Dispersal vs. vicariance in the Mediterranean: historical biogeography of the Palearctic Pachydeminae (Coleoptera, Scarabaeoidea)

Aim The geological evolution of the Mediterranean region is largely the result of the Tertiary collision of the African and Eurasian Plates, but also a mosaic of migrating island arcs, fragmenting tectonic belts, and extending back‐arc basins. Such complex paleogeography has resulted in a ‘reticulate’ biogeographical history, in which Mediterranean biotas repeatedly fragmented and merged as dispersal barriers appeared and disappeared through time. In this study, dispersal‐vicariance analysis (DIVA) is used to assess the relative role played by dispersal and vicariance in shaping distribution patterns in the beetle subfamily Pachydeminae Reitter, 1902 (Scarabaeoidea), an example of east–west Mediterranean disjunction. Location The Mediterranean region, including North Africa, the western Mediterranean, Balkans–Anatolia, Middle East, Caucasus, the Iranian Plateau, and Central Asia. Methods A phylogenetic hypothesis of the Palearctic genera of Pachydeminae in conjunction with distributional data was analysed using DIVA. This method reconstructs the ancestral distribution in a given phylogeny based on the vicariance model, while allowing dispersal and extinction to occur. Unlike other methods, DIVA does not enforce area relationships to conform to a hierarchical ‘area cladogram’, so it can be used to reconstruct ‘reticulate’ biogeographical scenarios. Results Optimal reconstructions, requiring 23 dispersal events, suggest that the ancestor of Pachydeminae was originally present in the south‐east Mediterranean region. Basal splitting within the subfamily was caused by vicariance events related to the late Tertiary collision of the African microplates Apulia and Arabia with Eurasia, and the resultant arise of successive dispersal barriers (e.g. the Red Sea, the Zagros Mountains). Subsequent diversification in Pachydeminae involved multiple speciation events within the Middle East and Iran–Afghanistan regions, which gave rise to the least speciose genera of Pachydeminae (e.g. Otoclinius Brenske, 1896). Finally, the presence of Pachydeminae in the western Mediterranean region seems to be the result of a recent dispersal event. The ancestor of the Iberian genera Ceramida Baraud, 1987 and Elaphocera Gené, 1836 probably dispersed from the Middle East to the Iberian Peninsula across North Africa and the Gibraltar Strait during the ‘Messinian salinity crisis’ at the end of the Miocene. Main conclusions Although the basal diversification of Pachydeminae around the Mediterranean appears to be related to vicariance events linked to the geological formation of the Mediterranean Basin, dispersal has also played a very important role. Nearly 38% of the speciation events in the phylogeny resulted from dispersal to a new area followed by allopatric speciation between lineages. Relationships between western and eastern Mediterranean disjuncts are usually explained by dispersal through Central Europe. The biogeographical history of the Pachydeminae corroborates other biogeographical studies that consider North Africa to be an alternative dispersal route by which Mediterranean taxa could have achieved circum‐Mediterranean distributions.     

L'intensité lumineuse,son influence sur la teneur en phosphore des carapaces d'Ostracodes

Actual and fossil shells of ostracodes have beenanalyzed with an electronic microprobe: all of them contain phosphorus. X—pictures teach us that the repartition of this chemical element is homogeneous in all the points of the shell. There is no relationship between phosphorus contents in the shells of ostracodes and salinity. In return, there is a relationship between phosphorus contents and temperature and, chiefly, organic phosphorus contents in the environment. However, it is interesting to observe that the analyzed shells of fresh-water ostracodes contain less phosphorus that the generality of the shells of marine ostracodes. The contents of organic phosphorus, in the biotope, depends on the growth of phytoplancton in the upper layers of water which needs a minimum light intensity.     

History of West Mediterranean newts,Pleurodeles (Amphibia: Salamandridae), inferred from old and recent DNA sequences

MtDNA sequences (396 bp cytochrome b and 369 bp 12S rRNA) from recent material and old museum specimens indicate Pleurodeles poireti and P. waltl form independent clades with 7.76% genetic divergence. Within P. poireti, populations from Djebel Edough, NE Algeria are very distinct with 6.12% genetic divergence from the remainder and may deserve separate species status. Away from Djebel Edough, P. poireti consists of three distinct clades (coastal NW Tunisia; central N Algeria; Constantine plus inland NW Tunisia) with a maximum genetic divergence of only 1%. P. waltl contains two clades with 2.96% genetic divergence, one in SE and E Spain plus north Morocco, the other in Portugal and SW and central Spain. Pleurodeles probably invaded NW Africa from SW Europe during the Messinian Salinity Crisis, when land contact was first established at 5.6 Ma, and then interrupted at 5.3 Ma. Molecular clocks, calibrated in the assumption that the latter event separated P. waltl and P. poireti, suggest that Pleurodeles diverged from its sister taxon, Tylototriton, at about 8.6–10 Ma. Djebel Edough P. poireti separated at about 4.2 Ma, perhaps through isolation on a temporary, now ‘fossil’, island initiated by the Messinian crisis. Differentiation in remaining P. poireti may have been caused by Pleistocene climatic fluctuations, while bifurcation in P. waltl appears to have taken place in the Pliocene approximately between 3.2 and 2 Ma. This species reached Morocco very recently, perhaps as a result of human introduction. Use in Pleurodeles of the slower divergence rates estimated in some other salamandrids results in a less parsimonious historical hypothesis that does not fit known geophysical events.     

Messinian‐Zanclean vegetation and climate in North‐Central Italy

A palynological study was carried out on four Italian Miocene‐Pliocene sections ranging in age from uppermost Tortonian to Zanclean located on the Adriatic side of the North‐Central Apennines. The study documents the Mediterranean isolation, the salinity crisis (s.s), the “lago‐mare”; event and the re‐establishment of open‐marine conditions in the Mediterranean at the beginning of the Pliocene. From a climatic point of view, a transition from subtropical/warm‐temperate conditions during the Messinian to warm‐temperate/temperate conditions during the Zanclean is recorded. The presence of a lower thermic level, with respect to the Messinian, the re‐establishment of open‐marine conditions and the uplift of the Apennines were major factors controlling paleoenvironmental variations during the Zanclean. The latter is also characterized by cyclic temperature oscillations. Correlations with coeval sections in the Mediterranean area confirm the existence of latitudinal climatic gradients within the studied area.     

The influence of the Miocene Mediterranean desiccation on the geographical expansion and genetic variation of Androcymbium gramineum (Cav.) McBride (Colchicaceae)

Chloroplast DNA (cpDNA) restriction site and isozyme data were combined to explore the spatial-temporal influence of the Messinian desiccation in the Mediterranean on the disjunct distribution of Androcymbium gramineum in Almería and Morocco (north and south of the straits of Gibraltar, respectively). Lack of evidence for different selective pressures, divergence time estimates based on the calibration of the isozyme molecular clock with the cpDNA data, the basal position of Almerian populations in the A. gramineum clade, and the much higher isozyme polymorphism in Almería suggest that (i) only a southern European range of A. gramineum existed before the Messinian [ approximately 11.2 million years ago (Ma), in the middle Miocene] and (ii) the desiccation of the Mediterranean basin about 5.5-4.5 Ma induced the migration of A. gramineum from Almería to Morocco (between 4.9 and 4.6 Ma, according to our time estimates). After the split into two allopatric units following the refilling of the Mediterranean, the major influence of drift associated with Plio-Pleistocene recurrent glaciation cycles and range expansions/contractions probably fostered the substantial interpopulation genetic differentiation observed within Almería (CGST = 0.41, average DNei = 0.185) and, to a lesser extent, within Morocco (CGST = 0.24, average DNei = 0.089), but did not hinder the maintenance of considerable levels of genetic variation in either geographical area (A = 2.14, HE = 0.230 and A = 1.90, HE = 0.213, respectively).     

The east‐west‐north colonization history of the Mediterranean and Europe by the coastal plant Carex extensa (Cyperaceae)

Coastal plants are ideal models for studying the colonization routes of species because of the simple linear distributions of these species. Carex extensa occurs mainly in salt marshes along the Mediterranean and European coasts. Variation in cpDNA sequences, amplified fragment length polymorphisms (AFLPs) and simple sequence repeats (SSRs) of 24 populations were analysed to reconstruct its colonization history. Phylogenetic relationships indicate that C. extensa together with the South American Carex vixdentata and the southern African Carex ecklonii form a monophyletic group of halophilic species. Analyses of divergence times suggest that early lineage diversification may have occurred between the late Miocene and the late Pliocene (Messinian crisis). Phylogenetic and network analyses of cpDNA variation revealed the monophyly of the species and an ancestral haplotype contained in populations of the eastern Mediterranean. The AFLP and SSR analyses support a pattern of variation compatible with these two lineages. These analyses also show higher levels of genetic diversity and differentiation in the eastern population group, which underwent an east‐to‐west Mediterranean colonization. Quaternary climatic oscillations appear to have been responsible for the split between these two lineages. Secondary contacts may have taken place in areas near the Ligurian Sea in agreement with the gene flow detected in Corsican populations. The AFLP and SSR data accord with the ‘tabula rasa’ hypothesis in which a recent and rapid colonization of northern Europe took place from the western Mediterranean after the Last Glacial Maximum. The unbalanced west‐east vs. west‐north colonization may be as a result of ‘high density blocking’ effect.     

Paleoenvironmental conditions preceding the Messinian Salinity Crisis in the Central Mediterranean: Integrated data from the Upper Miocene Trave section (Italy)

Integrated data of calcareous plankton and benthic foraminifers from the pre-evaporitic interval of Trave section (Central Italy) allowed the reconstruction of surface and bottom-water conditions in the Central Mediterranean during the interval from 7.61 to 6.33 Ma, preceding the Messinian Salinity Crisis.Our data point out a three-step paleoenvironmental evolution. During the first stage (7.61-7.02 Ma) benthic foraminiferal assemblages depict stable, well-oxygenated and ventilated bottom-water conditions, while the surface water records variable temperature and high nutrient conditions, probably associated with strong seasonality. The second stage (7.02-6.70 Ma) points to unfavourable bottom-water condition, triggered by deep-sea stagnation. This is witnessed by a significant decrease in oxygen concentration and biotic diversity, and by the presence of stress-tolerant taxa. A general warming of the surface water and a strongly stratified water column, characterized by an expanded mixed layer, are also recorded.From 6.70 Ma onwards (third stage), a prominent change to more restricted, low-oxygenated, hypersaline conditions at the sea floor is testified by the total disappearance of deep-dwelling planktonic foraminifers and the increasing abundance of stress-tolerant species. Calcareous plankton reflects high instability of the surface water in terms of nutrients, temperature and salinity. During this stage the environmental deterioration reaches intermediate depths in the water column.The initial change toward a step-wise isolation of the Central Mediterranean bottom-waters is probably related to a general warming, responsible for a first slowing-down of the vertical circulation, favouring stratification of surface and intermediate waters and stagnation of bottom-waters. This warming is related to the restricted connection between the Mediterranean Sea and the Atlantic Ocean, which occurred since 7.146 Ma.In the Trave section, the isolation of bottom-waters most likely occurred at the same time as in other Mediterranean sections. However, due to the presence of a hiatus it cannot be excluded that it occurred with a delay of ~ 100 kyr, probably related to the shallower paleodepth of the basin.     

Messinian Lago-Mare mollusc fauna from the Gorgona Island slope, Tyrrhenian Sea

Late Miocene Lago-Mare macrofossiliferous sediments were recovered in the northeastern Tyrrhenian Sea by dredging the continental slope off Gorgona Island, Tuscan Archipelago, at 300-470 m depth. The fossil assemblage consists of a rich lymnocardiid bivalve fauna dominated by Pontalmyra ex gr. P. incerta (Deshayes), associated with Dreissena ex gr. D. rostriformis (Deshayes), Pontalmyra cf. partschi (Mayer), “Limnocardium” sp., the gastropods Melanopsis narzolina (D’Archiac), Melanopsis sp. and cf. Saccoia sp. All bivalve taxa recognized at species level are of Paratethyan (Pontian) affinity and widespread in the Late Miocene of the Mediterranean Basin while M. narzolina has so far only recorded from the Mediterranean Basin. This finding represents the most diverse Lago-Mare macrofauna reported thus far from any submerged location in the Mediterranean Basin and documents that the post-evaporitic Cusercoli Formation contributes to the syn-rift neoautochthonous units of this sector of the Northern Tyrrhenian Sea.     

Messinian Prolagus (Ochotonidae, Lagomorpha) of Italy

A systematic revision of Italian Messinian Prolagus findings has been performed, evidencing the presence of four different species. Comparative analyses allowed their separation in two groups corresponding to two different palaeobioprovinces characterized by distinct faunal affinities, source areas and tempos and modes of colonization. Such considerations allowed to cast a new light on the palaeobiogeography and the migration pathways of continental vertebrate faunas in the Mediterranean area during Messinian, and to give a new importance to the genus Prolagus as a palaeobiogeographical marker.     

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.