Holoplanktonic gastropoda (Mollusca) from the Miocene of Cyprus: systematics and biostratigraphy

Abstract: Eleven randomly chosen outcrops in the Miocene Pakhna Formation of Cyprus were sampled for holoplanktonic Mollusca. Four species of Heteropoda were found, and 24 of Pteropoda, a substantial increase from the two species recorded until now from the Miocene of Cyprus. One pteropod species, Peracle charlotteae sp. nov. (Gastropoda, Pseudothecosomata), is introduced. Age assignments based on holoplanktonic molluscs for the 11 localities are as follows: Langhian (Alassa 1–4), (Serravallian?) Tortonian to Messinian (Episkopi 1), Tortonian (Agios Tychon, Tokhni and the Maroni Marlstone of Khirokitia 1–2) and Tortonian to Early Messinian (Episkopi 2). These age determinations in some cases are at odds with those from previous publications based on calcareous nannofossils and Foraminifera. The sediments underlying the Amathus Channel outcrop yielded insufficient fauna for definitive dating, but we suggest are younger than Late Serravallian. At some localities, particularly in the Alassa area, pteropod assemblages are strongly variable on a bed‐by‐bed basis, and this offers possibilities for future refined biostratigraphical interpretations. This is the first substantial holoplanktonic mollusc fauna described from the eastern Mediterranean basin and allows correlation with assemblages in the central Mediterranean and elsewhere.     

Paleoenvironmental evolution of the eastern Mediterranean during the Messinian: Constraints from integrated microfossil data of the Pissouri Basin (Cyprus)

Integrated data of calcareous nannofossils, as well as planktonic and benthic foraminifera from the Pissouri Motorway section on Cyprus allow the reconstruction of surface- and bottom-water paleoenvironments of the eastern Mediterranean during the interval preceding the Messinian salinity crisis (MSC). Contrary to deeper-water locations, where benthic foraminifera faunas are suppressed or absent just after the Tortonian–Messinian boundary, sediments deposited at intermediate water depths do contain benthic assemblages. From the earliest Messinian onwards, a development towards increasingly unfavourable paleoenvironments is reflected in the planktonic and benthic microfossil records of the Pissouri section and proceeds with rather discrete time steps that can be correlated to sequences throughout the Mediterranean. Shortly after the Tortonian–Messinian boundary a transition is recorded in the sedimentology and the open marine, deeper-water taxa disappear from the benthic foraminifera assemblages; subsequently, the diversity of all fauna groups diminishes. The changes recorded at species level in both surface-water and sea-floor dwelling taxa suggest decreasing circulation of the bottom waters, associated with changes in the surface waters, most likely due to increasing stratification. From ∼6.73 Ma onwards, our data indicate a prominent change to more restricted conditions and increasing salinity at the sea floor together with intermittently rising surface water salinity. The dominance of oligotypic and monospecific assemblages and the frequent shifts in assemblage compositions of all microfossil groups indicate severely stressed environments after ∼6.4 Ma, probably related to increased salinity. The major changes in paleoenvironmental conditions, including oxygen deprivation due to stagnation and hypersalinity, can be explained by hydrographical changes in the Mediterranean basin, which are probably caused by tectonic movements in the Rif Corridor acting in concert with astronomical cyclicity. Evaluation of the paleodepth proxies indicates that the depth of the Pissouri Basin remained rather constant at ∼300–500 m, with a minimum of 200 m, until deposition of the “barre jaune”, the transitional interval towards the evaporites and that early shallowing to neritic depths, as was proposed before, is highly unlikely.     

Messinian bottom-grown selenitic gypsum: An archive of microbial life

Primary gypsum deposits, which accumulated in the Mediterranean Basin during the so-called Messinian salinity crisis (5.97–5.33 Ma), represent an excellent archive of microbial life. We investigated the molecular fossil inventory and the corresponding compound-specific δ¹³C values of bottom-grown gypsum formed during the first stage of the crisis in four marginal basins across the Mediterranean (Nijar, Spain; Vena del Gesso, Italy; Heraklion, Crete; and Psematismenos, Cyprus). All studied gypsum samples contain intricate networks of filamentous microfossils, whose phylogenetic affiliation has been debated for a long time. Petrographic analysis, molecular fossil inventories (hydrocarbons, alcohols, and carboxylic acids), and carbon stable isotope patterns suggest that the mazes of filamentous fossils represent benthic microbial assemblages dominated by chemotrophic sulfide-oxidizing bacteria; in some of the samples, the body fossils are accompanied by lipids produced by sulfate-reducing bacteria. Abundant isoprenoid alcohols including diphytanyl glycerol diethers (DGDs) and glycerol dibiphytanyl glycerol tetraethers (GDGTs), typified by highly variable carbon stable isotope composition with δ¹³C values spanning from ?40 to ?14‰, reveal the presence of planktic and benthic archaeal communities dwelling in Messinian paleoenvironments. The compound inventory of archaeal lipids indicates the existence of a stratified water column, with a normal marine to diluted upper water column and more saline deeper waters. This study documents the lipid biomarker inventory of microbial life preserved in ancient gypsum deposits, helping to reconstruct the widely debated conditions under which Messinian gypsum formed.     

Messinian palaeoenvironments and hydrology in Sicily (Italy): The dinoflagellate cyst record

Hydrological conditions prevailing before, during and after the Messinian salinity crisis in Sicily have been approached using dinoflagellate cyst records. The synthetic sequence considered is based on five classical sections from the Caltanissetta Basin. Our interpretations are based on the recognition of autochthonous, allochthonous and reworked population among the dinocyst assemblages. For the first time, sea-surface temperatures and seasonal salinity contrasts were tentatively reconstructed using a “Mutual Climatic Range Method”. Sicilian late Tortonian deposits correspond to marine environment with significant terrestrial inputs favourable to eutrophic dinocyst species. Immediately after the beginning of the Messinian Stage, euryhaline assemblages took place, followed by meso-hyperhaline taxa, within a general trend to shallowing. At the end of the Tripoli diatomitic Formation, environment appears confined, with regular oceanic inflows. Such marine inflows remain persistent during the deposition of the salt Member, witnessing the probable persistence of nearby normal marine sea-surface water conditions in the Mediterranean Sea but with possible reduced hydrological circulation and/or low nutrient component. Such inflows are slightly decreasing up to the top of the Sicilian Upper Evaporites. At the same time, river inputs appear weak during the salt deposition, as the consequence of a rather dry climatic context. As shown by reworking activity, terrestrial inputs increase progressively from the base of the Upper Evaporites. During the Lago Mare period, while climate remains rather dry (absence of freshwater algae inputs, very low amount in trees requiring humid conditions), local deposition environment is confined with mesohaline to hypohaline sea-surface waters, in a context with relatively high seasonal sea-surface salinity contrast (up to 6‰). The very high reworking observed in the Arenazzolo silts, in addition to slightly increasing water depth, led us to consider the presence of a discontinuity between the Lago Mare and the Arenazzolo Fms. We consider the Arenazzolo Formation as a transgressive facies following a deep downcutting period. The Arenazzolo Formation presents a two-step development. The first one, correlative with the presence of a G. etrusca (a species with Paratethyan affinities), corresponds to a relative high-stand sea-level with oceanic influxes, low seasonal sea-surface salinity contrast and probable more humid context, as revealed by the important freshwater algal inputs. During the second step, salinity becomes much more variable, with a clear increase of seasonal sea-surface salinity contrast, a possible slight mean sea-surface temperature increase (only few degrees) and a clear weakening of the river inputs. Taking the proposed Zanclean position of the Arenazzolo Fm. into consideration, we state that mean sea-surface temperature did not change significantly from the base of the Messinian to the earliest Zanclean. At 5.33 Ma, the suddenly achieved flooding restored a fair, deep oceanic environment characterized, at the beginning, by a clear mean sea-surface temperature cooling (up to 6-7 °C) and a nutrient depletion, associated with the basins starvation. Sea-surface salinities were normal, with very low seasonal contrast. Hydrodynamics then nutrient supply became then quite normal from c.a. 5.08 Ma. The status of the Sicilian Caltanissetta Basin as a marginal basin although fastly deepening and the stratigraphical location of the Messinian discontinuity at the base of the Arenazzolo is the scenario that best matches our dinocyst record.     

The Primary Lower Gypsum in the Mediterranean: A new facies interpretation for the first stage of the Messinian salinity crisis

The detailed facies and physical stratigraphic analysis of the Primary Lower Gypsum in the Mediterranean indicates a surprising bed-by-bed correlation at basin-scale (Spain, Italy, Hellenic arc and Cyprus arc), that is tuned to the orbital calibration for the first stage of the Messinian salinity crisis from 5.96 to 5.61 Ma ago. A total of 16, precessionally-controlled, gypsum cycles were deposited rapidly in less than 350 ka, forming sequences up to 300 m thick. The lack of subaerial exposure features and the common facies associations and stacking pattern for sections located thousands of kilometers apart in different geological settings indicates a modest depositional depth, not extremely shallow. Selenite deposition occurred only at the bottom of restricted marginal basins less than 200 m deep, while no gypsum could precipitate in the deeper euxinic Mediterranean portions where only thin and barren shale/dolostone couplets formed. The lowermost selenite beds pass laterally to dolomite-rich limestones interbedded with barren euxinic shales in poorly oxygenated settings, indicating that the gypsum sedimentation was diachronous and did not necessarily mark the onset on the Messinian salinity crisis.Evaporite facies sequences (EF1 to 8) within individual gypsum beds show small-scale, subaqueous sedimentary cycles that mimic regressive-transgressive cycles: a) initial evaporite precipitation at relatively low supersaturation produced the massive selenite (facies EF3) in a relatively deep setting; b) continuous evaporation and drawdown by oscillating brine level formed the banded selenite (EF4) at the aridity acme of the precessionally-controlled cycle; c) general progressive brine level rise with strong brine flow led to the formation of large selenite supercones branching laterally (supercones in Spain and branching selenite, EF5, in the rest of the Mediterranean); and d) flooding by undersaturated continental water terminated gypsum precipitation with the deposition of argillaceous sediments (EF1, Northern Apennines), and/or limestone (EF2, Sicily and Spain) during the humid phase in the precession climate cycle.The stacking pattern and selenite facies associations suggest an overall shallowing-upward trend with a basin-wide hydrologic change starting from the 6th cycle (5.84 Ma), which is marked by the appearance of the branching selenite facies (supercones) in Spain and indicates that the brines became current-dominated. The Sr-isotope stratigraphy suggests that in the Northern Apennines the brines were strongly modified by continental waters (⁸⁷Sr/⁸⁶Sr = 0.708893 to 0.708998), and received direct pulses of Atlantic seawater (⁸⁷Sr/⁸⁶Sr = 0.70900 to 0.709024) only in the upper part of the section. In areas away from the mainland, such as Sicily, the continental input was by far less important.     

Sierra del Colmenar 1A,a new late Messinian (Late Miocene) locality in the Bajo Segura Basin (SE Spain): biostratigraphic and palaeoenvironmental implications

Abstract

In this paper, we have studied a new micromammal site from the Sierra del Colmenar section (Elche, SE Spain), named Sierra del Colmenar 1A (SCO-1A), representing the uppermost levels of Messinian age of the Bajo Segura Basin. The sedimentary context of this locality corresponds to a costal lagoon with marine influence. The fossil site has yielded remains of Apodemus aff. gorafensis, Paraethomys meini, Apocricetus alberti, Occitanomys alcalai, Ruscinomys sp., Eliomys cf. truci, Muscardinus sp., Parasorex ibericus, Prolagus michauxi and Soricidae indet. Based on this assemblage, we propose a Late Miocene age (MN13) for the locality SCO-1A, matching the previously inferred age for the stratigraphic unit in which the fossil site is situated. According to its faunal composition, we infer warm and humid climate conditions in the moment of formation of this locality, agreeing with the environmental conditions of the Messinian age in a parallic sedimentary context.     

Phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs have been determined by geological processes and climate change in the Late Cenozoic

AIM: Our aims were to assess the phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs and to estimate divergence times using different geological scenarios. We related divergence times to past geological events and discuss the relevance of our data for the systematics of eastern Mediterranean water frogs. LOCATION: The eastern Mediterranean region. METHODS: Genetic diversity and divergence were calculated using sequences of two protein-coding mitochondrial (mt) genes: ND2 (1038 bp, 119 sequences) and ND3 (340 bp, 612 sequences). Divergence times were estimated in a Bayesian framework under four geological scenarios representing alternative possible geological histories for the eastern Mediterranean. We then compared the different scenarios using Bayes factors and additional geological data. RESULTS: Extensive genetic diversity in mtDNA divides eastern Mediterranean water frogs into six main haplogroups (MHG). Three MHGs were identified on the Anatolian mainland; the most widespread MHG with the highest diversity is distributed from western Anatolia to the northern shore of the Caspian Sea, including the type locality of Pelophylax ridibundus. The other two Anatolian MHGs are restricted to south-eastern Turkey, occupying localities west and east of the Amanos mountain range. One of the remaining three MHGs is restricted to Cyprus; a second to the Levant; the third was found in the distribution area of European lake frogs (P. ridibundus group), including the Balkans. MAIN CONCLUSIONS: Based on geological evidence and estimates of genetic divergence we hypothesize that the water frogs of Cyprus have been isolated from the Anatolian mainland populations since the end of the Messinian salinity crisis (MSC), i.e. since c. 5.5-5.3 Ma, while our divergence time estimates indicate that the isolation of Crete from the mainland populations (Peloponnese, Anatolia) most likely pre-dates the MSC. The observed rates of divergence imply a time window of c. 1.6-1.1 million years for diversification of the largest Anatolian MHG; divergence between the two other Anatolian MHGs may have begun about 3.0 Ma, apparently as a result of uplift of the Amanos Mountains. Our mtDNA data suggest that the Anatolian water frogs and frogs from Cyprus represent several undescribed species.     

The western edge of the Mediterranean Pelagian Platform: A Messinian mixed siliciclastic–carbonate ramp in northern Tunisia

The marine Messinian deposits of Tunisia cover a narrow littoral strip some 300 km long between the northern Bizerte and Cap Bon areas and the central–eastern Sahel region. Litho- and biofacies analysis of six stratigraphic sections reveals the distinctive features of these deposits.The lower Messinian deposits are characterized by ubiquitous siliciclastics and abundant oolitic/bioclastic limestones organized in an eastward facing ramp. Westward (landward), this ramp changes into coastal lagoons, sometimes containing evaporites. Eastward, the ramp passes to the reefal Pelagian Platform extending as far as Lampedusa.Two main sedimentary cycles are distinguished: 1) an early Messinian siliciclastic retrogradational then oolitic/bioclastic progradational cycle (Beni Khiar Formation and lower Oued bel Khedim Formation); 2) a late Messinian brackish to continental cycle that probably accumulated in rapidly subsiding lagoons (Oued el Bir Formation and upper Oued bel Khedim Formation). The Tunisian early Messinian cycle is partly eustatically controlled, but the late Messinian cycle cannot be confidently correlated to other well-known Messinian series because of tectonic movements.The lower Messinian deposits of Tunisia are also characterized by abundant suspension-feeding organisms (molluscs and bryozoans) and rare corals, calcareous algae, echinoids, and larger benthic foraminifers. The proposed palaeoenvironmental model shows that the lower Messinian ramp of Tunisia was located on a current-protected margin and subjected to continent-derived sediment and nutrient supply. Eastward, nutrient influx diminished and a shallow-water isolated carbonate platform with coralgal facies developed between the western and the eastern Mediterranean basins. The main hydrological connection between these two basins occurred through a narrow seaway situated to the northeast of the Pelagian Platform, south of Sicily and Malta.     

Latest Messinian “Lago-Mare” Lymnocardiinae from Italy: Close relations with the Pontian fauna from the Dacic Basin

Rich hypo- to mesohaline molluscan assemblages characterising the latest Messinian “Lago-Mare” biofacies, composed of prosobranch gastropods (Neritidae, Thiaridae, Melanopsidae, Hydrobiidae) and bivalves of the families Cardiidae (subfamily Lymnocardiinae) and Dreissenidae, are widespread in shallow water basins characterized by low salinities within the Mediterranean realm, during the post-evaporitic phase in the time-span 5.5-5.3 Ma. Several genera and species are recorded in the Italian uppermost Messinian sediments. While the gastropods show endemic character being linked to continental water-systems, Lymnocardiinae and Dreissenidae have strong Paratethyan affinity. New records of significant species of Lymnocardiinae from the uppermost Messinian sediments of Tuscany, Marches and Sicily and the systematic review of the old literature data point out close relations of the Italian fauna with that from the Pontian sediments of the Dacic Basin. The palaeobiogeographical data referred to Messinian and Pontian Lymnocardiinae suggest that the Aegean Basin could be an intermediate basin from whence the Pontian Paratethyan-type fauna migrated into the Mediterranean area during the latest Messinian. Since the ecology of Lymnocardiinae is mainly tied to oligo- and mesohaline water, spreading of suitable habitats in depositional systems of marginal settings characterized by increasing freshwater influx after the “salinity crisis” favoured their dispersal into the Mediterranean area from the Paratethys realm.     

Paracamelus (Mammalia,Camelidae) remains from the late Messinian of Italy: insights into the last camels of western Europe

In this paper we describe fossil remains of an indeterminate species of the genus Paracamelus (Artiodactyla, Camelidae) from the Messinian post-evaporitic deposits (5.55–5.40 Ma) of Verduno (Piedmont, NW Italy). Camelins dispersed into Eurasia from North America in the late Miocene and almost instantaneously spread in western Europe and Africa. The size and morphology of the fossils found at Verduno are consistent in with those of Paracamelus, the earliest Old World camelin. Up to now, the only fossil camels recovered in the Neogene of Western Europe have been found at Venta del Moro and Librilla in Spain at 6.2 Ma. The remains from Verduno represent the first evidence of a camelin in the Neogene of Italy and they considerably expand the paleobiogeographic range of the Old World early camelins. The presence of a camelid at Verduno reinforces and confirms the importance of the fossiliferous deposits of NW Italy in defining the complex paleobiogeographic patterns of Europe during the Messinian, at the end of the Messinian salinity crisis.     

Did the Mediterranean marine reflooding precede the Mio–Pliocene boundary? Paleontological and geochemical evidence from upper Messinian sequences of Tuscany,Italy

The events related to the Messinian salinity crisis are among the most deeply investigated of Earth's history. According to the current hypothesis of Neogene paleogeographic evolution, approximately 5.5 Ma ago, after evaporitic sedimentation and before the Mio–Pliocene boundary, the Mediterranean was characterized by the widespread development of non-marine environments inhabited by molluscs and ostracods of brackish affinity. The Messinian post-evaporitic deposits that testify such a dramatic environmental change are commonly referred to as ‘Lago-mare’ and have been reported from several outcrops and boreholes throughout the entire Mediterranean basin. The origin of ‘Lago-mare’ conditions is commonly interpreted as the result of the synergistic effect of both the humid climatic conditions and change of the drainage patterns at the Mediterranean scale, with the capture of Paratethyan brackish waters. A few recent studies, however, suggest that such a scenario probably represents an oversimplification of the original context, from both a paleogeographical and paleoenvironmental point of view. Unfortunately, the results of these studies have never been commonly accepted and the proposed evidences have been considered questionable. In this paper we describe the fish assemblages from the ‘Lago-mare’ deposits of two localities, Cava Serredi and Podere Torricella, located in the Neogene hinterland basins of Tuscany, central Italy. These assemblages consists of a mixture of marine euryhaline and stenohaline taxa suggesting that the depositional environments were characterized by permanently open connections with a marine basin filled with normal marine waters. In order to better define the paleontological significance of these upper Messinian fish assemblages, the oxygen, carbon and strontium isotopic composition of fish otoliths and other fossils has been measured. These isotopic compositions are strongly indicative of the presence of normal marine conditions close to the depositional environments testified by the sedimentary sequences, thereby implying that the interpretation of the geochemical results are consistent with those derived from the paleoecological analyses of the fish assemblages. Based on the integrated paleoichthyological–geochemical approach discussed in this paper it is possible to unambiguously demonstrate that normal marine rather than fresh- or brackish waters were present in the Mediterranean at least during the upper part of the ‘Lago-mare’ event, providing an unquestionable evidence that the marine refilling of the basin preceded the Mio–Pliocene boundary.     

Coevolution between crossbills and black pine: the importance of competitors,forest area and resource stability

Studies of predator‐prey interactions have found that geographically structured coevolution has played an important role in the adaptive diversification of crossbills (Loxia spp.). We extend those studies by considering common crossbills (L. curvirostra) in the Mediterranean where they rely on seeds in the cones of black pine (Pinus nigra). On the continent, where tree squirrels (Sciurus vulgaris) are present, enhanced defenses against crossbills were most evident in larger areas of pine forest. On islands in the absence of tree squirrels, crossbills and black pine have coevolved in a predator‐prey arms race on Cyprus but not Corsica. In contrast to other conifers that island endemic crossbills rely upon, black pine does not hold seeds in its cones year round. Consequently, key to the strong crossbill–pine interaction on Cyprus is likely the presence of an alternative conifer that provides seeds during early summer when black pine seeds are scarce.     

Integrated stratigraphy and paleoceanography of the Messinian (latest Miocene) across the North Atlantic Ocean

The Messinian was a time of major climatic and paleoceanographic change during the late Cenozoic. It is well known around the Mediterranean region because of the giant anhydrite/gypsum sequence and the suggested desiccation of the Mediterranean Sea. However, this interval is less constrained outside the Mediterranean region, where several paleoceanographic changes could have taken place because of the desiccation. Hence, we present an integrated stratigraphic framework for future Messinian paleoceanographic studies, determination of the effect of the Mediterranean desiccation on deep-water paleoceanography, and comparison of intra-Mediterranean paleoceanographic changes with those in the open oceans during the Messinian Stage.Four DSDP/ODP Holes (552A, 646B, 608, and 547A) from the North Atlantic Ocean and one land borehole from Morocco have been studied to integrate bio-, magneto-, and stable isotope Messinian stratigraphy with high resolution sampling. Our results produce the best assessment of the Tortonian/Messinian boundaries in all holes because they do not rely on any one signal.In paleomagnetic Subchronozone CSAn.lr in the Salé borehole and DSDP Site 609, a S/D coiling direction change of Neogloboquadrina pachyderma/acostaensis appears to indicate PMOW entering the northeastern Atlantic Ocean, at least reaching 50°N. Diachrony and synchrony of some important Messinian planktic foraminifera from these Atlantic DSDP/ODP holes and the Salé borehole, such as the LO of Gq. dehiscens, the LO of Gt. lenguaensis, the FO and LO of Gt. conomiozea, the FO of Gt. margaritae s.s., the FO of Gt. puncticulata, and the FO of Gt. crassaformis are discussed for understanding some of the paleoceanographic changes. This integrated stratigraphie framework presented here allows much better North Atlantic correlations at this critical point in Messinian geologic history.     

The Messinian marine molluscs record and the dawn of the eastern Atlantic biogeography

The biological impact of the Messinian Salinity Crisis (MSC) and its bearing on the Pliocene Mediterranean marine molluscan fauna has been analyzed on the basis of the biogeographic and stratigraphic distributions of the taxa of 16 early Messinian outcrops. The extinction of the last paleoendemic Proto-Mediterranean taxa is historically significant but it appears to be numerically less important than the extinction of the Tortonian and Messinian neoendemic taxa. The available data suggest that the MSC caused a regional mass disappearance but only a limited number of extinctions. It is also emphasized that the Late Pliocene (Monegatti et al., 2002) extinctions were far heavier than those caused by the MSC. We suggest that the greatest Messinian extinctions were triggered, during the salinity crisis, in the Atlantic “sanctuary” by the Messinian glacial events TG22, TG20, TG14, and TG12, of Shackleton et al. (1994), dated between 5.79 and 5.55 Ma by Krijgsman et al. (2004). A comparison between the Mediterranean Messinian and the Redonian molluscan faunas was also carried out. Finally, the possible latitudinal stability of the climatic thresholds, despite the shifting of the climatic zones throughout the Neogene along the European coast, is pointed out.     

Tarbellastraea (Scleractinia): A new stable isotope archive for Late Miocene paleoenvironments in the Mediterranean

Geochemical proxy records of sea surface temperature (SST) or sea surface salinity (SSS) variability on intra- and interannual time-scales in corals from geological periods older than Pleistocene are extremely rare due to pervasive diagenetic alteration of coralline aragonite. Very recently, however, stable isotope data (δ¹⁸O, δ¹³C) from specimens of Porites of Late Miocene age (10 Ma) have been shown to preserve original environmental signatures. In this paper we describe new finds of the zooxanthellate corals Porites and Tarbellastraea in exceptional aragonite preservation from the island of Crete in sediments of Tortonian (～ 9 Ma) and Early Messinian (～ 7 Ma) age. Systematic, comparative stable isotope analysis of massive Tarbellastraea and Porites sampled from the same beds and localities reveal identical stable isotope fractionation patterns in both genera. Therefore, extinct Tarbellastraea represents an additional environmental archive fully compatible and mutually exchangeable with Porites. Provided that seasonal variations in δ¹⁸O reflect SST changes only, seasonal SST contrasts of 7.3 °C for the Tortonian and 4.8 °C for the Early Messinian are inferred, implying warmer summer and cooler winter SSTs during the Tortonian than during the Messinian. However, reduced δ¹⁸O seasonality (1.1‰ in the Tortonian and 0.7‰ in the Messinian) and slightly less negative mean δ¹⁸O in Messinian corals (? 2.4‰) compared to Tortonian specimens (? 2.7‰) may not necessarily indicate a long-term fall in SSTs, but changes in surface water δ¹⁸O, i.e. global ice build-up or enhanced evaporation during summer or increased precipitation/river discharge during winter and changes in insolation. On the other hand, coral communities of Tortonian and Messinian age in central Crete are identical, and compatible annual extension rates indicate similar average SSTs during the two investigated time periods. In addition, lithological and paleobotanical data from Central Crete document a change from humid to dry climatic conditions during the Late Miocene. Therefore, a likely explanation for the observed shift in coral mean δ¹⁸O and reduced δ¹⁸O seasonality from the Tortonian to the Early Messinian is a change in ambient seawater δ¹⁸O caused by a change in the hydrological balance towards high evaporation/high salinity during summer.     

Cyprus wheatears Oenanthe cypriaca likely reach sub‐Saharan African wintering grounds in a single migratory flight

Long‐distance migratory flights with multiple stop‐overs, multiple wintering sites, and small‐scale connectivity in Afro‐Palearctic migrants are likely to increase their vulnerability to environmental change and lead to declining populations. Here we present the migration tracks and wintering locations of the first six Cyprus wheatears to be tracked with geolocators: a species with high survival and a stable population. We therefore predicted a non‐stop flight from Cyprus to sub‐Saharan wintering grounds, a single wintering area for each individual and a wide spread of wintering locations representing low migratory connectivity at the population level. The sub‐Saharan wintering grounds in south Sudan, Sudan and Ethiopia were likely reached by a single flight of an average straight‐line distance of 2538 km in ca 60 h, with an average minimum speed of 43.1 km h^–1. The high speed of migration probably ruled out stop‐overs greater than a few hours. Cyprus wheatears migrated from Cyprus in mid‐late October and most probably remained at a single location throughout winter; three out of five birds with available data may have used a second site < 100 km away during February; all returned between 7–22 March when accurate geolocation data are not possible due to the equinox. Wintering locations were spread over at least 950 km. There were no tag effects on survival. Cyprus wheatears showed a migratory strategy in accordance with their observed high survival rate and demonstrated a routine flight range that allows much of the Mediterranean and the Sahara to be crossed in a rapid two and a half‐day flight.     

Late Miocene paleoenvironmental changes in North Africa and the Mediterranean recorded by geochemical proxies (Monte Gibliscemi section,Sicily)

The astronomically tuned marls of the Monte Gibliscemi section, Sicily, constitute an archive to trace the late Miocene palaeoenvironmental conditions (~ 9.7–7.0 Ma) in North Africa. Here we have utilised carbonate content and Al-normalised geochemical proxies to trace changes in terrigenous source area and bottom-water ventilation. The terrigenous input into the section is dominated by North African river systems draining into the Eastern Mediterranean. The proxy parameters indicate that the palaeoenvironmental conditions in North Africa were humid from 9.5 Ma onward with high fluvial input to the Mediterranean. Increases in the Si/Al and Mg/Al ratios occurred from 8.4 to 8.2 Ma and from 8.05 to 7.75 Ma, with maximum values similar to those of Messinian diatomites. These peaks indicate conditions of enhanced biosiliceous productivity and the presence of authigenic clay formation. Sluggish water circulation in the Mediterranean during those times is inferred from the Mn/Al and V/Al behaviour. Late Miocene changes of the Betic (southern Spain) and Rifian (Morocco) Mediterranean–Atlantic gateways are interpreted as the driving force for the changes in water circulation and the transgression associated with the opening of the Rifian corridor is interpreted to occur at 7.8 Ma.     

Historical biogeography of European leuciscins (Cyprinidae): evaluating the Lago Mare dispersal hypothesis

Aim To test the importance of the Lago Mare stage of the Messinian Salinity Crisis for the dispersal and diversification of European leuciscins (Cyprinidae: Leuciscinae). Location Europe. Methods Cytochrome b sequences of European leuciscins were employed to investigate phylogenetic relationships among species, using Bayesian inference, and to estimate times of diversification, using a relaxed molecular clock. The distributions of 190 European leuciscins were compiled, and regional species compositions were compared using a taxonomic similarity index and an area cladogram. Results Leuciscins restricted to the Iberian and Italian peninsulas and the West and South Balkan regions are phylogenetically more closely related to northern European species than to species from another southern European area. Application of a relaxed molecular clock to a Bayesian phylogeny indicates that most southern clades originated and diversified prior to the Messinian. Southern European regions are taxonomically distinct from one another, and from a more taxonomically homogeneous group of areas that includes Anatolia, East Balkans, Middle East, North Europe and West Russia. Main conclusions The scenario of a Messinian period of dispersal of Paratethyan fauna into Mediterranean regions, via the Lago Mare, predicts a rapid period of diversification and a pattern of close association among southern European faunas. Phylogenetic relationships among leuciscins, the timing of cladogenic events, and the taxonomic similarity among geographical regions do not conform to this expectation. The depth of clades endemic to southern Europe, together with the high levels of endemism in these regions, suggests that the faunas in these regions diverged prior to the Messinian and have evolved largely in isolation from one another. Our results support a model of gradual colonization of Mediterranean regions since the Oligocene. Subsequent connections between adjacent areas may have occurred in the Messinian or Pleistocene.     

Evolutionary dynamics of serpentine adaptation in Onosma (Boraginaceae) as revealed by ITS sequence data

Plant life on serpentine soils has been a topic of research for decades, but the evolutionary dynamics of adaptation to such a stressful habitat is still incompletely understood, especially in old-world groups. We present a study addressing this issue using Onosma (Boraginaceae) as the model system and a molecular phylogenetic approach. Original ITS sequences were generated for all the obligate endemics allopatrically distributed on the ophiolitic ??islands?? of the southeastern Euro-Mediterranean region, in addition to most of the species facultatively growing on ultramafics and a broad sample of non-serpentine species. Parsimony and Bayesian reconstructions showed that obligate endemics belonged to six distantly related clades, five continental and one insular in the Aegean sea (Cyprus). Lack of a common ancestor and of correlation between geographic and genetic distances between the endemics suggested polyphyletic and polytopic evolution on the different outcrops. Preference for non-serpentine habitats appeared as the ancestral condition, but constitutive preadaptive traits such as drought tolerance and ability to cope with high soil concentrations of magnesium have probably favoured multiple events in the colonization of ultramafics. Tree topology and absolute age estimations suggest that xerophytic Onosma underwent a rapid radiation in correspondence with the Messinian salinity crisis of the Mediterranean (6?C5.3?mya), and that the endemic lineages originated at the beginning of the Pleistocene. Serpentine ??islands?? may have acted as refugial habitats during the cold climatic phases, and then as major determinants of adaptive speciation due to isolation of populations and the selective pressure of soil constraints.     

“Earliest Zanclean age for the Colombacci and uppermost Di Tetto formations of the « latest Messinian » northern Apennines: New palaeoenvironmental data from the Maccarone section (Marche Province, Italy)” by Popescu et al. (2007) Geobios 40 (359-373)

Two possible alternative interpretations of the claimed Zanclean age (Popescu et al., 2007) of two historical lithostratigraphic units of the Northern Apennines, usually referred to as Late Messinian in age and recording the so called Lagomare final event of the Messinian salinity crisis (MSC), are here discussed. The wrong age attribution of the Colombacci and “tetto” Fms. is ruled out based on data from the Maccarone and other sections showing that the Colombacci-Argille Azzurre Fm. boundary is basin wide synchronous and coincident with the Miocene-Pliocene boundary as far as it has been formally defined in the Eraclea Minoa GSSP. Alternatively, the opportunity of emending the Zanclean GSSP to a stratigraphically lower horizon recording the first evidence of marine influences in the Mediterranean following the MSC peak, seems not suitable, as (1) the marine signature of uppermost Messinian deposits is weak and still controversial and (2) no significant bio- and magnetostratigraphic events, well chronologically defined and recognizable at a global scale appear to be available to such a purpose.