Paleontological Journal, Vol. 36, Suppl. 3, 2002, pp. S185ÐS259. Original Russian Text Copyright © 2002 by Popov, Akhmetiev, Bugrova, Lopatin, Amitrov, Andreyeva-Grigorovich, Zaporozhets, Zherikhin, Krasheninnikov, Nikolaeva, Sytchevskaya, Shcherba. English Translation Copyright © 2002 by MAIK “Nauka /Interperiodica” (Russia).

Abstract—This is the second part of the monograph “Biogeography of the Northern Peri-Tethys from the Late Eocene to the Early Miocene.” The main section, Biogeography of the Early Oligocene, considers the stratig- raphy of the Lower Oligocene Paratethys, the biogeographic distribution of the planktonic and benthic marine biotas and the fresh-water and terrestrial fauna and flora in the Lower Oligocene, as well as the biogeographic zonation of water bodies and land areas. In the sections Latitudinal Climatic Zonation and Major Events in the Late EoceneÐEarly Oligocene, the climatic history of the Early Oligocene is discussed and the major EoceneÐ Early Oligocene tectonic, eustatic, hydrologic, and biotic events are considered.

INTRODUCTION 1986; Geologicheskie i bioticheskie…, 1996); i.e., these This study is aimed at the most complete reconstruc- beds are distinguished based on the their correlation tion of the pattern of the distribution of marine and ter- with Northwest European stratotypes rather than with restrial flora and fauna over Western Eurasia from the the Mediterranean stratigraphic scale, which has been Terminal Eocene, through the Oligocene and to the used for the Priabonian. Early Miocene against the background of major geo- As a geochronological unit, the Rupelian was pro- logical events and paleogeographic and climatic posed by Dumont in 1849 based on outcrops in the changes (the objectivies of this study were discussed in Rupel cuesta (along the Rupel River), northern Bel- greater detail in the first part of this monograph). The gium. There, the major lithostratigraphic unit is the first part also covered the approaches and methods that Boom Clay. However, as a stage, the Rupelian sensu are used in the biogeographic analysis of the Recent lato also includes the underlying Upper Tongrian beds, flora and fauna. The present section is devoted to the Berg Sands, and clay beds with Nucula comta. There biogeographic distribution of planktonic and benthic are no outcrops with a complete sequence of these beds. marine biotas and freshwater and terrestrial fauna and The lowermost layers of the Rupelian Stage are flora in the Early Oligocene, as well as to the biogeo- exposed in eastern Belgium (Leuven Area), while the graphic zonation (integrated wherever possible) of outcrop of the Boom Clay lies farther to the east (Boom water bodies and land areas. At the end of this section, Area). The Upper Tongrian clays yield dinocysts of the the latitudinal-climatic zonation of the Early Oligocene Phthanoperidinium aomoenum Zone, which are typical is discussed and the major EoceneÐEarly Oligocene fossils of the Sannois Beds, Stampian (Cavelier and events are considered. The evolution of various floral Pomerol, 1986), whereas the sands include nanno- and faunal groups and the dynamic aspects of the evo- plankton of Zone NP22. Planktonic foraminiferal lution of biostratigraphic subdivisions will be discussed assemblages that contain no zonal species but include in the final section of this study, after the biotas of the species typical of Zones P18ÐP21, nannoplankton of Late Oligocene and Early Miocene are considered. Zones NP23ÐNP24, and rich dinocyst assemblages of The sections Lower Oligocene Stratigraphy of the the Wetzeliella gochtii Zone, including the W. symmet- Northern Peri-Tethys and Major Late EoceneÐEarly rica assemblage (Stover and Hardenbol, 1993), have Oligocene Events were written by S.V. Popov; the sec- been found in the Boom Clay. Rich molluskan assem- tion Paleogeography of the Paratethys, by S.V. Popov blages (Glibert and Heinzelin, 1954; Glibert, 1957) and and I.G. Shcherba; and the section Latitudinal Climatic assemblages of ostracodes, fishes, spores, and pollen Zonation of the Early Oligocene, by M.A. Akhmetiev were also recorded there. and S.V. Popov. Data on flora are presented by In deep-sea cores from the warm-water area, two M.A. Akhmetiev; on dinocysts, by M.A. Akhmetiev, planktonic foraminiferal zones, P18 and P19 (Blow, A.S. Andreyeva-Grigorovich, and N.I. Zaporozhets; on 1971), or three zones and one subzone, P18ÐP21a smaller benthic foraminifers, nummulitids, and disco- (Berggren et al., 1995), and three (or four) nannoplank- cyclinids, by E.M. Bugrova; on terrestrial vertebrates, ton zones, the upper part of Zones NP21, NP22, and by A.V. Lopatin; on nannoplankton, by A.S. Andreyeva- NP23 and the lower part of Zone NP24, are considered Grigorovich; on planktonic foraminifers, by V.A. Kra- to be equivalents to the Rupelian. Three dinocyst zones, sheninnikov; on ostracodes, by I.A. Nikolaeva; on mol- Phthanoperidinium amoenum (D13), Wetzeliella sym- lusks, by O.V. Amitrov and S.V. Popov; on corrals, by metrica (D14a), and W. gochtii (D14b) correspond to E.I. Kuzmicheva; on ichthyofauna, by E.K. Sytchev- the Rupelian succession based on the sequential skaya; and on insects, by V.V. Zherikhin. appearance of zonal species in the most complete Early Oligocene outcrops (table). BIOGEOGRAPHY OF THE EARLY OLIGOCENE The stratotype of the PriabonianÐRupelian bound- ary has been established in the northeastern Apennines Lower Oligocene Stratigraphy (the Massignano Section). There, in the homogenous of the Northern Peri-Tethys sequence of pelagic marly beds, this boundary is We consider the Lower Oligocene beds as equiva- defined based on the change in the planktonic foramin- lents to the Rupelian sensu lato (Cavelier and Pomerol, iferal zones (P17/P18) and the disappearance of Hanth-

S185

S186 POPOV et al. Regression, stagnation Lesser Caucasus orogenesis Eastern Paratethys Main events in the Full marine conditions 2-nd Paratethvs isolation Short vast transgression Peak of anoxy Turbiditic activity Paratethys isolation, 1-st crisis of salinity Boreal immigrants Eastern Paratethys separation sion/ +– Transgres- Regression ‰ , 2001, modified) 30 et al. salinity Shallow 10 20 nian bulak Kutan- Chilikta- Chagraian Sulte pre-

Aralian Suite

B. antegressa Aralian Anom. munda N. Usturt/ Cheganian TARKHANIAN CHOKRAKIAN Bishtyubjanian Baigubekian Suite Karato- makian Asche- airyk Tamdy Suite Kintykchenian Suite Upper Lower

Suite Suite Georgia Suite Uplistsikhe Akhaltsikhe Otskhe Beds Corbula Beds Abastumanian TARKHANIAN KOZAHURIAN CHOKRAKIAN SAKARAULIAN Globigerina tarchanensis G. tropicalis-

Gl. corpulenta Gl. turcmenica Foraminifera

Cibic. salensis Bol. antegressa benthos plankton costata P. parvus Florilus Saccammina zuramakensis Neobulimina elongata Cibicides omatus– Elphidium onerosum Sp. terekensis Virgulinella Troch. florifera Sp. carinata oligocaenica Lentic. herrmanni EASTERN PARATETHYS

gochtii H. floripes nium draco Rhombodi- D. spinulosa

Dinocysts Beds with

Wetzeliella Microdinium C. partispinatum C. Ph. amoenum W. symmetrica Rh. perforatum clathrata anquiosa Zaporozhec, 1998 Charlesdowniella Andr.-Grig., 1994 stage GANIAN KUMIAN regional KARADJAL- KALMYKIAN PSCHEKHIAN SOLENOVIAN TARKHANIAN KOZAHURIAN CHOKRAKIAN BELOGLINIAN SAKARAULIAN Clay Clay Kiscellian Upper Tard Lower Tard Central

Buda Marl EGERIAN

C. lipoldi Beds (Hungary) Paratethys BADENIAN KISKELIAN Bryazoan Marl KARPATIAN

OTTNANGIAN

Nummulitic Marl EGGENBURGIAN

Krosno Serie Krosno

Carpathians Menilithic Serie Menilithic Lower Cherts ra Marl Shesho- ., 1995 et al NN2 NP19–20 Coccolithus Sph. hetero- Sphenolithus Sphenolithus Martini, 1971 morphus NN5 Sp. ps.-lst. rec. distentus NP24 Nannoplankton reticulata NP22 ampliaperta NN4 T. carinatus NN1 Ch. oamar. NP18 ciperoensis NP25 subdistihus NP21 Sph. praed. NP23 Discoaster druggi S. belemnos NN3 Helicopontosphera Helicopontosphera PLANKTON ZONE N9 1979 sellii P19 sicanis N8 Globigerina Globigerina Globigerina

Blow, 1969, involuta P15

Foraminifera dissimilis N6 Globiger. am- dehiscens N5

T. cocoa. P16 Globig. semi- Globogerinita

G. trilobus N7 ma opima P21 Gl. kugleri N4 pliapertura P20 Globoquadrina correlation acc. to Berggren

tapuriensis P18

ciperoensis P22 Globigerinoides T. cen.–G.g.P17 Globorotalia opi- Stage Langh. Burdigal. Aquitan. Chattian Rupelian Priabon.

Epoch OEEOIOEEMIOCENE OLIGOCENE EOCENE Age(Ma) 20 25 30 35 Correlation of the Upper EoceneÐLower Miocene of the Paratethys and major events in the Eastern Paratethys (after Popov (after Popov in the Eastern Paratethys and major events Miocene of the Paratethys Correlation of the Upper EoceneÐLower

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002

BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S187 keninidae (Hantkenina and Cribrohantkenina), as well ary of the Kiscellian is drawn at the NP20/NP21 bound- as Turborotalia cunialensis and T. cocoaensis. Outside ary, whereas its upper boundary is the lithological the tropical zone, this boundary is difficult to determine boundary between the Kiscel Clay and the finer-grained because of the poor species composition of the Late and shallower Egerian deposits. Eocene Hanthkeninidae assemblages and dominance of globigerinids (Rögl, 1996). No major events in the evo- In Transylvania, the EoceneÐOligocene boundary is lution of nannoplankton occurred at this boundary, based on planktonic data and passes through the upper which is drawn in the lower part of Zone NP21 (after part of the Brebi Marl (Bombita and Rusu, 1981; Rusu Martini, 1971) or within Zone CP16a (after Okada and et al., 1993). However, in the Early Oligocene, shallow- Burky, 1980). The volcanic biotite from the Massig- water carbonate sediments (Hoia Limestone) still accu- nano Section is dated by the KÐAr method approxi- mulated there. These sediments yield warm-water mately at 34.3 ± 0.3 Ma (Premoli Silva et al., 1988). benthic assemblages, including typical Tethyan taxa. Dating of deep-sea cores at the P17/P18 boundary indi- The overlying sandyÐclay Mera Beds, which are fol- cates the age of 33.7 Ma (after Berggren et al., 1995). lowed by brackish-water interbeds higher in the succes- Based on dinocysts, the boundary is drawn at the base sion, yield the same fauna. For these beds, the Merian of the local Glaphyrocysta semitheca Zone, thus virtu- Regional Stage (Moisescu, 1975) was proposed. The ally coinciding with the base of Zone P18, and through lower boundary of the Merian Stage is drawn at the the middle part of Zone NP21. The Rupelian of the NP21/NP22 boundary, whereas the upper boundary is Mediterranean includes the local dinocyst interval- defined by the appearance of brackish-water mollusks zones, Areosphaeridium dictyoplokus, Reticulasphera (“Cardium” lipoldi and Janschinella). In the Getic actinocoronata (in the middle part of which Wetzeliella Depression, the change from the Eocene planktonic gochtii first appears), Chiropteridium inornatum, Hys- assemblages (Globigerina corpulenta and Coccolithus trichokolpoma pullillum, and Chiropteridium lobospi- subdistihus) to the assemblage with Pseudohastigerina nosum, at the top of which the RupelianÐChattian micra is observed in sandy facies at the base of the con- Boundary is drawn. This boundary runs through the glomerate and sandstone beds (Cheia and Gorbi after middle part of Zone P21 according to planktonic fora- Bombita and Rusu, 1981). minifer data (after Berggren) and through the upper In the Flysch Carpathians, many researchers assign part of the combined NP23ÐNP24 Zone according to the upper part of the Sheshora Marl to the Lower Oli- nannoplankton data. Based on nummulites, the Oli- gocene based on the appearance of planktonic foramin- gocene of the Mediterranean (Rupelian and lowermost ifers of Zone P18 and nannoplankton of Zone NP21 Chattian) corresponds to the Nummulites intermedius (Krhovsky et al., 1995). This is overlain by the Sub- (= N. fichtelli)–N. vascus Zone with the following typ- chert Member of the Menilithic Formation, dark non- ical species: Nummulites intermedius, N. vascus, N. fich- calcareous or slightly carbonate clay beds (6Ð10 m telli, and N. sublaevigatus (Colloque…, 1968; Cavelier thick) that were formed under low-oxygen conditions and Pomerol, 1985). and are considered by certain paleontologists to be the Western Paratethys. Within the Intra-Carpathian basal Oligocene beds (Andreyeva-Grigorovich et al., Region, in the Hungarian Paleogene Basin, the 1986). Carbonate components of the sediment are rep- EoceneÐOligocene boundary is drawn in the upper part resented there by coccoliths of the Helicopontosphera of the Buda Marl where the extinction of the group reticulata (NP22) assemblage or by smaller planktonic Globorotalia cerroazulensis, Globigerina gortanii, etc. foraminifers of Globigerina officinalis in association and the appearance of species typical of the Globiger- with the species that allow the correlation of the base of ina tapuriensisÐPseudohastigerina micra (P18) assem- the Menilithic Formation with Zone P18 and of the blage are observed at the boundary between the carbon- upper horizons with the transitional assemblage of the ate and terrigenous facies (Tard Clay). Based on nanno- P18/P19 boundary. Dinocysts of the Subchert Member plankton data, the base of the Tard Clay approximately are represented by the assemblage of the Wetzeliella corresponds to the NP20/NP21 boundary, whereas the symmetrica Zone (Andreyeva-Grigorovich et al., 1986). upper part of these clay beds belongs to Zone NP23, The Lower Oligocene Phthanoperidinium amoenum includes brackish-water mollusks, and corresponds to Zone is usually difficult to distinguish, because its the brackish-water period of the Paratethys. A thick zonal species is extremely rare in occurrence. In the horizon of the Kiscel Clay is distinguished higher in the middle and upper parts of the Subchert Member, inter- succession. This horizon is characterized by planktonic beds with abundant Limacina (= Spiratella) are foraminifers of the Globorotalia opima opima Zone observed, as well as horizons showing a decrease in and by nannoplankton of the Sphenolithus distentus salinity indicated by the composition of nannoplankton (NP24) Zone (Nagymarosy and Báldi-Beke, 1988). (the proportion of Braarudosphaera bigelowii increased) The deposits that correlate with the Rupelian (the and diatoms (the recrystallized cherts were predomi- uppermost part of the Buda Marl and Tard Clay) and nantly composed of a single fresh- to brackish-water with the lowermost part of the Chattian in the Pannon- species Melosira granulata). Marl beds at the top of the ian Region of the Paratethys were proposed as the Kis- chert member yield nannoplankton of Zone NP23 and cellian Regional Stage (Báldi, 1969). The lower bound- correlate well with the lower part of the Solenovian

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002

S188 POPOV et al.

Horizon in the Eastern Paratethys by the presence of EoceneÐOligocene boundary that is considered as the typical taxa (Krhovsky et al., 1995). Turborotalia centralis/Globigerina tapuriensis P17/P18 This brackish-water level of the Paratethyan salin- border passes in the vicinity of the abrupt lithological ity, which is characteristic of the second half of the change from the biogenous-carbonate (Beloglinian Rupelian and is well determined by the presence of Formation) to terrigenous sedimentation. The Lower brackish-water mollusks, monospecific rock-forming Oligocene is represented almost everywhere by dark- assemblages of nannoplankton, diatoms, and dinocysts colored noncarbonate or slightly carbonate clay of the of a typical composition, is a marker level for the Oli- Pshekhian Horizon (lower part of the Maykopian For- gocene. This is traced everywhere (Voronina and mation). Zonal species of planktonic foraminifers usu- Popov, 1984; Báldi, 1986; Rusu, 1988; Nagymarosy ally are absent in the Maykopian Beds. In the dinocyst and Voronina, 1993) from Bavaria (Heller Mergelkalk) scale, the EoceneÐOligocene transition is correlated and Slovenia (Sotzca Beds) via Hungary (upper part of with the boundary between the Wetzeliella clathrata the Tard Clay), Transylvania (Bizusa, Ileanda, and Cio- angulosa and Phthanoperidinium amoerum zones. The cmani), the Carpathians (Dynow Marl, Tylawa, and latter corresponds to the lower part of the Pshekhian Tarcau), southern Ukraine (Molochnaya Beds), Ciscau- Horizon, overlain the vast Wetzelliella gochtii sensu casia (Polbian Horizon), up to the northern Aral Sea lato Zone, which includes beds of the Solenovian Hori- coastal area and Central Asia (“ostracode layer” of the zon. Its lower half occasionally may be considered as Solenovian Horizon). the W. simmetrica Zone (Zaporozhets, 1993). Based on benthic foraminifers, the Lenticulina herrmanni regional In the Carpathians, the Upper Rupelian includes the Zone is distinguished in the lower part of the Pshekhian overlying sediments of the Menilithic Formation (the Horizon (Reshenie…, 1970). Higher, but not every- Sitborice Member in Czech Republic, the Lower where, the Spiroplectammina oligocenica (= S. cari- Menilithic Subformation in the Ukrainian Carpathians, nata oligocenica) Zone is distinguished. This zone and the Lower Dysodilic Shales in Romania). These includes Caucasina schischkinskayae, Cyclammina beds include both brackish-water and polyhaline sedi- constrictimargo, Bolivina mississippiensis, and a few ments, of which the latter are defined by the presence of sand-dwelling taxa. The boundary between them is not nannoplankton (NP23/NP24), planktonic foraminifers exactly synchronous, has a facial character, but is of Zone P21a, and diatoms and dinocysts of the Wetze- clearly traced in the majority of regions. liella gochtii Zone (Andreyeva-Grigorovich et al., 1986; Rusu et al., 1996; Krhovsky et al., 1995). Opinions on the nannoplankton zonation differ. The difficulties in the recognition of Zone NP21 in this In the Alpine Foreland Basin, the Upper Marine place were discussed above. In the upper half of the Molasse (UMM), represented by shallow-water facies Pshekhian Horizon, all researchers distinguish the and sands, is mainly dated as the Rupelian. The sedi- Helicopontosphere reticulata (NP22) Zone, whereas, at ments are characterized by planktonic foraminifers of the base of the Solenovian (Polbian) Horizon, an indi- Zones P17ÐP20, nannoplankton (NP21ÐNP24), and vidual assemblage of endemic mollusks, nannoplank- dinocysts and mammals (MP-12 after Berger, 1992, ton, and dinocysts was discovered, thus indicating a 1996). These are traced in the Jura Mountains, as well as change in salinity regimes. The Polbian Marl is usually in the Rhine Graben System, where their upper part dated at the Sphenolithus praedistentus NP23 Zone already belongs to the Chattian (NP25 after Berger, 1992). (Nagymarosy and Voronina, 1993) or at the NP22/NP23 Outside the Western Paratethys, within the study transition (Muzylev and Tabachnikova, 1987). The area, calcareous plankton assemblages that are repre- lower boundary of the Wetzeliella gochtii sensu lato sentative enough for reliable recognition of zonal closely corresponds to the base of the Spiroplectam- Rupelian subunits are only found south of the Lesser mina oligocenica Zone. In the Eastern Paratethys, the Caucasian land, in Armenia. In the reference section of composition of dinocyst assemblages markedly vary the Biralu Borehole, the level of disappearance of the across the single Wetzeliella gochtii Zone because of rosette-shaped discoasters (at the NP20/NP21 bound- the Solenovian drop in salinity. The lower assemblage ary) almost corresponds to the change of the Turbotota- is defined by mainly polyhaline taxa, whereas the upper lia centralisÐGlobigerina gortanii Zone to the Globige- assemblage is defined by euryhaline or brackish-water rina tapuriensis Zone (P17/P18) dominated by small- taxa. In the West European sections, the zones proposed sized species of newly appeared globigerinids. The correspond to Zones D13 and D14 (the latter zone con- Globigerina sellii Zone is recognized higher in the suc- sists of two subzones, D14a and D15a). cession, where closely similar species in association After the Solenovian drop in salinity, the salinity in with rare specimens of the index species are found the Eastern Paratethys began gradually to increase to (Geologicheskie i bioticheskie…, 1998). This zone is restore the polyhaline groups of organisms. Usually, overlain by sand beds with an impoverished assem- this event was dated based on mollusks and benthic for- blage of foraminifers, which probably belong to the aminifers at the beginning of the Chattian (Ter-Grigo- Globigerina ampliapertura Zone. ryants, 1964; Stolyarov, 1991; Popov et al., 1993). Eastern Paratethys. In the Crimea, Ciscaucasia, However, recent data on phytoplankton indicate that Transcaucasia, Mangyshlak, and Turkmenistan, the this event should be assigned to the terminal Rupelian,

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002

BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S189 as in the Carpathians. Nannoplankton of the upper part (Popov et al., 2001). Eocene nannoplankton and radi- of Zone NP23 was recorded by Khrovsky, whereas olarians were also reported (Muzylev et al., 1996). dinocysts of Zone D14a were identified by Zaporozhets Nannoplankton of the I. recurvus (NP19) Zone and from carbonate interbeds at the base of the Upper Sub- Zones NP18ÐNP20 was recorded by Andreyeva-Grig- formation of the Morozkina Balka Formation in the orovich (see Starshinin, 1997) from the upper part of Belaya River succession (Akhmetiev et al., 1995). the Kushan Horizon; however, an impoverished Oli- In Transcaucasia, there are more complete assem- gocene assemblage with Reticulofenestra umbilicata, blages of planktonic foraminifers with Globigerina R. scussira, Cyclocargolithus floridanus, Cyclococco- tapuriensis, G. ampliapertura, and Globorotalia per- lithus formosus, and Sphenolithus prenistanius (NP22) micra (P18); nannoplankton of the Coccolithus subdis- was found in the upper part of the Sanglak Horizon in tichus (NP21) Zone; mollusks of the typical Rupelian the Manzob Section. assemblage; and numerous remains of Limacina (= Spi- A benthic foraminiferal assemblage from the Sang- ratella and Planorbella). The Lower Oligocene beds in lak Horizon have quite dissimilar composition com- this zone are relatively thin, a few tens of meters thick pared to the Late Eocene assemblages, including Loithos- (up to 100Ð150 m in the Kura–Iori interfluve). tomata macrura, Asianella subbotinae, Brotzenella In Transcaspia, the Lower Oligocene marine beds munda, Cibicidoides khanabadensis, and a few new defined by nannoplankton of Zone NP22 and dinocysts species of the genus Brotzenella, which are unknown are traced up to the Western Siberia and Tajik Depres- from the other regions of Central Asia prior to the Oli- sion (see the section Paleogeography of the Paratethys gocene (Davidzon et al., 1982). Asianella subbotinae, for more detail). Brotzenella munda, Cibicidoides khanabadensis and In shallower-water Lower Oligocene facies of sand-dwelling species of the genus Trochammina southern Ukraine, Volga-Don Region, Mangyshlak, (Varzyk Section after Bugrova) described by Davidzon Ustyurt, northern Aral Sea coastal area, and Kopet- from the Sanglak Horizon were also recorded from the Dagh, fossil planktonic assemblages are either rare or Sumsar red-colored beds in Ferghana. A closely similar represented by the transient species Globigerina offici- assemblage was reported by Minakova (Paleogen- nalis. However, the Lower Rupelian marine beds are ovaya…, 1975, p. 275) from the crimson clay imbedded rather reliably recognized by the typical benthic taxa: in a horizon with Ostrea pygmaea in Ferghana. This mollusks, including species zonal for the Rupelian assemblage was compared with the fauna of the Lenti- sensu stricto of the Stratotype Region (Pecten stetten- culina hermanni Zone. It is impossible to compare this ensis), and foraminifers in common with the Rupelian fauna with the Beloglinian Assemblage, because it fauna. The Solenovian Horizon may be also used as a developed after the crisis at the EoceneÐOligocene marker level at the top of these beds. This horizon is boundary, according to Bugrova, and is discussed in easily distinguished by having a lighter color of rocks this section of the present study. and the presence of carbonate clay imbedded in the dark-colored noncarbonate Maykopian Formation, benthic brackish-water molluskan and ostracode faunas Paleogeography of the Paratethys (Popov et al., 1985), and of a specific nannoplankton assemblage with Reticulofenestra ornata and Transver- The Late Eocene Orogeny of the Alpine folded belt sopontis fibula. The dinocyst assemblage becomes and the eustatic regression transformed the Early Oli- more impoverished. Two or three species predominate, gocene Paratethys into a much more isolated sea basin comprising 95% of the total number of specimens. The compared to the Eocene marginal sea basins in this zonal species Wetzeliella gochtii is extremely rare. area. The uplift of the Alps, Dinarides, Anatolia, and the A break in the marine sedimentation that corre- Lesser CaucasusÐKopet Dagh formed a continental sponds to the Oligocene base is usually observed in barrier that separated the Paratethys from the Mediter- peripheral areas of the basin. Dinocyst assemblages ranean area of the Tethys. In the Early Oligocene, the from the Ergeni Highland, Mangyshlak, Ustyurt, and Paratethys still was connected with this area in the west Turgay are abnormal in composition due to river water via the Alpine Foreland Basin (Berger, 1996) and Slov- influxes and unstable facial environments. The Lower enian Corridor, and probably in the east via the VardarÐ Oligocene Phthanoperidinium amoenum and Wetze- Rhodope Area (Fig. 1). It also was connected with the liella gochtii zones are easily recognized there, whereas North Sea Basin (based on faunal data, see below) via the W. simmetrica Zone may be only preliminary distin- the Rhine Graben, as well as via the DnieperÐDon and guished because of the absence of the index species. Prypat depressions. The available data on the stratigraphic position and The Early Oligocene Paratethys markedly decreased age of the Sumsar Horizon in Ferghana and the Sanglak in area along the entire periphery compared to the Late Horizon in the Tajik Depression are rather contradic- Eocene basin, whereas the deepwater part of the Parat- tory. On the one hand, the molluskan and ostracode ethys inherited its major character of the paleobathymet- assemblages are closely similar to the Chegan assem- ric zonation. Its bathyal and shelf depressions became blages and were discussed in the first part of this study even deeper (Stolyarov, 1991; Shcherba, 1993).

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S190 POPOV et al. ) mountains; ) continental India 6 14 Central Depression Land High North Pamir Abbreviation: Pamir Basin Sea Depression Tarim ALB-Alboran Land CAL-Calabria DH-Donets High EBD-East Black Sea KAB-Kabilia LC-Lasser Caucasus MC-Massif Central PEL-Peloritanian WBD-West Black ) limestones; ( 5 OCEAN Dzungar Basin Land INDIAN Central ) deepwater shelf; ( ) deepwater Afghanistan 13 Basin Tajik ) marls; ( High Ferghana-

4

d d d

North Tienshan n n n

a a a

L L L

h h h

) clays; (

g g g

3

a a a

High

d d d

t t t

n n n

e e e

a a a

p p p

r r r

n n n

I I I o o o i i i

) shallow shelf; ( ) shallow

s s Kazakhstanian s

l l l

K K K

SEA

a a a

- - -

a a a

12

s s s

r r r

B B B r r

r ) sands; ( t t t

u u

u Basin

2 n n n

o o o

e e e

b b n b

l l

i l C C C

TURAN E E s E Strait Turgay

a Iran Land

B

n

a

i ) shifts.

s

a 20 c High

u Ural

a

C ) conglomerates; (

r 1 LC

e Mesopotamian

t

a 0 500 km

e ) thrusts; (

r

G 19 EBD Land DH LAND 19 20 Anatolian RUSSIAN WBD ) lagoons episodically covered by the sea; ( ) lagoons episodically covered Basin ) normal faults; ( ) normal faults; 11 18 h

g 16 17 18 i

H Land d

n n Moesian a a i n L y

Basin s h l e o d i V r 13 14 15 a East Mediterranean n

i basins; ( ) lake Basin ) volcanoes; ( ) volcanoes;

D Sirt 10 17 Alpine-Carpathyan Apulia High 10 11 12 High ) flysh; ( Bohemian Alps Fenno-Scandian 16

) lowlands; ( ) lowlands; h

9 g

u

o CAL r

7 8 9 T

r

e SEA

PEL

o

r

NORTH

e

a F KAB MC

4 5 6

h

g

u

o r

) terrigenous deposits; (

T ALB

l l

15 a 1 2 3

Atlas k High c

) nondifferentiated land; ( ) nondifferentiated o

Land R 8 Iberian Bay of Biscay Paleogeographic map of the Tethys and Northern Peri-Tethys in the Rupelian. Explanations: ( and Northern Peri-Tethys Tethys map of the Paleogeographic 50° 40° 30° 20° ) highlands; ( 7 slope and bathyal; ( slope and bathyal; ( Fig. 1. Fig.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S191

In the Rupelian, the depositional environments and iments were represented by thin (a few ten meters the faunal living conditions in the Paratethys markedly thick) black-colored and dark-grayish noncarbonate changed due to the progressive isolation, overdeepen- mud layers. Strong fluxoturbidites arrived to these ing of the basin, water cooling, and the presence of a troughs from the East European Platform, as well as strong river influx. This resulted in the formation of the from the Silesian Cordillera and Inner Eastern Car- thermal and salinity water stratification, impairment of pathians (Fusaru Sandstone and Otrit Sandstone). Two gas exchange regime, hydrogen-sulfide contamination, deep depressions, the Magura Trough and the Szolnok and accumulation of dark-colored clay sediments with Trough, were still located southwest of the cordillera high content of undecomposed biogenous material that bordered the axial trough. These depressions were (Tard Clay, menilithic beds in the Carpathian Area and compensated for by black-colored mud and diatomites. the Maykopian Facies in the CaucasusÐKopet Dagh In the Rupelian, the Szolnok Trough decreased in area Area of the Paratethys). due to its shallowing and movement of the Bukk Volca- At the end of the Rupelian, this isolation resulted in nic Ridge in the southward direction (Nagymarosy and a change in the hydrological regimes of the basin and in Báldi-Beke, 1993). the accumulation of specific sediments containing The southern shelf occurred within two large endemic fauna characteristic of brackish-water envi- blocks, the Alcapa (Tatran Unit) and the Tisza-Dacia ronments. Marine environments were restored in these (Bihor Unit). The eastern area of the Alcapa was occu- basins in different times. In the Carpathian Basin, pied by the central Carpathian Paleogene Basin and its marine fauna already appeared in the second half of the continuation to Slovakia. This area increased in depth Rupelian (in the Early Menilithic Epoch), whereas, in toward southeast, where the Tard Clay (60 to 100 m the Eastern Paratethys, the appearance of polyhaline thick) accumulated in a basin of the euxine type with marine fauna is dated at the Terminal RupelianÐEarly traces of hydrogen-sulfide contamination. This basin Chattian. still continued into the North Italian Sea via the Slove- nian Corridor (Báldi, 1986; Nagymarosy, 1990). This AlpineÐCarpathian Basin connection was interrupted in the second half of the Rupelian. At the beginning of the Rupelian, the hydrological The eastern part of the Tisza-Dacia Block was occu- regimes of the central area of the residual marginal sea pied by the Transylvanian Shelf Sea with carbonate basin in the Carpathians remained relatively favorable sedimentation (Hoja Limestone and sandy-clay Mera for life. This area was inhabited by marine benthic Beds). Shallow-water facies occurred in the western fauna, although impoverished. These beds yield fora- area of this basin and bordered the Apuseni Land, from minifers, sponges, traces of echinoderms and mud-eat- which terrigenous and biogenous material arrived. In ers (possibly worms), as well as fossil abyssal fishes the southeast, the Transylvanian Basin joined the shelf with photophores, which indicate the sea depths of at area of the southern part of the deepwater Carpathian least a few hundred meters. The increasing isolation of Depression. In the southwest, this basin rapidly became this basin subsequently resulted in salinity stratifica- shallower and clay was replaced by sand and then by tion, periodical anoxic environments, and the disap- conglomerate of the Getic Depression, the products of pearance of the deepwater fauna. Growing hydrother- erosion of the South CarpathianÐMoesian Land. mal activities that were accompanied by the supply of nutrients (nitrogen, phosphorus, and silica) resulted in The South CarpathianÐMoesian Land and the the acme of diatoms in the Middle Rupelian (at the bor- Dinarides were probably separated by straits that con- der between the nannoplankton zones NP23 and NP24). nected the Carpathian Basin and the Rhodopian Fore- The diatoms were originally represented by marine land Basin, a gulf of the Ancient Mediterranean. This is taxa (Krhovsky et al., 1993) and subsequently by indicated by the available biogeographic data, the com- brackish-water taxa, whereas their assemblage became position of benthic fauna of Mediterranean origin that monospecific (Melosira granulata and Aulacosira). persisted in Transylvania up to the Solenovian drop in Subsequently, the dominance of diatoms changed to the the Paratethys salinity, as well as by the common geo- mass development of typical Paratethyan nannoplank- logical evolution of Transylvania and northeastern ton (Reticulofenestra ornata and Transversopontis fib- Greece, where the Lower Rupelian marine beds are also ula), while the sediments were carbonate everywhere overlain by brackish-water sediments with Corbula. (Nagymarosy and Voronina, 1993). At the end of the These straits are also traced in Serbia (And– elkovic′ et al., Rupelian, marine environments were restored and accu- 1991). In the Oligocene, they became probably even mulation of dark-colored noncarbonate mud (Lower deeper compared to the Eocene. The same data indicate Menilithic, foliated, bituminiferous shale beds) started. that the Inner (Eastern) Dinarides represented a slightly In the Early Oligocene, the Skole-Zdanice-Silesian elevated land with depressions compensated for by (Tarcau) troughs and the Skibas-Pokut Trough were freshwater, lagoonal, and volcanic sediments. still the deepest-water depressions (Beer and Shcherba, In the Oligocene, the Carpathian Foreland mainly 1984), in which condensed sediments accumulated represented a denudation area (Kul’chitskii and below the level of carbonate compensation. These sed- Sovchik, 1986). The northern shelf was probably very

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S192 POPOV et al. narrow and joined the Sadomir Uplift, which was sub- In the Rupelian, the southern shelf was strongly ject to erosion. The sediments of this shelf were only affected by a newly formed land. The uplift and drying found in the Alpine Foreland Basin. These are repre- of the Lesser-CaucasusÐIranian Island Arc resulted in sented by the lower marine molasse, i.e., by shallow- the accumulation of thick, coarse, clastic beds that bor- water clay beds. Farther to the northwest, the clay beds der the modern-day Lesser Caucasian Arc and repre- are replaced by sandyÐcarbonate lagoonal and lake sent clay, sandstone, and conglomerate formations of beds (Berger, 1996). According to Berger, there was no the Akhaltsikhe Depression (up to 1000 m thick), marine connection between this shelf and the North Sea Gyandja Region in Azerbaijan (400 m thick), and the Basin in the Rupelian. However, the available faunal Araks Depression (up to 700 m thick). These beds data (on mollusks) indicate that there existed at least occasionally bear traces of sand streams, mudflow episodic connections (see below). The existence of a intrusions, and lagoonal environments (Akhaltsikhe). certain connection between the Carpathian Shelf and The Lesser CaucasusÐIranian Land represented a the Greater Caucasian basin probably via the Gelati low-mountain structure that was occasionally subjected Strait is also indicated by the biota in common, espe- to the early orogenic volcanism (of the acid or interme- cially by the endemic taxa of the Solenovian Epoch diate types) and was occupied by laurelÐbeech forests (mollusks, ostracodes, and nannoplankton). with a participation of palms. The plant fossils were preserved in tuffÐsandstone beds of the Negram Plateau Greater CaucasusÐKopet Dagh Basin (Akhmetiev and Zaporozhets, 1989). At the beginning of the Oligocene, the deepwater gulf of the Central Ira- The Greater CaucasusÐKopet Dagh Basin (Eastern nian Basin reached this land in the south via the Yere- Paratethys) was connected in the first half of the Rupe- vanÐOrdubad Depression. The coastal zone with nor- lian with the Carpathian Basin in the region of the Moe- mal marine environments was inhabited by various sian Plate and was open to the North Sea Basin (Fig. 1). colonial corals, numerous warm-water mollusks, small Concurrently, the uplift of the Lesser CaucasusÐIranian foraminifers, nummulitides, echinoids, ostracodes, and Land entirely separated this basin from the Mediterra- nannoplankton; and sandy-mud sediments (up to 1000 m neanÐMesopotamian Basin already by the Late Eocene. thick) accumulated there. In the second half of the The closure and shallowing of the Eastern Paratethys at Rupelian, the sea left this region. the beginning of the Solenovian Epoch resulted in its short-term transgression, improvement of gas exchange The northern shelf was very wide and included a regime, formation and wide distribution of endemic complex system of deep and wide depressions that benthic fauna over the entire Paratethys, and the wide were not yet compensated for by the Oligocene sedi- accumulation of biogenous (nannoplankton) carbonate mentation (Stolyarov, 1961, 1991). These depressions mud. In the Late Solenovian, the basin again decreased had their individual pseudo-oceanic structure, a shal- in area. Nevertheless, its connection with the World low-water shelf, and fairly steep slopes. The TerekÐ Ocean became wider and euryhaline representatives of Mangyshlak Depression was the deepest, where thin the marine fauna could reach the Paratethys, but its gas carbonate clay beds (40Ð20 m thick, or smaller) accu- exchange regime was deteriorated and the sediments mulated with remains of planktonic foraminifers and were saturated with organic material. bathyal fishes with photophores. The second largest depression was the IndolÐKuban Depression separated The deepwater zone still occurred within the East- from the TerekÐMangyshlak Depression by the ern and Western Black Sea depressions, LazarevskoeÐ Stavropol Uplift. This depression was situated much Kobystan Trough, and the South Caspian Depression closer to the sources of terrigenous material; thus, the (Shcherba, 1993; Kopp and Shcherba, 1998). The Oli- Lower Oligocene beds were there 300Ð400 m thick. goceneÐEarly Miocene sediments of this part of the Farther to the west, in the region of the Kerch Strait, the basin are represented by homogenous clay beds that are IndolÐKuban Depression was opened into the axial area difficult to subdivide. At the beginning of the Oligocene, of the basin. the LazarevskoeÐKobystan Deep-water Trough was approximately 200 km wide, similarly to that in the Spore and pollen data and leaf flora from Ciscauca- Eocene (Shcherba, 1989). Farther south of the Schatsky sian outcrops indicate an increase in the land area in the Swell, the deepwater sediments accumulated in the western part of the Greater Caucasus (Zaporozhets, Western and Eastern Black Sea depressions (Tugolesov 1993; Akhmetiev, 1995). This is also indicated by the et al., 1985), where these are known only based on geo- occurrences of coastal molluskan assemblages at two physical data as a component of the undivided Mayko- stratigraphic levels, at the base of the Oligocene and in pian Formation. There, the Oligocene sediments are the lower part of the Solenovian Horizon (sections of very shallower-water in character compared to the the Belaya, Fars, and Kuban rivers). Similar uplifts Eocene for the reason of the Lesser Caucasus Orogen- probably existed in the vicinity of the southwest flank esis and the decrease in the trough wide. The deepwater of the TerekÐMangyshlak Depression, where shallow- Talysh Foredeep is an equivalent to the Eastern Black water sandy sediments with molluskan assemblages of Sea Depression in the east, however, separated from it by the Solenovian Horizon were found (Northern Osetia, a shallower-water plateau (central and eastern Georgia). Fiagdon and Tsraudon rivers). These depressions were

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S193 also bordered in the north by fairly steep slopes, as indi- sediments. The maximal sea transgression was cated by the sharp changes in facies and thickness. This observed in the Middle Ascheairyk when the sea con- is the Mangyshlak Island, as well as the uplifts west of tinued via Turgay into the Middle Trans-Uralian Astrakhan and the Rostov Uplift (Stolyarov, 1991). depressions and the Omsk Depression (marine Rupe- In the western part of the Greater-Caucasus Basin, lian sediments were found along the southeast-facing on the slope of the Moesian Platform, in the outer shelf slope of Trans-Urals, in the vicinity of the town of Kur- environments, noncarbonate mud with pteropods, typi- gan and farther east, Akhmetiev et al., in press). In the cal Paratethyan molluskan species (“assemblage with Terminal Ascheairyk (the lower part of the Wetzeliella Lucina batalpaschinica”) and fish fossils accumulated. gochtii D14a Zone), the sea retreated, and the sandy In the south, the Moesian Shelf was bordered by the coastal facies of this epoch are known in the Northern zone of Balkan thrusts. In their southern flank, in the Ustyurt. Sredna Gora and Rhodopian Upland, the Lower Oli- gocene is represented by quite different facies: the At the Eocene-Oligocene boundary, the sea coastal beds with a rich warm-water fauna represent retreated from the territory of Kyzyl Kum and south- sediments of Mediterranean gulfs (Karagyuleva, 1964). eastern Kara Kum. In the Kyzyl Kum, the denudation Available data on marine biota indicate that, starting cutoff reached 100Ð150 m (Pinkhasov et al., in Geolo- from the Terminal Eocene, these basins were separated gicheskie i bioticheskie…, 1996). Subsequently, in the by a continental barrier rather than connected with each first half of the Early Oligocene, the Kara KumÐKyzyl other. This land continued into the modern-day south- Kum Lowland was occupied by a shallow-water sea, western area of the Black Sea Basin and probably whereas the red-colored and variegated silty-clay beds served as a continental bridge, via which Asian verte- of the Sarybatyr and Murghab formations accumulated brates could reach southeastern Europe (see below). and were a few ten meters thick. At that period of time, Farther east of the Moesian Land, the shelf sharply the short-term sea transgression could probably reach increased in width. In the northwestern area of the the Ferghana and Tajik depressions, where Oligocene Black Sea Basin and the South Ukrainian Depression, foraminifers were found, as well as nannoplankton of typical coastal facies occur near the south slope of the the Helicosphaera reticulata (NP22) Zone with a char- Ukrainian Shield, which had transformed from an acteristic set of species (Manzob Section). This nanno- archipelago to the land area by the Oligocene. This land plankton was determined by Andreyeva-Grigorovich area separated the South Ukrainian Depression from a from the southern area of the Hissar Upland. As in the shallow-water epicontinental basin that occupied the Carpathians, the acme of the species Cyclocargolithus DnieperÐDon and Pripet depressions, via which the floridanus was found there. A narrow shallow-water Paratethys was connected with the North Sea Basin. marine strait markedly affected by a fresh-water influx The entire vast area of marine sedimentation in probably continued farther east into the Tarim Depres- Transcaspia, i.e., the Turanian Sea Basin, represented a sion. Here, red clay and fine-grained terrigenous beds shelf zone with depressions that were mainly compen- of the Bashibulake Formation with the endemic mol- sated for by sedimentation. Only the axial area of the lusks Platigena, Ferganea, Cubitostrea, and Donax Southern Mangyshlak Depression, Buzachi Depres- and with the cosmopolitan species Chlamys, Anomia, sion, and the DaryalykÐDaudan Depression may be and Crassatella accumulated (Lan and Wei, 1991). The assigned to the deep shelf zone. In the DaryalykÐ tree lower members of this formation are dated by these Daudan Depression, fossil material is only represented researchers at the Late Eocene, whereas the two upper by globigerinids and fish fossils, thus indicating the gas members are assigned to the Early Oligocene. exchange regime unfavorable to the benthos and the sea depths of over 150Ð200 m. The southwestern part of the Farther to the northeast, the coastal beds of this zone Krasnovodsk Peninsula and the Western Kopet Dagh were gradually imbedded by terrigenous and alluvial also represented a relatively deep area of the Turkme- sediments and then became entirely terrigenous. The nian Basin, where planktonic foraminifers accounted presence of gypsum in some red-colored interbeds indi- for up to 65% of the paleobiocoenoses (Nevmirich et al., cates a hyperhaline lagoonal paleoenvironment. The in Geologicheskie i bioticheskie…, 1996). Mugodzhar Hills Land, Kazakhstanian Shield Land, The rest of the area belonged to the shallow-water and the Tien Shan, Altai, Hissar, and Kopet Dagh shelf with gas exchange regimes relatively favorable for formed a zone of steady uplift in Transcaspia and rep- benthic fauna. Its richest assemblages were found in the resented a source of terrigenous material. north, in the Northern Ustyurt and northern Aral Sea coastal area (Merklin, 1974; Amitrov, 1993). In the Ter- The tectonic rearrangement and the orogenesis also minal EoceneÐEarly Oligocene, the sea regression and started farther east, in Central Asia, as indicated by the lowering of the erosion basis resulted in deep erosion changes in the lithological structure of beds at the level (up to 40Ð60 m in the northern Aral Sea coastal area). of the Buranian Formation in the Zaisan Depression, as Subsequently, the depressions formed were compen- well as by the transformation of its freshwater ichthyo- sated for by freshwater, terrigenous, and marine sandy fauna and molluskan fauna.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S194 POPOV et al.

Biogeography of Water Bodies the foraminiferal assemblages and facial environments, Planktonic Foraminifers but investigations of this type have not yet been per- formed. The Oligocene planktonic foraminiferal fauna dif- fers markedly from the Eocene assemblages in having The second factor that makes difficult the analysis of a poorer composition. This fauna includes representa- the biogeographic distribution of planktonic foramini- tives of globigerinids and globorotaliids of mainly fers in the Oligocene is associated with geological evo- small and medium size. The taxonomic diversity seems lution in this epoch. The development of eustatic and to increase when we use the modern-day classification, tectonic processes resulted in a wide accumulation of which diminishes the volume of generic units. The regressive terrigenous (clayey and sandy-clay) sedi- assemblage of planktonic foraminifers includes ten ments over continents and in the formation of the Parat- genera: Globigerina, Subbotina, Cassigerinella, Tenui- ethys and anoxic hydrochemical regimes. As a result, it tella, Tenuitellinata, Catapsydrax, Paragloborotalia, is very difficult to differentiate which factor affects the Globorotaloides, Pseudohastigerina, and Chiloguem- composition of foraminiferal assemblages, the primary belina. The species composition of the Oligocene general factor (climatic zonation), the local factor (bio- planktonic foraminifers is of limited diversity (about chemical environments for the sedimentation), or the 40 species), but it should be emphasized that the spe- secondary taphonomic factor (selective dissolution of cies of Globigerina, Subbotina, and Paragloborotalia the fossils in non-carbonate or slightly carbonate sedi- predominate, while the representatives of Classiger- ments). inella, Globorotaloides, Tenuitellinata, and Chiloguem- The available data on the distribution of the Oli- belina are rare, and the species of Pseudohastigerina gocene planktonic foraminifers only allow an estimate are entirely restricted to the Lower Oligocene. of the general biogeographic pattern of this group of Such a decrease in the taxonomic diversity of the microfauna but not always the factors that form the planktonic foraminifers is usually associated with the individual taxonomic composition. climatic changes (climatic cooling) at the EoceneÐOli- In the study area, we can distinguish two provinces, gocene boundary. Actually, a number of typical Eocene the Southern (Mediterranean) and the Northern genera and groups of species, (Cribrohatkenina, Hant- (CrimeanÐCaucasian Region, Carpathians Foreland, kenina, Globigerapsis, Globigerinatheca, and the and northwest Europe). group “Globorotalia” cerro-azulensis), disappear (or decrease in number) at the lower boundary of the Para- In the Southern (Mediterranean) Province, rich globorotalia centralisÐSubbotina gortanii Zone, which assemblages of planktonic foraminifers occurred in corresponds to the transitional EoceneÐOligocene beds. facies of true white chalk (northeastern Syria) or in car- At this boundary, the terrestrial flora (spore and pollen bonate clay facies (Cyprus and Italy). The Oligocene assemblages) indicates a climatic change toward cool- sections include the entire set of the foraminiferal ing. At the upper boundary of the Paragloborotalia zones, Subbotina tapuriensis, Subbotina selii, Globige- centralisÐSubbotina gortanii Zone (i.e., EoceneÐOli- rina ampliapertura, Paragloborotalia opima, Globige- gocene boundary), the last elements of the Eocene rina ciperoensis, and Paragloborotalia kugleri. A dis- fauna of planktonic foraminifers disappear, and the tinctive character of the foraminiferal assemblages is Eocene assemblages are replaced by a typical Oli- the constant presence of these index species in combi- gocene foraminiferal assemblage of the Subbotina nation with Subbotina gortanii (Bors.), S. galavisi tapuriensis Zone. At this level, a further temperature (Berm.), S. pseudovenezuelana (Blow et Bann.), S. pra- decrease occurred, as indicated by the terrestrial flora. saepis (Blow), S. tripartita (Koch), Globigerina anguliof- It is not improbable, however, that such a transfor- ficinalis Blow, G. officinalis Subb., G. angulisuturalis mation of the planktonic foraminiferal fauna at the Bolli, G. euapertura Jenk., and Tenuitellinata angus- EoceneÐOligocene boundary was associated with the tiumbilicata (Bolli). The total taxonomic composition individual evolution of this group of microfauna. In of these assemblage is closely similar to that from the subsequent Neogene epochs, the climatic conditions Oligocene of the classic tropical Caribbean Region and were very unstable and the temperature regime was the adjacent regions in the Atlantic Ocean. Possible dis- often more severe compared to that in the Oligocene, tinctions between the assemblages may be only found but the taxonomic composition of the Neogene plank- based on the proportion of species, as mentioned above. tonic foraminifers was much richer compared to the In the territory of the former Soviet Union, southern Oligocene assemblages of this group of microfauna. Armenia (Yerevan Basin) may be probably assigned to The factor indicated (the impoverished taxonomic the northern periphery of the Mediterranean Basin. composition) makes very difficult the reconstruction of There, the Lower Oligocene beds (the Subbotina tapu- the Oligocene biogeographic zonation based on plank- riensis and Subbotina sellii zones) usually include tonic foraminifers. The biogeographic zoning at the S. tapuriensis (Blow et Bann.), S. tripartita (Koch), generic taxonomic level is not efficient. It is certainly S. prasaepis (Blow), and Globigerina ampliapertura possible to reconstruct the paleobiogeographic zona- Bolli, as well as few specimens of Subbotina sellii, but tion with due consideration of specific proportions in globigerinids of small size start to predominate in quan-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S195 tity. Unfortunately, more recent Oligocene beds are rep- composition of the genera Sphenolithus, Heli- resented there by continental facies. cosphaera, Reticulofenestra, etc. changed. In the Oligocene of the Northern Province (Parat- Northern Mediterranean. In the Mediterranean, ethys), noncarbonate and slightly carbonate clayey sed- the Early Oligocene nannofloral assemblage mainly iments (Maykopian and Menilithic formations) with consisted of species of the genera Discoaster, Spheno- very impoverished assemblages of planktonic foramin- lithus, Helicosphaera, Reticulofenestra, and Cyclicar- ifers predominate. More precisely, these assemblages golithus. The Early Oligocene nannoplanktonic assem- occur in the basal part of the Oligocene, whereas, in the blages from central and northern Italy include 38 spe- overlying beds, the foraminifers are very rare or cies (Roth et al., 1971). Their distinctive character is entirely absent. A similar pattern is observed in the Oli- the presence of warm-water discoasters, D. rufus, gocene of the Northern Caucasus (Russia), northern D. woodringi, D. adamanteus, D. cubensis, and D. din- Aral Sea coastal area (Kazakhstan), Turkmenistan, quarensis, as well as Sphenolithus, including S. pseu- Azerbaijan, Georgia, the Crimea, and the Fore-Car- doradians, S. distentus, S. predistentus, S. ciperoensis, pathians (Ukraine). Clearly, it is impossible to use this etc. This assemblage indicates southern subtropical group of microfauna in the zonal division of the Oli- environments. gocene beds. The planktonic foraminiferal assemblage At the beginning of the Early Oligocene, in the is composed of small Globigerina officinalis Subb., Armenian Gulf of the TurkishÐIranian Basin, the nan- G. ouachitaensis Howe et Wall., G. praebulloides noplankton assemblage included 24 species (Andreyeva- Blow, G. postcretacea Myatl., Tenuitella danvillensis Grigorovich, 1981; Krasheninnikov et al., 1985). Almost (Howe et Wall.), T. liverovskae (Bykova), T. evoluta all of the species were in common with the Italian spe- (Subb.), T. brevispira (Subb.), Paragloborotalia permi- cies, with the exception of the species of the genus Dis- cra Blow et Bann., Pseudohastigerina micra Howe, coaster. The latter are represented by isolated speci- P. barbadoensis Blow, and Chiloguembelina cubensis mens in the Armenian assemblages. One more tropical (Palm.). The presence of separate representatives of genus, Sphenolithus, was abundant in the Armenian Globigerina ampliapertura Bolli, Subbotina prasaepis Basin, except for the species S. ciperoensis, which (Blow), and a few other species does not change the appeared in the Late Rupelian and was absent in Armenia. general pattern. Eastern Mediterranean. The Late Rupelian Meso- Similar assemblages of planktonic foraminifers potamian Basin (Tavda Section, Syria) was dominated were also found in the Oligocene clayey sediments in by the species Sphenolithus distentus and S. ciperoen- northwestern Europe (Germany, Belgium, and the sis, as well as Helicosphaera recta and Cyclicar- Netherlands). According to some studies on the Oli- golithus abisectus (Müller, 1971). The nannofloral gocene of Europe, the assemblages of small globigerin- assemblage was almost entirely similar to the Early ids and globorotaliids should be considered as a general Oligocene flora from Trinidad Island (Bramlette and characteristic of the Oligocene fauna of planktonic for- Wilcooxon, 1967); i.e., it was of the circumtropical type. aminifers, but this is contrary to facts. North Sea Basin. The development of the coccoli- Supposedly, the major factor for such essential dif- thophorid flora in the basins of western and northern ferences between the planktonic foraminiferal assem- Europe was strongly affected by Atlantic cold waters. blages from the Oligocene of the Southern and the There, thermophilic discoasters were almost absent, Northern provinces is the climatic zonation; however, representatives of Sphenolithus were rather rare, while the unfavorable chemical regimes in the Northern Prov- boreal species and species of the genera Reticulofenes- ince and the subsequent secondary dissolution of cal- tra, Cyclicargolithus, Coccolithus, and some other spe- careous foraminiferal tests represent eliminating fac- cies tolerant with respect to temperature predominated. tors that further increased the differences between these The assemblages from stratotype Rupelian sections Oligocene assemblages. (Berg Sand, clay beds with Nucula, Boom Clay, Herm- dorf and Framnvald septarian clay beds, and the North- ern Germany cores Schach Profile Rossenray I and Nannoplankton Hoerstsen) include 27 species, 13 of which are in com- mon with the Latdorfian and Tongrian assemblages. The climatic and paleogeographic changes in the A distinctive character is the wide development of the Terminal Eocene—at the beginning of the Early Oli- warm-water Sphenolithus species, S. predistentus and gocene—resulted in a marked change in the coccolitho- S. distentus, and the species Braarudosphaera big- phorid assemblages. In the southern basins, where the elowi, which forms large accumulations at shallow carbonate sedimentation continued, these changes were shelf areas and is rather tolerant with respect to salinity. gradual. The extinction of typical tropical Eocene spe- The same assemblage is typical of the Early Rupelian cies of discoasters started in the Late Eocene, when basins of southern Germany (Rhine Graben, Mainz their subtropical assemblages appeared. Representa- Basin, and Sessishe senke). In the Late Rupelian basins tives of the genus Chiasmolithus (except for Ch. oama- of northern Germany, the species Cyclicargolithus abi- ruensis) almost entirely became extinct. The species sectus and Helicosphaera recta were abundant (Müller,

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S196 POPOV et al.

1971). Twenty-eight species were reported from the ovich, 1986; Muzylev and Tabachnikova, 1987; Ana- Rupelian of the northwestern basins of Germany (Gar- niashvili and Minashvili, 1990; Nagymarosy and towÐKothe cores, 1986). Species of the genera Reticu- Voronina, 1993). lofenestra, Coccolithus, and Cyclicargolithus were The normal marine nannoplanktonic flora again very numerous. Species of Sphenolithus were absent appeared and dominated the central Paratethys only with an exception of the sole S. moriformis. This from the end of the Early Oligocene (at the base of assemblage indicates the relatively deepwater boreal Zone NP24). In the eastern Paratethys, it also appeared environments. in the Middle Maykopian and was recorded from geo- By the end of the Rupelian, the warm-water and rel- logical sections in the Northern Caucasus and Georgia atively shallow-water assemblage had been replaced by (Andreyeva-Grigorovich, 1980; Ananiashvili and Mina- a deeper water and colder water assemblage. Spheno- shvili, 1990; Minashvili, 1992; Nagymarosy and lithus were entirely absent, Braarudosphaera bigelowi Voronina, 1993). and Cyclicargolithus abisectus were rare, whereas the Thus, the major factors that controlled the distribu- species Helicosphaera bramlettei was abundant. tion and the productivity of nannoflora in the Oligocene Paratethys. The Early Oligocene Paratethyan nan- basins of the Paratethys were the proportion of terrige- nofloral assemblages were closely similar and are char- nous material and the salinity regimes. A partial isola- acterized by cold-water species 15 to 25 in quantity. tion of the Paratethys in the middle part of the Early Among the most typical species were Isthmolithus Oligocene resulted in a decrease in salinity and in the recurvus, Reticulofenestra umbilica, Cyclicargolithus appearance of endemic flora. floridanus, Zyrhablithus biyugatus, Discoaster tani nodifer, Corannulus germanicus, etc. The assemblages Organic-Walled Phytoplankton are closely similar to those from the Early Oligocene basins of Germany. The qualitative and quantitative At the beginning of the Oligocene, the organic- compositions were controlled by the basin depths, pro- walled phytoplankton was half-new compared to the portion of terrigenous material, temperature regimes, Late Eocene assemblage. In the Terminal Eocene, as and by a few other factors. Thus, in relatively shallow- well as at the beginning of the Early Oligocene, in more water shelf areas, the assemblages with Isthmolithus northerly regions, Rottnestia borussica, Areosphaerid- recurvus, Lanternithus minuthus, and Zygrhoblithus ium diktyoplokum, A. arcuatum, Tytthodiscus beloglin- bifugatus (from the base of the Kyzyldzhar Beds in the ensis and the Eocene species of the genera Phthanope- Crimea) formed. The deeper-water areas were charac- ridinium, Spiniferites, Hystrichokolpoma, Deflandrea, terized by abundant Cyclicargolithus floridanus (acme- Thalassiphora, Wetzeliella, etc., disappeared. New zone at the base of the Lower Menilithic Formation in taxa, Phthanoperidinium amoenum, Areosphaeridium the Carpathians). pectiniforme, Membranophoridium aspinatum, etc., A marked deterioration of the qualitative composi- appeared and were widespread in the Early Oligocene. tion of nannoplankton assemblages and an acme of Northern Mediterranean. Over 100 species of endemic floras is observed in the Oligocene basins of alga Dinophyta were found in the Oligocene of the the entire Paratethys from the upper part of Zone NP22. Mediterranean Basin. The most typical genus, Chirop- The species Reticulofenestra ornata and Transversopo- teridium, was almost globally distributed. The species ntis fibula were of rock-forming importance, whereas of the genera Spiniferites, Operculodinium, Systemato- Braarudosphaera bigelowi, Orthozugys aureus, and a phora, and Deflandrea increased in quantity. In the few other species tolerant with respect to salinity were Early Oligocene, species of the genera Homotryblium present in small quantities (Nagymarosy, 1983; and Wetzeliella, as well as the species Hystrichokol- Andreyeva-Grigorovich and Gruzman, 1994). The nor- poma pussillum, Areosphaeridium pectiniforme, Chi- mal marine species, including the zonal species Sphe- ropteridium lobospinosum, and Corrudinium incom- nolithus distentus (NP23), were extremely rare (Báldi- positum were very abundant. Beke, 1977; Nagymarosy and Báldi-Beke, 1988; A high biodiversity (over 100 species) was still Andreyeva-Grigorovich, 1994). observed in Armenian assemblages tolerant with This level with endemic nannoplankton is traced respect to high salinity that was associated with hydro- throughout the Paratethys, from the Bavarian Piedmont logic and climatic factors. Within the YerevanÐOrdubad of the Alps and basins of the central Paratethys (base of Depression, the organic-walled phytoplanktonic assem- Tard Clay after Nagymarosy, 1983; upper part of Brebi blage mainly included stenohaline taxa, with many of Marl, Romania, after Rusu et al., 1993; Lower them (especially chorate taxa) being descended from Menilithic Formation of the Carpathians) to the Aral the Eocene. Sea (Nagymarosy and Voronina, 1993). In the eastern North Sea Basin. The dinocyst assemblage Paratethys, the acme of the endemic flora was assigned included 75Ð80 species. A mixture of various ecologi- to the Early Solenovian (Polbian and Molochnaya hori- cal groups was typical similarly to that in the Late zons), but the changes had started earlier, in the Pshek- Eocene, but the qualitative and quantitative composi- hian (Veselov and Lyulieva, 1980; Andreyeva-Grigor- tions markedly changed. The assemblage was domi-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S197 nated by the neritic, normal marine genus Impleto- salinity are better represented than the corresponding sphaeridium, especially in the Late Rupelian. The rela- assemblages from its northern areas (Voronezh Antec- tively deepwater and cold-water species Cribropteridi- lise and Ustyurt). In the peripheral eastern area of the nium spp. was abundant (up to 35% in the upper part of basin, in conditions of periodical brackishing, the eury- Zone NP23), as well as the group Spiniferites/Achomo- haline taxa developed that today inhabit estuaries and sphaera. The groups of species Glaphyrocysta, Areo- lagoons. In the peripheral northeastern area of the ligera, Cordosphaeridium, Homotryblium, as well as basin, as well as within Ergeni Region, the composition Thalassiphora spp. had almost equal proportion. The of the dinocyst assemblages was abnormal and much quantity of representatives of the latter genus markedly closer associated with changes in facial environments, increased in the Early Rupelian Basin and in the Termi- probably because of continuous inflow of fresh water nal Rupelian (in the vicinity of the RupelianÐChattian from the land. This allows the definition of a sequence boundary). The deflandrians and Wetzeliella spp. had of dinofacies here with the alternating predominance of small proportions. Wetzeliella, Deflandrea, Hystrichokolpoma, Phthano- The algal composition indicates an increase in the peridinium, and of the group CyclonepheliumÐ basin depths, especially in the Middle Rupelian, and a Adnatosphaeridium or Micrhysridium. In the entire marked increase in the proportion of cold-water taxa eastern peripheral area of the Paratethys, especially in and cosmopolites with respect to temperature. regressive phases, green algae and acritarches mark- Carpathian Basin. In the Flysch Carpathian Basin, edly increased in importance. Their quantitative distri- the proportion of alga Prasinophyta increased, which bution corresponds well to the sedimentation cycles. was also typical of the terminal Eocene. However, their The diversity and the number of taxa in the assemblage acme was observed at the beginning of the Early Oli- increased during the sea transgression in the gocene. The West Carpathian assemblages were closely Ascheairyk Epoch. Species of Deflandrea, Wetzeliella, similar to those from northern Germany. In the terminal Phthanoperidinium, and the group GlaphyrocystaÐ Rupelian, the acme of the species Wetzeliella symmet- CyclonepheliumÐAdnatosphaeridium were still the rica, W. spp., and Thalassiphora pelagica was leading in combination with Areosphaeridium pectini- observed, as well as the acme of Rhombodinium draco forme, Membranophoridium aspinatum, species of in the Eastern Carpathians. In the southern shelf (Tran- Lejeunecysta, Hystrichokolpoma, Spineferites, Dapsil- sylvanian Basin), the acme of the cosmopolitan species idinium, Dinopterigium, etc. Other Paratethyan assem- Deflandrea phosphoritica is reported. blages (from the Mangyshlak, Northern Ciscaucasia, southern area of the East European Platform, and the Eastern Paratethys. At the beginning of the Oli- gocene, the composition of the organic-walled phy- Carpathian Region) are characterized by the closely toplanktonic assemblage was usually impoverished similar taxa but the higher diversity. because of a general decrease in the subsurface water The compositions of the dinocyst assemblages temperature (possibly by over 5Ð6°C) and a slight within the single Zone Wetzeliella gochtii markedly dif- decrease in salinity, especially in the peripheral areas of fer due to the basin brackishing in the Solenovian the basin. In shallow-water sediments of the peripheral Epoch; the more ancient assemblage is characterized areas (Voronezh Anteclise and Ustyurt), numerous fos- by the presence of mainly stenohaline taxa, whereas the sil organisms redeposited from underlying Eocene beds more recent assemblage includes brackish-water taxa. were found, which disguise the Oligocene age of the The latter assemblage is almost monospecific, includes assemblage. two to three species, which make up 95% of the total The taxonomic diversity of the organic-walled phy- quantity of specimens. The zonal species Wetzeliella toplankton in the assemblages tolerant with respect to gochtii is extremely rare. Thin-walled Hystrichokol- high salinity reached 60Ð80 species, which is less than poma (up to 7 morphotypes) and species of Batiacas- that in the Late Oligocene (120 species, or more). phaera (B. sphaerica, B. baculata, and B. micropapil- Anoxic environments that occasionally formed in the lata) are common. The accompanying taxa were Gerdi- basin markedly affected the composition of the Mayko- ocysta, Parallecaniella, Selenopemphix, Palaeocysto- pian assemblages and possibly the biota of the euphotic dinium golzowense, Horologinella, Leberidocysta, and zone. These epochs were characterized by lower biodi- Chiropteridium. A few assemblages, for instance the versity, less quantity of taxa in the assemblage and Kuban Assemblage, were dominated by cysts of the facial changes (accumulation of black-colored noncar- group CyclonepheliumÐGlaphyrocystaÐAdnatosphaerid- bonate clay beds with concretions and dispersed sul- ium. Generally, the composition of dinocysts from the fides). The biodiversity also markedly decreased in Solenovian Horizon (sensu lato) indicates the periodic brackish-water facies (up to 40Ð60 species) with a pre- fluctuations of the subsurface water salinity in the basin dominance of 2Ð3 species (up to 90Ð95% of the assem- and the absence of entire separation from the open blage). ocean, because the polyhaline taxa, although in small Climatic and local factors also markedly affected quantities, were constantly found in mixture with the the biodiversity. The assemblages of open areas of the euryhaline species and the species tolerant with respect Scythian Basin (Northern Ciscaucasia) with high water to low salinity.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S198 POPOV et al.

In the EarlyÐLate Oligocene boundary beds, the members of the gadiform genus Palaeogadus, distinc- composition and biodiversity of the organic-walled tive for their broad dispersal and ecological diversity,1 phytoplankton markedly increased again. However, and osmerids. The first appearance of boreal elements this part of the Maykopian Formation should be proba- and decline of tropical and subtropical forms are bly assigned to the Upper Oligocene, because the spe- recorded in a number of the earliest Early Oligocene cies of Chiropteridium and Rhombodinium draco are faunas of the Paratethys known from the Belgian and very numerous. Caucasian basins. In general, the faunal events of the Early Oligocene Plankton-based Biogeographic Zonation are most fully documented in the Caucasus and Tran- scaucasia (Bogachev, 1933, 1938, 1964; Danil’chenko, In the Early Oligocene, the distribution and the 1950, 1960; Menner and Ryabinin, 1949; Rozhdestven- diversity of planktonic foraminifers, dinoflagellates, skii, 1950; Aslanova, 1960; Novitskaya, 1961; Fedotov, and nannoflora of the Mediterranean–Mesopotamian 1976; Switchenska, 1979; Danil’chenko et al., 1980; and Paratethyan basins also depended upon the latitudi- Fedotov and Bannikov, 1984; Bannikov, 1985, 1986, nal climatic zonation and water circulation. Two 1988, 1990, 1991, 1993; Bannikov and Danil’chenko, regions are still distinguished: the Ancient Mediterra- 1985; Dzhafarova, 1988; Bannikov and Tyler, 1992; nean Region and the North European Region. The first Tyler and Bannikov, 1994; Sytchevskaya, 1998; region is characterized by a presence of a full set of Prokofiev, 2002), in the Lower Menilite Formation of zonal species of planktonic foraminifers and nanno- the Carpathians (Kalabis, 1948, 1957, 1981; Jerz- plankton. Its distinctive character compared to the man′ ska, 1960, 1968, 1974; Gorbatch, 1961a, 1961b; North European Region is the wide development of Jerzman′ ska and Kotlarczyk, 1968, 1976, 1978, 1979, warm-water discoasters as well as Sphenolithus. This 1981, 1983; Kotlarczyk and Jerzman′ ska, 1976, 1980, region may be considered as the southern subtropical 1988; Gregorová, 1988), and in the Brown Marles and

subzone of the tropical zone.

^ Disodylic Schists of^ Romania (Cosmovici, 1889, 1913, In the North European Region, the planktonic fora- 1922, 1966; Pauca , 1934; Cosmovici and Pauca , 1943; miniferal assemblages and warm-water discoaster Jonet, 1958; Ciobanu, 1970, 1975, 1977, 1978). Of fur- assemblages were very impoverished or even absent, ther importance are also the data on the Early Oli- the representatives of Sphenolithus were rare, whereas gocene fish faunas of Iran (Arambourg, 1967), Belgium the boreal species and the species of the genera Reticu- (Leriche, 1910), and the basins of Balkans, and Western lofenestra, Cyclicargolithus, and Coccolithus, as well and Central Europe (Wettstein, 1887; Weiler, 1922, as some other taxa tolerant with respect to temperature 1928, 1931, 1932, 1938, 1953, 1963, 1966; Böhm, predominated. Thus, this region may be considered as 1941; Frohlicher and Weiler, 1952; And– elkovic′ , 1970, the boreal by its temperature regime. The border 1989; Pharisat, 1991; Micklich and Parin, 1996; Phari- between these regions went along uplands of the Alps, sat and Micklich, 1998). Dinarides, and Rhodope (Fig. 2) and then along the Pontides, Lesser Caucasus, and the Elburs, that sepa- A general analysis of the history of the Early Oli- rated the Tethyan Region (including the Armenian gocene ichthyofauna of the Paratethys is complicated Gulf) from the North European basins and the Parat- for many reasons, which primarily include the contro- ethys. The subsurface water circulation in Paratethys versial dating of some local assemblages; the low reso- was probably closed and affected mainly by North lution of their discrimination (which results in that the Atlantic waters (Atlas…, 1997; Fig. 2). Only southern one and the same assemblage may be reported to con- areas of the Carpathian Basin were slightly affected by tain the taxa of different age) and, finally, the doubtful the Tethys. taxonomic attribution of a large number of forms described from the Early Oligocene. Central Iranian Basin. A rich Rupelian ichthyo- Marine Ichthyofauna fauna of Iran is known to include 31 genera (Aram- With the onset of the Oligocene, the run of develop- bourg, 1967), showing predominance of tropical and ment of the marine ichthyofauna in the Paratethys subtropical forms. The abundance of mesopelagic ele- underwent fundamental changes. In Europe, a number ments, such as Praewoodsia mesogeae Aramb., Scope- of more or less isolated basins of various depth and loides glarisianus (Ag.), Maurolicus morgani Aramb., salinity arose at that time. The composition of the fish assemblages that lived here during the Early Oligocene 1 According to Danil’chenko (1950), the Palaeogadus species of the subgenus Rupelianus (P. intergerinus Dan. and P. aram- reflected the changes in the degree of isolation of par- bourgi) found together with the deepwater fishes obviously ticular basins, the dynamics of their shallowing or dwelled at the great depths, whereas other members of the same deepening, as well as climatic changes. Most local genus could have populated more shallow zones, which is evi-

communities were dominated by both the autochtho- denced by their finds in association with clupeids, syngnathi- forms, and scombrids. Finally, the members of the subgenus^ Loti- nous genera inherited from the Middle to Late Eocene morpha (P. simionescui Dan. and P. athanasiui Pauca ), known to and the groups of boreal origin, which came at that time be accompanied by Ammodites and Capros, most likely populated from the Atlantic. The bulk of the latter was formed by the lower part of the sublittoral and were the bottom-dwellers).

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S199

50° Kazakhstanian Land Russo-Scandia

N O R T 40° H E U R O P E A N R E G I O N

30° Anatolia Iran-Afghanistan Land ANCIE NT ME DITER RANE AN RE GION Subtropical Africa India 20° Arabia

Legend: 5 7 1 6 8 2 4 3 9 Tropical

Fig. 2. Superficial circulation of water in the Rupelian basins (after Paleogeographic..., 1997, modified) and the pelagial latitudinal zones. Explanations: (1) Temperate and Boreal zones with impoverished planktonic associations and the presence of specialized cold-water genera and species: (2) extremely impoverished assemblages of planktonic foraminifers with the predominance of small globigerinids (Globigerina officinalis) and globorotaliids; (3) nannoplanktonic assemblages with the absence of discoasters and the predominance of tolerant species of the genera Reticulofenestra, Cyclicargolithus, and Coccolithus; (4) boreal dinocyst assem- blages, including Wetzelliella simmetrica, Deflandrea phosphoritica, Enneadocysta pectiniforme, etc.; (5) Subtropical and Trop- ical zones with complete zonal warm-water planktonic assemblages; (6) nannoplanktonic assemblages with abundant Discoaster, Sphenolithus, and Helicosphaera; (7) dinocyst assemblages of the southern type with the predominance of chorate cysts of the gen- era Spiniferites, Cordosphaeridium, etc.; (8) complete assemblages of planktonic foraminifers with Subbotina, Globigerina, and Paragloborotalia; and (9) Transitional Zone.

Astronesthes simus Aramb., Proserrivomer mecquen- the Caucasian Basin (Danil’chenko, 1960; Danil’chenko emi (Priem.), Bregmaceros filamentosus (Priem.), and et al., 1980; Bannikov, 1985, 1990; Sytchevskaya, 1998). Myroconger roustami Aramb., gives evidence of rich These are documented in ascending order by the finds mesopelagic life and suggests considerable depth of the from the Pshekhian Horizon (which contain the sub- basin. The genera survived from the Eocene include Planorbella, Planorbella and Aeoliscus, or “Amphisile” Astronesthes, Urosphen, Bregmaceros, Sphyraena, Pris- Beds), Polbian Horizon, and the basal part of the tigenys, Palaeorhynchus, Scomberomorus, etc. The Ira- Morozkina Balka Horizon (and its equivalents), with nian Ichthyofauna contained a number of genera and the total range extending from the NP21 Zone to the species shared with the coeval fish communities of the lower part of NP24 in terms of the nannoplankton zona- Caucasian, Carpathian, and Alpine basins. The indica- tion. In this succession, four to five fish assemblages of tions of exchange with these basins are traceable for different ages are detected (Sytchevskaya, 1998). both the Early Rupelian (as evidenced by the presence The earliest assemblage comes from the sub-Plan- of Bregmaceros filamentosus (Priem.), Scopeloides orbella and Planorbella beds of the Pshekhian Horizon glarisianus (Ag.), Palaeorhynchus, Isurichthys, Scomb- (Ericsonia subdisticha NP21 Zone) and includes in eromorus, etc.) and the Late Rupelian, as seen, for turn a number of successive assemblages whose differ- instance, from the occurrence of Thyrsitoides zaratous- ences reflect the change in the characteristics of the trae (Aramb.) known elsewhere from the ichthyozones basin and the climatic conditions. The latest assem- IPM3ÐIPM4 of the Carpathian basin (Kotlarczyk and ′ blage is ecologically closely linked to the next follow- Jerzman ska, 1988). ing assemblage of the Aeoliscus Beds (see below). Greater Caucasian Basin. In the Paratethys, as a The assemblage of the sub-Planorbella level was whole, the main stages of development of the Early Oli- impoverished and included Lyrolepis sp. (Elopidae) gocene ichthyofauna are most clearly distinguished in and Thunnus sp., (Thunnidae), i.e., the relicts of some

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S200 POPOV et al.

Late Eocene dominant genera (Sytchevskaya’s data). considerable warming. Here is recorded the rise and The assemblage is traceable in Ciscaucasia and Tran- coming to predominance of tropical and subtropical scaucasia (Dzhafarova and Aslanova, 1974) and bears forms, such as trachychthyids [Gephyroberyx robustus resemblance to the basal fauna of the Early Rupelian (Bog.)], turkmenids (Analectis pala Dan.), digoriids (Upper Tongrian) of Belgium. The latter also shows the (Digoria ambigua Dan.), syngnathids [Doryrhamphus decrease in diversity accounted for by extinction of pre- squalidus (Dan.)], serranids (Serranus comparabilis ceding (Early Tongrian) tropical and subtropical forms, Dan.), priacanthids (Pristigenys spinosus Dan.), car- and the survival of only a limited part of the Late angids (Caranx daniltshenkoi Ban.), acanthurids Eocene components (Leriche, 1910). (Caprovesposus parvus Dan.), stromateids (Pinichthys The subsequent stage in the history of the fish fauna pulcher Ban.), nomeids (Rybapina caucasica Ban. and of the Caucasian Basin refers to the Planorbella Beds. Psenicubiceps alatus Dan.), scombrids (Scombrosarda It is represented by two essentially different assem- cernegurae (Ciob.) and Sarda rara Ban.), thunnids blages. The earlier of them was dominated by the clu- (Thunnus sp.), and balistids (Oligobalistes robustus peids of the subfamilies Clupeinae (Sardinella rata Dan.), along with the development of clupeids (appear- Dan.) and Alosinae (Pomolobus curtus Dan.), the ance of Paretrumeus avitus Dan.), congrid eels (Paveli- gadiforms of the genus Palaeogadus showing a great chthys daniltschenkoi Ban. et Fed.), and paleorhynchids species diversity [P. crassus (Ag.), P. intergerinus Dan., (Danil’chenko, 1960; Fedotov, 1976; Danil’chenko P. rarus Novit., P. eximius Novit., and P. germanus et al., 1980; Fedotov and Bannikov, 1984; Bannikov Fed.],2 the Trichiuridae (Anenchelum angustum (Dan.) and Tyler, 1992; Tyler and Bannikov, 1994; Sytchev- and A. glarisianum Blainv.), and the Zeidae (Zenopsis skaya, 1998; Prokofiev, 2001, 2002), while the clarus Dan.). Detected is also the presence of osmerids gadiforms decrease in their role. By the predominant (unpublished author’s data), and the development of adaptation patterns and composition, the Upper Plan- argentinids (Proargentina inclinata Dan. and Glossan- orbella Beds Subassemblage is close to the immedi- odon sp.), paralepidids [Holosteus mariae (Menner)], ately succeeding Aeoliscus (= “Amphisile”) Beds Assem- brotulids (Protobrotula sobijevi Dan.), and euza- blage, which allows one to consider the both communi- phlegids (Palimphyes chadumicus Dan.)3 (Danil’chenko, ties as a single faunal stage. This, in terms of the 1960; Fedotov, 1976; Sytchevskaya, 1998; Prokofiev, development of the marine fish fauna, the biochrono- 2002). logical demarcation between the Early and Late Pshek- hian Time evidently lies within the Planorbella Beds The presence of the four listed families testifies to time span rather than between it and the Aeoliscus inter- the abundant mesopelagic life and the considerable val (as was generally accepted). depth of the basin. The fish assemblages of its neritic The next succeeding assemblage comes from the zone were dominated by trichiurids and zeids, while the Aeoliscus Beds and corresponds to the Helicop- gadids populated both the mesopelagic and sublittoral ontosphaera reticulata NP22 Zone. It was a subtropical zones. Some characters, primarily the occurrence of community dominated by the nearshore, nerito-pelagic, osmerids and the abundance of gadids, point to a more and mesopelagic fishes. A wide distribution of the lat- moderate temperature of the basin in comparison with ter, especially of the distribution of photophore-bearing both the Late Eocene and the subsequent Late Pshek- fish, shows that, at that time, the bottom of the Parat- hian Time. ethys in the Caucasian area descended to a depth of By the end of the Early Pshekhian (marked by upper about 1000 m. In the community discussed, a great role levels of the Planorbella Beds), the composition of the was played by the species inherited from the Late Plan- fish community shows marked changes, which indicate orbella Beds Association. In contrast to their scarcity there, in the Aeoliscus Beds Assemblage, some of them 2 The occurrence in this assemblage of P. latebrosus Dan. grew in numbers, which occurred in parallel to the (Fedotov, 1976) seems doubtful. This species is known only by increase in the generic and species diversity of tropical the type specimen from the lower part of the Miatlin Horizon of the Gumista River (Danil’chenko, 1960). and subtropical groups. At the same time, the role of the 3 The fish assemblage of the Planorbella Beds was traditionally moderately thermophilic groups was reduced. In partic- considered as a single faunal unit (Danil’chenko, 1960; Fedotov, ular, the Palaeogadus species were much reduced in 1976; Bannikov, 1985, 1990; Bannikov and Parin, 1997; Sytchev- abundance and diversity while the osmerids vanished. skaya, 1998). However, with the increase in knowledge of its composition, it became clear that the upper part of the beds con- In all, at the Late Pshekhian Time, the Great Cauca- tains a considerable number of the deepwater and warm-requiring sian Basin was a subtropical sea with normal salinity, forms, which is unusual for the typical community of these beds. open for exchange with the marine faunas of the Medi- This justifies distinguishing the older and younger Planorbella Beds fish subassemblages, the latter bearing resemblance with terranean, Carpathian, and West European basins. It the Later Aeoliscus Beds Assemblage with respect to the ecology contained a fish community displaying a wide diversity of the predominant components. It should be noted that even of syngnathiforms, including the Centriscidae [Aeolis- Danil’chenko (1960) detected scarce finds of some tropical and cus heinrichi (Heckel)], Urosphenidae (Oligosphenop- deepwater species, such as Archaeolicus (Palaeotroctes) strictus Dan. (Sternoptychidae) and Doryrhamphus (Acanthognathus) sis gracilis Par.), Fistulariidae (Fistularia contermina squalidus (Dan.) (Syngnathidae), in the upper part of the Planor- Dan.), and the Syngnathidae (Syngnathus incertus bella Beds. Dan., Doryrhamphus squalidus (Dan.), D. incolumnis

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S201

Par., and Maroubrichthys serratus Par.). These taxa autochtonous forms, along with a contribution from the formed a rich fauna of the nearshore zone along with more boreal taxa of Atlantic origin, such as the the littoral and sublittoral dwellers, such as trachinids gadiforms and osmerids. Only later, with the warming (Trachinus rusticus Ban.), leiognathids [Leiognathus of both the superficial and deepwater masses at the end altapinnus (Weiler)], turkmenids (Analectis pala Dan.), of the Early Pshekhian, the basin was populated by the sparids (Sparus priscus Swich.), champsodontids tropical and subtropical ichthyofauna, which most [Champsodon (= Myersiscus, = Pseudoscopelus) likely came from the southern refuges of the Tethys. grossheimi (Dan.)], platacids (Paraplatex absyrtus This run of events obviously accounts for the enormous Bog.), ammodytids (Ammodytes antipai Paucá), balis- amount of those genera in the Pshekhian Fish Fauna tids (Oligobalistes robustus Dan.), tetraodontids that were inherited from the Tethyan Eocene Ichthyo- (Archaeotetraodon winterbottomi Tyler et Ban.), fauna. In the course of development of the Pshekhian lophotids (Oligolophotes fragosus Ban.), and caproids Basin, it shows the change in the dominants of the [Capros radobojanus (Kramb.)]. The stromateids (Pin- mesopelagic communities. In the Early Pshekhian ichthys pulcher Ban.) and nomeids (Rybapina cauca- Time, this part was played by euzaphlegids, paralepid- sica Ban. and Psenicubiceps alatus Dan.) continued to ids, brotulids, trachychtyids, and mesopelagic gadids of develop. the genus Palaeogadus, while the Late Pshekhian The pelagic zone was abundantly populated by (Aeoliscus Beds) Assemblage was dominated by the members of the clupeid subfamilies Clupeinae (Sar- photophore-bearing forms: phosicththyids, gonostoma- dinella sardinites Dan.) and Alosinae (Pomolobus faci- tids, sternoptychids, myctophids, and bregmacerotids. lis Dan.), along with scombrids (Scombrosarda cerne- The development of the rich mesopelagic fish fauna gurae (Ciob.) and Sarda rara Ban.) and palaeo- enables one to assume that both the Early and Late rhynchids [Palaeorhynchus zitteli (Kramb.) and Pshekhian basins could attain the depths about 1000 m. Homorhynchus colei (Ag.)]. For the mesopelagic zone A significant part of the deepwater genera of the Aeolis- detectable is the further development of euzaphlegids cus Beds Fish Fauna of the Caucasus was also repre- (Palimphyes chadumicus Dan. and P. longirostratus sented in the Carpathian Basin. Dan.) and a great role played by the photophore-bear- The next following stage of the history of the ing forms: phosicththyids [Eovinciguerria (Vinciguer- Ciscaucasian Marine Ichthyofauna is poorly known. ria) obscura Dan. and E. talgiensis Dan.], gonostoma- It is represented by the Polbian Horizon Assemblage tids [Scopeloides glarisianus (Ag.)], sternoptychids (NP23 Zone) showing a severe reduction in systematic (Argyropelecus cosmovicii Cosm. et Pauca),4 and myc- diversity which indicates the shallowing and brackish- tophids (Eomyctophum menneri Dan., E. koraense ing of the basin. The only taxa recorded from here are Dan., and E. limicola Dan.). The bregmacerotids (Breg- the clupeid Sardinella sp. and the scarce nearshore syn- maceros filamentosus), first recorded in the Middle gnathid Syngnathus sp. (Danil’chenko, 1960). Since Eocene of Georgia and known to predominate in the both these forms were inherited from the Aeoliscus Middle and Late Eocene fish assemblages of the East- Beds Assemblage, they can be regarded as its eurybion- ern Paratethys, appeared again (Bogachev, 1933, 1938, tic component, which survived after the drop of salinity. 1964; Menner and Ryabinin, 1949; Rozhdestvenskii, The youngest Early Oligocene assemblage is that 1950; Danil’chenko, 1960; Fedotov, 1976; Switchen- from the basal level of the Morozkina Balka Horizon ska, 1979; Danil’chenko et al., 1980; Bannikov, 1985, and its equivalents (lower part of the NP24 Zone). 1986, 1988, 1993; Dzhafarova, 1988; Sytchevskaya, It corresponds to the end of the Rupelian and indicates 1998; Prokofiev, 2001, 2002, in press). In a whole, the reestablishing of normal salinity in the basin. The characteristic of the Pshekhian Ichthyofauna is a great sections of this stratigraphic level exposed in western role played by the members of the autochtonous Ciscaucasia (Belaya River) yield the following taxa: Tethyan genera retained from the MiddleÐLate Eocene: the Clupeidae [Sardinella sardinites (Heckel)], Argen- Lyrolepis, Sardinella, Glossanodon, Archaeolicus, tinidae (Glossanodon sp.), Syngnathidae (Hypposyng- Eovinciguerria, Eomyctophum, Bregmaceros, Analec- nathus), Merlucciidae (Palaeogadus simionescui Dan. tis, Pristigenys, Caprovesposus, Palimphyes, Scombro- and P. latebrosus Dan.), Serranidae (Serranus sp. nov.), sarda, Sarda, Rybapina, Anenchelum, Thunnus, Palae- Percichthyidae (Repropca sp. nov.), Priacanthidae (Pri- orhynchus, Homorhynchus, and Oligobalistes. acanthus sp. nov.), Caproidae [Capros radobojanus (Kramb.)], and Scombridae (Sarda remota Dan.). This To summarize the pattern of faunal events of the nerito-pelagic community existed in the water space Pshekhian Time, the following points are to be noted. with considerable depths under moderately warm cli- The temperature conditions that existed in the Cauca- matic conditions (Sytchevskaya, Bannikov, and Prokofiev, sian Basin at the beginning of the Early Oligocene in press). obviously restricted the dispersal of tropical and sub- tropical forms, so that the fish community of the basin Carpathian Basin. The Early Oligocene marine was built up in many respects at the cost of survived fish fauna of the Carpathians is known from the locali- ties of Romania, Czech, Poland, and Ukrainian Cis- 4 The presence here of Argyropelecus cosmovicii is in question Carpathians (Cosmovici, 1889, 1913, 1922, 1966; (Jerzman′ ska and Kotlarczyk, 1983). Paucá, 1934; Cosmovici and Paucá, 1943; Rozh-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S202 POPOV et al. destvenskii, 1950; Jonet, 1958; Jerzman′ ska, 1960, lution is represented by the ichthyozones IPM5 and 1968, 1974; Gorbatch, 1961a, 1961b; Ciobanu, 1970, IPM6 (Late OligoceneÐEarly Miocene). 1977, 1978). The composition of this fauna and the Correlation of these events revealed in the Car- stratigraphic position of many its constituent elements 5 pathian Rupelian with the evolution of the Early Oli- need revision. A correlation of the Carpathian assem- gocene ichthyofauna of the Caucasus and Ciscaucasia blages is also hampered by controversial assessments of ′ 6 remains somewhat unclear. According to Jerzman ska data on the accompanying nannoplankton microfossils. and Kotlarczyk (1980, 1983), the Carpathian Ichthyo- An important piece of data on the Rupelian history zone IPM1 (the lower bathypelagic assemblage) can of the regional marine ichthyofauna is provided by the correspond to the both of main faunal levels of the MenilithicÐKrosno Series of the Polish Carpathians. In Pshekhian Horizon in Caucasia, i.e., the Planorbella this unit, there were distinguished the six ichthyozones, and Aeoliscus Beds assemblages. Judging from current which span a sequence of stratigraphic levels from the datings of these faunas in terms of the nannoplankton Oligocene to Early Miocene, (Jerzman′ ska and Kotlarc- zonation (NP21ÐNP22 for Ciscaucasia, and NP21 to zyk, 1980, 1981, 1983; Kotlarczyk and Jerzman′ ska, the base of NP23 for Polish Carpathians), such a corre- 1988). The lowermost of them, Ichthyozone IPM1, or lation looks plausible. However, it is not immediately the lower bathypelagic assemblage is characterized by corroborated by the comparison of these regional faunas. the following index taxa: Scopeloides glarisianus (Ag.) In fact, the above viewpoint would imply that the (Gonostomatidae), Eomyctophum limicola Dan. (Myc- time range of the lower bathypelagic fish assemblage of tophidae), Aeoliscus heinrichi (Heckel), A. teleajensis, the Carpathian Basin should correspond to three differ- and A. longispinus (Centriscidae). According to the lat- ent events in the history of the Caucasian Basin (see est interpretation by Kotlarczyk and Jerzman′ ska above): (1) the existence of the impoverished relict (1988), this faunal unit is correlated with the interval post-Eocene community (just before the “Planorbella” ranging from the upper part of Zone NP21 to the lower Time); (2) the formation of community with admixture half of NP23. of moderately thermophilic forms and the boreal forms of Atlantic origin (Early “Planorbella” Time); only In the younger Ichthyozone IPM2, attributed to the after that (3), the appearance of subtropical communi- upper part of NP23, the typical forms are the gadid spe- ties with the mesopelagic component dominated by cies Paleogadus simonescui and the argentinoid Glos- phosicththyids, gonostomatids, myctophids, and ster- sanodon musceli. The assemblage is estimated as the noptychids (Late Pshekhian Time, spanning the Late neritic-sublittoral community devoid of deepwater “Planorbella” and “Aeoliscus” intervals). components, which is believed to evidence the shallow- ing of the basin in comparison to the preceding epoch. In this sequence, the main faunal replacement, first The next following two ichthyozones, IPM3 [with the noted by Danil’chenko (1960), is expressed as a differ- ence between the Early Pshekhian Fish Assemblage, index species Bregmaceros filamentosus (Priem), Kot- yielded by the Planorbella Beds (Early “Planorbella” larczykia bathybia Jerzm., and Polyipnus brevis Gorb.] Subassemblage in our interpretation), and the mesope- Polyipnus sobniovensis and IPM4 (with Jerzm.), tenta- lagic assemblage of the Aeoliscus Beds. It is only the tively equated to the lower and upper parts of the NP24 ′ latter that is directly comparable with Ichthyozone IPM1 Zone respectively (Kotlarczyk and Jerzman ska, 1988), of the Carpathians. Correspondingly, all the parallels are designated as the upper bathypelagic assemblage ′ between this zone and the Pshekhian Ichthyofauna (Jerzman ska and Kotlarczyk, 1983), whose further evo- stressed by Kotlarczyk and Jerzman′ ska (1988) to 5 This refers, in particular, to the richest assemblage of Peatra ground their concept of correlation, such as the pres- Neamt in Romania which is reported to include (along with Early ence of Scopeloides glarisianus (Ag.), Eomyctophum Oligocene forms) some taxa common for the Miocene. For exam- limicola Dan., Aeoliscus heinrichi (Heck.), etc., refer ple, the set of the local deepwater taxa is said to include Vinciq- only to the “Aeoliscus” Assemblage. uerria merclini Dan., which is known from the Tarkhan deposits of the Caucasus and Crimea (Ciobanu, 1977). To account for the difference between the Early and 6 For instance, this is the case for the ichthyozones distinguished in ′ Late Pshekhian ichthyofaunas, the above-cited authors the Polish Carpathians by Kotlarczyk and Jerzman ska (1980, referred to the strictly ecological factors, on presump- 1988; Jerzman′ ska and Kotlarczyk, 1981). At different times, they equated the lower ichthyozones IPM1 and IPM2 with either the tion that in the “Planorbella” Time the Caucasian nannoplankton NP22 Zone, based on the data by Martini and Basin was still too shallow to enable the wide distribu- Lebenzom (1971), or the end of NP21 plus NP22, or, finally, con- tion of the deepwater forms typical of the “lower bathy- cluded that most of the IPM1 Ichthyozone corresponds to the end pelagic assemblage” (Jerzman′ ska and Kotlarczyk, of NP21 plus NP22, while its upper part belongs to the lower half of NP23 (cf. Kotlarczyk, Jerzman′ ska, 1988). In addition, it 1983). However, this hypothesis does not explain the remains unclear which particular Ichthyozone within the contrast between the abundance of the boreal taxa in the sequence established for the Polish Carpathians actually con- Early Pshekhian Assemblage (six species of Palaeoga- forms to the end of the Early Oligocene. Formerly, this age was dus, along with the presence of osmerids) and their implied for Ichthyozone IPM4, but later, this zone was suppos- edly assigned to the upper part of NP24 which should mean that it scarcity in both the Late Pshekhian Fauna and Car- belongs to the Late Oligocene (cf. Jerzman′ ska and Kotlarczyk, pathian Ichthyozone IPM1. The same concerns the con- 1981; Kotlarczyk and Jerzman′ ska, 1988, p. 348). trast between subordinate role of the tropical elements

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S203

in the Early Pshekhian Assemblage and abundance of stratigraphic distribution of fish fossils has not been

^ them (both in diversity and numbers), especially the studied in detail. The fauna is known^ from a number of syngnathiforms and deepwater photophore-bearing localities (Homoric u, Suslanes ti, Peatra Neamt , etc.) taxa, in both the “Aeoliscus” Assemblage and the basal and comprises a rich set of the littoralÐsublittoral, Ichthyozone IPM1 of the Carpathian Oligocene. In two pelagic, neritic, and mesopelagic fishes (Cosmovici, latter communities the above-named groups include a 1889, 1913, 1922, 1966; Paucá, 1934; Cosmovici and number of shared species, such as Aeoliscus heinrichi Paucá, 1943; Jonet, 1950, 1958, 1971; Ciobanu, 1970, (Heckel), Eovinciguerria (Vinciguerria) obscura, 1973, 1975, 1977). In the Early Rupelian (Brown Marls Eomyctophum menneri, E. koraense Dan., and E. limi- of the Peatra Neamt locality), there were abundantly cola Dan. (Danil’chenko, 1960; Jerzman′ ska, 1968; represented the various pelagic clupeids: Sardinella Jerzman′ ska and Kotlarczyk, 1983). rata Dan., S. denticulata Ciob.*,7 Alosa sculptata Weiler, Pellonula gracionescui Ciob.*, and Pomolobus In the light of these facts, it seems very likely that, facilis Dan.; neritic argentinoids [Glossanodon musceli in reality, Carpathian Ichthyozone IPM1 conforms only (Pauca)], trichiurids (Anenchelum glarisianum Bla- to the Aeoliscus Beds Assemblage (NP22 Zone) as was inv.); caproids [Capros radobojanus (Kramb.)*]; pria- formerly assumed by Polish authors (Kotlarczyk and canthids [Pristigenys spinosus (Blainv.)], leiognathids Jerzman′ ska, 1976). The equivalents of the earlier Cau- [Leiognathus altapinnus (Weiler)]; and a number of casian ichthyofaunas, reflecting the invasion of the mesopelagic groups such as brotulids (Propteridium boreal taxa and gradual increase in the role of tropical profundae Ciob.*), merlucciids (Palaeogadus interger- forms, could also occur there, but they have not been inus Dan.), gonostomatids [Scopeloides glarisianus found. This view is in accord with the fact that, in Ich- (Ag.)], sternoptychids [Archaeolicus? (Vinciguerria) thyozone IPM1, the mass finds of the genus Aeoliscus macarovicii (Ciob.)],8 and various myctophids of the fall on the middle part of the zone but are lacking in its genera Myctophum* (M. weileri Ciob., M. praeptero- basal part (Kotlarczyk and Jerzman′ ska, 1976; Jerz- tum Ciob., M. robustus Ciob., and M. antelaternatum man′ ska and Kotlarczyk, 1983, pp. 4Ð5). Ciob.), Lampanictus (L. longaevus Ciob.), and Diaphus The next point to be discussed with respect to the (D. duosensitivus Ciob.) (Ciobanu, 1977). This assem- Early Oligocene fish faunas of the Caucasus and Polish blage was largely formed by inhabitants of a deep warm Carpathians is a comparison of their terminal assem- marine basin with oceanic salinity. It displays certain blages. If one follows the dating of the Carpathian ich- relationships to the Caucasian Planorbella Beds Ich- thyozonal succession (in terms of NP zones) accepted thyofauna by the presence of Sardinella rata Dan., by Kotlarczyk and Jerzman′ ska (1988), the Caucasian Pomolobus, Glossanodon, Palaeogadus intergerinus fish assemblages of the Polbian Horizon and of the Dan., Anenchelum glarisianum Blainv., and Pristigenys lower part of the Morozkina Balka Horizon should be spinosus (Blainv.) but differs by the abundance of deep- equated to the Carpathian icthyozones IPM2 and IPM3, water photophore-bearing taxa (which in the Caucasus and, possibly to a part of IPM4. If this is the case, the first appear in the younger Aeoliscus Beds) and by a episode of shallowing and brackishing distinguished in lower diversity of the gadid genus Paleaogadus, which the Caucasian Basin as the Polbian Horizon can proba- is represented here by a single species, P. intergerinus bly be correlated with the nerito-pelagic assemblage Dan. These features may point to a mixed origin of the IPM2 containing the index species Palaeogadus simio- assemblage, which appears to combine some typical nescui Dan. and Glossanodon musceli (Paucá). Like- mesopelagic components of the “Aeoliscus” commu- wise, the younger community of the basal part of the nity with those of the earlier assemblage from the basal Morozkina Balka Horizon may correspond to a more Oligocene. deepwater Carpathian assemblage of Ichthyozone IPM3 with Bregmaceros filamentosus (Priem.), Kot- The second half of the Rupelian (the Lower Diso- larczykia bathybia Jerzm., and Polyipnus brevis Gorb. dylic Schists) in the Romanian Carpathians is docu- Hence, on the strength of evidence provided by the fish mented by a mixed fish assemblage, which embraces fauna, at the end of the Early Oligocene, the Carpathian the late Early Oligocene communities of different age. Basin was deeper and did not suffer freshening, which Here is recognized the presence of the index taxa of was so markedly expressed in the Caucasian Basin dur- Ichthyozone IPM1 of the Polish Carpathians (Aeoliscus ing the Polbian Time. The data on the Early Oligocene heinrichi, Scopeloides glarisianus, and Eomyctophum fauna of Moravia (Gregorová, 1988), showing the pres- limicola), which are associated with a number of typi- ence there of at least two lower ichthyozones that were cal species of the mesopelagic, pelagic, and nearshore recognized for the Polish Carpathians, corroborate communities known in the Aeoliscus Beds Assemblage these conclusions. 7 The taxa marked with the asterisk need to be revised. In the Romanian Carpathians, the Early Oligocene 8 According to the data by Prokofiev (2002), the attribution of Vin- ichthyofauna is known from the Brown Marls and the ciguerria macarovicii to the Phosichthyidae is doubtful. This form appears to be close to the genus Maurolicus (OligoceneÐ overlying Lower Disodylic Schists, which are assigned Recent) or Archaeolicus (Oligocene) and belongs to the Sternop- together to zones NP21ÐNP23 and the base of NP24 tychidae. V. praeattenuata is possibly a synonym to Eovinciguer- (Melinte, 1995; Rusu et al., 1996). In these units, the ria obscura.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S204 POPOV et al. of the Caucasus: Eovinciguerria (Vinciguerria) obscura predominate; the presence of Chimaera testifies to (Dan.)*, E. talgiensis (Dan.), Eomyctophum koraense some cooling (Leriche, 1910). E. limicola E. menneri Pomolobus Dan., Dan., Dan., The later assemblage of the Boom Shales shows fur- facilis Scombrosarda cernegurae Aeolis- Dan., (Ciob.), ther diversification and, in particular, the appearance of cus heinrichi Syngnathus incertus (Heckel), and Dan. the abyssal forms (Macrurus), which indicates the Balkan Basin. In this basin, the Early Oligocene deepening of the basin. At that time, a great variety of ichthyofauna is known from several localities of Slove- the subtropical genera existed in the pelagic and littoral nia and Serbia (Gorjanovic′ -Kramberger, 1882a; zones, such as Sarda, Cybium, Neocybium, Xiphias, And– elkovic′ , 1989). In Slovenia, the Rupelian contains Sphyraenodus, Scombramphodon, Glyptorhynchus, an impoverished marine fish assemblage, which is Sparus, Eutrichiurides, and Trigla. Their presence pro- recorded in the Sava River basin (Inner Dinarids) and vides evidence of warming. However, distinct from the stratigraphically positioned above the beds with Carpathian and Caucasian basins, the development of Anthracotherium. It includes Clupea alta Steind., the rich deepwater mesopelagic fauna is not observed C. mucronata Kramb., Sardinella (Clupea) sardinites and the photophore-bearing taxa are absent. Likewise, (Heckel), Alosa sagorensis (Steind.) (Clupeidae), Mor- there is no tropical syngnathiforms that constituted one one lata (Kramb.), M. elongata (Kramb.), M. mojsisov- of leading components in the Caucasian and Carpathian icsi (Kramb.), M. stiriaca (Rolle), M. sagorensis communities of the same age. In general, the Belgian (Kramb.) (Moronidae), Anenchelum? (Lepidopus) cau- Rupelian Ichthyofauna differed from other coeval datum (Trichiuridae), Scomberomorus sp. (Scom- European assemblages in showing a more important bridae), Thunnus planovatus (Kramb.) (Thunnidae), role of the boreal and Arctic forms, which undoubtedly Palaeorhynchus riedly Kramb., P. deschmani (Palaeo- reflected a greater influence of the Atlantic Ichthyo- rhynchidae), Serranus validus Kramb., S. stiriacus fauna. On the other hand, the relationships of the Bel- Kramb. (Serranidae), Caranx gracilis Kramb., Lichia gian Fauna with other Rupelian fish communities of alta Kramb. (Carangidae), Sparus hertlei (Kramb.), Europe are proved by the presence in them of the Sparnodus inflatus Kramb. ?Pargus sp. (Sparidae), shared genera of sharks and scombrids. The present Chaetodon hoeferi Kramb. (Chaetodontidae), Zeus data are not enough to substantiate the establishment of robustus (Zeidae), Rhinoptera aff. studeri (Mylio- the Belgian Rupelian Fauna as a separate biogeo- batidae), Gadus? sp. (Gadidae), Saurodon cf. phlebot- graphic unit, but such a possibility is not ruled out with omus (Cope) (Saurocephalidae), and Isurus sp. the increase in knowledge of its composition. ′ (Isuridae) (Gorjanovic -Kramberger, 1882, 1884, 1895; Basins of Central and Western Europe. The ′ 9 And– elkovic , 1989). The assemblage populated rela- Rupelian fish faunas of Central and Western Europe tively shallow warm sea with some zones of reduced need revision. The bulk of pertinent data come from the salinity, as indicated by abundance of moronids (prefer- fish schists of the BavarianÐSwitzerland Molasse, ring the fresh- and brackishwater settings) and by a pos- Rhine Valley, and northeastern France. These beds are sible admixture of cyprinids (Barbus crenatus). Most of a different age and stratigraphic position (Rath, forms were the nearshore and pelagic inhabitants. 1859; Weiler, 1922, 1928, 1931, 1932, 1953, 1963, Some of the genera are also known from the Rupelian 1966; Leriche, 1927, 1927-1928, 1941; Theobald, of Western and Central Europe and (or) the Caucasus 1934; Frohlicher and Weiler, 1952; Hess and Weiler, (Clupea, Alosa, Sardinella, Serranus, Caranx, Morone, 1955; Micklich and Parin, 1996; Pharisat and Micklich, Sparus, Lepidopus, Thunnus, Scomberomorus, and 1998). Of the assemblages yielded by them, the earliest Palaeorhynchus). Some degree of freshening evi- one apparently comes from the Glarus Canton in Swit- denced by the Slovenian Ichthyofauna enables one to zerland and is distinguished in that it contains deepwa- suggest its proximity to the poorly known assemblage ter elements. For most other faunal assemblages (usu- of the Polbian Horizon of the Eastern Paratethys. ally assigned to the Middle Rupelian), typical is the presence of the beds with Aeoliscus (“Amphisile”) and Belgian Basin. In this area, the fish community of clupeid beds with “Meletta”, attributed to zones NP23 the Upper Tongrian (the basal Oligocene) demonstrates and NP24, respectively (Martini and Müller, 1971). the loss of most of the Late Eocene tropical forms. On the other hand, it comprises Odontaspis acutissima and In Switzerland, the Rupelian fish assemblage Myliobatis sp., which survived from the Lower Ton- includes the mesopelagic gonostomatids [Scopeloides grian (Eocene). The increase in taxonomic diversity is glarisianus (Ag.)], euzaphlegids (Palimphyes cha- demonstrated by the younger assemblage of the Berg dumicus Dan.), and merluccids [Palaeogadus crassus Sands comprising Squatina angeloides, Odontaspis (Ag.)], along with trichiurids (Anenchelum glari- acutissima, O. cuspidata, Notidanus primigenius, sianum Blainv.), fistulariids (Fistularia koenigi Ag.), Lamna rupeliensis, Oxyrhina desori, O. benedeni, gempylids (Thyrsitocephalus), scombrids, palaeo- Galeus latus, Chimaera gosseleti, Myliobatis aguila, rhynchids [Palaeorhynchus, Homorhynchus colei (Ag.)], and Cybium dumonti. The tropical and temperate forms ariommids [Isurichthys macrurus (Ag.)], balistids (Acanthoderma), etc. (Rath, 1859; Wettstein, 1886). 9 Most taxa of this ichthyofauna need revision. This community is indicative of the warm deep basin

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S205 and may be compared to Ichthyozone IPM1 of the Pol- The teleosts comprise elopids (cf. Elops sp.), congrids ish Carpathians as well as to the entire Pshekhian Fauna (Conger sp.), clupeids (Sardinella sardinites and Alosa of the Caucasus, since it contains a number of forms sculptata?), phycids (Eophycis sp.), merlucciids common for both the Early Planorbella Beds Subas- (Palaeogadus emarginatus and P. cf. intergerinus), semblage and the Aeoliscus Beds Assemblage. hemiramphids (Hemiramphus jerzyi), caproids (Capros The one more Early Rupelian assemblage might be longispinatus and C. radobojanus), aulostomids represented by that of Aschwil in Central Europe (Aulostomus medius and Frauenweilerstomus synarc- (Weiler, 1955). It is close to the Aeoliscus Beds Assem- ualis), urosphaenids (Oligosphenopsis cf. O. gracilis), blage of the Pshekhian Horizon of the Caucasus and to centriscids (Aeoliscus heinrichi and A.distinctus), syn- Ichthyozone IPM1 of the Carpathians by the presence gnathids (Doryrhamphus sp., ?Microphis sp., and Syn- of the genera Pristigenys, Leiognathus, Palimphyes, gnathus incompletus), moronids (Dicentrarchus sp.), Sarda, and Palaeorhynchus (apart from common syng- serranids (Properca sabbai and Serranus budensis), nathiforms). However, the absence here of the deepwa- priacanthids (Pristigenis spinosus), eheneids (cf. Ehe- ter elements, in contrast to the CarpathianÐCaucasian neis), carangids (Caranx cf. glarisianus), bramids Early Rupelian fish faunas, precludes their unequivocal (Pterycombus sp.), sparids (Sparus cf. S. schopii), tra- correlation. chinids (?Trachinus minutus), ephipids (?Archaephip- Of the apparently younger Rupelian ichthyofaunas pus sp.), sphyraenids (Sphyraena sp.), gempylids of Western Europe, the most comprehensively studied (cf. Gempylus sp. and cf. Epinnula sp.), trichiurids are those from the northeastern France (Froidefontaine, [Anenchelum (Lepidopus) glarisianum and Eutrichiu- the Belfort area) and Germany (Frauenweiler, south of rides (Trichiurides) delbeidi], scomprids [Scomber sp., Heidelberg) (Weiler, 1931, 1966; Theobald, 1934; Scomberomorus (Cybium) lingulatus, Sarda brachyceph- Pharisat, 1991, 1992; Micklich and Parin, 1996; Phari- ala, Scombrosarda sp., Scombramphodon sp., and sat and Micklich, 1998). Both are thought to corre- Sphyraenodus sp.], palaeorhynchids (Palaeorhynchus spond to zones NP23–NP24 (Müller, 1970). glarisianus), xiphiids (Xiphias rupeliensis?), nomeids In France, the Froidefontaine Assemblage includes (?Psenicubiceps sp.), stromateids (Pinichthys cf. 36 species of sharks and teleosts which dwelled in a P. pulcher), Psettodoidei indet., and Pleuronectoidei shallow and warm sea of normal salinity (Pharisat, indet. 1991). Two successive assemblages are distinguished here, both having the same species composition, with The core of this assemblage was formed by the taxa the younger one (that of “Meletta” Beds) being distinct of southern origin, which appear to have come from the in smaller body size of the forms recovered. In addition, southeastern Paratethys (Pharisat and Micklich, 1998). the earlier assemblage was dominated by the syng- Along with that, there was a limited group of immi- nathiform Aeoliscus heinrichi, while the later assem- grants from the Atlantic indicating links with that blage was dominated, by the clupeid Sardinella sar- region. The occurrence of a few mesopelagic forms dinites (Heckel). The shallow nearshore zone was pop- points to a moderate depth; on the other hand, the abun- ulated by the Aulostomidae and Syngnathidae, along dance of syngnathiforms and littoral forms character- with the acanthopterygians Capros, Serranus, Pro- izes the rich nearshore communities. The presence of perca, Caranx, Sparus, Pristigenys, Leiognathus, the taxa shared with the faunas of the Carpathians and Trachinus, etc.; the epipelagic component included Caucasus (Danil’chenko, 1960; Jerzman′ ska, 1968; Anenchelum glarisianum Blainv., Palaeorhynchus gla- Danil’chenko et al., 1980; Pharisat, 1992; Micklich and risianus (Blainv.), and the Scombridae (Sarda, Cybium, Sphyraenodus, and Glyptorhynchus) (Pharisat, 1991). Parin, 1996; Pharisat and Micklich, 1998) reflects the On the whole, by the set of the dominants contained, interconnections with these basins. It worth noting that this assemblage is most close to that of Frauenweiler the above-surveyed principal West European assem- (see below) and, along with the latter, characterizes the blages of the Middle Rupelian (Froidefontaine and seas of the epoch of the Early Oligocene transgression. Frauenweiler) were devoid of the bathypelagic photo- phore-bearing forms, which differs them from the Car- In Germany, the Rupelian assemblage of Frauen- pathian and Caucasian Aeoliscus (= “Amphisile”) fau- weiler (Micklich and Parin, 1996; Pharisat and Miklich, 1998) similarly displays a broad scope of taxa, includ- nas synchronized with Zone NP22. Judging by their ing over 40 genera that pertain to 32 families and datings in terms of the nannoplankton zonation (NP23Ð 13 orders. The sharks and rays include Physogaleus NP24), these assemblages should correspond in age to (Eugaleus) latus (Storms), Galeocerdo sp., ?Rhizopri- the poorly known terminal Rupelian faunas of the East- onodon sp., Synodontaspis acutissima, S. cuspidata, ern Paratethys , i.e., to those of the Polbian Horizon and Alopias (Alopecias, Vulpecula) exigua, A. latidens, the basal part of the Morozkina Balka Horizon. In a Cetorhinus parvus, Isurus desori, Lamna rupeliensis, broad sense, all these faunas inhabited relatively shal- Charcharocles (Charcharodon) angustidens, Noto- low seas. By contrast, the coeval faunas of the Car- rhynchus primigenius, Squalus alsaticus?, Squatina pathians (icthyozones IPM2ÐIPM4?) belong to the angeloides?, Myliobatis oligocaena, and M. serratus. basins of considerable depth.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S206 POPOV et al.

Discussion. Heterogeneity and marked differentia- (Polbian Horizon) and not so much in the Carpathian tion of the marine fish communities observable in the Basin. Some evidence of brackishing is recorded for the Carpathian and Caucasian realms during the Early Oli- Balkan Basin. The coeval West European basins were gocene make one very cautious when comparing the of moderate depth, with the fish fauna including a great faunas of that age known from various regions of the variety of the nearshore dwellers. Paratethys, since the distinctive features of particular (5) By the end of the Rupelian, a full salinity of the basins reflected by them may be related to the different Paratethys was restored. This was accompanied by the time intervals. Hence, the analysis of spatial differenti- deepening of some basins, which was moderately ation of the Paratethyan Rupelian fish faunas heavily expressed in the Eastern Paratethys and much more depends not only on the knowledge of their composi- strongly in the Carpathian Basin (which resulted in tion but also on the dating accuracy for the assemblages reappearance here of the mesopelagic fauna). under comparison. At present, the data on the distribu- tion patterns of the regional fish faunas are not enough In summary, it may be concluded that, based on the to elaborate a clear picture of the overall biogeographic present evidence, the Paratethyan marine fish faunas of structure of the Paratethys in the Early Oligocene. Only the Early Oligocene appear to have developed within a a few more or less common events of different scale, single biogeographic province. It is not ruled out that characterizing the history of the fish fauna of the realm with further increase in data it will be possible to under discussion may be currently outlined. They are as ground the biogeographic discrimination of the West follows: European, Carpathian, and Caucasian fish faunas of the terminal Rupelian. One more possibility is the distin- (1) The development of the Rupelian marine fish guishing in the future of a separate Iranian Province. faunas of Europe started on the basis of the authoch- tonous Tethyan Ichthyofauna whose local communities were abundantly represented by the assemblages of the Smaller Benthic Foraminifers Caucasus, Carpathians, and the Mediterranean. During the Rupelian, the faunal evolution was mostly con- Abrupt paleogeographic, climatic, and facial trolled by the changes in salinity and depth of the local changes in the terminal EoceneÐat the beginning of the basins along with changes in climatic conditions. The Oligocene resulted in similar abrupt changes in the faunal exchange detectable for the Rupelian Time dem- composition of small benthic foraminiferal assem- onstrates that the particular Paratethyan basins were blages throughout the Northern Peri-Tethys. The taxo- open for interconnection to a considerable extent. nomic diversity and the faunal abundance markedly decreased, whereas the tests became small and medium (2) The earliest faunal event of the Rupelian Time, in size. However, such changes did not mean substan- most readily detected in the Belgian Basin (Upper Ton- tial evolutionary changes in this group, because, in grian) and Caucasian Basin (sub-Planorbella level), warm seas, where carbonate sedimentation continued was the appearance of the impoverished fish assem- (Mediterranean Sea and Armenian Gulf), numerous blages formed mostly by the survivors of the preceding speciesÐdescendants from the Eocene were found in Late Eocene fauna. Subsequently, they were enriched the Oligocene assemblages (up to 46%, after Krash- by the boreal (Atlantic) newcomers and the further evo- eninnikov, in Geologicheskie i bioticheskie..., 1998). lution of autochtonous elements. It was established that the evolutionary changes of (3) As early as the first half of the Rupelian (at the benthic foraminifers in oceanic basins were very grad- end of the Early Pshekhian Time), with the increase in ual, without catastrophic extinction (Geologicheskie i temperature in both surface and deep waters of the Cau- bioticheskie..., 1998). Certain groups of organisms casian Basin, there was started a wide expansion of the were simply replaced by others. Such changes are tropical and subtropical elements that most likely came observed at different levels, in the terminal Eocene and from the Tethyan refuges, judging by the presence in at the EoceneÐOligocene Boundary. the Pshekhian Fauna of a great amount of the genera Northern Mediterranean Region. Tropical and inherited from the Tethyan Eocene Ichthyofauna. These subtropical fauna of the Mediterranean Region is trends were accompanied by the deepening of most known from northern Italy, and Armenia, where, apart basins, which resulted in the appearance in some cases from small foraminifers, nummulitid assemblages were of a great variety of photophore-bearing forms and the distributed and reef facies developed. The composition overall development of the rich mesopelagic fish biota. of the foraminiferal assemblages indicates the shelf This condition persisted into the Mid-Rupelian (Aeolis- environments closely similar to those in the terminal cus Beds time span) and is best typified by the fish com- Eocene. Thus, in northeastern Italy, a coral reef biofa- munities of the Caucasian and Carpathian basins and, cies was found (Frost, 1981), in which the foraminiferal to some extent, by that of the Alpine Basin. assemblages markedly differed from the North Euro- (4) The succeeding event, recorded to various pean and Chadumian assemblages in generic composi- degrees in some parts of the Paratethys, was the shal- tion, including Halkyardia, Chapmanina, Asterigerina, lowing and (or) brackishing of the basins. This was and the Gypsinidae. Northern Italy and northern Spain most markedly expressed in the Eastern Paratethys were also inhabited by thermophilic genera. Halk-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S207 yardia, Chapmanina, Borelis, Haddonina, Gyroidinella, Numerous miliolids (Quinqueloculina, Articulina, Asterigerina, and Sphaerogypsina are known from the and Spiroloculina), Polymorphinidae (Globulina and northern Adriatic coasts, where reef facies are present Guttulina), and the genera Discorbis, Nonion, Cibici- (Pavlovec et al., 1986). doides, and Planorbulina widespread in the Lower Stampian of the Paris Basin (see Syntese geologique..., The Oligocene foraminiferal assemblage from 1980) indicate facial individuality of the assemblage, Armenia is very rich and diverse and includes over which do not include the species characteristic of the 70 species of 53 genera. This assemblage differs in North Sea and Paratethyan faunas. One hundred and both specific and generic composition from the Parat- ten species of benthic foraminifers were recorded there ethyan Assemblage, because it includes many Tethyan (after LeCalvez). species (Bugrova, 2001). These are species of the gen- era Almaena, Queraltina, Herondenia, Pseudoplan- North Sea Basin. In the basin of northwestern ulinella, Sphaerogypsina, Asterigerina, Rotalia, and Europe (Alsace, Belgium, Holland, and Germany), the Pararotalia. In addition, this assemblage includes a Rotaliatina bulimoidesÐCibicidoides ungerianus assem- few species that are known from West European basins blages were distributed (Doppert and Neele, 1983; (Valvulineria petroleyi, Cibicidina amphisyliensis, Andreae, 1884). Many species from these assemblages, Svratkina perlata, Rotaliatina mamillata, Turrilina including Lenticulina herrmanni, were described from alsatica, Angulogerina gracilis, Bitubulogerina kasse- the Alsace Basin (Andreae, 1884). Rotaliatina buli- lensis, Bolivina melettica, and Spiroloculina canalicu- moides, Svratkina perlata, and Turrilina alsatica are lata). Some of these species, as well as Melonis dosu- present only in these assemblages, whereas Cibici- larensis, Heterolepa almaensis, Vsegeina nana, and doides ungerianus, C. sulzensis, Ceratobulimina con- Falsoplanulina, sp. nov., were also found in the assem- traria, and Hoeglundina elegans appeared there and blages from the CrimeanÐCaucasian area of the Pshek- were widespread in the Eastern Paratethys. The North hian (Chadumian) Basin. In addition, this area is inhab- Sea Basin was inhabited by some of these species, ited by endemic species and species that were only Svratkina perlata, Rotaliatina bulimoides, and Turril- found in Azerbaijan (species of the genera Bolivia, ina alsatica, as well as Ceratobulimina contraria (Gra- Chilostomella, and Pararotalia). Such a mixed compo- mann and Hiltermann in Colloque..., 1964). sition of fauna indicates the presence of contacts of the In the region of the English Channel (Isle of Wight Yerevan Basin with the open Mediterranean Basin and in the southern area of England), the fauna is similar to with the neighboring Azerbaijan Basin. There was that from the Belgian Basin. Sixty-two species of probably no direct connection with the Greater Cauca- 33 genera are known from this region (Bhatia, 1958). sus Basin. The foraminiferal assemblages are facially changeable, Eastern Mediterranean. Benthic foraminifers but usually dominated by secreting (Miliolidae and the dominate the Oligocene assemblages from Syria. Three genera Nonion, Rotalia, Valvulineria, Bulimina, and zones were distinguished based on the differences in Buliminella). the temporal compositions of the benthic foraminiferal In the NorwegianÐGreenland Basin (Knipovich assemblages (Krasheninnikov, 1969). The fauna from Ridge), the foraminiferal assemblages were similar to the lower zone, the Almaena taurica Zone, included those from the North Sea basins but slightly impover- typical Late Eocene species and certain species that ished. This basin was inhabited by Pullenia quinque- appeared in the Oligocene of the Crimea and Caucasus. loba, Nonionella aff. odiliae, Alabamina tongentialis, The majority of species of the middle zone, the Cibici- Pseudoparrella oveyi, Cibicidoides sulzensis, Glob- doides pseudoungerianus Zone, including the zonal ocassidulina aff. oblonga, G. subglobosa, Bulimina cf. species Cibicidoides oligocenicus, Cibidina amphi- elongata, Buliminella elegantissima, Fursenkoina syliensis, Melonis dosularensis, Angulogerina gracilis, schreibersiana, Bolivina melettica, and B. variabilis etc., are typical of the assemblage of the Lenticulina (the total of 17 species after Bugrova et al., 2001). For- herrmanni Local Zone from the CaucasusÐKopet Dagh aminifers were small in size and thin-walled, which Basin (Reshenie kollokviuma..., 1970). The thermo- indicates environments unfavorable for the secreting philic species that were only found in the Tethyan part forms. Representatives of the orders Nodosariida and of the basin are absent from the two upper zones. Polymorphinida, which, in association with miliolids Eastern Atlantic. In the Oligocene, the Aquitaine and nonionids, predominated in the assemblages from Basin was still characterized by very warm water, as Belgium, Germany and England, were absent from this indicated by the presence of the Tethyan genera Bulla- basin. lveolina, Sphaerogypsina, Amphistegina, Halkyardia, The foraminiferal assemblage that inhabited the Queraltina, Victoriella, Falsocibicides, Carpenteria, western area of Belgium includes 49 species of 27 gen- Eorupertia, etc. (Boulanger and Poignant, 1971; Bou- era (Willems, 1972). Among foraminifers, nodosariids langer et al., 1970), which are often associated with (five Dentalina species and five Lagena species), poly- reef beds. This fauna has no equivalents among the morphinids (seven species of three genera), and noni- assemblages from the northeastern marginal areas of onids (five species) were numerous. Heterolepa dutem- the Paratethys. plei, Cibicidoides tenellus, and Cribrononion subno-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S208 POPOV et al. dosum were most frequently found. This assemblage nated by local species of the genera Cibicidoides and differs from those in other areas of the North Sea Basin Elphidium. in the species and generic composition, which may be The southern shelf of the Carpathian Basin (Kiscell considered as facial dissimilarities. The association Clays, northern Hungary) was inhabited by littoral and also markedly differs from the Paratethyan assem- lagoonal assemblages. Of 160 forms registered there blages. The only species in common are Cibicidoides (Hantken, 1875; Koreczne-Laky and Nagyne-Gellai, sulzensis and Grammostomum teretum. In other areas 1985), 69 species of 27 genera belong to the arenaceous of the Belgian Basin, the assemblages typical of the group; less than half of these genera are of a primitive Rupelian, including Svratkina perlata, Rotaliatina morphotype. The lagoonal inhabitants, miliolids, are bulimoides, Turrilina alsatica, Angulogerina gracilis, rather diverse (six genera and 16 species); the Polymor- and Bolivina melettica, were found (Batjes, 1958; phinidae are represented by 18 species of five genera. Drooger, see Colloque..., 1964). Some species are also present in the Chadumian and In the North German Basin, the Early Rupelian North European assemblages, including Pararotalia fauna was impoverished and still similar to the Late canui, Epistomina elegans, Ceratobulimina contraria, Eocene fauna in composition (Spiegler, 1966). This Svratkina perlata, Alabamina tongentialis, Grammos- included arenaceous foraminifers of the genera Rhab- tomum teretum, Quinqueloculina seminula, and dammina, Rhizammina, Ammodiscus, Usbekistania, Spiroloculina canaliculata (Nagy-Gellai, 1968). The Haplophragmoides, Cyclammina, Spiroplectammina, generic composition of other secreting foraminifers and Karreriella (Rupel 1). Calcareous forms were rep- (47 genera) markedly differs from that of the assem- resented by 21 species of 14 genera and by a group of blages that inhabited the CaucasusÐKopet Dagh Basin polymorphinids. Species in common with the fauna in the presence of the Tethyan genera Amphicoryna, from the northeastern area of the Paratethys were Trilo- Lankesterina, Bolivinella, Sagrina, Escornebovina, culina enoplostoma, Spiroloculina canaliculata, Cibic- Patellina, etc. idoides pseudoungerianus, C. sulzensis, Ceratobulim- Eastern Paratethys. In the Paratethys, at the ina contraria (= ? C. intrusa), Rotaliatina bulimoides, EoceneÐOligocene boundary, the foraminiferal group Alabamina tongentialis, Svratkina perlata, Vsegeina experienced profound changes, which resulted in the sp., Epistomina elegans, Turrilina alsatica, Bulimina disappearance of highly specialized taxa, general alsatica, and Angulogerina oligocenica. In other areas impoverishment of the composition, and the appear- of this basin, the assemblages were less representative ance of new groups with a simpler morphotype. and included 35 species of 21 genera (Jung and Langer, 1990). A total of 167 Rupelian species was reported In the Priabonian, the foraminifers were represented (Spiegler, 1966). by over 150 species of 80 genera of all orders; this gen- erally indicates the biological progress of their evolu- In Poland in the Late Rupelian (NP24 Zone), an tionary development. In the terminal Priabonian, the assemblage that was similar to the West European changes were primarily associated with shallowing; the assemblages rather than to the Chadumian assemblage quantity of plankton decreased, whereas the proportion was widespread. This assemblage included Lenticulina of arenaceous foraminifers increased and they included herrmanni, Ceratobulimuna contraria, Rotaliatina a great number of primitive forms. Nevertheless, the bulimoides, Svratkina perlata, Pararotalia canui, Tur- evolutionary changes, in particular, the appearance of rilina alsatica, Robertina germanica, etc. (Odrzywol- new genera and species, were also observed. Some new ska-Bienkowa and Pozaryska, 1981; Odrzywolska- taxa appeared shortly before the crisis, originally occu- Bienkowa, 1983). Forty-one species were in common pied subordinate positions in communities, but became with the East German assemblages. This similarity more representative and widespread after the crisis; decreased further west to 30 species. Twenty-one spe- others existed only during the short-term crisis phase, cies were in common with the Belgian Assemblage, in which new elements also appeared. and only nine species (including Rotaliatina buli- In the Early Maikopian (Pshekhian) Basin, before moides) were in common with the Ukrainian Assem- its brackishing in the Solenovian Time, the species of blage. The mediterranean genera were absent from the two assemblages, i.e., the earlier assemblage with Len- Polish area of the basin. ticulina herrmanni and the later assemblage with AlpineÐCarpathian Basin. In the Rupelian, the Spiroplectammina oligocenica, were widespread. These fauna from the deepwater (Skibas) zone of the Eastern assemblages are well traced throughout the Early Carpathians was mainly represented by endemic spe- Maikopian Basin and are designated together as the cies that were unknown further east (Mjatliuk, 1970). Chadumian Assemblage. In the basal Rupelian, the This assemblage markedly differed from those from composition of assemblages abruptly changed and a other Paratethyan areas, even in generic composition. marked decrease in the diversity occurred. However, in The comparison between the Oligocene East Car- particular sections, assemblages of a mixed type were pathian Assemblage and the North Caucasian assem- often found. In addition, the tests were redeposited blages made by Dabagyan needs a revision; neverthe- through the bottom erosion; as a result, the assemblages less, it may be said that these assemblages were domi- include the EoceneÐOligocene species. The composi-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S209 tion, structure, andecological type of these assemblages widespread. This assemblage includes the species from considerably depended upon the facies; nevertheless, the North German and Chadumian basins, Lenticulina benthic forms started to predominate everywhere. The herrmanni, Cibicidoides oligocenicus, Cibicidina impoverished composition and the morphology of the amphisyliensis, Caucasina schischkinskayae, Angulo- foraminifers indicate more unfavorable environments gerina oligocenica, lagenids, miliolids, and nonionids compared to those of the Late Eocene. (Yartseva, 1959). The impoverished foraminiferal The typical species of the earlier assemblage (Lenti- assemblage from other areas of the DnieperÐDonets culina herrmanni Local Zone) are Lenticulina her- Depression includes Cibicidoides oligocenicus, Melo- rmanni, Plectofrondicularia volgensis, Heterolepa nis dosularensis, and several species with the calcare- almaensis, Cibicidoides aff. pseudoungerianus, C. sulzen- ous test. In the Rostov Region (Belyaeva, 1964), the sis, C. extremus, C. oligocenicus, Cibicidina amphi- foraminiferal assemblage includes species of the Lenti- syliensis, Melonis dosularensis, Bolivina mississippi- culina herrmanni Zone: the zonal species Cibicidoides ensis (Maikopskie otlozheniya..., 1964; Reshenie kol- oligocenicus, C. aff. pseudoungerianus, Cibicidina lokviuma..., 1970), Haplophragmoides deformabilis, amphisyliensis, etc. Further to the north (Belarus and Gaudryinopsis gracilis, etc. In the lower part of this Lithuanian Depression), the Oligocene foraminifers are zone, transitional species and the species that appeared absent. at the end of the Eocene and occurred in the boundary The Early Oligocene assemblage from the Crimea beds (Robulus diaphanus, Almaena taurica, Svratkina (from the carbonate siltstones of the Kyzyldzhar Hori- perlata, Vsegeina nana, Tergrigorjanzaella sectile, Angu- zon) is rather diverse compared to the fauna from other logerina gracilis, etc.) were found. regions and includes 35 species of 24 genera. Of the Foraminifers from the Solenovian Basin are either arenaceous taxa, only Spiroplectammina azovensis was unknown or represented by rare alien forms. found; nodozariids were absent, except for the zonal species Lenticulina herrmanni. Each genus is usually Scythian Shelf. In the western area of Moldova represented by a single species, except for Cibici- (Pechenkina in Maikopskie otlozheniya..., 1964), the doides, which is represented by six species, including assemblage from the Lower Oligocene noncarbonate C. oligocenicus, C. extremus, C. crimensis, C. pseudoun- beds is composed mainly of the agglutinating forms gerianus, and C. tahtaensis. Heterolepa almaensis and and dominated by the primitive genera Rhabdammina, Bolivina mississippiensis, which also occur beyond the Saccammina, Reophax, Ammobaculites, and Haplo- Crimea, are present there. In the beds adjacent to the phragmoides. An assemblage of the primitive arena- Priabonian, Almaena taurica and Vsegeina nana are ceous foraminifers was also distributed in the Azov area known. Higher in the succession, the assemblage of the of the Early Oligocene Basin. These assemblages of Spiroplectammina oligocenica Zone occurs; it includes arenaceous foraminifers represented a facies variety of Caucasina schischkinskayae and Melonis dosularensis. the fauna from the Lenticulina herrmanni Zone of the Silty calcareous sediments in the plain territory of the Early Chadumian Basin. Crimea were inhabited by agglutinating forms of the At the beginning of the Oligocene, the northern area genera Ammodiscus (A. tenuiculus) and Haplophrag- of the Black Sea Depression was inhabited by arena- moides (Pechenkina, 1971). ceous foraminifers (“the zone of arenaceous foramini- In Ciscaucasia, in the deepwater western, central fers,” after Kraeva, 1960, 1961; Kraeva and Pechen- (Kuban River), and eastern areas of the Pshekhian kina, 1965; Maikopskie otlozheniya..., 1964), mainly (Early Chadumian) Basin, rare impoverished fauna of the primitive genera Rhabdammina, Proteonella, occurs. There are about 20 genera there; each is usually Saccammina, Hyperammina, Reophax, Ammodiscus, represented by one species. In addition to the wide- Ammomarginulina, Ammobaculites, Haplophragmoides spread species (such as Heterolepa almaensis, Cibici- (H. deformabilis and H. stavropolensis), Spiroplectam- doides aff. pseudoungerianus, C. oligocenicus, Cibi- mina (S. azovensis), and Gaudryinopsis gracilis. How- cidina amphisyliensis, Melonis dosularensis, and ever, rare calcareous frorms (about ten genera), includ- Bolivina mississippiensis), rare findings of Pararotalia ing Lenticulina herrmanni, Cibicidoides pseudoungeri- canui and Cassidulina sp. have been registered (Kuban anus, and C. oligocenicus, were also found. The above River). The foraminifers are usually small-sized and species were typical of the Early Oligocene (Lenticu- thin-walled, indicating unfavorable environments. lina herrmanni Zone) of the northern Caucasus. Noncarbonate sediments were inhabited by the arena- The later assemblage (Spiroplectammina oligoce- ceous species of the genera Saccammina, Rhizammina, nica = S. carinata Zone) includes Cyclammina con- Trochamminoides, Ammodiscus, Haplophragmoides, strictimargo, Verneuilina rasilis, Caucasina schisch- and Cyclammina. kinskayae, and C. oligocaenica (Kraeva and Pechen- Further to the north, in shallower-water facies of the kina, 1965); these species also occur in the Caucasus Stavropol Region, Kuban Lowland, and the Trans- and in Transcaspia. Terek Lowland, an increase in the content of arena- On the southeastern slope of the DnieperÐDonets ceous foraminifers is observed, whereas the calcareous Depression, an assemblage of the L. herrmanni Zone is forms decrease in number. Haplophragmoides fidelis

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S210 POPOV et al. and H. deformabilis zones, which correspond to the inhabited by a more representative fauna (18 genera of Lenticulina herrmanni Zone, were established there calcareous foraminifers) having much in common with (Ter-Grigorjants, 1965; Reshenie kollokviuma..., 1970). the Rupelian fauna of Western Europe (Kacharava in In association with the species typical of this zone and Maikopskie otlozheniya..., 1964). At the same time, it widespread in the Crimea and northern Caucasus, two includes the species of the Chadumian Assemblage; new index species and Haplophragmoides stavropolen- this suggests the presence of connections between these sis, Ammomarginulina foliacea, Trochamminoides sp., basins. Ammobaculites grossecameratus, Trochammina parva, The foraminiferal assemblage discovered in the T. caucasica, Verneuilina aff. risilis, Gaudryinopsis gracilis, Pseudogaudryina trigona, Plectofrondicu- Early Oligocene of Azerbaijan is extremely impover- laria volgensis, Heterolepa almaensis, Pararotalia ished in both composition and abundance. This region was inhabited by the fauna with a large proportion of canui, Asterigerina gurichi, Caucasina aff. schischkin- endemic forms. This fauna markedly differed in generic skayae, Tergrigorjanzaella sectile, Uvigerina sp., etc. composition from the assemblages of the Pshekhian (a total of 16 genera, mostly of the agglutinating group) and Mediterranean basins; the species composition are also present. Occasionally, species of Planorbella depended upon the facial characteristics of sediments. were found in association with foraminifers. In Western Thirteen genera were recorded there, including Elphid- Ciscaucasia and in the IndolÐKuban Depression, the ium Nonion Non- Haplophragmoides deformabi- (three species), (three species), and assemblages including ionella lis, H. stavropolensis, Ammobaculites grossecamera- , which presence indicates the existence of lagoonal, possibly brackish-water areas (Khalilov, 1962; tus, Melonis dosularensis, and Lenticulina herrmanni Khalilov and Mamedova, 1984). In addition, local spe- occur; they are also typical of the Stavropol Region. cies of the genera Cibicidoides, Pararotalia (five spe- The foraminiferal assemblages from the SalÐManych cies), Caucasina, Bolivina (four forms), Chilostomella, and Don–Sal interfluves are transitional. These assem- and Ammosphaeroidina were distributed. Only the spe- blages include species known from the Stavropol cies Melonis dosularensis, Rotalia canui, and Cibicid- Region (from the Haplophragmoides fidelis and ina amphisyliensis were in common with those from H. deformabilis zones) and from more widely distrib- the Chadumian Assemblage. uted fauna of the Lenticulina herrmanni Zone, i.e., the zonal species, Heterolepa almaensis, H. pileola (abun- Turanian Sea. In the northeastern area of the Fore- dant), Melonis dosularensis, Plectofrondicularia vol- Caspian Lowland, species with calcareous tests (Het- gensis, Bolivina mississippiensis, etc. (Maikopskie erolepa almaensis, Cibicidoides ex gr. oligocenicus, otlozheniya..., 1964; Nikitina, 1972). Cibidina amphisyliensis, Melonis dosularensis, and Bolivina mississippiensis) typical of the Lenticulina In the northwestern area of the Fore-Caspian herrmanni Zone were found. the index species of the Depression, the foraminifers that characterize the Cha- Brotzenella munda Zone that was distinguished in the dumian Assemblage, Cibicidoides oligocenicus and AralÐTurgai Region appeared. In the Southern Emba Cibicidina amphisyliensis, are present (Maslun et al., Region, an assemblage of a mixed type was also found: 1984). A rather impoverished assemblage, which miliolids (of the genus Quinqueloculina) and Brotzenella included both arenaceous (six genera) and calcareous munda were present in association with the species of (seven genera) forms, is known from the central area of the Lenticulina herrmanni Zone, including the index this depression. The northeastern assemblages of a species, Spiroplectammina azovensis, Cibicidoides mixed type include species of the Lenticulina her- expertus, C. kugultaensis, C. aff. pseudoungerianus, rmanni and H. deformabilis zones (Grachev et al., and Bolivina mississippiensis (Paleogenovaya sistema…, 1971; Paleogenovaya sistema…, 1975). Melonis dosu- 1975; Tengiz, collected by S.K. Nikolaeva). larensis is abundant in this region. In addition, the first miliolids (Quinqueloculina errmanni) typical of the In the Buzachi area of the Turanian Sea (Bykova and Oligocene of Western Europe and of the more easterly Azbel’, 1962), at the beginning of the Oligocene (Bur- areas of the Pshekhian Basin, but unknown further lin or Uzunbas Formation), a rather abundant benthos west, appear there. In the North Ergeni Upland, Haplo- of the Heterolepa almaensis Zone (26 genera) with the phragmoides fidelis and Cibicidoides oligocenicus presence of Eocene species was found. Arenaceous were found (Geologicheskie i bioticheskie..., 1996). forms (three genera) were rare and only reported from The proportion of foraminifers gradually decreases the base of this formation. The major part of the assem- from north to south down to their disappearance. blage was composed of calcareous foraminifers. Of them, Heterolepa almaensis, H. pileola, and Lenticu- Transcaucasia. In the western area of Georgia lina herrmanni were of particular importance. in the (Abkhazia), dark-colored silty sediments were inhab- Oligocene of Mangyshlak, a series of foraminiferal ited by a very impoverished foraminiferal assemblage assemblages were recorded (Paleogenovaya sistema..., with Melonis dosularensis and Bolivina mississippien- 1975) that included polymorphinids and miliolids, Len- sis; it was also distributed in the Ciscaucasian area of ticulina herrmanni, Heterolepa almaensis, Melonis this basin (Archvadze and Kacharava in Maikopskie dosularensis, Bolivina mississippiensis, Pararotalia otlozheniya..., 1964). The Akhaltsikhe Depression was canui, and Pseudoparrella caucasica, typical of the

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S211

Lenticulina herrmanni Zone, as well as Cassidiella 34 species of 29 genera were found there (data from mustoni, known from the Rupelian of Northwestern boreholes in the vicinity of the towns of Nukus and Europe. In the northern area of Fore-Kara Bagaz Gol, Tashauz). Among the arenaceous forms (seven genera), an assemblage of the Lenticulina herrmanni Zone was Haplophragmoides deformabilis, Gaudryinopsis gra- found, which included this species, Spiroplectammina cilis, and Pseudogaudryina trigonia (typical of the Oli- oligocenica, Quinqueloculina errmanni, Spirolocu- gocene of the Stavropol Region) were found there. Len- lina canaliculata, Melonis dosularensis, Cibicidoides ticulina aff. herrmanni (in the lower part), L. herrmanni, pseudoungerianus, C. oligocenicus, Loithostomata Cibicidoides pseudoungerianus, C. oligocenicus, Cibi- macrura, Planorbulina hadlei, etc. (Bugrova, 1986). cidina amphisyliensis, Melonis dosularensis, and These species were widespread in the Aral Region of Bolivina mississippiensis, known from the deeper- the basin. In the northwestern area of the Ustyurt Pla- water area of the Caucasus-Kopet Dagh Basin, also teau, foraminifers were represented by species in com- occurred Planorbells were recorded. mon with those of the Kyzyldzhar Assemblage from the In addition, this region was inhabited by Brontz- Crimea (Nikolaeva and Prusova, 1978). enella munda ssp. assakensis, miliolids (Triloculina The problem of changes in the foraminiferal assem- enoplastoma, Quinqueloculina errmanni, Spirolocu- blages at the EoceneÐOligocene boundary in the Aral- lina canaliculata, etc.), Ceratobulimina cf. intrusa, Turgai Region deserves special consideration Loithostomata macrura, and polymorphinids typical of (Bugrova, 2001). Bondareva established the “Anoma- the AralÐTurgai Region of this basin. Thus, the fora- lia” (= Brotzenella) munda Zone there and placed the miniferal assemblage from the South Aral Region was EoceneÐOligocene boundary at the base of this zone. Mel- a fauna of the transitional (mixed) type. In the southwest- onis dosularensis, Ceratobulimina intrusa (= ?C. con- ern Aral Region, an assemblage comprising 26 species traria), and Tergrigorjanzaella sectile appeared in the of 21 genera (borehole 495, Torangly) was found, lower part of this zone (in the beds with Cibicidoides almost half of which was represented by typical Rupe- salensis, which probably belonged to the Eocene), lian species of northwestern Europe, including Cibici- although, in other regions, these species occur at the doides pseudoungerianus, C. sulzensis, C. kiliani, Cer- EoceneÐOligocene boundary. atobulimina contraria, Rotaliatina bulimoides, Val- The current territory of the North Ustyurt and Chag- vulineria petroleyi, Alabamina tongentialis, Svratkina rai plateaus was inhabited by an impoverished assem- perlata, Hoeglundina elegans, Angulogerina oligoce- blage that was similar to the Caucasian assemblages in nica, Grammostomum teretum, and, in the upper part of composition (Bronevoi et al., 1967). This assemblage the zone, Robertina sp. In addition, the taxa in common included arenaceous forms, Brotzenella munda, Cera- with those from the Ciscaucasian area of the basin were tobulimina intrusa (= ? C. contraria), and Lenticulina found, including Cibicidoides oligocenicus, C. extre- herrmanni. This is similar to the European North Sea mus, and Melonis dosularensis, and the subspecies assemblages in composition but has not been found in Brotzenella munda assakensis, which was restricted to other areas of the Paratethys; therefore, it is difficult to the eastern marginal area of the basin (to some area of trace its migration. Kazakhstan and Central Asia). The fauna from the North Aral Region of this basin Central and northern Turkmenistan, the western markedly differs from the typical Chadumian Assem- area of the DariyalykÐDaudan Depression, and the blage and includes representatives of the order Miolida, Zaunguzskie Kara Kum were inhabited by an assem- 16 species and subspecies of the genera Quinquelocu- blage of the Lenticulina herrmanni Zone that was lina, Triloculina, and Spiroloculina (Serova, 1962). mixed in composition. This assemblage included spe- Besides the endemic taxa, of 13 taxa that first appeared cies of the Chadumian Basin and miliolids typical of in the Oligocene, eight were in common with those of the Aral Region: Lenticulina herrmanni, Haplophrag- the West European assemblages, including Quinquelo- moides deformabilis, Cibicidoides aff. pseudoungeri- culina akneriana (three subspecies), Q. selene, Q. gra- anus, C. oligocenicus, Heterolepa almaensis, Cibicid- cilis, Tritoculina enoplastoma, and T. austriaca. Out- ina amphisyliensis, Caucasina schischkinskayae, side the AralÐTurgai Region, only individual species of Quinqueloculina errmanni, Q. akneriana, and Spirolo- this assemblage were found. In addition to miliolids, culina canaliculata. In the Zaunguzskie Kara Kum, species of 13 genera from the Brotzenella munda Zone Brotzenella munda assakensis appeared (Paleogen occurred there, including species that had wide geo- Turkmenii, 1975). graphical ranges (Melonis dosularensis and Turrilina In the Western Kopet Dagh area of the basin, over alsatica). 35 species of at least 25 foraminiferal genera were Foraminifers from the southern part of the Turanian found. The Lenticulina herrmanni Zone was distin- BasinÐSouth Aral Region, southeastern Ustyurt, and guished based on these taxa. Among the arenaceous northeastern area of the Zaunguzskie Kara Kum have forms (eight genera), the following are of special inter- already been studied in detail (Ivanova, 1971; Paleogen est: Haplophragmoides fidelis, H. deformabilis, Turkmenii, 1975). Foraminifers of the Lenticulina her- H. stavropolensis, and Gaudryinopsis gracilis, the spe- rmanni Zone (with the zonal species) represented by cies that also inhabited shallow-water areas of the

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S212 POPOV et al.

Ciscaucasian Basin (Stavropol Region, etc.). Among Tajik-Ferghana Basin was an isolated peripheral other typical species of the Chadumian Assemblage, area of an Early Oligocene basin that was inhabited by there were Lenticulina herrmanni, Heterolepa almaen- endemic fauna. The majority of species found there are sis, Cibicidoides expertus, C. oligocenicus, Planorbu- unknown outside the TajikÐFerghana Region (David- lina ammonoida, Cibicidina amphisyliensis, Melonis zon et al., 1982). The assemblage includes many aren- dosularensis, Caucasina schischkinskayae, Anguloger- aceous forms, local species of the genera Haplophrag- ina gracilis, Uvigerinella californica, and Bolivina moides and Trochammina, Spiroplectammina tuaevi, mississippiensis. This region was also inhabited by and species of the genera Nonion and Cribrononion. Brontzenella munda assakensis reported from the Aral Brotzenella munda, B. postacanica, B. multa, B. schura- Region of the basin and by the species in common with bica, Asianella subbotinae, and Cibicidoides khana- the TajikÐFerghana Region (Asianella subbotinae, badensis had wider ranges (including Turkmenistan). Brotzenella postacanica, B. multa, and B. schurabica). According to Tsatsir, Loithostomata macrura occurred In addition, the genera Nonion, Astrononion, and Non- in the area of the basin that today belongs to the south- ionella, which were absent or rare in other regions, western offshoots of the Hissar Mountains. Some of the were found there. Shallower-water assemblages from above species were recorded from Ferghana (Reshenie the far southwestern area of the Kopet Dagh included kollokviuma..., 1970, data by Bugrova) and from north- species from the TajikÐFerghana Region and miliolids ern Afghanistan. (Quinqueloculina errmanni and Triloculina enoplas- toma) typical of the Pre-AralÐTurgai Region (Geolog- icheskie i bioticheskie..., 1996). In another area of this Nummulites and Discocyclinids basin (Gyaurli), a unique assemblage was found; it was dominated by miliolids but had different composition In the Oligocene, the geographical range of large compared to the Pre-AralÐTurgai assemblages. In par- foraminifers sharply decreased. In the Northern Num- ticular, the genus Pyrgo, unknown in other regions, was mulite Province, these organisms became entirely found there. extinct, and only at the end of the Oligocene (in the Chattian), sole species of the genera Lepidocyclina, Further to the east, in the Central Kopet Dagh and Miogypsina, and Miogypsinoides entered this region. Gyaurs Dagh areas of this basin, the fauna was impov- In the southern province, large foraminifers still erished and included endemic forms and Brotzenella existed, but the diversity of the assemblages substan- munda assakensis (Paleogen Turkmenii, 1975), although tially decreased. The basin cooling affected nummu- species recorded from Ciscaucasia (Haplophragmoides lites; their tests became smaller and nongranulated; aff. fidelis, Gaudryinopsis gracilis, Plectofrondicularia and, in the terminal Oligocene, this genus became volgensis, and Uvigerinella californica) were also extinct. recorded there. Further east, (in the piedmont area of the Eastern Kopet Dagh), an assemblage with Uviger- At the EoceneÐOligocene boundary, marked inella californica, Cibicidoides oligocenicus, and non- changes in the taxonomic composition of this foramin- ionids occurred. iferal group occurred. Only a small number of species The Middle Amu Darya and Mary regions were of the genus Nummulites (subfamily Nummulitinae) inhabited by a mixed fauna (Paleogen Turkmenii, 1975; survived this boundary and coexisted with rare opercu- Bugrova, 1987, data by L.E. Nevmirich). This fauna lins for a short time in transitional layers. In the other included the species of the Chadumian Assemblage subfamily, Heterostegininae, the genus Grzybowskia (Lenticulina herrmanni, Melonis dosularensis, Cibici- disappeared, species of the genera Spiroclypeus and doides pseudoungerianus, C. expertus, C. oligoceni- Heterostegina became rare, and a new family, Mio- cus, Heterolepa almaensis, Cibicidina amphisyliensis, gypsinidae (two genera), appeared. In the Early Oli- Angulogerina gracilis, and Bolivina mississippiensis) gocene, among orbitoidids, one or two residual disco- and miliolids typical of the Pre-Aral regions (Quinque- cyclinid species still existed; in the upper part of the loculina errmanni, Q. akneriana, Q. subungeriana, P18 or the lower part of the P19 zones, the family Dis- Q. rotunda, Q. romboidea, and Spiroloculina canalicu- cocyclinidae was replaced by the Lepidocyclinidae, lata). In addition, both these regions were inhabited by including the typical Oligocene genus Lepidocyclina the species usual for the TajikÐFerghana Region was represented by a series of subgenera (or, possibly (Brotzenella postacanica, B. multa, B. schurabica, Bia- by separate genera). pertorbis asiaticus, and Caucasina sp. nov.). The Eastern and Southern Mediterranean. Syria may assemblages include at least 15 genera. be considered to be a typical region where the Oli- In the western Kyzyl Kum, foraminifers are repre- gocene fauna of the MediterraneanÐMesopotamian sented by species having wide ranges (Lenticulina her- Basin developed (Krasheninnikov and Ptukhyan, 1973; rmanni, Spiroplectammina oligocenica, Cibicidoides Krasheninnikov and Nemkov, 1975). Here, sandy sedi- aff. pseudoungerianus, Loithostomata macrura, and ments and bioherms were inhabited by abundant num- Unigerinella californica) and by a small number of mulites (five species) and lepidocyclines that were also endemic species (Makarova and Tsatsir, 1964). found in Iraq and Iran (Adams et al., 1983). Lepidocy-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S213 clinids were abundant in the southern coastal areas of At the beginning of the Oligocene, the Armenian Turkey (Krasheninnikov and Ptukhyan, 1973). Basin was still inhabited by Priabonian nummulites (three species), operculines, and discocyclines, as well In a narrow belt in North Africa, nummulites were as by four new species of the genus Nummulites (Grig- reported from Morocco (Ennouchi, in Colloque..., oryan, 1986; Geologicheskie sobytiya..., 1988). In the 1964) and Tunis (two species). In the shallow-water basins of Georgia, scarce nummulites were also found zone of Libya (in carbonate silt), five nummulites spe- (Mrevlishvili, 1978). Local species of this genus inhab- cies, two species of Eulepidina, and Miogypsinoides ited the Rupelian basins in both Transcaucasian were found in association with sea urchins, oysters, and regions, while representatives of other foraminiferal other mollusks (Schaub, 1981; Terminal Eocene Events, groups were absent. 1986). Nummulites reached Somali and Madagascar. Carpathian Basin. At the beginning of the Oli- Eastern Atlantic. Shallow-water carbonate muds in gocene, large foraminifers inhabited the southern shelf the southwestern area of the Aquitaine Basin were area of this basin: Nummulites intermedius, N. vascus, inhabited by small nummulites (four species), opercu- and three other species of this genus were recorded lines, large lepidocyclines, and the genus Miogypsi- from Hungary (Sztrakos, 1971, in Cavelier, 1979). noides, which appeared later (Schaub, 1981; Veillon, Transitional forms between N. fabiani and N. interme- cited by Cavelier, 1979). Lepidocyclina, Miogypsina, dius occurred in Transylvania, and Lepidocyclina and Miogypsinoides (one species in each genus) proba- (Nephrolepidina) and miogypsinides were in Zagorje, bly reached the North European Basin in the terminal Slovenia (Papp, 1954, 1955). Oligocene (in the Chattian) and inhabited northwestern Germany (Van-der-Flerk, 1966; Drooger, in Col- Five species of Late Eocene nummulites were loque..., 1964). recorded from the shallow-water area of the Central Carpathian Cordillera (Slovakia), as well as a form Northern Mediterranean. Northern provinces of transitional to N. intermedius (Samuel and Salaj, 1968). Italy were inhabited by large foraminifers. Their assem- The northern shelf area of the Carpathians (Galizia) blages were more diverse compared to fauna from other was inhabited by six nummulite species, including areas. In the northwestern area (Piedmont), nummulites N. intermedius, N. vascus, and N. fichteli (Bieda, 1963; (three species) occurred; Eulepidina and Nephrolepid- Uhlig, 1886). ina (two species) appeared later and were followed by It is worth mentioning that the Intra-Carpathian Miogypsina and Miogypsinoides (Lorenz, cited by basins and the northern regions of the Mediterranean Cavelier, 1979). Northwest of Verona (Monto Baldo), (Hungary, Romania, Bulgaria, northern Turkey, and nummulites (four species, including N. intermedius and Armenia) were only inhabited by nummulites and rare, N. vascus) and Operculina were found (Cita and Sci- disappearing operculines and discocyclines. Such polo, 1961). In some areas where water was probably impoverishment is most probably associated with the deeper (Possagno and Brendola), nummulites were effects of boreal basins. The presence of species with absent (Herb and Hekel, 1975; Broglio, cited by Cave- narrow geographical ranges (for instance, in Georgia) lier, 1979). The occurrence of N. vascus was reported cannot serve as a guide for recognizing a specific pale- from the Oligocene of southeastern France (French obiogeographic unit. However, nummulitids could Alps). occur there only at the beginning of the Oligocene, Shallow-water areas of southern Spain (Malaga) before the appearance and expansion of lepidocyclinids were inhabited at the beginning of the Oligocene by and miogypsinids. Priabonian discocyclinids and Pellatispira, and, subse- quently, by Lepidocyclina (Molina et al., cited after Cavelier, 1979). Ostracodes The eastern area of the Northern Mediterranean and Terminal EoceneÐEarly Oligocene events consisted the northeastern areas of the Adriatic coasts were in great paleogeographic and climatic changes, rear- inhabited by Nummulites (four species), including rangement of the oceanic circulation, and the formation N. fichtelli, N. vascus vascus, and the Late Eocene spe- of the psychrosphere; consequently, they evidently cies N. germanicus from the northern province (Termi- affected the evolution and distribution of ostracodes. In nal Eocene Events, 1986). In the coastal area of the the Oligocene, the ostracode biota was represented by Dinarides (Albania), nummulites, as well as lepidocy- all its types: continental, fresh-water, brackish-water, clines and Miogypsinoides that appeared later (in the lagoonal, marine shelf, and deepwater, as well as deep- P19 and P20 planktonic zones, respectively) were water fauna of psychrosphere. found (Cavelier, 1979). Nummulites intermedius and Northern Mediterranean. At the beginning of the N. vascus were recorded from the northern coastal area Oligocene, shelf areas of the north Italian basins (the of Turkey (Terminal Eocene Events, 1986). According sections of Priabona, Brendola, and Bressana) were to data by Belmustakov (1959), the Bulgarian Shelf still inhabited by numerous Eocene ostracode species; was inhabited by seven species of Oligocene nummu- however, the taxonomic diversity and abundance of the lites and operculines. assemblages decreased compared to the Eocene

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S214 POPOV et al.

(Barbin and Guernet, 1988). The quantity of ostracode Ducasse and Peypoquet, 1978, 1979, 1986; Ducasse species in the Oligocene was halved (20 compared to and Roussele, 1978, 1979a, 1979b; Peypoquet, 1979; 40 in the Late Eocene). The generic composition also Peypoquet et al., 1981). The changes included a partial became less diverse. In the terminal Eocene, 14 genera, change in the species composition and structural rear- including Eopaijenborchella, Protoargilloecia, and rangement of the populations of certain species of the Phacorhabdotus, which are indicators of depths of the genera Quadracythere, Schuleridea, Cytheretta, Poko- outer shelf, as well as Triebelina and Callistocythere, rnyella, and Hammatocythere. Lagoonal and marine which are indicators of warm water regimes, disap- environments, which first appeared in northern Aquita- peared from the shelf area of northern Italy. The Oli- ine in the terminal Eocene, were dominated in the Oli- gocene basin was probably of colder water and less gocene by Hemicyprideis helvetica and Cyamocytheridea deep-water. The appearance of any new species has not infanta. been reported. North Sea Basin. Shallow-water shelf areas that In the Armenian Gulf in the Oligocene, there was no were situated south of the central part of the North Sea marked change in the ostracode assemblages. Against (Central Graben) and occupied the entire northern area the background of still existing deep-water, totally of Western Europe contained especially abundant ostra- blind fauna, the genus Trachyleberidea disappeared code fauna (Gramann, 1988; Keen, 1989; Uffenorde, and the new genus Buntonia from the family Buntoni- 1989). The ostracode assemblages were extremely idae, the typical inhabitants of the Mediterranean diverse there. Ostracodes typical of the water salinity of Region, appeared. The first appearance of sighted ostra- 17–32‰ inhabited the Isle of Wight in the Late Hem- codes (genus Costa, also of Mediterranean origin) is stead during the accumulation of Sannois Limestones assigned to the formation of the Globigerina selii Zone; in the Paris Basin and beds with Nucula comta in the this likely indicates shoaling of the basin in the Oli- Belgian area of the shelf zone, as well as the Rhine Gra- gocene. ben in the first half of the Rupelian (Keij, 1957; Goer- lich, 1958). Eastern Atlantic. The terminal EoceneÐEarly Oli- gocene events are traced differently in different bathy- The Rupelian ostracode assemblages are character- metric zones. In the lower bathyal and abyssal zones in ized by a mixture of brackish-water and polyhaline the northern area of the Bay of Biscay, representatives groups. The first group includes the genera Neocypri- of deep-water, so-called psychrosperic cold-water deis, Hemicyprideis, Cytheromorpha, Cytheridea, ostracode fauna (genera Poseidonamicus, Bradleya, Cyamocytheridea, and Loxoconeha. The second group and Agrenocythere) appeared in the eurythermal ostra- comprises the genera Cytherella, Cytherelloidea, code assemblages since the terminal Eocene. Such Cytheretta, Cytherura, Echinocythereis, Konarocythere, composition of assemblages was mainly associated Leguminocythereis, and Pontocythere. In the Paris with oceanic currents and the formation of a two-lay- Basin, the proportion of marine groups was much ered structure of water masses, with the lower layer higher. The genera Hammatocythere, Pokornyella, being formed by cold Arctic waters and the upper layer Trachyleberidea, and Pterygocythereis predominated being formed by Mediterranean waters. North of the the shelf zone of the Armorica Massif and the north Bay of Biscay (Rockall Plateau), ostracodes were rep- German shelf area. The generic composition indicates resented in the Early Oligocene by the genera Cyther- moderately warm water temperature regimes. The ella, Bairdia, Bythoceratina, Argilloecia, Atlanti- Rupelian ostracode assemblage was generally charac- cythere, Trachyleberidea, and Cardobairdia in associa- terized by a low proportion of endemic forms. Many tion with the newly appearing psychrospheric genera species, such as Paracyprideis rarefistulosa, Legumi- Poseidonamicus and Bradleya. In the lower bathyal nocythereis scrobiculata, Echinocythereis hispida, zone, in the southeastern area of the Bay of Biscay, the Cuneocythere marginata, Cyamocytheridea punctatella, majority of the ostracode fauna was composed in the and Hemicypideis helvetica, that first appeared in the terminal Eocene and, beginning with the Oligocene of Oligocene, had very wide geographical ranges. the genera Cytherella and Argilloecia, the numbers of Ostracodes tolerant of high water salinity appeared slowly evolving groups. The assemblage was generally in the Belgian shelf area during the formation of Boom uniform. Clays. This assemblage is characterized by a small pro- In the littoral zone that occupied a large part of portion of early transitional species and by the first Aquitaine, the evolution of the ostracode fauna tolerant appearance of Henryhowella. In the deeper-water areas with respect to salinity was normal. The Oligocene of the north German and Danish shelf zones, shallow- ostracode assemblage in the Aquitaine Basin included water groups are absent. The ostracode assemblage 58 ostracode species of 36 genera, with 44 species and is composed of the genera Argilloecia, Krithe, 32 genera descending from the Late Eocene. Four gen- Cytheropteron, and Henryhowella; however, no psy- era did not continue into the Oligocene. Seven genera, chrospheric groups have been registered. Neocyprideis, Bythoceratina, Trieblina, Costa, Callis- As a result of the Rupelian sea transgression in the tocythere, Hemicyprideis, and Eucythere newly appeared North Sea Basin, marine waters reached lagoons and (Deltel, 1962; Ducasse, 1969, 1974, 1975, 1981, 1983; intracontinental water basins; ostracodes penetrated the

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S215 neighboring zones, and a large number of mixed ostra- disappearance of this group by the final stage of the for- code faunas appeared. At the beginning of the Rupe- mation of the lower beds of the Tard Clays. lian, fresh-water and brackish-water ostracodes of the lake and lagoonal types inhabited Isle of Wight, the Ostracodes that appeared during the accumulation Paris Basin, and Rhine Graben (Middle Hemstead, of beds enclosing Cardium lipoldi composed an Middle Sannois, and Middle Pechelbronner assem- entirely new polyhaline assemblage of shallow-water blages). These assemblages included species of the and deepwater forms (Monostori, 1985), accompanied transitional genera Cypris, Ilyocypris, Eucypris, and by ecologically tolerant fresh-water and brackish-water Cypridopsis; representatives of Moenocypris and Vir- groups. This assemblage included the genera Cyther- gatocypris, the genera that first appeared in the Late etta, Schuleridea, Pterygocythereis, Megahemicythere, Eocene; and one species of the Oligocene genus Lineo- Leguminocythereis, Echinocythereis, and Cytheridea. cypris. The taxonomic composition of the assemblage indi- cates a shallow basin with wide paleogeographical con- AlpineÐCarpathian Basin. The Oligocene evolu- nections. This basin could periodically experience tion of ostracodes in the Bavarian and Swiss molassa marine invasions. The formation of the upper beds of depressions displaced certain unique features. The Tard Clays and the Lower Kiscell Clays in the Rupelian Rupelian assemblage included 43 species in the Swiss occurred in the environments unfavorable for ostra- Depression and at most 10 species in the Bavarian codes, possibly also in anoxic environments. Depression (Oertli, 1956; Witt, 1967). Polyhaline assemblages of the infralittoral zone (20Ð80 m) had In the Transylvanian sShelf, the change from the rather narrow geographical ranges and inhabited Late Eocene to the Oligocene assemblages was gradual clayey-marly facies, where the occasionally formed (Olteanu, 1980). The generic composition experienced mass accumulations. The shallow-water species Hemi- only slight changes (above the horizon containing Pyc- cyprideis helvetica, Cyamocytheridea punctatella, and nodonte gigantica); the genus Argilloecta disappeared; Cytheridea pernota were widespread, whereas inhabit- however, the species composition was nearly halved ants of the deep shelf zone, the genera Henryhowella, (67 and 35 species in the terminal part of the Breb Argilloecta, etc., were absent. Marls). At the same time, certain species, mainly from the family Cytherideidae, increased in number. Subse- In the Western Carpathians (Moravia), in the Early quently (during the formation of the Hoja Marls), no Oligocene that corresponds to the initial stage of the marked changes in the quantity and quality of the ostra- formation of the Menilithic Beds, deepwater ostracode code composition were registered (27 species of fauna still occurred, the genus Krithe predominated, closely similar taxonomic composition). The assem- whereas the genera Abyssocypris and Agrenocythere blage became more diverse in the Middle Rupelian were rare. The morphology of the representatives of the (Mera Beds); the number of species increased to 47. genus Krithe indicates rather good aeration of bottom Mediterranean groups appeared, including the genera waters, whereas the increase in the size of species that Triebelina, Costa, and Callistocythere. The assemblage came from the Eocene indicates cooling (Pokorny, also included the forms that were widespread in the 1980). However, the temperature of water in the West- Rupelian of northern Europe, i.e., Schuleridea perfo- ern Carpathians in the Oligocene was still much higher rata, Cytheropteron gulincki, Echinocythereis ex gr. than that of the oceanic water of the psychrosphere. scabra, and Leguminocythereis scrobiculata. The Rupelian ostracode fauna of the Hungarian In the Late EoceneÐEarly Oligocene, the composi- Basin was clearly associated with the structure of the tion of the ostracode assemblages only sligtly changed basin, the increase in sea depths from west to east and in the percentage of eurybathic and shallow-water shelf the marked fluctuations of the sea level. At the begin- polyhaline groups (approximately 25Ð20% and 75Ð80%, ning of the Oligocene, the structure of the basin was respectively). Along with the marine assemblages, spe- almost the same as in the Eocene (Monostori, 1985, cies of the brackish-water groups Hemicyprideis hel- 1986). The final stage of the Buda Marls formation and vetica and Cytheridea pernota (Curtuius^ Formation) the initial stage of the Tard Clays formation coincided were abundant in some brackish-water areas of the with the major deepening of the eastern area of this Early Rupelian basin; this indicates the effects of river basin. The ostracode assemblages included the deepwa- inflows. ter genera Henryhowella, Agrenocythere, Argilloecia, Cytherella, Cardobairdia, and Parakrithe in associa- A sharp impoverishment of the polyhaline ostracode tion with epinerithic groups (Cuneocythere marginata) assemblages was associated with the basin brackishing and the fresh-water genera Candona, Cypridopsis, and in the Middle Rupelian (during accumulation of the Moenocypris, which were brought there by the turbid Bizusa Marls, and especially of Ileanda Clays). The flows. A decrease in the oxygen content dissolved in number of species in the assemblage decreased to 7Ð10, water that was observed since the terminal Eocene and including euryhaline Cytheridea ventricosa, Bosque- continued in the Rupelian affected the ostracode assem- tin sp., Cyamocytheridea bouendensis, and “Thracella blages and resulted in a marked decrease in abundance apostolescui”, as well as the fresh-water species Can- and taxonomic diversity of ostracodes, and in subsequent dona sp. and Ilyocypris sp.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S216 POPOV et al.

Greater CaucasusÐKopet Dagh Basin. Ostra- formation, ostracode assemblages were especially codes inhabited the shallow shelf area of this basin and abundant and represented by a large number of brack- have not been found in its deepwater area, in typical ish-water taxa (the Polbian, Molochanian, Solenovian, Maikopian facies. Ostracodes from the shelf zone of and the Middle Daryalyk assemblages of shallow areas Eastern Paratethys markedly differ from the Rupelian of the basin). Disopontocypris oligocaenica, D. majko- ostracode fauna of Western Paratethys. piensis, Cyprinotus medianus, and Haplocytheridea (1) The assemblage was uniform throughout the solenica occurred throughout the basin. The absence of area of its distribution (the Rubanovka, Kyzyldzhar, ostracodes in deepwater areas of the basin (the section Ascheairyk, Uzunbas, Kujulus), and Lower Sarybatyr of the Kheu River in Ciscaucasia and the Caspian assemblages, as well as the assemblages of Lower Region) is typical. The ranges of the Solenovian ostra- Rupelian ostracodes from the South Aral Region and code biota became indistinct in the eastern area of the Central Turkmenistan only slightly differed from each basin. The South Aral Region was the eastern boundary other (Gramm, 1963; Sheremeta, 1969; Nikolaeva, 1985). of the genus Disopontocypris and Cyprinotus medianus and Haplocytheridea solenica. At the final stage, the (2) In these assemblages, over 30 euryhaline genera Solenovian Basin was filled by noncarbonate Maiko- (including Bairdia, Argilloecia, Xestolebers, Eopaijen- pian-type clayey beds that formed unfavorable condi- borchella, etc.) that inhabited the Western Paratethyan tions for ostracodes. The North Ustyurt Plateau was the Region from the Eocene to the beginning of the Rupe- sole region where a relatively numerous ostracode lian, and later, were absent. assemblage was found. The species diversity was rather (3) The oceanic genus Henryhowella and the Medi- low, and the assemblage was formed by transitional terranean genera Costa, Callistocythere, and Trie- marine species. Of the brackish-water ostracodes, only belina, which are rather typical for Western Paratethys, Disopontocypris kasachstanica was present. The first were absent from the Pshekhian Basin. The Pshekhian representative of Paracyprideis, P. punctata, appeared Assemblage included 20Ð25 species of 16 mainly poly- that differs this assemblage from the Early Solenovian haline genera, with a predominance of cythereidids and Assemblage. The Solenovian Basin was inhabited by trachyleberidids of the subfamily Echinocythereidinae. over 30 species of 17 ostracode genera. Cuneocythere marginata, Cytheropteron steinmanni and species of the genus Leguminocythereis ex gr. stri- atopunctata were found everywhere. The decrease in Mollusks the proportion of stenohaline groups from west to east (the absence of the genus Krithe in the Ascheairyk North Africa. Available data on the Early Oli- Assemblage) was registered. gocene molluskan fauna from North Africa are very Solenovian Basin. In the Solenovian Time, a scarce, unlike those on the Priabonian mollusks. The unique ostracode assemblage was formed. It comprised impoverished assemblages recorded in Algeria (Cuvil- elements of inherited marine fauna and the fauna that lier, 1930; Flandrin, 1938) indicate that the African appeared in shallow brackish-water basin. Among coast had lost its peculiarity by the onset of the Oli- Solenovian marine ostracodes, there were Cytheridea, gocene. All the 13 bivalve genera reported were wide- Cuneocythere, Cyamocytheridea, Eucytheridea, Hap- spread in Europe, almost a half of the species being locytheridea, Schuleridea, Pterygocythereis, Echi- known from southern Europe. The assemblage nocythereis, Bosquetina, and Cytheropteron. The spe- included typical Mediterranean taxa (Chlamys biar- cies of these genera (that had wide geographical ranges ritzensis, Pecten arcuatus, and Pycnodonte brongn- in the majority of cases) composed over 75% of taxa of iarti) that were abundant in Europe from the Eocene, the Solenovian Assemblage. However, regarding the but were absent in the Eocene of Egypt. abundance, certain brackish-water species of the gen- Northern Mediterranean. Representative assem- era Disopontocypris, Cyprinotus, and Haplocytheridea blages of the Rupelian molluskan fauna of the southern predominated. The type species Disopontocypris oligo- type were recorded from the French Alps, northern caenica of the genus Disopontocypris was widespread Italy, Transylvania, southern Bulgaria, and Armenia in the lower part of the Solenovian Horizon and in the (Popov, 1995; Amitrov, 1996; Fig. 3). upper part of the Rupelian in the North Sea Basin and in the Western Paratethys (Veselov and Sheremeta, These data indicate that molluskan fauna from the 1966; Popov et al., 1985; Jiricek and Riha, 1991). Other coastal area of the MediterraneanÐMesopotamian brackish-water species of ostracodes were similar in Basin was mainly inherited from the Eocene and distribution. Thus, Moenocypris olmensis found in the included numerous warm-water tropicalÐsubtropical Polbian Assemblage was common in the Middle Pech- elements (Trisidos, Crassatella sensu stricto, Cardium elbronner of the Mainz Basin. Along with marine and sensu stricto, Vepricardium, Corculum, strombids, and brackish-water genera, the species of the widely eury- diastomas), as well as many typical Bartonian and Pri- biont genera Loxoconcha and Cytheromorpha were abonian genera and species that retained their ranges found in the Solenovian Basin; however, they were not unchanged and were widespread in the Rupelian. The pro- numerous. At the initial stage of the Solenovian Basin portion of the newly appearing species is less than 30%.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S217

50° 40 Kazakhstanian Land Russo-Scandia 31 Germania Province 39 30 33 36 32 N O 29 R T H 38 18 E U R O 41 42 20 P E A 19 N R 37 40° 25 E G I O 24 N 21 22 35 c e 44 34 a n P r o v i n 23 K h a d u m i 43 9 26 27 14 11 28 10 12 15

30° 13 17 8 Anatolia ANCIE NT M 16 Iran-Afghanistan Land EDITE 7 1 RRAN 2 EAN REGIO 3 N 6

6 5 11 Arabia 1 7 12 2 8 4 3 13 4 9 14 Africa 5 10 15

Fig. 3. Complex benthos-based zoogeographic zonation of the shelf of the Rupelian basins. Explanations: (1) North European Region with the presence of moderately warm-water benthic fauna of the Rupelian type: (2) molluskan assemblages, including Arc- tica, Cyrtodaria, Scalaricardita, and lacking the most warm-water taxa; (3) richer transitive molluskan assemblages, including the warm-water taxa, but lacking the typical Tethyan species; (4) impoverished assemblages of large foraminifers with Nummulites, Operculina, and Discocyclina; (5Ð6) ostracode assemblages: (5) assemblages of smooth-walled groups descended from the Priabon- ian, including new “psychrospheric” genera Poseidonamicus, Brodleya, and Agrenocythere; (6) assemblages with high proportion of euryhaline and brackish-water groups; (7Ð8) assemblages of small foraminifers: (7) of the northern type; (8) of the transitional (mixed) type; (9) boundary zones; (10) Ancient Mediterranean Region with the benthic fauna including various assemblages of colonial corals, nummulitids, and benthic foraminifers with large sculptured tests: (11) assemblages of the Mediterranean ostracode genera with typical sculptured forms and with the presence of Buntoniidae of the Mediterranean origin; (12) assemblages of small foraminifers with Pararotalia, Herondenia, and Queraltina; (13) reef-building corals; (14) assemblages of large foraminifers with nummulitids, lepidocyclines, and miogypsinids; (15) molluskan assemblages of the southern type with tropical taxa.The main local- ities of the benthic fauna from the Rupelian: (1) Morocco; (2) Algeria; (3) Tunis; (4) Libya; (5) Egypt; (6) Syria; (7) Iraq; (8) southern Spain; (9) Aquitaine; (10) Coastal (Nummulite) Alps; (11) northern Italy; (12) Slovenia; (13) Albania; (14) southern Bulgaria; (15) northwestern Turkey; (16) southern Turkey; (17) Armenia; (18) southern England; (19) central France; (20) Mainz Basin; (21) Switzerland; (22) Austria; (23) Hungary; (24) Slovakia; (25) Ukrainian Carpathians; (26) Transylvania; (27) western Georgia; (28) southern Georgia (Akhaltsykhe); (29) Belgium; (30) the Netherlands; (31) Denmark; (32) northern Germany; (33) Poland; (34) Crimea; (35) Northern Caucasus (Kuban River); (36) Ergeni; (37) Mangyshlak; (38) northern Ustyurt; (39) Pre-Aral Coastal Area; (40) southern part of Western Siberia (Kurgan Beds); (41) Southern Aral Coastal Area; (42) Kysyl Kum; (43) Kopet Dagh; (44) Tajikistan, Ferghana.

Nevertheless, the available data on bivalves indicate (Popov 1994, 1995). This was probably caused by dif- substantial changes as compared to the Late Eocene. ferential extinction in different areas of the basin. How- (1) Both the generic and species compositions of ever, it is not inconceivable that the compared assem- molluskan faunas were impoverished (41 genera were blages are unrepresentative. found in northern Italy as opposed to 59 Priabonian (3) The species of north European origin, such as genera, and 24 genera were recorded in Bulgaria as Barbatia nysti, Anadara sulcicosta, Glycymeris lunu- opposed to 63 Priabonian genera). lata, G. obovata, Astarte kickxi, Nemocardium (2) The similarity between the assemblages decreased, tenuisulcatum, Glossus subtransversus, and Pelecyora and the fauna lost its integrity; the comparison of the westendorpi, appeared in all south European assem- generic compositions of bivalves indicates that the Pre- blages. ston’s coefficients of dissimilarity increased from 0.40– The gastropod composition also indicates very 0.69 for the Priabonian to 0.64Ð0.91 for the Rupelian warm-water environments in the Northern Mediterra-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S218 POPOV et al. nean. Judging from the gastropod faunas, it was not luskan assemblage (from the Etampes Sands) supports colder in this region in the Rupelian than in the Pria- this fact. Mediterranean species and genera were absent bonian. Much more species (580 versus 190) and fam- in this fauna. Endemic species constituted a large part ilies (63 versus 51) were found in the Rupelian of of the assemblage; the other species were widespread northern Italy compared to the Priabonian. The quantity and the same as in the Rupelian of northern basins. The of the most warm-water families also increased; strom- bivalve assemblages were almost as diverse as in the bids, diastomatids, etc., were accompanied by tonnids, Aquitaine and included 112 species (Cossmann and vasids, and harpids. The Early Oligocene molluskan Lambert, 1884; Furon and Soyer, 1947); however, the assemblage from the Yerevan Gulf (Shorakbyur Forma- warm-water groups decreased in diversity. They tion in Armenia) is also more diverse than the Late included 33 species in common with those from the Eocene assemblage (from the Rind Horizon, Late Mainz Basin and only 10 species in common with Bel- Eocene). Only assemblages from Bulgaria and the gium; this indicated the direct faunal connections with French Alps decreased in diversity. The number of new the Rhine Basin via Alsace (Saubade, 1970). gastropod species is substantially greater than those of Strombids, Velates, and some other warm-water bivalves; the proportion of gastropod species that are gastropod groups (neritopsids and harpids) were absent unknown in the Eocene is 70% (mainly at the expense from the Stampian of the Paris Basin, although diasto- of the large North Italian Assemblage). Of 180 species matids were present. For the Oligocene, this assem- in common with the Eocene, 36 were recorded from the blage may be considered to be a standard transitional Middle Eocene and were not registered in the Priabon- assemblage. This assemblage is clearly more cold ian. The similarity of the Rupelian and Priabonian gas- water than the transitional Eocene assemblages (for tropod assemblages did not decrease, unlike that of the example, the Mandrikovka Assemblage and the Barto- bivalve assemblages. nian assemblage from the Paris Basin); in the family Similar to bivalves (although not so clearly), the and generic compositions, it is similar to the Eocene ranges of certain gastropod species decreased, and assemblages of the northern type (Latdorfian Assem- some species possibly migrated from the north to the blage, etc.). The quantity of species in common with south. Of species known from both the Priabonian and the Eocene assemblages is rather small, 44 of 206 the Rupelian, 98 species were only found in southern (21.3%). areas, 55 species were only recorded in northern assem- The molluskan composition indicates that there blages, nine species occurred everywhere in the Pria- existed a faunal exchange between the Aquitaine and bonian and only in southern areas in the Rupelian, and the Paris basins via the Atlantic; however, it was com- 16 species were found in northern assemblages in the plicated by a climatic barrier. This is indicated by the Priabonian and only in southern assemblages in the small number of species in common and by the transi- Rupelian. tional pattern of the fauna found in Brittany (Renn). Eastern Atlantic. Molluskan assemblages from the As distinct from the Paris Basin in the southern area Aquitaine shelf of Atlantic were very rich and diverse of England, the molluskan habitats became unfavorable and included 140 bivalve species and 200 gastropod in the Early Oligocene compared to the Late Eocene. species (Stampian of Aquitaine after Cossmann, 1921; Only 24 bivalve species were found in the Hemstead Saubade, 1970). Although this fauna included warm- Beds of the Hempshire Basin. This is mainly endemic water genera and species in common with the Mediter- fauna with fresh-water taxa. Some marine species are in ranean, this assemblage is rather unique; the Stampian common with those from the Paris Basin. The Hem- molluskan assemblages were dominated by the species stead Gastropod Assemblage includes 56 species of that were first discovered there. 19 marine families in addition to fresh-water species. Similar to the situation in bivalves, the Rupelian In the family and species compositions, this assem- gastropod assemblage from Aquitaine was more blage is similar to the northern Oligocene type. It virtu- diverse compared to that from the Priabonian (about ally lacks moderately psychrotolerant groups (ampull- 200 species versus 60 and 62 families versus 30). inids are the most warm-water forms of Hemstead gas- Strombids were absent in the Priabonian of Aquitaine. tropods). Of 56 gastropod species from the Hemstead The Rupelian assemblage in this region was of the Beds, only 18 were reported from other Rupelian southern type and included two strombide species and assemblages; 18 species are in common with those representatives of other warm-water families. The from the Eocene, with 8 of them being found only in the absolute quantity of the species in common (34) and the Headon Beds in the Eocene of England. dissimilarity estimated by the Preston’s coefficient North Sea Basin. Of molluskan assemblages from indicate that the Rupelian assemblage from Aquitaine this region, the assemblage from sandy facies of the is most similar to that from Italy. Mainz Basin is the most abundant (Sandberger, 1856Ð In the Paris Basin, normal marine environments 1863; Neuffer, 1973; Kuster-Wendenburg, 1973, 1982; were restored after a brackishing in the Ludian Time Gürs, 1995; Fig. 3, locality 20). It includes 128 bivalve (Late Eocene) and in the Sannois Time (beginning of species and 212 gastropod species (hereinafter, conti- the Rupelian). The composition of the Stampian mol- nental groups that are occasionally described in associ-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S219 ation with marine taxa are not considered). A recent coefficient for the Mainz Basin and Belgium–the Neth- revision by Gürs has markedly extended the list of mol- erlands (0.65) is almost as low as that for the Mainz and lusks from this assemblage, especially of gastropods. In Paris basins. The assemblages of the northern areas addition to new species, this list today includes the gen- under consideration differ more markedly from those of era and families previously unknown there. Naturally, the Paris Basin than from those of the Mainz Basin; this revision somewhat changed the values of the Pre- however, several common species absent from the ston’s coefficients; nevertheless, this supported rather Mainz Basin, were registered in the Paris Basin and the than changed previous conclusions (Amitrov, 1993, northern areas (Amitrov, 1996). This indicates the 1996). probable existence of an occasional direct connection The Mainz Basin had the maximum species in com- between the Paris Basin and the North Sea Basin (in a mon with the Paris Basin (93, the Preston’s coefficient narrow sense), not via the Mainz Basin. is 0.64; Fig. 3, locality 19). The family and generic The Alsace Region generally may be assigned to the compositions indicate that these two basins had similar North Sea Basin. Only 14 gastropod species were deter- temperature regimes. As distinct from the Paris Basin, mined there with certainty. Alsace is situated south of in the Mainz Basin diastomas were absent, but there the Mainz Basin; however, its assemblage is closer to were many groups that were absent in both the Eocene the northern assemblages based on both the family assemblages and the Oligocene northern assemblages (only psychrotolerant groups) and the species composi- (cocculinids, littorinids, omalogyrids, vasids, philinids, tion (12 species in common with Belgium and the atyids, umbraculids, siphonariids, and juliids), as well Netherlands, the Preston’s coefficient is 0.57). as the groups that are known from northern Eocene The proportion of gastropods that were inherited assemblages, but are absent or very rare in northern Oli- from the Eocene fauna is about 30% in the Mainz gocene assemblages (pleurotomariids, patellids, tur- Basin; 55% in Belgium and the Netherlands; 37% in binids, lacunids, cypraeids, etc.). Denmark, northern Germany, and Poland; and about 57% in Alsace. In more northern regions, certain distinctions in the A representative molluskan assemblage is known patterns of bivalve and gastropod assemblages are from the Fore-Alpine Depression, namely, from Swit- observed. The bivalve assemblage from the Rupelian of zerland, Bern Jurassic (Kissling, 1896). In the Early Belgium (Berg Sands, clay with Nucula comta and Oligocene, this depression was a strait, which con- Boom Clays) is rather unique. About 60 bivalve species nected Western Mediterranean and the Carpathian were recorded there (Gilbert and Heinzelin, 1954). Basin. Although the assemblage was composed of Warm-water elements of this fauna are much more rather warm-water groups, it almost lacked typical diverse than those of other molluskan assemblages Mediterranean genera and species. It was similar to the from the Lower Oligocene of the North Sea Basin. Gly- cymeris (Tucetilla), Claidornites, Arcopagia, Isogno- fauna from the Mainz Basin (of 55 bivalve and 29 gas- tropod species, 41 and 23 are in common, the Preston’s mon, and Spondylus showing certain similarity coefficient is 0.65); this indicates direct connections between this fauna and the faunas from the Mainz between these basins, probably via a system of Rhine Basin and Switzerland were found only there. This grabens. This is a transitional assemblage, like the diversity may be determined by the shallow-water Mainz Assemblage. As concerns of gastropods, it facies that were only known in Belgium or by the prox- includes ampullinids, patellids, and vermetids. Only imity of a strait that flew through the Rhine Graben, three species are in common with those from the neigh- Mainz Basin, and Alsace (Saubade, 1970, fig. 3). boring region of the French Alps. It was mentioned In more northern area of the basin (in the Nether- above that the assemblage from the French Alps is lands, Denmark, northern Germany, and Poland), a uni- assigned to the southern type and we consider this form fauna (about 40 bivalve species) typical of the region as a component of the Northern Mediterranean Rupelian clay facies was found. This assemblage was Region. dominated by paleotaxodonts, thyasirids, astartes, and In the Rupelian of Bavaria, turritellids are the most pectinids. However, any true comparison is impossible, warm-water gastropod group, and this assemblage may because these features were primarily determined by be only conventionally considered to be transitional. Its facial conditions in which the sediments accumulated species composition is most similar to those of the and bivalves occurred. Mainz Assemblage (16 species in common, the Pre- All the gastropod assemblages in northern areas, ston’s coefficient is 0.68) and the Belgian–Netherlands from Belgium and the Netherlands to Denmark, north- Assemblage (17 species in common, the Preston’s coef- ern Germany and Poland, are closely similar to each ficient is 0.53). other, assigned to the northern type, and have similar The assemblage from the Swiss Alps includes distinctions from the Mainz Assemblage. The above- 15 species in common with the Priabonian Assemblage listed relatively warm-water families that are known (52%). from the Mainz Basin are absent there. At the same Carpathian Basin. In the Czech and Polish Car- time, their species composition is closely similar to that pathians, the Lower Rupelian deepwater facies do not of the Mainz Assemblage. The Preston’s dissimilarity yield mollusks or include very impoverished assem-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S220 POPOV et al. blages that are very difficult to compare. The shallow- northern and eastern shelf areas (from southern water assemblage from the Pokut (Ukrainian) Car- Ukraine, Crimea, northern Caucasus, Ergeni Highland, pathians was transported to deepwater facies by turbid and the entire Transcaspian) in water temperature flows and still requires a revision. However, prelimi- regimes. Among the southern shelf assemblages, the nary data indicate that this assemblage clearly belongs assemblage from southern Georgia is most diverse to the North European Faunal type (Studencka et al., (Akhaltsikhe, about 100 bivalve species). The species 1999) based on its generic composition (the presence of of Tethyan origin are very rare there (Modiolus rutim- Astarte and Scalaricardita and the absence of specific eyeri, Pecten arcuatus, and Chlamys meissonieri), with Mediterranean taxa). At the same time, the presence of all of them probably from the Eocene. Only eight spe- such gastropod families as the Turritellidae, Volutidae, cies are in common with the Armenian Assemblage. Conidae, and Marginellidae indicates more warm- Their presence does not support the existence of any water regimes in the North Carpathian shelf than in the direct connections between Transcaucasia and Tethys. northern shelf of the Eastern Paratethys. The marked impoverishment of fauna indicates that In the southern shelf of the Carpathian Basin, an these connections could disappear as early as the Mid- abundant molluskan fauna was only found in Transyl- dle Priabonian. Approximately a half of the bivalve vania (Lower Rupelian Hoja and Mera beds). The species was in common with those from more northern bivalve assemblage is not very rich but rather represen- assemblages of southern Ukraine, Mangyshlak, and tative. Although the content of the warm-water ele- Ustyurt. ments is relatively low, this assemblage is characterized Of gastropods from relatively warm-water groups by the presence of genera and species specific for the that were absent in northern Oligocene assemblages, Mediterranean Region (Chlamys biarritzensis, Spondy- diastomas, melongenids, and large ampullinids were lus cisalpinus, Macrosolen hollowaysi, etc.). The gas- found in Georgia. To characterize the species composi- tropod assemblage is rather large, including 117 spe- tion, it is of interest to compare the dissimilarities of the cies of 41 families. This is assigned to the transitional assemblages from Georgia and other regions of western type; the warm-water groups include only diastomas. Eurasia in the Rupelian and Priabonian. The similarity Recall that the Priabonian assemblage from Transylva- of the Georgian and southern assemblages (from Italy, nia included both strombids and Velates and was Bulgaria, and Armenia) evidently decreased with time, assigned to standard southern assemblages. Despite i.e., the Preston’s coefficients increased from 0.77–0.84 this transition from the southern to the intermediate in the Priabonian to 0.82Ð0.93 in the Rupelian. At the type, in the Rupelian, southern assemblages (northern same time, the similarity between the Georgian and Italy, southern Bulgaria, and Armenia) were still most northern assemblages (from BelgiumÐNetherlands, similar to the Transylvanian assemblage in species northern Germany, and Transcaspian) increased; the composition. This indicates the persistence of direct Preston’s coefficients decreased from 0.91–0.93 to connections between the Transylvanian shelf and the 0.82Ð0.84 (Amitrov, 1996). However, in gastropods, Mediterranean Region at the beginning of the Oligocene. this tendency is not as clear as in bivalves; in particular, Nevertheless, the Preston’s coefficients increased from in the Rupelian, the similarity to the Armenian Assem- 0.71Ð0.75 to 0.83Ð0.86. Eighty-six gastropod species blage remained high. Unlike bivalves, there were gas- in common are reported for the Rupelian and Eocene of tropod species there unknown in the Eocene and found Transylvania (in the Priabonian, 11 of them were only in the Rupelian of Armenia, Georgia, and Azer- absent), i.e., 63% species in the assemblage were in baijan (Benoistia korobkovi Zotova), or in Armenia and common and 37% were not known in the Eocene. The Azerbaijan (Melongena azerbaidjanica Alizade et Bag- proportion of bivalve species that first appeared in the manov). A small assemblage from Azerbaijan (village Oligocene is only 14%. of Mirlyar, Kubatly District) resembles the Georgian At the end of the Rupelian, the Carpathian Basin, as Assemblage. Gastropod fauna, as benthic foraminiferal the entire Paratethys, was of a closed or semiclosed assemblages, indicates that at the beginning of the Oli- type, inhabited by endemic brackish-water fauna gocene, an indirect connection could exist in Transcau- (Baldi, 1986; Rusu, 1988). This molluskan assemblage casia between the Pshekhian and the Mediterranean is the richest in Transylvania, where Janschinella basins, possibly, via the Armenian Gulf. In the Rupe- garetzkii, Ergenica sp., Urbnisia lata, Merklinicardium lian of Georgia, 41% of species in common with the apostolovense, Korobkoviella sp., Cerastoderma serog- Eocene species were found. osicum, and C. lipoldi (determination by Popov from Pshekhian mollusks from the northern shelf area the material collected by Rusu) were found (Popov, (southern Ukraine, Crimea, Ciscaucasia, VolgaÐDon 1995). In Hungary, Janschinella sp., Urbnisia sp., and Region, Mangyshlak, and Ustyurt) are rather diverse ?Cerastoderma lipoldi were found (Baldi, 1986). (about 130 bivalve species of 80 genera and 60 gastro- CaucasusÐKopet Dagh (Pshekhian) Basin. In the pod species) and similar to each other, despite an first half of the Early Oligocene, normal marine impoverishment of some assemblages. The molluskan regimes existed in the basin. The molluskan assem- assemblages from these regions (even if considered to blages in the southern shelf area (recorded from Geor- be an integral fauna) are more markedly impoverished gia and Azerbaijan) markedly differed from those in than the more northern Eocene assemblages (for exam-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S221 ple, the Chegan Assemblage) and the Oligocene assem- An increase in the dissimilarity is very natural, blages from the northern areas of Western Europe. Con- nearly all the species in common with a more distant cerning gastropods, not only tibiids, melongenids, mar- assemblage are also present in a closer located assem- ginellids, and turritellids, but also trochids, xenophorids, blage. olivids, volutids, conids, etc., which were found in A molluskan assemblage of the same (northern) some Rupelian assemblages of northern Europe are type inhabited the Fore-Kopet Dagh Gulf of the Tura- absent. A good preservation of fauna (for example, in nian Sea; 33 bivalve species and 15 gastropod species the Uzunbas Formation in Mangyshlak) indicates that were recorded in the upper part of the Torymbeur For- these assemblages are complete; i.e., the impoverish- mation (Lower Oligocene), Western Kopet Dagh (data ment is characteristic of the assemblage rather than a by Amitrov; Popov et al., 1993). result of secondary changes. Despite the low proportion of warm-water elements (the proportion of subtropical Thus, the comparison of molluskan distribution in taxa was at most 15Ð20%), this fauna was similar to the Early Oligocene and Late Eocene clearly indicates that from the Transcaucasian shelf in the species com- the impoverishment of assemblages and the appearance position (about 40% of species and 70% of genera in of new species; this is especially well pronounced in the common). Outside the Eastern Paratethys, this fauna northern regions. An increase in the temperature gradient was most similar to the assemblages from the North is especially well traced in the gastropod distribution. Sea Basin. The ranges of many species were restricted CaucasusÐKopet Dagh (Solenovian) Basin. In the to the Eastern Paratethys (about 20% of this fauna) a second half of the Early Oligocene, numerous bivalve large part of these species were dominants and contrib- species and genera endemic to the Paratethys appeared uted to the general pattern of the assemblage (Pterolu- in Paratethyan lagoons in the Late Pshekhian after the cina batalpaschinica, Yoldiella chadumica, etc.). closure and brackishing. Among these forms, there Of 60 gastropod species, 34 (57%) are in common were Ergenica, Urbnisia, Janschinella, Corbula (Len- with the Eocene fauna, although in some regions, the ticorbula), and some species of Cerastoderma. At the proportion is higher, i.e., ten species of 13 in the north- same time, gastropods of this basin were mainly the ern Caucasus and 20 species of 25 in the Crimea. lagoonal forms that had wide geographic and strati- The composition of molluskan assemblages from graphic ranges (Popov et al., 1985). the platform part of Ukraine deserves special consider- The integrity of the molluskan fauna of the entire ation. More warm-water Eocene taxa were probably Paratethys, as well as the distinctions in the composi- included into the list of assemblages from the Nikopol tion of the western, northern, and southern shelf areas, Formation of the South Ukrainian Depression. The sit- are more prominent in the taxonomic composition of uation is even more complicated in the Zhukovtsy and the impoverished brackish-waterÐlagoonal Early Sole- Bishki Beds of the DnieperÐDonets Depression. Some novian Molluskan Fauna (18 bivalve species and researchers deny their Oligocene age. The poor preser- 22 gastropod species) than in that of the marine fauna. vation of mollusk fossils does not allow any justified The majority of molluskan genera were in common, but assumption. However, even if the presently known the species composition was much more diverse in assemblages do not belong to the Rupelian, the analysis Transcaucasia. The species composition of mollusks of the molluskan distribution in adjacent regions indi- from the AlpineÐCarpathian Basin has not yet received rectly but strongly supports the connection between the a detailed study. At present, nearly all the taxa identi- sea basins in the southern areas of the former USSR and fied there are in common with those from Eastern the North Sea Basin in the Rupelian via a strait in north- Paratethys. Only ten bivalve species were found in the ern Ukraine and Belarus, rather than via the Carpathian northern shelf area (from southern Ukraine to the Basin. In the CrimeanÐTranscaspian and PolishÐBel- Ustyurt Plateau), including the most characteristic gian assemblages, species in common are present that Ergenica and Janschinella garetzkii, which are unknown were absent from both the Eocene and Rupelian of Cen- further south. A more diverse assemblage (20 species) tral Europe, as well as from the Mainz and Paris basins. was typical of the southern (Transcaucasian) area of the Compare the gastropod assemblages from the CrimeaÐ basin; it was dominated by the forms that were rare or Transcaspian and northern Europe. unknown in the northern regions, including Urbinisia lata, Janschinella vinogrodskii, Merklinicardium, Number of species in Preston’s coef. Korobkoviella, and Corbula (Lenticorbula) mefferti. common The northern shelf of Caucasian Island (sections of the With the eastern area of 27 0.76 rivers of Fars, Tsraudon, and Fiagdon) was probably the North Sea Basin (Po- land, northern Germany, inhabited by impoverished fauna of a southerner type. Denmark) With the western area of 21 0.79 the North Sea Basin Scleractinian Corals (NetherlandsÐBelgium) Transcaucasia. Environments favorable for the With the Mainz Basin 22 0.82 development of coral reefs were retained in the Early With the Paris Basin 17 0.87 Oligocene only in the Yerevan Gulf that continued into

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S222 POPOV et al. the Central Iranian Basin. The coral assemblage was on scanty data on nummulitids). Nummulitids and, to closely similar in taxonomic composition to the Late certain extent, gastropods indicate that the Mediterra- Eocene and Oligocene assemblages from the southern nean influence was also rather marked in the Georgian regions of Europe. It included representatives of the area of the Paratethys, which should certainly be suborders Archeocoeniina, Faviina, Meandriina, and included in the North European Region. Fungiina. Representatives of the families Acroporidae, The North European Region is distinctly divided Stylophoridae, Faviidae, Heliastraeidae, Astrangiidae, into smaller units: however, the patterns of differentia- Agariciidae, Agathiphyllidae, and Poritidae are most tion varies depending on the groups involved in the frequent. About 35 species of 20 genera of the Arme- analysis. This region includes (1) the southener zone nian Assemblage are assigned to the colonial taxa. inhabited by impoverished assemblages of large fora- minifers, Nummulites, Operculina, and Discocyclina Complex Benthos-based Zoogeographic Zonation and by diverse subtropical-boreal molluskan assem- blages; and (2) the colder-water impoverished zone At the onset of the Oligocene, the faunal distinctions with mainly boreal biota inhabiting the northern and between the North African and Eurasian coastal areas eastern shelf areas. This division by the water tempera- of the MediterraneanÐMesopotamian Basin probably ture regimes is supported by the data on nummulites decreased because of an exchange between benthic and gastropods; in the case of mollusks, the Oligocene shallow-water faunas; the Ancient Mediterranean assemblages are divided into transitional and northern Region became integral, since it was not divided into (Amitrov, 1996, fig. 3). However, other groups indicate subregions. that this zonation is complicated by the paleogeo- Distinctions between the Ancient Mediterranean graphic peculiarities of particular basins. Thus, a clear and the North Sea regions still existed. The tropical and integrity of the Eastern Paratethyan Fauna is indicated subtropical fauna of the former included the reef facies by the compositions of bivalves (Popov, 1995), benthic inhabited by nummulites of the Southern Nummulitid foraminifers, and ostracodes; this allows one to con- Zone. Among small benthic foraminifers, there were sider these assemblages to be components of an inte- the specialized genera typical of only the Mediterra- grated province. These assemblages commonly desig- nean assemblages, such as Pararotalia, Herondenia, nated as the Chadumian assemblages are similar to the Queraltina, Pseudoplanulinella, and Sphaerogypsina, North Sea assemblages in species and generic compo- which usually had large, thick-walled, and sculptured sitions. Benthic foraminifers of these assemblages tests. Although the faunal differentiation within the included the same species of the genus Cibicidoides Ancient Mediterranean Region is rather high, the dif- (C. pseudoungerianus and C. sulzensis) and Cibicidina ferences appear to be occasional and do not allow for amphisyliensis, Ceratobulimina contraria, Rotaliatina the subdivision into provinces. bulimoides, Svratkina perlata, Turrilina alsatica, etc. In the Atlantic coastal area of Europe, the boundary In the shallow-water areas, an important role in the fau- was between the Aquitaine and the Paris basins (Fig. 3), nal composition was played by several species of aren- similarly to that in the Priabonian. The northern coastal aceous foraminifers (Haplophragmoides fidelis, area of the Carpathian Basin was included in the North H. deformabilis, Gaudryinopsis gracilis, etc.) that were European Region. In the south, in the region of the widespread in only the Pshekhian Basin and local taxa Transyvanian shelf, the basin was still opened in the that mainly had primitive tests. Similar relationships of first half of the Rupelian and inhabited by a fauna of the the molluskan and ostracode faunas from the Eastern Mediterranian type. The fauna of the Hungarian Paleo- Paratethys and northern Europe combined with clear gene Basins was mixed, which indicates the climatic differences between them allow one to designate the nature of this boundary in the Carpathian Basin. This Eastern Paratethys as the Eastern Chadumian Province area was inhibited by both the Tethyan species and gen- of the North European Paleobiogeographic Region. era, and the characteristic taxa of the North European Based on this assumption, the distinctions between the basins. This border (that was probably determined by a faunas of the northern and southern (Transcaucasian) continental barrier) further continued to the Balkanides coastal areas that are also very marked should be and isolated the Varna Gulf of the Eastern Paratethys regarded as minor (subprovincial) differences. from the Fore-Rhodope Basin, in which the Mediterra- The biogeographical position of the Azerbaijan nean Gulf entered. Similarly obvious faunal distinc- shelf of the CaucasusÐKopet Dagh Basin, with its tions were observed in Transcaucasia; Armenia was endemic fauna of benthic foraminifers, remains uncer- still inhabited by the Tethyan fauna with reef-building tain. It may only be mentioned that, judging by the corals, whereas the southern area of Georgia generic composition, this assemblage belonged to the (Akhaltsikhe) was a component of the Eastern Parat- North European Region and included certain rare spe- ethys assigned to the North European Region. The cies of the Chadumian Province. The faunal distinc- boundary between the Balkanides and the Lesser Cau- tions of the benthic foraminiferal assemblages from the casus probably extended across the Black Sea Basin, Turanian Subprovince were probably inherited from since the northern coastal area of the Anatolian Land the Eocene and gradually disappeared in the Rupelian. belonged to the Ancient Mediterranean Region (based The complicated stratigraphy of the TajikÐFerghana

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S223

Region does not allow for the discussion of biogeo- cies were found only there. In the category of ende- graphic importance of disctinctions of the unique fau- mism, this fauna is comparable to the modern-day Cas- nas from this region. pian Assemblage; the zoogeographic status is very A puzzle is the biogeographic similarity of the Syr- high, at a level of a separate region (Starobogatov, ian area of the MediterraneanÐMesopotamian Basin to 1970). However, among gastropods, only one species the Eastern Paratethys, which was inhabited by the for- was endemic; 75% of ostracodes were marine species aminiferal assemblages closely similar to the CrimeanÐ and endemic species were rare (Popov et al., 1985). Caucasian assemblages, as in the Priabonian (Krash- The Solenivian endemic genera and species of mol- eninnikov, 1965). The routes of exchange between lusks, ostracodes, and nannoplankton occurred from these faunas remain uncertain. the Western Carpathians to Kysyl Kum and Kopet Dagh. For this reason, we consider the entire Paratethys The faunas of the Paris Basin also appear to be of the second half of the Rupelian as the separate Sole- unique; this, along with the distinctions of the more novian Province (= Proto-Paratethyan Province after warm-water Aquitaine Fauna, probably displayed the Rusu, 1988). The Solenovian endemic species and gen- peculiarity of the Atlantic biota. Although the micro- era of fauna and microflora probably appeared in fauna and the macrofauna of the Aquitaine Basin lagoons of the Eastern Paratethys; subsequently, they included warm-water genera and species in common occupied the entire Paratethys. Thus, the Carpathian with the Mediterranean, this assemblage is rather Basin, which at the beginning of the Oligocene was unique and dominated by the species that were origi- partially included in the German Province, whereas its nally described there. southern (Transylvanian) area was included in the Shallow-water areas of the Paris Basin were inhab- Ancient Mediterranean Region, was involved in the ited by numerous endemic miliolids (among benthic Solenovian Province of the Paratethys from the second foraminifers) and mainly endemic mollusks. The spe- part of the Rupelian, as indicated by biogeographical cies in common with the North European and Mediter- data. It is evident that the Solenovian Province is a ranean faunas were mainly the cosmopolitans. How- component of the North European Region and subdi- ever, we assign this region to the North European vided into two subprovinces, the northern (ScythianÐ Region based on the absence of characteristic Mediter- Turanian) subprovince and the southern (Caucasus) ranean warm-water genera and species and the transi- subprovince. The latter probably included the entire tional pattern of the Mainz and Alsace assemblages. shelf of Greater Caucasian Island. It remains uncertain Biota of the North Sea Basin (assemblages of the whether the Carpathian Basin fauna belongs to the North Sea, Belgium, Netherlands, Denmark, northern southern or northern subprovince. Germany, and Poland) is the type of the Rupelian. The composition of this fauna was mainly descended from the Latdorfian (Late Eocene). Of the highest category Terrestrial Biogeography of endemism is the assemblage of benthic foraminifers, Phytogeography in which species of the genus Cibicidoides were numerous; Lenticulina herrmanni, Ceratobulimina In the Early Oligocene, the transition from the Pale- contraria, Rotaliatina bulimoides, Cibicidina amphi- ocenophytic to the Neocenophytic was accomplished syliensis, Svatkina perlata, Turrilina alsatica, Angulo- and resulted in a drastic change in the terrestrial flora gerina oligocenica, and the genera of the order Miliol- and vegetation. Plants of the ancient Cenozoic flora ida (Quinqueloculina, Triloculina, and Spiroloculina) represented by form-genera were entirely lost from the were common. Tropical elements and nummulites were vegetative cover and replaced by the Recent genera, absent from these assemblages. which formed the major core of the new flora. At the The faunas of the North Alpine and Mainz basins are beginning of the Early Oligocene, the floras of all similar to each other and substantially differ from the regions were characterized by approximately equal assemblages of the North Sea. However, the transi- proportions of the Paleocenophytic and Neocenophytic tional pattern of the Belgian Fauna indicates that these forms. By the end of the Early Oligocene, the transfor- distinctions are possibly facial, and, therefore, these mation had been completed almost everywhere. This faunas may be components of the integral German transformation was especially prominent in the midlat- Province with two subprovinces, the northern subprov- itudes of western Eurasia and extended from the ince with the typical Rupelian fauna and the southern Tethyan Region to the Boreal Region or to the ecotone subprovince enriched by warm-water elements. zone at the end of the Early Oligocene. The distribution of biota over the Paratethys As the northern boundary of the subtropical zone changed in the Solenovian time. However, available and the entire ecotone zone, which divided the major data on the faunal peculiarity of this area with reference phytochores was displaced southward, plants also to different taxonomic groups are rather contradictory. migrated submeridionally from north to south so that Bivalves show an extremely high peculiarity; of ten the ranges of moderately thermophilic deciduous genera and 18 species that inhabited Paratethys in the plants, forming the core of the Boreal Region Flora, first half of the Solenovian Time, six genera and 11 spe- increased. This process resulted in the formation of a

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S224 POPOV et al. moderately thermophilic, deciduous, mesophilic Oli- vascus and a nannoplankton assemblage of the NP23 gocene flora designated by A.N. Krishtofovich the Tur- Zone. The Vicenza Assemblage was dominated by scle- gai Flora. rophyllous laurels, Myricaceae, Quercus accompanied North Africa and Arabia. The North African Trop- by Myrsinaceae, Sapotaceae, Ericaceae, Araliaceae, ical Flora was widespread in the southern coastal area Sapindaceae, Anacardiaceae, Myrtaceae, Rhamnaceae, of the Mediterranean Basin, similarly to that in the Rutaceae, and Leguminosae; the latter family com- Eocene, although the southern ecotone zone, in which posed at least 10% of the entire flora. Zizyphus zizy- the exchange of floras occurred, occupied this area. The phoides, Myrica lignitum, and Tetraclinuis brongniartii modern-day Eastern Mediterranean, Asia Minor, and were common. Certain early conifers, such as Dolios- Iran were components of the Tethyan Region inhabited trobus taxiformis and Athrotaxis couttsiae, remained. by relatively hydrophilic flora in the first half of the Early Exotic tropical plants, dragon-tree and even Zamites Oligocene and by a subarid flora in its second part. (Bonnieux Flora), were recorded. The deciduous taxa include Alnus, Ailanthus, Acer, Aralia, Betula, Carpi- Southern EuropeÐLesser Caucasus. In the north- nus, Carya, Corylus, Cornus, Crataegus, Fagus, ern coastal area of the Mediterranean Region, humid Gleditschia, Juglans, Nyssa, Ostrya, Platycarya, Popu- climate changed to arid type at the onset of the Oli- lus, Platanus, Pterocarya, Sassafras, Ulmus, and Zelk- gocene. These new environments and the correspond- ova. Quercus sect., Ilex, Pistacea, Arbutus, Ceratonia, ing type of xerophytic floras covered southern areas of and Smilax first appeared in the floras of this level (flo- France, Italy, and Balkan Peninsular and expanded to ras of Aix, Bonnieux, and Chiavon). The major occur- Anatolia and the Lesser Caucasus. The earliest floras of rences of the floras of the second half of the Oligocene this South European Oligocene Province were recorded in the arid province are the Aix en Provance, Camoins- from the Marseille and Manosque depressions in the les-Bains, Saint-Zacharie, and Bonnieux in the southern area of France, Liguria, and Serbia. Their age Provence Province; Chiavon, Salcedo, and Novale in is supported by the occurrences of assemblages from the Vicenza Province; and one occurrence in Pavia. The the MP21 and MP22 vertabrate zones. In the Marseille floras of the southern province are similar to those of Depression, Mai (1995) assumes the Saint-Jean-de- the northern province in the presence of Arcto-Tertiary Garquier Flora from the vicinity of Semeno in marly elements, differing by the presence of xerophilous and limestones and gypsum beds at the base of the Estak microphyllous evergreen elements. Limestones to be the stratigraphic type of this level (names of the main floras shown in Fig. 4 are under- In the second half of the Early Oligocene, this flora lined). This is a typical subxerophytic laurel-leaved was in direct contact with the belt of subarid floras in flora, in which laurels were accompanied by pines, Transcaucasia (Akhaltsikhe, Lesser Caucasus). Rich legumes, and Tetraclinuis. The Taxodiaceae are repre- Lower Oligocene subtropical floras were studied by sented by Glyptostrobus europaeus and Sequoia abiet- Kasumova on the northern slope of the Lesser Caucasus ina. This flora and floras of similar age are dominated (Zeyva and Nizhnii Adzhakend in the Gyandzha Dis- by Laurophyllum primigenium, Daphnogene lan- trict, Azerbaijan). A small collection from these out- ceolata, Persea princeps, Myrica longifolia, Compto- crops were subsequently treated by Akhmetiev. The nia schrankii, Zizyphus zizyphoides, and Pinus paleo- Lauraceae and Fagaceae (Eotrigonobalanus, Lithocar- strobus. Doliostrobus taxiformis is common. Palms and pus, Cinnamomum, and Daphnogene) were the major ferns are frequent. The deciduous plants Carpinus cus- forest-forming taxa; they were accompanied by the pidens, C. grandis, Acer primaevum, Betula oblongata, Taxodiaceae, Myricaceae, and Ericaceae (Myrica agd- Platanus leucophylla, Pterocarya parasidiaca, Craigia jakendica and Leucothoe protogaea). europaea, and Cedrelosphermum leptospermum are Palibin described a subtropical flora from the basal relatively rare. Semievergreen sclerophyllous low- sandy member of the Shorakbyur Formation of the Yer- growing forests and brushes are the major components evanÐOrdubad Zone (Shorakbyur) underlain by the of the zonal vegetation. marly clayey Zovashen Formation with Nummulites In addition to the type flora, the group of the earliest fabiani and the Later Eocene planktonic foraminiferal Early Oligocene floras of the Marseille Depression assemblage of the P17 Zone. In addition to herbaceuos includes the Fenestrelle and Montespin floras, as well monocotyledons, the Shorakbyur Assemblage includes as the Cereste Flora in the Manosque Depression, the Litsea primigenia, Sapindus graecus, Ilex ambigua, Gargas and Sult floras in Vaucluse, and the Santa Gius- Myrica hakaefolia, etc. Based on planktonic foramini-

fers (analyzed by Krasheninnikov), this part of the sec- tina Flora in Liguria.^ In Serbia, the Ljiljanska Reka Flora from the Pc einja Depression is similar in general tion belongs to the Globigerina tapuriensis (P18) Zone. pattern. The spore and pollen assemblages from the basal The flora of the second half of the Early Oligocene Oligocene of Armenia were characterized by a high is represented by the Vicenza Flora found in the hom- proportion of pollen of the Pinaceae and an increased onymous province of Italy in the vicinity of Farra. Mai range of subtropical xerophytes (Nypa, Elaegnaceae, (1995) considered this to be the type flora. This flora Malvaceae, Bombacaceae, etc.). Frequent fluctuations was found in the littoral marl beds with Nummulites of the temperature and the climatic humidity were

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S225

50° Kazakhstanian

Province 33 N o r t h E u r o 34 35 p e a n P r o v i n c e 29 32 31 25 22 26 24 27 21 23 28 30 40° 14 Middle 12 13 C e n t r Asian 10 a l E u r o p e a 19 11 15 n P r o v i n c e Province 18 36 16 20 I O N 9 2 3 A N R E G 7 8 T E T H Y S o u t h 4 E u r o p 6 30° e a n P r 5 o v i n c e

1 2 3 N O R T 4 H A F R I C A 5 N R E G I O 0 500 km 6 N

Fig. 4. Phytogeographic zonation of the Boreal and Tethyan regions in the Rupelian. Explanations: (1Ð3) Boreal Region: (1) decid- uous mesophilic coniferÐbroad-leaved forests, including pinaceous, taxodiaceans, juglandaceans, betulaceans, and fagaceans; (2) xerophytic oakÐlaurel forests and sclerophyllous shrubs formed by legumes and ericaceans; tugai forests with poplar, alder, and ulmus in valleys; (3) shrubÐherb formations, including halophytes (Ephedra, Artemisia, and Salsola); (4Ð5) Tethyan Region; (4) subtropical mesophilic forests formed by taxodiaceans, cupressaceans, fagaceans, and lauraceans; (5) subxerophilous and xero- philous forests formed by lauraceans, legumes, fagaceans, and ericaceans; (6) North African Region: tropical forests formed by lauraceans, legumes, and, occasionally, by fagaceans; farther inland, these are replaced by open savannah woodlands with legumes and eophobic plants. The main Rupelian floral localities: (1) Aix en Provance; (2) Saint Jean de Garquier; (3) Vicenza; (4) Edirne; (5) Shorakbyur; (6) Darry-Dagh; (7) Akhaltsikhe; (8) Tori; (9) Zeyva; (10) Lobstann; (11) Vivey; (12) Nerhau; (13) Valec; (14) Haselbach; (15) Csillaghegy (NP22); (16) Sotzca (NP23); (17) Kiscell; (18) Mera; (19) Belaya River; (20) Mayakovski; (21) Devonshire; (22) Sieblos; (23) Dormach; (24) Haselbach; (25) Svetlogorsk; (26) Volhynia; (27) Starodubka; (28) Pasekovo; (29) Kyzyltobe; (30) Ashcheairyk; (31) Zhabyske; (32) Kumbulak; (33) Altyn Chokysy; (34) Chelkar Nura; (35) Tortmolla; (36) Tajik Depression. favorable for rapid disintegration of the Eocene flora. The floras of the second half of the Early Oligocene The frequent humidity fluctuations in the spore and pol- of Transcaucasia were more xerophilous; this indicates len assemblages of Armenia are supported by the oppo- a new increase in aridity. Among these floras, there is site phase distribution of cryptogams as indicators of the well dated Tori Flora from the Corbulla Beds, humid regimes and pollen of grasses, Ephedra, the Borzhomi District, of the Solenovian age, although, in Cupressaceae, and certain other plants that characterize addition to the xeromorphic sclerophyllous plants this the arid regime. Zaporozhets indicated that the frequent flora includes large-leaved taxa. This flora is polytopic, alternation of humid and arid climatic phases in the sec- dominated by Dryophyllum curticillense and Myrica ond half of the Late Eocene in the southwestern regions lignitum. Avakov found fruits of legumes in Tori. The of the former USSR was also typical of the Early Oli- floras from the Solenovian level of the north-facing gocene. The first humid phase in Armenia corresponds slope of the Lesser Caucasus are far more xenomorphic to the epoch of accumulation of beds of the lower (but after Kasumova. for the basal beds) part of the Globigerina tapuriensis Central Europe and Northern Caucasus. The ear- (P18) Zone, whereas the arid phase corresponds to the liest Early Oligocene floras in the midlatitudes of west- lowermost part of this zone. The second part of the ern Eurasia correlates with the NP21ÐNP22 nanno- G. tapuriensis Zone was characterized by a more plankton zones, the Globigerina tapuriensis Planktonic humid climate, followed by an arid phase that corre- Foraminiferal Zone, the Phthanoperidinium amoemum sponds to the transition area of the P18 and P19 zones. Dinocyst Zone, Grande Caupture, and the MP21 Mam-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S226 POPOV et al. malian Zone. These floras differed little from the flora meyeri was found in the adjacent Brez^ ani Flora of from Bembridge Beds of the terminal Eocene; there- approximately the same age. The taxodiaceae are rep-

fore, some paleobotanists assigned them to the Early resented by the early Doliostrobus and Athrotaxis, as Oligocene.^ Mai (1995) considers the western Bohe- well as by Sequoia abietina. The Leguminosae are rare mian Valec Flora from the beds containing mammals of because of the humid type of flora; however, all of them the MP21 Zone to be the type flora of this level. This belong to evergreen plants. Legumes dominate only the flora and the floras of similar age from the northern more xerophytic Sotzca Flora in Slovenia from the beds coastal area of the Paratethys display the vegetation of with Rzhehakia. The proportions of laurels, legumes, the mesophilic forests formed by the Lauraceae and conifers in this flora are 25 : 51 : 23.8. Palms occur (including Daphnogene), oaks with the entire leaves, in the floras from the Red Molassa in Switzerland and magnolias, etc. Among thermophytes, an important are even more usual in the Balkan floras. Deciduous ele-

role was played by the Eocene Doliostrobus certus, ments mainly inhabited river valley facies, as in Turgai. Zizyphus zizyphoides, Myrica banksiaefolia^ , and Lygo- Further to the east, the floras of this type are traced dium kaulfusii. However, the Valec Flora also includes in the CrimeanÐCaucasian Region. The Belaya River many deciduous plants, such as Alnus, Carpinus, Flora from the Planorbella Beds (transitional interval Ulmus, Zelkova, Celtis, Carya, Cercidiphyllum, etc. between the NP21 and NP22 zones) with the presence The subsequent subtropical floras of Central Europe of evergreen Myrica, Fagaceae, Lauraceae, and Taxodi- are the following. The Nagybatony Ujlak Flora (Hun- aceae (Akhmetiev, 1995) is similar to the subtropical gary) and Mera Flora (Romania) come from the beds floras of Western Europe. Uznadze earlier reported the with the NP22 nannoplanktonic assemblage. The floras presence of palm fossils in western Ciscaucasia. She of Csillaghegy, Budapest, Kiseged (found in the vicin- also recorded a similar but more taxomonically impover- ity of Eger), and many Romanian floras [in the vicinity ished assemblage from the Lower Oligocene beds in the

vicinity of the town of Mayakovski (western Georgia). of Cluj; Aghires, Muereasca^ des Sus, and Sulanesti (in the vicinity of Pites ti), and Clit, Jac, and Ileanda (in the Western and Northern Europe. Evergreen and vicinity of Calau)] come from the NP23 zone. In Slov- deciduous plants were jointly found in the majority of enia, the Sotzca Flora (from the beds of the NP23 Zone taphocenoses from the northern margin of the epeiric and the MP23 Vertabrate Zone) and the floras from sea; this allows for their inclusion in the ecotone zone. Savine, Trbovlje, and Zagorje are of the same age. The Subtropical plants dominated the coastal zone in west- same type of flora was found in the Zenica-Sarajevo ern regions, whereas plants of moderately warm cli- Basin in Bosnia and in Reit, Horw (near Bern), and mates inhabited the eastern regions. Vevey (Lake of Geneva). In the southeastern area of the

The West European floras from the Rhine Graben

^ Balkan ^ Peninsula, the floras from the Rhodope Massif (Momc ilovic i) and Edirne are assigned to the same (Sieblos Flora) and Mainz Basin (Dormach Flora)

include numerous plants with the entire leaves typical type. In addition to the^ Kiscell Flora, the latest flora of this type is the Bystric e Flora from Bohemia; it comes of the Leguminosae. The abundance of Nyssa and from the NP24 Zone. Glyptostrobus in the Sieblos Flora indicates the devel- opment of swampy forests at that time. The type spore According to Andreansky and Givulescu (1980), the and pollen assemblage of this level is recorded from a majority of the Early Eocene floras from Central section in the vicinity of the town of Marburg (Heskem- Europe indicate the luxuriant mesophilic forests. The Bild, PG19 Pollen Zone); it includes abundant and major forest-forming plants were Eotrigonobalanus diverse pollen of deciduous plants exceeding coeval (Castanopsis), evergreen oaks, and laurels accompa- megafossil assemblages in diversity. This is associated nied by the conifers Doliostrobus and Athrotaxis. The with a high pollen productivity of conifers and the Arcto-Tertiary forms are usually scarce; their propor- Amentiflorae and a low polen productivity of laurels tion in the taphocenoses only rarely increases to 5Ð10%; and other evergreen plants pollinated mainly by insects. only in the Sulanesti Flora (Romania) it is 18%. The Heskem Assemblage is dominated by the Gymno- Mai considers the Kiscell Clays Flora (Budapest, sphermae and includes abundant pollen of Carpinus. Obuda) from beds transitional from the Lower to the Pollen of Tsuga, Juglans, Carya, Pterocarya, Fagus, Upper Oligocene (NP24 Zone) as the type Early Oli- and Betula play a significant part. Boehlensipollis gocene flora of Central Europe. This flora is large- absent in the Eocene is very typical, as well as Milfor- leaved and dominated by Eritrobalanus, Daphnogene, dia and pollen allied to the Chenopodiaceae. In addi- Laurophyllum, Zizyphus zizyphoides, and species of tion to the above-enumerated floras, this group also Myrica. The Fagaceae are represented by Trigonobala- includes the floras of Hagenau and Habsheim located in nus, Lithocarpus, and Quercus; the Lauraceae include the vicinity of Mulhouse in the Rhine Graben. Litsea, Ocotea, Persea, Laurus, and Sassafras. Plata- In the eastern continuation of the ecotone zone, nus neptunei, Comptonia shrankii, Palaeocarya mac- Portnyagina considers the transition EoceneÐOligocene roptera, Cedrelospermum leptospermum, and Sterculia Sheshora Beds in the Carpathains (beds with Globige- labrusca are common. Mai considers this assemblage rina vialovi) to be the Tricolporopollenites villensisÐ to be a typical mastixial association, because Mastixia Tricolporopollenites cingulum Pollen Zone with a high

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S227 proportion of pollen of the Taxodiaceae and the This resulted in the formation of cypressÐnyssa bogs Pinaceae and a lower pollen diversity of thermophytes within the dried areas with the presence of abundant compared to that in the underlying Upper Eocene beds. evergreens, ericaceous Epacridicarpum rossicum and The spore and pollen assemblage from the lower beds deciduous Acer tricuspidatum, Nyssa zhilinii, Populus of the Mezhigorian Formation of the DnieperÐDon populina, and Sorbus praealnifolia. The appearance of Depression is characterized by the miospores assem- new migration tracts after the sea regression resulted in blage including Sciadopitys vertillaciformis and Retitri- the formation of wide connections of the Pasekovo colpites foraminatus (Stotland, 1984). Flora with the Svetlogorsk Flora that occurred further north, as well as with the western (Haselbach) and east- The next level of the Early Oligocene floras from the ern (Kazakh) floras. This is supported by both the sim- ecotone zone of Europe was designated by Mai the ilarity in age and the commencement of active floristic Haselbach Flora after the type locality Haselbach found exchange. Taxodium balticum is a typical species of the in the vicinity of Altenburg south of Leipzig. Although Pasekovo, Svetlogorsk, and Haselbach floras. Nyssa this flora has not been aged with certainty based on zhilinii with the dentate leaves was described from other fossil groups, its assignment to the Early Rupe- Pasekovo and Kazakhstan, whereas its vicarious spe- lian is beyond question. The Haselbach Assemblage cies was found in the Altenburg Flora coeval with the displays a mixed mesophilic forest vegetation with a Haselbach Flora. The entire leaves of N. punctata, as high proportion of hydrophilic holophytes. This assem- well as Apocynophyllum unite the Pasekovo and Svetl- blage includes deciduous Arcto-Tertiary elements, as ogorsk floras; the large-leaved A. helveticum is known well as laurel-leaved subtropical plants composing at from these floras and the Haselbach and Aral floras. least 50% of the assemblage. The latter include rela- The Pasekovo flora is similar to the Kazakh floras in the tively early taxa, in particular, Aquilaria, Caricoidea, presence of lobe-leaved red oak Quercus pseudoalex- Litsea phoenica, Myrica longifolia, Phoebe, etc. At the eevii. The absence of Quercus pseudoneriifolia, which same time, representatives of the so-called young mas- was wide spread in all floras of the northern subtropical tixian flora, i.e., Aralia haselbachensis, Ampelopsis hibschii, Manglietia zinkeisenii, Mastixia meyeri, province of the Tethyan Region in the Late Eocene, is noteworthy. This species remained in the Early Oli- Myrica, Nyssa, Meliosma, Ilex, etc., play a significant gocene floras, but only in typical subtropical floras. part in this group. However, this flora also includes numerous typical deciduous plants. Some of them The floristic assemblages of Flörsheim and Nerhau occurred to the Early Miocene (Acer haselbachense, belong to the NP23 Nannoplankton Zone and the lower Carpinus mediomontana, Carya quadrangula, Comp- part of the Wetzeliella gochtii Dinocyst Zone (after tonia goniocarpa, Populus hermanica, Taxodium balti- Mai, 1995). Flörsheim is the type locality found in fish- cum, Rosa lignitum, etc.); however, the species known bearing shales of the Mainz Graben; Nerhau is in from the earlier beds, such as Alnus kefersteinii, Carpi- clayey beds of the Weisselster Depression, in the vicin- nus cordataeformis, Ceratophyllum lusaticum, Disan- ity of Grimma, southwest of Leipzig. Both these thus bavaricus, Liquidambar europaea, etc., were also assemblages are dated to the beginning of the Late reported. The flora-bearing beds include pollen of Boe- Rupelian (NP23 Zone). The Nerhau Flora is a typical hlensipollis and other taxa that allow for the assignment laurel-leaved flora with Arcto-Tertiary accessories; of the Haselbach Assemblage to the PG20 Pollen Zone. therefore, among the ecotone floras, it is closer to sub- tropical floras. Its major components are Eotrigonobal- The Haselbach Flora from the ecotone zone of anus furcinervis, Zizyphus zizyphoides, Myrica longifo- Western and Central Europe may be correlated with the lia, Comptonia schrankii, Trigonobalanus rhamnoides, Devonshire Flora (Boney Tracey, Heathfield), Alt- and Sterculia labrusca. Based on available carpological enrath Flora in Siebengebirge along Rhine, and with the data, Mai considers this flora among young mastixial floras from the Baltic coastal areas (Svetlogorsk and floras, which include Tetrocarya nerchauensis, Mas- Otradnoe), Voronezh Anteclise (Pasekovo), Ustyurt tixia boveyana, Symplocus anglica, etc. The Nerhau (Ashcheairyk), and the Aral Region (Kumbulak). In the Populus zaddachii, Flora includes the following Arcto-Tertiary deciduous Svetlogorsk Flora, the deciduous Acer angustilobum, A. intergerrimum, Betula Alnus heerii, A. pseudogracilis Pinus thomasiana taxa: , and ; brondniartii, B. prisca, Cercidiphyllum crenatum, Athrotaxis couttsiae Sequoia abiet- the evergreen and Craigia europaeum, Fagus deucalions, Pterocarya ina; the palm Calamus daemonorops, etc. are common paradisiaca, etc. An extensive group of floras of this as well as the above-enumerated taxa (after Budantsev age with various proportions of paleotropical and and Sveshnikova). Arcto-Tertiary elements is known in Europe. The floras The Pasekovo Flora, occurring in the ecotone zone, from the volcanic-detrital beds of the Bohemian Sredna is more similar to those of the Tethyan Region, while Gora are distinguished by the abundance of the entire- the others are similar to the Boreal Flora. According to leaved laurophilous plants combined with a smaller Vikulin (1991), the Pasekovo assoications (compared proportion of deciduous Arcto-Tertiary forms (60 : 40 to the Eocene groups including laurels, castanopses, in the Berand and Markvartice floras and 30 : 70 in the and the other primitive Fagaceae) belong to a unique Suletice Flora). In the spore and pollen assemblages, type that developed in the course of regression of sea. the typical taxa are Boehlensipollis hohlii, Slovakipollis

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S228 POPOV et al. hyppophaeoides, Tricolpopollenites henrici, and pollen porata. In the southern part of Western Siberia and in of the Chenopodiaceae. After the type sections, this Northern Turgai, a typical transitional assemblage with spore and pollen assemblage is designated the Ber- abundant gymnospermous pollen and a high proportion gischÐGladbach or the Calau Assemblage (PG20aÐb of thermophilic pollen of the Fagaceae (Quercus graci- Pollen Zone) from the type sections. This group also lisÐQ. graciliformis group) was recorded from the Kur- includes certain other floras from the Mainz Depression gan Beds; Zaporozhets identified Phthanoperidium and the Weisselster lignite-bearing basin, DittelsheimÐ amoenum in the same locality. Kloppberg (east of Else), and Niederwalluf (in the The spore and pollen assemblages from the Pasek- vicinity of Eltville). The deciduous taxa are more ovoÐKumbulak level of the ecotone zone (correlated numerous and diverse in the Bechleijovice and Hrazeny with the NP22 Nannoplankton Zone and the Wetzeliella floras. However, they existed later and were aged to the symmetrica Dinocyst Zone of marine beds) are domi- transition between the NP23 and NP24 nannoplankton nated by conifer pollen (pine or the Taxodiaceae) and zones. contain a high proportion of the Amentiflorae, espe- Northwestern Asia. Further to the east, in the eco- cially the Juglandaceae. According to Panova and tone zone, the Rupelian floras are well known based on Zaporozhets, in Kazakhstan, the pollen floras of this spore and pollen data and, to a lesser extent, on type are combined into the Carya spackmaniaÐTilia megafossils. The floral composition at the beginning of grandireticulataÐJuglans sieboldianiformis Pollen Zone. the Oligocene is mainly judged based on the analysis of This zone corresponds to the peak of the Ashcheairyk pollen assemblages from the beds that are assigned to Transgression (beds with Cypris) and to the accumula- the Phthanoperidinium amoemum Dinocyst Zone. Of tion of the upper beds of the Kutanbulak and Urkimbai the megafossil assemblages, only the Kyzyltobe Flora formations. According to Stotland, in the Ukrainian from Mugodzhary, which was studied by Rayushkina Shield, the upper beds of the Mezhigorian Formation (1979), is conventionally assigned to the floras of this with Pinus silvestris, Carya spackmania, etc., corre- level. This flora includes derivatives of both the sub- spond to this zone. tropical flora (Torreya and Sequoia with the typical The spore and pollen assemblages from the Solen- unguiform leaves, Myrica, Magnolia, and an unidenti- ovian Horizon of the East European Platform and Turan fied genus of the Lauraceae) and the temperate decidu- Plate were assigned by Boitsova (1973) to an integral ous flora. The spore and pollen assemblages from the Juglans sieboldianiformisÐBetula gracilis Pollen East European Platform and the northern area of the Zone. This zone was recorded in the Northern Ustyurt Turgai Platform (Northern Ustyurt, Aral Region, and beds, the lower beds of the Chilikty Formation in the Turgai) differ in the predominance of the Taxodiaceae North Aral Region, and the lower beds of the Betpak- and the Pinaceae, a rather high proportion of the Dala Formation of the Chu-Sarysu Depression. For this Juglandaceae and Betulaceae pollen that dominate over level, Zaporozhets (Akhmetiev and Zaporozhets, 1992) the Fagaceae pollen, and in the presence of the Hama- reported a decrease in the proportion of gymnosper- melidaceae, Myricaceae, Ulmaceae, etc. The propor- mous pollen and an acme of the Qurcus pollen (includ- tion of pollen of evergreen plants is low. Judging from ing the fine pollen of the Quercus gracilisÐQ. gracili- the above facts, this is associated, on the one hand, with formis group); the researcher treated this as a result of a high pollen productivity of the anemophilous Amen- climatic warming in the Solenovian Time, increased tiflorae, and, on the other hand, with a better represen- seasonality, and aridity of climate, especially in Turgai. tativeness of pollen of plants that inhabit riparial areas, where deciduous forms appear earlier than on low and The floras with Ergenica cimlanica from the Solen- relatively dry flat interfluve. ovian Horizon of Northern Ustyurt were mainly com- posed of deciduous species, although Myrica longifolia The earliest Early Oligocene flora is the Kumbulak was still found. In the Chagrai Plateau, in the Zhabyske Flora (Kutanbulak Formation, North Aral Region); it is locality, the flora was also formed by deciduous taxa dominated by moderately thermophilic forms with ref- but also included Myrica longifolia. The Kenkous Flora erence to the number of specimens, whereas the ever- recorded by Budantsev (1959) from this plateau green microphyllous forms with marked xeromorphic included certain evergreen plants (M. longifolia and characteristics are more diverse taxonomically (Sequoia, Apocynophyllum), along with the broad-leaved genera Cinnamomum, Myrica, Apocynophyllum, etc.). Myrica Juglans, Fagus, Ulmus, Liquidambar, etc. lignitum and Apocynophyllum are solitary in the Kutan- The distribution of mesophitic and xerophitic plants bulak Formation of the Altyn-Chokysy outlier and in in the Late Rupelan floras of the Aral Region and Turgai the Urkimbai Formation of Turgai (Chelkar-Nura). (correspond to the Solenovian Horizon sensu lato) is of A high proportion of pollen of pine and the Taxodi- great interest. The floras from the flat interfluves aceae in the floral assemblages is typical of the basal (assigned to this group based on the phytocenoses of beds of the Ashcheairyk Formation of the Northern temporal streams) were dominated by the derivatives of Ustyurt Plateau (Phthanoperidinium amoenum Dinocyst the Eocene evergreen xerophitic flora, Palibinia and Zone). Panova designated these beds as the beds with Rhus turcomanica, microphyllous sclerophyllous lau- the Pinaceae, Quercus graciliformis, and Juglans poly- rels, etc. (Tortmolla, Zhaman-Kaindy, Mynyske-Suiek,

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S229 etc). At the same time, coeval riparian floral assem- (1) In the Early Oligocene, as in the Late Eocene, blages that characterize taphocenoses of the oxbow western Eurasia was included in two regions of the Hol- facies were dominated by macrophyllous deciduous arctic Kingdom, the Boreal Region occupied by a mod- poplars, elms, alders, and ashes. In the Late Oligocene, erately thermophilic deciduous flora and the Tethyan these trees formed the core of the Turgai vegetation. Region with subtropical flora. The regions are isolated Hypoautochthonous burials include both the fossils of from each other by a wide ecotone zone. The latter xenomorphic leaves and the larger dentate leaves of replaced the Late Eocene North European Province of deciduous species. According to Kornilova (1963), the Tethyan Region because of the development of such a mixed type of taphocenoses allows for the rec- hydrophilous subtropical flora and included the entire ognition of floras of a transition type between the sub- territory of central and northwestern Europe. tropical and the temperate (so-called Shintuzsai) floras (2) Climatic cooling in the terminal Eocene and the in the Lower Oligocene of Kazakhstan. However, the beginning of the Oligocene resulted in the displacement xeromorphic pattern of the Turgai floras may indicate of the boundary between the Boreal and Tethyan not only the arid habitats but also physical adaptation to regions by at least 10°Ð15° to the north. The southern halophytic and littoral facies, taking into account the boundary of the ecotone zone, which should be occupation by flora of coastal lowlands formed as a assigned to the Boreal Region, extended across Western result of recent sea regression. In the Solenovian, the Europe, the Carpathian Basin, the DnieperÐDon Turgai flat interfluves were mainly bald, being gradu- Depression, and continued across the Eastern Parat- ally occupied by shibliak, low grasses and dwarfshrubs. ethys. A decrease by 4Ð6°C in the average annual tem- This inference follows from the presence of abundant peratures (the average winter temperatures decreased, pollen of the Chenopodiaceae, including Artemisia, while the average summer temperatures remained con- and the Compositae in the spore and pollen assem- stant) in the Early Oligocene compared to those in the blages from the Chelkar Nura beds with indricotheres Late Eocene was the major factor of the changes in veg- (after Zaporozhets). etation. Central Asia. At the beginning of the Oligocene, the climate in Central Asia became more humid (3) In the Early Oligocene, the Tethyan Region because of a climatic cooling; however, arid regimes (sensu stricto) significantly decreased in area down to returned in the second half of the Oligocene. The com- only two former Late Eocene provinces, the Central mencement of tectonic destruction of the Epihercynian European Province and the South European Province. Platform resulted in the development of heterogenous It retained the division into two provinces; however, vegetative cover in different areas of the province, as their characteristics substantially changed. The Central indicated by spore and pollen data obtained in the Tajik European Province was inhabited by hydrophilous Depression (data obtained by M.Z. Pulatova and flora, whereas the South European Province was occu- A.M. Pen’kova). In the Baisun District, desertÐsteppe pied by subarid assemblages. The boudary between formations with the participation of the Gramineae, these provinces differed from that in the Eocene Artemisia, Ephedra, the Compositae, and the Chenop- (Fig. 4). Despite marked climatic and humidity odiaceae were widespread. Southern slopes of the changes, the entire northern coastal area of the Mediter- Hissar Mountains were occupied by coniferÐbroad- ranean Basinretained the provincial status in the Early leaved forests with laurels. The forests alternated with Oligocene Tethyan Region; in the Late Eocene, it was open areas. This zone was most humid within the included in the South European Province with flora of depression. In the Vakhsh District, steppe grassy forma- the transition type between subtropical and humid trop- tions of flat interfluves alternated with the bush forma- ical floras. At the onset of the Oligocene, this province tions. In valleys, the intrazonal forest communities experienced aridification. Further expansion of the arid were formed by Populus, the Juglandaceae, and the zone occurred in the Solenovian Time, as indicated by Fagaceae. In the eastern area (Pyandzh River Basin), a wide development of xerophilic forms in certain Bal- the steppe and meadow formations were dominated by kan assemblages. This flora occupied the Iberian Pen- the Chenopodiaceae, Compositae, Artemisia, Gramineae, insular, southern France, the major area of Italy, and Leguminosae, and low-growing Ephedra. The main partially the Balkan Mountains and extend to Trascau- changes in the floral assemblages from the Tajik casia. Depression occurred in the Late Sumsar Time, when To the contrary, the climate in the northern area of the majority of the Eocene forms disappeared and the the Tethyan Region became more humid with cooling. spore and pollen assemblages were dominated by the The Late Eocene xeromorphic flora of Central Europe Taxodiaceae, Juglandaceae, and Oleaceae. was replaced by the humid type, as indicated by the macrophillous character of the majority of floras. The Early Oligocene floral assemblages were rather homo- Phytogeographic Zonation geneous in composition in both western and eastern The study of the Early Oligocene flors and vegeta- areas, where this flora occupied the entire northern part tion and their comparison with the Late Eocene pattern of the CrimeanÐCaucasian Region and the Scythian allow the following conclusions (Fig. 4): Platform.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S230 POPOV et al.

(4) The ecotone zone extended from Western climatic parameters of the Late Rupelian floras with Europe to the Aral Region and covered the entire north- those of earlier floras from the Northern Province of the ern coastal area of the Paratethys. This zone was inhab- Tethyan Region and from the ecotone zone. The ited by the broad-leaved mesophilic forests with lau- increase in yearly average temperatures in the west was rels; the Eocene relicts played a significant part. The associated with the increase in average winter tempera- flora of this zone markedly changed during the Rupe- tures combined with unchanged average summer tem- lian. peratures. Conversely, in the east (Turgai), this was (5) The predominance of deciduous forms in the associated with an increase in average summer temper- taphocenoses of Ustyurt and the Aral Region, and a low atures combined with increased seasonality of climate proportion of evergreens allow for the exclusion of and the development of continental climate. these regions from the ecotone zone, beginning from the second half of the Early Oligocene, and their reli- able assignment to the Boreal Region. In the Aral Insects Region and Turgai, the floras from riparial facies are The Early Eocene entomofaunas of Western Eurasia represented by deciduous plants, whereas the floras have not yet been studied in detail. Several large local- from the sediments accumulated by temporal streams ities that have been long considered to be of the Lower of the flat interfluves are composed of evergreens. Only Oligocene age most likely belong to the terminal at the end of the Solenovian, the flora of northern and Eocene; therefore, they were discussed in the first part central Kazakhstan took on the Turgai characteristics; of this work (Zherikhin in Popov et al., 2001). It should this allows one to assign with certainty this flora to the be emphasized that the entomofaunas (at least those of Boreal Region. North America and Africa) had probably taken the Oli- (6) In contrast to the floras of the Northern Ustyurt gocene pattern before the commencement of marked and North Aral Region, which insignificantly differed cooling at the EoceneÐOligocene boundary. For this from the floras of westerner regions of the ecotone zone reason, it is very difficult to distinguish between the in the Early Oligocene, the Turgai floras were more Eocene and Oligocene insect assemblages at the unique. We have good grounds to consider this territory present state of study. This task necessitates consider- as a descendant of the Late Eocene Kazakh Province. ation of other stratigraphically more important groups (7) In the Early Oligocene, the Central Asian Prov- of organisms from the same beds, especially plant fos- ince of the Tethyan Region retained the same bound- sils, which usually accompany insects in the burials. aries; however, its characteristics changed. The differ- The major features of the terminal Eocene assem- entiation of floras in this territory probably indicates the blages, such as the Brembridge Marls Fauna in England reactivation of the Alpine Orogeny in Central Asia. and the Célas Fauna and fauna from the salt-bearing beds of Alsace in France, may probably be extrapolated (8) Two stages may be distinguished in the develop- to the Early Oligocene entomofaunas of Western ment of the Late Oligocene floras. They may be con- Europe. ventionally designated as the earlier (Presolenovian) stage and the later (Solenovian) stage. At the first stage, The most important Early Oligocene entomofaunas the floral rearrangement was especially well pro- are known from Italy (amber from Sicily and the north- nounced due to a commencement of active expansion ern Apennines), southern France (Camoins-les-Bains in of the Arcto-Tertiary deciduous elements into the sub- the Bouches du Rhone Department), and Germany tropical flora and the elimination of Eocene elements (Sieblos in Bavaria). Other regions of Western Europe from the vegetative cover. At the second stage, the arid (from Spain, central France, and Menilithic Formation zone expanded, especially in the eastern area of the in the Carpathians, in particular) yield only isolated Tethyan Region; this resulted in complete formation of findings. These specimens insufficiently characterize the southern subarid province, which included the entomofaunas of this age. The occurrences of termitaria southern province of the Tethyan Region in the west in Fayum (Egypt) are also noteworthy. (this was earlier involved in a more humid zone) and the In North Africa, the Early Oligocene insects are southern area of the Kazakh and Central Asian prov- only known by ichnolites. In Egypt, soil termitaria of inces in the east. This wide zone of subarid floras rather diverse types (five ichnospecies of four ichno- became more distinct from the Late Rupelian (Solen- genera) were found in the Jebel Quatrani Formation of ovian) Time (NP23). This zone extended from southern the Lower Oligocene. They include large and complex France via Italy, the Balkan Uplift, and Transcaucasia termitaria indicating families of large size (Genise and to Kazakhstan and Central Asia. In Turgai, it expanded Brown, 1994). The taxonomy of termites that built northward, where the arid flora of the Tethyan Region these nests remain uncertain. It is not improbable that and flora of the Boreal Region met. some of them belong to the tropical Termitidae. It (9) A climatic warming that occurred in the Eastern should be emphasized that two ichnospecies are in Paratethys in the Solenovian time (Akhmetiev and common with those from the underlying Upper Eocene Zaporozhets, 1992) was also observed in Central Qasr el Sagha Formation, in which termitaria are rela- Europe. This became evident when we compared the tively rare and less diverse.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S231

Of particular interest is a fauna from the Italian and bushes) is typical. These taxa include, in particular, amber that was generally described by Kohring and the Pompilidae and large Sphecidae (of the Hymenoptera) Schlüter (1989) and by Skalski and Veggiani (1990). and the Asilidae and Bombyliidae (of the Diptera). These Until recently, the Sicilian amber (known only in rede- features differ the Camoins-les-Bains Fauna from all posited state) was aged as the Miocene. However, it is the Late Eocene faunas of the more northern regions of now established that this amber is closely similar to that France (Languedoc and Alsace), the Early Eocene from the Arenarie di Ranzano (Lower Oligocene; Ran- entomofaunas of which have received only fragmentary zano Sandstone, northern Apennines) in both chemical study. The generic composition of ants (which have not properties and in faunal composition (Skalski and Veg- been described) was determined (if correctly) by giani, 1990). The fauna from the Italian amber is of par- Timon-David (1943) and is also consistent with the ticular interest, because it has no equivalents among assumption of the landscapes of open woodlands. Cenozoic insect assemblages of western Eurasia and A preliminary study of material on the Coleoptera closely resembles the faunas from the Lower Miocene (without a description) from the National Museum of amber from the Dominican Republic and southern Natural History (Paris) supports the similarity to the Mexico (Chiapas). This naturally does not indicate any Late Eocene faunas from Provence. In our opinion, the direct faunal connections. The similarity is evidently Camoins-las-Bains Fauna occurred in warm and sea- associated with similar landscapes and climates. The sonally arid (possibly summer-dry) climates and was most distinctive feature in all three cases is the abun- most likely an ancestor of the extant Mediterranean dance of timber beetles of the subfamily Platypodinae. entomofaunas. Autochthonous aquatic entomofauna is This indicates the production of these rezins by absent (insects were probably buried in a brackish- angiosperms rather than by conifers; morevoer, in the water lagoon), and the finds of imagoes of allochtho- American amber, the rezins were produced by the Cae- nous insects that develop in aquatic environment are salpiniaceae. The other distinctive feature is abundant rare. The presence of the Sieblosiidae represented by stingless bees of the subfamily Meliponinae. Both these the genus Stenolestes, which was also found in Sieblos groups are common in subfossil copals in South Amer- (Nel, 1986), is noteworthy. ica and Africa. Their abundance is a reliable indicator The general characteristic of the Sieblos Fauna was of a very warm climate. Dlussky (1981) reported that made by Martini (1971a) and Willmann (1988). Unfor- the ant fauna from the Sicilian amber closely resembles tunately, a number of taxa described from this locality the extant forest faunas of southern Asia. The absence need revision. The terrestrial insect assemblages are of wasps of the family Ichneumonidae is also notewor- dominated by the taxa that inhabit forest landscapes. thy. The presence of such a thermophilic assemblage in The high proportion of beetles of the family Bupres- the Early Oligocene of Europe is somewhat unex- tidae and the presence of termites indicate a mild (pos- pected. However, certain facts indicate that the Italian sibly warm temperate) climate. It is very difficult to faunas were not of the true tropical type, in contrast to make clear the forest patterns because neither insects the Dominican and Mexican faunas. This inference fol- nor floras have received a detail study. The leaf floral lows from the presence of a few aphids, which are very assemblages from Sieblos include certain evergreen rare in tropics, in a small sample, and the absence of ter- elements (Martini, 1967). The composition of terres- mites of the family Termitidae, which are typical of trial entomofauna does not directly indicate a markedly tropical regions and abundant in the Dominican and warmer climate than that in the terminal Eocene of Mexican amber (some other termites are rather usual in southern England and northern France (Alsace). How- the Italian amber; this supports the supposition of a ever, a direct comparison of these faunas is impossible warm climate). The islands that were located in the because they do not include common taxa of a lower Early Oligocene in the territory of the modern-day Italy rank than families. The seasonal climatic aridity pro- were supposedly faunalally connected with Africa posed by some researchers (Lutz, 1997) was not cor- rather than with the residual Europe. The climate in roborated. The aquatic entomofauna includes certain these regions was a very warm (and supposedly humid) obviously taphonomically autochthonous lake ele- version of the subtropical type that was possibly com- ments; however, the imagoes of aquatic insects appar- parable only to the modern-day climate of humid low ently include the species that developed beyond the mountains of Mexico and Yunnan, rather than with that lake, in which the sedimentation occurred. In particular, of tropical coastal areas. the dragonfly “Petalura” acutipennis Hagen probably The Camoins-les-Bains Insect Fauna is quite differ- belongs to the family Gomphidae mainly associated ent in general pattern and resembles the well-studied with flowing water bodies, rather than to the Petalu- Late Oligocene faunas from southern France (Aix-en- ridae (Nel et al., 1998). The autochthonous assemblage Provence, Cereste, and some other localities). A gen- is unusual, because of the abundance of aquatic weevils eral characteristic of this fauna was presented by “Pissodes” effossus Heyden [actually belonging to Théobald (1937) and Timon-David (1943). Unfortu- Bagoini or to Tanysphyrini rather than to Pissodini nately, this was only partially described, and some taxa (Willmann, 1988)], probably inhabiting floating plants need revision. The significant role of taxa that inhabit (possibly, the Nymphaeaceae), and the presence of well-insolated landscapes (probably open woodlands nymphs of dragonflies of the family Lestidae. These

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S232 POPOV et al. two groups have not been found in comparable quantity cidae, and Oligopithecinae, composed the autochtho- in other localities. However, it is unclear whether or not nous evolutionary lineages. The Macroscelidea this peculiarity of the aquatic entomofauna from Sieb- (Metoldobotes) and the unique family Ptolemaiidae los is mainly determined by the biogeographic individ- (Pantolesta) also represented the Early Oligocene uality of the region, or by the local features of the lake endemic groups of North Africa (Simons, 1995a; Sav- basin. According to Lutz (1997), Sieblos Lake experi- age and Russel, 1983). enced marked fluctuations of the water level and occa- A marsupial that was described as Peratherium afri- sional increases in water salinity. The presence of drag- canum from the Jebel Quatrani Formation actually onflies of the family Sieblosiidae, the only extinct belongs to the endemic genus Quatranitherium and is insect family known from the Oligocene, is noteworthy. assigned to an individual group of the African Peradec- This famity existed to the end of the Neogene, and its tidae that independently developed at least from the range was probably always restricted to western Eur- beginning of the Eocene (Crochet et al., 1992). The asia (the easternmost finds of the Sieblosiidae were in assignment of this form to the genus Peratherium has the Miocene of northern Caucasus). been rejected; therefore, the presence of this animal The Sieblos Fauna does not markedly differ in gen- should not be considered as an argument (Bonis et al., eral pattern from the terminal Eocene Bembridge Marl 1985) in favor of the invasion of North Africa by Euro- Fauna from England and possibly inhabited similar pean mammals. However, available data on other landscapes. groups indicate the presence of the Early Oligocene Data on the Early Oligocene insects of Eastern immigrants of European origin in North Africa. In par- Europe, Caucasus, and the western area of Asia are ticular, the artiodactyls Botriogenys, Brachyodus almost absent. (Anthracotheriidae), Rhagatherium (Haplobunodon- tidae), and Mixtotherium (Mixtotheriidae); creodont Thus, the three major Early Oligocene entomofau- Metasinopa; and turtles of the family Testudinidae nas in Europe are markedly distinct. The Bavarian were recorded in the Early Oligocene of Egypt (Simons Fauna indicates a warm temperate and probably a and Wood, 1968; Savage and Russell, 1983; Bonis rather humid climate and forest landscapes. The south- et al., 1985). At the EoceneÐOligocene boundary, the ern French fauna demonstrates a warm temperate or a Embrithopoda from western Asia or southeastern subtropical summer-dry climate of the Mediterranean Europe probably penetrated Africa via the Sicilian Cor- type and landscapes of the open woodlands and bushes, ridor (Bonis et al., 1985) and gave rise to the highly whereas the Italian Fauna indicates a humid subtropical specialized giant Arsinoitherium. climate and forest landscapes. On the Arabian Peninsula, the Early Oligocene mammals were found in the Ashwaq Formation of Terrestrial Vertebrates Dofar Province (Sultanate of Oman). They included the marsupials Quatranitherium aff. africanum, primates North Africa and Arabia. The Early Oligocene (adapids, tarsioids, propliopithecid anthropoids Moerip- mammal fauna of North Africa generally inherited the ithecus markgrafi, and Oligopithecus rogeri), the pro- Late Eocene composition. This is especially well illus- boscidean Barytherium, embriothopods, hyracoids, the trated by the example of the Qasr el Sagha and Jebel creodont Masrasector ligabuei, and rodents of the fam- Quatrani formations in Egypt (Fayum), where the pro- ily Phiomyidae (Thomas et al., 1988, 1989, 1991; Cro- boscidean Moeritherium and the creodonts Hyaenodon chet et al., 1990, 1992; Gheerbrant et al., 1993, 1995). and Apterodon from the Eocene to the Oligocene and No immigrants of European origin have been found in were represented by closely related species (Holroyd Arabia. In the Early Oligocene, North Africa and the et al., 1996). Rapid development of local faunal ele- Arabian Peninsula probably belonged to the same zoo- ments was still observed. In addition to Moeritherium geographic region: the European elements could pene- and Barytherium, the primitive mastodonts Palaeo- trate North Africa from Western Europe. mastodon and Phiomia (Palaeomastodontidae) appeared among proboscideans. Hyracoids of the family Plio- Western Europe. The beginning of the Oligocene hyracidae represented by a large quantity of genera and in Europe was associated with the so-called Grande species were at the acme in the Early Oligocene. Some Coupure (Great Cutting), the greatest rearrangement of of them probably occupied the ecological niches of the West European Fauna. By that time in Europe, four large Holarctic ungulates; even highly specialized cur- artiodactyl families (Choeropotamidae, Mixtotheri- sorial forms were present. A marked adaptive radiation idae, Xiphodontidae, and Haplobunodontidae) had of endemic rodents of the family Phiomyidae (Phiomys become extinct; the other five families continued to the and Metaphiomys) occurred, and they became widely Oligocene with a sharp decrease in taxonomic diversity distributed in North Africa and Arabia (Fejfar, 1987b; (Dichobunidae, Cebochoeridae, Dacrytheriidae, Ano- Thomas et al., 1989). Thryonomyids represented by the plotheriidae, and Cainotheriidae). The number of gen- genus Gaudeamus deviated from this family. The cre- era and species of palaeotheres (Perissodactyla), which odont Masrasector (Proviverrinae) and numerous dominated the ungulate associations in the Late anthropoid primates, the Parapithecidae, Propliopithe- Eocene, decreased; the Lophiodontidae disappeared.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S233

The diversity of creodonts decreased; miacids, Cynod- theridomyids of the BlainvillimysÐTaeniodus lineage, ictis, and almost all primates disappeared. In the termi- the extinction of the genus Pseudoltinomys, and the nal Eocene, ischyromyoid rodents became extinct; the appearance of Issiodoromys. The second phase (MP24) generic composition of the theridomyoid rodents is associated with immigration of the cricetids Melissi- changed; in particular, the brachyselenodont taxa (pseu- odon and Paracricetodon, and the third phase (MP25) dosciurids) disappeared (Russell and Tobien, 1986). is associated with a new diversification of the Theri- At least 20 mammalian families migrated to Europe domyinae (Archaeomys) and immigration and (or) from Asia at the beginning of the Oligocene, including adaptive radiation of new cricetids and eomyids. the Erinaceidae, Plesiosoricidae, Heterosoricidae, Sori- Artiodactyls were represented by descendants of cidae, Lagomorpha, Cricetidae, Aplodontidae, Sciu- certain archaic lineages (Dichobune, Tapirulus, and ridae, Eomyidae, Castoridae, Felidae, Viverridae, Mus- Diplobune) in association with new cainotheriids telidae, Procyonidae, Ursidae, Entelodontidae, Tayas- (Plesiomeryx and Caenomeryx) and rare Asian immi- suidae, Suidae, Leptomerycidae, Tragulidae, and the grants (Entelodontidae, Suidae, Anthracotheriidae, Rhinocerotoidea, as well as a new assemblage of and Ruminantia). Among perissodactyls, autochtho- anthracotheriid genera, the so-called neoanthracotheres nous palaeotheriids still existed (the last Plagiolophus (Sigé, 1977; Sigé and Vianey-Liaud, 1979). Appar- disappeared in MP23); rhinoceroses and chalicotheres ently, this migration was gradual and included several appeared (in MP23 and MP25, respectively). successive stages occurring in the Rupelian (MP21Ð MP25 zones); however, the majority of the groups The earliest Oligocene mammals of Spain were mentioned above appeared at the onset of the Oli- recorded from the localities of Saintpedor (Ebro) and gocene (MP21 Zone). Among the factors for these Ciruelos (Betic Cordillera). The Saintpedor Fauna events were new continental connections that formed includes the rodents Theridomys aff. aquatilis, Pseudol- between Europe and Asia at the EoceneÐOligocene tinomys gaillardi, Atavocricetodon atavus, and Glira- boundary, the absence of climatic barriers, and the vus aff. priscus; the Palaeotheriidae; and the Ano- higher level of evolutionary dynamism of the immi- plotheriidae (Agustí et al., 1987). In Ciruelos, the grants as compared to the local taxa and, hence, their rodents Blainvillimys cf. langei, Atavocricetodon ex. higher competitive ability (Brunet, 1979). gr. atavus, Eomys sp., and Gliravus sp. were found (Ferriz et al., 1993). Glamys olallensis was a typical The mammalian composition indicates that, in the species of the Gliridae of this level (Freudenthal, 1996). Early Oligocene, the periods of European land consol- In Spain, there are 20 more localities of the Early Oli- idation alternated with the periods of isolation of indi- gocene mammals (especially rich in rodents); they cor- vidual regions; this is seen in the zoogeographic zona- respond to the MP22, MP23, and MP25 zones (Agustí tion of Western Europe. et al., 1987; Cuenca et al., 1992). In the first half of the In southern France (Quercy), some Late Eocene lin- Oligocene, the fauna of the Iberian Peninsula was not eages of marsupials persisted after the Grande Coupure only affected by Asian elements but also occasionally (Remy et al., 1987). At the beginning of the Early Oli- experienced autochtonous development due to tempo- gocene, insectivores were dominated by the Nyctitheri- ral geographical isolation. This is indicated by an idae in association with the gymnura Tetracus nanus. impoverishment of the Iberian fauna, as compared to The Carnivora also markedly changed, Cynodictis dis- other European faunas, and by a large number of appeared, whereas Eusmilus (Felidae), Amphicynodon endemic species among rodents, including the Theri- (Ursidae), and Mustelictis (Mustelidae) appeared. The domyidae, Gliridae, Cricetidae, and Eomyidae (Alva- number of creodont genera decreased down to two rez Sierra et al., 1987a; Daams et al., 1989; Pelaez- (Hyaenodon and Thereutherium), with Hyaenodon Campomanes, 1995; Freudenthal, 1996). being represented by a large number of species; how- The presence of mammalian taxa typical of conti- ever, this was associated with successive immigrations nental Europe in the Early Oligocene of Majorca (Cain- rather than with cladogenesis. Primates became extinct. otheriidae, Anthracotheriidae, Tayassuidae, Tragulidae, Primitive lagomorphs (Desmatolaginae) appeared at Theridomyidae, Pteromyidae, and Cricetidae) indicates the MP21 Zone; however, they were rather rare and soon the periodical faunal contacts of this territory with other disappeared from Western Europe (MP22ÐMP23). European regions (and primarily with the Iberian On the extinction of the Ischyromyidae and the Land). Judging from the findings of anthracotheriids, majority of the Pseudosciuridae and Sciuroidea in the the Balearic Massif was also connected to the CorsicaÐ terminal Eocene, theridomyids differentiated widely Sardinia Block, Kabilia, and Calabria. The mammalian and rapidly. Representatives of this family dominated assemblage was substantially impoverished in taxo- the rodent associations of Western Europe, whereas the nomic composition compared to the continental faunas, groups that appeared in the Oligocene (Aplodontidae, similar to that in the Late Eocene (Bruijn et al., 1979; Cricetidae, and Eomyidae) were minorities. After the Hugueney and Adrover, 1982). The isolation of the Grande Coupure, three phases in the evolution of Early Balearic Land resulted in the appearance of individual Oligocene European rodents are recognized. The first endemic rodents. Thus, Moissenetia paguerensis phase (MP23) is characterized by the cladogenesis of (MP23ÐMP24) has no equivalents among the Oli-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S234 POPOV et al. goceneÐMiocene Gliridae in the structure of the grind- 1987; Gad, 1987). These facts may be explained by the ing surface of teeth and is an example of the diachronic existence of a physical barrier (marine strait) in place of parallelism with the extant genus Eliomys (Hugueney the Rhine Graben. In the German Province, immigrants and Adrover, 1995). Of great interest is the presence of gradually replaced the autochthonous elements. The pseudosciurids of the genus Suevosciurus in the large palaeotheres Palaeotherium and anoplotheres Majorca Fauna that became extinct in the terminal became extinct at the very beginning of MP21; the Eocene everywhere in Europe, except for southern Ger- majority of primitive pseudosciurids and the last many. Thus, we should assume the existence of direct palaeotheres, dichobunids, and anoplotheres disap- faunal connections between the Balearic and German peared in MP23; and the last Suevosciurus disappeared zoochores via the Alpine Land or consider these in MP25 (Heissig, 1987). Certainly, many species and regions as parts (refugia) of an integral Eocene range genera of immigrants also became extinct; however, of Suevosciurus, which became isolated in the Oli- almost all their families persisted to the Late Oligocene gocene. In our opinion, the second assumption is pref- (Aplodontidae, Pteromyidae, Castoridae, Eomyidae, erential (Hugueney and Adrover, 1982), although the Rhinocerotidae, Suidae, etc.). The exception is pro- similarity of the Balearic taxon and the southern Ger- vided by lagomorphs (Shamolagus franconicus) and man species S. ehingensis is also beyond question. entelodontids, which disappeared at the end of MP22 or Approximately in the middle of the Oligocene, rodents at the beginning of MP23. of the family Thryonomyidae, immigrants from North Africa, appeared in Majorca; they are represented in the The easternmost Early Rupelian mammalian fauna Paguera Fauna (MP23ÐMP24) by the endemic genus of the West European type was recorded in northwest- and species Sacaresia moyaeponsi (Hugueney and ern Bohemia, Czechia (Fejfar, 1987a). This fauna Adrover, 1991). This species dominate the Rodentia in includes the marsupial Amphiperatherium sp.; erina- abundance. For the first time, these occurrences reli- ceids cf. Neurogymnurus sp.; talpid cf. Paratalpa sp.; ably indicate the existence of trans-Tethyan migrations heterosoricid Qurcysorex sp.; pseudosciurid Suevosciu- from Africa to Europe in the Early Oligocene. In the rus ehingensis; sciurid Palaeosciurus sp.; cricetids latest Oligocene fauna of Majorca (Sineu, MP25), no Paracricetodon cf. dehmi, Atavocricetodon cf. muri- descendants of the Thryonomyidae were found. These nus, and Pseudocricetodon montalbanensis; eomyid probably yielded to new European immigrants, Eomys cf. zitteli; zapodid cf. Parasminthus sp.; glirid Pseudocricetodon and Paracricetodon (Cricetidae) Gliravus sp.; aplodontid Plesispermophilus angus- and the advanced Theridomyidae of the BlainvillimysÐ tidens; ruminants Pseudogelocus laubei, Bachitherium Archaeomys lineage (Hugueney and Adrover, 1990). cf. curtum, and Lophiomeryx cf. pomeli; cainothere Paroxacron sp.; entelodontid Entelodon cf. antiquus; The Asian elements in the Early Oligocene of anthracotheres Anthracotherium sp. and Elomeryx cf. England (Hampshire Basin; MP21) were represented woodi; suid Propalaeochoerus cf. paronae; carnivores by the anthracotheres Anthracotherium alsaticum, Cephalogale and Pseudocynopsis cf. antiquus; cre- Elomeryx porcinus, and Bothriodon spp.; rhinoceros odont Hyaenodon sp.; and rhinocerotoid Ronzotherium Ronzotherium sp.; entelodont Entelodon magnum; eri- cf. filholi. This assemblage is generally typical of the naceid Tetracus sp.; castorid Steneofiber sp.; cricetid Atavocricetodon atavus; and the eomyid Eomys sp. The MP21 Zone and rather similar to the southern German marsupial Amphiperatherium; anoplotheriid Tapirulus; Fauna. On the other hand, it clearly differs from the and the rodents Pseudoltinomys, Isoptychus, Bran- westerner faunas by the absence of the Theridomyidae; Paracricetodon, Pseudocric- satoglis, and Gliravus were inherited from the Late a very early appearance of etodon Eocene. The Early Oligocene endemic taxa of England , and the Zapodidae; and by unusually large sizes Palaeosciurus, Suevo- were represented by the primate Leptadapis stintoni of representatives of the genera sciurus, Paracricetodon, Atavocricetodon, Gelocus (one of the last in Europe) and the pantolestid Dys- , Lophiomeryx peterna woodi (Hooker, 1987). The Early Rupelian and . These distinctions are possibly mammalian fauna of Belgium (Hoogbutsel) was closely accounted for by paleogeographical and paleoecologi- similar to the English Fauna. cal factors (Fejfar, 1987a). This region was most prob- ably a continuation of the German Province and played The Early Oligocene rodent fauna of southern Ger- the role of an ecotone zone, i.e., the same role that the many markedly differed from those of the western German Province played with reference to the entire localities. Only there, brachyselenodont theridomyids Western Europe. of the genus Pseudosciurus remained (Hartenberger, 1973b). Among other distinctions of the southern Ger- The South German mammalian faunas is similar to the man fauna is the persistence of the pseudosciurid Sue- Rupelian faunas of Austria and Hungary. Anthracothe- vosciurus, the endemic theridomyid lineage Theri- rium magnum, Protaceratherium albigense, Diplobune, domys brachydensÐT. ludensis (from MP21 to MP24, and the endemic rodents Meteomys noskyi and migrated to France in MP25), early appearance of the Pseudotheridomys (Anomegodus) baconicus (Eomy- eomyid Pseudotheridomys, and the extreme rarity of idae), and Heterocricetodon (Alsocricetodon) telonii cainotheres, which were widespread in western and (Cricetidae) were recorded from Hungary (Russell southern provinces (Hartenberger, 1973b; Heissig, et al., 1982).

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S235

Thus, in the Early Oligocene, the faunal distinctions dremotherium); and Pan-Holarctic taxa (carnivores between Western Europe and Asia markedly decreased. Nimravus, Amphicyon, Cephalogale, and Plesictics Of 15 endemic European families of the Late Eocene and creodont Hyaenodon). The Asian taxa also include mammals, only eight existed up to the onset of the Oli- the rodents Parasminthus sp. (Zapodidae) and Yindirt- gocene (with the Mixtotheriidae and Haplobunodon- emys sp. (Ctenodactylidae), which were identified by tidae migrating at the EoceneÐOligocene boundary to Lopatin based on Benara material from the collection Africa and persisting there up to the Early Oligocene). of the Paleontological Institute of the Russian Academy By the end of the Early Oligocene, only two endemic of Sciences. families (Theridomyidae and Cainotheriidae) still At the beginning of the Oligocene, Greece and the existed. By that time, the number of families in com- European area of Turkey (Thrace) was widely inhabited mon with the Asian families had increased almost by the anthracotheres Anthracotherium prealsaticum, threefold. Migrations occurred throughout the Early A. magnum, A. cf. mosvialense, Eomeryx woodi, and Oligocene. After MP20/MP21, the expansion of the E. cf. porcinus. The Early Oligocene rodents from the Asian elements was more gradual and well traced in Turkish Thrace, and especially the Cricetidae, are very each Suevian (Rupelian) mammalian zone. diverse. The Kavakdere locality (MP23ÐMP24) Southeastern Europe and Lesser Caucasus. In includes the pseudosciurid Suevosciurus fraasi; crice- the Early Oligocene, the giant rhinoceroses Indrico- tids Paracricetodon kavakderensis, Trakamys saratji, theriidae first reached southeastern Europe. The earliest Pseudocricetodon cf. moguntiacus orientalis, P. phil- member of the group, ?Urtinotherium sp. was found at ippi, and Kerosinia variabilis; eomyid Eomys sp.; fly- the base of the Mera Beds (Lower Rupelian) in the ing squirrels Oligopetes cf. lophulus and O. aff. vicinity of Cluj, Transylvania (Codrea, 1993; Codrea obtusus; glirids Glis aff. guerbuezi and Bransatoglis and Suraru, 1995). Benaratherium gabuniae was sjeni; castorid Steneofiber sp.; and Ctenodactyloidea described from the later beds (Upper Stampian, MP24 indet. The Kocayarma Fauna (MP25) includes Suevo- Zone); it was more primitive than B. callistrati from the sciurus ehingensis, Paracricetodon kodjayarmensis, Oligocene of Georgia. In addition, the rhinoceros Pseudocricetodon orientalis, P. philippi, Lignitella sue- “Ronzotherium” kochi, anthracothere Anthracothe- mengeni, Edirnella sinani, Eucricetodon aff. dubius, rium magnum, entelodontids Entelodon aff. deguilhemi Heterocricetodon cf. schlosseri, Eomys fahlbuschi, and E. magnum, embrithopod Crivadiatherium mack- Eomys sp., Glis guerbuezi, and Bransatoglis sjeni ennai, and the pantolestid Kochictis centenii were (Ünay and Bruijn, 1987; Ünay-Bayraktar, 1989; found in Romania (Râdulesko et al., 1976; Russell Freudental et al., 1994). Unlike the West European fau- et al., 1882; Codrea, 1993). In Bulgaria (Dragovica), a nas, this assemblage lacked theridomyids and was typical representative of the North Asian (Kazakh) dominated by the Muroidea. The above facts allow one giant rhinoceroses, Indricotherium transouralicum, to assign the East Mediterranean Region and Transcau- was found (Russell et al., 1982). casia to the BalkanÐCaucasian Zoogeographic Prov- The widely known Benara Fauna from Georgia is ince, which was probably isolated from the typical usually dated to the Late Oligocene (Gabunia, 1964; European faunas. One can conclude that in the Early Russell and Zhai, 1987). However, this supposition Oligocene, as in the Late Eocene, the southeastern lacks support from the other taxonomic groups and the Europe, Anatolia, and Transcaucasia were components position in the geological sections support this fact; of the North Asian Subregion of the Asian Zoogeo- moreover, the presence of Brontotheriidae indet., graphic Region. Ardynia (Hyracodontidae), and Hyaenodon dubius Northwestern and Central Asia. The Early Oli- (Creodonta), found in Europe in zones MP21ÐMP23, gocene evolution of Asian mammals is usually divided and Bothriodon cf. velaunus (Anthracotheriidae), into two ages, the Ergilian and the Shandgolian (Rus- recorded from the MP21 Zone, allows for the assign- sell and Zhai, 1987). The Ergilian is correlated with the ment of this fauna to the Early Oligocene (Codrea, European MP21 Mammalian Zone. The Shandgolian is 1993). The generic composition of the fauna is gener- usually correlated with the MP22ÐMP27 mammalian ally consistent with this dating. The Benara Fauna is zones (Vislobokova, 1996) and, consequently, includes characterized by a large proportion of endemic genera a part of the Late Oligocene (the uppermost horizons of and species and includes elements of different origin: the Shand-Gol Formation correspond to the Upper Oli- the true European taxa (rhinoceros Eggysodon and gocene, but this epoch is usually designated the Taben- antracothere Elomeryx); native Asian taxa (chalico- bulukian Age). Dashzeveg (1996b) has recently proposed there Borissiakia; hyracodontid Ardynia; rhinoceros the division of the Shand Gol Formation (= Hsanda-Gol Meschotherium; suiforms Paraentelodon, Hyoboops, Formation) into two members, the Tatal Beds (below and Telmatodon; ruminant Iberomeryx; and the giant the basalt bed that is dated as 31Ð32 Ma) and the Shand rhinoceroses Indricotheriidae); Asian-American taxa Beds (above this bed), with the Shandgolian Fauna (Brontotheriidae, tapirid Colodon, and rodents Cylin- being generally correlated with the MP22 Zone. drodontidae); Pan-Paleoarctic taxa (chalicothere Schizotherium, anthracotheres Bothriodon and Anthra- In Central Asia, faunas from a number of localities cotherium, and ruminants Lophimeryx and Pro- in Mongolia (Ergilin-Dzo, Khoer-Dzan, Dzamyn-Ude,

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S236 POPOV et al. the upper part of the Ergilin-Dzo Formation, or Ergilin Didymoconus (= Tshelkaria, and Archaeomangus) Beds; Shand-Gol, Tatal-Gol, Menkhen-Teg, Ikhe-Arga- (Huang, 1992; Dashzeveg, 1993; 1996b; Spassov and luntu-Nuru, the lower part of the Shand Gol Formation Lange-Badré, 1995; Vislobokova, 1996; Lopatin, 1997a). below the basalt bed dated at 31Ð32 Ma, or the Tatal The giant rhinoceros Indricotherium parvum was Beds) and in northern China (Urtyn Obo, Tsaganbulag replaced by I. transouralicum and the lagomorph Des- formations, etc.) are assigned to the Ergilian Age. This matolagus ardynensis was replaced by D. gobiensis and epoch is generally characterized by the persistence of D. robustus. Ruminants were represented by more certain archaic Eocene mammalian genera, including advanced taxa than in the Ergilian and included Eum- the swamp rhinoceros Gigantamynodon, hyracodontids eryx imbellis, Lophiomeryx cf. mouchelini, and Pseudo- Ardynia and Prohyracodon, deperetellid Teleolophus, meryx gobiensis (Vislobokova et al., 1996). The com- chalicothere Eomoropus, brontothere Embolotherium, mencement of the later stage of the Shandgolian fauna didymoconid Ardynictis, lagomorph Gobiolagus, and development (MP24ÐMP27), which continued to the carnivore Stenoplesictis, as well as by the appearance Late Oligocene, was characterized by the appearance of of a number of new taxa, including the carnivore Nim- large progressive Lophiomeryx of the L. chalantiae ravus; perissodactyls Cadurcodon, Hypsamynodon, type; lagomorph Sinolagomys; and the rodents Tachyo- Symphysorrachis, Ronzotherium, Allocerops, and Indri- ryctoides, Eomys, Pseudotheridomys, and Eomyodon cotherium; suiforms Entelodon and Bothriodon; rumi- (Wand and Ermy, 1991; Vislobokova, 1996). Almost all nants Miomeryx, Eumeryx, and Palaeohypsodontus; the Early Shandgolian mammalian genera and species and rodents Pseudocylindrodon, Eucricetodon, Allos- still persisted. minthus, Sinosminthus, and Heosminthus (Wang, 1985; In Kazakhstan, the Ergilian includes the Upper Wang and Meng, 1986; Dashzeveg, 1993, 1996a, Aksyir Subformation and the Kusto and Tuzkabak for- 1996b; Vislobokova, 1997). Ruminants were repre- mations of the Zaisan Depression (eastern Kazakhstan). sented by the most primitive (for the Oligocene) taxa, including Lophiomeryx angarae, L. gobiae, Miomeryx The Upper Aksyir Formation includes the hedge- cf. altaicus, Gobiomeryx dubius, and Eumeryx culminis hogs Amphechinus sp., Tupaiodon sp., Ictopidium sp., (Vislobokova, 1996; Vislobokova et al., 1996). Fossils Galerix sp., and Galericinae gen.; moles Talpidae gen.; of true Asian leptictids (Ongghonia dashzevegi) has lagomorphs Gobiolagus, Zaissanolagus, and Desmato- been recently discovered in the lower part of the Shand- lagus; didymoconid Ardynictis sp.; ruminant Archae- Gol Formation in the locality of Tatal-Gol (Kellner and omeryx sp.; anthracotheres Brachyodus and Bothri- McKenna, 1996). However, the primitive tooth struc- odon; brontotheres Brontotheriidae indet.; amynodon- ture of Ongghonia indicates that the terminal Pale- tids; and the rodents Symplokeomys zaysanicus oceneÐthe beginning of the Eocene rather than the (Eomyidae); Sinosminthus sp., Allosminthus cf. ernos, Early Oligocene should be considered to be the time of Allosminthus sp., and Heosminthus primiveris (Zapo- the leptictid dispersion in Asia. The pantolestid Gobio- didae); Eucricetodon sp. (Cricetidae); Ardynomys pithecus khan found in the Khoer-Dzan locality glambus and Ardynomys sp. (Cylindrodontidae), and (Dashzeveg and Russell, 1992) indicate the possibility cf. Karakoromys sp. (Ctenodactylidae). Amphibians, of migration of certain European (in origin) mammals turtles, lizards, and snakes are abundant; crocodiles and (Dyspterninae in this case) to Asia in the terminal birds were also found (Shevyreva, 1972; Gabunia, Eocene. 1977; Gabunia and Gabunia, 1987; Emry et al., 1997; Erbajeva and Tyutkova, 1997; Gabounia and Chkhik- The Central Asian faunas of the Shandgolian Age vadze, 1997). are rather abundant in Mongolia and northern China. The Kusto Formation yields the pantolestid Oboia The first stage of the development of the true Shandgo- argillaceus (Gabunia, 1989); swamp rhinoceros Cadur- lian faunas is considered to occur 32 to 29.5 Ma BP and codon ardynensis (= C. tuskabakensis); entelodontid comprises Shand-Gol, Tatal-Gol, and Shunkht in Mon- Entelodon gobiensis (Ermy et al., 1997); hedgehogs golia and Ulantatal, Sanshenggon (Saint Jacques), Tupaiodon sp. and Galericinae gen.; moles Talpinae Shuidonggong and Chanlishan in China; these ages gen.; rodents Cylindrodontidae indet., Cricetidae indet., correspond to the European MP22ÐMP23 zones (Vis- Ctenodactylidae indet., Zapodidae indet., and Eomy- lobokova, 1996). This period is characterized by the idae indet.; anthracotheres Bothriodon sp., cf. Brachyo- extinction of the majority of the Late Eocene Ergilian dus; leptomerycids Archaeomeryx sp., cf. Miomeryx sp., relicts and many true Ergilian taxa, as well as by the and Gobiomeryx dubius; and the cervid cf. Eumeryx sp. appearance of many new taxa (erinaceids Amphechi- (Gabunia, 1977; Russell and Zhai, 1987). nus, Tupaiodon, and Exallerix; heterosoricid Gobisorex; rodents Tsaganomys, Cyclomylus, Anomoemys, Prosci- The assemblages from the Chilikty (= Chelkarnura; urus, Selenomys, Tataromys, Karakoromys, Cricetops, = Indricothere) and the Betpak-Dala formations in Parasminthus, Shamosminthus, and Gobiosminthus; western and central Kazakhstan, as well as the Buran carnivores Amphicticeps, Palaeogale, Plesictis, Palae- and Aktau formations in eastern Kazakhstan, are oprionodon, Proailurus, Amphicynodon, Asiavorator, assigned to the Shandgolian Asian Land Mammal Age. and Stenogale; gelocid Pseudomeryx; bovids Palaeo- The Chelkarnura (Indricothere) Fauna of terrestrial hypsodontus and Hanhaicerus; and the didymoconid vertebrates in central Kazakhstan (Turgai Depression, a

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S237 number of localities in the vicinity of Chelkar-Teniz latter are represented by the marsupial Asiadidelphis Lake) includes the amphibian Pelobatidae gen., turtles tjutkovae; chiropterans cf. Myotis sp.; Tupaiidae gen.?; Planiplastron tatarinovi, Ocadia turgaica, Chrysemys erinaceids Tupaiodon cf. morrisi, Ictopidium cf. tatal- lavrovi, “Trionyx” turgaicus, and “T.” ninae; and var- golensis, Ictopidium sp., Galerix sp., Pseudoneurogym- ious birds and mammals. Rodents are represented by nurus shevyrevae, P. zchickvadzei, Galericinae gen., the castorid Steneofiber kazachstanicus; cylindrodon- Amphechinus cf. rectus, and Amphechinus sp.; talpids tids Ardynomys kazachstanicus, Cyclomilus turgaicus, Uropsilinae gen., Desmaninae? indet., Desmanella sp., and Polinaomys shintuzsaensis; aplodontids Prosciu- Urotrichini indet., Talpinae gen., and Talpidae indet.; rus arboraptus and Plesispermophilus sp.; eomyid heterosoricids Gobisorex cf. kingae and Heterosori- Eomys sp.; zapodid Parasminthus quartus; cricetids cidae gen.; lagomorphs Gobiolagus cf. major, Gobiol- Cricetops aenius, C. dormitor, C. elephantus, Eucrice- agus sp., Procaprolagus sp., Desmatolagus cf. gobien- todon caducus, and E. asiaticus; and the ctenodactiloid sis, D. aff. robustus, and Desmatolagus sp.; carnivore Karakoromys decessus (= Woodomys chelkaris = Ter- cf. Cynodictis sp.; creodont cf. Hyaenodon sp.; didy- rarboreus arcanus: Wang, 1994). Among other small moconid Didymoconus (= Tshelkaria); swamp rhinoc- mammals, there are the lagomorphs Desmatolagus eros Cadurcodon ardynensis (= C. zaissanensis); and gobiensis and Desmatolagus sp. and insectivore Icto- the tragulid Miomeryx sp. (Shevyreva, 1995; Emry et al., pidium sp. In addition, Lopatin (1999) described a new 1995; Lucas and Emry, 1996; Gabounia and Chkhik- galericine species, Neurogymnurus indricotherii, and a vadze, 1997). Rodents (Shevyreva, 1995) include cas- new talpid genus and species, Pseudoparatalpa torids Agnotocastor devius, Steneofiber kazakhstani- shevyrevae (Urotrichini), from the Donguztau locality. cus, S. zaissanensis, S. shevyrevae, and S. aff. shevyre- Large mammals include the creodont Hyaenodon vae; cylindrodontids Cyclomylus mashkovae and dubius; carnivore Cynodictis minor; didymoconids Beatomus gloriadei; cricetids Cricetops collator and Didymoconus rostratus (= Tshelkaria rostrata) and Leidymys asybaevi; and ctenodactyloids Karakoromys D. gromovae; tapiroid ?Colodon sp.; chalicothere dimetron, Muratkhanomys velivolus, M. kulgayniae, Schizotherium turgaicum; hyracodontid Ardynia kaz- M. djanarae, Roborovskia collega, Tataromys raeda, achstanensis; amynodontid Cadurcodon kazakade- and T. boreas. mius; giant rhinoceroses Indricotherium transourali- cum and I. minor; rhinoceroses ?Allacerops turgaica, In the southern area of Eastern Siberia (Olkhon ?Allacerops sp., Aceratherium aralense, and Acerathe- Island, Lake Baikal; Ular’ Formation), the Oligocene rium sp.; entelodontids Entelodon diconodon, E. major, lagomorphs Desmatolagus cf. gobiensis and Leporidae and Entelodon sp.; anthracotheres Brachyodus sp., indet. and rodents Cricetops cf. dormitor, which are Hemimeryx turgaicus, Hyoboops sp., and Anthraco- closely similar to the Mongolian species, are known theriidae gen.; suid Propalaeochoerus sp.; and rumi- (Pokatilov and Nikolaev, 1986). nants Lophiomeryx turgaicus, Gobiomeryx dubius, Thus, the Early Oligocene fauna of large mammals Prodremotherium flerovi, P. longipes, and Eumeryx in North Asia was rather homogenous. Territories from culminis (Flerov and Yanovskaya, 1971; Russell and western Kazakhstan to Ordos in China and even to Zhai, 1987; Bendukidze, 1994, 1997; Kordikova, 1994; Japan were inhabited by the species of perissodactyls Lopatin, 1997a; Tyutkova, 1997). In the opinion of and artiodactyls that were the same, or closely similar, Kordikova (1994), this fauna belongs to at least two to those from Mongolia (Tomida, 1986; Russell and stratigraphic levels of the Lower Oligocene. In the Zhai, 1987; Vislobokova, 1996; Lucas and Ermy, opinion of Shevyreva (1995), this fauna existed later 1996). The most important distinction of the Early Oli- than the Buranian Fauna but slightly earlier than the gocene Kazakhstan Fauna as compared to that from Shandgolian Fauna. Central Asia is the presence of certain groups of small The Aktau Formation (Aktau Mountains, Dzhungar mammals clearly associated with closed and rather Alatau) includes Ardynia kazachstanensis (Russell and humid biotopes and marsupials, gymnures, moles, and Zhai, 1987; Kordikova and Mavrin, 1996), Ceratomor- beavers, which are unknown in Mongolia and northern pha indet., Suiformes indet., Creodonta indet.; the tur- China. The available shared genera of the majority of tles Paraplastromenus cf. mlynarskii and Emydidae small and medium-sized mammals are represented by indet.; and crocodiles Crocodylidae indet. (Kordikova different species. In particular, among didymoconids, and Mavrin, 1996). the genus Didymoconus is represented by D. colgatei The Buranian Fauna (numerous localities in the Zai- (= Tshelkaria robusta) and D. berkeyi in Mongolia and san Depression) is generally dated as the end of the first D. rostratus and D. gromovae in Kazakhstan (Lopatin, half and the beginning of the second half of the Oli- 1997a). Rodents of the genus Ardynomys are repre- gocene (Shevyreva, 1995) and corresponds to the Euro- sented by A. olseni and A. vinogradovi in Mongolia, pean MP22ÐMP23 zones. This fauna includes various A. kazachstanicus in Central Kazakhstan, and by vertebrates: fishes, amphibians Pelobatidae gen. indet., A. glambus in eastern Kazakhstan; the same is typical turtles (Zaisanemys, Chelidropsis, Rafetus, Ergilemys, of the genera Cynodictis, Cyclomylus, Beatomus, Kar- Chrisemys, Plastomenus, Trionyx, Planiplastron, and akoromys, Cricetops, and Tataromys (Yanovskaya, Ocadia), lizards, snakes, and numerous mammals. The 1970; Shevyreva, 1995).

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S238 POPOV et al.

These faunal distinctions are most probably associ- North Asian Subregion ated with ecological and geographical factors, the BalkanÐCaucasian Province lower climatic aridity compared to that in Central Asia Kazakhstan Province and the localization of the Kazakhstan faunas in the coastal areas of two water basins (the lagoonal-lake West Kazakhstan Subprovince system of Turgai and the Aral Sea, in the coastal low- East Kazakhstan Subprovince lands of the Eastern Paratethys, on the one side, and Central Asian Province large Zaisan Lake, on the other side), which were sepa- Afrotropical Kingdom rated by a higher and more arid area. African Region North AfricanÐArabian Province Zoogeographic Zonation At the beginning of the Oligocene, three routes of The disappearance of the Turgai Strait and the con- migration between Asia and Europe were possible: solidation of the major part of European Land at the (1) from northwestern Asia via the Turgai Depression EoceneÐOligocene boundary resulted in a marked and farther along the northern coastal area of the Parat- decrease in the faunal distinctions between the Euro- ethys; (2) from northwestern and central Asia via Asia pean and Asian zoogeographic regions. Nevertheless, Minor and farther along the old tract of an island sys- they remained separate regions. tem that had existed as early as the Late Eocene and After the mass immigration of Asian mammals at separated Paratethys from Tethys; and (3) from western the beginning of the Oligocene, the European fauna and Asia via the continental bridges of the East Mediterra- the individual island and peninsular zoochores again nean Region. The most important area of emigration became isolated. In the Later Rupelian, the provincial probably was the North Asian Biochore. The faunal distinctions of the European theriofauna still persisted exchange between the Iberian Land and North Africa and became even more pronounced. There existed at could occur in the region of Gibraltar at the beginning least four provinces, the EnglishÐFrenchÐBelgian of the Early Oligocene and via Kabilia and the Balearic Province in the northeast; the FrenchÐSwiss Province continental block at the end of the Early Oligocene. and the Iberian Province, which differentiated the latter, in the southwest; and the German Province in Central Europe. Continental bridges were occasionally formed Freshwater Ichthyofauna between the neighboring provinces; however, the pro- Most data on the composition of the Early Oli- portions of endemic taxa indicates their independence. gocene freshwater fish faunas of Europe and Asia were The BalkanÐCaucasian Province was a component obtained from the revision of fossil assemblages of of the North Asian Subregion of the Asian Zoogeo- France, Germany, and Czech (Obrhelová, 1975; Gaud- graphic Region (like that in the Late Eocene). Within ant, 1978a, 1978b, 1978c, 1979a, 1979b, 1979c, 1981a, this subregion, there are two more provinces, the Cen- 1981b, 1982, 1984, 1985a, 1985b, 1987, 1988, 1993) tral Asian Province and the Kazakhstan Province, or in and the material from eastern and central Kazakhstan, a wider sense, the Central Asian Biochore may be con- Far East, Mongolia (Lebedev, 1959; Sytchevskaya, sidered to be an equivalent to the entire Northern Asian 1968, 1973a, 1973b, 1974, 1976, 1981, 1984, 1986, Subregion (Shevyreva, 1995), with the exclusion of the 1989a, 1989b, 1998; Khisarova, 1971, 1974), Turkey BalkanÐCaucasian Province. (Rückert-Ülkümen, 1963), and China (Tang, 1959; Cheng, 1962; Wang et al., 1981; Zhang et al., 1985; Since mammals of the Shandgolian Age of western Chang and Chow, 1986; Chang and Zhou, 1993; Chang and eastern Kazakhstan markedly differ from those of and Chen, 2000; Chang et al., 2001). Central Asia and from each other by the taxa of species rank, these territories may be distinguished as separate Europe. In Europe, the Early Oligocene freshwater zoochores, namely, the subprovinces of the Kazakhstan faunas display further development of the genera inher- Province. ited from the Eocene. These were primarily forms of marine origin, such as the gonorhynchids, chandids, Thus, in the vicinity of the Tethys in the Early Oli- atherinids, percichtiids, gobiids, and members of some gocene, the following zoochores are distinguished Early Paleogene relict families: the Amiidae, Thauma- (Fig. 5). turidae, Umbridae, and the Lepisosteidae. By the onset Holarctic Kingdom of the Oligocene, only a moderate part of the Early Paleogene forms went extinct. This includes the esoci- European Region form Palaeoesox and the double-armored herrings EnglishÐFrenchÐBelgian Province unknown from the succeeding faunal record. The FrenchÐSwiss Province reports on the extinction of the Thaumaturidae by that Iberian Province time (Gaudant, 1993) are invalidated by the finds of the Thaumaturus otoliths in the Early Oligocene of the German Province Hessen Depression (Weiler, 1961, 1973). The persis- Asian Region tence into the European Early Oligocene Freshwater

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S239

50° 15 Kazakhstanian Province Russo-Scandia English-French- Belgian Province E U R 12 O P E Germanian A N 40° R Province E G 14 I French-Swiss 13 O N Province 10 11

Balkan- Central Asian Iberian 8 9 ASIAN 5 7 Caucasian Province Province Province 30° 4 6 3 REGION

Iran-Afghanistan Land

1 2 3 N Arabian 4 o r t h A f r i c a n P r o v i 5 n c e 6 1 0 500 km 2 7 AFRICAN REGION

Fig. 5. Zoogeographic zonation of the Tethyan Region in the Rupelian based on terrestrial vertebrates. Explanations: (1) Land lack- ing vertebrate finds; (2) European Region penetrated by over 20 mammalian families from Asia throughout the Rupelian, but keep- ing its independence and provincial subdivisions: of 15 endemic families, 12 still existed; (3Ð5) Asian Region, extending up to China and Japan, characterized by homogenous fauna with the same species of Rhinoceratoidea and Entelodontidae: (3) Kazakhstanian Province, differing from the Central Asian Province (4) in the presence of small mammals associated with forest, humid biotopes; (5) BalkanÐCaucasian Province, experiencing migration from Asia but being still separated from the European faunas; (6) African Region, North AfricanÐArabian Province, retaining distinctions of the Late Eocene faunas in the composition of proboscideans, hyracoids, subhuman primates, and endemic rodents; (7) the main vertebrate localities. The localities and group of localities con- sidered in the text (1) Fayum Depression; (2) Dofar, Sultanate of Oman; (3) Ciruelos; (4) Ebro; (5) Quercy; (6) Chantras (Greek Thrace); (7) Dragovica, South Bulgaria; (8) Turkish Thrace; (9) Benara, Akhaltsikhe; (10) Szapar and Pustavam, Hungary; (11) Cluj, Transylvania; (12) Hoogbutsel, Belgium; (13) Bohemia; (14) Austria; (15) Chelkar-Teniz, Northern Kazakhstan.

Fauna of the gonorhynchids, amiids, and lepisosteids the cyprinodontid Prolebias, which became wide- (Gaudant, 1978c, 1981a, 1987b; Martini, 1988; Weiler, spread in the European basins by the Middle Stampian 1973), along with further evolution here of umbrids, (Gaudant, 1981b, 1982, 1988). The distribution of the chandids, and gobiids (Gaudant, 1979a, 1979b, 1981a, short-bodied Amphiatlantic pikes of the subgenus 1987a, 1993; Martini, 1965, 1988), point to a consider- Kenoza also occurs in the Middle Stampian, as docu- able degree of continuity of this fauna with reference to mented by the finds of Esox primaevus and E. sp. in the Late Eocene fauna. France and Switzerland, respectively (Gaudant, 1978a, 1979b); this subgenus is now known to widely populate Throughout the Early Oligocene, the European fish the east of North America. fauna showed only gradual changes, which did not essentially affect its structure. At that time, some In the Rupelian lacustrine basins of Europe, the var- Characidae emerged (Gaudant, 1979c). Since they are ious Chandidae, first recorded in the Middle Eocene represented by forms other than those known from the (Gaudant, 1993), continued to develop (And– elkovic′ , Eocene, this fact testifies to a short-term link of the 1989; Gaudant, 1985b, 1987a, 1989). At that time, their European fauna with the African fauna. Another coeval range covered Spain, France, Serbia, Germany, and event is the continuing inhabitation of fresh waters by Romania. In particular, the Rupelian of Serbia (And– el- new marine invaders such as the osmerid Enoploph- kovic′ , 1989) is characterized by a wide species diversi- thalmus (Gaudant, 1985a, 1993). The spreading of this fication of the chandid genus Dapalis (“Smerdis”), form to the southern regions of Europe suggests the which included D. macrurus, D. minutus, D. sieblosen- cooling. At the same time, there is recorded the rise of sis, and D. microcanthus, in association with the cyp-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S240 POPOV et al. rinid ?Palaeoleuciscus (Leuciscus) papyraceus and short-bodied pikes of the subgenus Kenoza, known cyprinodontids (Prolebias goreti). from the Middle Stampian, obviously reached there The invasion of cyprinids into European fresh from Rosso-Scandia. Similarly, the rare Leuciscinae waters started as late as the Early Oligocene. This event belonging to the genera Palaeoleuciscus and Varchos- is evidenced by the occurrence of “Chondrostoma” tichthys (with the earliest finds from the Stampian of France: Gaudant, 1977, 1984) and problematic Barbi- elongata Kramb. in the Lower Oligocene beds of Bohe- – ′ mia (Gaudant, 1993), along with the first appearance of nae (Rupelian of Slovenia: And elkovic , 1989) came unquestionable Asiatic immigrants, Palaeoleuciscus from Asia; and the characids came from Africa. As and Varhostichthys (V. cf. brevis), in the European noted above, previous reports on the appearance of cyp- freshwater assemblages of the LowerÐMiddle Stam- rinids in the PaleoceneÐEocene of Europe have been pian (Gaudant, 1977, 1979b, 1984), and also by the recently disproved. The earliest known cyprinid records reports on the problematic Barbinae (?Barbus sp.: fall on the Eocene of China (Wang et al., 1981), eastern And– elkovic′ , 1989) in the Rupelian of Slovenia. Kazakhstan (Sytchevskaya, 1984, 1986), and Turkey (Rückert-Ülkümen, 1963). Former reports on the earliest cyprinid finds from the PaleoceneÐEocene of Europe proved to be errone- Northern Asia. The bulk of data on the composition ous. In particular, the genus Chela known from isolated and pattern of the freshwater fish fauna of the Early Oli- pharyngeal teeth is not a cyprinid (Patterson, 1975); the gocene of northern Asia was obtained from Kazakh- same was demonstrated for “Blicca” croydonensis stan. For the Late Eocene to the Early Oligocene, the White from the Lower Eocene of England on the most informative fossil-bearing levels (with respect to strength of its scale structure (Gaudant, 1993). Like- both the fauna and flora) have been detected in eastern wise, isolated teeth from the Priabonian (Late Eocene) Kazakhstan. On the evidence from the plant fossils and of Isle of Wight, once attributed to the cyprinids Phyl- mammals, it is usually believed that the Early Oli- lodens and Diplodens (Rutte and van de Weerd, 1980), gocene is covered here by the upper part of the Aksyir- as well as teeth from the Eocene of Turkey designated skaya Formation, and the Kustovskaya and Buranskaya as the cyprinid Caryophyllodens (Rutte and Becker- formations. According to an alternative viewpoint (Tol- Platen, 1980), in fact, belong in fact to rodents. stikova, 1976, 1980), the EoceneÐOligocene boundary in this section occupies a higher position. Cyprinids gradually progressed in their role through When accepting the predominant dating of the the OligoceneÐMiocene faunal succession of Europe, above units, followed in the present volume, one should in parallel with spreading of the long-bodied north Asi- conclude that in northern Asia the earliest Oligocene atic pikes. At the same time, most of former autochtho- (judging by data from the Upper Aksyirskaya and Kus- nous Paleogene dominants came to decline and disap- tovskaya formations) was characterized by the further peared. Among them, the last amiids, lepisosteids, and evolution of the Late Eocene dominants. Specifically, thaumaturids are still traceable by the scales and these were represented by amiids (genus Cyclurus with otoliths (Weiler, 1973), and the gonorhynchids by the a number of species), hiodontids (subgenus Gobihi- skeletal remains (Schraft, 1979), until the Early odon), and a great variety of catostomids inherited from Miocene. By contrast, the umbrids attained the peak of the Late Eocene: Amyzon, Cycleptus, Carpiodes, their abundance by that time (Gaudant, 1979b; Martini, Erimyzon, Minytrema, Xyrauchen, Moxostoma, and 1988; Weiler, 1973) and survived into the present Euro- Catostomus.10 The percichthids were represented by pean Fish Fauna. In general, the Early Oligocene Euro- the endemic genus Kiinkerishia (K. kazakhstanica). pean Freshwater Fish Fauna remained rather poor in The cyprinids, with the earliest local record in the Early terms of diversity. Most of its localities yield impover- ished assemblages comprising a few forms dominated 10It is surprising that, in spite of unquestionable evidence of wide as a rule by the derivatives of marine families, such as occurrence of catostomids in the Eocene–Oligocene fish commu- the Gobiidae, Chandidae, Cyprinodontidae, Moron- nities of northern Asia (Hussakof, 1932; Sytchevskaya, 1986), idae, and Osmeridae that penetrated freshwater bodies repeated attempts were made to disprove this fact in a quite arbi- trary way, without any reexamination of the fossils in question (Gaudant, 1977, 1981a, 1985a). These were accompa- (Nelson, 1949; Smith, 1992; Yakovlev, 1961). In reality, the abun- nied by a small admixture of the autochtonous relicts dance of catostomids in the continental EoceneÐOligocene of and of immigrants from adjacent land areas. eastern Kazakhstan is proved beyond any doubt by the mass finds of their most diagnosable remains, i.e., the pharyngeal teeth, The primarily freshwater forms belonging to the which in turn are accompanied by numerous fragments of typical Umbridae, Esocidae, Characidae, Cyprinidae, Thau- skull bones, vertebrae, and scales. To this end, it is worth noting maturidae, Amiidae, and Lepisosteidae were scarce and that, in the samples obtained from washing of the bone-bearing rocks of the Aksyirskaya and Kustovskaya formations, the few in number. Substantially, the freshwater ichthyo- catostomid pharyngeal teeth by far outnumber all other remains fauna of the European Early Oligocene remained (in contrast to the samples from the succeeding Buranskaya For- formed by the insular dwellers and had very restricted mation, in which the material is dominated by the cyprinid pha- links to other land regions from where some new ele- ryngeal teeth). The reality of a broad dispersal of catostomid fishes in the Paleogene of northern Asia is further corroborated by ments might occasionally have come. Such faunal con- the finds of their articulated skeletons in the EoceneÐOligocene of tacts were quite episodic and resulted in the invasion of the Maritime Province (Sytchevskaya, 1986) and in the Eocene of a few natively freshwater newcomers. For instance, the China (Chang et al., 2001).

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S241

Eocene (Obailinskaya Formation), contained Parabar- In China, the paucity of data on the Oligocene fish bus sp. and still retained their status of a scarce and fauna complicates very much the analysis of faunal poorly diversified group (Sytchevskaya, 1986). events of that time. The cyprinids, known from there since the Eocene (Wang et al., 1981) continued to Additional data are provided by the Oligocene ich- develop in this area during the Early or Middle Oli- thyofauna of Mongolia (yielded by the Ergiliin-Dzo gocene. In eastern China, these were dominated by the Formation, Ergiliin Member, Zone MP21). This fauna subfamily Leuciscinae (Zhang et al., 1985). The Chi- was impoverished and included scarce amiids (Cyclu- nese catostomids, also showing the earliest record in rus sp.), cyprinids (Rutilus sp.), catostomids (Catosto- the Eocene (Amyzon hunanensis), so far remain midae gen. indet.), and bagrids (Gobibagrus hoerdzan- unknown in the Oligocene (Chang et al., 2001). icus Sytch.). The fish fauna of the Maritime Province, similar in age, included catostomids (Vasnetzovia In sum, compared with the run of events in the Early artemica Sytch.) and cyprinids of the subfamily Oligocene of Europe, the contemporaneous history of Gobioninae (Sytchevskaya, 1986). the freshwater fish fauna of northern Asia reveals a much more marked transformation. In the most spec- Compared to these features, the younger fauna of tacular way, it is expressed in the faunal turnover the Buranskaya Formation of eastern Kazakhstan (involving both fishes and accompanying organisms) (dated on mammals as the end of the first half, or begin- seen in the transition from the Kustovskian to Buran- ning of the second half of the Early Oligocene, zones skian time in eastern Kazakhstan. In contrast to the Late MP22ÐMP23?), displays a quite different pattern. It is Aksyirskian and Kustovskian assemblages (associated almost devoid of amiids and catostomids, which had with the Ergilian Mammalian Fauna of Zone MP21), dominated the preceding Paleogene communities which retained a rich set of the catostomid taxa typical (Sytchevskaya 1986, 1998). Specifically, the diversity of the Paleogene Amphipacific Ichthyofauna, the of catostomids was reduced to a single genus, with their Buranskian community demonstrates a sharp switch finds (as well as these of amiids) becoming exceedingly toward the domination of cyprinid fishes that later ulti- scarce, whereas the hiodontids disappear at all. On the mately conquered the fresh waters of northern Eurasia. other hand, the cyprinid fishes for the first time attained the role of dominating group and showed a marked Analysis of Biogeographic Structure. Of the stud- increase in both diversity and abundance. These com- ies that most closely touched upon the history of dis- prise the Leuciscinae (Rutilus sp., Tribolodon sp., and persal of freshwater fishes in Eurasia, one should men- Zissanotinca cristidens), Barbinae (Parabarbus sp.), tion at first the works by Yakovlev (1961, 1964) who and Schizothoracinae (Eodiptychus longidens and analyzed the Neogene history of the Holarctic ichthyo- Schizothorax sp.). Another new dominants include the fauna. Demonstrating the lack of zoogeographic integ- Esocidae (Esox sp.), Siluridae, and Percidae. Similar rity of Holarctic in the Neogene, he believed this con- assemblages occur in Central Kazakhstan and Fore- dition to be inherited from the preceding epoch and Aral Region (in Kenderlykskaya and Chelkarnurin- stated on these grounds that the exchange between the skaya formations respectively); they also yield the Eso- Eurasian and North American freshwater fishes was cidae (Esox aralensis Sytch.) and Cyprinidae (Rutilus sp., much complicated during the whole Cenozoic. Based Tribolodon sp., Palaeotinca turgaica, Zaissanotinca on comparison of the Neogene families recorded from cristidens, and Parabarbus mynsayensis) (Sytchev- these two continents, Yakovlev concluded that in the skaya, 1986, 1998). Cenozoic the faunal exchange between them took place only twice: in the Paleogene, via the North Atlantic This dramatic faunal turnover observed in the tran- land, and in the Pleistocene, across the Bering Bridge. sition from the Kustovskian to Buranskian time evi- However, the extensive studies of the Paleogene ichthy- dently reflects the large-scale changes in the geological ofauna of northern Asia (Chang and Chow, 1986; history, climate, and paleogeography of northern Eur- Chang and Chen, 2000; Sytchevskaya, 1986) actually asia during the Early Oligocene. The approach of early revealed its close similarity to the coeval fish communi- phase of the Alpine orogeny obviously affected and ties of North America. With further progress of such destabilized the regular hydrological regime that had studies, the unity of both discussed Paleogene biotas been established in the north Asiatic freshwater basins becomes more and more evident, which led us to distin- during the Eocene. The new unstable conditions in guish them as a single biogeographic unit, the Amphipa- aquatic environment, perhaps devoid of clearly cific Province (Sytchevskaya, 1986, 1998, 2001). expressed cyclicity, gave the advantage to more eury- biontic fish groups, primarily the cyprinids, allowing However, the arguments used by various authors to them to oust and replace the Early Paleogene domi- explain the reasons for the similarity of the Amphipa- nants, such as the Catostomidae, Hiodontidae, and cific ichthyofaunas much differ. The most common is Amiidae. These changes in the pattern of the fish fauna, the assumption that the Amphipacific fish groups along with changes in the accompanying freshwater known in Asia had immigrated to there via different mollusk fauna, also indicate the relative cooling during routes from Western Nearctic. Chang and Chow (1986), the Buranskian Time (Tolstikova, 1976, 1980; Sytchev- Chang and Zhou (1993), and Chang and Chen (2000) skaya, 1986). suggest the spreading of some of these groups during

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S242 POPOV et al. the Eocene across a broad land connection between sitive to the mudding and pollution of the aquatic envi- Asia and North America provided by the Bering ronment (Scott and Crossman, 1973). Bridge. In northern Asia, the onset of tectonic activity dur- Grande (1994) proposed another way of transpacific ing EoceneÐOligocene transition resulted in the break- migration, based on the so-called Pacifica hypothesis, down of the Kazakh Plain and adjacent areas (Gross- which implies the existence in South Pacific of an early geim, 1957). This in turn caused the break of the stable continent, whose disintegrated fragments later drifted hydrologic regime in the local basins, so that they to, and finally collided with, the Pacific rims. became less suitable for the existence of catostomids. In contrast to these hypotheses based on the idea of The intensification of irregular fluvial erosion led to an transpacific migrations, Sytchevskaya (1986, 1998, increase in the transport of the terrigenous material and, 2001) put forward the above-mentioned concept of the in consequence, to occasional mudding of water and the integral Amphipacific Province, according to which appearance of swamps and ephemeral shallow pools. western Nearctic and northern Asia had a common All of these changes gave the advantage to eurybiontic biota and developed as a single biogeographic unit from cyprinids and stimulated their rapid ecological expan- the Late Cretaceous to Early Paleogene. During the sion, including the rise of forms adjusted to life in shal- Early Paleogene, the Amphipacific Province remained low, muddy, and poorly oxygenated water setting with isolated from Europe by the West Siberian Sea and dense vegetation. It seems most likely that the replace- from the Eastern Nearctic Realm by the epicontinental ment of the Asiatic catostomid dominants by cyprinids submeridional Nearctic Gulf, which is fully consistent that occurred in the Oligocene basically proceeded with the available paleogeographic data. under such conditions. As indirect evidence of this sce- Disintegration of the Amphipacific Province, as well nario of faunal events, it may be pointed out that the as the Amphiatlantic Province (which is presumed to only Recent survivor of Asiatic catostomids, the Chi- have embraced Europe and Eastern Nearctic: Sytchev- nese genus Myxocyprinus, which is most likely of the skaya, 1986, 1998, 2001), proceeded during the Oli- autochtonous origin, dwells in exceedingly muddy gocene and was largely caused by the paleogeographic waters of the Yangtze River basin. rearrangement of the northern landmasses. This In spite of the removal of marine barrier in the Tur- resulted in a gradual transition to a new, Neogene pat- gai area (western Kazakhstan), which had prevented tern of biogeographic differentiation of the freshwater free dispersal of the natively freshwater fishes in Eur- ichthyofauna. The decisive paleogeographic changes asia during the Early Paleogene, the integration of the involved in this process may be summarized as follows: freshwater ichthyofaunas of Europe and northern Asia (1) Integration of Europe and northern Asia (via still remained much hampered during the Oligocene Turgai Bridge); and partially Miocene. As is seen from the paleontological (2) Isolation of Asia from North America; data, the exchange between the fish faunas of both conti- (3) Broad unification of the western and eastern nents progressed rather slowly, so that the high degree of parts of Nearctic (North America); consistency in their systematic composition was most (4) Break of the land connection between North likely attained as late as the end of the Neogene. America and Europe. In northern Asia, the breakdown of the previous bio- Of these four events, the first two most directly geographic realm known as the Amphipacific Province underlay the disintegration of the Amphipacific Prov- was accompanied by the rise of the new dominants: the ince and then vanishing of its western (Asiatic) part, Esocidae, Cyprinidae, Siluridae, and Percidae. This with the spreading there of the newly arisen fauna of the event fell on the EoceneÐOligocene transition or, fol- Palearctic type. The first event, i.e., the integration of lowing other datings, on the Early Oligocene. At the Europe and Asia, is thought to fall on the EoceneÐOli- same epoch, these new dominants made the first steps gocene boundary (or on the Early Oligocene, following in their spreading into Europe across the Turgai Bridge some alternative datings), taking into account both the and along the southern coast of the Paratethys. This lat- geological data and the most probable time of replace- ter route might have been followed by some Leucisci- ment of the dominants within the north Asiatic cyprini- nae, judging by their fragmentary finds in the Eocene forms. and Oligocene of Turkey. (The same way could equally The reasons for which the Asiatic catostomid fishes have been used by some European groups, such as the became ousted by cyprinids in the Late Paleogene are Cyprinodontidae and Gobiidae, for counter migration of a special interest. The cyprinids evidently originated to Asia.) The earliest cyprinids (Leuciscinae) in Europe in Asia, where they were first recorded in the Eocene. are recorded immediately after that time, in the Middle Ecologically, they are rather extensively diversified and Stampian (Gaudant, 1977, 1979b, 1984). so pertain to the eurybiontic groups. By contrast, the To summarize, it may be stressed again that Early catostomids, judging by the biology of their Recent Oligocene rearrangement of the freshwater fish fauna in North American members, were mainly the stenobion- Asia was much more clearly expressed than in Europe. tic benthophages that populated the lakes and streams In Asia, this resulted in the decline of amiids and nearly with hard, usually stony, bottom and clean, well-oxy- entire extinction of catostomids and hiodontids as early genated water. This mode of life makes them very sen- as the Middle Oligocene. By contrast, in Europe, the

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S243 extinction of the local Early Paleogene dominants, such regimes. Nevertheless, a comparison of these determi- as amiids, thaumaturids, and gonorhynchids, was more nations with those made for the Late Eocene reliably gradual, and it was completed only in the Miocene. indicates that at the EoceneÐOligocene boundary the It seems very likely that these events were preceded by average annual temperatures dropped by over 4Ð6°C in the break of the Bering Land connection between Asia midlatitudes, mainly due to a decrease in the average and North America, because the Oligocene expansion winter temperatures. of cyprinids in Asia did not result in their broad pene- Less marked changes occurred in the Tethyan tration to Nearctic. Realm, where environments that are closely similar to The Early Oligocene faunal turnover, which closed those in the late Eocene still persisted in the shelf area. the existence of the former Amphipacific Province and Even the north European shelf area was inhabited by the succeeding expansion of the new North Asiatic Fish reef-building corals, numerous tropical and subtropical Fauna to Europe (with the integration of both Palearctic benthic genera and species inherited from the Eocene, continents on the close of the marine Turgai Strait), led as well as by fairly complete planktonic assemblages. to a rise of a new biogeographic unit, the Palearctic These changes resulted in the poorer taxonomic com- Subregion. The core of the Palearctic Freshwater Fish position of the majority of groups and in the appearance Fauna, dominated by esocids, cyprinids, silurids, and of individual species, which, in the Priabonian, were percids, began to form in northern Asia during the Early restricted to the northern regions of Europe. In the early Oligocene; its composition was later extended to incor- Oligocene, the European coast of the Mediterranean porate some European elements. probably was a part of a tropicalÐsubtropical transition It should be noted that, by their ancestry, the above- zone, where the water was constantly heated to a tem- listed principal dominants of the new North Asiatic perature of 18Ð20°C, which is required for a normal Fauna, which appeared in the course of the “Oligocene development of reef-building corals. Revolution,” are autochthonous to the Amphipacific The terrestrial vegetation of this coastal area area. In particular, cyprinids and silurids are known changed its character. The microphyllous evergreen from this area since the Eocene (China); esocids, since, plants indicate an advance of arid environments, and the Cretaceous (northern Nearctic); and the percids, the appearance of certain Arcto-Tertiary forms indi- since the Eocene. As mentioned above (with respect to cates a slight climatic cooling. Floristic data indicate cyprinids), the discussed faunal changes did not affect that the climate of the southern slopes of the Lesser the eastern part of the Amphipacific Province, North Caucasus was of a subtropical type without any tropical America, which seems to suggest that by the end of the indications. Eocene its land connection with Asia had already been lost. In connection with this, no broad invasion of Asi- SouthÐCentral EuropeanÐLesser Caucasian atic cyprinids ever took place to America, such that Coastal Area of the Paratethys was characterized by until now this group plays only a subordinate role in the a marked impoverishment of the benthic and planktonic North American Fish Fauna. On the other hand, some assemblages in the early Oligocene and by the disap- Amphipacific dominants, such as the amiids, hiodon- pearance of the most thermophilic taxa before their tids and catostomids, have survived into the North evolutionary extinction. The colonial corals did not American Fish Fauna until present, due to which this build any reefs. As during the Eocene, this coastal area fauna still retains its relict Amphipacific (Paleogene) belonged to a subtropical zone. The plant fossils indi- pattern in many respects. cate the existence of luxuriant mesophilic forests along the central European coastline. For the Kiscel Flora, the As a whole, on the evidence from freshwater fishes, average annual temperatures were 21Ð23°C, the aver- the Early Oligocene biogeographic structure of Holarc- age winter temperatures were 15°C, and the average

tic is outlined as follows: ° summer temperatures were^ 23 C. The average annual Holarctic Region temperatures for the Brez ani Flora were estimated at 20Ð23°C (Palamarev, 1967); those for the Sotzka Flora,

Palearctic Subregion ° at 17.6 C (Unger, 1861);^ and the mean annual precipi- European Province tation for the Brez ani and Kudratice floras, at 1200– Asiatic Province 1800 and 700Ð1300 mm, respectively. The majority of Nearctic Subregion heavy rainfalls occurred in summer (Andreansky, 1964). North Sea Basin and the northern coastal area of the Paratethys were most significantly affected by the LATITUDINAL CLIMATIC ZONATION global EoceneÐOligocene climatic cooling, as well as OF THE EARLY OLIGOCENE by regional rearrangements of the paleogeographic Even those numerous determinations of the early connections and currents. The character of the distribu- Oligocene climatic parameters that are based on similar tion and migration of benthos reliably indicate that the initial data (predominantly on the floras of western and Eastern Paratethys was most strongly affected at the central Europe) show a wide scatter, because they use beginning of the Oligocene by the water exchange with different methods for reconstructing the paleoclimatic the North Sea Basin. The taxonomic diversity of

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S244 POPOV et al. benthic fauna sharply decreased, corals and nummulit- and determined, to a large extent, the character of the ids were almost absent, and the molluskan assemblages global changes at the EoceneÐOligocene boundary. The were dominated by boreal taxa. In planktonic assem- trigger mechanism for large-scale paleogeographic, cli- blages, the majority of warm-water zonal species of matic, and biotic events during the late EoceneÐearly planktonic foraminifers and nannoplankton were also Oligocene was a tectonic rearrangement that resulted in absent. Thus, the composition of marine biota indicates changes in the patterns of oceanic and climatic circula- that in the early Oligocene this area was assignable to tions, which conspicuously affected the composition the boreal low-latitudinal zone. Its temperature param- and geographic distribution of biota. eters may be estimated as follows: the water remained Tectonics. A collision of the African and Arabian heated to 15Ð18°C for three or four months, whereas in plates with the European Plate was the determining fac- winter, it could experience long-term cooling to tem- tor for changes in the Tethyan Realm. The approach of peratures below 10°C. these plates to each other resulted in a progressive clo- The floral composition, i.e., the joint occurrence of sure and separation of Tethyan relics, reduced connec- evergreen and deciduous taxa, allows the inclusion of tion between the Atlantic and Indian oceans, higher the coastal area of northwestern and eastern Europe relief, and in the formation of the land bridge that sep- into the ecotone SubtropicalÐBoreal Zone. Its more arated the Paratethyan basins. westerly regions were dominated by subtropical floras, The results of the collision are also traceable at a whereas farther to the east, these were replaced by flo- regional level throughout the Alpine Zone. In the ras of a moderately warm climate. The climatic param- Alpine-Carpathian Region, the collision resulted in a eters of this zone are estimated as follows: the Hasel- large-scale horizontal displacement, decrease in the bach Flora existed at average annual temperatures of basin area, and rearrangement of paleogeographical 10Ð15.5°C, average winter temperatures of 2Ð4°C, connections: shallowing and closure of the Slovenian average summer temperatures of 20Ð22°C, and the aver- Corridor, a strait between the Alps and the Dinarides. age annual precipitation ranged from 1000 to 3000 mm The Pyrenean Orogeny resulted in drying and folding (Mai and Walter, 1978). For comparison, the climatic of the South CarpathianÐBalkanian area of the Paleo- parameters for the late Eocene Zéitz Flora located in the gene basin and in the formation of depressions that same Weisse-Elster Depression were as follows: the were compensated for by land sediments. average annual temperatures were 15Ð20°C, the aver- age winter temperatures were 6Ð13°C, the average At the beginning of the late Eocene, the system of summer temperatures were 15Ð23°C, and the average the Balkanids, Pontides, Lesser Caucasus, Elburz, and annual precipitation was over 2000 mm (Mai and Kopet Dagh still represented an archipelago. Subse- Walter, 1983). The Nerhau Flora was characterized by quently, this system occasionally formed a continuous the average annual temperatures of 13Ð16.5°C, average land massif, which already in the late Eocene served as winter temperatures of 6°C, average summer tempera- a land bridge to southeastern Europe for large Asian tures of 25°C, and the average annual precipitation over vertebrates and separated the Eastern Paratethys from 2000 mm. The subarid climatic zone became more the Tethyan Realm. In the Oligocene Paratethys, the sea humid compared to that in the Eocene and was displaced bottom became much more differentiated to form deep farther south, to the CrimeanÐCaucasian Region. depressions not compensated for by sedimentation. The temperate climatic zone became markedly Large land massifs also formed in that epoch farther wider. Its southern border was displaced farther south. south (Eastern Taurus and the Ararat Massif). This This zone covered the entire East European Platform, resulted in the closure of water circulation in the Med- most of the Turan Plate, and the territory of northern iterranean, and in the cessation of biogeographic isola- and central Kazakhstan and Western Siberia. This zone tion of benthic fauna that inhabited the North African occupied the northern, most humid area of the late coastal area. Eocene subtropical zone. The Kyzyltobe Flora, the The activation of the Tian Shan zone first resulted in most ancient of the early Oligocene Kazakhstan floras, the loss of connections and drying of the Tarim Basin developed in climatic conditions closely similar to and subsequently in the closure of the Ferghana and those in the modern-day Colchis Refugium with aver- Tajik basins and accumulation of continental deposits age January temperatures of 2°C, average July temper- there. Previously, at the end of the middle Eocene, the atures of 20°C, and average annual precipitation of continental environments had already formed in the 1500Ð2000 mm (Rayushkina, 1979). Central Pamir Basin. Eustatic motions. Three transgressive phases of the Late Eocene (Beloglinian) Basin have been recorded in MAJOR EVENTS IN THE LATE the sections of the northern passive margin of the EOCENEÐEARLY OLIGOCENE Paleo-Paratethys where marine facies are wedged in The studied region, extending from northern Africa land sediments (Geologicheskie i bioticheskie..., 1998, and the Atlantic coastal area of Europe to Central Asia, p. 197). The first transgressive phase has been dated as covers a considerable part of the Earth’s Northern nannoplanktonic zone NP18; the second phase, as Hemisphere. The events that occurred there reflected zones NP19 and NP20. At the boundary between these

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S245 phases, a short-term decrease in the sea level (by 50 m, (coral and nummulitid) carbonates were widespread on or more) occurred. This was accompanied by climatic the shelf. cooling and anoxic events, which are indicated by the In the Northern Peri-Tethys of the first half of the sharp increase in the abundance of Uvigerina (Marzuk, Priabonian, the eustatic sea level was high and the 1992). The sea level was even higher during the second water salinity was still closely similar to that of the transgression, the transgressive sediments of which ocean. The entire Alpine-Carpathian Basin and the overlie more ancient beds (the Mandrikovka Beds in southern Transcaucasian shelf area of the Caucasian Ukraine, the Balyklei Beds in the Volga Region, and the Basin experienced strong effects of warm Tethyan Sumsar Beds in Tajikistan) (Muzylev et al., 1996). water, whereas no such effects are known in the more A marked sea advance and a coarser-grained com- easterly regions, the Fore-Carpathian shelf area of the position of the sediments is indicated in the northern basin inhabited by endemic fauna. In basin facies, fine shelf area, starting from the second half of the Priabon- biogenous-carbonate silt dominated to be replaced by ian (Bolivina antegressa Zone). The peak of sea regres- terrigenous sediments only in the shallow shelf zone. sion is observable in the EoceneÐOligocene boundary The water exchange via the Turgay Strait and the beds, where shallow-water benthic fossils and leaf flora cyclonic current in the West Siberian Basin served as a were even recorded from deepwater marly facies. radiator that heated the adjacent coastal areas (Geolo- These fossils indicate the appearance of islands in the gicheskie i bioticheskie..., 1998). western area of the Greater Caucasus (Akhmetiev, As a result of the separation of the Paleo-Paratethys 1995). In the northern Aral Sea coastal area and Turgay, in the second half of the Priabonian, the entire Parat- the sea level decreased by over 60Ð80 m, as indicated ethys came under progressively stronger influence of by the depths of ravines that eroded the underlying the North Sea. The only exception was the southern Chegan Formation. The denudation in the Kyzyl Kum (Transylvanian) area of the Carpathian shelf and the Desert is estimated at 100Ð150 m in depth (Geolo- southern (Turkish) coastal area of the Euxine Basin, gicheskie i bioticheskie..., 1996, 1998). Even in the cen- where the benthic fauna indicates that the Tethyan tral areas of the CaucasusÐKopet Dagh Basin still occu- effects persisted into the early Oligocene. Marked pied by sea, breaks in sedimentation and erosive forms deepening of basins, the appearance of isolated deep- are observable in the EoceneÐOligocene boundary water basins, partially blocked connections with the beds. These are often indicated in argillaceous facies world ocean, an increase of the temperature gradient only by the active redeposition of fossils of the Eocene with latitude, and an active inflow of fresh water all microfauna and flora. resulted in the formation of a distinct temperature and salinity stratification. The appearance of stagnation Although the scale of the subsequent sea transgres- conditions and the accumulation of undecomposed sion in the first half of the early Oligocene of the Parat- biogenous material in bottom sediments resulted in the ethyan region fell short of the scale of the late Eocene formation of anoxic facies very typical of the Car- transgression, this transgression involved fairly large pathian (Menilithic) Basin, as well as of the CaucasusÐ areas, including the DnieperÐDonets Depression, cen- Kopet Dagh (Maykopian) Basin. The productivity of tal areas of the Volga Region, the northern area of the calcareous nannoplankton and planktonic foraminifers, Turan Plate, the Turgay Strait, as well as the major area which in the Eocene were rock-forming organisms, of the West Siberian Basin, the Kyzyl Kum area of the sharply decreased. Turanian Basin (Fig. 1), and possibly the FerghanaÐ Tajik and even the Tarim basins. The subsequent Solenovian closure of the basin resulted in the first brackishing of the Paratethys, The subsequent Solenovian Closure was determined changes in the character of sediments, extinction of by tectonic rather than eustatic factors. It was some- marine fauna, and development of the individual times accompanied by sea regressions and basin shal- endemic brackish-water fauna and microflora to restore lowing (in the DnieperÐDonets Depression, Volga for a short period of time the favorable gas exchange Region, and Ciscaucasia) sometimes by deposition of regimes and the biogenous-carbonate character of sed- transgressive sediments and erosion of underlying iments (Polbian Beds). beds. The Late Solenovian was undoubtedly an epoch Climate. Despite the very warm climate of the Mid- of sea regressions. dle Cenozoic, the latitudinal-climatic zonatation was Hydrology and sedimentology. The normal marine even in the Late Eocene defined sufficiently well to conditions and the warm current that went from the determine its biogeographic subdivision into provinces southeast (Indian Ocean) and was inherited from the (Popov et al., 2001). Middle Eocene still persisted in the Priabonian in the A marked, although gradual, climatic cooling Tethyan Realm. The distribution of biota in the second started in the second half of the late Eocene, with the half of the PriabonianÐEarly Rupelian indicates an peak of the temperature drop recorded at the EoceneÐ increased separation of this region and the formation of Oligocene boundary. In the northern Peri-Tethys, this the Circum-Mediterranean Current (Fig. 2). Since the global cooling became even more marked because of supply of terrigenous material was limited, biogenic the eustatic sea regression and paleogeographic rear-

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S246 POPOV et al. rangements that partially blocked Tethyan connections ids became extinct, whereas Hantkenina became rare. to increase the boreal effects of the Northern Atlantics. At the same time, the development of typical Oligocene This cooling is well traced by the change of the sub- species started later, at the EoceneÐOligocene bound- tropical biota to the boreal type and by the abundance ary, which is considered as a base of the Globigerina of conifer pollen in the spore and pollen assemblages tapuriensis (P17/P18) Zone. from the beds with Propeamussium fallax from Ciscau- The appearance of the modern species of marine casia and the South Ukrainian Depression, as well as mollusks and the formation of benthic assemblages from the basal Oligocene beds of these regions, south- typical of the MioceneÐPleistocene was gradual and ern areas of the East European Platform, and western not contemporary in different climatic zones. In the late Kazakhstan. The major factors of the changes in the EoceneÐearly Oligocene, a gradual impoverishment of vegetative cover were a decrease in the average annual typical Eocene assemblages was widespread in the temperatures in the early Oligocene by 4Ð6°C com- warm-water zone. Starting from the second half of the pared to those in the late Eocene (the average winter Oligocene, the composition of benthic assemblages temperatures decreased, while the average summer was closely similar to that in the Miocene. At the same temperatures were the same). The shelf water tempera- time, the boreal communities were very closely similar ture similarly decreased (by 3Ð5°C after Marzuk, to each other and distinctive in the taxonomic composi- 1992), as indicated by isotopic data. tion throughout the Oligocene. The abrupt changes in Although the southern cooling was not so important, the composition of boreal assemblages followed the some floristic data show that it was accompanied by paleogeographic and climatic changes and occurred at wide fluctuations in atmospheric humidity (for more the EoceneÐOligocene boundary. detail see the previous section). The change in the dinoflagellate assemblages, Biotic events. The very marked abiotic environ- which was accompanied by the extinction of more than mental changes considered above also determined the a half of the Eocene taxa (at the base of the Phthanope- character of the geographical distribution of biota and ridinium amoenum Zone), approximately corresponds its changes at the EoceneÐOligocene boundary. The cli- to the EoceneÐOligocene boundary. Major changes in matic cooling and the eustatic regression in the terminal the composition of nannoplanktonic assemblages Eocene and at the beginning of the Oligocene resulted occurred much later, in the middle of the Rupelian at in the most important of the Cenozoic rearrangements the base of the Sphenolithus praedistentus (NP23) of biota, which persisted throughout the Rupelian. The Zone, when the impoverishment of assemblages was direct result of this cooling was an extinction of biotas over, and a new thermophilic group, Sphenolithus, of the early Paleogene type and formation of communi- appeared. ties with a quite different taxonomic composition. The evolutionary changes in biota were accompanied by The most substantial change in flora, the transition local northward displacements of geographical ranges from the Paleocenophytic to the Neocenophytic, which and of the majority of biogeographical borders follow- was characterized by the predominance of modern-day ing the paleogeographic, climatic, or facial changes, assemblages and modern taxa, was gradual (Geolo- which are often difficult to trace in available data. gicheskie i bioticheskie..., 1998; Akhmetiev, 1999). This started in the middle of the late Eocene. The level The response of different groups of biota to this cri- of the most marked elimination of thermophilic taxa sis was much the same and amounted to the partial approximately corresponds to the EoceneÐOligocene extinction of richer and taxonomically more diverse boundary. Frequent fluctuations in temperature and Eocene association of species and formation of a new atmospheric humidity contributed to more rapid disin- typical Oligocene assemblage that was based on the tegration of the Paleocenophytic flora. The presence of remaining opportunist species. Northern taxa pene- such fluctuations in the spore and pollen assemblages trated assemblages of the southern type during the cli- of Armenia is supported by mutually exclusive ranges matic cooling, but not in large quantity. For this reason, of cryptogams as indicators of humid regimes and pol- the main biogeographic subdivisions were generally len of grasses, Ephedra, Cupressaceae, and other taxa closely similar to those in the Eocene, although their characteristic of arid regimes. The change in flora borders and taxonomy markedly changed. occurred in both major phytochores of the Holarctic In different groups of biota, the evolutionary trans- Land, the Boreal and the Tethyan regions, as well as in formation occurred at basically different times and was the ecotone zone that separated them. At the beginning not catastrophic in character. Thus, the most marked of the early Oligocene, the floras of each of these changes in the composition of planktonic foraminiferal regions were characterized by more or less equal pro- and nummulitid assemblages occurred as early as the portions of the Paleocenophytic and Neocenophytic terminal Priabonian (inside the Turborotalia centralisÐ representatives. This process was over in the second Globigerina gortanii (P17) and Nummulites fabianii half of the Rupelian and resulted in the formation of the retiatus zones). At the bases of these zones, highly spe- moderately thermophilic, deciduous, mesophilic Oli- cialized groups Globigerapsis and Gribrohantkenina, a gocene flora, which was designated by A.N. Krishto- family of discocyclides, and three genera of nummulit- fovich as the Turgay Flora.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S247

The evolutionary transformation of terrestrial vert- Andreae, A., Ein Beitrag zur Kenntnis des Elsasser Tertiärs, abrates was also gradual. For the vertebrates, the EarlyÐ Teil 2, Abh. Geol. Special-Karte Elsass-Lothr., 1884, vol. 2. Late Oligocene boundary, when the Indricotheriidae no. 3, pp. 1Ð239. fauna formed, and the active radiation of rodents and Andreansky, G., Zur FlorenÐund Vegetationgeschichte des insectivores started (Reshetov in Geologicheskie i bio- Ungarishen Tertiärs, Sitz.ÐBer. Öster. Acad. Wissensch., ticheskie..., 1998), was of greater importance. Wien, Math.-Naturwiss. Rl., 1964, vol. 173, pp. 8Ð10 and 351Ð368. Andreyeva-Grigorovich, A.S., Distribution of Cysts of ACKNOWLEDGMENTS Dinoflagellates in the Maykopian Formation of the Northern Stavropol Region, Paleontol. Sb. L’vovsk. Univ. (Lvov), This study was first supported by the Interna- 1980, no. 17, pp. 74Ð79. tional Science Foundation and, subsequently, by the Andreyeva-Grigorovich, A.S., Nannoplankton from the Russian Foundation for Basic Research, projects Transitional EoceneÐOligocene and Oligocene Deposits of nos. 94-04-11187, 97-04-49870, and 01-05-65448. Armenia and Northern Caucasus, Paleontol. Sb. L’vovsk. Univ. (Lvov), 1981, no. 18, pp. 57Ð62. REFERENCES Andreyeva-Grigorovich, A.S., Regional Paleogene Strati- graphic Chart for Southern Regions of the Commonwealth of Adams, C.G., Gentry, A.W., and Whybrow, P.J, Dating the Independent States Based on Cysts of Dinoflagellates, Terminal Tethyan Event, Bull. Utrecht. Micropaleontol., Al’gologiya, 1994, vol. 4, no. 2, pp. 66Ð75. 1983, no. 30, pp. 273Ð298. Andreyeva-Grigorovich, A.S., Gruzman, A.O., Reifman, F.M., Agustí, J., Anadón, P., Arbiol, S., Cabrera, L., Colombo, F., and Smirnov, S.E., Biostratigraphic Character of the Menil- and Sáez, A., Biostratigraphical Characteristics of the Oligocene itic Formation Standard Section of the Chechva River (the Sequences of Northeastern Spain (Ebro and Campins Basins), Ukrainian Carpathians), Paleontol. Sb. L’vovsk. Univ. Münchner Geowiss. Abh. (A), 1987, vol. 10, pp. 35Ð42. (Lvov), 1986, no. 23, pp. 83Ð89. Akhmetiev, M.A., Palaeobotanical Data on the Geological Andreyeva-Grigorovich, A.S. and Gruzman, A.D., The Bio- Age of the Greater Caucasus, Paleontol. J., 1995, vol. 29, stratigraphic Basis of the PaleogeneÐNeogene Boundary in no. 1A, pp. 172Ð181. the Central and Eastern Paratethys, Geol. Carpathica, 1994, Akhmetiev, M.A., Popov, S.V., and Krhovsky, J., Excursion vol. 45, no. 6, pp. 333Ð342. Guidebook. Paleontology and Stratigraphy of the EoceneÐ Arambourg, C., Résultats scientifiques de la Mission Miocene Sections of the Western Pre-Caucasia, Field Sym- C. Arambourg en Syrie et en Iran (1938Ð1939). Les poissons posium 2Ð10 August, MoscowÐKrasnodar, 1995, pp. 1Ð16. oligocènes de l’Iran, Notes mém. Moyen-Orient, 1967, vol. 8, Akhmetiev, M.A. and Zaporozhets, N.I., New Palm Occur- pp. 9Ð210. rences from Oligocene Beds of the Darrydag (Nakhichevan Aslanova, S.M., Some Data on Fossil Ichthyofauna from the ASSR), Byull. Mosk. OÐva Ispyt. Prir., Otd. Geol., 1989, Shemakha Region, Izv. Akad. Nauk Az. SSR, Ser. Geol., vol. 64, no. 6, pp. 57Ð67. Geogr., 1960, no. 5, pp. 77Ð84. Akhmetiev, M.A. and Zaporozhets, N.I., Oligocene Climates Báldi, T., On the Oligocene and Miocene Stages of the Cen- in the Eastern CrimeaÐCaucasus Region and Kazakhstan, tral Paratethys and on the Formation of the Egerian in Hun- Izv. Akad. Nauk, Ser. Geol., no. 3, pp. 5Ð23. gary, Ann. Univ. Sci. Budapesttensis, Sect. Geol., 1969, Albrecht, I. and Valk, W. Oligozäne Invertebraten von Süd- vol. 12, pp. 19Ð28. Limburg, Med. Geol. Stichting Maastricht, Ser. C, 1943, Báldi, T., The Terminal Eocene and Oligocene Events in vol. 4, no. 13, pp. 1Ð163. Hungary and the Separation of an Anoxic, Cold Paratethys, Ecl. Geol. Helv. Alvarez Sierra, M.A., Daams, R., Lacomba, J.I., López Mar- , 1984, vol. 77, no. 1, pp. 1Ð28. tínez, N., and Sacristán Martin, M.A., Succession of Micro- Báldi, T., Mid-Tertiary Stratigraphy and Paleogeographic mammal Faunas in the Oligocene of Spain, Münchner Evolution of Hungary, Budapest: Akad. Kiadó, 1986. Geowiss. Abh. (A), 1987, vol. 10, pp. 43Ð48. Báldi-Beke, M., Stratigraphical and Faciological Subdivi- Amitrov, O.V., Istoriya gastropod paleogenovykh morei sions of the Oligocene as Based on Nannoplankton, Földt. zapada Evrazii (The History of Gastropods from the Paleo- Közl., 1977, vol. 107, pp. 59Ð89. gene Seas of Western Eurasia), Moscow: Nauka, 1993. Bannikov, A.F., Fossil Scombrids in the USSR, Tr. Paleontol. Amitrov, O.V., Paleogeography and Climatic Zonation of the Inst. Akad. Nauk SSSR (Moscow), 1985a, vol. 210, pp. 1Ð111. Early Oligocene Seas in Western Eurasia Based on Gastro- Bannikov, A.F., Fossil Stromateoids (Teleostei) from the pods, Paleontol. Zh., 1996, no. 1, pp. 3Ð12. Caucasus, Paleontol. Zh., 1985b, no. 3, pp. 77Ð83. Ananiashvili, G. and Minashvili, T., New Data on the Bio- Bannikov, A.F., Occurrence of the Genus Trachinus (Teleo- stratigraphy of the Maykopian Formation of Georgia, Dokl. stei) in the Oligocene of the Caucasus, Paleontol. Zh., 1986, Akad. Nauk SSSR, 1990, vol. 311, no. 4, pp. 922Ð924. no. 3, pp. 126Ð129. And– elkovic′ , J., Tertiary Fishes of Yugoslavia: A Strati- Bannikov, A.F., A New Species of Stromateids (Perciformes) graphicÐPaleontologicÐPaleoecological Study, Palaeontol. from the Lower Oligocene of the Caucasus, Paleontol. Zh., Jugosl., 1989, no. 38, pp. 1Ð121. 1988, no. 4, pp. 108Ð113. And– elkovic′ , M., Eremija, M., Pavlovic′ , M., And– elkovic′ , J., Bannikov, A.F., Fossil Carangids in the USSR, Tr. Paleontol. and Mitrovic′ -Petrovic′ , J., Paleogeography of Serbia—Ter- Inst. Akad. Nauk SSSR (Moscow), 1990, vol. 224, pp. 1Ð108. tiary, Beograd: Univ. Beograd, Inst. Region. Geol., Paleon- Bannikov, A.F., A New Family of the Oligocene Perciformes, tol., 1991. Paleontol. Zh., 1991, no. 4, pp. 88Ð94.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S248 POPOV et al.

Bannikov, A.F., Fossil Stromateoids (Perciformes) of the Bieda, F., Duzún e otwornice eocenu tatrza′ skiego, Prace Inst. Families Nomeidae and Ariommidae, Paleontol. Zh., 1993, Geol., 1963, vol. 37, pp. 1Ð215. no. 4, pp. 87Ð96. Blainville, H.M., Sur les ichthyolites ou les poissons fossiles, Bannikov, A.F. and Danilchenko, P.G., A New Family of the Nouv. Diet. Hist. Natur. (Paris), 1818, vol. 27, pp. 310Ð395. Cenozoic Beryciformes, Dokl. Akad. Nauk SSSR, 1985, Bogachev, V.V., Materials on the Tertiary Ichthyofauna of the vol. 283, no. 3, pp. 707Ð709. Caucasus, Tr. Azerb. Neft. Inst., Otd. Geol., 1933, vol. 15, Bannikov, A.F. and Parin, N.V., Description of a New Species pp. 1Ð62. of a Hairtail (Aphanopinae, Trichiuridae) from the Middle Bogachev, V.V., A New Fish Genus and Species from the Eocene of the Northern Caucasus; Reestablishment of the Paleogene of Abkhazia, Byull. Gos. Muz. Gruzii, 1938, Validity of the Fossil Genus Anenchelum, Vopr. Ikhtiol., vol. 9a, pp. 1Ð5. 1995, vol. 6, pp. 725Ð731. Bogachev, V.V., Paleoichthyological Notes, Tr. Azerb. Neft. Bannikov, A.F. and Parin, N.N., A List of Marine Fishes from Inst., 1964, vol. 13, pp. 126Ð133. the Cenozoic (Upper PaleoceneÐMiddle Miocene) Localities of the Southwestern Area of Russia and Adjacent Territories, Böhm, B., Die fossilen Fische von Konvaszna und Kom- Vopr. Ikhtiol., 1997, vol. 37, no. 2, pp. 149Ð161. mando in Siebenbürgen, Mitt. Jahrb. Kgl. Ungar. Geol. Anstalt (Budapest), 1941, vol. 35, pp. 179Ð203. Bannikov, A.F. and Tyler, J.C., Caprovesposus from the Oli- gocene of Russia: the Pelagic Acronurus Presettlement Stage Boitsova, E.P., Miospores and Stratigraphy of the Paleogene of a Surgeonfish (Teleostei: Acanthuridae), Proc. Biol. Soc. Beds in the Eastern Regions of Western Kazakhstan, Doc- Wash., 1992, vol. 105, no. 4, pp. 810Ð820. toral (Geol.ÐMineral.) Dissertation, 1973, pp. 1Ð35. Barbin, V. and Bignot, C., New Proposal for an EoceneÐOli- Bombita, G. and Moisescu, V., Données actuelles sur le gocene Boundary According to Microfacies from the Pria- Nummulitique de Transylvanie, Mém. B.R.G.M., 1968, bonian-type Section, Terminal Eocene Events, Elsevier Sci. no. 58, pp. 693Ð729. Publ., 1986, vol. 5, pp. 49Ð52. Bombita, G. and Rusu, A., New Data on the Eocene/Oli- Barbin, V. and Guernet, C., Contribution to the Study of the gocene Boundary in the Romanian Carpathians Palaeo- Priabonian of the Type Region (North Italy): The Ostracods, geogr., Palaeoclimatol., Palaeoecol., 1981, vol. 36, nos. 3Ð4, Rev. Micropaleontol., 1988, vol. 30, no. 4, pp. 209Ð231. pp. 213Ð222. Batjes, D.A.J., Foraminifera of the Oligocene of Belgium, Bondareva, T.P., Ivanova, V.A., and Samodurov, V.I., Corre- Inst. R. Sci. Nat. Belgique Mém., 1958, vol. 143, pp. 1Ð188. lation of Paleogene Marine Sections of Northern and South- Beer, M.A. and Shcherba, I.G., The Late Alpine Evolution of ern Aral Regions, Geologiya i perspektivy neftegazonosnosti Eastern Carpathians, Byull. Mosk. OÐva Ispyt. Prir., Otd. nekotorykh raionov SSSR (Geology and Prospects for Pres- Geol., 1984, vol. 59, no. 4, pp. 47Ð62. ence of Oil and Gas in Some Regions of the USSR), Mos- cow: Nedra, 1964, pp. 266Ð290. Belmustakov, E., Fosilite na B”lgariya (Fossils of Bulgaria), vol. 4: Paleogen. Golemi foraminiferi (Paleogene: Larger Bonis, L. de, Bouvrain, G., Buffetaut, E., Denys, Ch., Ger- Foraminifers), Sofia: B”lg. Akad. Nauk, 1959. aads, D., Jaeger, J.-J., Martin, M., Mazin, J.-M., and Rage, J.-Cl., Contribution des Vertébrés à l’histoire de la Téthys et des Belyaeva, M.M., Kharkov Beds in Rostov Region, continents péritéthysiens, Bull. Soc. Géol. France, 1985, Geologiya i mineral’nye resursy territorii Nizhnego Dona i sér. 8, vol. 1, no. 5, pp. 781Ð784. Nizhnego Povolzh’ya (Geology and Mineral Resources of Lower Don Region and Lower Volga Region), Rostov-on-Don: Boulanger, D., Debourle, A., and Deloffre, R., Le recif Oli- 1964, pp. 60Ð69. gocène du Tuc de Saumon (Aquitaine, France Sud-Ouest), Bull. Centre Rech. Pau Bendukidze, O.G., Rodent Fauna from Karakoin and Its , 1970, vol. 4, no. 3, pp. 9Ð37. Stratigraphic Significance, Materialy Simpoziuma Perekhod Boulanger, D. and Poignant, A., Sur quelques foraminifères ot oligotsena k miotsenu v Severnom polusharii Mater. Simp. fixes de l’Éocène supérieur et de l’Oligocène en Aquitaine, (Proc. Symp. Transition from Oligocene to Miocene in Rev. Micropaleontol., 1971, vol. 14, no. 9, pp. 167Ð176. Northern Hemisphere), Alma-Ata, 1994. Bronevoi, V.A., Bryzzheva, N.N., Vasiliev, I.V., et al., The Bendukidze, O.G., The Oligocene Rodents of Central and Upper Paleogene Beds of the Northern Ustyurt and Chagrai Western Kazakhstan and Their Stratigraphic Significance, Plateaus, Tr. Vses. Geol. Inst., Nov. Ser. (Leningrad), 1967, Mém. Trav. E.P.H.E., Inst. Montpellier, 1997, no. 21, vol. 123, pp. 33Ð61. pp. 205Ð208. Bruijn, H. de, Sondaar, P.Y., and Sanders, E.A.C., On a New Berger, J.-P., Correlative Chart of the European Oligocene Species of Pseudoltinomys (Theridomyidae, Rodentia) from and Miocene: Application to the Swiss Molasse Basin, Ecl. the Paleogene of Mallorca, Proc. Kon. Neder. Akad. Wetensch., Geol. Helv., 1992, vol. 85, no. 3, pp. 573Ð609. 1979, ser. B, vol. 82, no. 1, pp. 1Ð10. Berger, J.-P., Cartes paléogéographiques–palinspasthiques Brunet, M., Les grands Mammifères chefs de file de l’immi- du bassin molassique suisse (Oligocène inférieur–Miocène gration Oligocène et le problème de la limite Éocène-Oli- moyen), Neues Jahrb. für Geol. Paläontol. Abh., 1996, gocène en Europe, Paris: Singer-Polignac, 1979. vol. 202, no. 1, pp. 1Ð44. Budantsev, L.Yu., Oligocene Flora from the Northern Aral Berggren, W.A., Kent, D.V., Swisher, C.C., and Aubry, M.-P., Coastal Area, Problemy paleobotaniki (Problems in Paleobo- A Revised Cenozoic Geochronology and Chronostratigra- tanics), MoscowÐLeningrad: Akad. Nauk SSSR, 1959, phy, Soc. Econ. Paleontol. Mineral., Spec. Publ., 1995, pp. 190Ð252. no. 54, pp. 129Ð212. Bugrova, E.M., Detailed Elaboration of the Biostratigraphic Bhatia, S.B., The Foraminiferal Fauna of the Late Palaeo- Division of the Eocene of the Krasnovodskii Peninsula and gene Sediments of the Isle of Wight, England, J. Paleontol., the Kara Bogaz Region Based on Foraminifers, Tr. 28 sessii 1955, vol. 29, no. 4, pp. 665Ð693. Vsesoyuznogo Paleontologicheskogo Obshchestva (Proc.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S249

28 Session of All-Union Paleontol. Soc.), Leningrad, 1985, Colloque sur l’Éocène (Paris, mai 1968), Mém. Bureau Rech. pp. 41Ð49. Géol. et Minièr., 1968, no. 58, pp. 1Ð742. Bugrova, E.M., Paleogene Stratigraphy of Maryi Region, Cossmann, M. Synopsis illustré des mollusques des l’Éocène Sov. Geol., 1987, no. 8, pp. 53Ð63. et de l’Oligocène en Aquitaine, Mém. Soc. Géol. France, Bugrova, E.M., Stratigraphic and Geographic Distribution of Palaeontol., 1921, vol. 23, nos. 3Ð4, pp. 1Ð112. the Upper Eocene Foraminifers in the Northern Areas of the Cossmann, M. and Lambert, I., Étude paléontologique et Tethyan Basin, Stratigr. Geol. Korrelyatsiya, 2001, vol. 9, stratigraphique sur le terrain Oligocène marin aux environs no. 2, pp. 92Ð104. d’Étampes, Mém. Soc. Géol. France, 1884, ser. 3, vol. 3, Bugrova, E.M., Gusev, E.A., and Tverskaya, L.A., Oligocene pp. 1Ð187. Beds in the Knipovich Ridge, Tezisy Dokl. Mezhdunarodnoi Crochet, J.-Y., Thomas, H., Roger, J., Sen, S., and Al-Sulaim- Shkoly morskoi geologii (Abstracts of Pap. Int. School of ani, Z., Première découverte d’un créodonte dans le pénin- Marine Geol.), Moscow, 2001 (in press). sule Arabique: Masrasector ligabuei nov. sp. (Oligocène Bykova, N.K. and Azbel’, A.Ya., Stratigraphic Division of inférieur de Taqah, Formation d’Ashwaq, Sultanat d’Oman, the Maykopian Beds in the Peninsula of Buzachi Based on C. R. Acad. Sci. (Paris), 1990, sér. 2, vol. 311, pp. 1455Ð1460. Foraminifers, Tr. VNIGRI, 1962, vol. 190, pp. 375Ð397. Crochet, J.-Y., Thomas, H., Sen, S., Roger, J., Gheerbrant, É., Cavelier, C. La limite Éocène-Oligocène en Europe occidentale, and Al-Sulaimani, Z, Découverte d’un Péradectidé (Marsu- Mém. Univ. L. Pasteur Strasbourg, 1979, no. 54, pp. 1Ð280. pialia) dans l’Oligocène inférieur du Sultanat d’Oman: nou- velles données sur la paléobiogéographie des Marsupiaux de Cavelier, C. and Pomerol, Ch., Stratigraphy of the Paleogene, la plaque arabo-africain, C. R. Acad. Sci. (Paris), 1992, sér. 2, Bull. Soc. Géol. France, 1986, vol. 11, no. 2, pp. 255Ð265. vol. 314, pp. 539Ð545. Chang, M.-M. and Chen, Y.-Y., Late Mesozoic and Tertiary Cuenca, G., Canudo, J.I., Laplana, C., and Andres, J.A., Bio Ichthyofaunas from China and Some Puzzling Patterns of y cronostratigrafía con mamíferos en la Cuenca Terciaria del Distribution, Vertebr. Palasiat., 2000, no. 38, pp. 161Ð175. Ebro: ensayo de sintesis, Acta Geol. Ispan., 1992, vol. 27, Chang, M.-M. and Chow, C.-C., Stratigraphic and Geo- nos. 1Ð2, pp. 127Ð143. graphic Distributions of Late Mesozoic and Cenozoic Fishes Cuvillier, I., Révision du Nummulitique Égyptien, Mém. of China, Indo–Pacific Fish Biology: Proc. Second Int. Conf. Inst. d’Égypte, 1930, vol. 16, pp. 1Ð372. Indo–Pacific Fishes, Tokyo, 1986, pp. 529Ð539. Daams, R., Freudenthal, M., Lacomba, J.I., and Alvarez, M.A., Chang, M.-M., Miao, D., Chen, Y., Zhou, J., and Chen, P., Upper Oligocene Micromammals from Pareja, Loranca Suckers (Fish, Catostomidae) from the Eocene of China: Basin, Prov. of Guadalajara, Spain, Scripta Geol., 1989, Account for the Family’s Current Disjunct Distributions, Sci. no. 89, pp. 27Ð56. China, 2001, vol. 44, no. 7, pp. 577Ð586. Danil’chenko, P.G., Fishes of the Family Gonostomidae Chang, M.-M. and Zhou, J.-J., A Brief Survey of the Chinese from the Tertiary Beds of the Caucasus and Crimea, Izv. Eocene Ichthyofauna, Kaupia, Darmstadter Beiträge zur Akad. Nauk SSSR, Ser. Biol., 1946, no. 6, pp. 139Ð146. Naturgeschichte, 1993, vol. 2, pp. 157Ð162. Danil’chenko, P.G., Fishes of the Family Myctophidae from Cheng, C.C., Fossil Fishes from the Early Tertiary of Hsiang- the Oligocene of the Caucasus, Dokl. Akad. Nauk SSSR, Hsiang, Hunan, with Discussion of Age of the Hsiawanpu 1947, vol. 56, no. 2, pp. 193Ð196. Formation, Vertebr. Palasiat., 1962, vol. 6, pp. 333Ð343.

Danil’chenko, P.G., The Genus Palaeogadus and Its Evolu-

^ Ciobanu, M.,^ Date noi asupra pes tilor fosili din Oligocenul tion, Tr. Paleontol. Inst. Akad. Nauk SSSR, 1950, vol. 25, de la Piatra Neamt (II), Stud. Cercet. Muz. S¸ ti. Natur. Piatra pp. 1Ð25.

Neamt, 1970, vol. 1, pp. 67Ð90. ^ Danil’chenko, P.G., Teleosts from the Maykopian Beds of Ciobanu, M., Fauna fosila din Oligocenul de la Piatra the Caucasus, Tr. Paleontol. Inst. Akad. Nauk SSSR, 1960,

Neamt, Bucharest: Acad. R.S.R., 1977. vol. 78, pp. 1Ð208.

^ Ciobanu, M.,^ Date noi asupra pes tilor fosili din Oligocenul Danil’chenko, P.G., The Main Assemblages of Fish Fauna in de la Piatra Neamt , An. Muz. S¸ ti. Natur. Piatra Neamt, Ser. the Cenozoic Seas of the Tethys, Tr. Paleontol. Inst. Akad. Geol.-Geogr., 1978, vol. 4, pp. 185Ð192. Nauk SSSR (Moscow), 1980a, vol. 178, pp. 175Ð183. Cita, M.B. and Scipolo, C., Chapmanina gassinensis (Silves- Danil’chenko, P.G., The Orders Elopiformes, Clupeiformes, tri) dans l’Oligocène du Monto Baldo (Italie), Rev. Micropa- Salmoniformes, Myctophiformes, Lampridiformes, Beryci- leontol., 1961, vol. 4, no. 3, pp. 121Ð134. fomies, Zeiformes, Perciformes, Echeneifoimes, and Tetra- Cizancourt, M., Sur la stratigraphie et la faune Nummuli- odontiformes, Iskopaemye kostistye ryby SSSR (Fossil Teleo- tique du flysch de l’Albanie, Bull. Soc. Géol. France, 1930, sts in the USSR), Novitskaya, L.I., Ed., Moscow: Nauka, sér. 4, vol. 30, nos. 3Ð4, pp. 195Ð212. 1980b, pp. 6Ð28, 45Ð50, 91Ð96, 99Ð104, 115Ð169, 173Ð174. Codrea, V., Stratigraphical Significance of Ceratomorph Danil’chenko, P.G., Switchenska, A.A., Sergienko, N.I., Perissodactyls (Mammalia) from the Transylvanian Paleo- Sytchevskaya, E.K., and Fedotov, V.F., Fossil Teleosts of the gene, Stud. Univ. Babes-Bolyai. Geol., 1993, vol. 38, no. 2, USSR, Tr. Paleontol. Inst. Akad. Nauk SSSR (Moscow), pp. 71Ð76. 1980c, vol. 178, pp. 1Ð210. Codrea, V. and Suraru, N. New Remains of Indricotheriin Dashzeveg, D., Asynchronism of the Main Mammalian Fau- (Perissodactyla, Mammalia) in the Lower Oligocene at Fildu nal Events near the EoceneÐOligocene Boundary, Tertiary de Jos (Salaj District, NW Transylvania), Rom. J. Paleontol., Res., 1993, vol. 14, no. 4, pp. 141Ð149. 1995, vol. 76, pp. 81Ð84. Dashzeveg, D., A New Hyracodontid (Perissodactyla, Rhi- Colloque sur le Paleogène (Bordeaux, septembre 1962), nocerotoidea) from the Ergilin Dzo Formation (Oligocene, Mém. Bureau Rech. Géol. et Minièr., 1964, no. 28, vol. 1, Quarry 1) in Dzamyn Ude, Eastern Gobi Desert, Mongolia, pp. 1Ð544, vol. 2, pp. 547Ð1107. Am. Museum Novit., 1996a, no. 3178, pp. 1Ð12.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S250 POPOV et al.

Dashzeveg, D., Some Carnivorous Mammals from the Paleo- Ducasse, O. and Roussele, L., L’approche populationniste gene of the Eastern Gobi Desert, Mongolia, and the Applica- chez les Hammatocythere du Paléogène aquitain. Une manière tion of Oligocene Carnivores to Stratigraphic Correlation, de suivre leurs réponses aux fluctuations du milieu. Incidences Am. Museum Novit., 1996b, no. 3179, pp. 1Ð14. taxinomiques et biostratigraphiques, VII International Sympo- Dashzeveg, D. and Russell, D.E., Extension of Dyspternine sium on Ostracodes, Belgrade, 1979b, pp. 185Ð190. Pantolestidae (Mammalia, Cimolesta) in the Early Oligocene Dzhafarova, Zh. D., A Member of the Family Trichiuridae of Mongolia, Géobios, 1992, vol. 25, no. 5, pp. 647Ð650. with the Otolith from the Khadumian of the Apsheron Penin- Davidzon, R.M., Kreidenkov, G.P., and Salibaev, G.Kh., sula, Available from VINITI, 1988, Baku, no. 7581-V. Stratigrafiya paleogenovykh otlozhenii Tadzhikskoi depressii Dzhafarova, Zh.D. and Aslanova, S.M., Remains of the i sopredel’nykh territorii (Stratigraphy of the Paleogene Paleogene Tuna (Thunnus) from the Apsheron Peninsula, Deposits of the Tajik Depression and Adjacent Regions), Dokl. Akad. Nauk Az. SSSR, 1974, vol. 30, no. 12, pp. 44Ð47. Dushanbe: Donysh, 1982. Emry, R.J., Lucas, S.G., Szalay, F.S., and Tleuberdina, P.A., Deltel, B., Nouveaux ostracodes de l’Éocène et de l’Oli- A New Herpetotheriine Didelphid (Marsupialia) from the gocène de l’Aquitaine méridionale, Act. Soc. Lin. Bordeaux, Oligocene of Central Asia, J. Vertebr. Paleontol., 1995, 1962Ð1963, vol. 100, pp. 127Ð221. vol. 15, no. 4, pp. 850Ð854. Dlusskii, G.M., Murav’i pustyn’ (Ants from Deserts), Mos- Emry, R.J., Wang, B., Tjutkova, L.A., and Lucas, S.G., cow: Nauka, 1981. A Late Eocene Eomyid Rodent from the Zaysan Basin of Doppert, J.W.Chr. and Neele, N.G., Biostratigraphy of Kazakhstan, J. Vertebr. Paleontol., 1997, vol. 17, no. 1, Marine Paleogene Deposits in the Netherlands and Adjacent pp. 229Ð234. Areas, Med. Rijks Geol. Dienst, 1983, vol. 37, no. 2, pp. 3Ð79. Erbajeva, M.A. and Tyutkova, L.A., Paleogene and Neogene Ducasse, O., Étude micropaléontologique (ostracodes) de Lagomorphs from Kazakhstan, Mém. Trav. E.P.H.E., Inst. l’Éocène nord-aquitain. Interprétation biostratigraphique et Montpellier, no. 21, pp. 209Ð214. paléogéographique, Thèse de Doctorat Sc. (Naturelles), Bor- Fedotov, V.F., The Gadids from the PaleogeneÐNeogene deaux, 1969, pp. 1Ð381. Beds of the USSR, Tr. Paleontol. Inst. Akad. Nauk SSSR Ducasse, O., Quelques remarques sur la faune d’ostracodes (Moscow), 1976, vol. 157, pp. 1Ð80. des faciès profonds du Tertiaire aquitain, Bull. Inst. Géol. Fedotov, V.F., The Order Gadiformes, Iskopaemye kostistye Bassin d’Aquitaine (Bordeaux), 1974, vol. 16, pp. 127Ð135. ryby SSSR (Fossil Teleosts in the USSR), Novitskaya, L.I., Ducasse, O., Les associations fauniques d’ostracodes de Ed., Moscow: Nauka, 1980, pp. 62Ð82. l’Éocène moyen et supérieur dans le Sud du Bassin d’Aquita- Fedotov, V.F. and Bannikov, A.F., The First Find of the ine. Distribution schématique et valeur paléoécologique, Marine Eel (Congridae) in the Oligocene of the Caucasus, Bull. Inst. Géol. Bassin d’Aquitaine (Bordeaux), 1975, Byull. Mosk. OÐva Ispyt. Prir., Otd. Geol., 1984, vol. 59, vol. 17, pp. 17Ð26. no. 5, pp. 101Ð105. Ducasse, O., Étude populationiste du genre Cytherella Fejfar, O., A Lower Oligocene Mammalian Fauna from (ostracodes) dans les faciès bathyaux du Paléogène aquitain. Detan and Dvérce NW Bohemia, Czechoslovakia, Münchner Intérêt dans la reconstitution des paléoenvironnements, Bull. Geowiss. Abh. (A), 1987a, vol. 10, pp. 253Ð264. Inst. Géol. Bassin d’Aquitaine (Bordeaux), 1981, vol. 30, pp. 161Ð185. Fejfar, O., Oligocene Rodents from Zallah Oasis, Libya, Münchner Geowiss. Abh. (A), 1987b, vol. 10, pp. 265Ð268. Ducasse, O., Étude de populations du genre Protoargilloecia (ostracodes) dans les faciès bathyaux du Paléogène aquitain: Ferriz, F.J., Freudenthal, M., and Martin Suarez, E., The Old- deuxième test effectué en domaine profond. Comparaison est Mammals from the Betic Cordillera: the Oligocene of the avec le genre Cytherella, Géobios, 1983, vol. 16, no. 3, Internal Prebetic (Neppio, Albacete, SE Spain), Géobios, pp. 273Ð283. 1993, vol. 26, no. 5, pp. 625Ð631. Ducasse, O. and Peypouquet, J.P., Les ostracodes et l’évolu- Flandrin, I., Contribution à l’étude paléontologique du Num- tion paléogéographique et paléohydrologique des marges mulitique Algérien, Mater. Carte Géol. Algérie, Paléontol., continentales de l’Atlantique nord-oriental pendant le Céno- 1938, no. 8, pp. 13Ð15. zoïque, Bull. Soc. Géol. France, 1978, sér. 7, vol. 20, no. 4, Flerov, K.K. and Yanovskaya, N.M., The Ecological Mam- pp. 441Ð452. mal Assemblages from the Oligocene of Asia and Their Zoo- Ducasse, O. and Peypouquet, J.P., Cenozoic Ostracoda, Their geographic Characteristics, Tr. Paleontol. Inst. Akad. Nauk Importance for Bathymetry, Hydrology, and Biogeography, SSSR (Moscow), 1971, vol. 130, pp. 7Ð30. Initial Reports Deep Sea Drilling Project, 1979, vol. 48, Freudenthal, M., The Early Oligocene Rodent Fauna of Olalla pp. 343Ð363. 4A (Teruel, Spain), Scripta Geol., 1996, no. 112, pp. 1Ð67. Ducasse, O. and Peypouquet, J.P., Ostracods at the EoceneÐ Freudenthal, M., Hugueney, M., and Moissenet, E., The Oligocene Boundary in the Aquitaine Basin: Stratigraphy, Genus Pseudocricetodon (Cricetidae, Mammalia) in the Phylogeny, Paleoenvironments, in Terminal Eocene Events, Upper Oligocene of the Province of Teruel (Spain), Scripta Amsterdam: Elsevier Sci. Publ. B.V., 1986, pp. 265Ð273. Geol., 1994, no. 104, pp. 57Ð114. Ducasse, O. and Roussele, L., Hammatocythere oertlii Frochlicher, H. and Weiler, W., Die Fischfauna der unter- (Ducasse, ostracodes): espèce polymorphe de l’Éocène du stampischen Molasse des Eutlebuchs Kt. Luzern, und ihre Blayais, Bull. Inst. Géol. Bassin d’Aquitaine (Bordeaux), paläogeographische Bedeutung, Ecl. Geol. Helv., 1952, 1978, vol. 24, pp. 3Ð35. vol. 45, no. 1, pp. 1Ð37. Ducasse, O. and Roussele, L., Les Hammatocythere (ostra- Frost, S.N., Oligocene Reef Coral Biofacies of the Vicentin, codes) de l’Oligocène aquitain. Bull. Inst. Géol. Bassin Northeast Italy, Publ. Soc. Econ. Paleontol. Mineral., 1981, d’Aquitaine (Bordeaux), 1979a, vol. 25, pp. 221Ð255. no. 3, pp. 483Ð539.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S251

Furon, R. and Soyer, R., Catalogue des fossiles tertiaires du real (Province de Tarragona, Catalogne), Estudios Geol., bassin de Paris, Savoir. Hist. Natur., 1947, vol. 22, no. 240, 1982, vol. 38, pp. 95Ð102. pp. 1Ð240. Gaudant, J., Nouvelles recherches sur les Cyprinidae (pois- Gabounia, L.K. and Chkhikvadze, V.M., Aperçu sur les sons téléostéens) oligocènes des Limagnes, Géobios, 1984a, faunes de vertébrés du Paléogène de la region de Zaissan vol. 17, no. 6, pp. 659Ð666. (Kazakhstan de l’Est), Mém. Trav. E.P.H.E., Inst. Montpellier, Gaudant, J., Sur la présence de “Percichthyidae” (poissons 1997, no. 21, pp. 193Ð203. téléostéens) dans l’Éocène moyen du bassin du Duero (Province Gabunia, L.K., Benarskaya fauna oligotsenovykh poz- de Zamora, Espagne), Acta Geol. Hisp., 1984b, pp. 139Ð142. vonochnykh (Benara Fauna of Oligocene Vertebrates), Tbilisi: Metsniereba, 1964. Gaudant, J., Mise en évidence d’Osmeridae (poissons téléostéens, Salmoniformes) dans l’Oligocène lacustre Gabunia, L.K., Contribution à la conaissance des Mam- d’Europe occidentale, C. R. Acad. Sci. (Paris), 1985a, mifères paléogènes du Bassin de Zaissan (Kazakhstan cen- vol. 300, no. 2, pp. 79Ð82. tral), Géobios, 1977, Mém. Spéc., no. 1, pp. 29–37. Gaudant, J., Mise au point sur les vertébrés inférieurs de Gabunia, L.K., First Occurrence of Pantolestids (Pantoles- l’Oligocène de Sieblos (Hesse, Allemagne), C. R. Acad. Sci. tidae, Insectivora) from the Paleogene of the USSR, Soob- (Paris), 1985b, vol. 300, no. 2, pp. 185Ð188. shch. Akad. Nauk GSSR, 1989, vol. 136, no. 1, pp. 177Ð180. Gaudant, J., Mise en évidence de Chandidae (poissons Gabunia, L.K. and Gabunia, V.D., A Brief Review of the téléostéens, Percoidei) dans le Cénozoïque européen, C. R. Paleogene and Early Miocene Insectivore Faunas from the Acad. Sci. (Paris), 1987a, vol. 304, no. 2, pp. 1249Ð1252. Zaisan Depression (Eastern Kazakhstan), Izv. Akad. Nauk Gruz. SSR., Ser. Biol. (Moscow), 1987, vol. 13, no. 6, Gaudant, J., Mise au point sur certains poissons Amiidae du pp. 406Ð411. Cénozoïque européen: le genre Cyclurus Agassiz (Kindleia Gad, J., Theridomys brachydens n. sp., a New Theridomys Jordan), J. Paläontol., 1987b, vol. 61, pp. 321Ð330. Species from Möhren (South Germany), Münchner Geowiss. Gaudant, J., Les cyprinodontiformes (poissons téléostéens) Abh. (A), 1987, vol. 10, pp. 269Ð272. Oligocènes de Ronzon, le Puy-en-Velay (Haute-Loire): anat- Gaudant, J., Nouvelles observations sur l’ichtyofaune stamp- omie et signification paléoécologique, Géobios, 1988, vol. 21, ienne d’Oberdorf (Canton de Soleure), Ecl. Geol. Helv., no. 6, pp. 773Ð785. 1977, vol. 70, no. 3, pp. 789Ð809. Gaudant, J., L’ichtyofaune stampienne des environs de Char- Gaudant, J., Découverte du plus ancient représentant connu tres-de-Bretagne, près de Rennes (Ille-et-Vilaine): un réexa- de genre Esox L. (poisson téléostéen, Esocoidei) dans le men, Géol. France, 1989, nos. 1Ð2, pp. 41Ð54. Stampien moyen du bassin d’Apt (Vaucluse), Géol. Méditerr., Gaudant, J., The Eocene Freshwater Fish-Fauna of Europe: 1978a, vol. 5, pp. 257Ð268. From Palaeobiogeography to Palaeoclimatology, Kaupia, Gaudant, J., Nouvelles observations sur les poissons oli- Darmstädter Beitr. Natur., 1993, vol. 3, pp. 231Ð244. gocènes de Monteviale (Vicenza-Italie), Mém. Sci. Géol., Genise, J.F. and Bown, T.M., New Trace Fossils of Termites 1978b, no. 32, p. 9. (Insecta: Isoptera) from the Late EoceneÐEarly Miocene of Gaudant, J., Sur les conditions de gisement de l’ichtyofaune Egypt, and the Reconstruction of Ancient Isopteran Social oligocène d’Aix-en-Provence (Bouches-du-Rhône): essai de Behavior, Ichnos., 1994, vol. 3, pp. 1Ð29. définition d’un modèle paléoécologique et paléogéographique, Geologicheskie i bioticheskie sobytiya pozdnego eotsenaÐ Géobios, 1978c, vol. 11, no. 3, pp. 393Ð397. rannego oligotsena na territorii byvshego SSSR (Geological Gaudant, J., Sur la présence de Gobiidae (poissons and Biotic Events in the Late Eocene to the Early Oligocene téléostéens) dans l’Oligocène inférieur de Rouffach (Bas- in the Territory of the Former Soviet Union), vol. 1: Rhin), Sci. Géol. Bull., 1979a, vol. 32, no. 3, pp. 131Ð137. Regional’naya geologiya verkhnego eotsena i nizhnego oli- Gaudant, J., Contribution à l’étude des vertébrés oligocènes gotsena (Regional Geology of the Upper Eocene and Lower de Soulce (Canton du Jura), Ecl. Geol. Helv., 1979b, vol. 72, Oligocene), Krasheninnikov, V.A. and Akhmetiev, M.A., no. 3, pp. 871Ð895. Eds., Moscow: GEOS, 1996, 1998, vol. 2. Gaudant, J., Sur la présence de dents de Characidae (poissons Geologicheskie sobytiya na granitse eotsena i oligotsena na téléostéens, Ostariophysi) dans les “calcaires à bythinies” et territorii Armyanskoi SSR (Geological Events at the EoceneÐ les “sables bleutés” du Var, Géobios, 1979c, vol. 12, no. 3, Oligocene Boundary in the Territory of the Armenian SSR), pp. 451Ð457. Gabrielyan, A.A., Ed., Yerevan: Akad. Nauk Arm. SSR, Gaudant, J., Mise au point sur l’ichtyofaune paléocène de 1988. Menat (Puy-de-Dôme), C.R. Acad. Sci., 1979d, vol. 288, Gheerbrant, E., Thomas, H., Roger, J., Sen, S., and Al- pp. 1461Ð1464. Sulaimani, Z., Deux nouveaux primates dans l’Oligocène Gaudant, J., Nouvelles recherches sur l’ichtyofaune des inférieur de Taqah (Sultanat d’Oman): premiers adapiformes zones salifères moyenne et supérieure (Oligocène inférieur) (?Anchomomyini) de la Péninsule Arabique?, Palaeoverte- du bassin potassique Alsacien, Sci. Géol. Bull., 1981a, brata, 1993, vol. 22, no. 4, pp. 141Ð196. vol. 34, no. 4, pp. 209Ð218. Gheerbrant, E., Thomas, H., Sen, S., and Al-Sulaimani, Z., Gaudant, J., Un nouveau Cyprinodontidae (poisson Nouveau primate Oligopithecinae (Simiiformes) de l’Oli- téléostéen) de l’Oligocène inférieur de Kleinkems (Pays de gocène inférieur de Taqah, Sultanat d’Oman, C. R. Acad. Bade, Allemagne): Prolebias rhenanus nov. sp., Sci. Géol. Sci., 1995, sér. 2a, vol. 321, pp. 425Ð432. Bull., 1981b, vol. 34, pp. 3Ð12. Givulescu, R., Les progrès de l’investigation paléobotanique Gaudant, J., Prolebias catalunicus nov. sp., une nouvelle du Tertiaire de la Roumanie, Rev. Paleobot., Polynol., espèce de poissons Cyprinodontidae de l’Oligocène de Sar- (Amsterdam), 1980, vol. 29, pp. 35Ð48.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S252 POPOV et al.

Glibert, M., Pélécypodes et Gastropodes du Rupelien Hartenberger, J.-L., Étude systématique des Theridomyoidea supérieur et du Chattien de la Belgique, Mém. Inst. R. Sci. (Rodentia) de l’Éocène supériuer, Mém. Soc. Géol. France, Natur. Belg., 1957, no. 137, pp. 1Ð98. 1973b, nov. Ser., vol. 7, no. 117, pp. 1Ð76. Glibert, M. and Heinzelin, B.J., L’Oligocène inférieur Belge, Heissig, K., Changes in the Rodent and Ungulate Fauna in Bruxelles, 1954, vol. 2, pp. 281Ð438. the Oligocene Fissure Filling of Germany, Münchner Geow- Gorbach, L.P., Ichthyofauna and Environments for Forma- iss. Abh. (A), 1987, vol. 10, pp. 101Ð108. tion of Menilithic Beds in the Carpathians, Cand. Sci. Herb, R. and Hekel, H., Nummuliten aus dem Obereozän von (Geol.ÐMineral.) Dissertation, Lvov, 1956, pp. 1Ð16. Possagno, Schweiz. Paläontol. Abh., 1975, vol. 97, pp. 113Ð136. Gorbach, P.L., Fossils of Fishes of the Family Sternopty- Hess, H. and Weiler, W., Untersuchungen an Fischen aus chidae from the Menilithic Formation of the Carpathians, dem Alttertiär der Umgebung von Basel, Ecl. Geol. Helv., Paleontol. Zh., 1961a, no. 4, pp. 168Ð170. 1955, vol. 48, pp. 427Ð447. Gorbach, P.L., Fossil Fishes from the Upper Horizon of the Holroyd, P.A., Simons, E.L., Bown, Th.M., Polly, P.D., and Lower Menilithic Formation in the Vicinity of Chechva Kraus, M.J., New Records of Terrestrial Mammals from the River, Eastern Carpathians, in Geol. Sb. L’vovsk. Geol. O–va, Upper Eocene Qasr el Sagha Formation, Fayum Depression, 1961b, nos. 7Ð8, pp. 421Ð426. Egypt, Palaeovertebrata, 1996, vol. 25, nos. 2Ð4, pp. 175Ð192. Gorbunova, N.N., Taxonomy, Distribution, and Biology of Hooker, J.J., Mammalian Faunal Events in the English Vinciguerria (Pisces, Gonostomatidae), Tr. Inst. Okeanol. Hampshire Basin (Late EoceneÐEarly Oligocene) and Their Akad. Nauk SSSR, 1972, vol. 93, pp. 70Ð109. Application to European Biostratigraphy, Münchner Geow- iss. Abh. (A), 1987, vol. 10, pp. 109Ð116. Gorjanovic′ -Kramberger, D., Beiträge zur Kenntniss der fos- silen Fische der Karpaten, Palaeontograph. Abt. A, 1879, Huang, X., Zapodidae (Rodentia, Mammalia) from the Mid- vol. 26, pp. 51Ð68. dle Oligocene of Ulantatal, Nei Mongol, Verterbr. Palas., 1992, vol. 30, no. 4, pp. 249Ð286. Gorjanovic′ -Kramberger, D., Die jungtertiäre Fischfauna Kroatiens, Beitr. Paläontol. Geol. Österr.ÐUngarns, 1882, Hugueney, M. and Adrover, R., Le peuplement des Baléares vol. 2, pp. 86Ð135. (Espagne) au Paléogène, Géobios, 1982, Mém. Spéc. no. 6, pp. 439Ð449. Gorjanovic′ -Kramberger, D., Die jungtertiäre Fischfauna Karpatens, Beitr. Paläontol. Österr.ÐUngarns Orients., 1884, Hugueney, M. and Adrover, R., Rongeurs (Rodentia, Mam- vol. 3, pp. 65Ð85. malia) de l’Oligocène de Sineu (Baléares, Espagne), Paleon- tol. Evolucio, 1990, vol. 23, pp. 157Ð169. Grachev, N.V., Zhizhchenko, B.P., Kolykhalova, L.A., and Hugueney, M. and Adrover, R., Sacaresia moyaeponsi nov. Kholodilina, T.S., Paleogene Beds in the Central Interfluvial gen. nov. sp., rongeur Thryonomyidé (Mammalia) dans le VolgaÐUral Region, Tr. VNIIgaz, 1971, vol. 31/39Ð32/40, Paléogène de Majorque (Baléares, Espagne), Géobios, 1991, pp. 36Ð45. vol. 24, no. 2, pp. 207Ð214. Gramann, F., Ostracods in: The Nord-West European Ter- Hugueney, M. and Adrover, R., Moissenetia paguerensis n. tiary Basin, Geol. Jahrb., 1988, vol. 100, pp. 225Ð252. gen. n. sp. (Rodentia, Gliridae) de l’Oligocène des Baléares: Gramm, M.N., Some Fresh-Water Ostracodes from the Oli- première apparition du schéma dentaire Eliomys, C. R. Acad. gocene of the Southwestern Kyzyl Kum, vol. 2 of Strati- Sci. (Paris), 1995, sér. 2a, vol. 321, pp. 917Ð922. grafiya i paleontologiya Uzbekistana i sopredel’nykh Ivanova, V.A., Oligocene and Lower Miocene Beds of the raionov (Stratigraphy and Paleontology of Uzbekistan and Southeastern Ustyurt, Southern Aral Coastal Area, and Adjacent Regions), Tashkent: Akad. Nauk UzSSR, 1963, Northeastern Zaunguz Kara Kum, in Tr. VNIIgaz, 1971, pp. 282Ð287. vol. 31/39Ð32/40, pp. 152Ð182. Grande, L., Repeating Patterns in Nature, Predictability and Jerzman′ ska, A., Ichtiofauna l upkow Jaselskich z Sobniowa, “Impact” in Science, Interpreting the Hierarchy of Nature: Acta Palaeontol. Polon., 1960, vol. 5, no. 4, pp. 367Ð420. From Systematic Patterns to Evolutionary Process Theories, ′ Grande, L. and Rieppel, O., Eds., New York: Academic, Jerzman ska, A., Ichtyofaune des couches à Ménilite (Flysh 1994, pp. 61Ð84. des Karpates), Acta Palaeontol. Polon., 1968, vol. 13, no. 3,

pp. 379Ð488. Gregorová,^ R., Rybí fauna menilitového sovrství na lokalité ′ Litenc ice a její stratigrafická pozice, Acta Musei Moraviae, Jerzman ska, A., Oligocene Alepocephaloid Fishes from the Acta Palaeontol. Polon. Scientiae naturales, 1988, vol. 73, pp. 83Ð88. Polish Carpathians, , 1979, vol. 24, no. 1, pp. 65Ð76. Grigoryan, S.M., Nummulity i orbitoidy Armyanskoi SSR Jerzman′ ska, A. and Kotlarczyk, J., Kopalny zespól quasi (Nummulites and Orbitoids from the Armenian SSR), Yere- sargassowy z warstw menilitowych jednostki skolskiej Kar- van: Akad. Nauk Arm. SSR, 1986. pat polskich, Inst. Geol., Kwartalnik geol. (Warsaw), 1975, Grossgeim, V.A., Paleogeography, Paleogene System, Mos- vol. 19, no. 4, pp. 875Ð886. cow: Nedra, 1957, pp. 457Ð472. Jerzman′ ska, A. and Kotlarczyk, J., The Beginning of the Sar- Gruzman, A.D., Foraminifers from the Beds with Globigerina gasso Assemblage in the Tethys?, Palaeogeogr., Palaeocli- vialovi from the Lower Oligocene of the Ukrainian Carpathians, matol., Palaeoecol. (Amsterdam), 1976, vol. 20, pp. 297Ð306. Paleontol. Sb. Lvovsk. Univ., 1987, no. 24, pp. 53Ð59. Jerzman′ ska, A. and Kotlarczyk, J., Zmiany ewolucyjne ich- Gürs, K., Revision der marinen Molluskenfauna des Unteren tiofauny jako podstawa stratygrafii serii menilitowo-kros′- Meeressandes (Oligozän, Rupelium), Mainzer Beckens Dis- nen′ skei, Zemny′ Plyn a Nafta (Bratislava), 1981, vol. 26, no. 1. sertation, Mainz, 1995, pp. 1Ð314. Jerzman′ ska, A. and Kotlarczyk, J., Ichthyofauna Changes in Hantken, M., Die Fauna der Clavulina SzaboiÐSchichten, the Tertiary of the Carpathians and of the Caucasus, Ann. Jahrb. Ung. Geol. Anst., 1875, vol. 4, no. 1, pp. 1Ð93. Inst. Geol. Geoph., 1983, vol. 59, pp. 1Ð8.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S253

Jerzman′ ska, A., Kotlarczyk, J. and Krawczyk, A.J., Introduc- Kordikova, E.G., Oligocene Vertebrate Assemblages in Cen- tion to Quantitative Studies on Variation of Paleogene Ich- tral Kazakhstan: Faunas, Biogeography, and Paleoecology,

thyofauna of the Carpathians, Ann. Soc. Geol. Pol. (Kraków), Historical Biology, 1994, vol. 8, pp. 191Ð208.

1973, vol. 43, no.^ 1, pp. 83Ð117.

^ ^ Kordikova, E.G. and Mavrin, A.V., Stratigraphy and Oli- Jir ic ek, R. and R iha, J., Correlation of Ostracod Zones in the goceneÐMiocene Mammalian Biochronology of the Aktau Paratethys and Tethys, Proc. Shall. Tethys, Saito ho-on Kai Mountains, Dzhungarian Alatau Range, Kazakhstan, Palae- Spec. Publ., 1991, vol. 3, pp. 435Ð457. overtebrata, 1996, vol. 25, nos. 2Ð4, pp. 141Ð174. Jonet, S., Contribution à l’étude des schistes disodyliques Koreczne-Laky, I. and Nagyne-Gellai, A., Foraminiferal oligocènes de Roumanie: La faune ichtyologique de Homo- Fauna from the Oligocene and Miocene in the Borzsony raciu District de Prahova, Lisbon, 1958, pp. 1Ð100. Mountains, Ann. Hung. Geol. Inst., 1985, vol. 68, pp. 1Ð527. Jung, D. and Langer, W., Foraminiferen aus dem Oligozän Kornilova, V.S., The Major Stages in the Evolution of the des Schachtes Lohberg IV bei Hunxe a.d. Lippe (Nierderrhe- Cenozoic Floras in Kazakhstan, Doctoral (Geol.ÐMineral.) inische Bucht), N. Jb. Geol. Paläontol. Abb., 1990, vol. 180, Dissertation, Alma-Ata, 1963, pp. 1Ð51.

no. 1, pp. 75Ð96. ^ Kornilova, V.S., An Outline of the History of the Flora and Veg-

′ ^ Kalabis, V., Ryby se sve teln^ y mi orgány z moravského etation of Kazakhstan, Rastitel’nyi pokrov Kazakhstana (Vege-

paleogénu (menelitovy′ ch br idlic), Casopis Zem. Mus., 1948, tation of Kazakhstan), Alma-Ata, 1966, vol. 1, pp. 37Ð190.

vol. 32, pp. 131Ð174.^ Kotlarczyk, J. and Jerzman′ ska, A., Project to Subdivide the

Kalabis, V., R ád Syngnathiformes^ Berg, 1940 (Pisces) z Menilite and Krosno Beds in the Carpathians into Ichthyo- moravského^ paleogénu, Casopis pro Mineral., Geol., Nakla- faunal Zones, Materialy XI Kongressa Karpato-Balkanskoi datelství Ceskoslovenské Akad. (Prague), 1957, vol. 2, no. 3, geologicheskoi assotsiatsii po stratigrafii (Proc. XI Conf. Car-

pp. 261Ð274. pathian-Balkanian Assoc. Stratigr.), Kiev, 1980, pp. 107Ð115. ^

′ ^ ′ Kalabis,^ V., Historicky pr ehled studia ichthyofauny stars ích Kotlarczyk, J. and Jerzman ska, A., Ichtiofauna w stratygrafii tr etihor Moravy, Zemny Plyn a Nafta (Bratislava), 1981, Karpat, Prz. Geol., 1988, no. 6, pp. 346Ð352. vol. 26, no. 1, pp. 75Ð78. Kraeva, E.Ya., Foraminiferal Assemblages from the Upper Karagyuleva, Yu.D., Fosilite na B”lgariya (Fossils of Bul- Eocene and Oligocene Beds of the South Ukrainian Depres- garia), vol. 6a: The Paleogene Mollusks, Sofia: B”lg. Akad. sion (Western Regions), Paleogenovye otlozheniya yuga Nauk, 1964. Evropeiskoi chasti SSSR (Paleogene Beds in the Southern Keen, M.C., Oligocene Ostracod Biofacies from Onshore European regions of the USSR), Moscow: Akad. Nauk Areas of the North Sea Basin, Northwest European Micro- SSSR, 1960. palaeontology and Palynology, Brit. Micropalaeontol. Soc., Kraeva, E.Ya., Foraminifery Verkhneeotsenovykh i oligotse- Ellis Horwood, 1989, pp. 248Ð264. novykh otlozhenii severnogo kryla Prichernomorskoi Vpad- Kellner, A.W.A. and McKenna, M.C., A Leptictid Mammal iny (Foraminifers from the Upper Eocene and Oligocene from the Hsanda Gol Formation (Oligocene), Central Mon- Beds of the Upper Wing of the South Ukrainian Depression), golia, with Comments on Some Palaeoryctidae, Am. Mus. Kiev: Akad. Nauk Ukr. SSR, 1961. Novit., 1996, no. 3168, pp. 1Ð13. Kraeva, E.Ya. and Maslun, N.V., The Significance of Benthic Khalilov, D.M., Mikrofauna i stratigrafiya paleogenovykh Foraminifers for Division and Correlation of the Paleogene otlozhenii Azerbaidzhana (The Microfauna and Stratigraphy Deposits of the Ukraine, Geol. Zh., 1984, vol. 44, no. 4, of the Paleogene Deposits of Azerbaijan), Baku: Akad. Nauk pp. 107Ð112. Azerb. SSR, 1962. Kraeva, E.Ya. and Pechenkina, L.P., Do ogrunuvannya oli- Khalilov, D.M. and Mamedova, L.D., Zonal’noe podrazdele- gotsenovogo viku “zoni pishchanikh foraminifer” pivnich- nie paleogenovykh otlozhenii Azerbaidzhana (filogeniya nogo Prichernomor’ya (in Ukrainian), Geol. Zh., 1965, vidovogo sostava ryada vazhneishikh rodov foraminifer) vol. 25, no. 5, pp. 113Ð120. (Zonal Subdivision of the Paleogene Beds of Azerbaijan) Krasheninnikov, V.A., Stratigraphy of the Paleogene Depos- (Phylogeny of the Species Composition of Several Foramin- its of Syria, Tr. Geol. Ins. Akad. Nauk SSSR (Moscow), 1964, ifer Genera), Baku: ELM, 1984. vol. 133, pp. 28Ð75. Khisarova, G.D., The Fishes from Continental Deposits of Krasheninnikov, V.A. and Nemkov, G.I., The Relationship the Late Cretaceous and Cenozoic of Central and Eastern between the Faunas of Planktonic Foraminifers and Nummu- Kazakhstan, vol. 5 of Materialy po istorii fauny i flory Kaza- litids in the Paleogene Deposits of Syria, Vopr. Mikropaleon- khstana (Materials on the History of the Flora and Fauna of tol., 1975, vol. 18, pp. 179Ð211. Kazakhstan), Alma-Ata, 1971, pp. 57Ð62. Krasheninnikov, V.A., Muzylev, N.G., and Ptukhyan N.G., Khisarova, G.D., The Eocene Amiidae from the Southwest of Stratigraphic Division of the Paleogene Beds in Armenia Kazakhstan, vol. 6 of Materialy po istorii fauny i flory Kaza- Based on Planktonic Foraminifers, Nannoplankton, and Num- khstana (Materials on the History of the Flora and Fauna of mulitids, Vopr. Mikropaleontol., 1985, vol. 27, pp. 130Ð169. Kazakhstan), Alma-Ata, 1974, pp. 16Ð21. Krasheninnikov, V.A. and Ptukhyan, A.E., The Relationship Kissling, E., Die Fauna des Mitteloligozäns in Berner Jura, between the Faunas of Planktonic Foraminifers and Nummu- Abh. Schweiz. Paläont. Ges., 1896, vol. 22, pp. 1Ð74. litids in the Paleogene Deposits of Armenia, Vopr. Mikropal- Kohring, R. and Schlüter, T., Historische und paläontolo- eontol., 1973, vol. 16, pp. 146Ð182. gische Bestandaufnahme des Simetits, eines fossilen Harz Krasheninnikov, V.A. and Ptukhyan, A.E., Stratigraphic mutmasslich mio/pliozänen Alters aus Sizilien. Docum., Division of the Paleogene Deposits of Armenia Based on Naturae, 1989, vol. 56, pp. 33Ð58. Planktonic Microorganisms and Nummulitids (Regional Kopp, M.L. and Shcherba, I.G., The Caucasian Basin in the Stratigraphy, Zonal Scales, and Their Relationships), Vopr. Paleogene, Geotektonika, 1998, no. 2, pp. 29Ð59. Mikropaleontol., 1986, vol. 28, pp. 60Ð98.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S254 POPOV et al.

Krhovsky, J., Adamova, J., Hladikova, J., and Maslowska, H., Sbornik nauchnykh trudov (Transactions), Tashkent: Nauka Paleoenvironmental Changes across the EoceneÐOligocene^ Uzb. SSR, 1964, pp. 124Ð131. Boundary in the Zdanice and Pouzdrany Units, Knihovnic ka Martini, E., Die Fischfauna von Sieblos/Rhön (Oligozän), ZPN., 1993, vol. 14b, pp. 105Ð187. Senckenb. Lethaea, 1965, vol. 46a, pp. 291Ð314. Kul’chitskii, Ya.O. and Sovchik, Ya.V., The PaleogeneÐNeo- Martini, E., Die oligozäne Fossilfundstätte Sieblos an der gene Boundary in the Carpathians, Paleontol. Sb. (Lvov) Wasserkuppe, Natur. Mus., 1967, vol. 97, pp. 1Ð8. 1986, no. 23, pp. 89Ð99. Martini, E., Neue Insektenfunde aus dem Unter-Oligozän Kuster-Wendenburg, E., Die Gastropoden aus dem Meeres- von Sieblos/Rhön, Senckenb. Lethaea, 1971a, vol. 52, sand (Rupelium) des Mainzer Tertiarbeckens, Abh. Hess. pp. 359Ð369. Landesamt. Bodenforsch., 1973, no. 67, pp. 1Ð178. Martini, E., Standart Tertiary and Quaternary Calcareous Kuster-Wendenburg, E., Bestandsaufnahme der Gastropoden Nannoplankton Zonation, Proc. 2th Planktonic Conf., Roma, im “pra-aquitanien” Tertiär des Mainzer Beckens, Mainzer 1971b, vol. 2, pp. 739Ð785. Geowiss. Mitt., 1982, vol. 10, pp. 83Ð130. Martini, E., Fische aus dem Unteroligozän von Sieblos a.d. Lan, X. and Wei, J., On the Evolution of the West Tarim Gulf Wasserkuppe/Rhön, Beitr. Naturkunde Osthessen, 1988, Based on Late CretaceousÐEarly Tertiary Bivalve Faunas, vol. 24, pp. 149Ð160. Saito Ho-on Kai Spec. Publ., no. 3, pp. 229Ð237. Martini, E. and Müller, C., Das marine Alttertiär in Deut- Lebedev, V.D., The Neogene Freshwater Fish Fauna of the schland und seine Einordnung in die Standart Nannoplank- Zaisan Depression and West Siberian Lowland, Vopr. ton-Zonen, Erde und Kohle, 1971, issue 24. Ikhtiol., 1959, vol. 12, pp. 28Ð59. Marzuk, A.M., Foraminifers and the Evolution of Marine Leriche, M., Les Poissons oligocènes de la Belgique, Mém. Paleoenvironment in the Late Eocene of the Crimea, Cand. Musée R. d’Hist. Nat. Belg., 1910, pp. 233Ð263. Sci. (Geol.ÐMineral.) Dissertation, Moscow: Moscow State Leriche, M., Les Poissons de la molasse suisse, Mém. Soc. Univ., 1992, pp. 1Ð24. Pal. Suisse (Geneva), 1927, vol. 41, no. 1, pp. lÐ55. Melinte, M., Changes in Nannofossil Assemblages during Leriche, M., Les Poissons de la molasse suisse, Mém. Soc. the OligoceneÐLower Miocene Interval in the Eastern Car- Pal. Suisse (Geneva), 1927Ð1928, vol. 47, no. 2, pp. 57Ð119. pathians and Transylvania, Abstr. 10th RCMNS, 1995, Bucharest, Rom. J. Stratigr., 1995, vol. 76, suppl. no. 7, Leriche, M., Note sur les rapports entre la faune ichty- pp. 171Ð172. ologique de l’Argile à Septaria (Septarienton) du Bassin de Mayence et celle de l’Argile de Boom (Rupélien moyen), Menner, V.V. and Riabinin, A.N., Class Pisces, Atlas ruko- suivie d’observations nouvelles sur quelques-unes des vodyashchikh form iskopaemykh faun SSSR (Atlas of the espèces communes aux deux faunes, Bull. Soc. Belg. Géol., Index Forms of the Fossil Faunas of the USSR), vol. 12: Paléontol., Hydrol. (Brussels), 1948, pp. 1Ð57. Paleogen (Paleogene), Moscow: Gosgeolitizdat, 1949, pp. 300Ð311. Lutz, H., Taphozönosen terrestrischer Insekten in aquatis- chen SedimentenÐein Beitrag zur Rekonstruktion des Merklin, R.L. Opredelitel’ dvustvorchatykh mollyuskov olig- Paläoenvironments, Neues Jarhb. Geol., Paläont. Abh., otsena yuga SSSR (Handbook of Bivalve Mollusks from the 1997, vol. 203, pp. 173Ð210. Oligocene of the Southern Regions of the USSR), Moscow: Nauka, 1974. Lopatin, A.V., New Oligocene Didymoconidae (Mesony- chia, Mammalia) from Mongolia and Kazakhstan, Paleontol. Metodika sobytiinoi stratigrafii v obosnovanii korrelyatsii Zh., 1997a, no. 1, pp. 111Ð120. regional’nykh stratonov na primere nizhnego ordovika Severo-Zapada Rossii (Methods of Event Stratigraphy for Lucas, S.G. and Emry, R.J., Biochronological Significance Substantiation of Correlation of Regional Stratones by the of Aminodontidae (Mammalia, Perissodactyla) from the Example of the Lower Ordovician of the Northwest Russia), Paleogene of Kazakhstan, J. Paleontol., 1996, vol. 70, no. 4, St. Petersburg: VSEGEI, 1998. pp. 691Ð696. Micklich, N. and Parin, N., The Fishfauna of the Frauen- Mai, D., Tertiäre Vegetations geschichte Europas, Jena: weiler (Middle Oligocene, Rupelian; Germany): First G. Fischer, 1995. Results of a Review, Publ. Espec. Inst. Esp. Oceanogr., 1996, Mai, D.H. and Walter, H., Die Floren der Haselbacher Serie vol. 21, pp. 129Ð148. im Weisselster-Becken (Bezirk Leipzig, DDR), Abh. Staatl. Minashvili, T., Calcareous Nannofossil Biostratigraphy and Mus. Mineral. Geol. (Dresden), 1978, vol. 28, pp. 1Ð101. Correlation of Oligocene and Miocene Deposits of Georgia, Mai, D. and Walter, H., Die Oligozänen und Untermiozänen Mem. Sci. Geol., 1992, vol. 43, pp. 277Ð282. Floren NW Sachsen und des Bitterfelder Raumes, Abh. Staatl. Moisescu, V., Stratigrafia depozitelor paleogene s i miocen- Mus. Mineral. Geol. (Dresden), 1991, vol. 38, pp. 1Ð230. inferioare din regiunea Cluj-Huedin-Romanasi, An. Inst. Mai, D.H. and Walter, H., Die fossilen Floren des Weissel- Geol. Geofiz., 1975, vol. 47, pp. 1Ð211. ster-Beckens und seiner Randgebiete, Hall. Jahrb. Geowiss. Moisescu, V. and Meszaros, N., Faune à mollusque oli- (Gotha), 1983, vol. 8, pp. 59Ð74. gocènes des couches de Hoia et de Mera (N-W de la Transyl- Maikopskie otlozheniya i ikh vozrastnye analogi na Ukraine vanie) de la collection A. Koch, Bucharest, 1974. i v Srednei Azii (Maykopian Beds and Their Age Equivalents Monostori, M., Oligocene Ostracods from the Surroundings in the Ukraine and in Central Asia), Kiev: Naukova Dumka, of Budapest, Ann. Univ. Sci. Budapest, Sec. Geol., 1982, 1964. vol. 21, pp. 32Ð102. Makarova, R.K. and Tsatsyr E.F., Stratigrafiya paleogena Monostori, M., Ostracods of Eocene/Oligocene Boundary Yuzhnogo Priaral’ya i Kyzylkumov (Paleogene Stratigraphy Profiles in Hungary, Ann. Univ. Sci. Budapest, Sec. Geol., of the Southern Aral Coastal Area and Kyzyl Kum), vol. 3 of 1985, vol. 25, pp. 161Ð243.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S255

Monostori, M., Environmental Changes in Eocene/Oli- Nikolaeva, S.K. and Prussova, I.A., The Paleogene of the gocene Boundary Stratotypes in Hungary Based on Ostracod Northwestern Ustyurt Plateau, Byull. Mosk. OÐva Ispyt. Faunas, Ann. Univ. Sci. Budapest, Sec. Geol., 1986, vol. 26, Prir., Otd. Geol. (Moscow), 1979, vol. 54, no. 3, pp. 25Ð46. pp. 142Ð158. Novitskaya, L.I., The genus Palaeogadus (Gadidae) from the Mrevlishvili, N.I., Nummulity Gruzii i ikh stratigraficheskoe Khadumian Horizon of the Caucasus, Paleontol. Zh., 1961, znachenie (Nummulites from Georgia and Their Strati- no. 4, pp. 120Ð130.

graphic Significance), Tbilisi: Tbilisi Gos. Univ., 1978. Obrhelová, N., Osteologisches Bau von Thaumaturus furcatus Müller, C., Nannoplankton aus dem Mittel-Oligozän Deut- Reuss,^ 1844 (Pisces) aus dem nordböhmischen Süsswasserter- schlands, Belgiens und der Niederlande, Dissertation, Frank- tiär, Cas. Mineral. Geol., 1975, vol. 20, no. 3, pp. 273Ð290. furt am Main, 1970, pp. 44Ð45. Odrzywolska-Bienkowa, E., Wyniki mikrofaunistycznych Müller, C., Nannoplankton-Gemeinschatte aus dem W-deut- osadów Rupelu w Otworze Goleniów IG-2, Przegla d w schen Mittel-Oligozän, Notizbl. Hess., 1971, vol. 99, no. 8, Geol., 1983, vol. 31, no. 7, pp. 425Ð427. pp. 43Ð53. Odrzywolska-Bienkowa, E. and Pozaryska, K., An Outline Muzylev, N.G. and Tabachnikova, I.P., Zonal Subdivision of of the Tertiary Stratigraphy of the Polish Lowland (Based on the Lower Maykopian Beds of Ciscaucasia and Adjacent Foraminiferes), Bull. Acad. Polon. des Sci., Ser. Sci. Terre, Regions Based on Nannoplankton, Sov. Geol., 1987, no. 1, 1981, vol. 29, no. 1, pp. 19Ð22. pp. 65Ð74. Oertli, H., Ostracoden aus der Oligozänen und Miozänen Muzylev, N.G., Tabachnikova, I.P., and Voronina, A.A., The Molasse der Schweiz, Paläontol. Abh., 1956, vol. 74, pp. 1Ð119. Early Oligocene Stage of Brackishing in the Paratethys Okada, H. and Bukry, D., Supplementary Modification and Basins, Izv. Akad. Nauk, Ser. Geol., 1992, no. 5, pp. 39Ð51. Introduction of Code Numbers to the Low-Latitude Cocco- Myatlyuk, E.V., Foraminifery flishevykh otlozhenii Vostoch- lith Biostratigraphic Zonation, Marine Micropaleontol., nykh Karpat (melÐpaleogen) (Foraminifers from Flysch 1980, vol. 5, pp. 321Ð325. Beds of the Eastern Carpathians (CretaceousÐPaleogene), Olteanu, R., Evolution de la Communauté d’Ostracodes dans Leningrad: Nedra, 1970. l’Oligocène du NW de la Transylvanie, Rev. Roum. Géol., Nagy-Gellai, A., Foraminifères de l ’Oligocène dans les envi- Géoph., Géograph., ser. Geol., 1980, vol. 24, pp. 177Ð198. rons de Dorog, Ann. Inst. Geol. Publ. Hungar., 1968, vol. 55, Palamarev, E., Xerotenne Elemente in der Tertiarflora Bul- no. 3, pp. 421Ð617. gariens und Aspecte zum Problem der Formierung der med- diterranen Flora auf der Balkanhalbinsel, Abh. Zentr., Berlin: Nagymarosy, A., Mono- and Duospecific Nannofloras in Geol. Inst., 1967, vol. 10, pp. 167Ð175. Early Oligocene Sediments of Hungary, Palaeontol. Pr., 1983, vol. 86, no. 3, pp. 273Ð283. Paleogen Turkmenii (The Paleogene of Turkmenia), Ashkha- bad: Ylym, 1975. Nagymarosy, A., From Tethys to Paratethys, a Way of Sur- vival, Acta Geod. Geoph. Mont. Hungar., 1990, vol. 25, Paleogenovaya sistema. Stratigrafiya SSSR (Paleogene Sys- pp. 373Ð385. tem. Stratigraphy of the USSR), Moscow: Nedra, 1975. Nagymarosy, A. and Báldi-Beke, M., The Position of the Paleogeographic Atlas of Northern Eurasia, Moscow: Inst. Paleogene Formations of Hungary in the Standard Nanno- Tekton. Litosf. Plit, 1997, maps 40 Ma and 30 Ma. plankton Zonation, Ann. Univ. Sci. Budapest, Ser. Geol., Papp, A., Ljubljane Miogypsinidae aus dem Oligozän von 1988, vol. 28, pp. 1Ð25. Zagorje, vol. 2 of Geologija Rozprave in Porocila, Ljubljana, Nagymarosy, A. and Voronina, A., Calcareous Nannoplank- 1954, pp. 168Ð178. ton from the Lower Maykopian Beds (Early Oligocene, Papp, A., Lepidocyclinen aus Zagorje und Tuhinjska Dolina Union of Independent States), Nannoplan. Res., 1993, vol. 2, Ostlich von Kamrik (Slowenien), vol. 3 of Geologija Roz- pp. 189Ð223. prave in Porocila, Ljubljana, 1955, pp. 209Ð215. Nel, A., Sur la présence du genre Aeschna Fabricius, 1775 Parin, N.N., A New Genus and Species of the Syngnathidae dans les calcaires stampiens de Cereste (Odonata Aeschnidae), from the Lower Oligocene of the Northern Caucasus, Pale- L’Entomologiste, 1986, vol. 42, no. 4, pp. 195Ð198. ontol. Zh., 1992a, no. 2, pp. 132Ð135. Nel, A., Bechly, G., Jarzembowski, E., and Martínez-Delclòs, X., Parin, N.N., A First Find of the Urosphen Fish (Syngnathi- A Revision of the Fossil Petalurid Dragonflies (Insecta: Odo- formes) from the Oligocene of the Northern Caucasus, Pale- nata: Anisoptera: Petalurida), Paleontol. Lombarda, Nuov. Ser., ontol. Zh., 1992b, no. 4, pp. 85Ð89. 1998, vol. 10, pp. 3Ð68. Patterson, C., The Distribution of Mesozoic Freshwater Neuffer, O., Die Bivalven des unteren Meeressandes (Rupe- Fishes, Mém. Mus. Natn. Hist. Nat. Paris, 1975, Ser. A, lium) im Mainzer Becken, Abh. Hess. Landesanst. Bodenforsch., vol. 88, pp. 156Ð174. 1973, no. 68, pp. 1Ð41. Pauca^ , M., Vorläufige Mitteilung über eine fossile Fischfauna Nikitina, Yu.P., Biostratigraphy of the Paleogene Deposits of aus den Oligozänschiefern von Suslanes ti (Muscel), Bull.

Sect. Sci. Acad. Roum., 1929a, vol. 12, no. 4/5, pp. 112Ð120. the Southeastern Part of the Russian Platform and the Scyth- ^ ian Plate, Doctoral Sci. (Geol.ÐMineral.) Dissertation, Le- Pauca , M., Fossile Fische aus dem Rumänischen Tertiär ningrad, 1972, pp. 1Ð46. (vorläufige Mitteilung), Bull. Sect. Sci Acad. Roum., 1929b,

vol. 12, no. 7/10, pp. 208Ð212. Nikolaeva, I.A., Changes in Ostracode Assemblages at the ^ EoceneÐOligocene Boundary, Klimat i fauna kainozoya (Cli- Pauca , M., Die fossile Fauna und Flora aus dem Oligozän mate and Fauna in the Cenozoic), Leningrad: Zool. Inst. von Suslanes ti (Muscel) in Rumänien, Ann. Inst. Geol. Rom., Akad. Nauk SSSR, 1985, pp. 74Ð81. 1934, vol. 16, pp. 575Ð668.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S256 POPOV et al.

Pavlovec, R., Drobne, K., and Sikic, L., Upper Eocene and sion de Hat eg (Roumanie) et la géologie de la région, Oligocene in Jugoslavia, Terminal Eocene Events, 1986, N. Jahrb. Geol. Paläont. Mh., 1976, no. 11, pp. 690Ð698. pp. 109Ð112. Rath, J.G., Beitrag zur Kenntniss der fossilen Fische des Pechenkina, A.P., Micropaleontological Character of the Plattenberges im Kanton Glarus, Ztschr. Dt. Geol. Ges., Lower Oligocene of the Crimea, the South Ukrainian 1859, vol. 2, no. 1, pp. 108Ð132. Depression, and Central Ciscaucasia, Tr. VNIIGaz, 1971, Rayushkina, G.S., Oligotsenovaya flora Mugodzhar i Yuzh- vols. 31/39Ð32/40, pp. 104Ð115. nogo Altaya (Oligocene Flora of Mugodzhary and South Pelaez-Campomanes, P., Valdecollares: a Rodent Fauna from Altai), Alma-Ata, 1979. the Lower Oligocene of the Loranca Basin (Cuenca, Spain), Remy, J.A., Crochet, J.-Y., Sigé, B., Sudre, J., Bonis, L. de, Proc. Kon. Ned. Akad. Wetensch., 1995, vol. 98, no 3, Vianey-Liaud, M., Godinot, M., Hartenberger, J.-L., Lange- pp. 265Ð289. Badré, B., and Comte, B., Biochronologie des phosphorites Peypouquet, J.P., Ostracodes et paléoenvironnements. du Quercy: mise à jour des listes fauniques et nouveaux gise- Méthodologie et application aux domaines profonds du ments de mammifères fossiles, Münchner Geowiss. Abh. (A)., Cénozoïque, Bull. BRGM, 1979, ser. 2, vol. 4, pp. 3Ð79. 1987, vol. 10, pp. 169Ð188. Peypouquet, J.P., Ducasse, O., and Roussele, L., Morphogen- Reshenie kollokviuma po micropaleontologicheskomu obos- esis and Environment, Theoretical and Practical Aspects novaniyu granitsy eotsena i oligotsena na territorii yuga from Hammatocythere: Paleogene Ostracoda of the Aquita- SSSR (Moskva, 1969) (Decision of the Colloquium on the ine Basin, Inter. Symp. Concept, Meth. Paleontol., Barce- Micropaleontologcal Substantiation of the EoceneÐOli- lona, 1981, pp. 173Ð187. gocene Boundary in Southern Regions of the USSR), Mos- Pharisat, A., La paléoichtyofaune du Rupelien marin de cow: VNIGNI, 1970. Froidefontaine (Territoire de Belfort). Taxonomie et popula- Rögl, F., Palaeogeographic Considerations for Mediterra- tions, genèse du gisement implications paléo-biogéographiques, nean and Paratethys Seaways (Oligocene to Miocene), Ann. Ann. Sci. Univ. Fr.-Comté Geol., 1991, vol. 4, no. 2, pp. 13Ð97. Naturhist. Mus. Wien, 1996, vol. 99A, pp. 279Ð310. Pharisat, A. and Micklich, N., Oligocene Fishes in the West- Romanovskii, G.D., A New Genus of a Fossil Fish from the ern Paratethys of the Rhine Valley Rift System, Ital. J. Zool., Caucasus, Zap. Vseross. Mineral. OÐva, 1886, ser. 2, vol. 42, 1998, vol. 65, suppl. no., pp. 163Ð168. pp. 1Ð8. Pokatilov, A.G. and Nikolaev, V.G., Paleogene Beds of the Rozhdestvenskii, A.K., The Genus Centriscus from the May- Central Baikal, Dokl. Akad. Nauk SSSR, 1986, vol. 291, kopian Beds of the Caucasus, Tr. Paleontol. Inst. Akad. Nauk

no. 5, pp. 1203Ð1206. SSSR, 1950, vol. 25, pp. 26Ð55. Pokorny, V., The Genus Krithe (Ostracoda,^ Crust.) in the

Rückert-Ulkümen, N., Tertiäre Fische aus Thrakien und Dar- Paleogene Deep Sea^ Deposits of the Z dánice Unit, Moravia, danellen (Türkei), Rev. Fas. Sci. Univ. Istanbul, 1963, Czechsolovakia, Cas. Mineral. Geol., 1980, vol. 25, no. 4, vol. 28, nos. 1Ð2, pp. 65Ð108. pp. 337Ð349. Popov, S.V., Zoogeography of the Early Oligocene Basins of Russell, D.E., Hartenberger, J.-L., Pomerol, Ch., Sen, S., Western Eurasia Based on Bivalves, Stratigr. Geol. Korre- Schmidt-Kittler, N., and Vianey-Liaud, M., Mammals and lyatsiya, 1995, vol. 3, no. 2, pp. 83Ð97. Stratigraphy: the Paleogene of Europe, Palaeovertebr., 1982, Mem. Extr., pp. 1Ð77. Popov, S.V., Akhmetiev, M.A., Bugrova, E.M., Lopatin, A.V., Amitrov, O.V., Andreyeva-Grigorovich, A.S., Zherikhin, V.V., Russell, D.E. and Tobien, H., Mammalian Evidence Con- Zaporozhets, N.I., Nikolaeva, I.A., Krasheninnikov, V.A., cerning the Eocene-Oligocene Transition in Europe, North Kuzmicheva, E.I., Sytchevskaya, E.K., and Shcherba, I.G., America, and Asia, Terminal Eocene Events, Pomerol, C. and Biogeography of the Northern Peri-Tethys from the Late Premoli-Silva, I., Eds., Amsterdam: Elsevier Sci. Publ., Eocene to the Early Miocene, part. 1: Late Eocene, Paleon- 1986, pp. 299Ð307. tol. J., 2001, vol. 35, suppl. 1, pp. S1ÐS68. Russell, D.E. and Zhai, R.-J., The Paleogene of Asia: Mam- Popov, S.V., Il’ina, L.B., and Nikolaeva, I.A., Mollusks and mals and Stratigraphy, Mém. Mus. Nat. Hist. Natur., 1987, Ostracodes of the Solenovian Horizon of the East Paratethys, Sér. C, vol. 52, pp. 1Ð490. Paleontol. Zh., 1985, no. 1, pp. 28Ð41. Rusu, A., Oligocene Events in Transylvania (Romania) and Popov, S.V., Voronina, A.A., and Goncharova, L.A., Strati- the First Separation of Paratethys, D.S. Inst. Geol. Geofiz., grafiya i dvustvorchatye mollyuski oligotsenaÐnizhnego 1988, vols. 72Ð73, pp. 207Ð223. miotsena vostochnogo Paratetisa (Stratigraphy and Bivalves Rusu, A., Brotea, D., Ionescu, A., Nagymarosy, A., and of the OligoceneÐLower Miocene of the Eastern Paratethys), Wanek, F., Biostratigraphic Study of the EoceneÐOligocene Moscow: Nauka, 1993. Boundary in the Type Section of the Brebi Marls (Transylva- Premoli, Silva I., Coccioni, R., and Montanari, A., The nia, Romania), Rom. J. Stratigr., 1993, vol. 75, pp. 71Ð82. EoceneÐOligocene Boundary in the MarcheÐUmbria Basin Rusu, A., Popescu, G., and Melinte, M., Field Symposium: (Italy), Ancona (Italy), 1988. OligoceneÐMiocene Transition and Main Geological Events Prokofiev, A.M., The sternoptychiidae (Stomiiformes) in the in Romania, 1996, Excursion Guide, Roum. J. Paleontol., EoceneÐLower Oligocene of the Caucasus and Ukrainian 1996, vol. 76, suppl. no. 1, pp. 3Ð47. Carpathians, Vopr. Iktiologii, 2002, vol. 42, no. 1, pp. 23Ð31. Rutte, E., and Van de Weerd, A., Cypriniden Schlundzähne Prokofiev, A.M., The Phosichthyiidae (Pisces, Stomiiformes) west- und südosteuropäischer Tertiär-Lokalitaten, Newslett. from EoceneÐMiocene Deposits of Russia and Adjacent Stratigr., 1980, vol. 8, no. 3, pp. 223Ð231. Areas, Vopr. Ikhtiologii (in press). Samuel, O. and Salaj, J., Microbiostratigraphy and Foramin- Râdulesko, C., Iliesco, Gh., and Iliesco, M., Un Embritho- ifera (On the Slovak Carpathian Paleogene), Bratislava, pode nouveau (Mammalia) dans le Paléogène de la dépres- 1968.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S257

Sandberger, F., Die Conchylien des Mainzer Tertiärbeckens, Starshinin, A.D., Paleogene Beds of the ZeravshanÐGissar Wiesbaden, 1856Ð1863. Mountain Region, Available from NPITsentr, 1997, Du- Saubade, A.M., Remarques sur les relations entre les mol- shanbe, no. 19 (1110). lusques oligocènes de l’Aquitaine et ceux des bassins voisins, Stolyarov, A.S., A Case of Non-compensated Depression Bull. Inst. Géol. Bassin Aquitaine, 1970, no. 8, pp. 239Ð246. within Young Platform in the Oligocene of the Southern Savage, D.E. and Russell, D.E., Mammalian Paleofaunas of Mangyshlak Plateau, Byull. Mosk. OÐva Ispyt. Prir., Otd. the World, Reading: Addison-Wesley, 1983. Geol., 1961, vol. 36, no. 5, pp. 55Ð78. Schaub, H., Nummulites et Assillines de la Tethys Paléogène: Stolyarov, A.S., Paleogeography of Ciscaucasia, the VolgaÐ Taxonomie, phylogenèse et biostratigraphie, Schweiz. Palä- Don Region, and the Southern Mangyshlak Plateau in the ontol. Abh., 1981, vol. 104, pp. 1Ð236. Late Eocene and Early Oligocene, Byull. Mosk. OÐva Ispyt. Prir., Otd. Geol, 1991, vol. 66, no. 4, pp. 64Ð80. Schraft, A., Eine Fischfauna aus den Cerithienschichten bei Stotland, A.B., Spore and Pollen Character of the Mezhigo- Aufschluss Dexheim (Mainzer Becken), , 1979, vol. 30, rian Formation in the Kiev Area of the Dnieper Region, Pale- pp. 369Ð373. ontologiya i stratigrafiya fanerozoya Ukrainy (Phanerozoic Scott, W.B. and Crossman, E.J., Freshwater Fishes of Can- Paleontology and Stratigraphy of Ukraine), Kiev: Naukova ada, Bull. Fish. Res. Board Can., 1973, vol. 184, pp. 1Ð166. Dumka, 1984, pp. 144Ð149. Shcherba, I.G., Paleogeography and Tectonics of the Mayko- Stover, L.E. and Hardenbol, J., Dinoflagellates and Deposi- pian Basin, Caucasus, Dokl. Akad. Nauk SSSR, 1989, tional Sequences in the Lower Oligocene (Rupelian), Boom vol. 306, no. 5, pp. 1196Ð1200. Clay Formation, Belgium, Bull. Soc. Belge Géol., 1993, Shcherba, I.G., Etapy i fazy kainozoiskogo razvitiya vol. 102, nos. 1Ð2, pp. 5Ð77. Al’piiskoi oblasti (The Stages and Phases of the Cenozoic Studencka, B., Popov S.V., and Amitrov, O.V., Oligocene Development of the Alpine Region), Moscow: Nauka, 1993. Molluscan Fauna from the Skibas and BoryslavÐPokut Units Sheremeta, V.G., Ostrakody paleogena Ukrainy (Ostracodes (the Ukrainian Outer Carpathians), Preliminary Report, Car- from the Paleogene of the Ukraine), Lvov: Lvovsk. Gos. pathian Geol. (Bratislava), 1999, vol. 50, Spec. Issue, Univ., 1969. pp. 75Ð76. Shevyreva, N.S., Paleogenovye gryzuny Azii (Paleogene Swidnicki, J., Oligocene Zeiformes (Teleostei) from the Pol- Rodents of Asia), Moscow: Nauka, 1972. ish Carpathians, Acta Palaeontol. Polon. (Warsaw), 1986, vol. 31, nos. 1Ð2, pp. 111Ð135. Shevyreva, N.S., Age of Rodent Fauna (Rodentia, Mamma- lia) from the Buranian Formation of the Zaisan Depression Swidnicki, J., Juveniles of Some Oligocene Antigonia (Eastern Kazakhstan), Stratigr. Geol. Korrelyatsiya, 1995, (Caproïdae, Teleotei) from the Polish Carpathians, Acta Palae- vol. 3, no. 2, pp. 73Ð82. ontol. Polon. (Warsaw), 1988, vol. 33, no. 3, pp. 249Ð259. Switchenska, A.A., A New Species of the Genus Sparus Sigé, B., Les insectivores et chiroptères du Paléogène (Teleostei) from the Lower Oligocene of the Caucasus, Pale- moyen d’Europe dans l’histoire des faunes de Mammifères ontol. Zh., 1979, no. 3, pp. 139Ð141. sur ce continent, J. Paleontol. Soc. India, 1977, vol. 20, pp. 178Ð190. Synthèse géologique du bassin de Paris, vol. 3: Lexique des nom de formation, Mém. Bureau Rech. Géol. Minières, 1980, Sigé, B. and Vianey-Liaud, M., Impropriété de la Grande no. 103, pp. 1Ð467. Coupure de Stehlin comme support d’une limite Éocène-Oli- gocène, Newsl. Stratigr., 1979, vol. 8, no. 1, pp. 79Ð52. Sytchevskaya, E.K., The Fossil Umbridae from the Oli- gocene of Western Siberia, Essays on the Phylogeny and Sys- Simionescu, L., Sur quelques poissons fossiles du Tertiaire rou- tematics of Fossil Fishes and Agnathans, Moscow: Nauka, main, Bull. Arts. Univ. Jassy, 1905, vol. 2, no. 3, pp. 106Ð122. 1968, pp. 162Ð166. Simons, E., Skulls and Anterior Teeth of Catopithecus (Pri- Sytchevskaya, E.K., The Evolution and History of the Dis- mates: Anthropoidea) from the Eocene and Anthropoid Ori- persal of the Esocidae, Byull. Mosk. OÐva Ispyt. Prir., Otd. gins, Science, 1995a, vol. 268, no. 5219, pp. 1885Ð1888. Geol., 1973a, vol. 6, pp. 142Ð143. Simons, E.L. and Wood, A.E., Early Cenozoic Mammalian Sytchevskaya, E.K., The Role of the Beringian Land in the Faunas Fayum Province, Egypt, Peabody Mus. Nat. Hist. Dispersal of the Esocoidei, The Beringian Land and Its Sig- Yale Univ. Bull., 1968, vol. 28, pp. 1Ð105. nificance in the Development of the Holarctic Floras and Skalski, A.W. and Veggiani, A., Fossil Resin in Sicily and the Faunas in the Cenozoic, Khabarovsk, 1973b, pp. 33Ð34. Northern Apennines; Geology and Organic Content, Prace Sytchevskaya, E.K., The Genus Esox in the Tertiary Deposits Museum Ziemi, 1990, vol. 41, pp. 37Ð49. of the USSR and Mongolia, Fauna and Biostratigraphy of Spassov, N. and Lange-Badré, B., Asiovorator altidens gen. the Mesozoic and Cenozoic of Mongolia, Tr. Sovm. Sov.Ð et sp. nov., un mammifère carnivore nouveau de l’Oligocène Mongol. Paleontol. Eksp., 1974, vol. 1, pp. 221Ð234. supérieur de Mongolie, Ann. Paléontol. (Invertébr.ÐVertébr.), Sytchevskaya, E.K., The Fossil Esocids of the USSR and 1995, vol. 81, no. 3, pp. 109Ð123. Mongolia, Tr. Paleontol. Inst. Akad. Nauk SSSR (Moscow), Spiegler, D., Biostratigraphie des Rupels auf Grund von For- 1976, vol. 156, pp. 1Ð116. aminiferen im nördlichen Deutschland, Geol. Jarb., 1966, Sytchevskaya, E.K., The fossil Amiidae from the South of vol. 82, pp. 447Ð486. Mongolia, Tr. Sovm. Sov.ÐMongol. Paleontol. Eksp., 1981, Starobogatov, Ya.I., Fauna mollyuskov i zoogeograficheskoe vol. 15, pp. 7Ð12. raionirovanie kontinental’nykh vodoemov (Molluskan Fauna Sytchevskaya, E.K., Fishes from the Early Paleogene of the and Zoogeographic Division of Intracontinental Basins), Zaisan Depression, Flora and Fauna of the Zaisan Depres- Leningrad: Nauka, 1970. sion, Tbilisi: Metsniereba, 1984, pp. 51Ð61.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 S258 POPOV et al.

Sytchevskaya, E.K., Paleogene Freshwater Ichthyofauna of Age of the Iwaki Formation, Bull. Nat. Sci. Museum, 1986, the USSR and Mongolia, Tr. Sovm. Sov.ÐMongol. Paleontol. vol. 12, no. 4, pp. 165Ð170. Eksp., 1986, vol. 29, pp. 1Ð157. Tugolesov, D.A., Gorshkov, A.S., Meyener, L.B., Solovjev, V.V., Sytchevskaya, E.K., Freshwater Ichthyofauna from the Neo- and Khakhalev, E.M., Tectonics of the South Ukrainian gene of Mongolia, Tr. Sovm. Sov.ÐMongol. Paleontol. Eksp., Depression, Geotektonika, 1985, no. 6, pp. 3Ð20. 1989, vol. 39, pp. 1Ð144. Tyler, J.C. and Bannikov, A.F., A New Genus of Fossil Puff- Sytchevskaya, E.K., Zoogeographical History of the Ceno- erfish (Tetraodontidae, Tetraodontiformes) Based on a New zoic Freshwater Ichthyofauna of Inner Asia, The Main Species from the Oligocene of Russia and a Referred Species Results of the Research of the Joint SovietÐMongolian Pale- from the Miocene of the Ukraine, Proc. Biol. Soc. Wash., ontological Expedition, Moscow: Nauka, 1989, pp. 19Ð21. 1994, vol. 107, no. 1, pp. 97Ð108. Sytchevskaya, E.K., Evolution of the Ichthyofauna of North- Tyut’kova, L.A., New Cylindrodontid (Rodentia, Mamma- ern Eurasia in the Eocene and Oligocene, Geologicheskie i lia) of Indricotherium Fauna, Paleontol. Zh., 1997, no. 6, bioticheskie sobytiya pozdnego eotsenaÐrannego oligotsena pp. 96Ð101. na territorii byvshego SSSR (Geological and Biotic Events in Uffenorde, H., Ostracoden des Tertiär aus der Forschungs the Late Eocene to the Early Oligocene in the Territory of the bohrung Wursterheide (NW Deutschland), Höheres Mittel- Former Soviet Union), Akhmetiev, M.A. and Krasheninni- Ogligozän bis Ober-Miozän, Geol. Jahrb., vol. A111, 1989, kov, V.A., Eds., Moscow: GEOS, 1998, vol. 2, pp. 12Ð32. pp. 367Ð401. Tang, X., A New Fossil Fish from Linli, Hunan, Vertebr. Uhlig, V., Über eine Mikrofauna aus dem Alttertiär der west- Palasiat., 1959, vol. 1, pp. 211Ð213. galizischen Karpathen, Jahrb. D. K. K. Geol. Reichsanst, Ter-Grigorjantz, L.S., Foraminifers of the Maykopian 1886, vol. 36, no. 1, pp. 141Ð214. Deposits in the Stavropol Region and Their Stratigraphic Ünay-Bayraktar, E., Rodents from the Middle Oligocene of Significance, Maikopskie otlozheniya i ikh vozrastnye Turkisch Thrace, Utrecht Micropaleontol. Bull., Spec. Publ., analogi na Ukraine i v Srednei Azii (Maykopian Deposits 1989, no. 5, pp. 1Ð119. and Their Age Equivalents in the Ukraine and Central Asia), Kiev: Naukova Dumka, 1964, pp. 36Ð51. Ünay, E. and Bruijn, H. de., Middle Oligocene to Early Miocene Rodent Assemblages from Turkey, a Preliminary Report, Ter-Grigorjantz, L.S., The Beloglinian and Khadumian Hori- Münchner Geowiss. Abh. (A), 1987, vol. 10, pp. 203Ð210. zons of the Stavropol Region (Upper EoceneÐLower Oli- gocene), Tr. GrozNII (Grozny), 1965, vol. 18, pp. 261Ð285. Unger, F., Sylloge plantarum fossilum, IÐIII Denkschr., K. K. Acad. Wiss., Math.-Nat. (Wien), 1861, vol. L, no. 19, pp. 1Ð48. Terminal Eocene Events, vol. 9 of Developm. Paleontol. Stratigr., Pomerol, Ch. and Premoli Silva, I., Eds., Amster- Van-der-Flerk, I.M., Correlation of Tertiary Beds Based on dam: Elsevier. Sci. Publ., 1986. Biometric Study of the Genus Lepidocyclina, Vopr. Micropa- leontol., 1966, vol. 10, pp. 302Ð308. Théobald, N., Contribution à la paléontologie du Bassin oli- gocène du Haut-Rhin et du Territoire de Beifort. Les Pois- Veselov, A.A. and Lyul’eva, S.A., Age of the Polbian Hori- sons oligocènes, Bull. Serv. Cart. Géol. Als.-Lor. (Stras- zon, Oligocene, in Southern Regions of the USSR, Izv. Akad. bourg), 1934, vol. 2, no. 2, pp. 117Ð162. Nauk SSSR, Ser. Geol., 1980, no. 11, pp. 67Ð71. Théobald, N., Les insectes fossiles des terrains oligocènes de Veselov, A.A. and Sheremeta, V.G., Oligocene Ostracode France, Bull. Mens. Soc. Sci. Nancy, 1937, vol. 2bis, pp. 1Ð473. Fauna and Stratigraphic Occurrence of the Ostracode-Bear- Thomas, H., Roger, J., Sen, S., and Al-Sulaimani, Z., Décou- ing Beds in the Southern Ukrainian and CrimeanÐCaucasian verte des plus anciens “Anthropoides” du continent arabo- Regions, Paleontol. Sb., 1966, no. 3, issue 1, pp. 120Ð128. african et d’un Primate tarsiiforme dans l’Oligocène du Sul- Vikulin, S.V., Paleogene Floras from Tim and Pasekovo tanat d’Oman, C. R. Acad. Sci. (Paris), 1988, ser. 2a, vol. 306, (Southern Regions of the Central Russian Highland), Cand. pp. 823Ð829. Sci. Dissertaion, Leningrad: Inst. Botan., 1991, pp. 1Ð18. Thomas, H., Roger, J., Sen, S., Bourdillon-de-Grissac, C., Vislobokova, I.A., Age of Fauna from the Shand-Gol of and Al-Sulaimani, Z., Découvertes de vertébrés fossiles dans Mongolia and Evolution of Mammalian Fauna in Central l’Oligocène inférieur du Dhofar (Sultanat d’Oman), Géo- Asia in the Oligocene, Stratigr. Geol. Korrelyatsiya, 1996, bios, 1989, vol. 22, no. 1, pp. 101Ð120. vol. 4, no. 2, pp. 55Ð64. Thomas, H., Roger, J., Sen, S., and Al-Sulaimani, Z., The Vislobokova, I., EoceneÐEarly Miocene Ruminants in Asia, Discovery of Moeripithecus Markgrafi Schlosser (Propliop- Mém. Trav. E.P.H.E., Inst. Montpellier, 1997, no. 21, ithecidae, Anthropoidea, Primates) in the Ashwaq Formation pp. 215Ð223. (Early Oligocene of Dhofar Province, Sultanate of Oman), Vislobokova, I.A., Dmitrieva, E.L., and Trofimov, B.A., J. Hum. Evol., 1991, vol. 20, pp. 33Ð49. Ruminants at the PaleogeneÐNeogene Boundary of Mongo- Timon-David, J., Insects fossiles de l’Olgocène infériere des lia, Paleontol. Zh., 1996, no. 1, pp. 99Ð108. Camoins (basin de Marseille), I Diptèrs Brachecères. Bull. Voronina, A.A. and Popov, S.V., The Solenovian Horizon of Soc. Entomol. de France, 1943, vol. 48, pp. 128Ð134. the Eastern Paratethys, Izv. Akad. Nauk SSSR, Ser. Geol., Tolstikova, N.V., Molluscs of Ancient Lakes of the Zaisan 1984, no. 9, pp. 41Ð53. Depression, Fossil Freshwater Molluscs and Their Bearing Wang, B., Zapodidae (Rodentia, Mammalia) from the Lower on the Paleolimnology, Leningrad, 1976, pp. 51Ð256. Oligocene of Qujing, Yunnan, China, Mainzer Geowiss. Tolstikova, N.V., Paleolimnobios, Paleolimnology of Lake Mitt., 1985, no. 14, pp. 345Ð367. Zaysan, Leningrad: Nauka, 1980, pp. 69Ð80. Wang, B. and Emry, R.J., Eomyidae (Rodentia, Mammalia) Tomida, Y., Recognition of the Genus Entelodon (Artiodac- from the Oligocene of Nei Mongol, China, J. Vertebr. Pale- tyla, Mammalia) from the Joban Coalfield, Japan, and the ontol., 1991, vol. 11, no. 3, pp. 370Ð377.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002 BIOGEOGRAPHY OF THE NORTHERN PERI-TETHYS IN THE EARLY OLIGOCENE S259

Wang, B. and Meng, J., Eucricetodon (Rodentia, Mammalia) Weiler, W., Erster Nachweis fon Otolithen der Familie from the Lower Oligocene of Qujing, Yunnan, China, Ver- Umbridae (Pisces) in Tertiär des Mainzer Beckens, mit tebr. Palasiat., 1986, vol. 24, no. 2, pp. 110Ð120. Bemerkungen über die phyletischen Beziehungen innehalb Wang, J.K., Li, G.F., and Wang, J.S., The Early Tertiary Fos- der Unterordnung Esocoidei, Senckenb. Lethaea, 1973, sil Fishes from Sanshui and Its Adjacent Basin, Guandong, vol. 53, no. 6, pp. 455Ð467. Palaeontol. Sinica, Nov. Ser., 1981, vol. 22, no. 160, pp. 1Ð90. Weittstein, A., Über die Fischfauna des Tertiären Glam- Weiler, W., Beiträge zur Kenntnis der tertiären Fische des erschiefers, Abh. Schweiz. Paläontol. Ges., 1886, vol. 13, Mainzer Beckens, Abh. Hess. Geol. Landesanstalt zu Darm- no. 2, pp. 1Ð103. stadt , 1922, vol. 6, no. 2, pp. 71Ð135. Willems, W., Benthonic Foraminifera from the Type-locality Weiler, W., Beiträge zur Kenntnis der tertiären Fische des of the Sands of Grimmertingen (Lower Oligocene of Bel- Mainzer Beckens, Abh., Hess. Geol. Landesanstalt zu Darm- gium), Bull. Soc. Belge Géol., Paléontol., Hydrol., 1972, stadt, 1928, vol. 8, no. 3, pp. 1Ð633. vol. 81, nos. 1Ð2, pp. 27Ð51. Weiler, W., Revision der Fischfauna des Septarientones von Wiesloch bei Heidelberg, Sitz. der Heidelberger Akad. der Willmann, R., Der oligozäne Lebensraum von Sieblos/Rhön Beitr. Naturk. Osthessen Wiss. (Heidelberg), 1931, vol. 11, pp. 1Ð15. im Spiegel seines Insekten, , 1988, no. 24, pp. 143Ð148. Weiler, W., Die Fischfauna der unteren und oberen Meeres- molasse Oberbayerns, N. Jahrb. F. Min., 1932, vol. 68, Witt, W., Ostracoden der bayerischen Molasse (unter beson- part B, pp. 305Ð352. derer Berücksichtigen der Cytherinae, Leptocytherinae, Tra- Weiler, W., Zwei oligozäne Fischfaunen aus dem Königreich chyleberidinae, Hemicytherinae, Cytherettinae), Geol. Bavaria, Ungarn, Geol. Hungar. Ser. Palaeontol., 1933, vol. 5, pp. 1Ð54. 1967, vol. 57, pp. 5Ð120. Weiler, W., Neue Untersuchungen an mitteloligozänen Fis- Woodward, A.S., Catalogue of the Fossil Fishes in the Brit- chen Ungarns, Geol. Hungar. Ser. Palaeontol., 1938, vol. 15, ish Museum (Natural History), London: Brit. Museum Natur. pp. 1Ð31. Hist., 1901, vol. 4. Weiler, W., Die Verbindung des mitteloligozänen Rheintal- Yakovlev, V.N., Distribution of Freshwater Fishes of the grabens mit dem Mittelmeer, Jahresberichte und Mitteilun- Neogene of Holarctic and Zoogeographic Zonation, Vopr. gen des Oberrheinischen Geol. Vereins N.F., 1953, vol. 34, Ikhtiol., 1961, vol. 1, no. 2, pp. 209Ð220. pp. 21Ð29. Yanovskaya, N.M., New Cynodictis from the Middle Oli- Weiler, W., Untersuchungen an Fischen aus dem Alttertiär gocene of Mongolia and Kazakhstan and Evolution of Carni- der Umgebung von Basel, Palaeontol. Teil. Ed. Geol. Helv., vores of the Subfamily Gaminae, Materialy po evolutsii Baie, 1955, vol. 48, no. 2, pp. 430Ð447. nazemnykh pozvonochnykh (Materials on the Evolution of Weiler, W., Die Fischfauna des Unteroligozänen Melanien- Terrestrial Vertebrates), Moscow: Nauka, 1970, pp. 71Ð84. tons und des Rupeltons in der Hessischen Senke, Notizbl. Hess. Landesamt-Amt. Bodenforsch (Wiesbaden), 1961, Yartseva, M.B., Stratigraphy of Oligocene Beds on the vol. 89, pp. 44Ð65. Southeast-facing Slope of the Ukrainian Crystalline Shield (Based on Foraminifer Fauna), Geol. Zh., 1959, vol. 19, Weiler, W., Die Fischfauna des Tertiärs im oberrheinischen no. 3, pp. 25Ð36. Graben, des Mainzer Beckens, des unteren Maintals und der Wetterau unter besonderer Berücksichtigung des Untermi- Zaporozhets, N.I., Palinology and Phytoplankton from the ozäns, Abh., Senk. Naturw. Ges., Frankfurt am Main, 1963, Lower Oligocene of the Northern Caucasus, Stratigr., Geol. vol. 504, pp. 1Ð75. Korrelyatsiya, 1993, vol. 1, no. 1, pp. 128Ð136. Weiler, W., Die Bedeutung der Fischfunde im Rupelton der Zhang, M.-M., Zhou, J.-J., and Qin, D.-R., Tertiary Fish Fauna Tongrube Frauen-Weiler bei Wiesloch südlich Heidelberg, from Coastal Region of Bohai Sea, Mem. Hist. Vertebr. Pale- Zeit. Rhein. Natur. Ges. Mainz, 1966, vol. 4, pp. 17Ð26. ontol. Paleoanthropol. Acad. Sin., 1985, vol. 17, pp. 1Ð60.

PALEONTOLOGICAL JOURNAL Vol. 36 Suppl. 3 2002

View publication stats