Alpine Populations Research Articles

European geography of Alnus incana leaf variation

The intra- and inter-population and geographic differentiation of Alnus incana were examined biometrically on the basis of morphological character of the leaves. The study results only partly confirmed the assumption that Carpathian and Alpine populations of the species differ significantly. This may indicate that gene exchange continued during glacial periods as these regions were not sufficiently isolated from each other. The geographic variation of grey alder along the Balkans–Scandinavia axis is more eminent. It is debatable whether A. incana has migrated northwards from montane refugia, along Central European rivers, mostly the Vistula and Oder. It can be, however, noted that the Masurian, Sudetan and East Carpathian populations are similar to one another.

Natural selection and neutral evolution jointly drive population divergence between alpine and lowland ecotypes of the allopolyploid plant Anemone multifida (Ranunculaceae).

Population differentiation can be driven in large part by natural selection, but selectively neutral evolution can play a prominent role in shaping patters of population divergence. The decomposition of the evolutionary history of populations into the relative effects of natural selection and selectively neutral evolution enables an understanding of the causes of population divergence and adaptation. In this study, we examined heterogeneous genomic divergence between alpine and lowland ecotypes of the allopolyploid plant, Anemone multifida. Using peak height and dominant AFLP data, we quantified population differentiation at non-outlier (neutral) and outlier loci to determine the potential contribution of natural selection and selectively neutral evolution to population divergence. We found 13 candidate loci, corresponding to 2.7% of loci, with signatures of divergent natural selection between alpine and lowland populations and between alpine populations (Fst = 0.074–0.445 at outlier loci), but neutral population differentiation was also evident between alpine populations (FST = 0.041–0.095 at neutral loci). By examining population structure at both neutral and outlier loci, we determined that the combined effects of selection and neutral evolution are associated with the divergence of alpine populations, which may be linked to extreme abiotic conditions and isolation between alpine sites. The presence of outlier levels of genetic variation in structured populations underscores the importance of separately analyzing neutral and outlier loci to infer the relative role of divergent natural selection and neutral evolution in population divergence.

Local adaptation, refugial isolation and secondary contact of Alpine populations of the land snail Arianta arbustorum

Depressed-shelled populations of the land snail Arianta arbustorum occur in rocky Alpine habitats, whereas globular shell forms are ubiquitous in lowland habitats of central Europe. It has been proposed that Alpine, steep, rocky habitats are the ancestral habitat and depressed-shelled snails the ancestral form of A. arbustorum. According to this hypothesis, the globular shell form evolved when A. arbustorum adapted to damp lowland habitats during the Pliocene and depressed-shelled forms survived the Pleistocene glaciations in isolated Alpine habitats like nunataks throughout the Alps. Alternatively, the depressed shells could be a more recent adaptation to steep rocky habitats, derived from the widespread globular-shelled lowland A. arbustorum. To test these alternatives, we compared globular and depressedshelled morphotypes and their molecular differentiation between two geologically similar massifs in the

Differentiation of Floral Traits Associated with Pollinator Preference in a Generalist-Pollinated Herb, Trollius ranunculoides (Ranunculaceae)

Premise of research. In flowers that are specialized, floral traits often differentiate in association with pollinator shifts. However, in species of generalized pollination, the role of pollinators in local population differentiation of floral traits is difficult to test. A crucial line of evidence is to contrast the direction and strength of pollinator selection among populations.Methodology. Variation in floral traits and pollinator assemblages was investigated in two populations of a generalist-pollinated herb, Trollius ranunculoides (Ranunculaceae), to detect the potential role of pollinators in floral differentiation.Pivotal results. The alpine population was visited more by flies, and the subalpine population was visited more by bees. Flower manipulations suggested that large flowers were more attractive to both bees and flies, while more petals were favored only by nectar-feeding flies. Compared to the subalpine population, plants in the alpine population had more petals per flower but smaller flo...

Genetic changes in flowering and morphology in response to adaptation to a high-latitude environment in Arabidopsis lyrata.

The adaptive plastic reactions of plant populations to changing climatic factors, such as winter temperatures and photoperiod, have changed during range shifts after the last glaciation. Timing of flowering is an adaptive trait regulated by environmental cues. Its genetics has been intensively studied in annual plants, but in perennials it is currently not well characterized. This study examined the genetic basis of differentiation in flowering time, morphology, and their plastic responses to vernalization in two locally adapted populations of the perennial Arabidopsis lyrata: (1) to determine whether the two populations differ in their vernalization responses for flowering phenology and morphology; and (2) to determine the genomic areas governing differentiation and vernalization responses. Two A. lyrata populations, from central Europe and Scandinavia, were grown in growth-chamber conditions with and without cold treatment. A QTL analysis was performed to find genomic regions that interact with vernalization. The population from central Europe flowered more rapidly and invested more in inflorescence growth than the population from alpine Scandinavia, especially after vernalization. The alpine population had consistently a low number of inflorescences and few flowers, suggesting strong constraints due to a short growing season, but instead had longer leaves and higher leaf rosettes. QTL mapping in the F2 population revealed genomic regions governing differentiation in flowering time and morphology and, in some cases, the allelic effects from the two populations on a trait were influenced by vernalization (QTL × vernalization interactions). The results indicate that many potentially adaptive genetic changes have occurred during colonization; the two populations have diverged in their plastic responses to vernalization in traits closely connected to fitness through changes in many genomic areas.

The Swiss Vote on Limiting Second Homes

The Alps have an important role in defining the Swiss national identity, and play a major part in forging the country’s image from the beginning of tourism in the 18th and 19th centuries. In a federalist country which assigns large decision powers to the cantonal and communal levels, local and national interests as to how to develop this particular space can profoundly diverge. The vote held on March 11th, 2012 on the Franz Weber initiative introducing a ban on building new holiday residences in touristic communes was a blatant example of such an opposition. The success of the initiative, albeit with a tiny majority, was a great shock; its territorial imprint was evident, drawing a line between alpine and prealpine cantons on one side, and mainland cantons on the other side. This paper proposes a spatial analysis of this vote and put it in context through a historical perspective. Indeed, of all the votes held since 1950 with an environmental or landscape component, we can find only one with the same territorial pattern – in 1965, “against inflation in the building industry” – as the 2012 vote. In all the other votes held on those subjects, other splits – linguistic, religious, or regional – manifested themselves, allowing the alpine populations to find allies. This paper asks thus the question as to whether the long-lived solidarity of mainland Switzerland with the Alps tends to dissipate.

Determinants of the acoustic behaviour of red deer during breeding in a wild alpine population, and implications for species survey

A necessary step in the application of animal communication studies to population ecology involves quantitative descriptions of acoustic variation within individuals. Stereotypy is in fact fundamental to allow individual recognition based on call structure and, in turn, to establish the number of different calling individuals in a population (complementing censuses), or to track individuals through years by the acoustic characteristics of their signals (complementing monitoring). Here we centred on the acoustic behaviour of red deer stags during reproduction to analyze its ecological determinants and to test whether calls are individual specific. Scanty information is available on how day time and weather may affect red deer acoustic activity, and whether spectrotemporal analyses of calls could permit to discriminate among different individuals. We centred on roars, the loud repeated calls given by stags throughout the rut, in a mountainous forest habitat, over two rutting seasons. Significant inter-annual differences were found in the duration of the roaring period and in the magnitude of roaring rates. Roaring rate dropped in the wettest days, suggesting that counts by ear may be biased by changeable environmental conditions within and among breeding seasons. By analysing the spectrotemporal features of roars, we found that several spectro-temporal variables varied more among than within individuals. Discriminant analysis classified a posteriori the 72% of the calls to the individuals that uttered them, and the fundamental frequency was the acoustic parameter that best discriminated individuals. Therefore, in areas in which visual information is limited by habitat characteristics, surveys by ear should be performed in dry weather and over several nights to well cover the roaring period. This count method could be complemented by spectrografic analysis of stag call bouts to assist in censusing populations and reduce the probability of double counts when stags are moving.

Rock sparrow song reflects male age and reproductive success.

The evolution of mating signals is closely linked to sexual selection. Acoustic ornaments are often used as secondary sexual traits that signal the quality of the signaller. Here we show that song performance reflects age and reproductive success in the rock sparrow (Petronia petronia). In an Alpine population in south-east France, we recorded the songs of males and assessed their genetic breeding success by microsatellite analysis. In addition to temporal and spectral song features, we also analysed for the first time whether the sound pressure level of bird song reflects reproductive success. Males with higher breeding success sang at a lower rate and with a higher maximum frequency. We found also that older males gained more extra-pair young and had a higher overall breeding success, although they also differed almost significantly by having a higher loss of paternity in their own nests. Older males could be distinguished from yearlings by singing at lower rate and higher amplitudes. Our findings suggest that song rate may be used as a signal of age and together with song pitch as a signal of reproductive success in this species. Alternatively, younger and less successful males might try to compensate their inferior status by increased song rates and lower pitch. Independent of age and quality, high-amplitude songs correlated with paternity loss in the own nest, suggesting that in this species song amplitude is not an indicator of male quality but high-intensity songs may be rather a response to unfaithful social mates.

Comprehensive analysis of Phyteuma orbiculare L., a wild Alpine food plant

Plants which have been traditionally eaten by the alpine population may provide new opportunities for agricultural development in mountain regions. In this context we have investigated the chemical composition of Phyteuma orbiculare (Campanulaceae), a perennial herb whose leaves have been eaten as salad by rural populations in Valais (Switzerland). Extracts of different polarities were subjected to comprehensive metabolite profiling using a dereplication platform combining HPLC–PDA-MS, and offline NMR analysis. Twenty-three compounds, including various phenolic glycosides, a new dimeric phenylpropanoid glucoside, saponins, and fatty acids were identified online, or after targeted isolation. Selected phenolic constituents were quantitatively assessed by HPLC–PDA analysis. In addition, substances relevant for nutrition, such as β-carotene, fatty acids, ascorbic acid and minerals were quantified in leaves and flowers. The antioxidant capacity was determined with an ORAC assay, and total phenolic compounds were quantified. Finally, the phytochemical profile was compared to that of the related species P. spicatum, P. hemisphaericum and P. ovatum.

Global major histocompatibility Class II β (mh‐IIβ)‐polymorphism in Arctic charr Salvelinus alpinus

This study explored the use of the gene encoding the β subunit of the major histocompatibility (MH) receptor as a population marker in Arctic charr Salvelinus alpinus. The use of this polymorphic marker allowed differentiation of the S. alpinus lineages previously defined using mitochondrial DNA (mtDNA) but also allowed differentiation between the populations studied within those lineages. The majority of the variation observed here occurred prior to the last glaciation event. Nevertheless, all S. alpinus populations were differentiated using both MH Class II β (mh-IIβ) sequences and allelic frequencies. The fact that all the populations studied presented high rates of non-synonymous: synonymous substitutions and high levels of interpopulation variation, suggested mh-IIβ as an ideal marker to assess differentiation among S. alpinus populations in ways that may represent divergence both by genetic drift and natural adaptation to the local environment.

Refugia within refugia as a key to disentangle the genetic pattern of a highly variable species: The case of Rana temporaria Linnaeus, 1758 (Anura, Ranidae)

Two distinct lineages of Rana temporaria are known in the Palaearctic region, but it is uncertain whether this species persisted in one or more Pleistocene refugia. We resolved the phylogeographic history and genetic variability of R. temporaria in the Italian peninsula, a ‘traditional’ Pleistocene refugium, and related our findings to patterns described for other European populations.We sequenced the mitochondrial markers Cox I and cytochrome b. Phylogenetic reconstruction only indicated the presence of haplotypes belonging to the Western lineage in the Italian peninsula. Overall, the genetic variability of Italian populations was higher than other European populations, which shared haplotypes with the Alpine populations. We demonstrated subdivision into five main Italian sublineages, which was associated with a geographical structure of populations in two divergent groups. In particular, one Apennine group might have resulted from bottlenecks during the last interglacials ages. In contrast, Alpine populations were recently diverged and showed incomplete lineage sorting.Our data indicate that the Italian peninsula served as refugium for the Western lineage of R. temporaria. Dispersion towards Central Europe probably started only from the western slope of the Alps via a rapid leading edge expansion. The identified structure is partially congruent with traditional peripheral refugia identified for plants. This evolutionary scenario does not support any taxonomic distinction at the subspecific level for R. temporaria.

Pattern of genetic differentiation in Gentiana pannonica Scop.: did subalpine plants survive glacial events at low altitudes in Central Europe?

The molecular population structure of 20 populations of the subalpine plant Gentiana pannonica was studied by use of amplified fragment length polymorphism (AFLP) and sequencing of non-coding regions of plastid DNA. Of the populations sampled, 18 were native (11 were from the Eastern Alps, which is the distribution centre of the species, and seven were from the Bohemian Forest, which is on the margin of the distribution range), and two were from the Giant Mts and of unclear status. No plastid DNA polymorphisms were found within the entire 6,185 bp investigated. The AFLP data revealed grouping of populations at the regional level. However, differentiation at the regional level (10.3 %) and at the interpopulation level (14.2 %) was low. Even though current populations are isolated and contain small numbers of individuals, the within-population variation (75.511 %) was high. Genetic variation was higher for alpine populations than for Bohemian Forest populations, probably because of fundamental differences in historical changes in population size between these regions. Within-population variation was intermediate for populations in the Giant Mts. The results indicate the possibility of a large distribution of species in the unglaciated areas of Central Europe, irrespective of altitude, during the late Pleistocene and early Holocene. Our results do not confirm that G.pannonica was introduced in the Giant Mts, and native status in the Giant Mts is possible.

Using forensic microsatellites to decipher the genetic structure of linguistic and geographic isolates: A survey in the eastern Italian Alps

The study of geographically and/or linguistically isolated populations could represent a potential area of interaction between population and forensic genetics. These investigations may be useful to evaluate the suitability of loci which have been selected using forensic criteria for bio-anthropological studies. At the same time, they give us an opportunity to evaluate the efficiency of forensic tools for parentage testing in groups with peculiar allele frequency profiles. Within the frame of a long-term project concerning Italian linguistic isolates, we studied 15 microsatellite loci (Identifiler kit) comprising the CODIS panel in 11 populations from the north-eastern Italian Alps (Veneto, Trentino and Friuli Venezia Giulia regions). All our analyses of inter-population differentiation highlight the genetic distinctiveness of most Alpine populations comparing them either to each other or with large and non-isolated Italian populations. Interestingly, we brought to light some aspects of population genetic structure which cannot be detected using unilinear polymorphisms. In fact, the analysis of genotypic disequilibrium between loci detected signals of population substructure when all the individuals of Alpine populations are pooled in a single group. Furthermore, despite the relatively low number of loci analyzed, genetic differentiation among Alpine populations was detected at individual level using a Bayesian method to cluster multilocus genotypes. Among the various populations studied, the four linguistic minorities (Fassa Valley, Luserna, Sappada and Sauris) showed the most pronounced diversity and signatures of a peculiar genetic ancestry. Finally, we show that database replacement may affect estimates of probability of paternity even when the local database is replaced by another based on populations which share a common genetic background but which differ in their demographic history. These findings point to the importance of considering the demographic and cultural profile of populations in forensic applications, even in a context of substantial genetic homogeneity such as that of European populations.

Book review: An integrative overview on alpine habitats

news and update ISSN 1948-6596 book review An integrative overview on alpine habitats The Biology of Alpine Habitats, by Laszlo Nagy & Georg Grabherr Oxford University Press, 2009, 336 pp. ISBN13: 978-0-19-856703-5 http://www.oup.com/ Mountains, and specifically alpine habitats, are fragile and extremely sensitive to perturbations, because of intense gravity effects on flows of en- ergy and materials. As a consequence, they are highly exposed to global change impacts. Under- standing the abiotic and biotic patterns and proc- esses involved in alpine habitats is not only of ut- most importance for researchers interested in mountain regions, but also for those looking to gain deeper insights in current biological debates, such as the effects of nutrient limitations on spe- cies diversity, dispersal/colonization dynamics and niche space filling, the changing nature of species interactions in harsh habitats, or impacts of global change in “nowhere to go” habitats. Nagy and Grabherr’s The Biology of Alpine Habitats deals, among others, with those topics and it is a neat inspection of the upper part of mountains, their constituent elements and the challenges that they face under global change. The book is organized in eleven chapters. The first part (chapters 1, 2, and 3) focuses on de- fining alpine habitats, describing the different mountain regions across the globe from high lati- tudes to the tropics, and also the distribution of the main abiotic and biotic factors along eleva- tional gradients. The second part of the book (chapters from 4 to 8) describes the main abiotic factors constituting alpine habitats, such as en- ergy, climate, landforms, hydrology and soils, and the relation of these factors to alpine communi- ties. These first two parts of the book provide the necessary foundations to discuss global change impacts on climate, nitrogen deposition and land uses in alpine habitats, and to offer a brief sum- mary on conservation issues (chapters 9 and 10). The way the authors frame and organise the book provides the reader with the chance to surf smoothly across the key elements constituting alpine habitats and the relationships between these elements. This easy-to-follow flow also con- tinues when it comes to integrate the variety of spatial and temporal scales in which the drivers shaping current alpine habitats operate: from ex- planations of the evolutionary and biogeographic processes behind the formation of alpine floras and their traits, to the role of species interactions and their specific abilities in the colonization of new areas. This journey across scales allows Nagy and Grabherr to integrate knowledge that belongs to disciplines such as geology, climatology, bio- geography, community ecology or eco-physiology. Their definition of ecological communities (“…net outcome of all biotic interaction and abiotic con- straints in a given location at a given time, with a given background of history, species pool)” illus- trates the integrative perspective in which this book is framed. The role of historical biogeographic proc- esses to explain the formation of alpine floras and current alpine biodiversity patterns is explicitly discussed in Chapter 7. Readers will find that the authors highlight how biogeographical disciplines such as phylogeography (Box 7.1) are of utmost interest for linking past dynamics and structure of alpine populations with current biodiversity pat- terns. An excellent idea was to devote the last part of this chapter to the adaptation and survival of alpine organisms. In doing this, the authors rec- ognise the evolutionary foundations of issues that have been often perceived as pure ecological phe- nomena, such as eco-physiology or reproduction (adaptations to alpine existence across different time scales are summarized in Fig 7.12). This inte- gration of evolutionary and ecological aspects in a biogeographical framework to better understand adaptation and survival abilities of alpine habitats provides a robust venue to better discuss future responses of alpine biodiversity to global change. In summary, The Biology of Alpine Habitats, which is part of the Habitat Series published by Oxford University Press, intends to provide stu- frontiers of biogeography 2.2, 2010 — © 2010 the authors; journal compilation © 2010 The International Biogeography Society

An initial investigation of the taxonomic status of Saussurea esthonica Baer ex Rupr. utilising DNA markers and sequencing

Saussurea esthonica Baer ex Rupr. is a species of the genus Saussurea that is found in Estonia, Latvia and the Leningrad region in Russia. Only two populations have been identified in Latvia, and it is protected at the national level. The taxonomic relationships within this genus are unclear, and the majority of phylogenetic studies of this genus have been undertaken on species found in Asia. The relationship of S. esthonica to the other Saussurea species is equally unclear, with some suggestions that this species should be regarded as a subspecies of S. alpina. Two different molecular marker techniques, iPBS and ITS sequencing, were utilised to investigate the phylogeny of S. esthonica and its relationship with other Saussurea species, including three European species: S. alpina, S. pygmaea and S. discolor. The phylogeny obtained with the iPBS markers was similar to that obtained with the ITS sequences. The phylogenies obtained were broadly consistent with previous morphological classification, with S. alpina, S. esthonica and S. discolor clustering together. S. pulchella, S. albescens and S. candicans also formed one discrete cluster. The taxonomic status of S. esthonica was not unambiguously determined, with Norwegian and Italian S. alpina populations clustering with Latvian and Estonian S. esthonica populations, and the Austrian S. alpina populations clustering with Austrian S. discolor populations.

Genetic variability of relict Rhododendron ferrugineum L. populations in the Northern Apennines with some inferences for a conservation strategy

In this study, the genetic diversity of three Rhododendron ferrugineum L Apennine populations (AP1, AP2, and AP3) was analyzed and compared to three populations of the Maritime Alps and six populations of Central-Eastern Alps. Genetic variations across microsatellite markers revealed that the Apennine populations show some clonal individuals and the lowest genetic diversity values (AP1 and AP3 A[18] values are 2.46 and 2.31, respectively), as well as heterozygosity deficiency with respect to the Alpine populations. Genetic relationships among populations (Nei's genetic distance) showed that populations from the Central-Eastern Alps and from the Maritime Alps clustered in two separate groups. Unweighted pair group method with arithmetic averages as well as PCA analysis showed a clear separation of the three Apennine populations according to the high FST values detected (AP1–AP2 = 0.427; AP1–AP3 = 0.446; AP2–AP3 = 0.325). Mantel test revealed a significant correlation between genetic and geographical distance matrices (r = 0.314, P = 0.001, 999 permutations). Concerning the relationship between Apennines and alpine population, PCA analysis showed a clear genetic similarity among Maritime Alps populations and individuals of AP1 population. Considering the geographical and ecological peripheral condition of these populations and the high impact of the climatic changes on their habitat, we suggest the combination of in situ and ex situ conservation strategies to preserve the genetic diversity of this species in the Northern Apennines.

The dynamics of alternative male mating tactics in a population of Black Grouse Tetrao tetrix in the Italian Alps

The Black Grouse Tetrao tetrix is primarily a lekking species, although in some populations solitary display is also common. We analysed data from a 25-year-long survey of an increasing Black Grouse population in the western Italian Alps in order to test whether the numbers of males adopting different tactics conformed to patterns expected, based on the hypothesis that dominant birds occupy the leks and solitary displaying birds are subordinate, and that the tendency for birds to be excluded from leks increases with lek size. Four predictions were made to test this hypothesis: (1) the proportion of solitary males would increase as the overall population increases, (2) there would be a threshold at which there is little or no increase in the size of the lekking population, but continued increase in the population of solitary displaying birds, (3) available lek habitat would be occupied prior to an increase in solitary displaying males, and (4) there would be an increase in the use of less preferred habitat by solitary displaying males as the total population increases. Population trends varied according to display status, in that the lekking population reached an apparent threshold while the number of solitary males continued to increase, and there was an expansion of the range of solitary males into higher altitudes in years when the population was larger. However, there was no increase in the proportion of solitary males as the total population increased. Furthermore, there was no suggestion that suitable lek habitat was limiting, as the number of leks and the mean size of those leks continued to increase throughout the study period. Although two predictions were supported, the precise patterns of trends, and results of additional analyses, suggested that solitary display is more likely to be an alternative rather than a subordinate tactic. Several factors may promote this apparent dual display tactic in Black Grouse, but habitat structure in this and other Alpine populations is likely to be a key influence.

Evidence for source - sink dynamics in a regional population of arctic ground squirrels (Urocitellus parryii plesius)

Context Variable demographic rates can manifest themselves between habitat types in the form of source–sink dynamics where populations in sink habitats would not exist without the addition of migrants from source habitats. Aims Arctic ground squirrels (Urocitellus parryii pleisus (Osgood, 1900)) occupy a large geographic area in northern Canada and live in a variety of habitat types, including boreal forest, low-elevation meadows and alpine meadows, providing an opportunity to investigate the possible existence of source–sink dynamics. Methods We hypothesised that arctic ground squirrels in the south-western Yukon exhibit demographic characteristics indicative of source–sink dynamics. Boreal forest habitat could be a sink in spite of previous high squirrel densities, whereas meadows could be a source. We investigated this by mark–recapture live-trapping and radio-telemetry. Key Results In the boreal forest in the Kluane region, we found reduced recruitment, reduced population growth rates (λ), and reduced survivorship for radio-collared individuals that moved from low-elevation meadows into the boreal forest. There was no evidence from radio-collared juveniles of dispersal from high-density ground squirrel populations in alpine meadows down into boreal forest. Conclusions Boreal forest is a sink habitat for arctic ground squirrels. Source–sink dynamics observed between low-elevation meadow and boreal forest habitats appear to result from increased predation pressure in the boreal forest. The result has been a near extirpation of boreal forest arctic ground squirrels in the Kluane region since 1998. Implications Because the source areas of low-elevation meadows occupy only 7–9% of the lowland habitat, recolonisation of boreal forest sites has been very slow. Whereas alpine populations remain high in 2011, boreal forest populations remain near zero. Alpine populations do not appear to be a source for the boreal forest.

Status and distribution of the lynx (Lynx lynx) in the Swiss Alps 2005–2009

We evaluated the status of lynx in the Swiss Alps for the period 2005– 2009. Even though the number of lynx presence signs remained almost stable between the present (2,068 signs) and previous pentad (2,091), there was a 7.6% increase in the area occupied by the 5-km circular buffers around the confirmed lynx signs of presence over the five years period (12,637 km2 ). The north-western Swiss Alps (VI) remained the compartment with the highest number of chance observations. It was followed by compartments central Switzerland west (III) and north-eastern Switzerland (II). These sub-populations acted as source in the current pentad, as signs of reproduction were reported almost every year. The translocation to north-eastern Switzerland is still the only significant contribution to the spatial increase of the lynx range in the last 10 years in the Swiss Alps. The small and vulnerable north-eastern Switzerland lynx sub-population plays an important role for the Alpine population. There is hope that in the future this sub-population could act as stepping stone to the eastern Alps and together with individuals dispersing from the central Switzerland west (III) subpopulation would enable to found a new sub-population in central Switzerland east (IV). The status of the sub-population in the Valais (VII) is less clear. As only few signs of reproduction and mortalities were reported over the pentad, it acted more as sink than a source population. From the few signs of lynx presence reported in the remaining compartments (Grisons V, central Switzerland east IV and Ticino VIII) we concluded that only a few single lynx that did not yet establish the typical social organisation occur there. An occupancy-based population estimate from a parallel study resulted in about 111 (SE = 10) independent lynx for the period 2005–2009. This is higher than the 60–90 individuals estimated for the previous pentad.

Intricate variation patterns in the diploid‐polyploid complex of Alyssum montanum‐A. repens (Brassicaceae) in the Apennine Peninsula: Evidence for long‐term persistence and diversification

The Apennine Peninsula, one of the three peninsulas of the European Mediterranean, is an important hotspot of genetic and species diversity, but studies devoted to plant evolution are still very scarce in this region. We studied the diploid-polyploid complex of Alyssum montanum-A. repens, focusing on Apennine and adjacent southwestern Alpine populations from southeastern France, with the aim of examining their taxonomic position and evolutionary patterns. We employed AFLP markers and cpDNA sequences, along with cytotype determination using flow cytometry, and a multivariate morphometric approach. The Italian and French populations formed two well-delimited groupings within the studied complex that were, in contrast to previous taxonomic treatments, clearly distinct from A. montanum. Populations from southeastern France represent A. orophilum, a previously described but abandoned species. Those from central and southern Italy correspond to A. diffusum, exhibiting high, geographically structured variation (central Apennines, Gargano, and southern Apennines/Calabria). This pattern coincides with hotspot refugial regions, in congruence with the "refugia-within-refugia" hypothesis, and is reflected here in the recognition of three subspecies within A. diffusum. We provide evidence for the presence of Mediterranean refugia for the studied Alyssum montanum-A. repens complex located in central and southern Italy, which, however, did not contribute to the postglacial colonization of Central Europe. Past extinctions, genetic bottlenecks, and recent expansion were inferred in Central Europe, while long-term accumulation of diversity as well as polyploidization occurred in the Apennines.