Stratigraphy and Geological Correlation, Vol. 2, No. I, 1994, pp. 80 - 87. Translated from Stratigrafiya. Geologicheskaya Korretyatsiya, Vol. 2, No. 1, 1994, pp. 86-94. Original Russian Text Copyright © 1994 by Kuznetsova. English Translation Copyright © 1994 by Interperiodica Publishing (Russia).

Evolution of the Tethyan K. I. Kuznetsova Geobgical Institute, Russian Academy of Sciences, Pyzhevskii per. 7, Moscow, 109017 Russb Received January 31,1993

Abstract - The evolution of foraminiferal assemblages that inhabited the East Mediterranean basins during the Jurassic is discussed using the analysis of the material obtained during my work in Syria from 1986 to 1991 and of the Jurassic foraminifera collections from the adjacent countries, the Crimea, Caucasus, and southwest­ ern Europe. Analysis of the taxonomic composition of the assemblages and their changes through time and space revealed stages of the foraminifera development related to sea basin dynamics in the Jurassic. Faunal assemblages differing in composition and structure are confined to different facies-ecological environments and structural zones. The time range of some stages is nearly equal to the stratigraphic unit of age (stage). According to the biological peculiarities of the fauna, i.e., the dominant morphotypes, rates of evolution, tolerance, inten­ sity of speciation, distribution areas, degree of endemism, two large stages (megaphases) of the Tethyan fora­ miniferal evolution are recognized: the early-middle Jurassic and late Jurassic-early .

The evolution of foraminifera, like any other group of appear in the Jurassic, are rare, but they occur in the organisms, is a multicomponent process, rates and trends deposits of three of the seven studied stages. of which depend on a number of factors: biological In a biogeographic sense, these assemblages contain peculiarities of a group under discussion, its dependence tropical Tethyan endemics, widespread subtropical forms, on a complex of abiotic factors, species migrations, and and cosmopolites occurring outside of the Tethys in the others. Boreal realm (boreal-cosmopolitan elements) (Fig. 1). Most of these factors are unstable during even the Bearing in mind the foresaid, I have distinguished smallest stage in foraminiferal evolution, a zone, in this the main stages of the Jurassic foraminiferal evolution case. The wider the studied distribution area of the in Syria and the adjacent Mediterranean regions. group and the longer the analyzed period of its evolu­ tion, the more complex and reliable its evolutionary The first stage corresponding to the lower Jurassic, patterns. which cannot be divided into stages in Syria, is distin­ guishable in fauna from Morocco and southwestern The vast areas of the Mesozoic Tethys ocean inhab­ France. The stage is characterized by the occurrence of ited by the diverse and abundant Jurassic biota includ­ representatives of the Orbitopsellidae family restricted ing foraminifera are of exceptional interest for such in distribution by the tropical Tethyan area. Associated studies. Pseudocyclammina, Lituosepta, Haurania, and Pfen- Using my own abundant materials, mostly from derina, which are also endemics, have a wider strati­ Syria, partly from Israel, Crimea, Caucasus, and Western graphic distribution and are recorded in the middle and Europe, as well as publications on the other Tethyan partly upper Jurassic of the East Mediterranean. regions, such as the Fiench-Spanish Pyrenees, Algeria, In Syria, the lower Jurassic sediments were depos­ Morocco, Sinai, Saudi Arabia, Iran, and Iraq, I attempted ited in shallow, sometimes brackish-water basins, as to consider the Jurassic foraminiferal evolution in indicated by an impoverished composition of foramin­ the Tethys. iferal assemblages and by the presence of fresh-water Analysis of the foraminiferal assemblages included ostracodes. In die Crimea, this time was characterized studies on their taxonomic composition, facies and eco­ by a deposition of Tavriya Series lacking foraminifers. logical dependence, distribution area sizes of both the The second stage spans the stratigraphic interval of assemblages and principal components (mainly at the the Aalenian-lower Bajocian and is characterized in generic level), and morphological peculiarities of dom­ Syria by a predominance of boreal-cosmopolitan ele­ inant groups. ments (Epistomina, Lenticulina, and Citharina) and According to the dominant groups, the Jurassic less abundant Tethyan endemics (Timidonella). In Tethyan foraminifera belong to the cyclamminid-pfen- southwestern Syria, these assemblages were formed derinid type of Protozoa, (Gordon, 1970; Basov et al., under conditions of the mid-littoral zone. In southwest­ 1972; Basov, 1974; Kuznetsova and Gorbachik, 1985), ern Europe, identical assemblages existed in similar which inhabited the tropical and subtropical basins. settings (Pellissie et al., 1984). Ecologically, these are benthic forms, i.e., facies- The third stage corresponds in time to the Bajocian. controlled organisms. Planktonic foraminifera, which Secreting benthic foraminifera retain their predomi- EVOLUTION OF THE JURASSIC TETHYAN FOl oo sea water Open Shallow- Lagoon and boreal-cosmopolitan forms Tethyan, Subtethyan, 10 10 20 30 40 50 60 70 80 90100% and planktonic species Agglutinated and secreted benthic foraminifera, |o~ol4 S S 5 offoraminifera Families, genera, and species E22 i ESI 2 0 0 5 10 20 30 40 50 60 70 80 90 100 10 20 30 40 50 60 70

1 1 - families; 2 - genera; 3 - species; 4 - agglutinated benthic foraminifera; 5 - secretory benthic foraminifera; 6 - planktonic foraminifera; 7 - boreal-cosmopolitan forms; 8 - Sub­ tethyan forms; 9 -Tethyan forms. Fig. 1. Relations between different taxonomic, ecological, and paleobiogeographic components in the Jurassic foraminiferal thanatocoenoses ofSyria and the basin dynamics.

I < STRATIGRAPHY AND GEOLOGICAL CO] o nance in the assemblages, and the Tethyan elements are Sinai Peninsula were areas of terrigenous-carbonate still of secondary importance. The wide distribution of sedimentation, which preserved assemblages domi­ these assemblages over the Mediterranean enables nated by the boreal-cosmopolitan elements. Here less the correlation of the Bajocian deposits in Algeria, abundant Tethyan forms inhabited the shallow-water Morocco, Sinai, and southwestern Syria, revealing mediolittoral zones and the carbonate platform areas. a common stage in fauna evolution. Northwestern Syria, the French-Spanish Pyrenes, and The fourth (Bathonian) stage corresponds to a wide Morocco were occupied by the shallow-water Tethyan transgression. In the Mediterranean, it is marked by the endemics. Transitional type assemblages with abun­ occurrence of various foraminiferal assemblages, dant boreal-cosmopolitan elements appeared in the which evolved in different parts of the rapidly expand­ Crimea and Caucasus. The planktonic foraminifera ing basin. The boreal-cosmopolitan elements continue advanced in species diversity. to predominate in southern and southwestern Syria, Sinai, and northern Africa, but in central and north­ The sixth stage, marked by significant changes western Syria, southwestern France, and Algeria they in the Mediterranean foraminiferal assemblages, is were of minor importance. Here the Tethyan endemics related to the expansion of the marine transgression in increase sharply in abundance and taxonomic diversity. the Oxfordian. The taxonomic diversity of this group The planktonic foraminifera (Conoglobigerina and reaches its maximum; the secreting benthic forms Globuligerina) appear for the first time. The Syrian increase sharply in importance; and the planktonic for­ foraminiferal assemblages were renewed at the genus aminifera continue to develop. The boreal-cosmopoli­ level. Eighteen genera originated at either the begin­ tan to Tethyan elements ratio is, 9:1 , varying slightly in ning, or in the middle of the Bathonian stage (Fig. 2). different parts of the basin. In the remote regions, the The fifth stage of the foraminiferal development assemblages are characterized by a uniform composi­ spans the Callovian, when there was some contraction tion. The environmental uniformity seems to be charac­ of the marine transgression and an increased differenti­ teristic during the wide transgression that reaches ation of facies-ecological environments in the East the central part of Syria (the Palmyrides). New taxa Mediterranean. Two types of assemblages, inheriting in (Everticyclammina, Choffatella, Mesoendothyra, and general their composition from the Bathonian faunas, Alveosepta) appear and spread over the Crimea, Cauca­ are typical of this stage. Southwestern Syria and the sus, Syria, and Israel. These taxa existed till the end

Tropical Subtropical

Foraminifera Timidonella Torinosuella Charentia Satorina Choffatella Pfenderella Sanderella Steinekella Pseudocyclammina Everticyclammina Stomatostoecha Kilianina Paleopfenderina Feurtillia Paracoskinolina Praekurnubia Kurnubia Protopeneroplis Paravalvulina Pseudomarssonella Phenacophragma Riyadhella Redmondoides Bramkampella Mesoendothyra Melathrokerion Meyendorffina Labyrinthine Dhrumella Amijiella Alveosepta Anchispirocyclina

1 2 3 8 5 6 4 9 1 Haurania1 19 10 11 12 13 14 15 16 17 18 20 21 22 23 24 25 26 27 28 29 30 31 32 33 Kf n jb» 21 28 30 32 27 29 31 33 i r n ■ 19 22 24 26 j2* 1 N T 1 20 23 25 j2bj 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 jf 1 h

Fig. 2. Stratigraphic range of some foraminiferal genera (the tropical and subtropical types of the Tethyan fauna). 1 - one or two species; 2 - three - live species; 3 - more than five species. of the Jurassic, and some of them occurred in the their dominants, rates of evolution, area of distribution, Cretaceous. and intensity of speciations. The beginning of the seventh (Kimmeridgian) stage The first megaphase spans the long interval from the is related to a regressive phase in the development of early Jurassic (Pliensbachian-) to the end of the East Mediterranean basins accompanied by a gen­ the middle Jurassic (Callovian). Foraminiferal assem­ eral structural reorganization of the region. This blages of this megaphase are characterized by: (1) the resulted in an improverished fauna composition and occurrence of highly specialized forms with complex a mosaic pattern of the Kimmeridgian deposits in Syrian exo- and endoskeletal structures, a particular morpho- territory. The deposits occur mostly in its southwestern type, (2) a high degree of fauna endemism at the family part and are very similar to the coeval deposits of Israel level, (3) a reduced tolerance as a result of the high spe­ in composition and depositional environments. In the cialization of the fauna, (4) limited distribution areas of foraminiferal assemblages, the boreal-cosmopolitan genera and families, (5) high rates of genus evolution; elements predominate distinctly over the Tethyan genus life-intervals are 3.3 - 11 Ma, those for species forms. In northwestern Syria, the randomly preserved are, accordingly, 1 -3 Ma, which corresponds to one Kimmeridgian deposits yielded foraminiferal assem­ or two ammonite zones, and (6) highly intensive blages, which included Subtethyan elements (Torino- speciation. suella). The agglutinated benthic species dominated The second megaphase of the Tethyan foraminiferal over die secreted forms. No planktonic foraminifera evolution occurs from the late Jurassic to the beginning have been recovered in the coastal shallow-water sedi­ of the early Cretaceous, having such characteristics as: ments. (1) the occurrence of highly specialized forms along The eighth and final stage of the foraminiferal with more adapted, competitive groups, mainly of the development spans the end of the Jurassic and the family Lituolidae, which became dominants, (2) a dif­ beginning of the Cretaceous, indicating that the Titho- ferent morphotype of dominants, (3) a lesser degree of nian and Berriasian assemblages share a common evo­ endemism of the assemblages (at the genus level), lutionary stage. The deposits of this age have a limited (4) moderate tolerance, (3) broader distribution areas and mosaic distribution in the Near East. The assem­ covering the tropical and subtropical Tethyan regions blages show high taxonomic diversity and retain the (the East Mediterranean, Crimea, Caucasus, North previous relationships between the Tethyan, Sub­ Africa, Central Europe), and (6) moderate rates of evo­ tethyan and boreal-cosmopolitan groups, amid which lution. Generic taxa existed for 17 - 21 Ma. Worth not­ the Subtethyan and cosmopolitan elements dominate, ing, the rates of species evolution did not decrease in while the Tethyan endemics are less abundant. At the comparison to the previous stage, and the species life- beginning of the stage, the generic composition of the periods are 1 - 3 Ma. Tethyan fauna was renewed: six new genera appeared So, these are the peculiarities of evolution of the and were then distributed widely in the Cretaceous. foraminifera, which inhabited the Mediterranean in the These genera belong to the subtropical fauna wide­ Jurassic. spread in the Near East, Crimea, Caucasus, southwest­ The evolutionary trends discussed were revealed by ern Europe, and northern Africa. It is characteristic that thorough studies of the foraminiferal assemblages from the modifications in the foraminiferal composition Syria. occurring near the Jurassic-Cretaceous boundary are as adynamic in the Tethys area as they are in the Boreal Analysis of the composition, structure, and strati­ province. The fauna was renewed only at the species graphic range of the Syrian assemblages suggests the level. New Cretaceous elements appeared at the begin­ following conclusions. ning and in the middle of the late Tithonian. The most abundant and diverse assemblages are When analyzing changes in the foraminiferal confined to the deposits of the Bathonian, Callovian, assemblages in the context of their close relations to the Oxfordian, and Tithonian stages. The Liassic, Aalen- development of the Jurassic Tethyan basins, I refrained ian(?), Bajocian, and Kimmeridgian strata, which have from discussing such important factors as biological limited distribution in Syria, are characterized by less peculiarities, rates of evolution, and the spatial distribu­ abundant, although locally rather diverse and copious tion of this group. assemblages. The quantitative relationship between the benthic forms with secreted and agglutinated tests is Since a rate of evolution of a taxon (a genus in our variable. Most of the stages show a predominance of case, because we deal with the generic categories as far arenaceous over calcareous forms, and only in the as evolution is concerned) varies inversely in relation to Bajocian and Oxfordian did the calcareous foramin­ the duration of its existence, I attempted to define the ifera become dominant in the benthic thanatocoenoses. moments of the rate changes, the relationships between This is related to the occasional invasions of the boreal- the Tethyan foraminifera during the Jurassic, and some cosmopolitan forms, mainly of the Lenticulinidae and morphological trends in their evolution. Vaginulinidae families, dominant in the assemblages The Jurassic evolution of the Tethyan foraminifers from the Boreal provinces, Madagascar, western in the Mediterranean can be divided into two large Indostan, and the Atlantic shelf of Canada. As was pre­ stages, or megaphases, differing in the morphotypes of viously noted, the planktonic foraminifers are scarce and represented by single specimens of Globuligerina Vaginulidae, but no Tethyan endemics. The assem­ bathonicma (Pazdro), G. calloviensis K. Kuzn., G. meg­ blages show more diverse taxonomic composition and anomica K. Kuzn., and G. oxfordiana (Grigelis). include up to 30 species of 11 genera and 7 families. But these species aie of great stratigraphic and correla­ Noteworthy, the appearance of some Ceratobulimini- tive importance, being zonal index species for the dae (Praelamarckina, Lamarckella, and Reincholdella) foraminiferal zones of the Bathonian, Callovian, and and Epistominidae (Epistomina) in the Bajocian. These Oxfordian. They subglobally distributed and can be forms have a broad geographical distribution. How­ used to correlate sections from different paleobio- ever, being benthic organisms, they are indicative of chores: provinces, areas, and belts. facies-ecological environments and are related, gener­ ally, to shallow shelf sediments. Quite a different assemblage was recovered from the Bajocian deposits ASSEMBLAGE ANALYSIS of the Coastal Mountains. It is less diverse than that The earliest Jurassic strata in Syria are of Liassic from the Anti-Lebanon and consists mainly of rare age and contain ostracodes and rare foraminifera. They lenticulins (Lenticulina polymorpha (Terq.), L. veto are exposed in two sections, in the Anti-Lebanon, near Hoffman), marginulins (Marginulina solida Terq.), and the village of Ameh and in the Coastal Mountains, near nodosariids. the village of Jdaida. The first section yielded rare spec­ No fossiliferous Bajocian strata have been found in imens of Trochammina nana Brady known from the the region of Kurd-Dag and Palmyrides. Liassic of western Europe and an ostracode assemblage The Bathonian is marked by a wide marine trans­ including Procytheridea magnicoutensis Apostolescu, gression in the East Mediterranean and many other P. sermoisensis Apostolescu, P. triebel Klingler, P. ver- regions of the world. Deposits of this age are recorded m iculata Apostolescu, Procytheridea sp., Progono- in all the Jurassic sequences of Syria, and contain abun­ cythere strilla Sylvester-Bradley, Aphelocythere kuhni dant and diverse foraminiferal assemblages. Their Triebelel et Klinger, Isobythocypris ovalis Bate and composition differs considerably from that of the Bajo­ Coleman, Cytherelloidea sp., Limnocythere sp. nov. A, cian assemblage, being more diverse: these assem­ and Limnocythere sp. nov. B. The Jdaida section blages include up to 65 species of 42 genera and yielded tests of Involutina liassica (Jones), typical of 24 families. the Liassic strata in Europe and North America. The assemblages became dominated by forms with The middle Jurassic strata are represented by depos­ agglutinated tests, their secretory species are of second­ its of all four stages and contain abundant and diverse ary importance. There are new and scarce planktonic foraminiferal thanatocoenoses. foraminifera of a single species Globuligerina bathoni- The oldest assemblages include Timidonella sarda ana (Pazdro). According to their composition, the Bassoulet, Chabrier et Fourcade, Ammobaculites cob- Bathonian assemblages, which inhabited the Syrian bani Loeblish et Tappan, Lenticulina protracta (Bome- territory, can be classified into two main types. The first mann), Citharina chlatrata (Terquem), C. pauperata type found in the Anti-Lebanon includes abundant (Terquem), and Praelamarckina humilis Kaptarenko, forms of Lituolidae, Lenticulinidae, Polymorphinidae, which are characteristic of the Aalenian and Aalenian- Ophtalmidiidae, Trocholinidae, Ceratobuliminidae, Bajocian. The species list shows the presence of both and planktonic Globuligerinidae. The assemblage is typical Tethyan forms (Timidonella sarda) and boreal- similar in composition to the Bathonian foraminiferal cosmopolitan elements represented by the genera Len­ assemblage from the Sinai Peninsula, the Jebel ticulina, Astacolus, and Citharina, which occur in Maghara Range (Said and Barakat, 1958). The most abundance in western Europe, Madagascar, the Sinai typical forms are Reophax horridus (Schwager), Peninsula, and Morocco. This assemblage was found Recurvoides bartouxi Said et Barakat, Ammobaculites for the first time, present only in the sections of the fontinensis Terquem, A. suprajurassicus (Schwager), Jurassic strata in southwestern Anti-Lebanon, Syria A. hermonensis sp. n., Lenticulina polymorpha (the village of Ameh, the Hermon Mountains, and the (Terquem), Citharina proximo (Terquem), Lamarckella foothills of the Jebel Sheikh Range). The assemblage is antiqua Kapt., and Globuligerina bathoniana (Pazdro). characterized by a high diversity of abundant species. The Bathonian strata of the Coastal Mountains, For example, forms of Timidonella sarda Bassoulet, Kurd-Dag and Palmyrides contain another assemblage, Chabrier et Fourcade are represented in some beds which is dominated by the Tethyan endemics of by dozens of tests of micro- and megalospherical the families Cyclamminidae (Amijiella, Dhrumella), generation. The same is true for the Lituolidae family, (Haurania), Dorothiidae (Eomars- especially for abundant tests of some species o i Am m o­ sonella), Protolixoplectidae (Pseudomarssonella, Riy- baculites. Praelamarckina humilis Kapt., known from adhella), and Dictioconidae (Kilianina, Meyendorffina, the Aalenian in the platform part of the Ukraine and Paracoscinolina). The family Pfenderinidae is repre­ northwestern margins of the Donets basis, is represented sented by numerous species of the genera Paleo- by sporadically occurring small-sized specimens. pfenderina, Pfenderella, Praekumubia, and Kumubia, The Bajocian assemblages inherit particular fea­ which is of the utmost importance in this case. tures from their predecessors. They also include abun­ The succession of foraminiferal assemblages dant boreal-cosmopolitan forms of Lenticulinidae and enables us to subdivide the Bathonian strata of the Coastal Mountains, with the most abundant and diverse Bathonian, and contain representatives of 16 families, fauna into lower and upper substages consisting of sev­ 29 genera, and nearly 60 species. eral zones or fauna beds. A distinct sequence of the assemblages and steady taxonomic composition is Like the Bathonian, the Callovian is also character­ characteristic of the whole distribution area of this ized by two types of assemblages: in Anti-Lebanon, the fauna type. The typical early Bathonian species forms of wide geographic distribution are still domi­ include: Haurania deserta Henson, Amijiella amii nant, whereas the Coastal Mountains were inhabited (Henson), Redmondoides lugeoni (Septfontaine), mainly by the Tethyan endemics. The composition of Pseudomarssonella primitiva Redmond, Riyadhella the Callovian assemblages is insufficient to define two arabica Redmond, Pmtopenemplis striata Weyn schenk, units of this stage in Syria (as in the Crimea). In sec­ Paleopfenderina tmchoidea (Smout et Sugden), tions of different structural zones the stage varies, but in Dhrumella evoluta Redmond, and Globuligerina complete and continuous sections the main species bathoniana (Pazdro). Noteworthy is the almost com­ assemblage can be traced as essentially consistent from plete absence of boreal-cosmopolitan elements typical the uppermost Bathonian to the base of the Oxfordian. of the southern Hermon assemblage. In Anti-Lebanon, the most characteristic species are Lituotuba nodus Kos., Kumubia palastiniensis Henson, Thus, the beginning of the Bathonian is marked by Paleogaudryina varsoviensis Biel, et Poz., Nautilocu- significant modifications in the foraminiferal assem­ lina colithica Mohler, and Ammobaculites fontinensis blages, the appearance of new generic taxa of the (Terquem). Tethyan endemics (Fig. 1), and the decreasing impor­ tance of boreal-cosmopolitan elements. The assemblage from the Coastal Mountains The late Bathonian shows the same trends, four new includes: Ammodiscoides magharaensis Said et genera appear near the early-late Bathonian boundary, Barakat, Sanderella laynei Redm., Steinekella crusei in addition to the still existing genera known from Redm., Kumubia palastiniensis Henson, Paleopfende­ the early Bathonian (Paleopfenderina, Pfenderella, rina salemitana (Sart. et Cresc.), Astacolus pellucida Satorina, Sanderella, Dhrumella, Haurania, Amijiella, Said et Barakat, Frondicularia spissa (Terq.), and Rein- Protopenemplis, Pseudomarssonella, Riyadhella, and choldella dreheri Bart. Redmondoides). The boundary between the substages The Oxfordian was the expansion period of the is defined by the renewal of species composition, a less marine transgression, which resulted in wide distribu­ important role of the genera Haurania and Am ijiella, tion of deposits of this age in the East Mediterranean, and the appearance of species of the above listed new including Syria. These deposits contain extremely rich Tethyan genera. The most typical late Bathonian spe­ faunal communities, the most important components of cies include: Kilianina blanched Pfen., Meyendorffina which are foraminifera. Their assemblages are charac­ bathonica Auroze and Bizon (zonal index species), terized by high taxonomic diversity, a uniform compo­ Paracoscinolina occitanica Peyb., Praekumubia sition over vast areas (as a result of extensive fauna crusei Redm., Kumubia bramkampi Redm., Berthe- migrations), high density of most species populations linella paradoxa (Berth.), Limognella dufaurei Pell, reflecting the environmental stability of the paleoba- and Peyb., Paleopfenderina salemitana Sart. and sins, the occurrence of planktonic forms, a higher abun­ Cresc., Mesoendothyra croathica Gusi<5, Pseudomars­ dance of secreted benthic forms as compared to sonella bipatriata Redm., and Globuligerina bathoni­ agglutinated species and the dominance of widespread ana (Pazdro) (Pellisse et al., 1984). boreal-cosmopolitan components over the Tethyan The beginning of the Callovian is characterized by endemics. These peculiarities are typical of foramin­ a contracting transgression and impoverished foramin­ iferal assemblages from all areas of the Oxfordian iferal assemblages with still evolving Bathonian spe­ deposits in Syria. Their taxonomic diversity (28 fami­ cies and genera. The Callovian assemblages are clearly lies, 46 genera, and more than 100 species) is high inherited from the Bathonian foraminifers. Most of the enough to reveal successive assemblages, to subdivide late Bathonian species continued to exist in the Callov­ the Oxfordian into lower and upper substages, and to ian, when no new genera originated. The Tethyan distinguish their foraminiferal zones. The early Oxfor­ endemics are still of prime importance in the assem­ dian foraminiferal assemblages from the Anti-Lebanon blages, but there are also some forms of wide geo­ sections (Hadar, Rowda, and Wadi Al-Kam) contain the graphical distribution: Polymorphinidae (Eoguttulina, following species: Alveosepta jaccardi (Schrodt), Globulina) Ammodiscidae (Ammodiscoides), Ver- Ammobaculites coprolithiformis (Schwag.), Pseudocy- neuilinidae (Paleogaudryina), and Haplophragmoidii- clammina maynci Hott., Mesoendothyra sp., Lenticu- dae (Haplophragmoides). During the evolution of the lina brueckmanni (Mjatl.), L. quenstedti (Guemb.), planktonic foraminifera, Globuligerina bathoniana L. russiensis Mjatl., Astacolus vacellantes Esp. et (Pazdro) is replaced by the Gallovian species Globuli­ Sigal, Planularia beierana (Guemb.), Citharina enty- gerina meganomica K. Kuznetsova and G. calloviensis pom atus Esp. et Sigal, C. paralella Biel, et Poz., Mar- K. Kuznetsova. They are still rare, but occur both in ginulinopsis suprajurassicus (Schwager), Conorboides purely carbonate and in terrigenous-carbonate deposits. paravalendisensis Reiss, Spirillina kuebleri Mjatl., Callovian foraminiferal assemblages dominated by Epistomina nemunensis Grig., E. porcellanea agglutinated forms are less diverse than those of the Brueckm., and Globuligerina oxfordiana (Grig.). The assemblage consists mainly of widespread In the rest of the Syrian territory, there was either no secreted benthic forms known from the Boreal prov­ Kimmeridgian sedimentation (Kurd-Dag), or sediments ince, Madagascar, the Sinai Peninsula, western Europe, of this age were eroded as a result of the pre-Cretaceous Canada, and the Crimea. This permits us to correlate uplifts, as evidenced by ostracode and foraminiferal fau­ deposits of the lower Oxfordian Lenticulina brueck- nas redeposited in the Cretaceous strata of some local­ manni-Globuligerina oxfordiana zone in european ities (the Wadi Jahannam section, Coastal Mountains). Russia, the Crimea, and Caucasus {Stratigrafiya i Kor- relyatsiya..., 1985). The Tithonian deposits are preserved in only a few sections of Syria because the pre-Cretaceous uplifts The lower Oxfordian strata grade into the mid­ involved a major part of the territory. They were studied dle-upper Oxfordian beds. Their foraminiferal assem­ in the Anti-Lebanon (the Wadi Al-Kam, Sad Al-Kam, blages were clearly inherited from the earlier fauna. and Wadi Favuar sections) and the Coastal Mountains The species composition is renewed by the following (the Qadmous section), The deposits yielded abundant late Oxfordian forms: Reophax metensis Franke, assemblages of high species diversity. They contain the Haplophragmium lutzei Hanzl., Ammobaculites brun- widespread zonal species Anchispirocyclina lusitanica steini Cushm., Trocholina transversarii Paalz., Lenticu­ (Egger) and Melathrokerion eospiralis Gorb. The cor­ lina quenstedti (Guemb.), L audax Loebl. et Tapp., and responding zone is recognizable in Syria, the Crimea, Steinekella steineki Redm. The last-mentioned species Caucasus, and probably in Morocco. The incomplete­ belongs to the Paleopfenderinidae family, peculiar to ness of the Tithonian strata prevents a detailed subdivi­ the Tethys. The Tethyan and Subtethyan endemics are sion. The Tithonian foraminiferal assemblages from only represented by the genera Steinekella, Pseudocy- Anti-Lebanon are characterized by a mixed composi­ clammina, and Everticyclammina. The genera are tion including both the Tethyan and Subtethyan endem­ poorly diversified at the species level, but in some beds ics and cosmopolitan forms, namely: Gaudryina vadazi their forms are very abundant. Cushm. et Glaz., Stomatostoecha compressa Gorb., The taxonomic stability of the late Oxfordian fora­ Melathrokerion eospirialis Gorb., Alveosepta powersi miniferal assemblages is reflected in the widespread Redm., Vemeuilina angularis Gorb., Marssonella hehti zones distinguished in Syria, which can be considered Dieni et Mass., Anchispirocyclina lusitanica (Egger), as units of a common stratigraphic scale. Similar, Pseudocyclammina sp. 1, and Kumubia jurassica Red­ though less diverse foraminiferal assemblages were mond. The Tithonian strata of the Coastal Mountains recovered from the Oxfordian strata of the Palmyrides, (the Qadmous section) produced an assemblage differ­ despite the fact that previously they were thought to be ing in species composition and dominated by the missing here (Jarmakani et al., 1989). Tethyan endemics Bramkampella arabica Redm. and the Subtethyan species: Choffatella decipiens Schlum., In Syria, the Kimmeridgian is marked by the begin­ Anchispirocyclina lusitanica (Egger), Pseudocyclam­ ning of a structural reorganization, extensive uplifts, mina parvula Hott., Feurthillia frequens Maync, Tori- and sea basin contraction, which influenced the compo­ nosuella sp., and others. All these species are very sition, structure, and spatial distribution of foramin­ abundant. In addition, the assemblage includes the iferal assemblages. Tithonian forms, which appeared in the Tithonian and Occurrences of Kimmeridgian strata are restricted then evolved in die Cretaceous. This section is likely to mainly to Anti-Lebanon (the Wadi Al-Kam and Wadi exhibit the latest Jurassic strata (the late Tithonian) and Favuar sections). The shallow-water sediments here are the beds transitional to the Berriasian. predominantly clastic, oolitic, and stromatoporitic Thus, the analysis of the development of the Jurassic limestones, and contain impoverished foraminiferal foraminiferal assemblages allows the following conclu­ assemblages: Reophax hounstoutensis Lloyd., Tripla- sions. sia jurassica Mjatl., Marssonella doneziana Dain., M. hehti Dieni et Massari, Citharina lepida (Schwag.), (1) Study of the foraminifera evolution in relation to and Alveosepta personata (Tobler). The last form is the dynamics of the Mediterranean sea basins revealed an index species from a zone of the same name, which a number of stages different in the composition, struc­ occurs widely in the Mediterranean carbonate facies. ture, and distribution of their assemblages. Many of This form is the only “Tethyan” species in the assem­ these stages correspond in duration to such stratigraphic blage, whereas the rest are represented by widespread units as a stage. Biological peculiarities of the studied euryfacies forms. A similar assemblage with more foraminifera enable the recognition of two large Tethyan elements occurs in the deposits of the Coastal evolutionary stages (megaphases); early-middle Juras­ Mountains (the Nicola quarry), here defined as Kim­ sic and late Jurassic-early Cretaceous. These stages are meridgian. The assemblage contains Ammobaculites different in their morphotypes of dominants, degree of pellucida Said et Barakat, Pseudocyclammina sp., endemism, tolerance, rates of evolution, intensity of Alveosepta personata (Tobler), Nautiloculina oolithica speciation, and area distribution sizes of the foramini- Mohler, and Lenticulina subtilis (Wins). It includes in fers. The results show that the biological transforma­ general 14 families, 20 genera, and 25 species, most of tions of the fauna were slower than the changes in which are agglutinated forms; planktonic species are the abiotic factors that influenced the basins that were lacking. their habitat. (2) The most distinct steps in the fauna renewal are Pellissie, T., Peybemes, B., and Rey, J., The Larger Benthic recorded at the beginning of the Bathonian, in the late Foraminifera from the Middle/Upper Jurassic of SW France Bathonian, at the beginning of the Oxfordian, and at the (Aquitaine, Causses, Pyrenees), Biostratigraphic, Paleoeco- beginning of the Tithonian. All these boundaries are logic, and Paleobiogeographic Interest, Benthos*83, 2nd marked by appearance of new taxa of a generic rank. Symp. Int, Benthic Foraminifera, Pau, Bordeaux, 1984, pp. 479 - 489. (3) All genera appearing in the middle Jurassic (Aalenian-Callovian) are tropical endemics. Beginning Said, R. and Barakat, M.G., Jurassic Microfossils from Gebel Maghara, Sinai, Egypt, Micropaleontology, 1958, vol. 4, from the late Jurassic (Oxfordian-Tithonian) till the no. 3, pp. 231 - 272. early Cretaceous (Berriasian-Barremian), new elements of the assemblages are represented exclusively by Basov, V.A., On Some Characteristics of Foraminifera Geo­ Subtethyan genera, which had wider distribution areas graphical Spreading in Jurassic, Paleogeografiya Severa outside of the Tethys. Evrazii v Mezozoe (Mesozoic Paleobiogeography of North­ ern Eurasia), Moscow: Nauka, 1974, pp. 63 - 76. (4) In the Mediterranean, the changes in the fora- miniferal assemblages near the Jurassic-Cretaceous Basov, V.A., Vakhrameev, V.A., Krymgol’ts, G.Ya., Mesezh- nikov, M.S., Saks, V.N., and Shul’gina, N.I., A Problem of boundary were of the same nature as in the Boreal prov­ Continental Drift in the Jurassic and Cretaceous According to ince. The fauna was renewed at the specific level only, Paleobiogeographic Data, Paleontologiya, Mezhdunarodnyi the Cretaceous and Jurassic assemblages are very sim­ Geologicheskii Kongress, XXIV Sessiya, Doklady Sovetskikh ilar, new elements (genera) appeared mainly in the Geologov (Paleontology, XXIV Int. Geol. Congr.: Rep. Sov. Tithonian, and then evolved in the early Cretaceous. Geologists), Moscow: Nauka, 1972, pp. 104 -113. Kuznetsova, K.I. and Gorbachik, T.N., Stratigrafiya i Fora- REFERENCES minifery Verkhnei Yury i Nizhnego Mela Kryma (Upper Jurassic and Lower Cretaceous Stratigraphy and Foramini- Gordon, W.A., Biogeography of Jurassic Foraminifera, Bull fers of Crimea), Moscow: Nauka, 1985. . Am., 1970, vol. 81, no. , pp. 1689 - 1704. Geol. 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