Higher genetic diversity in recolonized areas than in refugia of Alnus glutinosa triggered by continent‐wide lineage admixture

Genetic admixture is supposed to be an important trigger of species expansions because it can create the potential for selection of genotypes suitable for new climatic conditions. Up until now, however, no continent‐wide population genetic study has performed a detailed reconstruction of admixture events during natural species expansions. To fill this gap, we analysed the postglacial history of Alnus glutinosa, a keystone species of European swamp habitats, across its entire distribution range using two molecular markers, cpDNA and nuclear microsatellites. CpDNA revealed multiple southern refugia located in the Iberian, Apennine, Balkan and Anatolian Peninsulas, Corsica and North Africa. Analysis of microsatellites variation revealed three main directions of postglacial expansion: (i) from the northern part of the Iberian Peninsula to Western and Central Europe and subsequently to the British Isles, (ii) from the Apennine Peninsula to the Alps and (iii) from the eastern part of the Balkan Peninsula to the Carpathians followed by expansion towards the Northern European plains. This challenges the classical paradigm that most European populations originated from refugial areas in the Carpathians. It has been shown that colonizing lineages have met several times and formed secondary contact zones with unexpectedly high population genetic diversity in Central Europe and Scandinavia. On the contrary, limited genetic admixture in southern refugial areas of A. glutinosa renders rear‐edge populations in the Mediterranean region more vulnerable to extinction due to climate change.     

Southern isolation and northern long‐distance dispersal shaped the phylogeography of the widespread,but highly disjunct,European high mountain plant Artemisia eriantha (Asteraceae)

We investigated the range dynamics of Artemisia eriantha, a widespread, but rare, mountain plant with a highly disjunct distribution in the European Alpine System. We focused on testing the roles of vicariance and long‐distance dispersal in shaping the current distribution of the species. To this end, we collected AFLP and plastid DNA sequence data for 17 populations covering the entire distributional range of the species. Strong phylogeographical structure was found in both datasets. AFLP data suggested that almost all populations were genetically strongly differentiated, with 58% of the overall genetic variation partitioned among populations. Bayesian clustering identified five groups of populations: Balkans, Pyrenees, Central Apennines, one southwestern Alpine population and a Widespread cluster (eastern Pyrenees, Alps, Carpathians). Major groups were supported by neighbor‐joining and NeighbourNet analyses. Fourteen plastid haplotypes were found constituting five strongly distinct lineages: Alps plus Pyrenees, Apennines, Balkans, southern Carpathians, and a Widespread group (eastern Pyrenees, northern Carpathians, Mt. Olympus). Plastid DNA data suggested that A. eriantha colonized the European Alpine System in a westward direction. Although, in southern Europe, vicariant differentiation among the Iberian, Italian and Balkan Peninsulas predominated, thus highlighting their importance as glacial refugia for alpine species, in temperate mountain ranges, long‐distance dispersal prevailed. This study emphasizes that currently highly disjunct distributions can be shaped by both vicariance and long‐distance dispersal, although their relative importance may be geographically structured along, for instance, latitude, as in A. eriantha. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 214–226.     

The molecular population structure of the tall forb Cicerbita alpina (Asteraceae) supports the idea of cryptic glacial refugia in central Europe

There is an ongoing debate about the glacial history of non‐arctic species in central and northern Europe. The two main hypotheses are: (1) postglacial colonization from refugia outside this region; (2) glacial survival in microclimatically favourable sites within the periglacial areas. In order to clarify the glacial history of a boreo‐montane tall forb, we analysed AFLPs from populations of Cicerbita alpina through most of its range (Scandinavia, the mountains of central Europe, the Alps, the Pyrenees and the Balkan Peninsula). We found a major differentiation between the Pyrenean population and all others, supported by principal coordinate, neighbour joining and STRUCTURE analyses. Furthermore, three populations from the central and north‐eastern Alps were genetically distinct from the bulk of populations from Scandinavia, central Europe, the Alps and the Balkan Peninsula. Most populations, including those from central and northern Europe, had moderate to high levels of genetic diversity (mean Shannon index H_Sh = 0.292, mean percentage of polymorphic loci P = 54.1%, mean Nei's gene diversity H = 0.195). The results indicate separate glacial refugia in the Pyrenean region and the Italian Alps. Furthermore, they provide evidence of glacial persistence in cryptic refugia north of the Alps, from where Scandinavia and most of the Alps are likely to have been colonized following deglaciation. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 142–154.     

Phytogeography of parasteppic vegetation in the high Friulian Plain (NE Italy)

The vascular flora of pedologically conditioned, extensive dry grasslands occurring in a very rainy area of northeastern Italy, was submitted to phytogeographic analysis. The distributional ranges of 144 species were digitized into two presence-absence matrices of species and Operational Geographic Units (quadrants), one covering Europe, the other, limited to 56 species with extra-European distributions, extended to the Northern Hemisphere north of the Tropic of Cancer. These matrices were submitted to numerical classification, obtaining clusters of species with similar distributional patterns (chorotypes). For each cluster of species, the percent occurrencies in each quadrant were processed by a program of automatic mapping, producing a series of isoporic maps showing the joint distribution of the species of each cluster. The species can be subdivided into six main phytogeographical groups: (1) Narrow-ranging (34% of the total), including the endemic, subendemic, Illyrian-Balcanic and NW submediterranean elements, (2) Southern European-Submediterranean (18.8%), (3) Wide-ranging European (16.6%), (4) Eastern-Pontic (7.6%), (5) Southern Eurasiatic (10.5%), (6) Northern Eurasiatic (12.5%). The phytogeographic originality of the investigated grasslands is high, with more than one third of the species having narrow distributional ranges. Particularly relevant are the connections with the Balkanic-Illyrian and the eastern Alpine regions.Nomenclature: Pignatti (1982).     

Common Distribution Patterns of Marsupials Related to Physiographical Diversity in Venezuela

The aim of this study is to identify significant biotic regions (groups of areas with similar biotas) and biotic elements (groups of taxa with similar distributions) for the marsupial fauna in a part of northern South America using physiographical areas as Operational Geographical Units (OGUs). We considered Venezuela a good model to elucidate this issue because of its high diversity in landscapes and the relatively vast amount of information available on the geographical distribution of marsupial species. Based on the presence-absence of 33 species in 15 physiographical sub-regions (OGUs) we identified Operational Biogeographical Units (OBUs) and chorotypes using a quantitative analysis that tested statistical significance of the resulting groups. Altitudinal and/or climatic trends in the OBUs and chorotypes were studied using a redundancy analysis. The classification method revealed four OBUs. Strong biotic boundaries separated: i) the xerophytic zone of the Continental coast (OBU I); ii) the sub-regions north of the Orinoco River (OBU III and IV); and those south to the river (OBU II). Eleven chorotypes were identified, four of which included a single species with a restricted geographic distribution. As for the other chorotypes, three main common distribution patterns have been inferred: i) species from the Llanos and/or distributed south of the Orinoco River; ii) species exclusively from the Andes; and iii) species that either occur exclusively north of the Orinoco River or that show a wide distribution throughout Venezuela. Mean altitude, evapotranspiration and precipitation of the driest month, and temperature range allowed us to characterize environmentally most of the OBUs and chorotypes obtained.     

Phylogenetic and phylogeographic relationships in the crayfish genus Austropotamobius inferred from mitochondrial COI gene sequences

The present study explores the utility of mitochondrial COI gene sequences to reveal phylogenetic and phylogeographic relationships for the entire European freshwater crayfish genus Austropotamobius. The two traditional taxa, Austropotamobius pallipes and Austropotamobius torrentium, were monophyletic, showing similar genetic diversity, with 28 and 25 haplotypes, respectively, and an uncorrected average pairwise divergence of 0.059 and 0.041. A third distinct haplotype clade, in sister relation to A. torrentium, was discovered at the Upper Kolpa drainage in the northern Dinaric area. All populations north and west of the Alps are genetically impoverished (nucleotide diversity (pi)=0.000-0.001), while southern populations are more diverse (pi=0.001-0.034). A. pallipes reaches the highest diversity in the region of Istra, probably its primary center of radiation. The genetic diversity center for A. torrentium is the southern Balkan peninsula. Other potential glacial refugia were identified in Southern France, Northwestern Italy, the Apennine Peninsula, and in the northern Dinaric area. The Iberian Peninsula has been stocked artificially from Northern Italy. Three main periods of radiation were tentatively identified: late Miocene/early Pliocene for the divergence of species and main lineages, the Pleistocene for the divergence within populations south from Alps, and a postPleistocene expansion north and west from Alps.     

Biogeographical analysis of species of the tribe Cytiseae (Fabaceae) in the Iberian Peninsula and Balearic Islands

Aim To analyse quantitatively the biogeographical distribution pattern of species of the Cytiseae Bercht. & J. Presl (= Genisteae Benth.) tribe in the Iberian Peninsula and the Balearic Islands, and to identify environmental variables related to the distributional patterns. Location Iberian Peninsula and Balearic Islands, using the 61 administrative provinces as operational geographical units. Methods In order to identify chorotypes (groups of species with similar geographical distribution), we performed a upgma classification based on the similarity index of Baroni‐Urbani & Buser. The method of McCoy et al. [Ecology 67 (1986), 749] enabled us to detect the significant groups and to differentiate them from those groupings that could be generated at random. Logistic regression analyses and environmental gradient analyses (DCA and CCA) were performed in order to find the relationships between the environmental variables and the observed distributional patterns. Results Sixteen chorotypes were obtained between the Cytiseae species of the Iberian Peninsula and the Balearic Islands. The thermal variables showed the greatest influence in species distribution. Specifically, temperatures (maximum, minimum and mean) of the coldest months were associated with the majority of the chorotypes. Main conclusions The species of the Cytiseae tribe were not randomly distributed in space, and can be classified in groups of species with common distributional patterns. The importance of cold tolerance in the distribution of these species, as well as their preference for acidic soils, was demonstrated. Certain general tendencies seem to exist with respect to the distribution of the biota in the Iberian Peninsula, and these seem to be independent of the taxonomic groups involved and to be determined by convergent macroclimatic factors.     

Chloroplast DNA phylogeography of European ashes, Fraxinus sp. (Oleaceae): roles of hybridization and life history traits

We investigated range-wide phylogeographic variation in three European ash species (Fraxinus sp., Oleaceae). Chloroplast DNA (cpDNA) microsatellites were typed in the thermophilous Fraxinus angustifolia and Fraxinus ornus and the observed haplotypes and the geographic distribution of diversity were compared to cpDNA data previously obtained in the more cold-tolerant Fraxinus excelsior. We found wide-ranging haplotype sharing between the phylogenetically close F. angustifolia and F. excelsior, suggesting hybridization (i) in common glacial refuges in the Iberian Peninsula, northern Italy, the eastern and/or Dinaric Alps and the Balkan Peninsula, and/or (ii) during postglacial recolonization. The data allowed us to propose additional glacial refuges for F. angustifolia in southern Italy and in Turkey, and populations from the latter region were particularly polymorphic. There was evidence for refuge areas in Italy, the Balkan Peninsula and Turkey for F. ornus, which did not share any single chloroplast haplotype with the other species. In both F. angustifolia and F. ornus, cpDNA diversity (h(S) = 0.027 and h(S) = 0.009, respectively) was lower and fixation levels (G(ST) = 0.964 and G(ST) = 0.983, respectively) higher than in sympatric F. excelsior (h(S) = 0.096, G(ST) = 0.870). These diversity patterns could be due to temperature tolerance or the demographic history.     

Taxonomy and ecology of <Emphasis Type="Italic">Castanea sativa</Emphasis> Mill. forests in Greece

Castanea sativa Mill. is an important species of the Balkan Peninsula with high ecological and economic value. This article contributes to a phytosociological synsystematic approach of the C. sativa plant communities in Greece that covers the southern part of the Balkan Peninsula and describes the ecological parameters involved in their distribution and taxonomy. Phytosociological data have been collected from 14 mountainous areas across continental Greece, which are considered representative of the main forest types of C. sativa in the area. The 14 mountains demonstrate a variety of environmental conditions. Five communities (associations) and seven sub-communities (sub-associations) have been identified and described using TWINSPAN, Corespondence Analysis and the Braun-Blanquet classification scheme. Castanea sativa (chestnut) forests of Greece can be distinguished as three broad groups, defined primarily by their geographical distribution. In northern and central Greece chestnut stands are characterized by their high floristic homogeneity, which is reflected in the identification of only one community per region. Chestnut forests in northern Greece, in particular, exhibit strong floristic similarities with those of the rest of the Balkans. In southern Greece, on the other hand, the southern distribution limit of chestnut in the Balkans, there is a greater floristic diversity between the sampled mountains which is reflected in the presence of three confirmed communities and two sub-associations. Several factors have been identified as important in determining the structure and floristic composition of chestnut forests in Greece. Whilst climate and the grazing regime are influential, the degree of silvicultural management appears to be the most important factor determining the floristic composition of chestnut forests and their long-term sustainability.     

Allopatric divergence and secondary contacts in <Emphasis Type="Italic">Euphorbia spinosa</Emphasis> L: Influence of climatic changes on the split of the species

Euphorbia spinosa, a perennial xerophilous shrub naturally distributed across the Italian peninsula, was selected for examination of the role of the Ligurian Alps and Apennines in glacial survival. The Italian Peninsula is considered to be one of the principal glacial refugia in Europe, but few plant population genetic and phylogeography studies have been undertaken in this region. The combined analysis of chloroplast and nuclear loci (ITS, cpSSR and ISSR) enabled us to detect extensive DNA variation and proved to be a very powerful tool for the reconstruction of the phylogeography. Molecular data support the hypothesis of a long-term separation of the Northwestern (Maritime Alps, Sardinia, Corsica, Northern Apennines) and Southeastern (Southern Apennines and Balkan area) lineages in glacial refugia. The existence of allopatrically fragmented lineages is most probably the result of isolation in different glacial refugia, possibly due to the Last Glacial Maximum cooling and the topographic complexity of the Italian peninsula. The most plausible hypothesis assumes the formation of two migration paths during more recent periods: the first one starting with southward migration and the second one moving northwards. The Central Apennines should be considered the confluence of migration routes radiating from separate refugia according to this hypothesis.     

Phylogeography of a subalpine tall‐herb Ranunculus platanifolius (Ranunculaceae) reveals two main genetic lineages in the European mountains

A phylogeographical analysis of Ranunculus platanifolius, a typical European subalpine tall‐herb species, indicates the existence of two main genetic lineages based on amplified fragment length polymorphism (AFLP) markers. One group comprises populations from the Balkan Peninsula and the south‐eastern Carpathians and the other includes the remaining part of the range of the species, encompassing the western Carpathians, Sudetes, Alps, Pyrenees and Scandinavia. The main phylogeographical break observed in this species runs across the Carpathians and separates the main parts of this range (western and south‐eastern Carpathians), supporting a distinct glacial history of populations in these areas. The high genetic similarity of the Balkan Peninsula and south‐eastern Carpathian populations could indicate a common glacial refugium for these contemporarily isolated areas of species distribution. The western and northern part of the species range displays an additional weak differentiation into regional phylogeographical groups, which could have been shaped by isolation in glacial refugia or even by a postglacial isolation. The observed weak phylogeographical structure could also be linked with ecological requirements, allowing survival along streams in relatively low, forested mountain ranges. © 2013 The Linnean Society of London     

Phytogeography of Italian deciduous oak woods based on numerical classification of plant distribution ranges

Abstract. 358 vascular plant species occurring in the deciduous oak woods of Italy were subdivided into 18 phytogeographical groups (chorotypes) through a numerical classification of their total ranges. The distribution patterns of the chorotypes are illustrated by chorograms, i.e. isopleth maps showing the joint distribution of species in Europe and surroundings regions. Except for Mediterranean species, which occur only occasionally in deciduous forests, the oak-wood flora of Italy consists of four main phytogeographical elements, which have a similar incidence (ca. 25 % each): (a) narrow-ranging, southern European species, (b) southeastern European species of submediterranean woodlands, (c) central-European species, (d) widely distributed European species. The oak woods of Italy differ from their central-European counterparts as to the relatively high floristic diversity, and in the presence of several narrow-ranging taxa, centred in different refugial areas for forest vegetation during the glacial period. There is a relation between the distribution patterns of the chorotypes and their relative frequencies in different communities; the latitudinal and longitudinal extensions of the ranges reflect the thermic and hygric conditions of the communities. The oak woods of northern and southern Italy are phytogeographically well-characterized; the former have a maximum of wide-ranging, central European species, the latter host several narrow-ranging taxa. The woodlands of central Italy have intermediate phytogeographical features, which is one of the main reasons for the difficulties in their syntaxonomical arrangement.     

Phylogeography of two European newt species--discordance between mtDNA and morphology

The newts Triturus vulgaris and Triturus montandoni are sister species that exhibit contrasting levels of intraspecific morphological variation. Triturus vulgaris has a broad Eurasiatic distribution encompassing both formerly glaciated and unglaciated areas and shows substantial morphological differentiation in the southern part of its range, while T. montandoni, confined to the Carpathians, is morphologically uniform. We analysed sequence variation of two mtDNA fragments of the total length of c. 1850 bp in 285 individuals of both species collected from 103 localities. Phylogenetic analysis of 200 unique haplotypes defined 12 major clades, their age estimated at c. 4.5-1.0 million years (Myr). Most of the older clades were found in the southern part of the range, and also in central Europe, mainly in Romania. The distribution of mtDNA clades points to the existence of several glacial refugia, located in the Caucasus region, Anatolia, the Balkan Peninsula, Italy, and more to the north in central Europe. The concordance between mtDNA based phylogeny and the distribution of T. vulgaris subspecies was weak. Triturus montandoni haplotypes did not form a monophyletic group. Instead they were found in six clades, in five of them mixed with T. vulgaris haplotypes, most likely as a result of past or ongoing hybridization and multiple introgression of mtDNA from T. vulgaris to T. montandoni. Patterns of sequence variation within clades suggested long-term demographic stability in the southern groups, moderate and relatively old demographic growth in the populations inhabiting central Europe, and high growth in some of the groups that colonized northern parts of Europe after the last glacial maximum.     

Phylogeography and Pleistocene refugia of the Little Owl Athene noctua inferred from mtDNA sequence data

Pleistocene glaciations greatly affected the distribution of genetic diversity in animal populations. The Little Owl is widely distributed in temperate regions and could have survived the last glaciations in southern refugia. To describe the phylogeographical structure of European populations, we sequenced the mitochondrial cytochrome c oxidase I (COI) and control region (CR1) in 326 individuals sampled from 22 locations. Phylogenetic analyses of COI identified two deeply divergent clades: a western haplogroup distributed in western and northwestern Europe, and an eastern haplogroup distributed in southeastern Europe. Faster evolving CR1 sequences supported the divergence between these two main clades, and identified three subgroups within the eastern clade: Balkan, southern Italian and Sardinian. Divergence times estimated from COI with fossil calibrations indicate that the western and eastern haplogroups split 2.01–1.71 Mya. Slightly different times for splits were found using the standard 2% rate and 7.3% mtDNA neutral substitution rate. CR1 sequences dated the origin of endemic Sardinian haplotypes at 1.04–0.26 Mya and the split between southern Italian and Balkan haplogroups at 0.72–0.21 Mya, coincident with the onset of two Pleistocene glaciations. Admixture of mtDNA haplotypes was detected in northern Italy and in central Europe. These findings support a model of southern Mediterranean and Balkan refugia, with postglacial expansion and secondary contacts for Little Owl populations. Central and northern Europe was predominantly recolonized by Little Owls from Iberia, whereas expansion out of the Balkans was more limited. Northward expansion of the Italian haplogroup was probably prevented by the Alps, and the Sardinian haplotypes remained confined to the island. Results showed a clear genetic pattern differentiating putative subspecies. Genetic distances between haplogroups were comparable with those recorded between different avian species.     

Allopatric distribution,ecology and conservation status of the Pilosella alpicola group (Asteraceae)

The Pilosella alpicola group comprises four morphologically distinct and geographically vicariant alpine taxa. We performed a thorough herbarium revision and literature survey to infer their distributional pattern(s). Pilosella alpicola s.s. occurs in the Alps in two disjunct areas: the Swiss Valais Alps and the Italian Dolomites. Historical records come also from the Austrian Alps (Gurktaler Alps and Hohe Tauern) and from one site from the Alpes Maritimes (Col de Larche), but the localities have not been recently confirmed. Pilosella rhodopea, a Balkan subendemic taxon, is quite widespread in Bulgaria (Stara planina Mts, Rila Mts and Pirin Mts), but is more rare in Albania, Greece and Macedonia. Interestingly, this species has also been recorded at two isolated sites in the Romanian southern Carpathians (the C?p??înii and Cozia Mts). This occurrence underlines the floristic affinities of this part of the Carpathians to the Balkan flora. Only two localities of P. serbica, based on voucher specimens, have been recorded so far; Kopaonik Mts in Serbia and the Prokletije Mts in Montenegro. The records from other ranges are related to P. rhodopea. Pilosella ullepitschii, the detailed distribution of which has already been published, is a Carpathian endemic with its core area of distribution in the western Carpathians (Slovakia and Poland). Three isolated localities are also known in the eastern Carpathians (Nemira Mts) and one locality in the southern Carpathians (Bucegi Mts). The possible causes of disjunctions between and within species ranges are briefly discussed. Based on the distributional data, population sizes and ecology, we evaluate the conservation status of the P. alpicola taxa and propose their inclusion in national Red Lists.     

Genetic legacy from past panmixia: high genetic variability and low differentiation in disjunct populations of the Eastern Large Heath butterfly

Glacial and interglacial cycles of the Pleistocene have led to severe range fluctuations of many species. These range shifts of the past often are reflected by extant genetic signatures. Retractions of distribution areas often have fostered splits into several small and isolated retreats as remnants of the formerly interconnected range. These processes often go in line with losses of intraspecific diversity. By contrast, large and interconnected distribution ranges mostly sustain high levels of genetic variability. The genetic impact of both scenarios strongly depends on the temporal scale. In the present study, we tested the genetic effects of an assumed long‐lasting widespread distribution during glacial periods and more short‐term population retractions to mountain archipelagos during warm stages. We analyzed polymorphic allozymes for individuals of the Eastern Large Heath butterfly, Coenonympha rhodopensis, including major parts of its distribution, such as central Italy and the Balkan Peninsula. Our data show extraordinarily high genetic diversity. The only remarkable genetic split is detectable between the central Apennines (Italy) and the Balkan mountain systems. The populations sampled over seven Balkan mountain systems (Jakupica, Shar Planina, Ossogovo, Pirin, Rila, Rhodopes, and Stara Planina) show low genetic differentiation. This low genetic differentiation and high genetic diversity diverges from the genetic structures frequently found in species with disjunct distributions. We therefore hypothesize that the obtained molecular structure is the product of down‐slope shift during the last cold stage and subsequent expansion over the lowlands of the Balkan Peninsula. The current mountain restriction most probably occurred with the beginning of the postglacial warming, which is too short a time span to be of evolutionary relevance. Therefore, the recent high genetic diversities and low differentiation may still reflect long‐lasting glacial panmixia but not (yet) the recent disjunction. The strong genetic differentiation between the Balkans and Italian Apennines must result from an earlier dispersal process, most probably from the Balkans to Italy. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110; , 281–290.     

Phylogenetic relationships and biogeography of Podarcis species from the Balkan Peninsula, by bayesian and maximum likelihood analyses of mitochondrial DNA sequences

Wall lizards of the genus Podarcis (Sauria, Lacertidae) comprise 17 currently recognized species in southern Europe, where they are the predominant nonavian reptile group. The taxonomy of Podarcis is complex and unstable. Based on DNA sequence data, the species of Podarcis falls into four main groups that have substantial geographic coherence (Western island group, southwestern group, Italian group, and Balkan Peninsula group). The Balkan Peninsula species are divided into two subgroups: the subgroup of P. taurica (P. taurica, P. milensis, P. gaigeae, and perhaps P. melisellensis), and the subgroup of P. erhardii (P. erhardii and P. peloponnesiaca). In the present study, the question of phylogenetic relationships among the species of Podarcis encountered in the Balkan Peninsula was addressed using partial mtDNA sequences for cytochrome b (cyt b) and 16S rRNA (16S). The data support the monophyly of Podarcis and suggest that there are three phylogenetic clades: the clade A (P. taurica, P. gaigeae, P. milensis, and P. melisellensis); the clade B (P. erhardii and P. peloponnesiaca), and the clade C (P. muralis and P. sicula). By examining intraspecific relationships it was found that extant populations of P. erhardii are paraphyletic. Furthermore, subspecies previously defined on the basis of morphological characteristics do not correspond to different molecular phylogenetic clades, suggesting that their status should be reconsidered. The distinct geographic distribution of the major clades of the phylogenetic tree and its topology suggest a spatial and temporal sequence of phylogenetic separations that coincide with some major paleogeographic separations during the geological history of the Aegean Sea. The results stress the need for a reconsideration of the evolutionary history of Balkan Podarcis species and help overcome difficulties that classical taxonomy has encountered at both the species and subspecies level.     

Haplotype richness in refugial areas: phylogeographical structure of <Emphasis Type="Italic">Saxifraga callosa</Emphasis>

This paper illustrates the phylogeographical structure of Saxifraga callosa in order to describe its genetic richness in refugial areas and to reconstruct its glacial history. S. callosa is a species spread throughout south-east France and Italy with a high distribution in the Maritime Alps. Four chloroplast microsatellite and AFLP markers were analyzed in populations of S. callosa. The size variants of all tested loci amount to 11 different haplotypes. Intrapopulational haplotype variation was found in two of the populations analyzed: on the Mt. Toraggio in the Maritime Alps, and in the Apuan Alps. On the other hand, no intrapopulational variation was found in 25 populations, most of which were sampled from isolated areas. Analysis of the haplotype distribution showed that population subdivision across all populations was high (G _ST = 0.899). Moreover, its genetic structure was studied using AMOVA and STRUCTURE analysis. The study legitimated inferred conclusions about the phylogeographical structure of the species and identified centers of diversity. Considerations concerning genetic structure and divergence among three major clades (Maritime Alps, Apuan Alps and Apennines), the patchy distribution of haplotypes, and the high number of private haplotypes support the proposal that S. callosa survived in some refugia within the Italian Peninsula refugium, and that mainly northern populations of refugia were involved in postglacial recolonization.     

Distribution of the Forest Dormouse Dryomys nitedula (Pallas, 1779) (Rodentia, Myoxidae) in Europe

A spot distribution map of Dryomys nitedula in Europe is presented, based on a critical evaluation of published information from the various countries of Europe. In addition, maps which are numbered to identify localities were prepared for regions where Dryomys nitedula distribution was largely unknown until now. The species has a unique distribution, ranging from the Alps in the west to the foothills of the Urals in the east, and from southern Latvia in the north as far south as Calabria and Peloponnisos. The species' reported vertical range is from sea level in the Balkans to the subalpine belt in the Alps. The presence of Dryomys nitedula in Europe can be traced back to the beginning of the Pleistocene, but its recent distributional area does not coincide with climatic factors, nor with the distribution of vegetation communities. Possible competition with Eliomys quercinus is discussed as a factor affecting the peripheral range distribution of Dryomys nitedula.     

Survival and long-term maintenance of tertiary trees in the Iberian Peninsula during the Pleistocene: first record of Aesculus L. (Hippocastanaceae) in Spain

The Italian and Balkan peninsulas have been places traditionally highlighted as Pleistocene glacial refuges. The Iberian Peninsula, however, has been a focus of controversy between geobotanists and palaeobotanists as a result of its exclusion from this category on different occasions. In the current paper, we synthesise geological, molecular, palaeobotanical and geobotanical data that show the importance of the Iberian Peninsula in the Western Mediterranean as a refugium area. The presence of Aesculus aff. hippocastanum L. at the Iberian site at Cal Guardiola (Tarrasa, Barcelona, NE Spain) in the Lower–Middle Pleistocene transition helps to consolidate the remarkable role of the Iberian Peninsula in the survival of tertiary species during the Pleistocene. The palaeodistribution of the genus in Europe highlights a model of area abandonment for a widely-distributed species in the Miocene and Pliocene, leading to a diminished and fragmentary presence in the Pleistocene and Holocene on the southern Mediterranean peninsulas. Aesculus fossils are not uncommon within the series of Tertiary taxa. Many appear in the Pliocene and suffer a radical impoverishment in the Lower–Middle Pleistocene transition. Nonetheless some of these tertiary taxa persisted throughout the Pleistocene and Holocene up to the present in the Iberian Peninsula. Locating these refuge areas on the Peninsula is not an easy task, although areas characterised by a sustained level of humidity must have played an predominant role.