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Palaeoclimatic implications of continental saline and fresh water mollusc communities of the Cenozoic Iberian Peninsula

By Thorsten Kowalke*

Ludwig-Maximilians-Universität, Department für Geo- und Umweltwissenschaften, Sektion Paläontologie, Richard-Wagner-Straße 10, 80333 Munich, Germany

Manuscript received June 15, 2006; revision accepted September 10, 2006.

Abstract salinarer und Süßwasser-Molluskenvergesellschaftungen aus dem Känozoikum der Iberischen Halbinsel untersucht. Einige Spatio-temporal distribution pattern of continental saline thermophile Vertreter der , welche rezent nicht and fresh water mollusc palaeocommunities of the Cenozoic mehr auf der Iberischen Halbinsel vorkommen und heute in Iberian Peninsula were investigated in order to detect palaeo- Nordafrika, dem südöstlichen Mittelmeer und Kleinasien ver- climatic implications. Several thermophilous representatives of breitet sind, zeigen warm subtropisches Klima an. Kontinentale the Cerithioidea, which indicate a warm subtropical climate, are aquatische Ökosysteme wurden zu unterschiedlichen Zeiten absent from the extant Iberian fauna and restricted to North des Känozoikums von verschiedenen Vergesellschaftungen Africa, the SE Mediterranean and Asia Minor. The fossil re- besiedelt: Verbreitungsmuster von Melanoides-Vergesellschaf- cord suggests that the different continental aquatic ecosystems tungen (Oligozän, Ebro Becken; Miozän, Albacete; Pliozän, were colonised at different times in the Cenozoic: distribution Barcelona; frühes Pleistozän, Guadix Baza Becken) und Pota- pattern of Melanoides palaeocommunities (Oligocene, Ebro mides-Vergesellschaftungen (Oligozän, Ebro Becken; Miozän, Basin; Miocene, Albacete; Pliocene, Barcelona; Lower Pleis- Duero Becken, Ebro Becken; frühes Pleistozän, Guadix Baza tocene, Guadix Baza Basin) and Potamides palaeocommunities Becken) und ihr sukzessiver Rückgang in den nordspani- (Oligocene Ebro Basin; Miocene, Duero Basin, Ebro Basin; schen Becken seit dem mittleren bis späten Miozän spiegeln Early Pleistocene, Guadix Baza Basin) and their successive die globale spätkänozoische Temperaturabkühlung wieder. disappearance since the mid- to Late Miocene in the northern Potamides-Vergesellschaftungen in der playa lake-Fazies des Spain basins refl ect the global Late Cenozoic temperature coo- Guadix Baza Beckens deuten auf ein semi-arides Klima hin, ling. Potamides palaeocommunities in playa lake-environments ähnlich dem heutigen Klima Nordafrikas. Phasen humideren of the Guadix Baza Basin indicate a semi-arid climate context, Klimas werden durch syntope Melanoides-Vergesellschaftun- similar to the modern North African climate. Humid intervals gen angezeigt, welche hyposalinare Ökosysteme besiedelten, are indicated by syntopic Melanoides palaeocommunities that ähnlich den aquatischen Ökosystemen zur Zeit der humideren colonised hyposaline environments, similar to the colonisation Klimaepisoden im Holozän Nordafrikas. -Verge- of aquatic ecosystems during the Holocene more humid climate sellschaftungen sind im Vergleich zu Melanoides- und Pota- episodes in North Africa. Melanopsis palaeocommunities are mides-Vergesellschaftungen durch höhere Temperaturtoleranz characterised by a greater temperature tolerance compared to gekennzeichnet. Die rezente Art Melanopsis praemorsa kommt the investigated Melanoides and Potamides palaeocommunities. in Spanien südlich des 42. Breitengrades vor; eine Verbreitungs- The extant species Melanopsis praemorsa is distributed in Spain grenze die etwa der +10°C-Januar-Isotherme entspricht. Das south of the 42nd degree of latitude, roughly coinciding with Verbreitungsgebiet kongenerischer miozäner Vertreter reichte the +10°C-January-Isotherme, while the area of distribution dagegen über die nordspanischen Beckensysteme hinaus. of the Miocene congeners extended the northern Spain basin systems. Schlüsselwörter: kontinentale Mollusken, Känozoikum, Spanien, Paläoklima Key words: continental molluscs, Cenozoic, Spain, pala- eoclimate 1. Introduction

Kurzfassung The Cenozoic Iberian Peninsula has a rich continental sedimentary record, preserved in large depressions and smal- Zur Klärung paläoklimatologischer Implikationen wurden ler restricted basins, which predominantly formed after the räumliche und zeitliche Verbreitungsmuster kontinentaler Priabonian (Late Eocene) regression. The basin-systems arose

*E-mail: [email protected] 76 induced by internal deformations of the Iberian micro-plate, Eocene). Evaporitic accumulations, which have been deposited coinciding with the formation of the Pyrenees and the Betic after the Pribonian regression, represent the terminal marine Orogene (BOCCALETTI et al. 1974; HSÜ 1977; RIBA et al. 1983; sedimentation. Oligocene and Neogene lacustrine fresh water PUIGDEFÁBREGAS et al. 1986; FRIEND & DABRIO 1996). A wide and athalasso-saline sedimentary successions were subsequent- variety of continental aquatic ecosystems and corresponding ly deposited in the context of an intracontinental basin without deposits established in these basins, which individual deve- marine connections (BATALLER 1929; RIBA et al. 1983; ANADÓN lopment was depending on the restriction of the basins or 1989). Similarly to the situation in the Ebro Basin, subsequent temporary or persisting connection with the Mediterranean to the Priabonian regression Oligocene and predominantly Sea, palaeoclimate, local ground water hydrology and the Miocene successions were deposited in the Duero Basin, the geochemical composition of the sedimentary basement. Apart south-western continuation, where the Neogene deposits from perennial and temporary fresh water palaeoenvironments, discordantly overly the Palaeozoic and Mesozoic basement athalasso-saline, i.e. continental saline ecosystems without (SÁNCHEZ-BENAVIDES et al. 1988, 1989). The northern margin marine connections, exhibited a broad spectrum of palaeoha- of the Duero Basin is characterised by fossiliferous athalas- bitats for the colonisation by fresh water and saline mollusc so-saline and fresh water deposits of Miocene age, which are palaeocommunities (ANADÓN 1989; KOWALKE 2006a, 2006b); particularly well-exposed in the Burgos/Castrillo del Val area see also BEADLE (1943), WILLIAMS (1981), and HAMMER (1986) (GONZÁLES-DELGADO et al. 1986; ANADÓN 1989; KOWALKE for general descriptions of the ecology of athalasso-saline 2006b). The Guadix-Baza Basin represents an intra-montane ecosystems. depression, which is situated above the contact of the Betic and Fossiliferous deposits are particularly well known from the Subbetic Zone of the Betic Orogene. While Upper Miocene Ebro and Duero Basins (N, NW and Central Spain) and the successions refl ect a marginal marine palaeoenvironment with Guadix-Baza Basin in S Spain (Fig. 1). The Ebro Basin formed marine-brackish faunas, Pliocene-Lower Pleistocene deposits along the southern margin of the Pyrenees. This large basin are characterised by alternations of fresh water and athalasso- was characterised by two major marine transgressive episodes saline faunas (AGUSTÍ et al. 1985; ANADÓN et al. 1986, 1987; with subsequent regressions in the course of the Ilerdian-Cui- ROBLES 1989; KOWALKE 2006a, 2006b). sian (Early Eocene) and Lutetian-Priabonian (Middle to Late Apart from rather uncharacteristic basommatophoran fresh

Figure 1: Location of the fossiliferous continental basin systems of the Iberian Peninsula. 77 water and archaeopulmonate paludinal faunas (Lymnaeidae- forms originated from the southern East Atlantic Province Planorbidae- and Carychium-palaeocommunities) with wide and exhibited a higher tolerance with regard to colder climatic European distribution, several thermophilous fresh water conditions. associations (Melanopsis- and Melanoides-palaeocommunities) With regard to the faunal composition, the Pyrenean mollu- and athalasso-saline faunas (Potamides-Corbicula-, Potamides- sc associations were very similar to the coastal faunas described Cerastoderma-, Potamides-Hydrobia-, and Melanoides-pala- from other European Eocene localities with tropical character, eocommunities) characterised the basin-systems in particular with several co-occurrences on genus level. For example, Cenozoic time intervals and were absent in others (KOWALKE corresponding faunas are well-known from the Lutetian of 2006b). The occurrences of these palaeocommunities may SW France (ROUAULT 1848), the Paris Basin (COSSMANN 1889, indicate time intervals during the Iberian history with conside- 1906a, 1906b) or W Hungary (SZÖTS 1953; STRAUSZ 1966; rably different distribution pattern of the characteristic faunas KECKSKEMÉTI-KÖRMENDI 1972; KOWALKE 2001, 2004). Similar than present, probably refl ecting different climatic conditions. Tethyan coastal palaeocommunities, distinguished on genus Furthermore the presence of perennial aquatic ecosystems level, have been described from South California, U.S.A. connected to riverine environments and the distribution of the (SQUIRES 1999). corresponding palaeocommunities as well as sedimentological Along with the decrease of the mangrove-ecosystems, features may be indicative of precipitation rates different from coinciding with the climate cooling in the course of the la- the situation in the subhumid West Mediterranean climate zone test Eocene/Early Oligocene (PLAZIAT et al. 2001), most of of Iberia today (see e.g., WALLÉN 1970 for comparisons with the associated molluscs such as Cerithidea became extinct the extant climate of the Iberian Peninsula). in Europe and migrated in the direction of the tropical bio- The aim of the present study is to assess whether spatio-tem- provinces (HOUBRICK 1884; HARZHAUSER & KOWALKE 2001; poral distribution pattern of fresh water and athalasso-saline KOWALKE 2003, 2005). A few coastal brackish water dwellers, molluscs of Oligocene to Quaternary palaeoenvironments e.g., Granulolabium, Terebralia, and Tympanotonos, exhibited refl ect the climatic development of the Iberian Peninsula. tolerance with regard to decreasing temperatures and colonised Comparisons with distribution patterns displayed by mar- the wet coastal ecosystems succeeding the mangrove palaeo- ginal marine Eocene faunas, contemporaneous faunas of the environments (e.g., MOISESCU 1969, 1972; GITTON et al. 1986, Mediterranean and Paratethys, and extant congeners have been PLAZIAT et al. 2001). carried out in order to document the mollusc faunal change Avicennia-dominated mangroves re-established during and its palaeoclimatic signifi cance. a temperature increase in the course of the Late Oligocene, with records in Spain, southern France and Hungary (NAGY & KÓKAY 1991) and probably far northward extensions to the 2. The Tethyan precursors – evidence Bavarian Molasse Sea (BARTHELT 1989; REICHENBACHER et al. from Eocene euryhaline coastal 2004). The associated molluscs (predominantly Tympanotonos palaeocommunities palaeocommunities) were restricted to the coastal brackish- marine ecosystems, since the mode of early ontogenetic deve- lopment, i.e. the planktotrophic veliger phase, prevented the Ilerdian and Cuisian (Lower Eocene) coastal sedimentary characteristic faunas from the colonisation of the continental sequences of the southern Pyrenean Tremp-Graus Basin, which saline ecosystems (KOWALKE 2006b). were deposited during the time interval encompassing the Early Eocene Climatic Optimum (EECO), are characterised by a rich record of mollusc associations. The coastal assemblages 3. The continental uppermost Eocene are composed of gradually changing mangrove/estuarine and and Oligocene record shallow subtidal palaeocommunities (DOMINICI & KOWALKE, in press). The mangrove and estuarine palaeocommunities are com- The Oligocene Ebro Basin was characterised by extended posed of characteristic tropical caenogastropod genera, which palaeo-lake environments, which resulted in the deposition are still dominating the mangrove ecosystems of the modern of thick successions of mudstones, limestones and coal se- faunal bioprovinces: the Neotropis, the West African coast, the ries. Lacustrine deposits from the Lower Oligocene of Calaf Western Indian Ocean and the Indo-Polynesian Province (VAN within the eastern Ebro Basin consist of grey mudstones and REGTEREN ALTENA 1942; KOWALKE 2001; PLAZIAT et al. 2001). intercalated layers of limestones, forming sequences which Several extant thermophilous genera of the families Cerithii- comprise up to one metre sediment. The marginal portions of dae (see HOUBRICK 1985; 1992), Potamididae (e.g., Cerithidea this depositional environment are represented by marls rich and Tympanotonos, cf. HOUBRICK 1984, 1991) and Thiaridae in plant detritus and coal layers; see CABRERA & SAEZ (1987) (Melanoides, cf. BROWN 1980; GLAUBRECHT 1996), and extinct for a detailed sedimentological reconstruction. The lacustrine genera of Pachymelaniidae and Pachychilidae (see KOWALKE ecosystem was characterised by an oligotypic freshwater fauna 2001, 2004), which are morphologically very similar to their dominated by the thiarid Melanoides albigensis (NOULET), a extant equivalents, document the tropical character of the species that is morphologically very similar to the modern faunas and the corresponding palaeoenvironments. Apart from type species M. tuberculata (MÜLLER) (e.g., BROWN 1980; these stenotherm elements, to lesser degree cerithioidean taxa GLAUBRECHT 1996; KOWALKE 1998, 2006a). The accompanying such as Granulolabium formed part of the palaeocommunities fauna is rather uncharacteristic, being composed of fragmentary (e.g., LOZOUET 1986; HARZHAUSER & KOWALKE 2001). These large pulmonates (Lymnaea sp.) and hydrobiids. 78 Melanoides represents a thermophilous element, which ration of 1000–2000 mm considerably exceeds this rate. The tolerates subtropical climates of N Africa and Asia Minor; precipitation and evaporation rates in the Oligocene climate for distribution data see e.g., BROWN (1980) and GLAUBRECHT context of the Calaf-ecosystems should have been comparable (1996). The extant type-species is pantropically distributed, in order to maintain in time an extended body of fresh water, but also occurs in North Africa and Asia Minor. Other extant see also CABRERA & SAEZ (1987) for a sedimentological-pala- thiarid genera are restricted to the tropics. While Melanoides eohydrological interpretation. represents a parthenogenetic direct developer which develops Upper Oligocene athalasso-saline successions of the Ebro within a brood pouch, related thiarids need a fair connection Basin are characterised by typical Potamides-Corbicula pa- of their habitats to the open sea, since they are characterised laeocommunities, which are predominantly exposed in the by a marine larval stage and enter the fresh water portions of plane of Zaragoza (VIDAL & DEPERET 1906; BATALLER 1929; rivers from estuarine habitats settled by juveniles after meta- ANADÓN 1989) (Figs 2–3). The associations of perennial morphosis, subsequent to the planktotrophic development molluscs indicate the presence of persisting saline aquatic (KOWALKE 1998). palaeoenvironments. The presence of palaeocommunities dominated by Mela- Apart from these ecosystems which refl ect stronger, probab- noides indicates a warmer subtropical climate than extant. The ly seasonal fresh water infl ux, during the Late Eocene common perennial character of the faunas and the presence of extended shallow saline lakes with characteristic micro-faunas existed in fresh water ecosystems (cf. CABRERA & SAEZ 1987; ANADÓN the Valldeperes region (Ebro Basin), composed of ostracods 1989) refl ect a considerably higher precipitation rate compared and foraminifers (ANADÓN 1978, 1989). The latter deposits lack to the modern climate of the Iberian Peninsula, which belongs a mollusc fauna, an absence that could indicate the temporary to the subhumid West Mediterranean climate zone (WALLÉN character of the water bodies in the context of temporarily and/ 1970). This climate zone is characterised by a mean annual or locally established semi-arid climate conditions inconvenient precipitation rate of 400–800 mm, while the possible evapo- for the colonisation by a perennial mollusc fauna.

Figure 2: Occurrences of Iberian Cenozoic thermophilous gastropod genera. Dotted symbols indicate occurrences of Eocene coastal taxa; black symbols = Oligocene athalasso-saline taxa; half fi lled symbols = Miocene-Pliocene athalasso-saline taxa; black symbols with white centre = Lower Pleistocene athalasso-saline taxa; white symbols = extant occurrences of Melanopsis. 79 Neogene distribution, they have neither been reported from 4. Evidence from Miocene athalasso-saline fossil European athalasso-saline ecosystems (see e.g., ANADÓN and fresh water faunas 1989), nor from corresponding extant tropical environments (KOWALKE 2005, and ecological data in HOUBRICK 1991). The entire spectrum of potamidids with planktotrophic larval de- While potamidid palaeocommunities dominated by Tym- velopment was hampered from the colonisation of continental panotonos, Terebralia and Granulolabium and indicative of saline ecosystems because larvae of gastropod could not suf- a warm subtropical climate, characterised marginal marine fi ciently develop in stagnant water bodies (cf. KOWALKE 1998, brackish to hypersaline environments with wide distribution 2005, 2006a, 2006b). in the Mediterranean and Paratethyan Upper Oligocene to Potamides represents the only known potamidid which mid-Miocene (e.g., GITTON et al. 1986; LOZOUET et al. 2001; lacked a planktotrophic larval stage and thus could colonise HARZHAUSER & KOWALKE 2001, 2002; HARZHAUSER et al. 2002), intracontinental saline lake-environments (PLAZIAT 1989, corresponding assemblages are not found in athalasso-saline 1993). A characteristic palaeocommunity occurred in Upper palaeo-ecosystems (KOWALKE 2005, 2006a, 2006b). Oligotypic Astaracian to Lower Vallesian (upper Middle to early Upper Terebralia-palaeocommunities were, for example, still present Miocene) athalasso-saline deposits of the Duero Basin (NW in the Upper Tortonian/Messinian partly hypersaline lagoon- Spain) in the Castrillo del Val/Burgos area (Figs 2–3) where systems with marine connections at Elche and Crevillente/Ali- Potamides gaudryi LARRAZET was associated with the hydro- cante, SE Spain (KOWALKE 2006a). The accompanying fauna biids Hydrobia deydieri DEPÉRET & SAYN and H. calderoni consisted of Granulolabium, Cerithium and juvenile bivalves ROYO-GOMEZ. Apart from the saline fauna of marine origin, of the genera Cerastoderma and Abra. Although Terebralia and primarily freshwater but salinity-tolerant elements (see REI- Granulolabium were characterised by a wide Mediterranean CHENBACHER et al. 2004; KOWALKE & REICHENBACHER 2005)

Figure 3: Spatio-temporal distribution pattern of the thermophilous genera Melanoides and Potamides in different continental facies and palaeoenvironments. Melanoides: black symbol = occurrence in Oligocene fresh water ecosystems; dotted = occurrence in Miocene fresh water ecosystems; white = occurrences in Pliocene and Early Pleistocene athalasso-saline ecosystems. Potamides: dotted = occurrences in Oligocene and Miocene athalasso-saline ecosystems; white = occurrence in Early Pleistocene playa lake ecosystems. 80 such as Lymnaea and Theodoxus formed part of the associa- In the course of the Early Pleistocene, the thermophilous tions. These elements predominantly lived in small rivers or element Melanoides was still characterised by a considerably creeks which were connected with the athalasso-saline lake more northern geographic distribution than extant. ANADÓN environment – an interpretation of the palaeo-habitat, which et al. (1987) and ROBLES (1989) reported probably hypohaline is also supported by sedimentological data. The euryhaline M. tuberculata associations from the Lower Pleistocene of the potamids established permanent populations that persisted for Guadix-Baza Basin. According to ESU (1980) contemporaneous several years. This interpretation of the life-spans of individuals faunas dominated by Melanoides occurred on the Balearic of P. gaudryi is probable, because the life-cyles of the closely Islands. Extant natural populations of M. tuberculata are related extant type species P. conicus (BLAINVILLE) from the absent from the Iberian Peninsula (ÁLVAREZ-HALCÓN 1995). South-East Meditteranean exceeded annual cycles and amounts The species occurs in N Africa and Asia Minor (BROWN 1980; several years (KOWALKE 2003). GLAUBRECHT 1996; KOWALKE 2005, 2006b). Artificially intro- Associations similar to the palaeocommunities from the duced Iberian populations of M. tuberculata are restricted to Duero Basin have also been documented from the Langhian hydrothermal vents, e.g., the Alhama de Aragón/Zaragoza (see and Serravallian of the Ebro Basin, particularly in the plane of ÁLVAREZ-HALCÓN 1995). Zaragoza (Muniesa, Belchite, Fuendetodos), where Potamides Potamides-Hydrobia palaeocommunities characterised a occurred in association with Cerastoderma (see ANADÓN 1989 shallow lacustrine playa lake- facies of the Lower Pleistocene and references therein). Guadix-Baza Basin in the vicinity of Orce (ROBLES 1989; The presence of the potamidid-dominated faunas in the KOWALKE 2006b) (Fig. 3). The occurrence of Potamides in pla- Duero and Ebro basins during the Middle Miocene refl ects ya-lakes is indicative of a warm subtropical semi-arid climate, a warm subtropical climate, since the closely related extant comparable to the Holocene climate in N Africa. Melanoides type-species P. conicus BLAINVILLE is restricted to the SE Me- faunas occurred well separated from the Potamides palaeocom- diterranean, distributed south of the 40th degree of latitude, munities in aquatic environments, which were characterised along the N African coast and on the E Mediterranean Islands by lower salinity levels (KOWALKE 2006b). The Melanoides (KOWALKE 2003). The persisting character of the palaeo-lake associations could refl ect episodes with at least seasonally environments is – similarly to the situation during the Iberian higher precipitation rates and possibly subsequent exposure Oligocene – indicative of a more humid climate compared to to progressive evaporation. Corresponding associations have the extant climate of the Iberian Peninsula, with precipitation been described from the Holocene of Algeria (KOWALKE 2006b) rates exceeding, or at most equalling the evaporation rates, and the Ethiopian Rift Valley (LENG et al. 1999). and leading to the formation of extended riverine systems The genus Melanopsis exhibited a higher tolerance with connected to the lacustrine palaeoenvironments with marginal regard to the Late Cenozoic temperature decrease: Melanopsis swamps (CABRERA & SAEZ 1987). praemorsa (LINNAEUS) is still distributed in extant fl uviatile Corresponding indications for the palaeoclimatic situation environments of the Iberian Peninsula, occurring south of are also evident from Miocene fresh water faunas: extended Up- the 42nd degree of latitude. The northern limit of the area per Miocene lacustrine habitats in SE Spain were colonised by of distribution roughly coincides with the +10°C-January- characteristic thermophilous associations. For example, MEIN Isotherme (see also GLAUBRECHT 1996 and ALTABA 1998 for et al. (1978) described freshwater deposits from the Turolian distribution data). of Tolosa (NE Albacete) that are dominated by “Melania” (= Melanoides) associated with Melanopsis. 6. Climatic implications of mollusc faunal dynamics in other European 5. The Plio-Pleistocene fossil record and continental sites the extant distribution pattern

Corresponding climatically induced faunal successions and During the Late Messinian, the genus Melanoides inhabi- distribution patterns are rarely documented from European ted hyposaline palaeoenvironments, which were previously continental palaeoenvironments. The marginal marine-bra- colonised by Potamides spp. (KOWALKE 2006b). The palaeo- ckish Iberian fauna was not dissimilar to (Proto-)Mediterrane- communities are dominated by the species Melanoides cur- an and Paratethyan Oligocene and Neogene Granulolabium-, vicosta V. MARTENS, which is morphologically very similar to Terebralia- and Tympanotonos palaeocommunities in being the modern type-species M. tuberculata. Apart from frequent prevented from the colonisation of athalasso-saline palaeo- occurrences of continental Melanoides-faunas in the typical environments by its indirect planktotrophic mode of early Upper Messinian Lago Mare-facies of Italy (ESU & GIROTTI ontogenetic development (e.g., HARZHAUSER & KOWALKE 2001, 1974; ESU 1980), similar hyposaline palaeocommunities 2002, and references therein). Potamides represents the only have been described from athalasso-saline deposits from the potamidid which changed its predominant mode of indirect Pliocene of Spain: Melanoides tournoueri (= M. curvicosta) development to lecithotrophic and direct development and occurred in a characteristic oligotypic association in Pliocene was thus able to colonise athalasso-saline habitats. Apart from deposits of Barcelona, accompanied by Neritina, Valvata, the Neogene records in Spain, continental palaeocommunities Melanopsis and Hydrobia (ALMERA et al. 1892; KOWALKE dominated by Potamides have also been described from SW and 2005). central France (REY 1965; PLAZIAT & GAUDANT 1984). 81 Thermophilous Melanoides faunas are well known from the Iberian Peninsula, where precipitation and evaporation had the Middle Eocene of W Hungary, where they occurred in comparable rates. Athalasso-saline Melanoides-faunas from marginal fresh water environments adjacent to, but well se- Barcelona indicate a Pliocene temperature peak. Lower Pleis- parated from, the tropical Tethyan brackish coastal swamps tocene occurrences are restricted to the southern Spain Guadix (KOWALKE 2001). Baza Basin. The lack of corresponding faunas in northern sites Melanoides assemblages are rarely documented from Up- of the former area of distribution refl ects the temperature per Oligocene deposits of the Lower Brackishwater Molasse cooling in the course of the Late Cenozoic. (Bavaria, S Germany) where the oligotypic fresh water pa- The potamidid genus Potamides represents a second indica- laeocommunities occurred well separated from the brackish tor of the Cenozoic climate development. Potamides-Corbicula Tympanotonos-Polymesoda- and Granulolabium associations associations frequently characterised Oligocene athalasso-sa- (REICHENBACHER et al. 2004) in a quiet water pond-facies line deposits of the Ebro Basin. Miocene athalasso-saline eco- (KOWALKE submitted). Ottnangian/Karpatian and Badenian systems were characterised by oligotypic Potamides-Hydrobia (Lower to lower Middle Miocene) brackish and fresh water (Duero Basin) and Potamides-Cerastoderma palaeocommu- deposits of the Upper Brackishwater Molasse and Upper nities (Ebro Basin). Similarly to the Pleistocene distribution Freshwater Molasse lack Melanoides faunas, although the pala- pattern of the Melanoides palaeocommunites, Potamides was eoenvironmental conditions would have been convenient (e.g., restricted to Andalusia during the Early Pleistocene, and was KOWALKE & REICHENBACHER 2005). The faunal composition of absent from the Iberian fauna since the Late Pleistocene. The the palaeocommunities of the marginal Molasse Sea and subse- Lower Pleistocene playa-lake environment, colonised by the quent fresh water palaeoenvironments point to a slight decrease Potamides palaeocommunities, indicates a semi-arid climate, in temperature, compared to the palaeoclimatological history similar to the modern North African climate. More humid of the Iberian Peninsula (KOWALKE 2005, 2006b). However, intervals are indicated by hyposaline environments colonised the conditions still refl ect warm climates, since Melanopsis, an by Melanoides palaeocommunities, comparable to Holocene element which exhibits a higher tolerance regarding slightly humid-climate episodes recorded in North Africa. colder temperate conditions, was still present in Ottnangian Melanopsis faunas exhibit a greater temperature tolerance deposits of the Bavarian Kirchberg Beds and the Oncophora compared to the potamidid and thiarid associations. The Beds (KOWALKE & REICHENBACHER 2005). A stronger decline extant melanopsid M. praemorsa is still present in fl uviatile of the temperature is indicated by the composition of the San- ecosystems of the Iberian Peninsula, distributed as far north delzhausen fauna (Early Badenian, Bavaria, S Germany), which as the 42nd degree of latitude, roughly coinciding with the faunal composition lacks any thermophilous elements, and has +10°C-January-Isotherme, whereas fossil melanopsids were apart from basommatophoran pulmonates even temperate to characterised by an area of distribution extending the basin colder water elements, such as Margaritifera (KOWALKE, own systems of northern Spain. observation). A corresponding development could not be ob- served from the Iberian faunas, where Melanoides was present until the Early Pleistocene (ROBLES 1989; KOWALKE 2006b). Acknowledgements Relic Melanoides palaeocommunities, which represented typical and ubiquitous associations of the Messinian Lago Silvio KELLER (Munich, Germany) assisted with the prepa- Mare-facies, have also been described from the Lower Pleis- ration of the fi gures. Michael KRINGS (Munich, Germany) im- tocene of southern Italy (ESU & GIROTTI 1974; ESU 1980). proved the English. The paper benefi ted from the constructive review by Stefano DOMINICI (Florence, Italy). The Deutsche Forschungsgemeinschaft (DFG) fi nancially supported my 7. Conclusions investigations of the evolution of brackish and inland saline molluscs (grant KO 2066/3-1). To all persons and institutions Apart from the marginal marine-brackish mollusc asso- involved I express my sincere thanks. ciations and their spatio-temporal distribution pattern, the continental athalasso-saline and fresh water palaeocommunities provide evidence for the palaeoclimate of the Iberian Peninsula 8. 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