Minuartia graminifolia (Caryophyllaceae), a south-east European species

A re-evaluation of Minuartia graminifolia has been undertaken based on comparative morphological studies. M. graminifolia subsp. rosani (an endemic of the south-central Apennines and Sicily) and subsp. hungarica (an endemic of the Banat region of Romania) are accepted infraspecific taxa. M. graminifolia subsp. hungarica is lectotypified. M. graminifolia subsp. clandestina is confirmed for Italy; the taxon is neotypified. The type subspecies is considered an endemic to the east-central Alps. Keys to the species of Minuartia ser. Graminifoliae and to the subspecies of M. graminifolia are provided.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 419–432.     

Biogeographical connections between the Maghreb and the Mediterranean peninsulas of southern Europe

The glacial–interglacial cycles have caused severe range modifications of species' distributions. In Europe, thermophilic species had to retreat into geographically distinct southern refugia during glaciations. This process produced strong genetic imprints, which are still detectable by the present pattern of genetic differentiation and the distribution of regional diversity. To reveal the biogeographical imprints in the western Mediterranean, we analysed 26 populations of the butterfly Maniola jurtina spread over large areas of its European and North African distribution range. The samples were analysed using allozyme electrophoresis. We detected three genetic groups, divided into Western Europe, Central/Eastern Europe, and Italy with the Maghreb. The North African samples randomly cluster within the Italian samples. Even the population sampled in Morocco is genetically closely related to these samples and not to the geographically neighbouring Iberian ones. Parameters of genetic diversity showed similar values over the whole study area. The observed genetic pattern reflects possible glacial refugia in Europe located in the Iberian Peninsula and the Balkans. For North Africa and Italy, our data reveal a colonization of Africa originating from Italy.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 693–703.     

Genetic variation of Saxifraga paniculata Mill. (Saxifragaceae): molecular evidence for glacial relict endemism in central Europe

Saxifraga paniculata Mill. is an amphi-atlantic species that is widely distributed in the Alps but also occurs with disjunct populations in central Europe. These isolated populations are considered to be glacial relicts. In the study presented here, we analysed the genetic variation within and between populations of Saxifraga paniculata in central Europe to test whether the molecular data give evidence for the supposed relict endemism. We used RAPD analysis and the analysis of molecular variance (AMOVA) technique to investigate 30 populations of Saxifraga paniculata from different locations in central Europe and adjacent geographical areas. In the RAPD analysis, 319 fragments could be amplified of which 91.2% were polymorphic. The percentage of polymorphic bands within populations correlated significantly with population size, indicating a higher level of inbreeding in small populations. AMOVA revealed 43.60% of the variation within populations, only 15.45% between populations within locations, and 40.95% between locations. Genetic distance between locations (Φ_ST) correlated significantly with the geographical distance between locations, giving evidence for geographical isolation. In an UPGMA cluster analysis, based on RAPD data, the populations were grouped together according to their location. The results exhibit a strong genetic differentiation which is obviously due to genetic drift in the isolated populations. Our study therefore gives evidence for glacial relict endemism of Saxifraga paniculata in central Europe.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80, 11–21.     

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.     

Morphological and geographic variation of the cycad Dioon edule Lindl. (Zamiaceae): ecological and evolutionary implications

The relationship in geographical distribution and morphological variation of leaflet width and length (diagnostic trait), between and within populations of Dioon edule Lindl., has been investigated throughout its known range in eastern Mexico (from the states of Nuevo León to Veracruz, north to south, respectively). A total of 1832 leaflets were measured for width and length from 154 plants distributed amongst five populations using four leaflet replicas from each of three leaves per plant. For leaflet width and length the variation among populations indicated significant stat-istical differences ( F _4,147 = 125.83; P  < 0.0001; R ² = 92.17% and F _4,147 = 9.04; P  < 0.001; R ² = 26.8%), respectively. With respect to leaflet width, the multiple range test showed three groups with a north to south distributional relationship along the range of the species. The correlation coefficient among paired populations, respect to geographical distance and the absolute value of the mean difference of leaflet width in each population, was positive, and different from zero ( r  = 0.82; P  = 0.013). A great variation of important ecological and evolutionary parameters was shown.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 465–470.     

A new violet species (Violaceae) from the south-west Alps

Viola laricicola Marcussen sp. nov. is described from collections from four French and Italian localities in the south-west Alps. It is closely related to V. riviniana (section Trigonocarpea ) but differs in a few morphological characters and by the chromosome number 2 n  = 20. Viola laricicola is endemic to the montane and subalpine coniferous forests in the continental parts of the Alps.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 142 , 119–123.     

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.     

Resource polymorphism in a Patagonian fish Percichthys trucha (Percichthyidae): phenotypic evidence for interlake pattern variation

Within-species differentiation in phenotypic characters related to resource use (resource polymorphism) is frequently thought to result from divergent natural selection in a heterogeneous environment with 'open niches'. In this study we found consistent resource-based polymorphism within three different populations of Percichthys trucha , a lake-dwelling fish native to the southern Andes. In each of three lakes we found two morphotypes that could be clearly distinguished by differences in gill raker length. However, the magnitude of the polymorphism, and the suite of phenotypic characteristics associated with the polymorphism, differed between lakes. Patterns of divergence were more similar between the two northern lakes which ultimately drain into a common river, than between these two lakes and a more southern, unconnected lake. The southern population, which had the largest divergence in gill raker length (32% vs. 16% and 19%), also showed substantial differences in diet. Evidence from the southern population suggests that polymorphism in P. trucha is present early during ontogeny. We conclude that while there are some strong parallels among lakes in the development of a trophic polymorphism, differences in environmental ­conditions and/or colonization history have led to substantial differences in the evolutionary history, resulting in ­different ecological roles of common morphotypes within different lakes.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 497–515.     

Glacial history of high alpine Ranunculus glacialis (Ranunculaceae) in the European Alps in a comparative phylogeographical context

There is considerable controversy concerning the fate of Alpine plants during Pleistocene glaciations. While some studies have found evidence for nunatak survival, others have explained the present genetic patterns by survival only in peripheral refugia. We investigated 75 populations of high alpine Ranunculus glacialis from its entire Alpine distribution. Phylogeographical analyses of AFLP data revealed four groups of populations. Two of them, located in the western Alps, were genetically isolated from each other and from the eastern groups, whereas the two eastern Alpine groups were genetically more similar to each other. This suggests longer isolation and/or lower levels of gene flow in the two western groups. As all groups are close to, or overlap with, presumed glacial refugia, invoking glacial survival on nunataks is unnecessary to explain the present genetic pattern. Similar to the phylogeographical patterns of R. glacialis , the previously investigated alpine Phyteuma globulariifolium and Androsace alpina , which are also confined to siliceous bedrock, showed strong geographical affinities to peripheral refugial areas and there were large-scale congruencies in the location of these refugia for all three species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 183–195.     

Phylogeography of the pool frog (Rana lessonae Camerano) in Europe: evidence for native status in Great Britain and for an unusual postglacial colonization route

The pool frog, Rana lessonae , has historically been considered an introduction into Britain, with Italy the most likely source. Recently, the possibility of native status for a Norfolk pool frog population was raised. We used random amplified DNA (RAPD) analyses to clarify the status issue. Nine arbitrarily designed primers detected 160 polymorphisms in 174 pool frogs from 11 European locations. Polymorphism levels varied widely, being lowest in the northern populations and highest in the south. As with many isolated populations, the peninsular and insular populations of Sweden, Norway and Britain showed relatively little polymorphism. Principal component and cluster analysis showed clear geographical groupings. The Norfolk, Norwegian and Swedish individuals formed a closely related group − a northern clade, substantiating native status for the Norfolk population, which, interestingly, had idiosyncratic features suggestive of a distinctive evolutionary history. Phylograms constructed from RAPD data were consistent with an unexpected postglacial colonization route, in which the northern clade derived from migration out of Italy, via Hungary and Poland, though cryptic glacial refugia in Eastern Europe are an alternative explanation. Our RAPD analyses concur with a parallel microsatellite investigation. Both genetic studies and bioacoustic and archaeozoological findings support native status for Norfolk pool frogs and have prompted a programme for re-establishing the northern clade in England.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 41–51.     

Arbuscular mycorrhizal fungi: genetics of multigenomic, clonal networks and its ecological consequences

Arbuscular mycorrhizal fungi are thought to have remained asexual for 400 million years, although recent studies have suggested that considerable genetic and phenotypic variation could potentially exist in populations. A brief discussion of these multigenomic organisms is presented.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 59–60.     

Phylogeography of five Polytrichum species within Europe

Using allozymes and microsatellites we have analysed the genetic structure among European populations for several Polytrichum species to infer relevant factors, such as historical events or gene flow, that have shaped their genetic structure. As we observed low levels of genetic differentiation among populations, and no decreasing levels of genetic variation with increasing latitude within most of the examined species, no genetic evidence was obtained for a step-wise recolonization of Europe from southern refugia after the latest glacial period for P. commune , P. uliginosum , P. formosum and P. piliferum . The near absence of population substructuring within these species does indicate that extensive spore dispersal is the most important factor determining the genetic structure among European Polytrichum populations. Gene flow levels have apparently been sufficient to prevent genetic differentiation among populations caused by genetic drift, and to wipe out any genetic structure caused by the postglacial recolonization process. On the other hand, increased genetic differentiation of alpine P. formosum populations suggests that mountain ranges might restrict gene flow significantly among Polytrichum populations. In contrast to most examined Polytrichum species, P. juniperinum showed high levels of genetic differentiation and a profound genetic structure. Assuming that gene flow is not more restricted in P. juniperinum , these findings suggest that this species has recolonized Europe after the latest glacial period from two different refugia, one possibly being the British Isles.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society 2003, 78, 203–213.     

Festuca inops and Festuca gracilior (Poaceae): are they two different species?

According to current systematics, Festuca inops and F. gracilior are two distinct species. However, they are hardly distinguishable from each other on the basis of their morphological characters. Festuca inops is considered a diploid species endemic to Italy, while F. gracilior has a discontinuous distribution area, apparently related to chromosomal levels: diploid populations in Italy and south-east France, tetraploid populations in north-east Spain. The diploid populations of both taxa from Italy and south-east France are investigated in the present study. Nearly 1000 exsiccata were examined and morphometric analysis was carried out on macro- and micromorphological features of 119 specimens (including type-specimens) and on 20 natural populations (including loci classici ). All these data showed that the two species should be referred to a single taxon, for which the rank of species seems to be appropriate. This result is supported by karyological, ecological and chorological data and was confirmed by the results of ISSR analysis. According to nomenclatural rules, the legitimate name for the species is Festuca inops De Not.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 239–258.     

Morphometric study and taxonomy of Genista pulchella Vis. s.l. (Fabaceae), a south European species

A morphometric study of the known populations of Genista pulchella has been undertaken, based on herbarium specimens and field research. This has made it possible to provide a new taxonomic outline as the disjunct populations of G. pulchella (western part of the Balkan peninsula, central Italy and southern France) seem to be distinct. A new taxon is described: G. pulchella ssp. aquilana ssp. nov. (central Italy), a new combination G. pulchella ssp. villarsiana comb. nov. is proposed for the population from southern France, while G. pulchella ssp. pulchella can be considered endemic to the Balkan Peninsula. Genista pulchella Vis., G. villarsii Clementi and G. villarsiana Jord. are lectotypified.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 245–254.     

Saponaria pumila (caryophyllaceae) and the ice age in the European alps

The polymerase chain reaction (PCR)-based amplified fragment length polymorphism (AFLP) technique was applied to elucidate the glacial history of the alpine cushion plant Saponaria pumila in the European Alps. Special emphasis was given to a dense sampling of populations. Our data support a survival of S. pumila during the last ice age in at least three refugia, which are characterized by unique marker sets. Patterns of genetic diversity and divergence can be explained by survival in peripheral refugia and additional in situ survival within the ice sheet on peripheral nunataks. A nunatak survival in interior parts of the Alps needs not be postulated to explain our results. The level of genetic diversity is dramatically different between populations (Shannon's diversity index: 0.87-19.86). Some peripheral populations are characterized by a high number of rare fragments indicating long isolation, but not necessarily by a high level of genetic diversity. Parts of the present distributional area were recolonized via recent long-distance dispersal, leading to severely bottlenecked populations lacking private or rare fragments. The combination of our data with palaeogeological and palaeoclimatological evidence allows us to confine Pleistocene refugia to certain regions and to draw a detailed scenario of the glacial and postglacial history of S. pumila.     

Several Pleistocene refugia detected in the high alpine plant Phyteuma globulariifolium sternb & hoppe (Campanulaceae) in the European Alps

Phyteuma globulariifolium is a high alpine plant species growing in the European Alps and the Pyrenees. In order to elucidate its glacial history, 325 individuals from 69 populations were analysed using the amplified fragment length polymorphism (AFLP) technique. A strongly hierarchical phylogeographical pattern was detected: Two major east-west vicariant groups can be separated along a gap in the distributional area. A further subdivision into at least four populational groups is in congruence with presumed peripheral glacial refugia. There is no indication for survival on unglaciated mountain tops (nunataks) in the interior of the Pleistocene ice shield covering the Alps. Our results favour glacial survival in peripheral, unglaciated or not fully glaciated areas. Populations of P. globulariifolium in the Pyrenees are the result of relatively recent long-distance dispersal. Within the Alps, there is strong differentiation among groups of populations, whereas within them the differentiation is weak. This suggests high levels of gene-flow over short to middle distances.     

Satyrinae butterflies from Sardinia and Corsica show a kaleidoscopic intraspecific biogeography (Lepidoptera,Nymphlidae)

The Mediterranean islands of Sardinia and Corsica are known for their multitude of endemics. Butterflies in particular have received much attention. However, no comprehensive studies aiming to compare populations of butterflies from Sardinia and Corsica with those from the neighbouring mainland and Sicily have been carried out. In the present study, the eleven Satyrinae species inhabiting Sardinia and Corsica islands were examined and compared with continental and Sicilian populations by means of geometric morphometrics of male genitalia. Relative warp computation, discriminant analyses, hierarchical clustering, and cross‐validation tests were used to identify coherent distributional patterns including both islands and mainland populations. The eleven species showed multifaceted distributional patterns, although three main conclusions can be drawn: (1) populations from North Africa and Spain are generally different from those belonging to the Italian Peninsula; (2) populations from Sardinia and Sicily often resemble the North Africa/Spain ones; Corsica shows transitional populations similar to those from France; and (3) sea barriers represent filters to dispersal, although their efficacy appears to be unrelated to their extension. Indeed, the short sea straits between Sardinia and Corsica and between Sicily and the Italian Peninsula revealed a strong effectiveness with respect to preventing faunal exchanges; populations giving onto sea channels between Corsica and Northern Italy and between Sicily and Tunisia showed a higher similarity. A comparison of island and mainland distributions of the eleven taxa have helped to unravel the complex co‐occurrence of historical factors, refugial dynamics, and recent (post‐glacial) dispersal with respect to shaping the populations of Mediterranean island butterflies. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 195–212.     

Comparative analysis of biological and ecological differentiation of Anemone palmata L. (Ranunculaceae) in the western Mediterranean (France and Spain): an assessment of rarity and population persistence

The western Mediterranean geophyte Anemone palmata L. is one of the most endangered angiosperm taxa in France. The biological and ecological characteristics of French populations are contrasted with those of populations from near the centre of the species distribution in east Spain. A Correspondence Analysis discriminates the French and Spanish populations according to substrate and the composition of the plant communities where they grow. The karyological study reveals that east Spanish plants are autotetraploid, whilst the French are all diploid. In addition, morphological differences were registered, Spanish plants generally being more vigorous. French populations consist mostly of senile plants, without or with very low rates of sexual reproduction, whereas the populations from Spain include all age groups with an important percentage of juvenile plants. The main factors which could affect the French populations are habitat destruction and modification of plant communities due to land-abandonment. Conservation measures should be supported by immediate protection of the French localities and habitats to prevent the total decline of this rare species.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 95–114.     

Some remarks on the genus Astragalus sect. Incani in Iran

New findings are given for the Astragalus sect. Incani , which occurs within the area covered by the Flora of Iran. Astragalus wojciechowskianus Ranjbar is described as a new species. A. procerus Boiss. & Hausskn . , A. cottonianus Aitch. & Baker, A. achundovii Grossh., A. orduabadensis Grossh. and A. glaucophyllus Bunge are reported as new records for the flora of Iran .   © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 443–447.