The Balearic Islands: a reservoir of cpDNA genetic variation for evergreen oaks

Aim To analyse the role of the Balearic Islands as a refuge area for evergreen Quercus (cork oak: Quercus suber L., holm oak: Q. ilex L., kermes oak: Q. coccifera L.), by using molecular, historical and palaeobotanical data. Location The Western Mediterranean Basin (Balearic Islands, eastern Iberia, Provence, Sardinia, Corsica, Sicily, Malta, Italy, Northern Africa). Methods We sampled 108 populations and used the PCR‐RFLP technique with five universal cpDNA primers to define haplotypes in the sampled populations. Diversity, differentiation parameters and spatial analysis of the populations, using a spatial version of amova , were linked to the geological history of the Western Mediterranean Basin in order to explain the present spatial pattern of the evergreen Quercus populations in the Balearics. Results Evergreen Quercus cpDNA shows a complex structure, with remnants of ancient diversity in the Balearics. Balearic populations of holm oak are related to Iberian populations, while for cork and kermes oaks, we found both Tyrrhenian and Iberian haplotypes. Main conclusions The complex spatial patterns of cpDNA in Balearic evergreen Quercus appears explicable in terms of a combination of physical (vicariance and long distance dispersal) and biological (introgressive hybridization) factors. The Balearics constitute a glacial refuge area and a reservoir of genetic variation with traces of ancient diversity from Messinian–Pliocene stages.     

Extensive gene flow blurs phylogeographic but not phylogenetic signal in Olea europaea L.

Genetic structure and evolutionary patterns of the wild olive tree (Olea europaea L.) were investigated with AFLP fingerprinting data at three geographic levels: (a) phylogenetic relationships of the six currently recognized subspecies in Eurasia and Africa; (b) lineage identification in subsp. europaea of the Mediterranean basin; and (c) phylogeography in the western Mediterranean. Two statistical approaches (Bayesian inference and analysis of molecular variance) were used to analyse the AFLP fingerprints. To determine the congruency and transferability of results across studies previous RAPD and ISSR data were analysed in a similar manner. Comparisons proved that qualitative results were mostly congruent but quantitative values differed, depending on the method of analysis. Neighbour-Joining analysis of AFLP phenotypes supported current classification of subspecies. At a Mediterranean scale no clear cut phylogeographic pattern was recovered, likely due to extensive gene flow between populations of subsp. europaea. Gene flow estimates calculated with conventional F-statistics showed that reproductive barriers separated neither populations nor lineages of O. europaea. Genetic divergence between eastern and western parts of the Mediterranean basin was observed only when geographical and population information were incorporated into the analyses through hierarchical analysis of molecular variance (AMOVA). Within the western Mediterranean, the highest genetic diversity was found in two regions: on both sides of the Strait of Gibraltar and in the Balearic archipelago. Additionally, long-lasting isolation of the northern-most populations of the Iberian Peninsula appeared to be responsible for a significant divergence.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Matching genetics with oceanography: directional gene flow in a Mediterranean fish species

Genetic connectivity and geographic fragmentation are two opposing mechanisms determining the population structure of species. While the first homogenizes the genetic background across populations the second one allows their differentiation. Therefore, knowledge of processes affecting dispersal of marine organisms is crucial to understand their genetic distribution patterns and for the effective management of their populations. In this study, we use genetic analyses of eleven microsatellites in combination with oceanographic satellite and dispersal simulation data to determine distribution patterns for Serranus cabrilla, a ubiquitous demersal broadcast spawner, in the Mediterranean Sea. Pairwise population F_ST values ranged between ?0.003 and 0.135. Two genetically distinct clusters were identified, with a clear division located between the oceanographic discontinuities at the Ibiza Channel (IC) and the Almeria‐Oran Front (AOF), revealing an admixed population in between. The Balearic Front (BF) also appeared to dictate population structure. Directional gene flow on the Spanish coast was observed as S. cabrilla dispersed from west to east over the AOF, from north to south on the IC and from south of the IC towards the Balearic Islands. Correlations between genetic and oceanographic data were highly significant. Seasonal changes in current patterns and the relationship between ocean circulation patterns and spawning season may also play an important role in population structure around oceanographic fronts.     

From Scilla to Charybdis – is our voyage safer now?

The genus Charybdis Speta (previously Urginea maritima agg.) was investigated karyologically and genetically throughout its geographic range in the Mediterranean. The different ploidy levels show a strong geographic pattern. Diploid populations are mainly found along the northern coast of Africa with C. pancration extending northwards from Tunisia to southern Italy. Tetraploid populations are most densely distributed in the eastern Mediterranean but are also found in North Africa, on the Balearic and Canary Islands. Hexaploid populations are restricted to the Iberian Peninsula and adjacent Morocco and Algeria. Chloroplast microsatellite data suggest that determination of ploidy levels alone is insufficient to adequately describe the existing populations. Especially the tetraploid and hexaploid populations exhibit additional genetic differentiation and geographic structuring. AFLP data indicate that tetraploid populations from southern Italy are of hybrid origin. Phylogenetic analysis further revealed that the genera Urginea Steinh. and Charybdis are not directly related to each other but have strong ties to genera from southern Africa. A possible colonization scenario of the Mediterranean via NW Africa and Iberia is discussed.     

Glacial refuges for three‐spined stickleback in the Iberian Peninsula: mitochondrial DNA phylogeography

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Neither connectivity nor genetic diversity matter in the conservation of a rare fern and a moss on insular erratic boulders

Erratic boulders provide habitat for rock-dwelling species and contribute to the biodiversity of landscapes. In the calcareous Swiss lowlands, siliceous erratic boulders are exclusive habitat islands for the regionally critically endangered fern Asplenium septentrionale, about 20 bryophyte species and numerous lichens. Focusing on island biogeographical processes, we analysed the conservation genomics of A. septentrionale and the moss Hedwigia ciliata on insular erratic boulders in the Swiss lowlands and the adjacent “mainland” in siliceous mountains. We genotyped both species using double digest restriction associated DNA sequencing (ddRAD). For the tetraploid A. septentrionale, abundant identical multilocus genotypes within populations suggested prevalent intragametophytic selfing, and six out of eight boulder populations consisting of a single multilocus genotype each indicated single spore founder events. The genetic structure of A. septentrionale mainland populations coincided with Pleistocene glacial refugia. Four genetic lineages of H. ciliata were identified, and populations consisting of a single multilocus genotype were less common than in A. septentrionale. For both taxa, multilocus genotype diversity on boulders was lower than in mainland populations. The absence of common genetic groups among boulder populations, and the absence of isolation by distance patterns, suggested colonisation of boulders through independent long-distance dispersal events. Successful boulder colonisation of A. septentrionale seems to be rare, while colonisation by H. ciliata appears to be more frequent. We conclude that pivotal principles of conservation biology, such as connectivity and genetic diversity, are of less importance for the studied cryptogams on insular erratic boulders because of long-distance dispersal, intragametophytic selfing and polyploidy.

     

The role of western Mediterranean islands in the evolutionary diversification of the spotted flycatcher Muscicapa striata,a long‐distance migratory passerine species

We investigated the evolutionary history of the spotted flycatcher Muscicapa striata, a long distance migratory passerine having a widespread range, using mitochondrial markers and nuclear introns. Our mitochondrial results reveal the existence of one insular lineage restricted to the western Mediterranean islands (Balearics, Corsica, Sardinia) and possibly to the Tyrrhenian coast of Italy that diverged from the mainland lineages around 1 Mya. Mitochondrial genetic distance between insular and mainland lineages is around 3.5%. Limited levels of shared nuclear alleles among insular and mainland populations further support the genetic distinctiveness of insular spotted flycatchers with respect to their mainland counterparts. Moreover, lack of mitochondrial haplotypes sharing between Balearic birds (M. s. balearica) and Corso‐Sardinian birds (M. s. tyrrhenica) suggest the absence of recent matrilineal gene flow between these two insular subspecies. Accordingly, we suggest that insular spotted flycatchers could be treated as one polytypic species (Muscicapa tyrrhenica) that differs from M. striata in morphology, migration, mitochondrial and nuclear DNA and comprises two subspecies (the nominate and M. t. balearica) that diverged recently phenotypically and in mitochondrial DNA and but still share the same nuclear alleles. This study provides an interesting case‐study illustrating the crucial role of western Mediterranean islands in the evolution of a passerine showing high dispersal capabilities. Our genetic results highlight the role of glacial refugia of these islands that allowed initial allopatric divergence of insular populations. We hypothesize that differences in migratory and breeding phenology may prevent any current gene flow between insular and mainland populations of the spotted flycatcher that temporarily share the same insular habitats during the spring migration.     

Evidence for reticulate palaeogeography: beetle diversity linked to connection-disjunction cycles of the Gibraltar strait

Aim Dispersal barriers between areas within some regions have appeared and disappeared throughout evolutionary time. Here we describe the distributional patterns displayed by three taxa living in such kind of regions. These patterns can be better explained considering a reticulated rather than a hierarchically branched palaeogeography. Location Western Mediterranean. Methods The taxa studied are Misolampus (Coleoptera, Tenebrionidae), Tentyria (Coleoptera, Tenebrionidae) and Thorectes (Coleoptera, Geotrupidae). All them are flightless and show a high degree of endemicity. The individual pattern of area relationships was determined separately for each genus by Brooks Parsimony Analysis (BPA). A theoretical general area cladogram was constructed based on the palaeogeographical history of the region. Finally, the general area cladogram is reconciled with the individual ones. Results The ancestor of Misolampus probably was North African. Land dispersal toward the Iberian Peninsula is proposed. Speciation within Iberia is related to specific vicariance events, and the presence of insular (Balearic Islands) populations is explained by sea‐surface or, more probably, human‐mediated dispersal. The ancestor of Tentyria was Iberic. The proposed hypothesis to explain the current species distribution mainly relies on the occurrence of specific vicariance events. However, the occurrence of some sea‐surface dispersal event is not discarded. Almost all possible vicariance events can be recognized in the first clade of the Thorectes genus. There is evidence for dispersal between Africa and Europe at different dates and in both directions. In spite of some uncertainties, the appearance of the second Thorectes clade can also be explained by the occurrence of specific historical events. An ancient dispersal toward the eastern Mediterranean and several dispersal events during the Messinian seem likely. Main conclusions The same historical events have specific outcomes in every tree (even in every branch within a tree) depending on the ability for dispersal and speciation of each taxon. Connection‐disjunction cycles of dispersal barriers have acted as diversity producers.     

Evolutionary history of Scinax treefrogs on land‐bridge islands in south‐eastern Brazil

Aim We investigated how Pleistocene refugia and recent (c. 12,000 years ago) sea level incursions shaped genetic differentiation in mainland and island populations of the Scinax perpusillus treefrog group. Location Brazilian Atlantic Forest, São Paulo state, south‐eastern Brazil. Methods Using mitochondrial and microsatellite loci, we examined population structure and genetic diversity in three species from the S. perpusillus group, sampled from three land‐bridge islands and five mainland populations, in order to understand the roles of Pleistocene forest fragmentation and sea level incursions on genetic differentiation. We calculated metrics of relatedness and genetic diversity to assess whether island populations exhibit signatures of genetic drift and isolation. Two of the three island populations in this study have previously been described as new species based on a combination of distinct morphological and behavioural characters, thus we used the molecular datasets to determine whether phenotypic change is consistent with genetic differentiation. Results Our analyses recovered three distinct lineages or demes composed of northern mainland São Paulo populations, southern mainland São Paulo populations, and one divergent island population. The two remaining island populations clustered with samples from adjacent mainland populations. Estimates of allelic richness were significantly lower, and estimates of relatedness were significantly higher, in island populations relative to their mainland counterparts. Main conclusions Fine‐scale genetic structure across mainland populations indicates the possible existence of local refugia within São Paulo state, underscoring the small geographic scale at which populations diverge in this species‐rich region of the Atlantic Coastal Forest. Variation in genetic signatures across the three islands indicates that the populations experienced different demographic processes after marine incursions fragmented the distribution of the S. perpusillus group. Genetic signatures of inbreeding and drift in some island populations indicate that small population sizes, coupled with strong ecological selection, may be important evolutionary forces driving speciation on land‐bridge islands.     

Phylogeography and genetic structure of the red-legged partridge (Alectoris rufa): more evidence for refugia within the Iberian glacial refugium

The Pleistocene climatic oscillations promoted the diversification in avian species during the last glacial period. The red‐legged partridge (Alectoris rufa, Family Phasianidae) has a large natural distribution extending from the Mediterranean to humid temperate zones. However, the genetic structure for this species is unknown. The present study investigates the phylogeography, genetic structure and demographic history of A. rufa across its distribution, employing both mitochondrial DNA control region sequences and nuclear microsatellite loci. Our results propose that this species was greatly affected by Pleistocene glaciations. The mismatch analyses suggest that the current populations resulted from post‐glacial expansion and subsequent differentiation resulting in five diagnosable genetic clusters: Southwestern, Central‐eastern, Northwestern, Balearic and French and Italian. Further, we found evidence of three glacial refugia within the currently recognized Iberian glacial refugium. The intraspecific structure revealed by both maternal and biparental phylogeographic analyses was not resolved in the phylogenetic analyses. Based on all considerations, we recommended that five management units be recognized.     

Genetic isolation of insular populations of the Maghrebian bat,Myotis punicus,in the Mediterranean Basin

Aim We investigate the population genetic structure of the Maghrebian bat, Myotis punicus, between the mainland and islands to assess the island colonization pattern and current gene flow between nearby islands and within the mainland. Location North Africa and the Mediterranean islands of Corsica and Sardinia. Methods We sequenced part of the control region (HVII) of 79 bats across 11 colonies. The phylogeographical pattern was assessed by analysing molecular diversity indices, examining differentiation among populations and estimating divergence time. In addition, we genotyped 182 bats across 10 colonies at seven microsatellite loci. We used analysis of molecular variance and a Bayesian approach to infer nuclear population structure. Finally, we estimated sex‐specific dispersal between Corsica and Sardinia. Results Mitochondrial analyses indicated that colonies between Corsica, Sardinia and North Africa are highly differentiated. Within islands there was no difference between colonies, while at the continental level Moroccan and Tunisian populations were highly differentiated. Analyses with seven microsatellite loci showed a similar pattern. The sole difference was the lack of nuclear differentiation between populations in North Africa, suggesting a male‐biased dispersal over the continental area. The divergence time of Sardinian and Corsican populations was estimated to date back to the early and mid‐Pleistocene. Main conclusions Island colonization by the Maghrebian bats seems to have occurred in a stepping‐stone manner and certainly pre‐dated human colonization. Currently, open water seems to prevent exchange of bats between the two islands, despite their ability to fly and the narrowness of the strait of Bonifacio. Corsican and Sardinian populations are thus currently isolated from any continental gene pool and must therefore be considered as different evolutionarily significant units (ESU).     

Mitochondrial differentiation and biogeography of Hyla meridionalis (Anura: Hylidae): an unusual phylogeographical pattern

Aim To study the patterns of genetic variation and the historical events and processes that influenced the distribution and intraspecific diversity in Hyla meridionalis Boettger, 1874. Location Hyla meridionalis is restricted to the western part of the Mediterranean region. In northern Africa it is present in Tunisia, Algeria and Morocco. In south‐western Europe it is found in the south of France, north‐western Italy and north‐eastern and south‐western Iberian Peninsula. There are also insular populations, as in the Canaries and Menorca. Methods Sampling included 112 individuals from 36 populations covering the range of the species. We used sequences of mitochondrial DNA Cytochrome Oxidase I (COI) for the phylogeographical analysis (841 bp) and COI plus a fragment including part of tRNA lysine, ATP synthase subunits 6 and 8 and part of Cytochrome Oxidase III for phylogenetic analyses (2441 bp). Phylogenetic analyses were performed with paup *4.0b10 (maximum likelihood, maximum parsimony) and Mr Bayes 3.0 (Bayesian analysis). Nested clade analysis was performed using tcs 1.18 and Geo Dis 2.2. A dispersal‐vicariant analysis was performed with diva 1.0 to generate hypotheses about the geographical distribution of ancestors. Results We found little genetic diversity within samples from Morocco, south‐western Europe and the Canary Islands, with three well‐differentiated clades. One is distributed in south‐western Iberia and the High Atlas, Anti‐Atlas and Massa River in Morocco. The second is restricted to the Medium Atlas Mountains. The third one is present in northern Morocco, north‐eastern Iberia, southern France and the Canaries. These three groups are also represented in the nested clade analysis. Sequences from Tunisian specimens are highly divergent from sequences of all other populations, suggesting that the split between the two lineages is ancient. diva analysis suggests that the ancestral distribution of the different lineages was restricted to Africa, and that an explanation of current distribution of the species requires three different dispersal events. Main conclusions Our results support the idea of a very recent colonization of south‐western Europe and the Canary Islands from Morocco. South‐western Europe has been colonized at least twice: once from northern Morocco probably to the Mediterranean coast of France and once from the western coast of Morocco to southern Iberia. Human transport is a likely explanation for at least one of these events. Within Morocco, the pattern of diversity is consistent with a model of mountain refugia during hyperarid periods within the Pleistocene. Evaluation of the phylogenetic relationships of Tunisian haplotypes will require an approach involving the other related hylid taxa in the area.     

Vicariant versus dispersal processes in the settlement of Pseudamnicola (Caenogastropoda,Hydrobiidae) in the Mediterranean Balearic Islands

The genus Pseudamnicola comprises a group of tiny springsnails inhabiting several continental and insular regions of the Mediterranean basin. Given the limited dispersal capabilities of these animals, it is difficult to explain the wide distribution range of the genus and, more specifically, its presence in isolated habitats, such as on islands. Thus, to investigate the process(es) that may explain these distribution patterns, we morphologically re‐described and genetically analysed the six Pseudamnicola (Pseudamnicola) species occurring in the Iberian Peninsula and the nearby Balearic Islands. Genetic relationships were explored by sequencing two mitochondrial (cytochrome c oxidase subunit I and 16S rRNA) and one nuclear (28S rRNA) gene in 19 populations. Our morphological study confirmed the presence of previously described species, whereas our phylogenetic results revealed three lineages within the subgenus: one clade grouping the species from Minorca Island with an Iberian Peninsula species, a second clade grouping the three species from Majorca Island, and a third clade that consists of a single species, which occurs in both the Iberian Peninsula and Ibiza Island. Calculated speciation times show that the cladogenetic events involving the insular species seem to have occurred after the current conformation of the Balearic Islands (c. 20 Mya). Therefore, the speciation process may have been related to subsequent transmarine colonizations, probably during the Messinian Salinity Crisis, and the Pleistocene glaciations when landmass corridors connected the islands with the continent. © 2014 The Linnean Society of London     

High levels of population subdivision in a morphologically conserved Mediterranean toad (Alytes cisternasii) result from recent,multiple refugia: evidence from mtDNA,microsatellites and nuclear genealogies

Pleistocene glaciations often resulted in differentiation of taxa in southern European peninsulas, producing the high levels of endemism characteristic of these regions (e.g. the Iberian Peninsula). Despite their small ranges, endemic species often exhibit high levels of intraspecific differentiation as a result of a complex evolutionary history dominated by successive cycles of fragmentation, expansion and subsequent admixture of populations. Most evidence so far has come from the study of species with an Atlantic distribution in northwestern Iberia, and taxa restricted to Mediterranean‐type habitats remain poorly studied. The Iberian Midwife toad (Alytes cisternasii) is a morphologically conserved species endemic to southwestern and central Iberia and a typical inhabitant of Mediterranean habitats. Applying highly variable genetic markers from both mitochondrial and nuclear genomes to samples collected across the species’ range, we found evidence of high population subdivision within A. cisternasii. Mitochondrial haplotypes and microsatellites show geographically concordant patterns of genetic diversity, suggesting population fragmentation into several refugia during Pleistocene glaciations followed by subsequent events of geographical and demographic expansions with secondary contact. In addition, the absence of variation at the nuclear β‐fibint7 and Ppp3caint4 gene fragments suggests that populations of A. cisternasii have been recurrently affected by episodes of extinction and recolonization, and that documented patterns of population subdivision are the outcome of recent and multiple refugia. We discuss the evolutionary history of the species with particular interest in the increasing relevance of Mediterranean refugia for the survival of genetically differentiated populations during the Pleistocene glaciations as revealed by studies in co‐distributed taxa.     

Modified dispersal‐related traits in disjunct populations of bird‐dispersed Frangula alnus (Rhamnaceae): a result of its Quaternary distribution shifts?

Many European tree species survived Pleistocene glaciations in Mediterranean refugia and rapidly recolonized temperate Europe afterwards. Inter‐ and postglacial migration processes are assumed to have catalized evolutionary optimizations of dispersal‐related traits, but up to now empirical evidence is lacking in vertebrate‐dispersed plants. We investigated if south Iberian glacial relict and central European “colonizer” populations of the bird‐dispersed tree Frangula alnus have experienced differentiations of dispersal‐related traits which increase the mobility of northern populations. A comparison of lifetime reproductive strategy, disperser guilds, ripening phenology, and fruit design revealed considerable differences. Compared to south Iberian conspecifics, central European plants were considerably smaller and experienced a highly accelerated generation turnover. In south Iberian populations seed dispersal was carried out almost completely by resident birds which occurred in constant abundances throughout the ripening season. In contrast, central European seeds were dispersed by migrants whose abundances changed considerably during the ripening season. Several bird species were involved in both study areas but rendered different importance for seed dispersal. The fruit ripening pattern was highly asynchronous throughout the ripening season in south Iberia, while central European trees showed a complex ripening sequence which resulted in a significant correlation between fruit abundance and changing disperser availability. Central European fruits were smaller and showed a considerably smaller seed load than south Iberian fruits, thus presumably being more attractive for their small‐sized main dispersers (Sylvia warblers). Chemical analyses revealed significant differences in contents of water, glucose, fructose, proteins, ash. and phenolic compounds. The extensive differentiation of dispersal‐related traits in F. alnus suggests that even weak selective pressures by frugivores may induce evolutionary adjustments of dispersal traits over large time scales. We suggest that the differences we observe today evolved during the species' distribution shifts in the Quaternary.     

Quaternary range dynamics of ecologically divergent species (Edraianthus serpyllifolius and E. tenuifolius,Campanulaceae) within the Balkan refugium

Aim We investigated Quaternary range dynamics of two closely related but ecologically divergent species (cold‐tolerant Edraianthus serpyllifolius and thermophilic Edraianthus tenuifolius) with overlapping distribution ranges endemic to the western Balkan Peninsula, an important yet understudied Pleistocene refugium. Our aims were: to test predictions of the ‘refugia‐within‐refugia’ model of strong genetic subdivisions due to population isolation in separate refugia; to explore whether two ecologically divergent species reacted differently to Pleistocene climatic fluctuations; and to test predictions of the displacement refugia model of stronger differentiation among populations in the thermophilic E. tenuifolius compared with the cold‐tolerant E. serpyllifolius. Location The western Balkan Peninsula. Methods We gathered amplified fragment‐length polymorphism (AFLP) data and plastid DNA sequences from two to five individuals from 10 populations of E. serpyllifolius and 22 populations of E. tenuifolius, spanning their entire respective distribution areas. AFLP data were analysed using a Bayesian clustering approach and a distance‐based network approach. Plastid sequences were used to depict relationships among haplotypes in a statistical parsimony network, and to obtain age estimates in a Bayesian framework. Results In E. serpyllifolius, both AFLP and plastid sequence data showed clear geographic structure. Western populations showed high AFLP diversity and a high number of rare fragments. In E. tenuifolius, both markers congruently identified a major phylogeographic split along the lower Neretva valley in central Dalmatia. The most distinct and earliest diverging chloroplast DNA (cpDNA) haplotypes were found further south in the south‐easternmost populations. North‐western populations, identified as a separate cluster by Bayesian clustering, were characterized by low genetic diversity and a low number of rare AFLP markers. Main conclusions Clear evidence for multiple Pleistocene refugia is found not only in the high‐elevation E. serpyllifolius, but also in the lowland E. tenuifolius, despite the lack of obvious dispersal barriers, in line with the refugia‐within‐refugia model. Genealogical relationships and genetic diversity patterns support the hypothesis that cold‐adapted E. serpyllifolius responded to climatic oscillations mostly by elevational range shifts, whereas thermophilic E. tenuifolius did so mainly by latitudinal range shifts, with different phases (and probably extents) of range expansion. In contrast to the displacement refugia hypothesis, the two elevationally differentiated species do not differ in their genetic diversity.     

Tracking the last elephants in Europe during the Würm Pleniglacial: the importance of the Late Pleistocene aeolianite record in SW Iberia

Abstract

In his short joint ventures across the world of vertebrate tracks, Richard Bromley recognized the aeolian sands as unsuitable soft substrates for their preservation. Only after his work in the Balearic Islands, a more systematic study of coastal aeolianites worldwide revealed that these depositional systems could preserve a highly important record of behavioural trace fossils for the evolution of vertebrates, especially in the Pleistocene, including extinct megafauna and the escalation of the Homo. Here we describe coastal aeolianites from the upper Pleistocene of SW Iberia, namely SW Portugal and Gibraltar, with trackways, tracks and trampled surfaces of the last elephants in mainland Europe. Photogrammetric 3?D modelling and analysis of the relevant proboscidean track levels allowed revision of and support for previous ichnotaxonomic identification to Proboscipeda panfamilia and behavioural interpretations of the producer. Smaller and very large trackways and footprints attributed to Palaeoloxodon antiquus are described and discussed according to new and recent, but always rare findings. The seeming progressive and definitive extinction of this species towards southern Iberia, following the same pattern for the replacement of the Neanderthals during the last initial Pleniglacial (until ca. 28 ka), suggests evidence for co-evolution.     

Genetic evidence for the origins of range disjunctions in the Australian dry sclerophyll plant Hardenbergia violacea

Aim The distribution of genetic variation in the Australian dry sclerophyll plant Hardenbergia violacea (Fabaceae) is examined in the context of Pleistocene climate change in order to identify likely refugia. Particular consideration is given to the origin of range disjunctions in South Australia and Tasmania, and to determining whether the Tasmanian population is indigenous or recently introduced from mainland Australia. Location Southeastern Australian mainland and Tasmania. Methods A combination of chloroplast polymerase chain reaction–restriction fragment length polymorphism and genomic amplified fragment length polymorphism (AFLP) marker systems was used to examine the genetic structure of 292 individuals from 13 populations across the range of H. violacea in southeastern Australia. Results Hardenbergia violacea populations in Tasmania and southern Victoria were characterized by low, almost monotypic chloroplast diversity. New South Wales showed higher haplotype diversity and haplotype sharing among widely distributed populations. Principal coordinates analysis (PCoA) of the AFLP data found a strong latitudinal cline in AFLP variation from northern New South Wales south to Tasmania. The Tasmanian population formed an isolated and somewhat disjunct genetic cluster at one end of this cline. However, the South Australian population was an exception to the clinal variation shown by all other populations, forming a highly disjunct cluster in the PCoA. Within‐population genetic diversity was low in both disjunct populations. Main conclusions The genetic evidence indicates that the Tasmanian population is likely to be indigenous and probably the product of vicariance, which was followed by range contraction at the Last Glacial Maximum or an earlier glacial event. The deep phylogenetic disjunction in South Australia is evidence of a much earlier separation on mainland Australia. The chloroplast structure indicates that, during the Pleistocene, H. violacea underwent broad‐scale recolonization in southern Victoria and Tasmania, possibly from a large continental refugium in eastern New South Wales. We conclude that H. violacea, and presumably the sclerophyll communities in which it occurs, have undergone multiple range contractions to large continental refugia during different Pleistocene glaciations in southeastern Australia.     

Are islands more susceptible to plant invasion than continents? A test using Oxalis pes‐caprae L. in the western Mediterranean

Aim We tested the relative vulnerability of islands to Oxalis pes‐caprae L. invasion compared to mainland regions. Oxalis pes‐caprae is a South African annual geophyte that reproduces via bulbils, and has spread in many Mediterranean and temperate regions of the world where introduced. Our study is one of the first detailed regional analyses of the occurrence and local abundance of a non‐native plant. Methods We conducted an extensive survey (2000 sampling points) to examine local and coarse‐scale patterns in both the occurrence and abundance of O. pes‐caprae on islands and in neighbouring mainland regions of Spain. Location We analysed occurrence (number of samples where present) and abundance (percentage cover) on two Balearic Islands (Menorca and Mallorca) and in two mainland administrative provinces of Spain (Murcia and València). Results Oxalis pes‐caprae occurrence was consistently higher on islands. Occurrence varied among habitats, being the highest in tree groves and the lowest in forests and shrublands. It was never found in these two habitats on the mainland. Mean O. pes‐caprae abundance was greatest in tree groves on the mainland, and in field margins and old fields on the islands. However, in general there were not significant differences in local abundances between island and mainland locations. Main conclusions These findings suggest that local processes (such as the biotic resistance of plant communities) are less important than coarse‐scale phenomena (such as environmental driving forces) in explaining differences in the invasion patterns observed between islands and adjacent mainland regions. We suggest that O. pes‐caprae has occupied a larger proportion of available habitats on islands due to: (1) its strong dependence on domestic animal and human‐mediated dispersal which are probably greater on the islands than in mainland areas, and (2) the smaller area encompassed by islands that, over a comparable period of time, enables a greater proportion of available habitats to be colonized (and hence higher occurrence) than equivalent larger mainland areas.     

Environmental gradients shape the genetic structure of two medicinal Salvia species in Jordan

• Environmental gradients, and particularly climatic variables, exert a strong influence on plant distribution and, potentially, population genetic diversity and differentiation. Differences in water availability can cause among‐population variation in ecological processes and can thus interrupt populations’ connectivity and isolate them environmentally. The present study examines the effect of environmental heterogeneity on plant populations due to environmental isolation unrelated to geographic distance.
• Using AFLP markers, we analyzed genetic diversity and differentiation among 12 Salvia spinosa populations and 13 Salvia syriaca populations from three phytogeographical regions (Mediterranean, Irano‐Turanian and Saharo‐Arabian) representing the extent of the species’ geographic range in Jordan. Differences in geographic location and climate were considered in the analyses.
• For both species, flowering phenology varied among populations and regions. Irano‐Turanian and Saharo‐Arabian populations had higher genetic diversity than Mediterranean populations, and genetic diversity increased significantly with increasing temperature. Genetic diversity in Salvia syriaca was affected by population size, while genetic diversity responded to drought in S. spinosa. For both species, high levels of genetic differentiation were found as well as two well‐supported phytogeographical groups of populations, with Mediterranean populations clustering in one group and the Irano‐Turanian and Saharo‐Arabian populations in another. Genetic distance was significantly correlated to environmental distance, but not to geographic distance.
• Our data indicate that populations from moist vs. arid environments are environmentally isolated, where environmental gradients affect their flowering phenology, limit gene flow and shape their genetic structure. We conclude that environmental heterogeneity may act as driver for the observed variation in genetic diversity.