasOelievcd to represent a P~~1otlc'tr.rte.ris known from the Vologda-Vyatka railwa!, 'nothosaur', identified as ."\.hthos~~uru.i;but the presence (Callovian or Oxfordian), Kostroma Kegion, and the of distinct foramina subcentralia, rare and irregular in Upper Jurassic of Hooker Island, 1:ranz-Josef Land stem-group nothosauriforms, suggests that the bone (R!lal)inin, 1000, 1936, 1939). Pliosaurids are also represents a primitive plesiosaur similar to Pi.rtoscrurtrs known from the Oxfordian of the Unzha River in the (Storrs, 1991). :I similar centrum (PIN 4466114) has Kostroma region. "fhatrmcrtosnuvu.r' has been reported heen reported I>!/ Novikov (1993) horn the late from the Callovian to Kimmeridgian of the hloscow Ladinian of the Pechora Iiiver Basin, Bolshaya Syny Region (13ogolyubov, 19 1 1 ), but Thaumnto.mtrrus is tributary, as ?L\rothosaurus. based on a single tooth from the German Jurassic and Plesiosaurs are also extremely rare from the Lower is a flonlc'a dubium. 'Po'olyptycL~ododhas been reported Jurassic of Russia with but a single report of a vertebra from the Cenomanian of Kursk and of Podolia, from the Middle Liassic of the Jurung-'l'umus Ukraine (Kipriyanov, 1883; Bogol!,ubol; 191 1; Peninsula, Nordnick Ray, of the north Siberian .irctic Hofstein, 1961; Kozhdestvenskii, 1973). (Ryabinin, 1939), hut in the uppermost Middle tindouhted rnemhers of the Elasmosauridae occur Jurassic, and especially the Upper Jurassic, they have in the Cenomanian of the Kursk and Saratov Iiegions been widely encountered. The so-called cryptoclid- and the Saratov luronian and Campanian, I>i~tnone ids, a possible fanlily of mesodiran, small-headed, is complete enough for generic identification many-toothed forms, have been reported fiorrl the (Kipriyanov, 1882; Praroslavlev, 191 8). Senonian elas- Callovian-Oxfordian of Gorodishche near mosaurs have come fiom the eastern I're-Urals Ul'yanovsk (Bogolyubov, 1909) and the Vologda (Novokhatskii, 1954) and from the regions of Penza Region, and may exist in the middle li)lgian of the and Orenburg (Chkalov), around h/lala!la Serdoba Mosconr Region (pers. obs., C.W.S., L:M.E.). (Campanian) and Konoplyanka, respectivel!~ Plesiomorphic elasmosaurs, reported as (Rogolyubov, 101 1, 1912; Pravoslavlev, 1918), and a ',llur~~~ttosnuvu.r'and 'Colj~mbos~~uvu.r',are known in the possible elasmosaur was identified by Ryabinin (191 5) Callovian of the Ryazan' Region, the Callovian or from the lower Senonian of Sakhalin Island. Oxfordian of Yaroslavl', and the Volgian of Moscow Maastrichtian occurrences at the .%dzhat Kiver in (especially hlnevniki and Schukino), the Kanla River north Kazakhstan and the hlugai River in the eastern Pre-Urals were noted t~yNovokhatskii (1954). Upper 1939), and the rilbian oFSaratov (hI.S..I., pers. ohs.). In Cretaceous (undifferentiated) elasmosaurs have been the Ihlga Basin near Ul'yanovsk, plesiosaurs range found along the Liska River, Don liegion (Pravo- fro~nCallovian to .\ptian. slavle\l, 1914, 1016, 1918), and in the Cuberlin ,I .lie Russian evidence of plesios;~urstratigraphic Mountains of the southern Pre-Urals distribution agrees well with that suggested 11y occur- (liozhdestvenskii, 1973). Elas~nosaur occurrences rences in Western Europe, Australia, and the from the Russian Lower Cretaceous are known from .\mericas. It appears that the Elasmosauridae, with the Iianterivian Vyatka-Ka~na phosphate deposit possil~lerepresentatives in the [.ate Jurassic, contin- (Dubeikovskii and Ochev, 1967) and the liptian of the ued to exist until the latest part of the Cretaceous, as Kara Sea (Kyabinin, 1930). l'ossible 'cirnoliasaurs', or did the polycotylids, although the latter did not make perhaps stern-group elasniosaurs, are reputed to occur their appearance until the latest Larly Cretaceous. in the LTolgian of the Moscow Basin at Schukino and Cryptoclidids, if a legitimate cladc, appear restricted Tartarovo, the Albian? of the Moscow Basin to the 1,ate Jurassic in Russia and elsewhere, ~~lthough (Berezniki), and the Senonian of Orenburg-Chkalov perhaps surviving locally into the Early Cretaceous. (Konoplyank:~, Cuberli), and the undifferentiated ?.I he giant Pliosaaridac thrived in the I,atejurassic, but Upper Cretaceous glauconitic sandstone of the continued into the Earl!, Cretaceous, seemingly Charkov Kegion (Ryabinin, 1909; Bogolyubo\.; 191 1). becoming extinct soon thereafter. Including GeurgiaJauru.r, small, longirostrine 'plio- saurs', or polycotylids, have been identified in the ICHTIIYOS.\URIrl dc Blainville, 1835 Cenomanian of the Voronezh (Devitcha), Saratov, Curtzvzeuts: The first Russian ichthyosaur remains were Volgograd, and Kursk regions, the 'Turonian of noted by Yazikhov (1832). Ichthyosaurs are cornmonly Saratov and the Tadzhik Depression of Uzbekistan encountered in the Upper Jurassic, largely Oxfordian (Talkhab Svita; Dzhalilov etal., 1086),the Santonian of to Volgian, and Lolver to lo~~~crn~ostUpper Penza (Ochev, 1976a), and the Senonian of Saratov Cretaceous (Cenomanian) rocks of the Russian (Campanian), Penza (Malaya Serdoba; Campaninn) Platform. As clsclvhere, their remains are more and Orenburg-Chkalov (Konoplyanka) (Bogolyubov, common in rocks of these ages than are those of ple- 191 1, 191 2; Sintsov, 1899). They are also known in the siosaurs, and hetter material is available. undifferentiated Upper Cretaceous of the Don Unfortunately, similar taxonomic chaos also exists. Region's Liska River (Pravoslavle\: 1014), the Over a dozen ichthyosaur species have been named Cuberlin Mountains (Rozhdestvenskii, 1073) and the from Iiussian specimens, and rnost have proved to l)e Lysogorsk District of the Saratov liegion (Ochev, indeter1nin:lte. I,ikewisc, historical works (e.g. 1976a). Most of these fragmentary specimens have Kipriyanov, 1881) have also made use of supposed been identified as Pulycotylus, but are generically inde- type series with the implicit, but unproven, assump- terminate. tion of specific identity The problematic taxa are In addition, indeterminate plesiosaurs are known included with potentiall!, valid species in 'Table 11.2.
from the UpperJurassic of the Vilyui River in Siberia Nesov (I nl. (1988) provide a stratigraphically orga- (Menner, 1948), the Volgian of east Siberia, near nized list of ichthyosaur localities for the former Zhigansk, the polar Urals of the Lapinsk Region Soviet Union, and 'l'riassic records map be found in (Yakovlev, 1903; Ryabinin, 1939), and the Priural'sk C:lllaway and Massare (1980). District of west Kazakhstan (Bazhanov, 1958). They Few remains of Russian ichthyosaurs are known are also known from the undilkrentiated Lower from the Triassic and these are all of poor quality. Cretaceous of the southwestern part of the Cissar Kyahinin (1046a) attrih~teda Norian vertebra from Mountains near Baisun (Kozhdestvenskii, 1973), the Kolynskii to Shastasnurus (S. .rievrr.ri), but Tatarinov I'alanginian of the Lyapin area, north Urals (Kyabinin, (1964) believed that it could more probably be Table 1 1.2. Compilatiotz of ichthyosau~agtaxa ba.red upolz material from thefovmev Soviet rgiotz. Holotypes igdicated by vepositovy abbveviatiog ovspecimew gumbev, wheve k~zo-wg
Taxon Holotype hlaterial Locality Horizon Status
Brachypterygius zhurilvleai Lirkllangel'skii, PIN 426160-76 partial fore limb Saratov Region middle Volgian ? Otschevia sp. 1998a Ichthyosaurus kurskeusis Eichwald, 1 85 3 teeth and I ertehrae Kursk Region Neocomian Ichthyosauria indet. Ichthyosaurus ~zasinzoviiFahren kohl, 1856 2 t ertebrae AZoscon. Region Volgian Plesiosauria indet. Ichthyosauws steleodo~zBogol yuhov, 1Y 09 frdg 41ertebrae Ul'yanovsk Region Hauteric-Barremian ? Plagptetygius steleodofz Ichthyo.iuurus volgensis Kazanskii, 1903 KG U fragmentary skeleton Ul'yanovsk Region Lhlgian ? Ophthalnzostrurus sp. Khudiukooia callovirn.ris Airkl~angel'skii, SGU l04a127 partial fore limb Saratov Region Call01 ian ? Ophthabuosaurus sp. 1999a lIlyopretygius kipri~a~iofiRoiner, 1968 ZIS.2 [Kipr., 18811 partial skull Kursk Region up. Albian-Cenom.
'Simbirtsit' No. 140 fragmentary skeleton Ul'yanovsk Region upper Kirninerigian presurned t alid UPAI No. 31100 partial skeleton Ll'yanovsk Region lower L'olgian presumed valid PhlK No. 2836 frag. postcrania Saratov Region middle Fhlgian Partropthal~tzostrurussp.
SGU 104a123 partial skull/skeleton lotver Volgian provisionally retained SGU 104a/24 partial skull Saratov Region middle Cenomanian provisionally retained - propodial fragment Criinea Neocomian Icl~thyosauriaindet. [Ryabinin, 1946bl UPhl No. 21740 skull/partial skeleton Ul'yanovsk Region upper Hauterivian Plntyptetygius hedengeusis ?Shustusaurus norde~zs2sPolubotko Sr SGU 104a/16 partial I ert. series Dyugadyak River Ladinian hlixosauridae indet. Ochev,1972 Basin Shustasatirus .rieversi Ryabinin, 1946a - tertebra Iganii River Basin Sorian Ichthyosauria indet. Sznzhivskiasaurus hi~~zikouiOcllev & SGU 104a122 frag. skull/skeleton Ul'yanovsk Region Hauterivian presumed valid Ehrnov, 1985 C'ndorosaurus gorodircheusis E fi mov, UPXZ EP-11-20 (527) fragmentary skeleton Ul'yanovsk Region middle Volgian presumed L alid 19Q6b C?zdolnsuurus un:rovz Efimov. 1Y96b Ul-'hI EP-11-24 (785) fragmentary skeleton Ul'yanovsk Region middle Volgian
Mesozoic marine reptiles assigned to Mixosauvus. Callaway and Massare (1989) Waldheirn, 1817; Kazanskii, 1903), but it is unlikely determined that this was not a mixosaur and that the that any of these specirnens truly represent species was a aomm dubium, reassigning the specimen Icbtbyosauvus, a LowerJurassic genus long treated as a to Shastasauridae incevtaesedis. Even this assignment is 'waste-hasket taxon'. Indeed, the English holotvpe of I. too specitic and the material is nondiagnostic beyond tvigo~zusOwen has been transferred (Huene, 1922) to Ichthyosauria. Similarly, fragmentary specimens from ,Macvoqtetygius, and I. pla[yoclof~ Conybeare to northeastern Russia (Okhotsk Sea, Russkaya River, Ternnodo~~tosauvusby hZcGowan (1074). Nevertheless, etc.) of the Carnian and Norian to Rhaetian discussed the Russian specimens assigned to these species by Ochev and Polubotko (1961) are generically inde- (Ryabinin, 191 2; Fischer von Waldheirn, 1817, respec- terminate. The Ladinian and Carnian Omolon Massif tively) do not allow confident identitication at even the specimens of Polubotko and Ochev (1972) on the generic level. lcbrhyo.sauvus nasilrrooii Fahrenkohl, 1856 other hand, may represent hlixosauvus (their was founded upon two indeterminate plesiosaur ?Sbastasauvus ~rovdefzsis,'I'able 11 .2) and Cymbo.rpo~~dylus, vertebrae and is thus a nonml duhium (Eichwald, 1868; respectively (Callaway, pers. conlm. 1996), although Kipriyanov, 1883; Bogolyubov, 191 1). Mazin (1983) considered the former Mixosauridae incevtae sedis. Nesov et a/. (1 988) also note problematic ICIITtlYOSAURID;l\~Ihmaparte, 1841 material ('Sbastasauvu.c and Mixosauvus') from the Opbtba1nrosauvu.r Seeley, 18741) Norian of the Magadan Region. Many very fragmen- Diagfaosis. Relatively large 'latipinnate' ichthyosaur, tary individuals have been found in the up to 4 rn, characterized by extremely large ort)its and undifferentiated Triassic of Chukotka District not far very weakly developed or absent dentition; teeth cer- from Alaska (V.M.E.). 'She earliest potential ichthyo- tainly present only in anterior tips of jaws; humerus saurian occurrences in the forrner Soviet Union may distinguished by presence of 3 distal facets of sub- be indeterminate .4nisian and Induan specimens from equal size. the Ornolon Massif, Volgograd Region (Ochev, 1976b; Polubotko and Ochev, 1972), although Shishkin and Opbtbalmosauvus u~rdovcnsi.~Efimov, 199 1 1,ozovskii (1979) note indeterminate material from the llolotype a?zd localilyy. 'Si~nbirtsit' 110, fragmentary 'Lower Triassic' of Russkii Island, as well as from the skeleton; Volga Killer shore, found in 1982 near the 'Middle Triassic' Daonella Shales of the Amur River village of Malye Undory, Ul'yanovsk Region, Russia. estuary. Hovizou. dulacostepba?~u.rmutahilis Foraminiferan Lone, Although only a felr ichthyosaur elements are Upper Kirn~neridgian. known from the Lower and Middle Jurassic (e.g. Comments. Etimov (1091) described this species on the 'Soarcian and Bajocian to Bathonian of Yakutia), basis of a very fragnlentary skeleton. Nevertheless, the Upper Jurassic finds abound. 'These latter are again character of the humerus clearly indicates that it primarily of Moscow and Volga Basin origin belongs to 0phthaltvosa1lvu.r. l'he material \\.as consid- (Arkhangel'skii, 1999b; Arkhangel'skii et al. 1997), ered to represent a new species partly on the basis of although indeterminate remains are known from the very short vertebrae and on inferred difkrences ill Volgian sediments in western Kazakhstan (Bazhanov, the scapula and forelin113 hetween it and 0. icc~~icus 1958), Ul'yanovsk (Kabanov, 19.50; Efimov, 1987), (Efimov, 1991 ). Provisionall!, retained here, this Moscow (Orlov, 1068), Saratov, and the Syzran' and species is poorly characterized and requires additional Ukhta Regions (Nesov et al., 1988). Various remains study. attributed to Icbtbyosauvus have heen reported from the .4 numher of Volgian-age ichthyosaur specimens Kimmeridgian of the Pechora River Basin (Ryabinin, perhaps referrable to Opbtbalmosauvus arc known. 1912) and the Volgian of Moscow (Pahrenkohl, 1856; Moscolr Region specimens (?Opbtbal~~~o.rn~~l.ussp.) Trautschold, 1861) and Ul'yanovsk (Fischer von include those noticed by Bogolyubov (1010a) and G.W. STORRS ptal.
Mitta (1984). Bogolyubov (1910a) also noted, but did Arkhangel'skii 1998a, is hased upon insufficient mater- not describe, a partial Opbtha1mo.muvu.r skeleton from ial for diagnosis and is considered Pavaopbtha1mo.rauvu.r the Volga River hetween Ul'yanovsk and Gorodische, ". which he provisionally assigned to the English species 0. icenicus. Zhuravlev (1 941, 1943) identified Pavaopbthalmo.rauvu.r,ya.~ykovi(Efi mov 1999a) Opbtha1mosauvu.r sp. in the Middle Volgian Dov.rop1anite.r Holotype aud locality. UPM No. EP-11-7 (12 35), skull pandevi and I7ivgatitt.r vivgatu.r zones of the Savel'evsk and partial skeleton; Detskii (Children's) Sanatorium oil shale mines near Pugachev, hut these may he area, Volga River, Ul'yanovsk Province. misidentified (M.S.A.). Another unique Russian Horizon. Cva.rpedite.r subdite.r Zone, upper Volgi an. species, Icbt/3yosauvus volp~.ri.rKazanskii, 1903 is based Comment.r. Described by Efimov (1999a) as a new upon a fragmentary skeleton from the K vivgatite.r zone genus, Yasykocia, this material is considered a junior near Ul'yanovsk that, although probahly specifically synonym of Pavaophtha1mo.rauvu.r hy ,4rkhangel'skii indeterminate, may represent Ophtha1mosauvu.r (but see (19991,). l'he species is provisionally retained here. helow). Indeed, Bogolyubov (1 91 0a) assigned I. volgen- Three other specimens descrihed hy Efimov (1 999a) as sir to the 'ophthalmosaurs'. Several species with possi- new species of Ya.rykovia are hased upon suspect mate- ble 'ophthal~nosaur'relationships have been described rial, two of which are synonymized with Pya.rykovi(see hy Arkhangel'skii (1997, 1998a, 1999a) and Efin~ov 'Table 1 1.2). (1999a). Lastly, a possihle Ophthalmosauvu.rvertebra has come from undifferentiated rock on the left bank of C'udovo.rauvu.r Efimov, 199911 the Volga near Kashpir and Syzran' (Bogolyubov, Diagn0.ri.r. Moderate to large sized ichthyosaur 1909). (46 m) with massive vertebrae and stout ribs; broad, pentadactyl forelimbs with three humeral distal facets. Pavaopbtba1mo.rauvu.r Arkhangel'skii, 1997 Seemingly distinguished from 0phthalmo.rauvu.r by its Diaguosi.r. Slnall ichthyosaur 2-2.5 m long with high large, strong teeth. skull, large orhits, and reduce dentition. Propodials with three distal facets as in Opbtbalmosauvu.rbut acces- Lrndovo.rauvu.r govodi.rchen.ri.r Efimov, 1999b sory facet reportedly bearing basal element of first Holo suggested an affinity with .I~~op~~~ygius,hut there is wedges het\vecn radius and ulna to narro\rly contact scepticism ahout the validity of this genus (McGowan, humerus. 1972), rnost material no\r being referred to Plut),ptt~~~~~ius. 0t.rchcaiu p.retrdo.rcjithicn Efi Inov, 1908 Ilolo~ypeulid lo~li[y.UPhl No. 31100, a partial skull S'I'F:NOPTEKYC;l1D:IL Kuhn, 1934 and skclcton with good forelimbs; Volga River right 0t.rchcvio Efi inlo\,, 1998 hank, 8 km cast of Novaya Bedenga, Ul'panovsk Diugfiosis. 'Longipinnate' ichthyosaur of moderate to Pro\,ince. large size with curved, stout tccth; cpipodial and Horizon. /lou,ai.rkg y~s p.r~i~do.rcythic~s Zone, lower mesopodial hones robust; irltcrmcdiuln large and Volgian. b1esozoic marine reptiles Com~rent.r. 'l'he seemingly unique forelimb suggests the basis of the available material. It is Campanian in that this is a distinct taxon with a relationship to the age and comes from the Liska River Basin, Don Region. Stcnopterygiidae. Bmc;c!ypteyygiu.r zbtrvnzdevi P/iopl(~tccnrpu.ris questionably present in Povolzh'e ,4rkhallgel'skii, 1908a has been referred to Otschez,in hy (Tsaregradskii, 1927; Khozatskii and Yur'ev, 1064; Arkhangel'skii (l900b), although represented by Yar kov, I 9 3 7,ylos~~unn.r (I,iodo~r) rhi/)/~a/u.s extremely poor material. ,-\rkhangel'skii (1999h) has (Bogolyuhov, 191 Oh) Lvas described from the also considered Ici~tbyosnurusvo1ge~~si.r Kazanskii, 1903 Campanian of Orenburg Province, but is also prohahlv to represent Otschtvia. not diagnostic of a valid taxon. Ty1o.ratrvtu.r has also been reported from the Campanian of Saratov, Volgograd, and the Konoplanka River near Orsk, and from the Crimeari Maastrichtian (GorOakh, 1067). Pvoguntl~odolon Indeter~ninateichthyosaur remains have been found is thought to he present in the Campanian of Saratov in the Lower Crcraceous of .izert)aijan (Aptian) and Penza, and in the blaastrichtian of the Cri~neaand (Bogachev, 1962), and the Sarato~; Samara Penza (Malaya Serdot~a) (h1.S.A.). Plntecnr/)us and (?\'alanginian), Kirov (I'alanginian), and Periza Clidu.rtcs reputedly have been found near Volgograd (Albian) Regions (Nesov et nl., 1988). F'1e.riosauvtu.r uord- (Yarkov, 1993), but these reports are considered prob- mnui Eichwald, 186.5 is hascd upon a limb fragment lematic at best. Similarly, C1idnste.r is questionably fro111 the Crimean (Biasala) Neoco~nian(Berriasi;~n?; reported froni the Campanian of Saratov and Penza Bogolyuhov, 1911) that Kyabinin (IY-Fhh) identified as (h1.S.A.). Indeterminate mosasaur remains have also ichth\7osaurian. In the Upper Cretaceous, ichth\lo- been found in the Lugansk and Serov regions, the saurs are known from Sakhalin Island, and from the eastern slope of the Urals, Karakalpakia, western Cenomanian of Saratov, the L'\~ov(Vinnitsa, Ukraine) Llzerl>aijan, arid Kazakhstan (Novokhatskii, 1954; and 'I'aml~ov Kegions, Cherkassia (Nesov eta/., 1088), Gabunia, 1958; Kozhdestvenskii, 1973; Kovaleva eta/., and bloldavia (hloldova) (Sukhov, 1950). The 1082; Yarkov, 1903). S~nolensk Region has produced material from undifferentiated Cretaceous sediments (Nesov et nl., Uol1o.rarrrtu.r Ya kovlel; 1001 1988). See Figure 11.5. l>iagno.ri~::I plioplatecarpine mosasaur characterized SQUAhI 41'A Oppel, 181 1 by no pre~naxillaryrostrum anterior to the dentition; -\NC U 1 hlO K P H i Furbr~nper,1900 recurved maxillary teeth with single carina on back hlOS -\S -\URII)IL Crervals, 1853 edge only; concave alveolar rim of dentary; strong Conz~re~1t.r.hlosasaurs of the former Soviet Union are symphysis; very large first t1vo dentary teeth; trans- known particularly from sediments of the Lower Volga versely broad cervical and anterior dorsal vertebrae; Basin in Russia (Saratov and Penza Regions) and range well developed zygosphenes and zygantra; co-ossified in ape from the Carnpanian to the Maastrichtian che\/rons(Yakovlel7, 1901;Tsaregradskii, 1935). ('iarkov, 1003). 'l'he genus Rlosusaurus has been reported from frag~nentar!, remains, primarily verte- Uol1o.raurtu.r lutugiui 'iakovlev, 1001 brae and teeth, from the I'olga 12egion hut also from the Ilolo~y/)ead loculi[y. fragmentary skull ;~ndskelcron basins of the Don, Ural and Pechora rivers, Turp~i,and (whereahouts unknown); Doncts River, I'oroshilov- Azerbaijan (Yakovlev, 1002, 1905; Khozatskii and grad Region, Ukraine. Yur'ev, 1964). Not all of these reports are reliable Hovizof~.Relc~~~~itelln ~~rrrcvouata Beds, Upper Cam- beyond hlosasauridae indet. The one uniquely Kussian panian. species of hlosasauvus, (If.dof~icu.r Pravoslavlev, 1914, Cotnnzefrts. This species represents the only genus of being extremely fragmentary, cannot he diagnosed on mosasaur that has been described solely on the basis of Figure 11.5. Dollo.sauru~.lutugz~~iYakovle~~,1901 from the uppcr Campanian of the Ukraine (Vroroshilovgrad Rcgion) (after Tsaregradskii, 1935). A, Right maxilla. B, Right dentary in labial aspect. Estimated Icngth ofdentary 300 mm; maxilla to scale. material from the forrner Soviet Union. Both Dollo tarsal of a metriorhynchid, perhaps Dakosau~~u.r,were (1 924) and Russell (1 967) accept Dol1osauru.r as a valid recovered froin the uppermost Jurassic or lowermost genus and have allied this animal with the Cretaceous of Khoroshevskii Island in the middle Plioplatecarpinae. Russell (1967) considers it close to Volga Region (Ochev, 1981), and an indeterminate Prognatbodon. thalattosuchian has come from the Kin~meridgianof the Moscow Region (Efimov and Chkhikvadze, 1987). SERP ENTES Linnaeus, 1758 :In unpublished partial thalattosuchian is also known Comment. Strictly marine fossil snakes are unknown frorn the Volgian of Gorotiishche near Ul'yanovsk from Russia, but a single occurrence of a primitive (V.M.E.). In Asia, teleosaurs have been reported, simoliophid is known from Albian- Cenomanian such as Steneosaurus tiom the Jurassic (A4alenian) lagoonal deposits in Kzyl-Kum (Khodzhakul Village), Karakhskaya Svita of South Ilagestan (Efimov, 1978), Central A& (Clikman etal, 1987). while T&osaurus reputedly occurs in the uppermost Jurassic or lowermost Cretaceous of the -4laiskii ARCHOSAURIA Cope, 1869 Ridge, Fergana Kasin (Efimov and Chkhikvadze, CROCODYLIFORMES BentonandClark, 1988 1987). Poekilopleuron scbmidti, from the Cenomanian of Comments: Few marine crocodyliform (crocodilian of the Kursk Region, was described by Kipriyanov traditional usage) remains are known ti-om Russia and (1883) as a marine crocodile, but Efimov and its former republics. Rare thalattosuchian fossils are Chkhikvadze (I 987) believe that these remains repre- known from the Russian Platform but are largely sent a dinosaur. undescribed. For example, a vertebra and fifth meta- Marine thoracosaurine remains are also rare in ,Ilosco\v-Brisrol Palaeontological Initiatiic v.Il~(-ii /II/I ..I/I~I/ / O/I 5.i. Oxford: spawned this project ant1 their invitation to ct~!lti-illaii. ~:l~lrcll~lo1l to this hook. \Ye are also indebted to Professor \i[.llii I:lai~l\~llt., 11.1). tlr 1 S3i. 11c\c1.il)tiondc cluelclues espi-ces Ochcv Ii)r his enthusiastic help and guidance i~itl~c tlc I-cptilcsde la <:;~liti)rnie,p1.6ctdte de I'a~ial\,sed'l111 pursuit and co~nl~letio~iof our studies. \\illi;l~i~ \ st61iic gGn6ral d'l:rl>erologie et d'.\~iipIiihiologie. .\ oi~~~i~//c,.i1117/11/c>.r dii I 11/.r?~/iiz(.\'nt;oii~zl) ~/'//i.stiiirc (;ollicr and Sinion Powell provided photogr.nl~Iiic. .\,t/i/ri~//~,,I),ir/! 3: 23 <-200. assistance, and we thank Mike 'l'aylor and the Iiltc.,l~ic.l, I:ogaclic\, \ .\. IOO1. I llisco\ crv of C;retaceol~s ichtlivo- (:allaway for their review comments. (;.U'.S. tll;11!J\5 s;lklrs in tlic(:auc:lsu~.l I'rii~ii11111962: 11'). tlie Royal Society of London and the N:ltu~-al l%ogol!~ul,~)\,N.S. 1000. Sur cll~elql~esrestcs dc dcus rcl>- Enviro~iment liesearch (:ouncil of the Lli iiis 1 i i c~ ( ( ~I:J~I/IJL/;11i/.i .I.~II//I;I:I.~~.I/.~~.~11. sp. et IO!I~,!~~JJI~LII~~I.~.stc2- financia1 support and the Universit! of I3rist1)l Ico~loiin. sl'.), rl.ou\ 6s par XI. le Profes. .I. I! 1'a\.lo\v sur Department of Earth Sciences and the hlusru~ii01' Ics I)ords de la \'oI+i dans les couches 1ii6sozoic1uesdc NaturaI tiistory & Science at Cincinnati h1~1scu111 hi nil~irsk.. I~~ILLII~I~~~(;?o/o~iq~i(~ cat .I li~i(;~~~ilog~q~~c~lc/u Center f;)r the use of their facilities. / fro111 the lower \i)lgian substage of tlie S;~~-;lro\. ,I 1~IIL~I~I~O~/~ZLO I/? 111 K~I.~.I.I(~ 1 1: 42-64. La\~olzh'e.]Puleo~ttcikogicl~c.~~kii %huniiil 199 7: 8 7-0 1 . -1 0 1 1 . 1 OIL 111c //z.rfo~~)~(11. /'/c~.rio.r~di~~r i7i / -1998b. LOri the genus of ichthyosaurs l'1ut~~itcy~~iii.1.~ \ 1 ~l/l'i'li//J~~~~~i~'dl' /il /t.rieur du (:rGraci. de la (;rimi.c. Uallcti~i d~z iclitlivosaurs and the ph\,logeny of the pup. I b/Ji-O.I:)~ 1: lcn(lb~iiic I~~ipt:t-inlci1~a.r Sci(,lice~; St. Pc;tc~~-hu~?7: I'nlt~o~itolociii ,St?,(itig~/!/ii 1999: 2 0--3 7. S.i.7--is. -, I\~a~iov,.I.\'. ;ind l'opo\,, k.1'. 1007. [On tlic first reli:il)lc (;a1 l:iway, I. and ?lassarc, J. 1080. SLn.st~i.surui-us~ilti.~pi?im.r discovery of remains of the ichthyosaur P//it)ptr~i:)yi~/.i (Iclitli\~osauri:~,Sliastasauridae) fro111 the Upper in louer .iptian deposits of Povolz1i'e.l 1'cl~eliie %iipi.rki Triassic of tlie 1'1 :\ntimonio district, northwestern Cc~oIogichr.rkogub2ic1rltetu Snmtuvskugu Gusr~d/dirvsti,e~i~iopo Sonora, hlesico. .~oIAI~I~/I/~/P/~li~o1itu/o~y63: Y 30-939. 1 izlacvsitc%a 1997: 57-50, (:ope, t1.11. 1800. Synopsis of the extinct Ratrachia, Reptilia Rardet, b. 1002. Stratigraphic evide~icefor tlie extinction and .Ives of North rimerica. 'li~u~z.rurtiu~is?/- the of the iclitliyosa~~rs.'/i.1-?-u fiiw4: 040-656. :Iiiici-icoli /'/li/n.io/lbicii/ Sut.ic<)l 14: 1-2 52. Bazlia~io\,,1:s. 1958. [Concerning a pliosaur and iclith\~o- Ilollo, L. 1024. (;lohi~/(~~i.ialiihii~~iie~isis, IIIOS~S~~I~~CII a~iitri- saur from the Ul>perJor;issic of westerri Kazak1ist:in. 1 cain rerrou\.t dans le Craie d'Obourg (Sbnonien .Ilute~~ial)ipoIstuvii F(zrnt SSR.1 [.\bsrr;lcr 1 ~j~i~/lr,/c~ii' I lri.~.ko;~.~.koxo O/~.s/)i./)(:rti,~ lii~/~ti-ir~/r,rhi .Zl(ltu~ulistr~.r dc .Ilo.icui, 20: 362-3 70. /.yytritr~Ic~iI'i-ii-ocll: Oti1c.l (;cologid)/:ikii5 3 : 1 7 7. t 'iirl~ri11j:c%r. 11. 1 VOO. Hr~itt~u~qzu~Sy.stc,inri~ik urid (;eiir~nlngi/~r1t.t- -1'288. [Fossil crocodiles and clian~psosaursof klonsoli;i l\'c!~iii!r~iiJenn:Gusrav b'isclier. and the USSR.] 7;,1idy ,Yooiiic:rtiioi Soz~t.rko- I lriz/gol:rkoi (;:~l~ulii;i.iL.li. 1058. [On the remains of mosasaurs from P(i/c~oiitol~~g~c/~~~.rI~niI:k.r/~r.rlit.~-ii 3 6, 1 - 1 OX. I. !.;XI. C:reraceous deposits of the Caucasus.] andClikhikvadze, \'.XI. 1087. ISurve!. OK tlie hncls of ,Sr~r~~~.sbcbcniyilyii. I.\' Gi,tizinskoi SSK 20: 56 1-564. fossil crocodiles ill the USSIl.1 Izvc,.rt/"~ri:l.V (;~-iiziii.rkoi (;cl,!,cr, I:.L. and Gekker, R.F. 105.5. / Rclnains of SSK, Sei-{)lri Hiulogicbr~skqyu1 3: 100-1 0 7. 'i'cutl)oidea from the \I.'hite Jura ancl Loucr Etimoy \'.&I. 1987. [hlarinc rcprilcs in Xlesozoic dCpcfiits (;l.c.taceous of the \.blga region.] lup~-ci.~:yI'rilc~oiitolo~qii of the Ui'yanorsk I 1999a. [Ichthyosaurs of rlic nclv sellus l[i.i.yko~,;rifrom I I:ilstead, 1..B. 1071. l.io/~lr~i/t-odui/I-o.s.sicu.c (No\,ozhilo\) - a the Upper Jurassic deposits of ICuropeal~ Russia.] l)liosaur from the loner \illpian of the hloscon. Basin. I'nlc~o~itulogicbt.skii%hi/,-iinl1999: 02-1 00. Pulnr~o~~tology14: 566-5 70. -199c)h. 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