Circum-Mediterranean Oligo^Miocene biogeographic evolution ... · Circum-Mediterranean Oligo^Miocene biogeographic evolution ^ the gastropods’ point of view Mathias Harzhausera,

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

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

a Naturhistorisches Museum Wien, Geologisch-Pala«ontologische Abteilung, Burgring 7, A-1014 Vienna, Austriab 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 Sivasbasins), in the Mesohellenic Basin, and in northeastern Egypt, a new palaeobiogeographic concept for the Oligoceneand Miocene in the circum-Mediterranean area with special emphasis on the distribution patterns of gastropod faunasis presented. A very strict biogeographic terminology is proposed to avoid the common fusion of geographic, faunisticand geodynamic terms in the palaeontologists’ jargon. Our main interest focuses on the affinities and the faunisticinteractions 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 Europeanmollusc faunas, as often vaguely indicated in the literature, is critically evaluated. This leads to the establishment of aMediterraneanÎranian Province and a Western IndianÊastern African Province as palaeobiogeographic subunits ofthe Western Tethys Region during the Oligocene. Due to major geodynamic and related biogeographic changes in theEarly Miocene the Western Tethys Region no longer existed and was replaced by the Proto-MediterraneanÂtlanticRegion. 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^Miocenenearshore mollusc faunas is demonstrated. Altogether this paper is a contribution to the discussion on the bio-geographic concepts, classification and nomenclature in palaeontology as initiated by Westermann [Palaeogeogr. Pa-laeoclimatol. Palaeobiogeogr. 158 (2000) 1^13; 163 (2000) 49^68]. @ 2002 Elsevier Science B.V. All rights reserved.

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

1. Introduction

During the 19th and 20th centuries a large da-taset for Cenozoic gastropods from the WesternMediterranean, the European Atlantic and Cen-

tral Europe has been produced (e.g. Ho«rnes,1856; Bellardi, 1872^1890; Ho«rnes and Auinger,1879^1891; Sacco, 1890^1904; Cossmann andPeyrot, 1909^1934). This led to an early under-standing of Eocene to Miocene biogeography ofthis area. In an eastern direction, however, infor-mation concerning the Eastern Mediterraneanand the Middle East was scanty. Gastropodswere documented as rare byproducts during inves-

0031-0182 / 02 / $ ^ see front matter @ 2002 Elsevier Science B.V. All rights reserved.PII: S 0 0 3 1 - 0 1 8 2 ( 0 1 ) 0 0 4 6 4 - 3

* Corresponding author.E-mail address: [email protected] (W.E. Piller).

PALAEO 2816 31-5-02 Cyaan Magenta Geel Zwart

Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133

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tigations of regional geology. Due to the strongpreservational bias related to the predominantlystudied calcareous facies, the mollusc studies fo-cused mainly on oysters and pectinids (Douglas,1927, 1928). On the other hand, scattered andinhomogeneous data on Eocene to Miocene gas-tropod faunas from Pakistan, Somalia, Kenya,Madagascar, Southern India, Burma, and Java(Noetling, 1901; Martin, 1914^1915, 1916^1917,1921^1922; Davies, 1923; Vredenburg, 1925^1928; Collignon and Cottreau, 1927; Cox,1930a; Azzaroli, 1958) allowed a glance at thecomposition and evolution of the early Indo-West Paci¢c faunas. The comparison of Mediter-ranean and Asian faunas, representing two major

biogeographic entities, was thus severely ham-pered by a palaeontological no man’s land span-ning the entire eastern Mediterranean and theconnecting Tethyan seaway. Therefore, modernpalaeobiogeographic interpretations of late Te-thyan mollusc faunas (Piccoli, 1984; Piccoli etal., 1986, 1991) were usually based on the well-studied Northern Italian and Pakistanian faunas.The vast area in-between was taken into consid-eration only cursorily since their poorly docu-mented and low diversity faunas were consideredto be of little palaeobiogeographic value.

To close this gap we focused on this poorlystudied area. Detailed investigations of Oligoceneand 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, Turkeyand Greece form the base for the present study, whereas the low diversity and poorly preserved faunas from Egypt and Omanare only supplementary. Additionally, the position of some selected European and Asian reference faunas is indicated. Amongthese, the Italian and Pakistanian gastropod faunas turned out to represent the cornerstones for biogeographic considerations inthe circum-Mediterranean area.


M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133104

hellenic Basin, the Turkish Mut and Sivas basinsand especially the Iranian Qom and Esfahan^Sir-jan basins (Fig. 1) yielded a high number of shal-low marine gastropod faunas. Hence, the faunisticgap in the Eastern Mediterranean was ¢lled anddata from Iran, which is identi¢ed as key areabetween Mediterranean and Paci¢c faunas, areavailable for the ¢rst time. Based on these dataa new concept of the palaeobiogeography of theOligocene and Early Miocene Western Tethys canbe developed.

The main interest focussed on the dawn of thefauna of the MediterraneanÎranian Province. Itsbirth from the Oligocene faunal-stock of the vastTethys Realm is outlined, and the in£uence of theProto-Indo-West Paci¢c Region on the EarlyMiocene gastropod faunas of the Iranian QomBasin and the faunas of the Mediterranean areais evaluated.

2. Biogeographic basics

A summary of the various concepts and hierar-chies in biogeography is given in Sedlag and Wei-nert (1987), who recommend a terminology corre-sponding to Lattin (1967). According to theseauthors the usually rather vaguely used term‘province’ should be con¢ned to a part of a sub-region. The latter term corresponds largely to theterm ‘province’ as used by Woodward (1856) andHedgpeth (1957). Finally, the subregions areunited into large biogeographic units ^ the re-gions.

A strict de¢nition of the various biogeographicunits based on percentages of endemisms is pre-sented by Kau¡mann (1973) and Kau¡mann andScott (1976). According to this de¢nition a realmis characterised by more than 75% endemism, aregion comprises 50^75%, a province 25^50%, asubprovince 10^25% and ¢nally an endemic centrereaches 5^10% endemism.

Generally, the term province as proposed bySedlag and Weinert (1987) is accepted by modernbiogeographers including Briggs (1995), Long-hurst (1998), and Westermann (2000a,b). None-theless the classi¢cation of higher ranks is in astate of £ux. Briggs uses ‘region’ as a synonym

to Lattin’s subregion whilst the subrealm of Wes-termann (2000b) seems to be more or less synon-ymous to ‘region’ of most biogeographers. Long-hurst (1998) unites the provinces in biomes.However, the geographic and biogeographicboundaries of the modern shallow-water subdivi-sions are more or less the same for all authors.

Recently, biogeographic principles and the de-nomination of units and their hierarchical rankingwere discussed by Westermann (2000a,b). In his‘Rules of Biogeographic Nomenclature’, Wester-mann (2000a) proposes a strict use of biogeo-graphic terms, with emphasis on homonyms andsynonyms as well as stability throughout geolog-ical time. Westermann (2000a) tries to establishthe general term ‘biochore’ for all biogeographicunits. The term ‘biochore’ is, however, highly con-troversially used: for many neontologists ‘bio-chore’ has a strong ecological component withoutgeographic restrictions (e.g. biochores are grass-land, forest, and desert) and even for palaeontol-ogists such as Lehmann (1996) ‘Biochorion’ is asmall area within a biotope (e.g. a treetrunck or acarcass). Thus a biochore in modern biology êspecially in botany ^ is a subdivision of the bio-cycle which contains a group of biotopes that re-semble one another. This diverging use of ‘bio-chore’ is based on the unclear translation of theGreek bio-‘MgbYc’ as living-‘space’ instead of liv-ing-‘place’. Therefore, not to prolong the confu-sions, we prefer ‘biogeographic unit’ instead ofthe term biochore.

Westermann (2000a,b) recommends the ranksSuperrealm, Realm, Subrealm, Province, and Sub-province as valid hierarchical categories for thede¢nition of his biochores. The term region isfor informal use only. We believe, however, thatthe well-established and widely used term ‘region’should be obligatory and not replaced by an op-tional term as ‘Subrealm’. Therefore, in this paperthe terms province and region are mainly usedsensu Briggs (1995). In contrast, we follow Wes-termann (2000a) in his proposed rule, that thereplacement of names is acceptable under certainconditions, e.g. extreme biochore expansion orcontraction, exceptional geologic change at re-gional or global scale, and at mass extinctionevents.


M. Harzhauser et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 183 (2002) 103^133 105

The hierarchical categories of biogeographicranks used in this paper are:

Realm ^ Region ^ Province.For re¢nement these biogeographic core-units

can be subdivided by adding the pre¢xes Super-or Sub- respectively.

3. Materials and methods

The shallow marine Late Oligocene Chattianfaunas from Mesolouri, Doutsiko and Kipourioin the Mesohellenic Trough (Greece) (Fig. 1)yielded more than 90 taxa, representing about50 species. An equal number of species was re-corded from Abadeh in Iran. The Early Miocenenearshore faunas from Turkey, Greece and Iranare documented by 145 taxa composed of 46,50 and 58 gastropod species respectively (Appen-dix 1). These faunas were compared and coin-cidence or di¡erences between the faunas on spe-cies level were determined. Due to the partlyrather small faunas, only presence/absence dataare used for the analysis, yielding percentagesof taxa occurring in two or more investigatedfaunas.

Finally, the Iranian and Eastern Mediterraneanfaunas were compared with those of the WesternMediterranean, the Atlantic, the Paratethys andespecially with those from Pakistan. Our knowl-edge of these reference faunas is rather inhomoge-neous. The small Eastern Mediterranean and Ira-nian faunas consisting of approximately 50^60species each are in contrast to the extremely richFrench and Italian faunas. For example, Coss-mann and Peyrot (1909^1932) describe morethan 600 Burdigalian and more than 500 Aquita-nian gastropods. Bellardi (1872^1904) report evenmore than 980 Early Miocene species. The Oligo-cene gastropod faunas of northern Italy consist ofat least 300 species, estimated from the variousmonographs of Fuchs (1870), Oppenheim (1900)and Bellardi (1872^1904). From the Late Oligo-cene of the Western Paratethys approximately 250species are recorded by Wol¡ (1896^1897), Baldi(1973) and Ho«lzl (1962), and more than 300 spe-cies are described from the Early Miocene Para-tethys by Steininger and Senes (1971), Ho«lzl in

Papp et al. (1973) and Harzhauser (2002). In con-trast, the Pakistanian faunas with 77 Oligoceneand 81 Early Miocene species described by Vre-denburg (1925^1928) are considerably smaller.The discrepancy is even more obvious when east-ern African and southern Indian gastropod fau-nas are considered. Thus, only 23 species are de-scribed from the Oligocene and Early Miocene ofSomalia (Azzaroli, 1958), 28 from the Miocene ofCeylon (Cox, 1930a), 13 from the Miocene ofMadagascar (Collignon and Cottreau, 1927),and only eight from the Miocene of Kenya (Da-vies, 1923).

The palaeobiogeographic interpretation of theseinhomogeneous faunas based on statistical meth-ods seems to be inappropriate. On the one hand,for many of the mentioned reference faunas mod-ern revisions are completely missing. Therefore areliable comparison of literature-based faunas isdi⁄cult, a fact that is obvious to every taxonomistafter a short look at the usually long and mani-fold synonymy lists in modern revisions. This isnot restricted to ‘di⁄cult’ groups, such as the nas-sariids or conids, but can also be documented for‘simple’ taxa such as the melongenids or strom-bids. E.g., the characteristic Oligocene Melongenasemseyiana (Erdo«s) ^ a seemingly endemic Para-tethyan species (Baldi, 1973) ^ was described atleast under three synonyms (see Mikuz, 1999).Now the ‘endemic’ Hungarian species turned upin Austrian, Croatian, Hungarian, Slovenian,Greek and maybe also in Turkish faunas.

Similarly, the endemism-index should not beoveremphasised, since endemism may partly de-pend on the quality of research. Recently, Lo-zouet (1998, 1999) and Valde¤s and Lozouet(2000) described 170 new species from the Oligo-cene and Lower Miocene of the Aquitaine Basin,indicating high endemism from the armchair.However, this high number of ‘endemic’ speciesseems mainly related to the lack of equivalent,elaborate studies in the Mediterranean area. Theinsu⁄cient study of Mediterranean faunas wasalso stressed by Studencka et al. (1998) for Mid-dle Miocene bivalve faunas. Thus, the lack ofmodern revisions caused a higher percentage ofbivalves common to the Atlantic and the Parate-thys (60%) than of species common to the directly



connected Mediterranean and the Paratethys(47%).

Furthermore, the taxonomic overcompletenessof some faunas, caused by the tendency of older

workers such as Ho«rnes and Auinger (1879^1891)to split ‘good’ species into morphs, is hard tohandle in statistics. These so-called subspecies orvariations tend to ‘evolve’ into species in subse-

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).



quent studies, resulting in several local ‘species’which mask palaeobiogeographic relations. Fi-nally, open nomenclature is hard to adapt to sta-tistics. This can be demonstrated very impres-sively by the otherwise highly valuable lists ofEarly Oligocene Paratethys faunas from Hungaryby Baldi (1986), which comprise about 37% oftaxa in open nomenclature. Within the easternAfrican gastropod faunas this percentage mayeven reach 61% (see Azzaroli, 1958).

Hence, in this paper the authors try to focus onthe qualitative and semiquantitative compositionof typical nearshore assemblages, which are sug-gested to characterise the biogeographic units invarious time slices. For biostratigraphic and geo-chronologic correlation see Fig. 2.

4. Circum-Mediterranean biogeography

Woodward (1856) was one of the ¢rst who triedto divide the marine mollusc faunas in bioprovin-ces with characteristic compositions. He proposedthe rule that 50% of the species within a provinceshould be peculiar (endemic) to it. Physical fac-tors such as geographic barriers and climate wererecognised by Woodward as dividing lines be-tween distinct provinces. He additionally takeshistorical factors into account such as ‘di¡erentdistribution of land and water over the surfaceof the globe’. Woodward’s mollusc-provinces arestill accepted by some authors working on mol-luscs such as Sabelli (1980). Nonetheless, the bio-province concept was improved during the 20th

century by authors such as Ekman (1953) orHedgpeth (1957), who attempted to integrate thelittoral fauna as a whole.

In the following the authors provide a brief re-view of selected recent and historical bioprovincesand their geographic pattern within the circum-Mediterranean area.

4.1. Modern bioprovinces

4.1.1. MediterraneanÂtlantic RegionAccording to Briggs (1995) this region com-

prises the Lusitania, Black Sea, Caspian, andAral provinces. Among them the Lusitania Prov-

ince is the most extensive, including the entireMediterranean Sea but also parts of the Atlanticcoast. Its westernmost extension are the Azores;in the north it includes the Bay of Biscay, and itreaches as far south as Madeira and the Canaries.It is bordered in the south by the Cape VerdeIslands, which are part of the Eastern AtlanticProvince ( = Senegalese Province sensu Wood-ward) and the English Channel in the north.

The term Lusitanian Region as used by Ekman(1953) has a quite di¡erent meaning. He con¢nesthe term to the Atlantic part north of the Straitsof Gibraltar. Consequently, he separates thesouthern part as Mauretanian Region. The Medi-terranean Sea^Black Sea Province introduced byLonghurst (1998) corresponds partly to the Lusi-tania Province of Briggs (1995). A clear di¡erenceis only that Longhurst treats the Bay of Biscay aspart of the Northeast Atlantic Shelves Province.The concept of Briggs (1995) concerns the shal-low-water fauna whereas the provinces introducedby Longhurst (1998) are based on pelagic organ-isms.

In contrast to Briggs (1995), who maintainsthat the Recent Caspian and the Aral Sea provin-ces have derived their biotas from the Mediterra-nean, these provinces are considered herein as bi-ogeographic and geologic descendants of theParatethys Sea. This is strongly supported bythe data given in Ekman (1953), who describesthe Caspian fauna to be ‘practically a pure Sar-matic fauna’. Ekman (1953) even emphasised thesmall percentage of species common to the Med-iterranean Sea and regards the Caspian fauna tobe of Paratethyan ( = Sarmatic sensu Ekman) ori-gin.

Similarly, the Black Sea Basin was part of theParatethys, but became part of the Mediterra-neanÂtlantic Region during the Pleistocene andthe Holocene by its connection to the Aegean Seavia the Marmara Sea.

4.1.2. Eastern Atlantic^Boreal RegionThe Region extends from the British Channel

north via the North Sea to the Baltic Sea andreaches from the Celtic Shelf as far north as theHebrides. It is limited to the south and the south-west by the Lusitania Province and to the north



and the north-east by the Arctic Region (Briggs,1995). In contrast to Woodward’s Celtic Province,the Eastern Atlantic^Boreal Region includes alsoIceland, the Norwegian coast and extends as farnorth as Murmansk.

Several terms are used in the literature as roughsynonyms for this Region. Especially Europeanpalaeontologists usually called the geographic an-cestor of this Region the ‘Boreal Province’ (Kaut-sky, 1925; Baldi, 1973), the ‘Scandic Region’(Ho«lzl, 1962), or simply the ‘northern province’(Senes, 1958), while Ro«gl and Steininger (1983,1984) introduced the modern term ‘Atlantic^Bo-real Bioprovince’.

4.1.3. Indo-West Paci¢c RegionThe term ‘Indo-West Paci¢c’ was introduced by

Ekman (1934), who united the Indo-Paci¢c, theWestern Paci¢c, and parts of the Central Paci¢cin his biogeographic unit. Later, Briggs (1995)demonstrated that the Region has to be extendedto the Easter Islands, thus introducing an areathat nearly extends to the coasts of America.This vast Region is divided into the Indo-Polyne-sian, the Western Indian Ocean, the Red Sea, theNorth-western Australian, the Hawaiian, the Eas-ter Island, and the Marquesas provinces. The mod-ern Western Indian Ocean Province extends fromthe Persian Gulf along the eastern African shelf încluding the Seychelles, Madagascar and theMascarene Islands ^ as far south as Kwa Zulunorth of Natal in South Africa (Kilburn and Rip-pey, 1982). The Red Sea is excluded from thisProvince, but forms a distinct Province due tothe high percentage of endemics (Briggs, 1995).In an easterly direction the Western Indian OceanProvince is bordered by the larger Indo-PolynesianProvince. This Province extends from the entranceof the Persian Gulf, via the Arabian Sea, alongthe Indian shelf to Sri Lanka and the Bay ofBengal and includes Malaysia, Indonesia andEastern Australia. To the north it reaches as faras the Japanese Amami Islands.

The border between these two provinces is notunequivocal. Even the de¢nition given in Briggs(1995) markedly di¡ers from his drawing since thesketch includes also the Iranian and Pakistaniancoasts of the Arabian Sea in the Western Indian

Ocean Province, while in the text the Indo-Poly-nesian Province reaches as far west as the entranceof the Persian Gulf. Other limits of the WesternIndian Ocean Province are given by Macnae(1968), who includes the entire coasts of thenorthern Arabian Sea as far as the Gulf ofKhambhat in north-western India. Additionally,the Western Indian Ocean Province as de¢ned byPor (1989) would reach to southern India, to thecoast between Mangalore and Calicut.

4.2. Historical biogeographic units

4.2.1. Tethys RealmThe term ‘Tethys’ was introduced by the Aus-

trian geologist E. Suess (1893) as a palaeogeo-graphic entity. Herein ‘Tethys Realm’ is used ina strict biogeographic sense and must not be in-termingled with the geographic term ‘Tethys’ asdescribed by Hsu« and Bernoulli (1978) and Sen-go«r (1998). This is of great importance, since thegeographic Tethys (sensu Hsu« and Bernoulli,1978) or the tectonically de¢ned Tethys (sensuSengo«r, 1998) vanished during the Eocene whenIndia collided with Laurasia.

Originally, the biogeographic Tethys Realm, asde¢ned by Kau¡mann (1973), was applied to Me-sozoic faunas. Kau¡man’s Tethys Realm is basedon Cretaceous bivalves and represents a vast, cir-cum-equatorial area. For the Cenozoic era thebiogeographic term Tethys is still used by Popov(1993) when de¢ning zoogeographic units for LateEocene bivalves. This Tethys Region sensu Popovis modi¢ed herein with respect to hierarchy andenlarged as Tethys Realm (Fig. 3(left, top)). Ac-cording to Popov, the Late Eocene Tethys ofwestern Eurasia can be divided into an Indo-Afri-can Region and a Mediterranean Region (subre-gions sensu Popov, 1993). At that time the south-ern Tethyan faunas of Egypt, Somalia and Indiadisplayed highest resemblance, but di¡ered clearlyfrom the northern Tethyan ones from Italy andsoutheastern France.

Simultaneously, with the narrowing of the Te-thys in the north of the Indian shield, the south-ern seaway and the ancestral Indian Oceanwidened. This seaway warranted a continuousconnection of the western and eastern tropical







Tethyan faunas, hence allowing to extend the bio-geographic term Tethys Realm also for Oligoceneand Early Miocene times. With respect to the di-vergent meaning of Tethys Ocean and TethysRealm, it might be more correct to name the bio-geographic unit the ‘Proto-MediterraneanÎndo-African^Western Paci¢c Realm’ ^ an obviouslyunwieldy term. Thus the term Tethys Realm isfavoured in this paper.

The post-Eocene Tethys Realm existed duringthe entire Oligocene and during the very EarlyMiocene without major interruption. The biogeo-graphic unit vanishes during the Burdigalian andwith the formation of the Gomphotherium land-bridge between Eurasia and the Arabian plate inthe Late Burdigalian (Ro«gl, 1998). The ¢nalbreakdown of the Tethys Realm is also evidentfrom a geographical viewpoint. However, as men-tioned above, the Tethys Sea vanished much ear-lier during the Eocene.

During the Oligocene and the Early Miocene,the Tethys Realm covered an extraordinarily largearea. Its main territory was an area, which coin-cides more or less with the modern Indo-WestPaci¢c Region, but in western direction it alsoincluded the area of the modern MediterraneanSea. A broad connection between the Iranian

and the African/Arabian plates enabled a fair ex-change of shallow-water faunas between theseareas. In the eastern direction, the Indonesian sea-way acted as a marine connection between the‘Indo-Paci¢c’ and the western Paci¢c areas whichlasted until the early Middle Miocene (Ogasawaraand Noda, 1996; Nishimura and Suparka, 1997).A tropical, equatorial current £owed through thisPaci¢cÎndian Ocean gateway in a westerly direc-tion (Tsuchi, 1997). Tethyan gastropods, such asTibia, Telescopium, Terebralia, or Vicarya, how-ever, were spread as far north as Southern Japanduring the Early and early Middle Miocene (Oga-sawara and Noda, 1996; Noda and Watanabe,1996).

Corresponding to the modern Indo-West Paci¢cRegion, a separation of this vast marine sphereinto several biogeographic units is to be expected.Due to inadequate data it is currently impossibleto designate any of these boundaries exactly.Nevertheless the representative Oligocene andMiocene faunas of Italy (Bellardi, 1872^1904;Fuchs, 1870), Greece, Central Iran, Pakistan (Vre-denburg, 1925^1928; Iqbal, 1969a,b, 1980), Bur-ma (Noetling, 1901), Java (Martin, 1916^1917,1921^1922), Celebes (Beets, 1942, 1943), and theSunda 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 biogeo-graphic 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 ma-rine sphere displays a distinct S^N trend expressed by a northern entity united in the North European Region and a southern en-tity, the Tethys Realm. The Tethys Realm can be divided into two Regions, which were named the ‘Mediterranean’ Region andthe ‘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 shoresof the Tethys facilitated the distribution of ‘European’ gastropods as far southeast as Pakistan. However, a distinct N^S gradientis indicated by the characteristic faunistic composition in the Western IndianÊastern African Province. In the Paratethys Sea, theDanubian Province and the Proto-Caspian Subprovince formed as a result of the reduced connections to the Tethys. The complexbiogeographic 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 MediterraneanÎranianProvince. 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: Theseparation of the Proto-MediterraneanÂtlantic Region from the Proto-Indo-West Paci¢c Region is completed by the formation ofthe 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.



compositions, which prove the existence of dis-tinctive regions and provinces. Some of themwill be proposed in the following.

4.2.1.1. Western Tethys RegionPaleogene : During the Oligocene, the Western

Tethys Region ( = Indo-European sensu Ekman,1953) represents part of the Tethys Realm. It cov-ered the area of the modern Mediterranean butprobably included also the European Atlanticcoast up to the Bay of Biscay in the west andreached to Pakistan, Somalia and Zanzibar inthe east and the south (Fig. 3(left, bottom)).Most characteristic for the Western Tethys Regionare the faunas of Northern Italy. The Mediterra-neanÎranian Province (see below) is designatedherein as the type unit for the Western TethysRegion ( = chorotype sensu Westermann, 2000a).

Its eastern border seems to have been locatedsomewhere along the coast of Southern India,since the south-eastern Asian faunas display nosimilarities either with those of Pakistan (Vlerk,1931) or with those of Europe. Such faunistic de-marcation lines have also been documented forthe Eocene by Cotter (1923) for the bivalves. Hestated that Burma, Sumatra, Java, and Borneoconstituted a distinct bioprovince, di¡erent fromthe Sind^Baluchistan and Egypt faunas. Someyears later Cox (1930b, 1936) concluded that thea⁄nities of the Bahrain and southern Iranian fau-nas with those of north-western India, Somaliaand Egypt hint at an Indo-African biogeographicentity ( = Indo-African fauna sensu Cox, 1930b).Correspondingly, Martin (1931) proposed anIndo-Malayian bioprovince based on the homoge-neity of Late Eocene faunas from eastern India toJava and Burma, and on their marked di¡erencefrom that of western India^Pakistan. This biogeo-graphic pattern ¢ts well the simulations of Barronand Peterson (1991) who demonstrated that Bur-ma and Java were separated from north-westernIndia during the Eocene by a strong westernboundary current, while the Indian Ocean circu-lation pattern and the Somali current are evidentonly from ‘middle’ Oligocene onwards (Barronand Peterson, 1991). Hence, in the east the Paleo-gene Western Tethys Region is bordered by a ma-jor adjacent biogeographic unit inhabited by these

Indo-Malayian faunas, called herein informallythe Proto-Indo-Polynesian Province of the Pro-to-Indo-West Paci¢c Region. The fauna of thisProto-Indo-West Paci¢c Region displays hardlyany resemblance with the Oligocene gastropodfauna of the Western IndianÊastern AfricanProvince. Neither the Pakistanian nor the easternAfrican faunas have species in common with theOligocene gastropod faunas of the Proto-Indo-Polynesian Province (e.g. Buton-fauna describedby Martin (1933, 1935) and Beets (1942, 1943)).

During the Oligocene a separation within theWestern Tethys Region in at least two provincesis obvious.

(1) MediterraneanÎranian Province. The hereinproposed MediterraneanÎranian Province becameestablished in the area of the modern Mediterra-nean Sea and had its south-easternmost knownextension in Central Iran (Fig. 3(left, bottom)).The new term is introduced instead of ‘Mediter-ranean’ Province, because of the di¡erent geo-graphic extension and moreover because of thesynonymous use of the term for Mesozoic andCenozoic biogeographic units with di¡erent bioticcharacteristics. Characteristic gastropod faunas ofthis province are those of Northern Italy (Fuchs,1870; Bellardi, 1872^1890; Sacco, 1890^1904), thefauna of the Mesohellenic Trough in Greece(Brunn, 1956; Harzhauser, 2000), and the faunaof the Esfahan^Sirjan Basin in Central Iran(Harzhauser, 2000; Fig. 4a,c). It is best character-ised by its rich strombid fauna which seems to bea heritage of the highly diverse, blooming Eocenestrombid fauna. None of the giant species of Oos-trombus or Dilatilabrum is known from any Oli-gocene section southeast of Iran. Characteristicgastropods of the MediterraneanÎranian Prov-ince are, among many others, Tectus lucasianus(Brongniart), Angaria scobina (Brongniart), Turri-tella asperula Brongniart, Turritella conofasciata(Sacco), Peyrotia strangulata (Grateloup), Campa-nile charpentieri (Basterot), Granulolabium plica-tum (Bruguie're), Tympanotonos stroppus (Brong-niart), Clava ampullosa (Brongniart), Clava vog-linoi (Michelotti), Cerithium calculosum Defrance,Gourmya romeo (Bayan), Diastoma costellatumelongatum (Brongniart), Proadusta parvitala (Sac-co), Ampullinopsis crassatina (Lamarck), Globu-



laria gibberosa (Grateloup), Sinum aquensis (Re-cluz), Bayania semidecussata (Lamarck), Oostrom-bus auriculatus (Grateloup), Oostrombus irregu-laris (Fuchs), Strombus radix (Brongniart),Strombus (Dilatilabrum) roegli Harzhauser, Xeno-phora cumulans (Brongniart), Cassis mamillarisGrateloup, Cassis vialensis Fuchs, Turbinella epi-soma (Michelotti), Volutilithes subambigua d’Or-bigny, Lyria anceps (Michelotti), Conus (Leptoco-nus) diversiformis Deshayes, and Terebra subtes-sellata d’Orbigny.

(2) Western IndianÊastern African Province.Towards the southeast the MediterraneanÎranianProvince is bordered by a very distinct biogeo-graphic unit, which is introduced herein as theWestern IndianÊastern African Province (Fig.

3(left, bottom)). The term ‘Indo-African’ shouldbe avoided, because of its quite di¡erent use in theliterature, e.g. Popov (1993) versus Cox (1930b).It is represented by the fauna of the PakistanianNari Formation, which has at least 40 species incommon with those of Northern Italy (Vlerk,1931) (Fig. 4). However, the high number of spe-cies unknown from European localities (about50^60 species) re£ects a rather independent devel-opment in this south-eastern termination of theWestern Tethys Region. This Western IndianÊastern African Province is documented by thePakistanian faunas (Vredenburg, 1925^1928; Iq-bal, 1969a,b, 1980), but can also be traced alongthe coast of Eastern Africa as con¢rmed by thefaunas of Somalia described by Azzaroli (1958)

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 thehomogeneity within the MediterraneanÎranian Province. During the Early Miocene (b,d) this homogeneity is strongly indicatedwithin the Proto-MediterraneanÂtlantic Region. In contrast, the Pakistanian gastropod faunas lost the similarities and have only4^7% 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 EarlyMiocene clearly witness the formation of a major biogeographic boundary between V30‡ and 35‡ N.



and Kenya (Davies, 1923). The Western IndianÊastern African Province includes also the area ofOman (Fig. 3(left, bottom)) as documented by asmall reefal gastropod fauna kindly donated byM. Bernecker (Erlangen). Characteristic gastro-pods of the Western IndianÊastern African Prov-ince are ‘Turbo (Senectus)’ naricus Vredenburg,‘Turbo (Olearia)’ protocepoides Vredenburg, Pro-toma retrodilatatum Vredenburg, Turritella naricaVredenburg, Semicassis oligocalantica (Vreden-burg), Harpa (Eocithara) narica Vredenburg, Bel-latara narica (Vredenburg), or Cerithium sindienseVredenburg.

Widespread species which are recorded fromboth provinces and are therefore diagnostic (typi-cal) of the entire Western Tethys Region are:Tectus lucasianus (Brongniart), Campanile char-pentieri (Basterot), Granulolabium plicatum (Bru-guie're), Tympanotonos stroppus (Brongniart), Ce-rithium ighinai (Michelotti), Turritella magnaspe-rula Sacco, Turritella asperula Brongniart, Turri-tella conofasciata (Sacco), Peyrotia desmarestinaBasterot, Peyrotia strangulata (Grateloup), Strom-bus radix (Brongniart), Cassis mamillaris Grate-loup, Cassis retusa Michelotti, Cassis nummuliti-phila (Sacco), Sconsia beyrichi (Michelotti),Proadusta parvitala (Sacco), Globularia gibberosa(Grateloup), Ampullinopsis crassatina (Lamarck),Amaurellina oweni (Archiac and Haime), Sinumaquensis (Recluz), Xenophora cumulans (Brong-niart), Ficus condita (Brongniart), Turbinella epi-soma (Michelotti), Conus diversiformis Deshayes,Conus carcarensis Sacco, Lyria anceps (Michelot-ti), and Terebra subtessellata d’Orbigny. Alongwith these gastropods the remarkable bivalve Avi-cularium carinatum (Bronn) may be regarded as acharacteristic element of the Western Tethys Re-gion.

The southeasternmost known gastropod faunaof the MediterraneanÎranian Province occurs inthe Late Rupelian and Early Chattian of the Es-fahan^Sirjan Basin in Central Iran. The Oligocenesection of Lordegan in the Iranian Zagros Moun-tains, located still further south, bears a strombidof the MediterraneanÎranian Province. In con-trast, the northwesternmost fauna of the WesternIndianÊastern African Province is actually repre-sented by the Pakistanian assemblages. Therefore

the boundary between these biogeographic areasalong the northern Tethys shores is located some-where between Central Iran and the Sind^Balu-chistan Region. Along the southern shores, thefaunas of Libya, Palestine and Egypt are consid-ered part of the MediterraneanÎranian Province(Fig. 3(left, bottom)), while the faunas of Omanrepresent the northern known extension of theWestern IndianÊastern African Province (Fig.3(left, bottom)). Similarities between the twoprovinces are high within the potamidids, turri-tellids and naticids but low amongst the volutids,turrids and other neogastropods, documenting thepeculiar character of the fauna. Only very few ofthese species, such as Ampullinopsis crassatina,Globularia gibberosa or Granulolabium plicatummanaged to colonise adjacent bioprovinces inhigh numbers. Others such as Turbinella episoma,Cassis mamillaris or Angaria scobina, which arevery common in Western Tethyan assemblages,are rare faunal elements within their northern-most distribution area in the Early Oligocene ofthe Paratethys. The connection between these bi-ogeographic entities was supported by a distincteasterly current along the northern shores of theTethys (Fig. 3(left, bottom); Barron and Peterson,1991). According to Barron and Peterson (1991) awestward-directed Eocene to Oligocene Tethyancurrent system along the northern shores as sug-gested by Berggren and Hollister (1974) has to berejected.

A third province in the western part might bebased on di¡erences in the composition of theOligocene faunas from southwestern France(Grateloup, 1840; d’Orbigny, 1852; Lozouet,1998, 1999) and the Italian and Greek localities.This is additionally supported by the northernin£uence in the French faunas as suggested byWelle (1998) and by numerous muricids, bucci-nids, nassariidis, or conids described only fromFrance (Lozouet, 1999).

Early Neogene : During the Early Miocene theWestern Tethys Region shrank drastically fromthe gastropods’ point of view. Still, the areafrom the Bay of Biscay to the Mediterranean ba-sins as far east as Central Iran display a rathersimilar fauna (Fig. 4). However, the Early Mio-cene brings about a drastic change for the West-



ern IndianÊastern African Province, which van-ishes as a distinct unit probably even before theclosure of the Tethyan corridor in the Late Bur-digalian. The data on Pakistanian molluscs fromthe Early Miocene Gaj Formation given by Vre-denburg (1925^1928) re£ect a strongly increasingin£uence by the fauna of South-East Asia, and arapid decrease of elements of the Western TethysRegion. Correspondingly, Azzaroli (1958) noticedin the Miocene faunas of Somalia some in£uencefrom Indonesia, although his fauna still has somespecies in common with Italian localities. Hence,the separation of the gastropod faunas of the east-ern African and the western Indian coastal areasfrom those of the Western Tethys Region alreadytook place during the Early Miocene (Burdigali-an?). Since the faunas of this area have a clear‘Indo-Paci¢c’ character and bear high percentagesof Indonesian species, they may be treated aswestern part of an early stage of the Indo-WestPaci¢c Region, which should be termed the Proto-Western Indian Ocean Province.

With the disconnection of the Indo-Africanfauna and the take-over by elements of the Pro-to-Indo-West Paci¢c Region in the area of theformer Western IndianÊastern African Provinceduring the Early Miocene the MediterraneanÎra-nian Province lost its southeastern counterpart.Finally, the point-of-no-return was reached bythe closure of the Eastern Mediterranean seawaybetween the Anatolian and the Arabian/Africanplates during the Late Burdigalian (Jones, 1999).Based on the dramatic geodynamic and biogeo-graphic changes and especially due to the evolu-tion of the advanced Miocene Mediterranean gas-tropod fauna, the term Western Tethys Regionseems hardly acceptable for the Early Mioceneat least for the Burdigalian (see also Wester-mann’s rules for replacement of names, Wester-mann, 2000a). Therefore, we propose the termProto-MediterraneanÂtlantic Region (Fig. 3(right, top and bottom)) for the descendant ofthe strongly reduced Paleogene Western TethysRegion.

4.2.1.2. Proto-MediterraneanÂtlantic RegionAs discussed above a marine faunal exchange

between the southeastern and the ‘Mediterranean’

faunas was strongly prevented during the EarlyMiocene. Therefore in the area of the RecentMediterranean Sea a biogeographic unit formedwhich might best be named Proto-Mediterra-neanÂtlantic Region (Fig. 3(right, top)). Accord-ing to the palaeogeographic situation as drawn byRo«gl (1998) this area roughly corresponds to thelimits of the modern MediterraneanÂtlantic Re-gion except for the Black Sea Province, which wasnot in existence. This area was part of the Para-tethys at that time.

The southwestern extension of the Proto-Medi-terraneanÂtlantic Region is not well de¢ned dueto the lack of available data on shallow marinefaunas along the shores of northwestern Africa.However, the limits along the European Atlanticcoast in northern direction were more or lessequivalent to those of the Recent Mediterra-neanÂtlantic Region (sensu Briggs, 1995). TheArtois axis formed a landbridge, which repeatedlyseparated the fauna from that of the North SeaBasin (Pomerol, 1982; Ziegler, 1990).

Unequivocal evidence for direct marine connec-tions to adjacent provinces exists for the CentralParatethys, which is strongly in£uenced by thefauna of the Proto-MediterraneanÂtlantic Regionduring that time (Ro«gl, 1998, 1999; Harzhauser,2002).

Within this area the data on gastropod faunasfrom France (Cossmann and Peyrot, 1917^1922),Italy (Bellardi, 1872^1904) and Iran (Harzhauser,2000) show good correspondence during the Bur-digalian, although a slight northwest-southeastshift in the composition is evident. However, theeastern Mediterranean faunas have 80% of speciesin common with Northern Italy and France, thusindicating a rather homogeneous distribution oftaxa within this area during the Burdigalian(Fig. 4d). A slight drop in resemblance is recordedin the Iranian fauna, which still contains 66% (38species) of ‘Mediterranean’ species (Fig. 4d). Toobtain a better understanding of the similaritiesand di¡erences between the Iranian and the Med-iterranean faunas it is necessary to focus on thefamily level, because the degree of resemblancestrongly £uctuates in the various families. In gen-eral the melongenids, cerithiids, strombids, and tosome extent also the naticids show highest a⁄n-



ities to the Mediterranean faunas. In contrast, theconids and the turrids display a considerablenumber of taxa which are known only from theQom Basin (10^15%); due to the poor informa-tion it is unclear whether these species are alsodistributed in the Indo-Paci¢c or if they representendemic forms, thus hinting at a distinct Iraniansubprovince. Some of the most characteristic taxawithin the Proto-MediterraneanÂtlantic Regionare: Turritella (Turritella) terebralis Lamarck,Protoma quadriplicata (Basterot), Peyrotia des-marestina (Basterot), Protoma cathedralis (Brong-niart), Terebralia bidentata (Defrance), Cerithiumvulgatum Bruguie're, Granulolabium bicinctum(Brocchi), Granulolabium plicatum (Bruguie're), Ti-bia dentata (Grateloup), Strombus (Lentigo) bo-nelli (Brongniart), Xenophora deshayesi (Michelot-ti), Sconsia striata miocenica Sacco, Melongenalainei (Basterot), Tudicla rusticula (Basterot), Ath-leta ¢culina (Lamarck), and Conus (Conolithus)dujardini Deshayes.

Aside from this core-area, the Danubian Prov-ince (see below) ^ spanning the western and east-ern part of the Paratethys Sea ^ is considered torepresent a distinct province of the Proto-Medi-terraneanÂtlantic Region at least during the lateEarly Miocene and the early Middle Miocene (seebelow).

This palaeogeographic framework of the Proto-MediterraneanÂtlantic Region based on gastro-pod data corresponds largely to the ‘Mediterra-nean Province’ as introduced by Jones (1999)based on larger benthic foraminifera. His prov-ince unites the entire Mediterranean area, theQom Basin, and the Atlantic coasts of the Aqui-taine Basin and of West Africa. Similarly, Ro«gland Steininger (1983, 1984) recognised this bio-geographic entity and called it the ‘Mediterranean(or Tethyan) bioprovince’.

The mollusc fauna of the Proto-MediterraneanÂtlantic Region su¡ered two major incisions. The¢rst one took place in the early Middle Miocenewhen a short-lived connection to the Indo-Paci¢c^ coinciding with a marked global warming (Mil-ler et al., 1991; Ro«gl, 1998; Nishimura and Supar-ka, 1997; Tsuchi, 1997) ^ gave rise to a highlydiverse Middle Miocene mollusc fauna. Untilnow the role of immigration of Indo-West Paci¢c

gastropods during the early Middle Miocene issomewhat dubious. Instead of the usually advo-cated immigrations, the autochthonous evolutionin a phase of climatic optimum seems to be un-derrated. Anyway, the short-lived connections ofthe Proto-MediterraneanÂtlantic Region to theIndo-West Paci¢c Region did not change the equi-librium gastropod faunas of the Proto-Mediterra-neanÂtlantic Region markedly. During the Mio-cene and Pliocene various reconnections arediscussed (Grecchi, 1978; Lozouet, 1992; Ro«gl,1998; Jones, 1999), however, the biogeographicboundaries between the Indo-Paci¢c and ‘Medi-terranean’ remained stable.

A second impact on Proto-MediterraneanÂt-lantic mollusc faunas was the Messinian salinitycrisis in the Late Miocene, when the marine faunaof the Mediterranean basins was strongly impov-erished (Hsu« et al., 1978). The gastropod fauna ofthe Mediterranean basins probably vanished com-pletely ^ except for some supposed shelters suchas the ‘Andalusian sanctuary’ (see Demarcq,1987) ^ due to desiccation throughout the Medi-terranean. The fauna of the Eastern Atlanticcoasts was less a¡ected. After the salinity crisisthe deep-water fauna was distinctly impoverished(Ra⁄ and Taviani, 1984), but it had not such acatastrophic e¡ect on shallow marine molluscs(Sabelli and Taviani, 1984), which recolonisedthe Mediterranean basins from the Atlantic.Nevertheless, the Messinian salinity crises causeda turnover in composition of the nearshore fauna.Post-Messinian gastropod faunas lack character-istic Miocene taxa, such as Terebralia bidentata(Defrance), Melongena cornuta (Agassiz), Polini-ces redempta (Michelotti) or Perrona jouanetti(Desmoulins). In fact, many of the conids, turridsand turritellids of the Proto-MediterraneanÂtlan-tic Region became extinct (cf. Dermitzakis andGeorgiades-Dikeoulia, 1987). Hence, the post-Messinian gastropod fauna represents rather theancestral, warm-water stock of the modern Med-iterraneanÂtlantic Region.

During the Late Pliocene and Pleistocene thechange from the Proto-MediterraneanÂtlanticRegion to the modern MediterraneanÂtlantic Re-gion was accelerated by distinct cooling events(Ro«gl and Steininger, 1983). Consequently, trop-



ical/subtropical taxa which contribute to the fau-na of the Proto-MediterraneanÂtlantic Regionare missing in the modern region (e.g. Olividaeas demonstrated by Davoli, 1989, or Strombidaeas emphasised by Abbott, 1960).

4.3. Biogeographic position of the Paratethys Sea

The Paratethys, as a distinct palaeogeographicand palaeobiogeographic unit, formed around theEocene/Oligocene boundary (Fig. 2) and lasteduntil the Pliocene. In fact, even the modern Araland Caspian provinces are a heritage of the hugeParatethys, based on its peculiar fauna and en-demism. During its maximum extent the Parate-thys Sea spread from the Rhone Basin in Francetowards Inner Asia. Its geographic subdivisionsare the Western Paratethys, comprising the RhoneBasin and the Molasse Basin of Switzerland andwestern Bavaria, the Central Paratethys, reachingfrom Bavaria in the west to the CarpathianMountains in the east, and the Eastern Parate-thys. Recently, the stratigraphy and geodynamicsof the latter area were studied by Popov et al.(1993) and Jones and Simmons (1996). A criticalup-to-date overview on palaeogeographic datahas been given and discussed by Ro«gl (1998,1999) and summarised in several palaeogeograph-ic sketches.

This region underwent an outstanding historyof total or partial isolation, re£ected in a highpercentage of endemics, alternating with variousconnections to the adjacent northern and/orsouthern seas.

The term Paratethys was introduced as early as1924 by Laskarev (1924). During the last decadesand especially due to the edition of the ‘Chrono-stratigraphie und Neostratotypen’ volumes thepalaeobiotic framework was distinctly improved(Cicha et al., 1967; Steininger and Senes, 1971;Baldi and Senes, 1975; Papp et al., 1973, 1974,1978, 1985; Stevanovic et al., 1990). Its geody-namic history was described by Senes and Ma-rinescu (1974) and Rusu (1988), who divided thegeodynamic development of the Paratethys intofour stages: the Proto-Paratethys formed in theLate Eocene to Early Oligocene, caused by ¢rstisolation from the open oceans. This stage is fol-

lowed by the Late Oligocene and Early MioceneEo-Paratethys, the late Early Miocene to earlyMiddle Miocene Meso-Paratethys and the Middleto Late Miocene Neo-Paratethys (see also Stei-ninger and Wessely, 2000).

To avoid an intermingling of geographic andbiogeographic terms, we reject the terms Western,Central and Eastern Paratethys for biogeographicuse in this paper. Instead we propose the newterms Danubian Province and Proto-Caspian Sub-province (Fig. 3(left, top; right, top and bottom)).Hence, the Danubian Province is characterised bythe total fauna of the Western, Central, and East-ern Paratethys. The peculiar faunistic inventory ofthe Eastern Paratethys allows a further seggrega-tion of a Proto-Caspian Subprovince. (A furtherdi¡erentiation of these biogeographic units anda detailed description of their evolution and ex-pansion will be published elsewhere.)

4.3.1. Danubian Province and Proto-CaspianSubprovince

The ¢rst endemic Paratethys fauna developedduring the Early Oligocene Solenovian, when the¢rst nearly complete closure of the Paratethys oc-curred. This event peaked in the blooming andrapid evolution of a highly endemic brackishwater bivalve fauna with genera such as Janschi-nella, Korobkoviella or Ergenica (Voronina andPopov, 1984; Popov et al., 1985; Nevesskaja etal., 1987; Ro«gl, 1998). By this event the Parate-thys was united in a single, very distinct biogeo-graphic entity ^ the Proto-Caspian Subprovince âs far as bivalves are concerned (Popov et al.,1993). Nonetheless, as shown by Popov et al.(1985) and Popov and Titova (1982), this endem-ism is much lower within the gastropods. Brackishor estuarine gastropods which are at that timewidespread in the entire Western Tethys Regioninhabited the coasts of the Paratethys; Melanopsisimpressa Krauss, Granulolabium plicatum (Bru-guie're), Theodoxus crenulatus (Klein), and Tympa-notonos margaritaceus (Brocchi) give evidence ofreduced marine conditions within this interval,but do not justify a separation of a biogeographicunit at the level of a region.

Aside from this short-lived extreme western ex-tension of the Proto-Caspian Subprovince, the



western part of the Paratethys was characterisedby the quite di¡erent fauna of the Danubian Prov-ince (Fig. 3(left, bottom)). In times of open sea-ways, the Early Oligocene Danubian Province con-sists of northern species accompanied by asurprisingly large number of Western Tethyan el-ements, giving evidence of the earliest stage ofseparation of these biogeographic entities.Amongst these the most characteristic WesternTethyan species are Turritella conofasciata Sacco,Turritella asperula Brongniart, Ficus oligo¢coidesSacco, Athleta italica Fuchs, Turbinella episoma(Michelotti), Cassis mamillaris Grateloup, Anga-ria scobina (Brongniart), and Babylonia caronisBrongniart (Baldi, 1986). Thus the later Early Oli-gocene, as documented by Baldi (1986) for thegastropod faunas of the Hungarian Kiscell Clay,displays about 31^35% endemic gastropods butalso more than 20% Western Tethyan taxa andapproximately 16% northern species.

During the Late Oligocene Egerian the south-ern £air decreased. The Danubian Province hosts amixture of northern elements, which made theirway from the North Sea Basin during the Rupe-lian, such as Cassidaria megacephala (Philippi),Drepanocheilus (Arrhoges) speciosus (Schlotheim),Ficus concinnus (Beyrich), ‘cosmopolitan’ Tethyanspecies such as Granulolabium plicatum (Bru-guie're), Tympanotonos margaritaceus (Brocchi)and Melanopsis impresssa Krauss, and a fair num-ber of endemics, such as Melongena incornuta(Ho«lzl), Bullia hungarica (Ga¤bor), Chicoreus tri-gonalis (Ga¤bor), Pisanella doboi (Noszky), andEgerea collectiva (Ga¤bor). These considerablenumber of endemics grow distinctly smaller dur-ing the Early Miocene Eggenburgian, when fau-nistic ingressions from the west and south led to aturnover in composition.

The heavy in£ux of species from the northernbioprovince ^ herein called the Proto-Eastern At-lantic^Boreal Region (Fig. 3(left, bottom; right,top and bottom)) ^ characterises also the Oligo-cene gastropod fauna of the Proto-Caspian Sub-province. Northern elements such as Drepanochei-lus (Arrhoges) speciosa, Cassidaria buchi/depressa,Ficus crassistria, Turritella planispira, Cancellariaevulsa, and Typhis pungens reach the Aral andCaucasus basins (Oveckin, 1956; Alizade, 1968),

and northern turrids such as Orthosurcula regula-ris contribute to the fauna of the Ustjurt area(Amitrov, 1973). Endemism, however, is muchstronger in the Proto-Caspian Subprovincethroughout the Oligocene.

The second phase of a rather uniform Parate-thys fauna, coinciding with the westward exten-sion of the Proto-Caspian Subprovince, developedduring the Early Miocene Kotsakhurian (corre-sponding to the Ottnangian). Again, the separa-tion of the Eastern Paratethyan Sea coincidingwith brackish water conditions is followed by asudden evolutionary peak in bivalves, resulting ina large number of endemic genera such as Limno-pagetia, Rzehakia, Lenticorbula, or Eoprosodacna(Kvaliashvili, 1962; Voronina and Popov, 1985).This peculiar, so-called ‘Rzehakia fauna’ reacheseven the Western and Central Paratethys Sea inthe Late Ottnangian. Corresponding to the ¢rstisolation stage in the Early Oligocene, the EarlyMiocene Proto-Caspian Subprovince is more dis-tinctly characterised by the bivalves than by thegastropod fauna (see also Ctyroky¤, 1972).

A complete disconnection of the Western/Cen-tral Paratethys and Eastern Paratethys began inthe latest Early Miocene (Fig. 3(right, bottom))and early Middle Miocene (Nevesskaja et al.,1987; Ro«gl, 1998). Good marine connections ofthe Western/Central Paratethys with the southernsea allowed immigration of ‘MediterraneanÂt-lantic’ gastropods. At times of high immigrationrates such as the late Early Miocene Karpatian orthe early Middle Miocene Badenian the ‘Mediter-ranean’ character became overwhelming. TheKarpatian gastropod fauna of Austria (Harzhau-ser, 2002) has only 25^30% Western Paratethysendemics ^ a percentage which is probably eventoo high, since several of the ‘endemic’ species areprobably hidden in poorly described Mediterra-nean species groups. Anyway, the remaining 70^75% of these latest Early Miocene gastropods aretypical inhabitants of the Proto-MediterraneanÂtlantic Region. In respect to biogeography thistake-over by ‘MediterraneanÂtlantic’ gastropodswithin the Danubian Province and the low endem-ism requires an integration of the Danubian Prov-ince into the Proto-MediterraneanÂtlantic Regionwhich thus consists of the MediterraneanÎranian



Province at its core and the Danubian Province.This development was reversed during the lateMiddle Miocene Sarmatian when the isolationof the Central Paratethys from the adjacent seasmay have caused a dramatic change of the waterchemistry (Pisera, 1996) leading to a severe im-poverishment of the marine fauna. Simultaneous-ly, its reconnection with the Eastern Paratethysallowed the establishment of a rather uniformSarmatian brackish gastropod fauna, being typi-cal for the entire Paratethys Sea. During the latestMiddle Miocene and earliest Late Miocene themarine cycle ended within the Paratethys and abrackish to freshwater lake system became in-stalled.

4.4. Transatlantic communication ^ witnessing anextensive Tethys Realm?

The Caribbean and adjacent tropical areas dis-play a characteristic faunistic composition at leastfrom the Late Cretaceous onwards (Kau¡mann,1973). During the Paleogene this is expressed inthe formation of a distinct large Central Americanbiogeographic unit, which roughly covers thegeographic area of the modern Eastern Paci¢cand the Western Atlantic regions sensu Briggs(1995). However, within the Central Americanbioprovinces a surprising £air of familiarity withthe Western Tethys Region is perceptible from theEocene up to the Miocene.

Especially during the Paleogene, the Tethyanheritage is obvious, since genera such as Ampulli-nopsis, Globularia, Terebralia, or Strombus indi-cate immigration from the Tethyan bioprovince.The faunistic exchange between the amphiatlanticbioprovinces was a two-way story for sure, butdue to the gaps in the fossil record, it is oftenunclear whether a taxon migrated from the OldWorld to the New World or vice versa. Althoughthe amphiatlantic relations were discussed severaltimes in the literature (e.g. Maury, 1902; Cooke,1924; Gardner, 1924; Woodring, 1924a,b, 1928;Kautsky, 1925; Vokes, 1964^1986; Savazzi, 1989;Lozouet et al., 1994), this important migrationpattern is yet hardly embodied in palaeobiogeo-graphic concepts.

Especially during the Eocene some major mi-

grations across the Atlantic took place. Amongstthe naticids the genera Ampullinopsis and Globu-laria, which both can be traced back to the West-ern Tethyan Eocene, reached the Americas duringthe Eocene. Ampullinopsis and the archaeogastro-pod Velates may be regarded as some of the mostsuccessful Tethyan gastropods during the Paleo-gene, since they are recorded from the easternPaci¢c in the west to as far east as Pakistan orBurma. Tethyan a⁄nities in the Eocene with theAmericas are documented by a large number oftaxa, usually on the generic level and rarely onspecies level. Palmer (1967) united some of thosetaxa of proposed Tethyan origin, e.g. the generaVelates, Eovasum, Gisortia, and Terebellum, in theso-called ‘Velates perversus Group’. This small listcan be easily enlarged by data of Clarke andVokes (1936), Givens (1978, 1989), Jung (1974),and Squires and Advocate (1986) to include thegenera Agaronia, Akera, Amaurellina, Ampullella,Ampullinopsis, Athleta, Buccinorbis, Calyptrapho-rus, Campanilopa, Chedevillia, Clavilithes, Cornu-lina, Crommium, Cryptochorda, Caricella, Ectino-chilus, Eocithara, Eocypraea, Eopleurotoma, Ga-leodea, Gilbertina, Hipponix, Keilostoma, Laevity-phis, Lyria, Mazzalina, Muricopsis, Paraseraphs,Pleurofusia, Sassia, Semicassis, Seraphs, Strepsi-dura, Streptochetus, Surculites and Volutilithes â list which is far from being complete. AnotherMiddle Eocene similarity is documented by theendemic American Dirocerithium, which showsclosest resemblance with the Tethyan genus Bella-tara hinting at a common Tethyan ancestry(Woodring, 1959).

An excellent example for the intensive transat-lantic relationship during the Paleogene are thestrombids. The family arises during the Eocenewhen it develops a fair number of new taxasuch as Dilatilabrum, Orthaulax and Oostrombus.The genus Oostrombus, well established in theTethyan Eocene by species such as Oostrombusauriculatus, displays one of the most striking dis-junct patterns. The genus has been recorded byOlsson (1931) as Oostrombus chiraensis from theLate Eocene Chira Formation of Peru, by Clarkeand Durham (1946) as Oostrombus cedroensisfrom the Eocene of Colombia, and by Woodring(1959) as Oostrombus a¡. chiraensis from the Mid-



dle or Late Eocene Gatuncillo Formation of Pan-ama. From the Eocene onwards, the New WorldÔld World route can also be documented for thetwo seemingly typical Western Tethyan generaTurbinella and Vasum. Turbinella ¢rst appearedin the Late Eocene of Peru with Turbinella peruvi-ana (Olsson). Thereafter, two lineages developedseparately in the New and the Old World (Vokes,1964). The latter is rooted in the earliest OldWorld species Turbinella episoma (Michelotti),which appears in the Western Tethyan Oligocene.Similarly, the genus Vasum developed in the NewWorld where it is recorded from the Eocene ofLouisiana. During the Oligocene the genus man-aged to pass the Atlantic barrier and, with Vasumsubpugillare (d’Orbigny), is documented from theOligocene of France. According to Vokes (1966)all Early Miocene Old World representatives ofVasum are very close to the ancestral EoceneNew World species Vasum humerosum Vaughan.Correspondingly, the genus Deromurex appears inthe Early Oligocene of the Mississippi Region,whilst its European counterpart is known onlyfrom the Late Oligocene of western France(Vokes, 1975a). The vicariating species pair is rep-resented by the New World Deromurex cookei(Vokes) and the Old World Deromurex cotteavi(Meunier).

The opposite route was taken by some westernAtlantic species of Cheilea. According to Vokes(1975b) the New World species Cheilea equestris(L.), which is ¢rst known from the Chipola For-mation (NW Florida), is a descendent of the Eo-cene Old World Cheila boutillieri (Cossman). Cor-respondingly, Cheila uncinata (Reeve) can betraced to the European Eocene where Cheila ber-nayi (Cossman) is considered to be the ancestralform. Another amphiatlantic connection of astrombid taxon has been documented by Savazzi(1989) for the genus Orthaulax. It ¢rst appearedwith Orthaulax dainelli in the early Middle Eoceneof northern Italy but thereafter was ubiquitous inthe Oligocene and Early Miocene of CentralAmerica. Correspondingly, the genus Strombusappeared in Tethyan Paleogene and crossed theAtlantic barrier during the Oligocene. Noticethat, according to Woodring (1959), the genus ^formerly mentioned from Eocene sediments in

Alabama as Strombus albirupianus by Dall(1890^1903) ^ has been only documented fromthe Late Oligocene of Georgia!

Correspondingly, the Western Atlantic repre-sentatives of Aspella originated in the EuropeanOligocene (Vokes, 1975a) and ‘Cerithium’ halenseof Dall (1917) has its closest relatives amongst theTethyan Oligocene potamidids usually assigned to‘Telescopium’ charpentieri or Tympanotonos strop-pus. This Oligocene migration from the Tethysinto the tropical Americas is obviously not re-stricted to the gastropods but can also be docu-mented for other molluscs. For example, thestrange bivalve Kuphus is very common in theentire Mediterranean Oligocene and gained afoothold in the Mid-American area at least inthe Late Oligocene (Mans¢eld, 1940). A cosmo-politan transatlantic character has been also re-ported for reef coral assemblages of the EoceneÔligocene (Budd, 2000).

Beside these Eocene/Oligocene migrations, a re-markable and well-documented cross-Atlantic mi-gration took place during the Burdigalian andprobably early Middle Miocene. As far as canbe judged from the fossil record the main routewas from the Western Tethyan Region to theWestern Atlantic, since many of the amphiatlantictaxa have their earliest record in the EuropeanCenozoic.

The most convincing evidence for those migra-tions are amphiatlantic occurrences at the specieslevel. Some of these species have been recognisedin the Chipola Formation of northwestern Flori-da. E.g., Eudolium (Galeodolium) subfasciatumSacco appears in the Burdigalian and early Mid-dle Miocene of the Mediterranean area and in theEarly Badenian of the Western Paratethys. In theBurdigalian the species also appears in the Can-taure Formation of Venezuela and Chipola For-mation of northwestern Florida (Vokes, 1986;Gibson-Smith and Gibson-Smith, 1988). Others,such as Lindapterys, are documented by highlysimilar species at both sides of the Atlantic (Lo-zouet et al., 1994). Similarly, Calyptraea (Trochi-ta) ornata Basterot and Calyptraea (Trochita) co-staria Grateloup have been recorded from theChipola Formation and the European Aquitanianand Burdigalian (Vokes, 1975b). High similarities



between European Atlantic Burdigalian faunasand the Chipola fauna was also recognised byDolin (1991) within the cypraeids. Trona leporinacalhounensis Dolin has its origin in the Old Worldand emigrated to the Western Atlantic coast. Thesame has been suggested by Dolin (1991) for Tal-paria dominiciensis (Gabb) and Mauritia camp-belliana (Pilsbry).

However, the New WorldÔld World route isalso documented. Following Dolin (1991) theBurdigalian Chipola species Siphocypraea chilonaDall seems to derive from a Caribbean stock andentered the Old World in the Early Miocene. Sim-ilarly, a migration from west to east is also sup-ported by Bernasconi and Robba (1982) based onpteropods. According to Bernasconi and Robba(1982) the faunistic interchange of pteropods be-tween the Caribbean and the Western Tethyanarea occurred mainly from the Oligocene up tothe Middle Miocene.

Less obvious at ¢rst sight, but very importantas indicators of transatlantic faunal relations aresome pairs that probably represent vicariatingspecies. Worth mentioning and recently studiedare pairs such as Bursa (Bufronariella) pelouatensis(Cossmann and Peyrot) from the Early Mioceneof the Aquitaine Basin and Bursa (Bufronariella)chipolana Schmelz from the Burdigalian of Flori-da (Schmelz, 1997). The similarities are so strikingthat Vokes (1973) erroneously even considered theFlorida species to be synonymous with the OldWorld gastropod. Vokes (1965a,b) also discussedthe high similarities between the Florida and theEuropean representatives of Chicoreus s.s. duringthe Late Burdigalian and the early Middle Mio-cene. According to her, the genus originated inthe Tethyan area, having its roots in Eocene OldWorld muricids, such as ‘Murex’ tricarinatus La-marck. Vicariating or at least very similar am-phiatlantic species pairs are Chicoreus folidodes(Gardner), Chicoreus lepidotus (Vokes) and Chi-coreus dujardinoides (Vokes) from the New Worldand Chicoreus aquitanicus (Grateloup), Chicoreusbourgeoisi (Tournouer) and Chicoreus dujardini(Tournouer) from the early Middle Miocene ofEurope. Similarly, Vokes (1974) mentioned Ptero-typhis wenzelidesi from the Badenian stage (earlyMiddle Miocene) of the Vienna Basin and Ptero-

typhis vokesae Gertman from the Chipola Forma-tion as closely related species. Two further specieswith a European ancestry are mentioned byVokes (1966, 1979) from the Early Miocene (Bur-digalian?) of the Dominican Republic. Especiallythe Burdigalian Vasum tuberculatum Gabb canhardly be distinguished from the Aquitanian Va-sum aquitanicum Peyrot from France and mighteven be conspeci¢c (Vokes, 1966). Similarly, Va-sum pugnus Pilsbry and Johnson seems to be adescendent of the French Vasum stephanensePeyrot, although a close relationship as discussedby Vokes (1966, 1979) seems to be unlikely due tothe large stratigraphic gap between the Late Oli-gocene French occurrence and the alleged Burdi-galian occurrence in the Americas (Vasum stepha-nense from St.-Etienne-d’Orthe is, according toLozouet (1998), of Chattian age, whilst Peyrot(1928) formerly assigned the species to the Aqui-tanian).

The latest but subordinate Miocene transatlan-tic migration seems to be documented for the LateMiocene. A remarkable example of this event isthe genus Vittularia, which is an Old World ge-nus, appearing in the Oligocene of France. Duringthe Early and Middle Miocene the genus, repre-sented by Vittularia linguabovis (Basterot), be-comes ubiquitous in the Mediterranean as wellas in the Paratethys. In the New World it ¢rstappears during the Late Miocene (Vokes, 1967,1977). Therefore, if a transatlantic migration isconsidered, some minor, occasional faunal ex-change occurred even as late as the Late Miocene.Generally, the Miocene transatlantic exchange ofbiota is still insu⁄ciently studied. For reef corals,for example, dispersal across the Atlantic fromthe Mediterranean to the Caribbean is consideredto have ceased at the end of the Oligocene (Frost,1977; Budd, 2000). Contrary, Riegl and Piller (inpress) reported the genus Mussismilia, which wasrecorded from the Late Miocene as a mere Car-ibbean element (Budd, 2000), from the MiddleMiocene of the Styrian Basin (Danubian Prov-ince).

However, the similarities between the WesternAtlantic and the European faunas are too poor atany time to allow a comparison on a hierarchylower than a biogeographic region. The Miocene



fauna of the ‘central’ part of this western Regionhas been very well studied, allowing a chie£y gas-tropod-based conception of the Miocene Mid-American bioprovinces as proposed by Woodring(1974). He de¢nes a Miocene Caribbean Region( = province sensu Woodring, 1974), which ischaracterised by the bivalve Clementia darienna(Woodring) and a large number of gastropods,such as Orthaulax, Strombina, Metula, Potamidessuprasulcatus, and Stigmaulax guppiana. This bio-province falls apart into the Mexican, West Indi-an, Central American^northern South American,Columbian^Venezuelan^Trinidad, Brazilian, andEcuadorian^Peruvian provinces ( = subprovincessensu Woodring, 1974). It is worth mentioningthat Woodring’s Brazilian Province is basedon the absence of taxa rather than on endem-isms.

Despite the amphiatlantic similarities, whichdocument some communication between two ma-jor biogeographic units, the Atlantic was a barrierfor many Tethyan genera, such as Pyrazus, Tibias.s., Sulcogladius, Tudicla, and Amalda. With re-spect to the large total gastropod faunas the per-centage of shared species is rather small and re-quires a distinct separation of both biogeographicunits on the level of a region. Many of theamphiatlantic taxa are only erratics within anotherwise independent Western Atlantic Region.Therefore, most authors studying the amphiatlan-tic distribution of tropical/subtropical molluscs inthe Cenozoic considered a more or less occasionaland accidental rafting of larvae as most probablemode of transportation, representing sweepstakesroutes in terms of modern biogeography. How-ever, there seem to be some distinct peaks of sim-ilarity between Western Tethyan and Americangastropod faunas, e.g. during the Middle Eoceneor the Burdigalian. Such peaks, if not an artefactdue to lack of investigations or adequate faunasin the intervening intervals, might indicatechanges in the Atlantic current systems, whichrepeatedly supported or suppressed the migrationof larvae. Correspondingly, the amphiatlanticsimilarities are explained by Berggren and Hollis-ter (1974) and Barron and Peterson (1991) by anearly gyre-like Gulf Stream with limited polewardextension. According to the simulations of Barron

and Peterson (1991) this early Gulf Stream in theNorth Atlantic is evident even during the Eoceneand Oligocene. In contrast, a ‘nearshore-hopping’along the northern coasts as documented forsome species in the Pleistocene can be excludedfor all the amphiatlantic Eocene to Miocenetaxa due to the strict climatic requirements ofthe warm-water faunas. Additionally, both fau-nas, especially the Mediterranean one, obviouslyreached a stage of equilibrium during the Miocene(Demarcq, 1987). Therefore, most of the intrudersprobably had little chance to gain a foothold.Despite the broad marine connections all thesetaxa rarely made their way in eastern directioninto the Paci¢c. The distribution via the Atlanticstrait is the only satisfying explanation, since evi-dence for a Paci¢c route is missing. The transat-lantic migration route during the Eocene was alsoemphasised by Clarke and Vokes (1936), whostated that the similarities between the faunas ofthe American west coast and those of India arelow, thus excluding an arrival via a Paci¢c migra-tion route, which was even at that time probablyobstructed by the East Paci¢c Barrier.

5. Conclusions

Several phases of major transatlantic faunal ex-change can be deduced from the gastropod dataon both sides. The ¢rst phase comprises the Eo-cene to Oligocene with an Eocene peak. It couldprobably be separated in several distinct migra-tion waves, if stratigraphic resolution would bebetter. A ¢rst westward migration wave is docu-mented for Early Eocene or even Late Paleocenetimes (Squires and Advocate, 1986). The secondphase spans the Burdigalian and the early MiddleMiocene. This phase is very well documented andagain might be composed of two minor waves.The ¢rst being characterised by migrations fromthe Old World into the Western Atlantic coastalfaunas mainly during the Burdigalian. The secondtook place somewhat later in the early MiddleMiocene, when American species or their closelyrelated forms suddenly appeared in European sec-tions. Finally, a third phase may be suggested forthe Late Miocene. However, data documenting



this phase are scanty and the impact on the fau-nas on both sides of the Atlantic was very low.

The western Atlantic coast ^ similar to theIndo-Paci¢c in late Early Miocene times for gen-era such as Tibia and Tudicla ^ obviously actedas refuge for several Tethyan genera such asAmpullinopsis and Orthaulax, which managed topass the Paleogene/Neogene boundary in theAmericas but became extinct in the Europeanbioprovinces.

Altogether the data furnish evidence for a cir-cum-equatorial, Paleogene Tethys Realm whichextended from Peru (Eastern Paci¢c!) throughthe early Caribbean Region across the Atlanticand the entire Mediterranean basins at least asfar east as Indonesia in the Eocene. During theOligocene this biogeographic unit still extendedfrom the Mid-Americas via Pakistan and easternAfrica to Indonesia with at least one centre oforigin in the Mediterranean basins. In contrast,an equivalent centre of origin is missing at thattime in the Proto-Indo-Polynesian Province.

The similarities of the Mediterranean, the Ira-nian, the Pakistanian and the eastern African gas-tropod faunas during the Oligocene requires tounite them in a Western Tethyan Region, whichotherwise can be split into a MediterraneanÎra-nian Province (Western and Eastern Mediterra-nean, Cyrenaica, Armenia, Libya, Syria, Pales-tine, Central Iran), and a Western IndianÊastern African Province (Oman, Pakistan, Soma-lia, Zanzibar, Kenya, Madagascar). A third prov-ince was probably established along the shores ofthe Eastern Atlantic (France).

The gastropod fauna of the Proto-Indo-WestPaci¢c Region developed in an area, which is sim-ilar to the geographic extension of the modernIndo-Polynesian Province but did not dominatethe Oligocene faunas of the Western IndianÊast-ern African Province. Therefore, the often advo-cated ‘Indo-Paci¢c’ in£uence on Oligocene andEarly Miocene Central European faunas seemsto be a myth. However, with the Early Miocenethe Indo-Paci¢c gastropod fauna established itselfwithin the area of the former Western IndianÊastern African Province and replaced the ‘Indo-European’ fauna completely.

The Paratethys Sea represents a distinct biogeo-

graphic entity ^ the Danubian Province. At least atthe subprovince level a separation of the easternProto-Caspian Subprovince is necessary. TheDanubian Province was always in£uenced byhigh immigration rates from the adjacent north-ern (Oligocene) and/or the southern (Oligocene^Miocene) bioprovinces. During phases of re-stricted marine connections and salinity crisesthe Danubian Province was repeatedly threatenedby the expansion of the Proto-Caspian Subprov-ince.

In the Early Miocene a rather homogeneousProto-MediterraneanÂtlantic Region formed(France, entire Mediterranean, Central Iran).During times of high emigration into the adjacentParatethys Sea, the Danubian Province has to betreated as part of the Proto-MediterraneanÂtlan-tic Region (latest Early Miocene through earlyMiddle Miocene). The biogeographic a⁄liationand hierarchical position of the Proto-CaspianSubprovince, however, is obscure with respect tothe poor data on gastropods from the EasternParatethys.

Generally, endemism in the Paratethyan gastro-pod fauna is low compared to that of bivalvesuntil the beginning isolation in the Sarmatian(late Middle Miocene). In contrast to bivalves,the composition of the gastropod fauna was oftenmassively in£uenced by immigrants rather than bylocal evolution.

Acknowledgements

The authors thank Fred Ro«gl and OrtwinSchultz (Museum of Natural History, Vienna)for fruitful discussions and for helpful commentson an earlier draft of this paper. We are alsograteful to Oleg Mandic (Institute for Palaeontol-ogy, Vienna) for information on bivalve distribu-tion. Our special thanks go to the members of theresearch projects on Late OligoceneÊarly Mio-cene circum-Mediterranean palaeobiological rela-tions, A. Kroh (Graz), F. Schuster (Tu«bingen),and U. Wielandt-Schuster (Frankfurt). This workwas supported by the Austrian FWF (P11886-GEO) and the Deutsche Forschungsgemeinschaft(STE 857/1-1).



Appendix 1

Observed occurrences of gastropod taxa instudied basins: 1 Greece ^ Mesohellenic Trough(Oligocene); 2: Iran ^ Esfahan^Sirjan Basin (Oli-gocene); 3: Iran ^ Esfahan^Sirjan Basin (Early

Miocene, Aquitanian); 4: Greece ^ MesohellenicTrough (Early Miocene, Aquitanian); 5: Iran ^Qom Basin (Early Miocene, Burdigalian); 6:Greece ^ Mesohellenic Trough (Early Miocene,Burdigalian); 7: Turkey ^ Mut Basin (Early Mio-cene, Burdigalian)

Gastropod taxa 1 2 3 4 5 6 7

Nerita (Theliostyla) plutonis Basterot o o oNerita (Theliostyla) asperata Dujardin oNerita caronis Brongniart oAgapilia picta (Fe'russac) o o o oTheodoxus? grateloupianus (Fe'russac) oGibbula buchi (Dubois) oGibbula a⁄nis (Eichwald) oClanculus (Clanculopsis) araonis (Basterot) oClanculus cerberi (Brongniart) oParoxystele amedei (Brongniart) o oJujubinus a¡. bucklandi (Basterot) oJujubinus a¡. subcarinatus (Lamarck) o oJujubinus sp. oAngaria scobina (Brongniart) o oTectus nov. sp. 1 oTectus? nov. sp. 2 oTectus sp. oTectus cf. rugosus (Grateloup) oTectus lucasianus (Brongniart) o oTurbo nov. sp. oTurbo cf. nov. sp. oTurbinidae Operculum indet. oLittorina? sp. oPeasiella cf. girondica Benoist oGreveniella mesohellenica Harzhauser and Kowalke oProtoma cathedralis (Basterot) o oProtoma quadriplicata (Basterot) o oPeyrotia tauroperturritus (Sacco) oPeyrotia strangulata (Grateloup) o o oPeyrotia desmarestina (Basterot) o o o oTurritella (Turritella) terebralis Lamarck o o oTurritella (Turritella) gradata Menke o oTurritella (Haustator) venus d’Orbigny o o oTurritella (Haustator) asperula Brongniart o oTurritella (Haustator) conofasciata (Sacco) oTurritella (Haustator) vermicularis Brocchi o oTurritella (Haustator) eryna d’Orbigny o oTurritella (Zaria) angulata Sowerby oTurritella cf. subtriplicata d’Orbigny oTenagodus cf. anguinus Linne' oMelanopsis impressa Krauss o oBellatara palaeochroma nov. ssp. oCampanile charpentieri (Basterot) oCampanile sp. oTerebralia bidentata lignitarum (Eichwald) o oTerebralia bidentata (Defrance) o o oTerebralia a¡. subcorrugata (d’Orbigny) o




Terebralia a¡. subcorrugata subinterrupta (d’Orbigny) oGranulolabium plicatum (Bruguiere) o o oGranulolabium (Tiaracerithium) pseudotiarella (d’Orbigny) oGranulolabium (Tiaracerithium) nov. sp. oGranulolabium (Tiaracerithium) cf. nov. sp. oGranulolabium pictum mitralis (Eichwald) oTympanotonos margaritacus (Brocchi) o oTympanotonos stroppus (Brongniart) o oPyrazus daemon (Oppenheim) oGourmya romeo (Bayan) oClava ampullosa (Brongniart) oClava voglinoi (Michelotti) oCerithium calculosum Defrance o o oCerithium vulgatum Bruguie're o oCerithium globulosum Deshayes oCerithium ex gr. granulinum (Bellardi and Michelotti) oCerithium (Ptychocerithium) bronni turritoplicatum Sacco oCerithiidae indet. 1 oCerithiidae indet. 2 oHemicerithium? sp. oDiastoma costellatum elongatum (Brongniart) o oDiastoma cf. alpinum (Tournouer) oAgapibittium greveniense Harzhauser and Kowalke oBayania semidecussata (Lamarck) nov. ssp. oAporrhais meridionalis (Basterot) oAporrhais uttingerianus (Risso) o oTibia dentata (Grateloup) o oRimella (Dientomochilus) decussata (Defrance) oRimella subrimosa d’Orbigny oOostrombus auriculatus (Grateloup) oOostrombus irregularis (Fuchs) oStrombus (Dilatilabrum) roegli Harzhauser o o oStrombus radix (Brongniart) o oStrombus bonelli (Brongniart) o o o oStrombus sp. oXenophora cumulans (Brongniart) o oXenophora deshayesi (Michelotti) o o oXenophora cf. deshayesi (Michelotti) oXenophora nov. sp. oCalyptraea cf. chinensis (Linne¤) oHipponix cf. interruptus Michelotti oCypraea cf. globosa (Dujardin) oCypraea cf. subovum (d’Orbigny) oCypraea (Zonaria) cf. fabagina Lamarck oCypraea sp. 1 oCypraea sp. 2 oCypraea sp. 3 oCypraea sp. 4 oCypraea sp. 5 oCypraea sp. 6 oCypraea sp. 7 oProadusta inquidens inaequilabiata (Sacco) oProadusta parvitala (Sacco) o oTrivia? sp. oAmpullinopsis crassatina (Lamarck) o o




Globularia gibberosa (Grateloup) o oGlobularia cf. callosa (Sowerby) oNatica sp. oNatica tigrina (Defrance) o o oNatica cf. subepiglottina d’Orbigny o oPolinices cf. redemptus (Michelotti) oEuspira catena helicina Brocchi o oNatica burdigalensis Mayer oNeverita olla (De Serres) o o o oAmaurellina (Euspirocrommium) oweni (Archiac and Haime) oAmaurellina (E.) a¡. sindiensis (Cossmann and Pissarro) oAmaurellina sp. 1 oAmaurellina sp. 2 oSinum michaudi (Michelotti) oSinum aquensis (Recluz) oSinum cf. striatum (De Serres) oSinum sp. oCassis mamillaris Grateloup o o oCassis vialensis Fuchs oCassis sp. oCassididae indet. oGaleodea? sp. oSemicassis cf. rondeleti apenninica (Sacco) oSemicassis grateloupi (Deshayes) oSconsia striata miocenica Sacco o oEudolium subfasciatum Sacco oCassidaria cf. tauropomum Sacco oSassia apenninica (Sassi) oSassia cf. £andrica (De Koninck) oCharonia sp. oCymatiidae indet. oCymatium? sp. 1 oCymatium? sp. 2 oFicus condita (Brongniart) o o o o oFicus cingulata (Bronn) oFicus oligo¢coides (Sacco) oFicus sp. oFicopsis (Fulguro¢cus) burdigalensis (Sowerby) oCerithiopsis sp. oMurex partschi ssp. oMurex (Haustellum) tchihatche⁄ Archiac and Haime oPterynopsis cf. tristichus (Beyrich) oEditharus polygonus (Lamarck) oCymia hellenica Harzhauser and Kowalke oEuthriofusus burdigalensis (Defrance) oDorsanum (Dorsanum) haueri excellens (Scha¡er) oNassarius (Hinia) nov. sp. oNassarius sp. 1 oNassarius sp. 2 oNassarius cf. semistriata (Brocchi) oNassarius cf. restitutianus (Fontannes) oNassarius sp. cf. limatus Chemnitz oMelongena lainei (Basterot) o o oMelongena semseyiana (Erdo«s) oMelongena cf. semseyiana (Erdo«s) o




Melongena cornuta (Agassiz) oMelongena subcarinata (Lamarck) oMelongena a¡. basilica (Bellardi) oMelongena sp. oPleuroploca tarbelliana (Grateloup) oLatirulus sp. oFusinus cf. costellatus (Grateloup) oStreptodictyon cf. subelongatus (d’Orbigny) oMitrella (Atilia) fallax (Ho«rnes and Auinger) oAcamptochetus submitraeformis (d’Orbigny) oSiphonalia sp. oTudicla rusticula (Basterot) o o oTurbinella episoma (Michelotti) oMitra sp. oAmalda (Baryspira) glandiformis Lamarck o o oAmalda (Baryspira) glandiformis anomala (Schlotheim) oOliva (Anazola) cf. clavula (Lamarck) o oOliva (Strephona) dufresnei Basterot oOlivella cf. longispira Bellardi oAthleta (Athleta) consanguinea (Bellardi) oAthleta (Athleta) ¢culina (Lamarck) o o o oVolutilithes subambigua d’OrbignyLyria magorum angustolonga Sacco oLyria anceps (Michelotti) o oLyria cf. anceps (Michelotti)Sveltia sp. oTerebra subtessellata d’Orbigny oTerebra pertusa Basterot oTerebra cf. pertusa subacuminata Peyrot oConus (Conolithus) antediluvianus Bruguie're o oConus (Conolithus) dujardini Deshayes o o o oConus (Lithoconus) antiquus Lamarck o oConus (Lithoconus) cf. mercati (Brocchi) o oConus (Lithoconus) nov. sp. oConus (Stephanoconus) cf. carcarensis (Sacco) oConus (Leptoconus) diversiformis Deshayes o oConus sp. 1 oConus sp. 2 oConus sp. 3 oGenota ramosa (Basterot) o o oAcamptogenotia intorta Brocchi oBathytoma cataphracta (Brocchi) oClavatula asperulata(Lamarck) o o oClavatula cf. glaberrima (Grateloup) oClavatula cf. carinifera Grateloup oClavatula calcarata Grateloup ssp. oClavatula sp. 1 oClavatula sp. 2 oClavus sp. oRaphitoma sp. 1 oRaphitoma sp. 2 oTurris cf. monilis (Brocchi) oTurris sp. 1 oTurris sp. 2 oTurridae indet. 1 o



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