Circum-Mediterranean Oligo/Miocene Biogeographic Evolution

Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 www.elsevier.com/locate/palaeo

Circum-Mediterranean Oligo^Miocene biogeographic evolution ^ the gastropods’ point of view

Mathias Harzhauser a, Werner E.Piller b;Ã, Fritz F.Steininger c

a Naturhistorisches Museum Wien, Geologisch-Pala«ontologische Abteilung, Burgring 7, A-1014 Vienna, Austria b Institut fu«r Geologie und Pala«ontologie, Universita«t Graz, Heinrichstrasse 26, A-8010 Graz, Austria c Naturmuseum Senckenberg, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany

Received 2 April 2001; accepted 24 October 2001

Abstract

Based on studies in Iran (Qom Basin, Esfahan^Sirjan Basin, Zagros Mountains), in Turkey (Mut and Sivas basins), in the Mesohellenic Basin, and in northeastern Egypt, a new palaeobiogeographic concept for the Oligocene and Miocene in the circum-Mediterranean area with special emphasis on the distribution patterns of gastropod faunas is presented.A very strict biogeographic terminology is proposed to avoid the common fusion of geographic, faunistic and geodynamic terms in the palaeontologists’ jargon.Our main interest focuses on the affinities and the faunistic interactions between the early ‘Mediterranean’ and the early ‘Indo-Pacific’ regions during the Oligo^Miocene. Especially the role of the early Indo-Pacific faunas as the mythic centre of origination of Early Neogene European mollusc faunas, as often vaguely indicated in the literature, is critically evaluated.This leads to the establishment of a MediterraneanÎranian Province and a Western IndianÊastern African Province as palaeobiogeographic subunits of the Western Tethys Region during the Oligocene.Due to major geodynamic and related biogeographic changes in the Early Miocene the Western Tethys Region no longer existed and was replaced by the Proto-MediterraneanÂtlantic Region.For the area of the Paratethys the palaeobiogeographic unit Danubian Province, which includes the Proto- Caspian Subprovince, is proposed.Furthermore, the underrated transatlantic communication of the Oligo^Miocene nearshore mollusc faunas is demonstrated.Altogether this paper is a contribution to the discussion on the biogeographic concepts, classification and nomenclature in palaeontology as initiated by Westermann [Palaeogeogr.Pa- laeoclimatol.Palaeobiogeogr.158 (2000) 1^13; 163 (2000) 49^68]. ß 2002 Elsevier Science B.V.Allrights reserved.

Keywords: biogeography; gastropods; Indo-Paci¢c; Mediterranean; Paratethys; Atlantic

1. Introduction tral Europe has been produced (e.g. Ho«rnes, 1856; Bellardi, 1872^1890; Ho«rnes and Auinger, During the 19th and 20th centuries a large da- 1879^1891; Sacco, 1890^1904; Cossmann and taset for Cenozoic gastropods from the Western Peyrot, 1909^1934).This led to an early under- Mediterranean, the European Atlantic and Cen- standing of Eocene to Miocene biogeography of this area.In an eastern direction, however, information concerning the Eastern Mediterranean * Corresponding author. and the Middle East was scanty.Gastropods E-mail address: [email protected] (W.E. Piller). were documented as rare byproducts during inves-

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 104 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 tigations of regional geology.Due to the strong biogeographic entities, was thus severely ham- preservational bias related to the predominantly pered by a palaeontological no man’s land span- studied calcareous facies, the mollusc studies fo- ning the entire eastern Mediterranean and the cused mainly on oysters and pectinids (Douglas, connecting Tethyan seaway.Therefore, modern 1927, 1928).On the other hand, scattered and palaeobiogeographic interpretations of late Te- inhomogeneous data on Eocene to Miocene gas- thyan mollusc faunas (Piccoli, 1984; Piccoli et tropod faunas from Pakistan, Somalia, Kenya, al., 1986, 1991) were usually based on the well- Madagascar, Southern India, Burma, and Java studied Northern Italian and Pakistanian faunas. (Noetling, 1901; Martin, 1914^1915, 1916^1917, The vast area in-between was taken into consid- 1921^1922; Davies, 1923; Vredenburg, 1925^ eration only cursorily since their poorly docu- 1928; Collignon and Cottreau, 1927; Cox, mented and low diversity faunas were considered 1930a; Azzaroli, 1958) allowed a glance at the to be of little palaeobiogeographic value. composition and evolution of the early Indo- To close this gap we focused on this poorly West Paci¢c faunas.The comparison of Mediter- studied area.Detailed investigations of Oligocene ranean and Asian faunas, representing two major and Early Miocene sections in the Greek Meso-

Fig.1.Geographic location of the herein reported Oligocene and Early Miocene gastropod faunas.The faunas from Iran, Turkey and Greece form the base for the present study, whereas the low diversity and poorly preserved faunas from Egypt and Oman are only supplementary.Additionally, the position of some selected European and Asian reference faunas is indicated.Among these, the Italian and Pakistanian gastropod faunas turned out to represent the cornerstones for biogeographic considerations in the circum-Mediterranean area.

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 105 hellenic Basin, the Turkish Mut and Sivas basins to Lattin’s subregion whilst the subrealm of Wes- and especially the Iranian Qom and Esfahan^Sir- termann (2000b) seems to be more or less synon- jan basins (Fig.1) yielded a high number of shal- ymous to ‘region’ of most biogeographers.Long- low marine gastropod faunas.Hence, the faunistic hurst (1998) unites the provinces in biomes. gap in the Eastern Mediterranean was ¢lled and However, the geographic and biogeographic data from Iran, which is identi¢ed as key area boundaries of the modern shallow-water subdivi- between Mediterranean and Paci¢c faunas, are sions are more or less the same for all authors. available for the ¢rst time.Based on these data Recently, biogeographic principles and the de- a new concept of the palaeobiogeography of the nomination of units and their hierarchical ranking Oligocene and Early Miocene Western Tethys can were discussed by Westermann (2000a,b).In his be developed. ‘Rules of Biogeographic Nomenclature’, Wester- The main interest focussed on the dawn of the mann (2000a) proposes a strict use of biogeo- fauna of the Mediterranean^Iranian Province.Its graphic terms, with emphasis on homonyms and birth from the Oligocene faunal-stock of the vast synonyms as well as stability throughout geolog- Tethys Realm is outlined, and the in£uence of the ical time.Westermann (2000a) tries to establish Proto-Indo-West Paci¢c Region on the Early the general term ‘biochore’ for all biogeographic Miocene gastropod faunas of the Iranian Qom units.The term ‘biochore’ is, however, highly con- Basin and the faunas of the Mediterranean area troversially used: for many neontologists ‘bio- is evaluated. chore’ has a strong ecological component without geographic restrictions (e.g. biochores are grass- land, forest, and desert) and even for palaeontol- 2. Biogeographic basics ogists such as Lehmann (1996) ‘Biochorion’ is a small area within a biotope (e.g. a treetrunck or a A summary of the various concepts and hierar- carcass).Thus a biochore in modern biology ^ chies in biogeography is given in Sedlag and Wei- especially in botany ^ is a subdivision of the bio- nert (1987), who recommend a terminology corre- cycle which contains a group of biotopes that re- sponding to Lattin (1967).According to these semble one another.This diverging use of ‘bio- authors the usually rather vaguely used term chore’ is based on the unclear translation of the ‘province’ should be con¢ned to a part of a sub- Greek bio-‘MgbYc’ as living-‘space’ instead of liv- region.The latter term corresponds largely to the ing-‘place’.Therefore, not to prolong the confu- term ‘province’ as used by Woodward (1856) and sions, we prefer ‘biogeographic unit’ instead of Hedgpeth (1957).Finally, the subregions are the term biochore. united into large biogeographic units ^ the re- Westermann (2000a,b) recommends the ranks gions. Superrealm, Realm, Subrealm, Province, and Sub- A strict de¢nition of the various biogeographic province as valid hierarchical categories for the units based on percentages of endemisms is pre- de¢nition of his biochores.The term region is sented by Kau¡mann (1973) and Kau¡mann and for informal use only.We believe, however, that Scott (1976).According to this de¢nition a realm the well-established and widely used term ‘region’ is characterised by more than 75% endemism, a should be obligatory and not replaced by an op- region comprises 50^75%, a province 25^50%, a tional term as ‘Subrealm’.Therefore, in this paper subprovince 10^25% and ¢nally an endemic centre the terms province and region are mainly used reaches 5^10% endemism. sensu Briggs (1995).In contrast, we follow Wes- Generally, the term province as proposed by termann (2000a) in his proposed rule, that the Sedlag and Weinert (1987) is accepted by modern replacement of names is acceptable under certain biogeographers including Briggs (1995), Long- conditions, e.g. extreme biochore expansion or hurst (1998), and Westermann (2000a,b).None- contraction, exceptional geologic change at re- theless the classi¢cation of higher ranks is in a gional or global scale, and at mass extinction state of £ux.Briggs uses ‘region’ as a synonym events.

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The hierarchical categories of biogeographic Papp et al.(1973) and Harzhauser (2002).In con- ranks used in this paper are: trast, the Pakistanian faunas with 77 Oligocene Realm ^ Region ^ Province. and 81 Early Miocene species described by Vre- For re¢nement these biogeographic core-units denburg (1925^1928) are considerably smaller. can be subdivided by adding the pre¢xes Super- The discrepancy is even more obvious when east- or Sub- respectively. ern African and southern Indian gastropod faunas are considered.Thus, only 23 species are described from the Oligocene and Early Miocene of 3. Materials and methods Somalia (Azzaroli, 1958), 28 from the Miocene of Ceylon (Cox, 1930a), 13 from the Miocene of The shallow marine Late Oligocene Chattian Madagascar (Collignon and Cottreau, 1927), faunas from Mesolouri, Doutsiko and Kipourio and only eight from the Miocene of Kenya (Da- in the Mesohellenic Trough (Greece) (Fig.1) vies, 1923). yielded more than 90 taxa, representing about The palaeobiogeographic interpretation of these 50 species.An equal number of species was re- inhomogeneous faunas based on statistical meth- corded from Abadeh in Iran.The Early Miocene ods seems to be inappropriate.On the one hand, nearshore faunas from Turkey, Greece and Iran for many of the mentioned reference faunas mod- are documented by 145 taxa composed of 46, ern revisions are completely missing.Therefore a 50 and 58 gastropod species respectively (Appen- reliable comparison of literature-based faunas is dix 1).These faunas were compared and coin- di⁄cult, a fact that is obvious to every taxonomist cidence or di¡erences between the faunas on spe- after a short look at the usually long and mani- cies level were determined.Due to the partly fold synonymy lists in modern revisions.This is rather small faunas, only presence/absence data not restricted to ‘di⁄cult’ groups, such as the nas- are used for the analysis, yielding percentages sariids or conids, but can also be documented for of taxa occurring in two or more investigated ‘simple’ taxa such as the melongenids or strom- faunas. bids. E.g., the characteristic Oligocene Melongena Finally, the Iranian and Eastern Mediterranean semseyiana (Erdo«s) ^ a seemingly endemic Para- faunas were compared with those of the Western tethyan species (Baldi, 1973) ^ was described at Mediterranean, the Atlantic, the Paratethys and least under three synonyms (see Mikuz, 1999). especially with those from Pakistan.Our knowl- Now the ‘endemic’ Hungarian species turned up edge of these reference faunas is rather inhomoge- in Austrian, Croatian, Hungarian, Slovenian, neous.The small Eastern Mediterranean and Ira- Greek and maybe also in Turkish faunas. nian faunas consisting of approximately 50^60 Similarly, the endemism-index should not be species each are in contrast to the extremely rich overemphasised, since endemism may partly de- French and Italian faunas.For example, Coss- pend on the quality of research.Recently, Lo- mann and Peyrot (1909^1932) describe more zouet (1998, 1999) and Valde¤s and Lozouet than 600 Burdigalian and more than 500 Aquita- (2000) described 170 new species from the Oligo- nian gastropods.Bellardi (1872^1904) report even cene and Lower Miocene of the Aquitaine Basin, more than 980 Early Miocene species.The Oligo- indicating high endemism from the armchair. cene gastropod faunas of northern Italy consist of However, this high number of ‘endemic’ species at least 300 species, estimated from the various seems mainly related to the lack of equivalent, monographs of Fuchs (1870), Oppenheim (1900) elaborate studies in the Mediterranean area.The and Bellardi (1872^1904).From the Late Oligo- insu⁄cient study of Mediterranean faunas was cene of the Western Paratethys approximately 250 also stressed by Studencka et al.(1998) for Mid- species are recorded by Wol¡ (1896^1897), Baldi dle Miocene bivalve faunas.Thus, the lack of (1973) and Ho«lzl (1962), and more than 300 spe- modern revisions caused a higher percentage of cies are described from the Early Miocene Para- bivalves common to the Atlantic and the Parate- tethys by Steininger and Senes (1971), Ho«lzl in thys (60%) than of species common to the directly

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Fig.2.Late Eocene to Late Miocene geochronology and biostratigraphy after Ro «gl (1996, 1998), Ro«gl et al.(1993), Steininger (1999) (mammal zones), Cahuzac and Poignant (1997) (larger foraminifera), and Hardenbol et al.(1998) (magnetostratigraphy).

connected Mediterranean and the Paratethys workers such as Ho«rnes and Auinger (1879^1891) (47%). to split ‘good’ species into morphs, is hard to Furthermore, the taxonomic overcompleteness handle in statistics.These so-called subspecies or of some faunas, caused by the tendency of older variations tend to ‘evolve’ into species in subse-

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 108 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 quent studies, resulting in several local ‘species’ ince is the most extensive, including the entire which mask palaeobiogeographic relations.Fi- Mediterranean Sea but also parts of the Atlantic nally, open nomenclature is hard to adapt to sta- coast.Its westernmost extension are the Azores; tistics.This can be demonstrated very impres- in the north it includes the Bay of Biscay, and it sively by the otherwise highly valuable lists of reaches as far south as Madeira and the Canaries. Early Oligocene Paratethys faunas from Hungary It is bordered in the south by the Cape Verde by Baldi (1986), which comprise about 37% of Islands, which are part of the Eastern Atlantic taxa in open nomenclature.Within the eastern Province (=Senegalese Province sensu Wood- African gastropod faunas this percentage may ward) and the English Channel in the north. even reach 61% (see Azzaroli, 1958). The term Lusitanian Region as used by Ekman Hence, in this paper the authors try to focus on (1953) has a quite di¡erent meaning.He con¢nes the qualitative and semiquantitative composition the term to the Atlantic part north of the Straits of typical nearshore assemblages, which are sug- of Gibraltar.Consequently, he separates the gested to characterise the biogeographic units in southern part as Mauretanian Region.The Medi- various time slices.For biostratigraphic and geo- terranean Sea^Black Sea Province introduced by chronologic correlation see Fig.2. Longhurst (1998) corresponds partly to the Lusi- tania Province of Briggs (1995).A clear di¡erence is only that Longhurst treats the Bay of Biscay as 4. Circum-Mediterranean biogeography part of the Northeast Atlantic Shelves Province. The concept of Briggs (1995) concerns the shal- Woodward (1856) was one of the ¢rst who tried low-water fauna whereas the provinces introduced to divide the marine mollusc faunas in bioprovin- by Longhurst (1998) are based on pelagic organ- ces with characteristic compositions.He proposed isms. the rule that 50% of the species within a province In contrast to Briggs (1995), who maintains should be peculiar (endemic) to it.Physical fac- that the Recent Caspian and the Aral Sea provin- tors such as geographic barriers and climate were ces have derived their biotas from the Mediterra- recognised by Woodward as dividing lines be- nean, these provinces are considered herein as bi- tween distinct provinces.He additionally takes ogeographic and geologic descendants of the historical factors into account such as ‘di¡erent Paratethys Sea.This is strongly supported by distribution of land and water over the surface the data given in Ekman (1953), who describes of the globe’.Woodward’s mollusc-provinces are the Caspian fauna to be ‘practically a pure Sar- still accepted by some authors working on mol- matic fauna’.Ekman (1953) even emphasised the luscs such as Sabelli (1980).Nonetheless, the bio- small percentage of species common to the Med- province concept was improved during the 20th iterranean Sea and regards the Caspian fauna to century by authors such as Ekman (1953) or be of Paratethyan ( = Sarmatic sensu Ekman) ori- Hedgpeth (1957), who attempted to integrate the gin. littoral fauna as a whole. Similarly, the Black Sea Basin was part of the In the following the authors provide a brief re- Paratethys, but became part of the Mediterra- view of selected recent and historical bioprovinces nean^Atlantic Region during the Pleistocene and and their geographic pattern within the circum- the Holocene by its connection to the Aegean Sea Mediterranean area. via the Marmara Sea.

4.1. Modern bioprovinces 4.1.2. Eastern Atlantic^Boreal Region The Region extends from the British Channel 4.1.1. Mediterranean^Atlantic Region north via the North Sea to the Baltic Sea and According to Briggs (1995) this region com- reaches from the Celtic Shelf as far north as the prises the Lusitania, Black Sea, Caspian, and Hebrides.It is limited to the south and the south- Aral provinces.Among them the Lusitania Prov- west by the Lusitania Province and to the north

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 109 and the north-east by the Arctic Region (Briggs, Ocean Province, while in the text the Indo-Poly- 1995).In contrast to Woodward’s Celtic Province, nesian Province reaches as far west as the entrance the Eastern Atlantic^Boreal Region includes also of the Persian Gulf.Other limits of the Western Iceland, the Norwegian coast and extends as far Indian Ocean Province are given by Macnae north as Murmansk. (1968), who includes the entire coasts of the Several terms are used in the literature as rough northern Arabian Sea as far as the Gulf of synonyms for this Region.Especially European Khambhat in north-western India.Additionally, palaeontologists usually called the geographic an- the Western Indian Ocean Province as de¢ned by cestor of this Region the ‘Boreal Province’ (Kaut- Por (1989) would reach to southern India, to the sky, 1925; Baldi, 1973), the ‘Scandic Region’ coast between Mangalore and Calicut. (Ho«lzl, 1962), or simply the ‘northern province’ (Senes, 1958), while Ro«gl and Steininger (1983, 4.2. Historical biogeographic units 1984) introduced the modern term ‘Atlantic^Bo- real Bioprovince’. 4.2.1. Tethys Realm The term ‘Tethys’ was introduced by the Aus- 4.1.3. Indo-West Paci¢c Region trian geologist E.Suess (1893) as a palaeogeo- The term ‘Indo-West Paci¢c’ was introduced by graphic entity.Herein ‘Tethys Realm’ is used in Ekman (1934), who united the Indo-Paci¢c, the a strict biogeographic sense and must not be in- Western Paci¢c, and parts of the Central Paci¢c termingled with the geographic term ‘Tethys’ as in his biogeographic unit.Later, Briggs (1995) described by Hsu« and Bernoulli (1978) and Sen- demonstrated that the Region has to be extended go«r (1998).This is of great importance, since the to the Easter Islands, thus introducing an area geographic Tethys (sensu Hsu« and Bernoulli, that nearly extends to the coasts of America. 1978) or the tectonically de¢ned Tethys (sensu This vast Region is divided into the Indo-Polyne- Sengo«r, 1998) vanished during the Eocene when sian, the Western Indian Ocean, the Red Sea, the India collided with Laurasia. North-western Australian, the Hawaiian, the Eas- Originally, the biogeographic Tethys Realm,as ter Island, and the Marquesas provinces.The mod- de¢ned by Kau¡mann (1973), was applied to Me- ern Western Indian Ocean Province extends from sozoic faunas.Kau¡man’s Tethys Realm is based the Persian Gulf along the eastern African shelf ^ on Cretaceous bivalves and represents a vast, cir- including the Seychelles, Madagascar and the cum-equatorial area.For the Cenozoic era the Mascarene Islands ^ as far south as Kwa Zulu biogeographic term Tethys is still used by Popov north of Natal in South Africa (Kilburn and Rip- (1993) when de¢ning zoogeographic units for Late pey, 1982).The Red Sea is excluded from this Eocene bivalves.This Tethys Region sensu Popov Province, but forms a distinct Province due to is modi¢ed herein with respect to hierarchy and the high percentage of endemics (Briggs, 1995). enlarged as Tethys Realm (Fig.3(left, top)).Ac- In an easterly direction the Western Indian Ocean cording to Popov, the Late Eocene Tethys of Province is bordered by the larger Indo-Polynesian western Eurasia can be divided into an Indo-Afri- Province.This Province extends from the entrance can Region and a Mediterranean Region (subre- of the Persian Gulf, via the Arabian Sea, along gions sensu Popov, 1993).At that time the south- the Indian shelf to Sri Lanka and the Bay of ern Tethyan faunas of Egypt, Somalia and India Bengal and includes Malaysia, Indonesia and displayed highest resemblance, but di¡ered clearly Eastern Australia.To the north it reaches as far from the northern Tethyan ones from Italy and as the Japanese Amami Islands. southeastern France. The border between these two provinces is not Simultaneously, with the narrowing of the Te- unequivocal.Even the de¢nition given in Briggs thys in the north of the Indian shield, the south- (1995) markedly di¡ers from his drawing since the ern seaway and the ancestral Indian Ocean sketch includes also the Iranian and Pakistanian widened.This seaway warranted a continuous coasts of the Arabian Sea in the Western Indian connection of the western and eastern tropical

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Tethyan faunas, hence allowing to extend the bio- and the African/Arabian plates enabled a fair ex- geographic term Tethys Realm also for Oligocene change of shallow-water faunas between these and Early Miocene times.With respect to the di- areas.In the eastern direction, the Indonesian sea- vergent meaning of Tethys Ocean and Tethys way acted as a marine connection between the Realm, it might be more correct to name the bio- ‘Indo-Paci¢c’ and the western Paci¢c areas which geographic unit the ‘Proto-Mediterranean^Indo- lasted until the early Middle Miocene (Ogasawara African^Western Paci¢c Realm’ ^ an obviously and Noda, 1996; Nishimura and Suparka, 1997). unwieldy term.Thus the term Tethys Realm is A tropical, equatorial current £owed through this favoured in this paper. Paci¢c^Indian Ocean gateway in a westerly direc- The post-Eocene Tethys Realm existed during tion (Tsuchi, 1997).Tethyan gastropods, such as the entire Oligocene and during the very Early Tibia, Telescopium, Terebralia, or Vicarya, how- Miocene without major interruption.The biogeo- ever, were spread as far north as Southern Japan graphic unit vanishes during the Burdigalian and during the Early and early Middle Miocene (Oga- with the formation of the Gomphotherium land- sawara and Noda, 1996; Noda and Watanabe, bridge between Eurasia and the Arabian plate in 1996). the Late Burdigalian (Ro«gl, 1998).The ¢nal Corresponding to the modern Indo-West Paci¢c breakdown of the Tethys Realm is also evident Region, a separation of this vast marine sphere from a geographical viewpoint.However, as men- into several biogeographic units is to be expected. tioned above, the Tethys Sea vanished much ear- Due to inadequate data it is currently impossible lier during the Eocene. to designate any of these boundaries exactly. During the Oligocene and the Early Miocene, Nevertheless the representative Oligocene and the Tethys Realm covered an extraordinarily large Miocene faunas of Italy (Bellardi, 1872^1904; area.Its main territory was an area, which coin- Fuchs, 1870), Greece, Central Iran, Pakistan (Vre- cides more or less with the modern Indo-West denburg, 1925^1928; Iqbal, 1969a,b, 1980), Bur- Paci¢c Region, but in western direction it also ma (Noetling, 1901), Java (Martin, 1916^1917, included the area of the modern Mediterranean 1921^1922), Celebes (Beets, 1942, 1943), and the Sea.A broad connection between the Iranian Sunda Islands (Vlerk, 1931) re£ect clear shifts in

Fig.3.Marine biogeography of the circum-Mediterranean area in four time slices.Di¡erent colours represent separate biogeographic units on the hierarchy of a Region or a Province.Arrows indicate hypothetical direction of surface currents.(Left, top) Late Eocene: The biogeographic pattern is based on bivalve genera as outlined by Popov (1993).At that time the Eurasian marine sphere displays a distinct S^N trend expressed by a northern entity united in the North European Region and a southern entity, the Tethys Realm.The Tethys Realm can be divided into two Regions, which were named the ‘Mediterranean’ Region and the ‘Indo-African’ Region by Popov (1993).(Left, bottom) Oligocene: The Western Tethys Region is divided into the Mediterra- neanÎranian Province and the Western IndianÊastern African Province.WÊ oriented surface currents along the northern shores of the Tethys facilitated the distribution of ‘European’ gastropods as far southeast as Pakistan.However, a distinct N^S gradient is indicated by the characteristic faunistic composition in the Western IndianÊastern African Province.In the Paratethys Sea, the Danubian Province and the Proto-Caspian Subprovince formed as a result of the reduced connections to the Tethys.The complex biogeographic development of these units is triggered by immigrations from the north and/or south as well as by local evolution. (Right, top) Early Miocene: The Western Tethys Region vanished as the Western IndianÊastern African Province ceased to exist. Hence, in this area the gastropod fauna of the Proto-Indo-West Paci¢c Region developed and the ‘European’ in£uence was lost. In contrast, the newly formed Proto-MediterraneanÂtlantic Region is a pure descendant of the Oligocene MediterraneanÎranian Province.Simultaneously, the Danubian Province, which now lacks the northern in£uence, contains southern species of the Proto- MediterraneanÂtlantic Region, and should probably be treated as part of this Region.(Right, bottom) Late Early Miocene: The separation of the Proto-MediterraneanÂtlantic Region from the Proto-Indo-West Paci¢c Region is completed by the formation of the so-called ‘Gomphotherium landbridge’, which connected Arabia with Eurasia in the Late Burdigalian (Ro«gl, 1998).The Proto- MediterraneanÂtlantic Region displays a rather homogeneous gastropod fauna, which nearly takes over the Danubian Province. In the west, the Proto-MediterraneanÂtlantic gastropod fauna spread along the Atlantic coast.

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 113 compositions, which prove the existence of dis- Indo-Malayian faunas, called herein informally tinctive regions and provinces.Some of them the Proto-Indo-Polynesian Province of the Pro- will be proposed in the following. to-Indo-West Paci¢c Region.The fauna of this Proto-Indo-West Paci¢c Region displays hardly 4.2.1.1. Western Tethys Region any resemblance with the Oligocene gastropod Paleogene: During the Oligocene, the Western fauna of the Western IndianÊastern African Tethys Region ( = Indo-European sensu Ekman, Province.Neither the Pakistanian nor the eastern 1953) represents part of the Tethys Realm. It cov- African faunas have species in common with the ered the area of the modern Mediterranean but Oligocene gastropod faunas of the Proto-Indo- probably included also the European Atlantic Polynesian Province (e.g. Buton-fauna described coast up to the Bay of Biscay in the west and by Martin (1933, 1935) and Beets (1942, 1943)). reached to Pakistan, Somalia and Zanzibar in During the Oligocene a separation within the the east and the south (Fig.3(left, bottom)). Western Tethys Region in at least two provinces Most characteristic for the Western Tethys Region is obvious. are the faunas of Northern Italy.The Mediterra- (1) MediterraneanÎranian Province.The herein neanÎranian Province (see below) is designated proposed MediterraneanÎranian Province became herein as the type unit for the Western Tethys established in the area of the modern Mediterra- Region ( = chorotype sensu Westermann, 2000a). nean Sea and had its south-easternmost known Its eastern border seems to have been located extension in Central Iran (Fig.3(left, bottom)). somewhere along the coast of Southern India, The new term is introduced instead of ‘Mediter- since the south-eastern Asian faunas display no ranean’ Province, because of the di¡erent geo- similarities either with those of Pakistan (Vlerk, graphic extension and moreover because of the 1931) or with those of Europe.Such faunistic de- synonymous use of the term for Mesozoic and marcation lines have also been documented for Cenozoic biogeographic units with di¡erent biotic the Eocene by Cotter (1923) for the bivalves.He characteristics.Characteristic gastropod faunas of stated that Burma, Sumatra, Java, and Borneo this province are those of Northern Italy (Fuchs, constituted a distinct bioprovince, di¡erent from 1870; Bellardi, 1872^1890; Sacco, 1890^1904), the the Sind^Baluchistan and Egypt faunas.Some fauna of the Mesohellenic Trough in Greece years later Cox (1930b, 1936) concluded that the (Brunn, 1956; Harzhauser, 2000), and the fauna a⁄nities of the Bahrain and southern Iranian fau- of the Esfahan^Sirjan Basin in Central Iran nas with those of north-western India, Somalia (Harzhauser, 2000; Fig.4a,c).It is best character- and Egypt hint at an Indo-African biogeographic ised by its rich strombid fauna which seems to be entity ( = Indo-African fauna sensu Cox, 1930b). a heritage of the highly diverse, blooming Eocene Correspondingly, Martin (1931) proposed an strombid fauna.None of the giant species of Oos- Indo-Malayian bioprovince based on the homoge- trombus or Dilatilabrum is known from any Oli- neity of Late Eocene faunas from eastern India to gocene section southeast of Iran.Characteristic Java and Burma, and on their marked di¡erence gastropods of the MediterraneanÎranian Prov- from that of western India^Pakistan.This biogeo- ince are, among many others, Tectus lucasianus graphic pattern ¢ts well the simulations of Barron (Brongniart), Angaria scobina (Brongniart), Turri- and Peterson (1991) who demonstrated that Bur- tella asperula Brongniart, Turritella conofasciata ma and Java were separated from north-western (Sacco), Peyrotia strangulata (Grateloup), Campa- India during the Eocene by a strong western nile charpentieri (Basterot), Granulolabium plica- boundary current, while the Indian Ocean circu- tum (Bruguie're), Tympanotonos stroppus (Brong- lation pattern and the Somali current are evident niart), Clava ampullosa (Brongniart), Clava vog- only from ‘middle’ Oligocene onwards (Barron linoi (Michelotti), Cerithium calculosum Defrance, and Peterson, 1991).Hence, in the east the Paleo- Gourmya romeo (Bayan), Diastoma costellatum gene Western Tethys Region is bordered by a ma- elongatum (Brongniart), Proadusta parvitala (Sac- jor adjacent biogeographic unit inhabited by these co), Ampullinopsis crassatina (Lamarck), Globu-

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 114 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133

Fig.4.(a,b) Species level similarities within gastropod faunas.Numbers in circles represent total numbers of species of basin; numbers in squares represent numbers of co-occurring species; the asterisk indicates reference faunas in Northern Italy.(c,d) Per- centages of N-Mediterranean species represented in the Greek, Turkish, Iranian, and Pakistanian gastropod faunas.The sketches (a) and (c) demonstrate the high faunistic similarities in the Western Tethys Region during the Oligocene and document the homogeneity within the MediterraneanÎranian Province.During the Early Miocene (b,d) this homogeneity is strongly indicated within the Proto-MediterraneanÂtlantic Region.In contrast, the Pakistanian gastropod faunas lost the similarities and have only 4^7% of species, which are typical of the Proto-Mediterranean Region, thus pointing to the beginning development of the Proto- Indo-West Paci¢c Region.The very signi¢cant drop in similarity between the Iranian and the Pakistanian faunas during the Early Miocene clearly witness the formation of a major biogeographic boundary between V30‡ and 35‡ N. laria gibberosa (Grateloup), Sinum aquensis (Re- 3(left, bottom)).The term ‘Indo-African’ should cluz), Bayania semidecussata (Lamarck), Oostrom- be avoided, because of its quite di¡erent use in the bus auriculatus (Grateloup), Oostrombus irregu- literature, e.g. Popov (1993) versus Cox (1930b). laris (Fuchs), Strombus radix (Brongniart), It is represented by the fauna of the Pakistanian Strombus (Dilatilabrum) roegli Harzhauser, Xeno- Nari Formation, which has at least 40 species in phora cumulans (Brongniart), Cassis mamillaris common with those of Northern Italy (Vlerk, Grateloup, Cassis vialensis Fuchs, Turbinella epi- 1931) (Fig.4).However, the high number of spe- soma (Michelotti), Volutilithes subambigua d’Or- cies unknown from European localities (about bigny, Lyria anceps (Michelotti), Conus (Leptoco- 50^60 species) re£ects a rather independent devel- nus) diversiformis Deshayes, and Terebra subtes- opment in this south-eastern termination of the sellata d’Orbigny. Western Tethys Region.This Western Indian^ (2) Western IndianÊastern African Province. Eastern African Province is documented by the Towards the southeast the MediterraneanÎranian Pakistanian faunas (Vredenburg, 1925^1928; Iq- Province is bordered by a very distinct biogeo- bal, 1969a,b, 1980), but can also be traced along graphic unit, which is introduced herein as the the coast of Eastern Africa as con¢rmed by the Western IndianÊastern African Province (Fig. faunas of Somalia described by Azzaroli (1958)

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 115 and Kenya (Davies, 1923).The Western Indian^ the boundary between these biogeographic areas Eastern African Province includes also the area of along the northern Tethys shores is located some- Oman (Fig.3(left, bottom)) as documented by a where between Central Iran and the Sind^Balu- small reefal gastropod fauna kindly donated by chistan Region.Along the southern shores, the M.Bernecker (Erlangen).Characteristic gastro- faunas of Libya, Palestine and Egypt are consid- pods of the Western IndianÊastern African Prov- ered part of the MediterraneanÎranian Province ince are ‘Turbo (Senectus)’ naricus Vredenburg, (Fig.3(left, bottom)), while the faunas of Oman ‘Turbo (Olearia)’ protocepoides Vredenburg, Pro- represent the northern known extension of the toma retrodilatatum Vredenburg, Turritella narica Western IndianÊastern African Province (Fig. Vredenburg, Semicassis oligocalantica (Vreden- 3(left, bottom)).Similarities between the two burg), Harpa (Eocithara) narica Vredenburg, Bel- provinces are high within the potamidids, turri- latara narica (Vredenburg), or Cerithium sindiense tellids and naticids but low amongst the volutids, Vredenburg. turrids and other neogastropods, documenting the Widespread species which are recorded from peculiar character of the fauna.Only very few of both provinces and are therefore diagnostic (typi- these species, such as Ampullinopsis crassatina, cal) of the entire Western Tethys Region are: Globularia gibberosa or Granulolabium plicatum Tectus lucasianus (Brongniart), Campanile char- managed to colonise adjacent bioprovinces in pentieri (Basterot), Granulolabium plicatum (Bru- high numbers.Others such as Turbinella episoma, guie're), Tympanotonos stroppus (Brongniart), Ce- Cassis mamillaris or Angaria scobina, which are rithium ighinai (Michelotti), Turritella magnaspe- very common in Western Tethyan assemblages, rula Sacco, Turritella asperula Brongniart, Turri- are rare faunal elements within their northern- tella conofasciata (Sacco), Peyrotia desmarestina most distribution area in the Early Oligocene of Basterot, Peyrotia strangulata (Grateloup), Strom- the Paratethys.The connection between these bi- bus radix (Brongniart), Cassis mamillaris Grate- ogeographic entities was supported by a distinct loup, Cassis retusa Michelotti, Cassis nummuliti- easterly current along the northern shores of the phila (Sacco), Sconsia beyrichi (Michelotti), Tethys (Fig.3(left, bottom); Barron and Peterson, Proadusta parvitala (Sacco), Globularia gibberosa 1991).According to Barron and Peterson (1991) a (Grateloup), Ampullinopsis crassatina (Lamarck), westward-directed Eocene to Oligocene Tethyan Amaurellina oweni (Archiac and Haime), Sinum current system along the northern shores as sug- aquensis (Recluz), Xenophora cumulans (Brong- gested by Berggren and Hollister (1974) has to be niart), Ficus condita (Brongniart), Turbinella epi- rejected. soma (Michelotti), Conus diversiformis Deshayes, A third province in the western part might be Conus carcarensis Sacco, Lyria anceps (Michelot- based on di¡erences in the composition of the ti), and Terebra subtessellata d’Orbigny.Along Oligocene faunas from southwestern France with these gastropods the remarkable bivalve Avi- (Grateloup, 1840; d’Orbigny, 1852; Lozouet, cularium carinatum (Bronn) may be regarded as a 1998, 1999) and the Italian and Greek localities. characteristic element of the Western Tethys Re- This is additionally supported by the northern gion. in£uence in the French faunas as suggested by The southeasternmost known gastropod fauna Welle (1998) and by numerous muricids, bucci- of the MediterraneanÎranian Province occurs in nids, nassariidis, or conids described only from the Late Rupelian and Early Chattian of the Es- France (Lozouet, 1999). fahan^Sirjan Basin in Central Iran.The Oligocene Early Neogene: During the Early Miocene the section of Lordegan in the Iranian Zagros Moun- Western Tethys Region shrank drastically from tains, located still further south, bears a strombid the gastropods’ point of view.Still, the area of the MediterraneanÎranian Province.In con- from the Bay of Biscay to the Mediterranean ba- trast, the northwesternmost fauna of the Western sins as far east as Central Iran display a rather IndianÊastern African Province is actually repre- similar fauna (Fig.4).However, the Early Mio- sented by the Pakistanian assemblages.Therefore cene brings about a drastic change for the West-

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 116 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 ern IndianÊastern African Province, which van- faunas was strongly prevented during the Early ishes as a distinct unit probably even before the Miocene.Therefore in the area of the Recent closure of the Tethyan corridor in the Late Bur- Mediterranean Sea a biogeographic unit formed digalian.The data on Pakistanian molluscs from which might best be named Proto-Mediterra- the Early Miocene Gaj Formation given by Vre- neanÂtlantic Region (Fig.3(right, top)).Accord- denburg (1925^1928) re£ect a strongly increasing ing to the palaeogeographic situation as drawn by in£uence by the fauna of South-East Asia, and a Ro«gl (1998) this area roughly corresponds to the rapid decrease of elements of the Western Tethys limits of the modern MediterraneanÂtlantic Re- Region.Correspondingly, Azzaroli (1958) noticed gion except for the Black Sea Province, which was in the Miocene faunas of Somalia some in£uence not in existence.This area was part of the Para- from Indonesia, although his fauna still has some tethys at that time. species in common with Italian localities.Hence, The southwestern extension of the Proto-Medi- the separation of the gastropod faunas of the east- terraneanÂtlantic Region is not well de¢ned due ern African and the western Indian coastal areas to the lack of available data on shallow marine from those of the Western Tethys Region already faunas along the shores of northwestern Africa. took place during the Early Miocene (Burdigali- However, the limits along the European Atlantic an?).Since the faunas of this area have a clear coast in northern direction were more or less ‘Indo-Paci¢c’ character and bear high percentages equivalent to those of the Recent Mediterra- of Indonesian species, they may be treated as neanÂtlantic Region (sensu Briggs, 1995).The western part of an early stage of the Indo-West Artois axis formed a landbridge, which repeatedly Paci¢c Region, which should be termed the Proto- separated the fauna from that of the North Sea Western Indian Ocean Province. Basin (Pomerol, 1982; Ziegler, 1990). With the disconnection of the Indo-African Unequivocal evidence for direct marine connec- fauna and the take-over by elements of the Pro- tions to adjacent provinces exists for the Central to-Indo-West Paci¢c Region in the area of the Paratethys, which is strongly in£uenced by the former Western IndianÊastern African Province fauna of the Proto-MediterraneanÂtlantic Region during the Early Miocene the MediterraneanÎra- during that time (Ro«gl, 1998, 1999; Harzhauser, nian Province lost its southeastern counterpart. 2002). Finally, the point-of-no-return was reached by Within this area the data on gastropod faunas the closure of the Eastern Mediterranean seaway from France (Cossmann and Peyrot, 1917^1922), between the Anatolian and the Arabian/African Italy (Bellardi, 1872^1904) and Iran (Harzhauser, plates during the Late Burdigalian (Jones, 1999). 2000) show good correspondence during the Bur- Based on the dramatic geodynamic and biogeo- digalian, although a slight northwest-southeast graphic changes and especially due to the evolu- shift in the composition is evident.However, the tion of the advanced Miocene Mediterranean gas- eastern Mediterranean faunas have 80% of species tropod fauna, the term Western Tethys Region in common with Northern Italy and France, thus seems hardly acceptable for the Early Miocene indicating a rather homogeneous distribution of at least for the Burdigalian (see also Wester- taxa within this area during the Burdigalian mann’s rules for replacement of names, Wester- (Fig.4d).A slight drop in resemblance is recorded mann, 2000a).Therefore, we propose the term in the Iranian fauna, which still contains 66% (38 Proto-MediterraneanÂtlantic Region (Fig.3 species) of ‘Mediterranean’ species (Fig.4d).To (right, top and bottom)) for the descendant of obtain a better understanding of the similarities the strongly reduced Paleogene Western Tethys and di¡erences between the Iranian and the Med- Region. iterranean faunas it is necessary to focus on the family level, because the degree of resemblance 4.2.1.2. Proto-MediterraneanÂtlantic Region strongly £uctuates in the various families.In gen- As discussed above a marine faunal exchange eral the melongenids, cerithiids, strombids, and to between the southeastern and the ‘Mediterranean’ some extent also the naticids show highest a⁄n-

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 117 ities to the Mediterranean faunas.In contrast, the gastropods during the early Middle Miocene is conids and the turrids display a considerable somewhat dubious.Instead of the usually advo- number of taxa which are known only from the cated immigrations, the autochthonous evolution Qom Basin (10^15%); due to the poor informa- in a phase of climatic optimum seems to be un- tion it is unclear whether these species are also derrated.Anyway, the short-lived connections of distributed in the Indo-Paci¢c or if they represent the Proto-MediterraneanÂtlantic Region to the endemic forms, thus hinting at a distinct Iranian Indo-West Paci¢c Region did not change the equi- subprovince.Some of the most characteristic taxa librium gastropod faunas of the Proto-Mediterra- within the Proto-MediterraneanÂtlantic Region neanÂtlantic Region markedly.During the Mio- are: Turritella (Turritella) terebralis Lamarck, cene and Pliocene various reconnections are Protoma quadriplicata (Basterot), Peyrotia des- discussed (Grecchi, 1978; Lozouet, 1992; Ro«gl, marestina (Basterot), Protoma cathedralis (Brong- 1998; Jones, 1999), however, the biogeographic niart), Terebralia bidentata (Defrance), Cerithium boundaries between the Indo-Paci¢c and ‘Medi- vulgatum Bruguie're, Granulolabium bicinctum terranean’ remained stable. (Brocchi), Granulolabium plicatum (Bruguie're), Ti- A second impact on Proto-MediterraneanÂt- bia dentata (Grateloup), Strombus (Lentigo) bo- lantic mollusc faunas was the Messinian salinity nelli (Brongniart), Xenophora deshayesi (Michelot- crisis in the Late Miocene, when the marine fauna ti), Sconsia striata miocenica Sacco, Melongena of the Mediterranean basins was strongly impov- lainei (Basterot), Tudicla rusticula (Basterot), Ath- erished (Hsu« et al., 1978). The gastropod fauna of leta ¢culina (Lamarck), and Conus (Conolithus) the Mediterranean basins probably vanished com- dujardini Deshayes. pletely ^ except for some supposed shelters such Aside from this core-area, the Danubian Prov- as the ‘Andalusian sanctuary’ (see Demarcq, ince (see below) ^ spanning the western and east- 1987) ^ due to desiccation throughout the Medi- ern part of the Paratethys Sea ^ is considered to terranean.The fauna of the Eastern Atlantic represent a distinct province of the Proto-Medi- coasts was less a¡ected.After the salinity crisis terraneanÂtlantic Region at least during the late the deep-water fauna was distinctly impoverished Early Miocene and the early Middle Miocene (see (Ra⁄ and Taviani, 1984), but it had not such a below). catastrophic e¡ect on shallow marine molluscs This palaeogeographic framework of the Proto- (Sabelli and Taviani, 1984), which recolonised MediterraneanÂtlantic Region based on gastro- the Mediterranean basins from the Atlantic. pod data corresponds largely to the ‘Mediterra- Nevertheless, the Messinian salinity crises caused nean Province’ as introduced by Jones (1999) a turnover in composition of the nearshore fauna. based on larger benthic foraminifera.His prov- Post-Messinian gastropod faunas lack character- ince unites the entire Mediterranean area, the istic Miocene taxa, such as Terebralia bidentata Qom Basin, and the Atlantic coasts of the Aqui- (Defrance), Melongena cornuta (Agassiz), Polini- taine Basin and of West Africa.Similarly, Ro«gl ces redempta (Michelotti) or Perrona jouanetti and Steininger (1983, 1984) recognised this bio- (Desmoulins).In fact, many of the conids, turrids geographic entity and called it the ‘Mediterranean and turritellids of the Proto-MediterraneanÂtlan- (or Tethyan) bioprovince’. tic Region became extinct (cf.Dermitzakis and The mollusc fauna of the Proto-Mediterranean^ Georgiades-Dikeoulia, 1987).Hence, the post- Atlantic Region su¡ered two major incisions.The Messinian gastropod fauna represents rather the ¢rst one took place in the early Middle Miocene ancestral, warm-water stock of the modern Med- when a short-lived connection to the Indo-Paci¢c iterraneanÂtlantic Region. ^ coinciding with a marked global warming (Mil- During the Late Pliocene and Pleistocene the ler et al., 1991; Ro«gl, 1998; Nishimura and Supar- change from the Proto-MediterraneanÂtlantic ka, 1997; Tsuchi, 1997) ^ gave rise to a highly Region to the modern MediterraneanÂtlantic Re- diverse Middle Miocene mollusc fauna.Until gion was accelerated by distinct cooling events now the role of immigration of Indo-West Paci¢c (Ro«gl and Steininger, 1983).Consequently, trop-

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 118 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 ical/subtropical taxa which contribute to the fau- lowed by the Late Oligocene and Early Miocene na of the Proto-Mediterranean^Atlantic Region Eo-Paratethys, the late Early Miocene to early are missing in the modern region (e.g. Olividae Middle Miocene Meso-Paratethys and the Middle as demonstrated by Davoli, 1989, or Strombidae to Late Miocene Neo-Paratethys (see also Stei- as emphasised by Abbott, 1960). ninger and Wessely, 2000). To avoid an intermingling of geographic and 4.3. Biogeographic position ofthe Paratethys Sea biogeographic terms, we reject the terms Western, Central and Eastern Paratethys for biogeographic The Paratethys, as a distinct palaeogeographic use in this paper.Instead we propose the new and palaeobiogeographic unit, formed around the terms Danubian Province and Proto-Caspian Sub- Eocene/Oligocene boundary (Fig.2) and lasted province (Fig.3(left, top; right, top and bottom)). until the Pliocene.In fact, even the modern Aral Hence, the Danubian Province is characterised by and Caspian provinces are a heritage of the huge the total fauna of the Western, Central, and East- Paratethys, based on its peculiar fauna and en- ern Paratethys.The peculiar faunistic inventory of demism.During its maximum extent the Parate- the Eastern Paratethys allows a further seggrega- thys Sea spread from the Rhone Basin in France tion of a Proto-Caspian Subprovince.(A further towards Inner Asia.Its geographic subdivisions di¡erentiation of these biogeographic units and are the Western Paratethys, comprising the Rhone a detailed description of their evolution and ex- Basin and the Molasse Basin of Switzerland and pansion will be published elsewhere.) western Bavaria, the Central Paratethys, reaching from Bavaria in the west to the Carpathian 4.3.1. Danubian Province and Proto-Caspian Mountains in the east, and the Eastern Parate- Subprovince thys.Recently, the stratigraphy and geodynamics The ¢rst endemic Paratethys fauna developed of the latter area were studied by Popov et al. during the Early Oligocene Solenovian, when the (1993) and Jones and Simmons (1996).A critical ¢rst nearly complete closure of the Paratethys oc- up-to-date overview on palaeogeographic data curred.This event peaked in the blooming and has been given and discussed by Ro«gl (1998, rapid evolution of a highly endemic brackish 1999) and summarised in several palaeogeograph- water bivalve fauna with genera such as Janschi- ic sketches. nella, Korobkoviella or Ergenica (Voronina and This region underwent an outstanding history Popov, 1984; Popov et al., 1985; Nevesskaja et of total or partial isolation, re£ected in a high al., 1987; Ro«gl, 1998).By this event the Parate- percentage of endemics, alternating with various thys was united in a single, very distinct biogeo- connections to the adjacent northern and/or graphic entity ^ the Proto-Caspian Subprovince ^ southern seas. as far as bivalves are concerned (Popov et al., The term Paratethys was introduced as early as 1993).Nonetheless, as shown by Popov et al. 1924 by Laskarev (1924).During the last decades (1985) and Popov and Titova (1982), this endem- and especially due to the edition of the ‘Chrono- ism is much lower within the gastropods.Brackish stratigraphie und Neostratotypen’ volumes the or estuarine gastropods which are at that time palaeobiotic framework was distinctly improved widespread in the entire Western Tethys Region (Cicha et al., 1967; Steininger and Senes, 1971; inhabited the coasts of the Paratethys; Melanopsis Baldi and Senes, 1975; Papp et al., 1973, 1974, impressa Krauss, Granulolabium plicatum (Bru- 1978, 1985; Stevanovic et al., 1990). Its geody- guie're), Theodoxus crenulatus (Klein), and Tympa- namic history was described by Senes and Ma- notonos margaritaceus (Brocchi) give evidence of rinescu (1974) and Rusu (1988), who divided the reduced marine conditions within this interval, geodynamic development of the Paratethys into but do not justify a separation of a biogeographic four stages: the Proto-Paratethys formed in the unit at the level of a region. Late Eocene to Early Oligocene, caused by ¢rst Aside from this short-lived extreme western ex- isolation from the open oceans.This stage is fol- tension of the Proto-Caspian Subprovince, the

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 119 western part of the Paratethys was characterised and northern turrids such as Orthosurcula regula- by the quite di¡erent fauna of the Danubian Prov- ris contribute to the fauna of the Ustjurt area ince (Fig.3(left, bottom)).In times of open sea- (Amitrov, 1973).Endemism, however, is much ways, the Early Oligocene Danubian Province con- stronger in the Proto-Caspian Subprovince sists of northern species accompanied by a throughout the Oligocene. surprisingly large number of Western Tethyan el- The second phase of a rather uniform Parate- ements, giving evidence of the earliest stage of thys fauna, coinciding with the westward exten- separation of these biogeographic entities. sion of the Proto-Caspian Subprovince, developed Amongst these the most characteristic Western during the Early Miocene Kotsakhurian (corre- Tethyan species are Turritella conofasciata Sacco, sponding to the Ottnangian).Again, the separa- Turritella asperula Brongniart, Ficus oligo¢coides tion of the Eastern Paratethyan Sea coinciding Sacco, Athleta italica Fuchs, Turbinella episoma with brackish water conditions is followed by a (Michelotti), Cassis mamillaris Grateloup, Anga- sudden evolutionary peak in bivalves, resulting in ria scobina (Brongniart), and Babylonia caronis a large number of endemic genera such as Limno- Brongniart (Baldi, 1986).Thus the later Early Oli- pagetia, Rzehakia, Lenticorbula,orEoprosodacna gocene, as documented by Baldi (1986) for the (Kvaliashvili, 1962; Voronina and Popov, 1985). gastropod faunas of the Hungarian Kiscell Clay, This peculiar, so-called ‘Rzehakia fauna’ reaches displays about 31^35% endemic gastropods but even the Western and Central Paratethys Sea in also more than 20% Western Tethyan taxa and the Late Ottnangian.Corresponding to the ¢rst approximately 16% northern species. isolation stage in the Early Oligocene, the Early During the Late Oligocene Egerian the south- Miocene Proto-Caspian Subprovince is more dis- ern £air decreased.The Danubian Province hosts a tinctly characterised by the bivalves than by the mixture of northern elements, which made their gastropod fauna (see also Ctyroky¤, 1972). way from the North Sea Basin during the Rupe- A complete disconnection of the Western/Cen- lian, such as Cassidaria megacephala (Philippi), tral Paratethys and Eastern Paratethys began in Drepanocheilus (Arrhoges) speciosus (Schlotheim), the latest Early Miocene (Fig.3(right, bottom)) Ficus concinnus (Beyrich), ‘cosmopolitan’ Tethyan and early Middle Miocene (Nevesskaja et al., species such as Granulolabium plicatum (Bru- 1987; Ro«gl, 1998).Good marine connections of guie're), Tympanotonos margaritaceus (Brocchi) the Western/Central Paratethys with the southern and Melanopsis impresssa Krauss, and a fair num- sea allowed immigration of ‘MediterraneanÂt- ber of endemics, such as Melongena incornuta lantic’ gastropods.At times of high immigration (Ho«lzl), Bullia hungarica (Ga¤bor), Chicoreus tri- rates such as the late Early Miocene Karpatian or gonalis (Ga¤bor), Pisanella doboi (Noszky), and the early Middle Miocene Badenian the ‘Mediter- Egerea collectiva (Ga¤bor).These considerable ranean’ character became overwhelming.The number of endemics grow distinctly smaller dur- Karpatian gastropod fauna of Austria (Harzhau- ing the Early Miocene Eggenburgian, when fau- ser, 2002) has only 25^30% Western Paratethys nistic ingressions from the west and south led to a endemics ^ a percentage which is probably even turnover in composition. too high, since several of the ‘endemic’ species are The heavy in£ux of species from the northern probably hidden in poorly described Mediterra- bioprovince ^ herein called the Proto-Eastern At- nean species groups.Anyway, the remaining 70^ lantic^Boreal Region (Fig.3(left, bottom; right, 75% of these latest Early Miocene gastropods are top and bottom)) ^ characterises also the Oligo- typical inhabitants of the Proto-Mediterranean^ cene gastropod fauna of the Proto-Caspian Sub- Atlantic Region.In respect to biogeography this province.Northern elements such as Drepanochei- take-over by ‘MediterraneanÂtlantic’ gastropods lus (Arrhoges) speciosa, Cassidaria buchi/depressa, within the Danubian Province and the low endem- Ficus crassistria, Turritella planispira, Cancellaria ism requires an integration of the Danubian Prov- evulsa, and Typhis pungens reach the Aral and ince into the Proto-MediterraneanÂtlantic Region Caucasus basins (Oveckin, 1956; Alizade, 1968), which thus consists of the MediterraneanÎranian

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 120 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133

Province at its core and the Danubian Province. grations across the Atlantic took place.Amongst This development was reversed during the late the naticids the genera Ampullinopsis and Globu- Middle Miocene Sarmatian when the isolation laria, which both can be traced back to the West- of the Central Paratethys from the adjacent seas ern Tethyan Eocene, reached the Americas during may have caused a dramatic change of the water the Eocene. Ampullinopsis and the archaeogastro- chemistry (Pisera, 1996) leading to a severe im- pod Velates may be regarded as some of the most poverishment of the marine fauna.Simultaneous- successful Tethyan gastropods during the Paleo- ly, its reconnection with the Eastern Paratethys gene, since they are recorded from the eastern allowed the establishment of a rather uniform Paci¢c in the west to as far east as Pakistan or Sarmatian brackish gastropod fauna, being typi- Burma.Tethyan a⁄nities in the Eocene with the cal for the entire Paratethys Sea.During the latest Americas are documented by a large number of Middle Miocene and earliest Late Miocene the taxa, usually on the generic level and rarely on marine cycle ended within the Paratethys and a species level.Palmer (1967) united some of those brackish to freshwater lake system became in- taxa of proposed Tethyan origin, e.g. the genera stalled. Velates, Eovasum, Gisortia, and Terebellum, in the so-called ‘Velates perversus Group’.This small list 4.4. Transatlantic communication ^ witnessing an can be easily enlarged by data of Clarke and extensive Tethys Realm? Vokes (1936), Givens (1978, 1989), Jung (1974), and Squires and Advocate (1986) to include the The Caribbean and adjacent tropical areas dis- genera Agaronia, Akera, Amaurellina, Ampullella, play a characteristic faunistic composition at least Ampullinopsis, Athleta, Buccinorbis, Calyptrapho- from the Late Cretaceous onwards (Kau¡mann, rus, Campanilopa, Chedevillia, Clavilithes, Cornu- 1973).During the Paleogene this is expressed in lina, Crommium, Cryptochorda, Caricella, Ectino- the formation of a distinct large Central American chilus, Eocithara, Eocypraea, Eopleurotoma, Ga- biogeographic unit, which roughly covers the leodea, Gilbertina, Hipponix, Keilostoma, Laevity- geographic area of the modern Eastern Paci¢c phis, Lyria, Mazzalina, Muricopsis, Paraseraphs, and the Western Atlantic regions sensu Briggs Pleurofusia, Sassia, Semicassis, Seraphs, Strepsi- (1995).However, within the Central American dura, Streptochetus, Surculites and Volutilithes ^ bioprovinces a surprising £air of familiarity with a list which is far from being complete.Another the Western Tethys Region is perceptible from the Middle Eocene similarity is documented by the Eocene up to the Miocene. endemic American Dirocerithium, which shows Especially during the Paleogene, the Tethyan closest resemblance with the Tethyan genus Bella- heritage is obvious, since genera such as Ampulli- tara hinting at a common Tethyan ancestry nopsis, Globularia, Terebralia,orStrombus indi- (Woodring, 1959). cate immigration from the Tethyan bioprovince. An excellent example for the intensive transat- The faunistic exchange between the amphiatlantic lantic relationship during the Paleogene are the bioprovinces was a two-way story for sure, but strombids.The family arises during the Eocene due to the gaps in the fossil record, it is often when it develops a fair number of new taxa unclear whether a taxon migrated from the Old such as Dilatilabrum, Orthaulax and Oostrombus. World to the New World or vice versa.Although The genus Oostrombus, well established in the the amphiatlantic relations were discussed several Tethyan Eocene by species such as Oostrombus times in the literature (e.g. Maury, 1902; Cooke, auriculatus, displays one of the most striking dis- 1924; Gardner, 1924; Woodring, 1924a,b, 1928; junct patterns.The genus has been recorded by Kautsky, 1925; Vokes, 1964^1986; Savazzi, 1989; Olsson (1931) as Oostrombus chiraensis from the Lozouet et al., 1994), this important migration Late Eocene Chira Formation of Peru, by Clarke pattern is yet hardly embodied in palaeobiogeo- and Durham (1946) as Oostrombus cedroensis graphic concepts. from the Eocene of Colombia, and by Woodring Especially during the Eocene some major mi- (1959) as Oostrombus a¡. chiraensis from the Mid-

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 121 dle or Late Eocene Gatuncillo Formation of Pan- Alabama as Strombus albirupianus by Dall ama.From the Eocene onwards, the New World^ (1890^1903) ^ has been only documented from Old World route can also be documented for the the Late Oligocene of Georgia! two seemingly typical Western Tethyan genera Correspondingly, the Western Atlantic repre- Turbinella and Vasum. Turbinella ¢rst appeared sentatives of Aspella originated in the European in the Late Eocene of Peru with Turbinella peruvi- Oligocene (Vokes, 1975a) and ‘Cerithium’ halense ana (Olsson).Thereafter, two lineages developed of Dall (1917) has its closest relatives amongst the separately in the New and the Old World (Vokes, Tethyan Oligocene potamidids usually assigned to 1964).The latter is rooted in the earliest Old ‘Telescopium’ charpentieri or Tympanotonos strop- World species Turbinella episoma (Michelotti), pus.This Oligocene migration from the Tethys which appears in the Western Tethyan Oligocene. into the tropical Americas is obviously not re- Similarly, the genus Vasum developed in the New stricted to the gastropods but can also be docu- World where it is recorded from the Eocene of mented for other molluscs.For example, the Louisiana.During the Oligocene the genus man- strange bivalve Kuphus is very common in the aged to pass the Atlantic barrier and, with Vasum entire Mediterranean Oligocene and gained a subpugillare (d’Orbigny), is documented from the foothold in the Mid-American area at least in Oligocene of France.According to Vokes (1966) the Late Oligocene (Mans¢eld, 1940).A cosmo- all Early Miocene Old World representatives of politan transatlantic character has been also re- Vasum are very close to the ancestral Eocene ported for reef coral assemblages of the Eocene^ New World species Vasum humerosum Vaughan. Oligocene (Budd, 2000). Correspondingly, the genus Deromurex appears in Beside these Eocene/Oligocene migrations, a re- the Early Oligocene of the Mississippi Region, markable and well-documented cross-Atlantic mi- whilst its European counterpart is known only gration took place during the Burdigalian and from the Late Oligocene of western France probably early Middle Miocene.As far as can (Vokes, 1975a).The vicariating species pair is rep- be judged from the fossil record the main route resented by the New World Deromurex cookei was from the Western Tethyan Region to the (Vokes) and the Old World Deromurex cotteavi Western Atlantic, since many of the amphiatlantic (Meunier). taxa have their earliest record in the European The opposite route was taken by some western Cenozoic. Atlantic species of Cheilea.According to Vokes The most convincing evidence for those migra- (1975b) the New World species Cheilea equestris tions are amphiatlantic occurrences at the species (L.), which is ¢rst known from the Chipola For- level.Some of these species have been recognised mation (NW Florida), is a descendent of the Eo- in the Chipola Formation of northwestern Flori- cene Old World Cheila boutillieri (Cossman).Cor- da. E.g., Eudolium (Galeodolium) subfasciatum respondingly, Cheila uncinata (Reeve) can be Sacco appears in the Burdigalian and early Mid- traced to the European Eocene where Cheila ber- dle Miocene of the Mediterranean area and in the nayi (Cossman) is considered to be the ancestral Early Badenian of the Western Paratethys.In the form.Another amphiatlantic connection of a Burdigalian the species also appears in the Can- strombid taxon has been documented by Savazzi taure Formation of Venezuela and Chipola For- (1989) for the genus Orthaulax.It ¢rst appeared mation of northwestern Florida (Vokes, 1986; with Orthaulax dainelli in the early Middle Eocene Gibson-Smith and Gibson-Smith, 1988).Others, of northern Italy but thereafter was ubiquitous in such as Lindapterys, are documented by highly the Oligocene and Early Miocene of Central similar species at both sides of the Atlantic (Lo- America.Correspondingly, the genus Strombus zouet et al., 1994). Similarly, Calyptraea (Trochi- appeared in Tethyan Paleogene and crossed the ta) ornata Basterot and Calyptraea (Trochita) co- Atlantic barrier during the Oligocene.Notice staria Grateloup have been recorded from the that, according to Woodring (1959), the genus ^ Chipola Formation and the European Aquitanian formerly mentioned from Eocene sediments in and Burdigalian (Vokes, 1975b).High similarities

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 122 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 between European Atlantic Burdigalian faunas typhis vokesae Gertman from the Chipola Forma- and the Chipola fauna was also recognised by tion as closely related species.Two further species Dolin (1991) within the cypraeids. Trona leporina with a European ancestry are mentioned by calhounensis Dolin has its origin in the Old World Vokes (1966, 1979) from the Early Miocene (Bur- and emigrated to the Western Atlantic coast.The digalian?) of the Dominican Republic.Especially same has been suggested by Dolin (1991) for Tal- the Burdigalian Vasum tuberculatum Gabb can paria dominiciensis (Gabb) and Mauritia camp- hardly be distinguished from the Aquitanian Va- belliana (Pilsbry). sum aquitanicum Peyrot from France and might However, the New World^Old World route is even be conspeci¢c (Vokes, 1966).Similarly, Va- also documented.Following Dolin (1991) the sum pugnus Pilsbry and Johnson seems to be a Burdigalian Chipola species Siphocypraea chilona descendent of the French Vasum stephanense Dall seems to derive from a Caribbean stock and Peyrot, although a close relationship as discussed entered the Old World in the Early Miocene.Sim- by Vokes (1966, 1979) seems to be unlikely due to ilarly, a migration from west to east is also sup- the large stratigraphic gap between the Late Oli- ported by Bernasconi and Robba (1982) based on gocene French occurrence and the alleged Burdi- pteropods.According to Bernasconi and Robba galian occurrence in the Americas (Vasum stepha- (1982) the faunistic interchange of pteropods be- nense from St.-Etienne-d’Orthe is, according to tween the Caribbean and the Western Tethyan Lozouet (1998), of Chattian age, whilst Peyrot area occurred mainly from the Oligocene up to (1928) formerly assigned the species to the Aqui- the Middle Miocene. tanian). Less obvious at ¢rst sight, but very important The latest but subordinate Miocene transatlan- as indicators of transatlantic faunal relations are tic migration seems to be documented for the Late some pairs that probably represent vicariating Miocene.A remarkable example of this event is species.Worth mentioning and recently studied the genus Vittularia, which is an Old World ge- are pairs such as Bursa (Bufronariella) pelouatensis nus, appearing in the Oligocene of France.During (Cossmann and Peyrot) from the Early Miocene the Early and Middle Miocene the genus, repre- of the Aquitaine Basin and Bursa (Bufronariella) sented by Vittularia linguabovis (Basterot), be- chipolana Schmelz from the Burdigalian of Flori- comes ubiquitous in the Mediterranean as well da (Schmelz, 1997).The similarities are so striking as in the Paratethys.In the New World it ¢rst that Vokes (1973) erroneously even considered the appears during the Late Miocene (Vokes, 1967, Florida species to be synonymous with the Old 1977).Therefore, if a transatlantic migration is World gastropod.Vokes (1965a,b) also discussed considered, some minor, occasional faunal ex- the high similarities between the Florida and the change occurred even as late as the Late Miocene. European representatives of Chicoreus s.s. during Generally, the Miocene transatlantic exchange of the Late Burdigalian and the early Middle Mio- biota is still insu⁄ciently studied.For reef corals, cene.According to her, the genus originated in for example, dispersal across the Atlantic from the Tethyan area, having its roots in Eocene Old the Mediterranean to the Caribbean is considered World muricids, such as ‘Murex’ tricarinatus La- to have ceased at the end of the Oligocene (Frost, marck.Vicariating or at least very similar am- 1977; Budd, 2000).Contrary, Riegl and Piller (in phiatlantic species pairs are Chicoreus folidodes press) reported the genus Mussismilia, which was (Gardner), Chicoreus lepidotus (Vokes) and Chi- recorded from the Late Miocene as a mere Car- coreus dujardinoides (Vokes) from the New World ibbean element (Budd, 2000), from the Middle and Chicoreus aquitanicus (Grateloup), Chicoreus Miocene of the Styrian Basin (Danubian Prov- bourgeoisi (Tournouer) and Chicoreus dujardini ince). (Tournouer) from the early Middle Miocene of However, the similarities between the Western Europe.Similarly, Vokes (1974) mentioned Ptero- Atlantic and the European faunas are too poor at typhis wenzelidesi from the Badenian stage (early any time to allow a comparison on a hierarchy Middle Miocene) of the Vienna Basin and Ptero- lower than a biogeographic region.The Miocene

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 123 fauna of the ‘central’ part of this western Region and Peterson (1991) this early Gulf Stream in the has been very well studied, allowing a chie£y gas- North Atlantic is evident even during the Eocene tropod-based conception of the Miocene Mid- and Oligocene.In contrast, a ‘nearshore-hopping’ American bioprovinces as proposed by Woodring along the northern coasts as documented for (1974).He de¢nes a Miocene Caribbean Region some species in the Pleistocene can be excluded ( = province sensu Woodring, 1974), which is for all the amphiatlantic Eocene to Miocene characterised by the bivalve Clementia darienna taxa due to the strict climatic requirements of (Woodring) and a large number of gastropods, the warm-water faunas.Additionally, both fau- such as Orthaulax, Strombina, Metula, Potamides nas, especially the Mediterranean one, obviously suprasulcatus, and Stigmaulax guppiana.This bio- reached a stage of equilibrium during the Miocene province falls apart into the Mexican, West Indi- (Demarcq, 1987).Therefore, most of the intruders an, Central American^northern South American, probably had little chance to gain a foothold. Columbian^Venezuelan^Trinidad, Brazilian, and Despite the broad marine connections all these Ecuadorian^Peruvian provinces ( = subprovinces taxa rarely made their way in eastern direction sensu Woodring, 1974).It is worth mentioning into the Paci¢c.The distribution via the Atlantic that Woodring’s Brazilian Province is based strait is the only satisfying explanation, since evi- on the absence of taxa rather than on endem- dence for a Paci¢c route is missing.The transat- isms. lantic migration route during the Eocene was also Despite the amphiatlantic similarities, which emphasised by Clarke and Vokes (1936), who document some communication between two ma- stated that the similarities between the faunas of jor biogeographic units, the Atlantic was a barrier the American west coast and those of India are for many Tethyan genera, such as Pyrazus, Tibia low, thus excluding an arrival via a Paci¢c migra- s.s., Sulcogladius, Tudicla, and Amalda.With re- tion route, which was even at that time probably spect to the large total gastropod faunas the per- obstructed by the East Paci¢c Barrier. centage of shared species is rather small and requires a distinct separation of both biogeographic units on the level of a region.Many of the 5. Conclusions amphiatlantic taxa are only erratics within an otherwise independent Western Atlantic Region. Several phases of major transatlantic faunal ex- Therefore, most authors studying the amphiatlan- change can be deduced from the gastropod data tic distribution of tropical/subtropical molluscs in on both sides.The ¢rst phase comprises the Eo- the Cenozoic considered a more or less occasional cene to Oligocene with an Eocene peak.It could and accidental rafting of larvae as most probable probably be separated in several distinct migra- mode of transportation, representing sweepstakes tion waves, if stratigraphic resolution would be routes in terms of modern biogeography.How- better.A ¢rst westward migration wave is docu- ever, there seem to be some distinct peaks of sim- mented for Early Eocene or even Late Paleocene ilarity between Western Tethyan and American times (Squires and Advocate, 1986).The second gastropod faunas, e.g. during the Middle Eocene phase spans the Burdigalian and the early Middle or the Burdigalian.Such peaks, if not an artefact Miocene.This phase is very well documented and due to lack of investigations or adequate faunas again might be composed of two minor waves. in the intervening intervals, might indicate The ¢rst being characterised by migrations from changes in the Atlantic current systems, which the Old World into the Western Atlantic coastal repeatedly supported or suppressed the migration faunas mainly during the Burdigalian.The second of larvae.Correspondingly, the amphiatlantic took place somewhat later in the early Middle similarities are explained by Berggren and Hollis- Miocene, when American species or their closely ter (1974) and Barron and Peterson (1991) by an related forms suddenly appeared in European sec- early gyre-like Gulf Stream with limited poleward tions.Finally, a third phase may be suggested for extension.According to the simulations of Barron the Late Miocene.However, data documenting

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart 124 M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 this phase are scanty and the impact on the fau- graphic entity ^ the Danubian Province. At least at nas on both sides of the Atlantic was very low. the subprovince level a separation of the eastern The western Atlantic coast ^ similar to the Proto-Caspian Subprovince is necessary.The Indo-Paci¢c in late Early Miocene times for gen- Danubian Province was always in£uenced by era such as Tibia and Tudicla ^ obviously acted high immigration rates from the adjacent north- as refuge for several Tethyan genera such as ern (Oligocene) and/or the southern (Oligocene^ Ampullinopsis and Orthaulax, which managed to Miocene) bioprovinces.During phases of re- pass the Paleogene/Neogene boundary in the stricted marine connections and salinity crises Americas but became extinct in the European the Danubian Province was repeatedly threatened bioprovinces. by the expansion of the Proto-Caspian Subprov- Altogether the data furnish evidence for a cir- ince. cum-equatorial, Paleogene Tethys Realm which In the Early Miocene a rather homogeneous extended from Peru (Eastern Paci¢c!) through Proto-MediterraneanÂtlantic Region formed the early Caribbean Region across the Atlantic (France, entire Mediterranean, Central Iran). and the entire Mediterranean basins at least as During times of high emigration into the adjacent far east as Indonesia in the Eocene.During the Paratethys Sea, the Danubian Province has to be Oligocene this biogeographic unit still extended treated as part of the Proto-MediterraneanÂtlan- from the Mid-Americas via Pakistan and eastern tic Region (latest Early Miocene through early Africa to Indonesia with at least one centre of Middle Miocene).The biogeographic a⁄liation origin in the Mediterranean basins.In contrast, and hierarchical position of the Proto-Caspian an equivalent centre of origin is missing at that Subprovince, however, is obscure with respect to time in the Proto-Indo-Polynesian Province. the poor data on gastropods from the Eastern The similarities of the Mediterranean, the Ira- Paratethys. nian, the Pakistanian and the eastern African gas- Generally, endemism in the Paratethyan gastro- tropod faunas during the Oligocene requires to pod fauna is low compared to that of bivalves unite them in a Western Tethyan Region, which until the beginning isolation in the Sarmatian otherwise can be split into a MediterraneanÎra- (late Middle Miocene).In contrast to bivalves, nian Province (Western and Eastern Mediterra- the composition of the gastropod fauna was often nean, Cyrenaica, Armenia, Libya, Syria, Pales- massively in£uenced by immigrants rather than by tine, Central Iran), and a Western Indian^ local evolution. Eastern African Province (Oman, Pakistan, Soma- lia, Zanzibar, Kenya, Madagascar).A third province was probably established along the shores of Acknowledgements the Eastern Atlantic (France). The gastropod fauna of the Proto-Indo-West The authors thank Fred Ro«gl and Ortwin Paci¢c Region developed in an area, which is sim- Schultz (Museum of Natural History, Vienna) ilar to the geographic extension of the modern for fruitful discussions and for helpful comments Indo-Polynesian Province but did not dominate on an earlier draft of this paper.We are also the Oligocene faunas of the Western IndianÊast- grateful to Oleg Mandic (Institute for Palaeontol- ern African Province. Therefore, the often advo- ogy, Vienna) for information on bivalve distribu- cated ‘Indo-Paci¢c’ in£uence on Oligocene and tion.Our special thanks go to the members of the Early Miocene Central European faunas seems research projects on Late OligoceneÊarly Mio- to be a myth.However, with the Early Miocene cene circum-Mediterranean palaeobiological rela- the Indo-Paci¢c gastropod fauna established itself tions, A.Kroh (Graz), F.Schuster (Tu «bingen), within the area of the former Western Indian^ and U.Wielandt-Schuster (Frankfurt).This work Eastern African Province and replaced the ‘Indo- was supported by the Austrian FWF (P11886- European’ fauna completely. GEO) and the Deutsche Forschungsgemeinschaft The Paratethys Sea represents a distinct biogeo- (STE 857/1-1).

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Appendix 1 Miocene, Aquitanian); 4: Greece ^ Mesohellenic Trough (Early Miocene, Aquitanian); 5: Iran ^ Observed occurrences of gastropod taxa in Qom Basin (Early Miocene, Burdigalian); 6: studied basins: 1 Greece ^ Mesohellenic Trough Greece ^ Mesohellenic Trough (Early Miocene, (Oligocene); 2: Iran ^ Esfahan^Sirjan Basin (Oli- Burdigalian); 7: Turkey ^ Mut Basin (Early Mio- gocene); 3: Iran ^ Esfahan^Sirjan Basin (Early cene, Burdigalian)

Gastropod taxa 1 2 3 4567 Nerita (Theliostyla) plutonis Basterot o o o Nerita (Theliostyla) asperata Dujardin o Nerita caronis Brongniart o Agapilia picta (Fe'russac) o o o o Theodoxus? grateloupianus (Fe'russac) o Gibbula buchi (Dubois) o Gibbula a⁄nis (Eichwald) o Clanculus (Clanculopsis) araonis (Basterot) o Clanculus cerberi (Brongniart) o Paroxystele amedei (Brongniart) oo Jujubinus a¡. bucklandi (Basterot) o Jujubinus a¡. subcarinatus (Lamarck) o o Jujubinus sp. o Angaria scobina (Brongniart) o o Tectus nov.sp.1 o Tectus? nov.sp.2 o Tectus sp. o Tectus cf. rugosus (Grateloup) o Tectus lucasianus (Brongniart) o o Turbo nov.sp. o Turbo cf.nov.sp. o Turbinidae Operculum indet.o Littorina? sp.o Peasiella cf. girondica Benoist o Greveniella mesohellenica Harzhauser and Kowalke o Protoma cathedralis (Basterot) o o Protoma quadriplicata (Basterot) o o Peyrotia tauroperturritus (Sacco) o Peyrotia strangulata (Grateloup) o o o Peyrotia desmarestina (Basterot) o o o o Turritella (Turritella) terebralis Lamarck o o o Turritella (Turritella) gradata Menke oo Turritella (Haustator) venus d’Orbigny o o o Turritella (Haustator) asperula Brongniart o o Turritella (Haustator) conofasciata (Sacco) o Turritella (Haustator) vermicularis Brocchi o o Turritella (Haustator) eryna d’Orbigny oo Turritella (Zaria) angulata Sowerby o Turritella cf. subtriplicata d’Orbigny o Tenagodus cf. anguinus Linne' o Melanopsis impressa Krauss o o Bellatara palaeochroma nov.ssp. o Campanile charpentieri (Basterot) o Campanile sp.o Terebralia bidentata lignitarum (Eichwald) o o Terebralia bidentata (Defrance) o o o Terebralia a¡. subcorrugata (d’Orbigny) o

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Gastropod taxa 1 2 3 4567 Terebralia a¡. subcorrugata subinterrupta (d’Orbigny) o Granulolabium plicatum (Bruguiere) o o o Granulolabium (Tiaracerithium) pseudotiarella (d’Orbigny) o Granulolabium (Tiaracerithium) nov.sp. o Granulolabium (Tiaracerithium) cf.nov.sp. o Granulolabium pictum mitralis (Eichwald) o Tympanotonos margaritacus (Brocchi) o o Tympanotonos stroppus (Brongniart) o o Pyrazus daemon (Oppenheim) o Gourmya romeo (Bayan) o Clava ampullosa (Brongniart) o Clava voglinoi (Michelotti) o Cerithium calculosum Defrance o o o Cerithium vulgatum Bruguie're oo Cerithium globulosum Deshayes o Cerithium ex gr. granulinum (Bellardi and Michelotti) o Cerithium (Ptychocerithium) bronni turritoplicatum Sacco o Cerithiidae indet.1 o Cerithiidae indet.2 o Hemicerithium? sp.o Diastoma costellatum elongatum (Brongniart) o o Diastoma cf. alpinum (Tournouer) o Agapibittium greveniense Harzhauser and Kowalke o Bayania semidecussata (Lamarck) nov.ssp. o Aporrhais meridionalis (Basterot) o Aporrhais uttingerianus (Risso) o o Tibia dentata (Grateloup) oo Rimella (Dientomochilus) decussata (Defrance) o Rimella subrimosa d’Orbigny o Oostrombus auriculatus (Grateloup) o Oostrombus irregularis (Fuchs) o Strombus (Dilatilabrum) roegli Harzhauser o o o Strombus radix (Brongniart) o o Strombus bonelli (Brongniart) o o o o Strombus sp. o Xenophora cumulans (Brongniart) o o Xenophora deshayesi (Michelotti) o o o Xenophora cf. deshayesi (Michelotti) o Xenophora nov.sp. o Calyptraea cf. chinensis (Linne¤)o Hipponix cf. interruptus Michelotti o Cypraea cf. globosa (Dujardin) o Cypraea cf. subovum (d’Orbigny) o Cypraea (Zonaria) cf. fabagina Lamarck o Cypraea sp.1 o Cypraea sp.2 o Cypraea sp.3 o Cypraea sp.4 o Cypraea sp.5 o Cypraea sp.6 o Cypraea sp.7 o Proadusta inquidens inaequilabiata (Sacco) o Proadusta parvitala (Sacco) o o Trivia? sp.o Ampullinopsis crassatina (Lamarck) o o

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Gastropod taxa 1 2 3 4567 Globularia gibberosa (Grateloup) o o Globularia cf. callosa (Sowerby) o Natica sp. o Natica tigrina (Defrance) ooo Natica cf. subepiglottina d’Orbigny o o Polinices cf. redemptus (Michelotti) o Euspira catena helicina Brocchi o o Natica burdigalensis Mayer o Neverita olla (De Serres) o o o o Amaurellina (Euspirocrommium) oweni (Archiac and Haime) o Amaurellina (E.) a¡. sindiensis (Cossmann and Pissarro) o Amaurellina sp.1 o Amaurellina sp.2 o Sinum michaudi (Michelotti) o Sinum aquensis (Recluz) o Sinum cf. striatum (De Serres) o Sinum sp.o Cassis mamillaris Grateloup o o o Cassis vialensis Fuchs o Cassis sp.o Cassididae indet.o Galeodea? sp. o Semicassis cf. rondeleti apenninica (Sacco) o Semicassis grateloupi (Deshayes) o Sconsia striata miocenica Sacco o o Eudolium subfasciatum Sacco o Cassidaria cf. tauropomum Sacco o Sassia apenninica (Sassi) o Sassia cf. £andrica (De Koninck) o Charonia sp.o Cymatiidae indet. o Cymatium? sp.1 o Cymatium? sp.2 o Ficus condita (Brongniart) o oooo Ficus cingulata (Bronn) o Ficus oligo¢coides (Sacco) o Ficus sp. o Ficopsis (Fulguro¢cus) burdigalensis (Sowerby) o Cerithiopsis sp.o Murex partschi ssp. o Murex (Haustellum) tchihatche⁄ Archiac and Haime o Pterynopsis cf. tristichus (Beyrich) o Editharus polygonus (Lamarck) o Cymia hellenica Harzhauser and Kowalke o Euthriofusus burdigalensis (Defrance) o Dorsanum (Dorsanum) haueri excellens (Scha¡er) o Nassarius (Hinia) nov.sp. o Nassarius sp.1 o Nassarius sp.2 o Nassarius cf. semistriata (Brocchi) o Nassarius cf. restitutianus (Fontannes) o Nassarius sp.cf. limatus Chemnitz o Melongena lainei (Basterot) o o o Melongena semseyiana (Erdo«s) o Melongena cf. semseyiana (Erdo«s) o

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Gastropod taxa 1 2 3 4567 Melongena cornuta (Agassiz) o Melongena subcarinata (Lamarck) o Melongena a¡. basilica (Bellardi) o Melongena sp.o Pleuroploca tarbelliana (Grateloup) o Latirulus sp. o Fusinus cf. costellatus (Grateloup) o Streptodictyon cf. subelongatus (d’Orbigny) o Mitrella (Atilia) fallax (Ho«rnes and Auinger) o Acamptochetus submitraeformis (d’Orbigny) o Siphonalia sp. o Tudicla rusticula (Basterot) o o o Turbinella episoma (Michelotti) o Mitra sp. o Amalda (Baryspira) glandiformis Lamarck o o o Amalda (Baryspira) glandiformis anomala (Schlotheim) o Oliva (Anazola) cf. clavula (Lamarck) o o Oliva (Strephona) dufresnei Basterot o Olivella cf. longispira Bellardi o Athleta (Athleta) consanguinea (Bellardi) o Athleta (Athleta) ¢culina (Lamarck) oooo Volutilithes subambigua d’Orbigny Lyria magorum angustolonga Sacco o Lyria anceps (Michelotti) o o Lyria cf. anceps (Michelotti) Sveltia sp. o Terebra subtessellata d’Orbigny o Terebra pertusa Basterot o Terebra cf. pertusa subacuminata Peyrot o Conus (Conolithus) antediluvianus Bruguie're o o Conus (Conolithus) dujardini Deshayes oooo Conus (Lithoconus) antiquus Lamarck oo Conus (Lithoconus) cf. mercati (Brocchi) o o Conus (Lithoconus) nov.sp. o Conus (Stephanoconus) cf. carcarensis (Sacco) o Conus (Leptoconus) diversiformis Deshayes o o Conus sp.1 o Conus sp.2 o Conus sp.3 o Genota ramosa (Basterot) o o o Acamptogenotia intorta Brocchi o Bathytoma cataphracta (Brocchi) o Clavatula asperulata(Lamarck) ooo Clavatula cf. glaberrima (Grateloup) o Clavatula cf. carinifera Grateloup o Clavatula calcarata Grateloup ssp.o Clavatula sp.1 o Clavatula sp.2 o Clavus sp. o Raphitoma sp.1 o Raphitoma sp.2 o Turris cf. monilis (Brocchi) o Turris sp.1 o Turris sp.2 o Turridae indet.1 o

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Gastropod taxa 1 2 3 4567 Turridae indet.2 o Stenodrillia obeliscus (Desmoulins) o Scaphander lignarius grateloupi (Michelotti) o o Sabatia a¡. utriculus (Brocchi) o Bulla? sp. o Architectonica cf. umbrosa (Brongniart) o Architectonica carocollata (Lamarck) o Architectonica sp. o Otopleura mitrula (Basterot) o Ringicula auriculata paulucciae Morlet o Limacina miorostralis (Kautsky) o Vaginella depressa Daudin o

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