Fossil coniferous wood from the Middle Jurassic of Liaoning Province, China

Silicified coniferous wood was collected from the Lanqi Formation (late Middle Jurassic in age) at Shebudaigou Village, Liaoning Province, China. Three taxa are identified, namely Pinoxylon dacotense Knowlton, Xenoxylon phyllocladoides Gothan, and Araucariopitys sp. Based on these new data, and those of other fossil plants reported previously from the same formation, we consider the climate during the deposition of the Lanqi Formation was subtropical, humid and seasonal. In this respect the Lanqi flora differs from the coeval Shimengou and Longmen floras from North China. The Longmen flora was deposited during more humid, subtropical conditions, while the Shimengou Formation indicates that the climate was warm temperate and dry. Our data would suggest that the Late Jurassic climatic pattern was initiated as early as the late Middle Jurassic.     

Sauropod diversity in the Late Cretaceous of southwestern Europe: The lessons of odontology

Recent discoveries in southern France and northern Spain suggest that the morphology of titanosaurian teeth shows much greater variations that previously thought. It is suggested that the different morphotypes are informative at specific or generic level and that titanosaurian genera may indeed be recognized by their isolated teeth. It is also confirmed that juvenile titanosaurian teeth have a rather uniform, cylindrical morphology. Four different morphotypes are described for the Ibero-Armorican Island in the Late Cretaceous.     

Fossil wood of Keteleerioxylon kamtschatkiense sp. nov. (Pinaceae) from the Cretaceous of the northwestern Kamchatka Peninsula

A new species, Keteleerioxylon kamtschatkiense, is described from the Cretaceous of the northwestern Kamchatka Peninsula on the basis of wood anatomy. Fossil wood showing anatomical characters of the modern genus Keteleeria is described from the Cretaceous of the Russian Far East for the first time.     

Lowermost occurrence of ostracod Cypridea species in East Asia and implications for the non-marine Jurassic/Cretaceous boundary

Cypridea Bosquet, 1852 is a non-marine ostracod genus of the Superfamily Cypridoidea, and the extinct Family Cyprideidae, which achieved high diversity in the Early Cretaceous. This genus plays an important role in the subdivision and correlation of strata, as well as in paleogeographic and paleoclimatic studies. Cypridea species are remarkably abundant and diverse, and are extensively distributed across East Asia. However, the lowermost occurrence of Cypridea species (LOOC) in East Asia is still being debated, varying from the Upper Jurassic to Lower Cretaceous. Here, we aim to conduct a preliminary investigation of the Cypridea-bearing strata of East Asia (China, Mongolia, Korea and Japan), and attempt to update the strata correlation, based on new progress of the ostracod correlations, as well as the progress of other research methods on the relative Jurassic/Cretaceous (J/K) strata of these areas. Results show that the LOOCs in most basins of East Asia are documented in Lower Cretaceous (～Berriasian) strata. However, Cypridea species are extremely rare near the J/K boundary interval but flourish until Valanginian–Hauterivian in most basins of East Asia. Therefore, we propose that the LOOC in each corresponding basin of East Asia can be used as an auxiliary marker of the non-marine J/K boundary. Cypridea species may have travelled first from Africa (Kimmeridgian) to Europe (J/K period), and then East Asia (Early Cretaceous), which resulted in the LOOC in East Asia later than in the other continents.     

Holocene distribution and extinction of the moose (Alces alces, Cervidae) in Central Europe

Moose (Alces alces L.) were among the first large mammals to recolonize Central Europe after the last glaciation. Already during the Allerød they established themselves in most parts of the area. In the early Holocene their distribution range extended from the Pyrenees to Denmark and from Austria to Great Britain and also covered eastern Central Europe where they still occur today. In the Preboreal, the moose slowly vanished from the southwestern parts of its distribution range, leading to its extinction in France and, later, in England. During the Atlantic period, the moose died out in large parts of Denmark and population densities apparently decreased in the rest of Central Europe as well. Around the birth of Christ only relict populations were left in western Central Europe, which finally became extinct in early medieval times. In Thuringia and in the region northeast of the river Elbe as well as in central Poland, some stocks persisted until the high and late Middle Ages. The causes of the gradual extinction in Central Europe during the Holocene are complex. Changes in vegetation, climate and sea-level, the increasing fragmentation of habitat through human activities and hunting were, at different times, important factors. In the recent past, however, moose have repeatedly migrated from the east towards the west. The development of its distribution range since the end of the Second World War as well as experiences with Scandinavian populations show that moose are able to thrive in close proximity to humans and that a future expansion of its distribution range towards the west seems possible.     

New and little-known grylloblattids of the family Geinitziidae (Insecta: Grylloblattida) from the Triassic and Jurassic of Europe,Asia, and South Africa

The fossil history of the family Geinitziidae is reviewed. New taxa are described in the family: Shurabia hissarica, sp. nov. (Lower Jurassic of Tajikistan), Sh. shartegica, sp. nov. (Upper Jurassic of Mongolia), Sh. serrata, sp. nov. and Ginitzia sagulensis, sp. nov. (both Lower Jurassic of Kyrgyzstan). The subfamily Stegopterinae Sharov, 1961 is synonymized under Geinitziinae Handlirsch, 1906. The genera Minesedes Fujiyama, 1973 and Ominea Fujiyama, 1973 (Upper Triassic of Japan) are synonymized under Shurabia Martynov, 1937 and Geinitzia Handlirsch, 1906, respectively. Geinitzia varia Bode, 1953 (Lower Jurassic of Germany) and Fletchizia picturata Riek, 1976 (Upper Triassic of South Africa) are redescribed from their holotypes. F. kapokraalensis Wappler, 2001 and F. aleda Wappler, 2001 (both Upper Triassic of South Africa) are transferred to the genus Shurabia.     

Catastrophic mass extinction and assemblage evolution in planktic foraminifera across the Cretaceous/Paleogene (K/Pg) boundary at Bidart (SW France)

A high-resolution biostratigraphic analysis of planktic foraminifers confirms that the Bidart section at the eastern margin of the Atlantic Ocean exhibits a continuous and complete Cretaceous/Paleogene (K/Pg) transition interval. The biozones and subzones recorded in this section are less expanded than their equivalent in Tunisian sections: El Kef (Global Stratotype Section and Point: GSSP for the K/Pg boundary) and Ellès (auxiliary section), but they are sufficiently thick to allow a detailed analysis of the evolution of the planktic foraminiferal assemblages across the K/Pg transition.Throughout the uppermost 4 m Maastrichtian, the planktic foraminiferal assemblages are highly diversified, containing up to 72 species. These Maastrichtian assemblages are rich in cosmopolitan taxa (70%), dominated by small biserial morphotypes which belong mainly to the genus Heterohelix which coexist with less abundant but highly diverse tropical and subtropical species.The extinction pattern at the Bidart section suggests a sudden catastrophic mass extinction at the K/Pg boundary which affected at least 53 out of 72 species. The species becoming extinct include globotruncanids (e.g. Contusotruncana spp., Globotruncana spp., Globotruncanita spp.) and complex heterohelicids (e.g. Racemiguembelina spp., Pseudotextularia spp., Gublerina spp.). At the Bidart section, only Archaeoglobigerina cretacea disappears 2 m below the K/Pg boundary event. Specimens of 18 small and even tiny Maastrichtian species, are found at the lowermost Danian. Only a few of these species belonging to the genera of Guembelitria, Hedbergella and Heterohelix are considered to be real “Cretaceous survivor species”, whereas the specimens belonging to the rest, are most probably reworked, because they differ in their preservation.Throughout lowermost Danian, the planktic foraminiferal assemblages are dominated by “opportunistic” species of the genus Guembelitria. These opportunists are associated to small and poorly diversified pioneer globigerinids (Palaeoglobigerina spp. and Parvularugoglobigerina spp.). These assemblages became progressively more diversified across the early Danian containing species with cancellate walls (Eoglobigerina spp., Parasubbotina spp., Subbotina triloculinoides and Praemurica spp.) and new taxa of biserial heterohelicids (Woodringina spp. and Chiloguembelina spp.) suggesting a paleoenvironmental recovery.     

Benthic foraminiferal turnover across the Cretaceous/Paleogene boundary at Agost (southeastern Spain): paleoenvironmental inferences

We studied Upper Cretaceous and Lower Paleogene benthic foraminifera from the Agost section (southeastern Spain) to infer paleobathymetrical changes and paleoenvironmental turnover across the Cretaceous/Paleogene (K/P) transition. Benthic foraminifera indicate uppermost bathyal depths at Agost during the Abathomphalus mayaroensis Biochron (from about 400 kyr before the K/P boundary) through the early Plummerita hantkeninoides Biochron (about 120–150 kyr before that boundary). The depth increased to middle bathyal for the remainder of the Cretaceous, and remained so for the Danian part of the studied section (Parasubbotina pseudobulloides Biochron, at least 200 kyr after the K/P boundary). There were no perceivable bathymetrical changes at the K/P boundary, where 5% of the species became extinct, and the species composition of the benthic foraminiferal fauna changed considerably. Below the boundary, infaunal morphogroups constitute up to 65–73% of the faunas. Directly above the boundary, in the black clays of the lower Guembelitria cretacea Biozone, benthic foraminifera are rare. Several opportunistic taxa (e.g. the agglutinant Haplophragmoides sp.) have short peaks in relative abundance, possibly reflecting low-oxygen conditions as well as environmental instability, with benthos receiving food from short-lived, local blooms of primary producers. Above the clays through the end of the studied interval, epifaunal morphogroups dominate (up to 70% of the assemblages) or there is an even mixture or epifaunal and infaunal morphogroups. Infaunal groups do not recover to pre-extinction relative abundances, indicating that the food supply to the benthos did not recover fully over the studied interval (about 200 kyr after the K/P boundary). The benthic foraminiferal faunal changes are compatible with the direct and indirect effects of an asteroid impact, which severely destabilized primary producers and the oceanic food web that was dependent upon them.     

Skin of the Cretaceous mosasaur Plotosaurus: implications for aquatic adaptations in giant marine reptiles

The physical nature of water and the environment it presents to an organism have long been recognized as important constraints on aquatic adaptation and evolution. Little is known about the dermal cover of mosasauroids (a group of secondarily aquatic reptiles that occupied a wide array of predatory niches in the Cretaceous marine ecosystems 92–65 Myr ago), a lack of information that has hindered inferences about the nature and level of their aquatic adaptations. A newly discovered Plotosaurus skeleton with integument preserved in three dimensions represents not only the first documented squamation in a mosasaurine mosasaur but also the first record of skin in an advanced member of the Mosasauroidea. The dermal cover comprises keeled and possibly osteoderm-reinforced scales that presumably contributed to an anterior–posterior channelling of the water flow and a reduction of microturbulent burst activities along the surface of the skin. Thus, hydrodynamic requirements of life in the water might have influenced the evolution of multiple-keeled body scales in advanced mosasauroids.     

New dinosaur tracksites from the Middle Jurassic of Madagascar: ichnotaxonomical,behavioural and palaeoenvironmental implications

Abstract: New dinosaur tracksites are described from the Bajocian–Bathonian Bemaraha Formation of western Madagascar. Two track‐bearing surfaces can be followed over a distance of at least 4 km, suggesting the existence of a hitherto unrecognized megatracksite. The track assemblage is theropod dominated, but sauropod tracks also occur at one site. Qualitative and quantitative analysis of the abundant theropod track material suggests that most, if not all, theropod footprints are attributable to a single trackmaker and are referred to Kayentapus isp. Although this ichnogenus, originally described from the Lower Jurassic of North America, has never been recorded from Gondwana nor from the Middle Jurassic, track morphology strongly suggests this attribution. Palaeogeographical, sedimentological and ichnological data suggest that the dinosaur tracks formed in an intertidal to supratidal setting where the coastline influenced the preferred walking direction of the animals.     

Snipe flies (Diptera: Rhagionidae) from the Daohugou Formation (Jurassic), Inner Mongolia,and the systematic position of related records in China

Abstract: Four new species referred to three new genera and one known genus of fossil snipe flies (Family Rhagionidae) from the Daohugou Formation of Chifeng City in Inner Mongolia, north‐eastern China, are described as Daohugorhagio elongatus gen. et sp. nov., Parachrysopilus jurassicus gen. et sp. nov., Sinorhagio sinuatus sp. nov. and Trichorhagio gregarius gen. et sp. nov. These taxa represent only a fraction of the rhagionid assemblage in the Daohugou biota. They contribute towards the high diversity and abundance of snipe flies during the latest Middle Jurassic – earliest Late Jurassic. The diagnosis of Sinorhagio K. Zhang, Yang and Ren, 2006 is revised based on the new results. Many genera and species from the Lower Cretaceous of China have been assigned to Rhagionidae. Most of these do not actually belong to this family: specifically Mesorhagiophryne incerta Hong and Wang, 1990, M. robusta Hong and Wang, 1990, Mesostratiomyia laiyangensis Hong and Wang, 1990, Stratiomyopsis robusta Hong and Wang, 1990, Longhuaia orientalis Hong, Wang and Sun, 1992, Basilorhagio venusius Ren, 1995. Pauromyia oresbia Ren, 1998 is transferred to an archisargid genus as Sharasargus oresbius (Ren, 1998) comb. nov.     

The petrosal and inner ear of the Late Jurassic cladotherian mammal Dryolestes leiriensis and implications for ear evolution in therian mammals

Dryolestes leiriensis is a Late Jurassic fossil mammal of the dryolestoid superfamily in the cladotherian clade that includes the extant marsupials and placentals. We used high resolution micro‐computed tomography (µCT) scanning and digital reconstruction of the virtual endocast of the inner ear to show that its cochlear canal is coiled through 270°, and has a cribriform plate with the spiral cochlear nerve foramina between the internal acoustic meatus and the cochlear bony labyrinth. The cochlear canal has the primary bony lamina for the basilar membrane with a partially formed (or partially preserved) canal for the cochlear spiral ganglion. These structures, in their fully developed condition, form the modiolus (the bony spiral structure) of the fully coiled cochlea in extant marsupial and placental mammals. The CT data show that the secondary bony lamina is present, although less developed than in another dryolestoid Henkelotherium and in the prototribosphenidan Vincelestes. The presence of the primary bony lamina with spiral ganglion canal suggests a dense and finely distributed cochlear nerve innervation of the hair cells for improved resolution of sound frequencies. The primary, and very probably also the secondary, bony laminae are correlated with a more rigid support for the basilar membrane and a narrower width of this membrane, both of which are key soft‐tissue characteristics for more sensitive hearing for higher frequency sound. All these cochlear features originated prior to the full coiling of the therian mammal cochlea beyond one full turn, suggesting that the adaptation to hearing a wider range of sound frequencies, especially higher frequencies with refined resolution, has an ancient evolutionary origin no later than the Late Jurassic in therian evolution. The petrosal of Dryolestes has added several features that are not preserved in the petrosal of Henkelotherium. The petrosal characters of dryolestoid mammals are essentially the same as those of Vincelestes, helping to corroborate the synapomorphies of the cladotherian clade in neural, vascular, and other petrosal characteristics. The petrosal characteristics of Dryolestes and Henkelotherium together represent the ancestral morphotype of the cladotherian clade (Dryolestoidea + Vincelestes + extant Theria) from which the extant therian mammals evolved their ear region characteristics. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 433–463.     

Crocodylomorph Trackways from the Jurassic to Early Cretaceous of North America and Europe: Implications for Ichnotaxonomy

Trackways described as Batrachopus (Batrachopodidae Lull, 1904 Lull, R. S. 1904. Fossil footprints of the Jura-Trias of North America. Memoirs of the Boston Society of Natural History, 5: 461–557.  [Google Scholar]) from the Lower Jurassic of Europe are rare and in some cases different from the type trackways from North America. Differences may be in part attributable to preservation, but current evidence suggests that there is inherent variability in Batrachopodidae morphotypes, beyond that attributable to differential preservation. Type Batrachopus is a stout-toed form, with minimal digit divarication (i.e., a long foot), whereas Antipus describes slender-toed forms with a wider foot and wider digit divarications.

Antipus is also similar to Crocodylopodus (ichnofamily Crocodylopodidae: Fuentes Vidarte and Meijide Calvo, 1999 Fuentes Vidarte, C. and Meijide Calvo, M. 1999. Primeras Huellas de Cocodrilo en el Weald de Cameros (Sria, Espana) Nueva Familia Crocodilopodidae: Nuevo icnogenero: Crocodylopodus Nueva icnoespecie: C. meijidei. Actas de las jornadas internacionales sobre paleontologia de dinosairios y su entorno. Sala de los infantes (Burgos, Espana). Collectivo Arqueologico-Paleontologico de Salas, : 329–338.  [Google Scholar]) from near the Jurassic-Cretaceous boundary in Spain. Crocodylopodus has a relatively large manus, and a less outwardly rotated trackway, but is not sufficiently different from Batrachopodidae to warrant its own ichnofamily. Manus-pes size (area) ratios (heteropody) may also be important in differentiating different crocodylomorph ichnotaxa, as is the case with other archosaurian ichnotaxa. However, heteropody may change with size, and be less pronounced in large individuals. Manus and pes rotation patterns, and trackway width are variable and may be of use for differentiation of ichnotaxa but may also be a function of speed.     

Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia

Atoposaurids are a group of small‐bodied, extinct crocodyliforms, regarded as an important component of Jurassic and Cretaceous Laurasian semi‐aquatic ecosystems. Despite the group being known for over 150 years, the taxonomic composition of Atoposauridae and its position within Crocodyliformes are unresolved. Uncertainty revolves around their placement within Neosuchia, in which they have been found to occupy a range of positions from the most basal neosuchian clade to more crownward eusuchians. This problem stems from a lack of adequate taxonomic treatment of specimens assigned to Atoposauridae, and key taxa such as Theriosuchus have become taxonomic ‘waste baskets’. Here, we incorporate all putative atoposaurid species into a new phylogenetic data matrix comprising 24 taxa scored for 329 characters. Many of our characters are heavily revised or novel to this study, and several ingroup taxa have never previously been included in a phylogenetic analysis. Parsimony and Bayesian approaches both recover Atoposauridae as a basal clade within Neosuchia, more stemward than coelognathosuchians, bernissartiids, and paralligatorids. Atoposauridae is a much more exclusive clade than previously recognized, comprising just three genera (Alligatorellus, Alligatorium, and Atoposaurus) that were restricted to the Late Jurassic of western Europe, and went extinct at the Jurassic/Cretaceous boundary. A putative Gondwanan atoposaurid (Brillanceausuchus) is recovered as a paralligatorid. Our results exclude both Montsecosuchus and Theriosuchus from Atoposauridae. Theriosuchus is polyphyletic, forming two groupings of advanced neosuchians. Theriosuchus (restricted to Theriosuchus pusillus, Theriosuchus guimarotae, and Theriosuchus grandinaris) spanned the Middle Jurassic to early Late Cretaceous, and is known from Eurasia and North Africa. Two Cretaceous species previously assigned to Theriosuchus (‘Theriosuchus’ ibericus and ‘Theriosuchus’ sympiestodon) are shown to be nested within Paralligatoridae, and we assign them to the new genus Sabresuchus. The revised phylogenetic placement of Theriosuchus has several implications for our understanding of eusuchian evolution. Firstly, the presence of fully pterygoidean choanae, previously regarded as a defining characteristic of Eusuchia, is not found in some basal members of Eusuchia. However, eusuchians can be distinguished from Theriosuchus and other basal neosuchians in that their choanae are posteriorly positioned, with an anterior margin medial to the posterior edge of the suborbital fenestra. This feature distinguishes eusuchians from Theriosuchus and more basal neosuchians. Secondly, our refined understanding of Theriosuchus implies that this taxon possessed only amphicoelous presacral vertebrae, and therefore fully developed vertebral procoely is likely to have evolved only once in Crocodylomorpha, on the lineage leading to Eusuchia. These and other findings presented herein will provide an important framework for understanding the neosuchian–eusuchian transition.     

Toarcian carbon isotope shifts and nutrient changes from the Northern margin of Gondwana (High Atlas, Morocco, Jurassic): Palaeoenvironmental implications

The Early Toarcian is marked by a global perturbation of the carbon cycle and major marine biological changes. These coincide with a general decrease in calcium carbonate production and an increase in organic carbon burial, and culminate in the so-called Toarcian Oceanic Anoxic Event. It is believed that the environmental crisis was triggered by the activity of the Karoo-Ferrar large igneous province. In order to further document the Early Toarcian palaeoenvironmental perturbations, carbon isotope, total organic matter, calcareous nannofossils and phosphorus content of the Amellago section in the High Atlas rift basin of Morocco were investigated. This section is extremely expanded compared to the well-studied European sections. Its position along the northern margin of the Gondwana continent is of critical importance because it enables an assessment of changes of river nutrient input into the western Tethyan realm. The carbon isotope curve shows two negative excursions of equal thickness and amplitude, at the Pliensbachian-Toarcian boundary and at the transition from the Polymorphum to the Levisoni Zone. This confirms the supra-regional nature of these shifts and highlights the possible condensation of the first “boundary” shift in European sections. Phosphorus content is used to trace palaeo-nutrient changes and shows that the two negative carbon isotope shifts are associated with increased nutrient levels, confirming that these episodes are related to enhanced continental weathering, probably due to elevated greenhouse gases in the atmosphere. In the High Atlas Basin, the increase in nutrient levels at the Pliensbachian-Toarcian boundary is moreover likely to be the main factor responsible for the coeval demise of the Saharan carbonate platform. A middle Toarcian event, centered on the boundary between the Bifrons and Gradata Zones, characterized by a positive carbon isotope excursion and nutrient level rise, is documented in the Amellago section.     

Diversity patterns amongst herbivorous dinosaurs and plants during the Cretaceous: implications for hypotheses of dinosaur/angiosperm co‐evolution

Abstract Palaeobiologists frequently attempt to identify examples of co‐evolutionary interactions over extended geological timescales. These hypotheses are often intuitively appealing, as co‐evolution is so prevalent in extant ecosystems, and are easy to formulate; however, they are much more difficult to test than their modern analogues. Among the more intriguing deep time co‐evolutionary scenarios are those that relate changes in Cretaceous dinosaur faunas to the primary radiation of flowering plants. Demonstration of temporal congruence between the diversifications of co‐evolving groups is necessary to establish whether co‐evolution could have occurred in such cases, but is insufficient to prove whether it actually did take place. Diversity patterns do, however, provide a means for falsifying such hypotheses. We have compiled a new database of Cretaceous dinosaur and plant distributions from information in the primary literature. This is used as the basis for plotting taxonomic diversity and occurrence curves for herbivorous dinosaurs (Sauropodomorpha, Stegosauria, Ankylosauria, Ornithopoda, Ceratopsia, Pachycephalosauria and herbivorous theropods) and major groups of plants (angiosperms, Bennettitales, cycads, cycadophytes, conifers, Filicales and Ginkgoales) that co‐occur in dinosaur‐bearing formations. Pairwise statistical comparisons were made between various floral and faunal groups to test for any significant similarities in the shapes of their diversity curves through time. We show that, with one possible exception, diversity patterns for major groups of herbivorous dinosaurs are not positively correlated with angiosperm diversity. In other words, at the level of major clades, there is no support for any diffuse co‐evolutionary relationship between herbivorous dinosaurs and flowering plants. The diversification of Late Cretaceous pachycephalosaurs (excluding the problematic taxon Stenopelix) shows a positive correlation, but this might be spuriously related to poor sampling in the Turonian–Santonian interval. Stegosauria shows a significant negative correlation with flowering plants and a significant positive correlation with the nonflowering cycadophytes (cycads, Bennettitales). This interesting pattern is worthy of further investigation, and it reflects the decline of both stegosaurs and cycadophytes during the Early Cretaceous.     

Rangewide phylogeography of a terrestrial slug in Europe: evidence for Alpine refugia and rapid colonization after the Pleistocene glaciations

Intraspecific phylogeographical patterns largely depend on the life history traits of a species. Especially species with a high degree of cold tolerance, limited requirements towards habitat preferences, and relatively low active dispersal capacities may have responded in a different way to the Pleistocene climatological fluctuations than the majority of taxa studied so far. To evaluate this possibility, we studied Arion fuscus (Muller, 1774), a common and widespread European terrestrial slug, from 88 locations (N = 964). Sequence variation was assessed for fragments of the mitochondrial 16S rDNA and COI genes by means of single-strand conformation polymorphisms (SSCP) and subsequent DNA sequencing. Additionally, eight allozyme loci were scored in 843 individuals. Phylogenetic analysis revealed the presence of two major evolutionary lineages, one in the Balkan region and another in the Alps and the rest of Europe. The sequence divergence between the two lineages was limited (3.3%), but gene flow between the regions was absent, suggesting that the two regions have been isolated since the late Pliocene or early Pleistocene. Allozyme differentiation among geographical regions and mitochondrial DNA (mtDNA) lineages was low. The geographical patterns observed in our data showed that (i) haplotype and nucleotide diversities are very low in northern Europe, suggesting that single haplotypes rapidly colonized large areas; (ii) recently expanded haplotype clades have restricted distribution ranges, suggesting that current gene flow is low; and (iii) genetic diversity in the Alps is much higher than in other regions and estimated past gene flow from the Eastern Alps to other regions was high, suggesting that this was a refugial zone during the Pleistocene. This full-range phylogeography suggests the existence of an alternative refugial zone, situated north of the refugial areas currently recognized in most other taxa.