Habitat-related genetic substructuring in a marine snail (Littorina fabalis) involving a tight link between an allozyme and a DNA locus

Metapopulation structure and genetic differentiation among subpopulations will be tightly related to patterns and processes of local adaptation and microevolution. Understanding the mechanisms behind genetic substructuring will aid in the interpretation of species' ecological performances and strategies. The marine gastropod Littorina fabalis occurs in two size morphs – a small and a large – found in microhabitats of different wave exposure, but overlapping in distribution where wave exposure is intermediate. Earlier studies have found substantial genetic differentiation linked to morph in one allozyme locus (arginine kinase), while 29 other allozyme loci reveal no or minute differences between morphs. Here we add new results showing DNA variation in a RAPD marker being tightly linked to the allozyme variation. Indeed, 97% of the snails homozygotic for one of the Ark alleles had a unique DNA band, while 89% of the snails homozygotic for the other Ark allele lacked the marker. We discuss alternative hypotheses explaining the genetic substructure and suggest that the linkage of size, allozyme and DNA traits might be due to a paracentric chromosomal inversion involving loci coding for these traits. A genetic linkage of traits might promote microhabitat specialization of this species, and such a chromosomal transformation may therefore be adaptive.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 301–306.     

Cryptic speciation, genetic diversity and gene flow in the California turret spider Atypoides riversi (Araneae: Antrodiaetidae)

Populations of the turret spider Atypoides riversi from eight central Californian sites were compared based on variation at ten allozyme loci. Multidimensional scaling of interpopulation genetic distances defined four population units (Coast Range, Sierran, Valley, Jenness Camp), corroborating the distinctness of Coast Range and Sierran populations indicated by a prior study. While the species status of these units has yet to be determined, Jenness Camp is the most likely to represent a new species, given its clear genetic uniqueness (two fixed allelic differences). Populations in all units were generally in Hardy–Weinberg equilibrium with no evidence of inbreeding, though variability was minimal (mean H _o = 2.8%, mean P  = 15.4%). Reduced variability in these populations may be the result of repeated bottlenecks, environmental homogeneity, and/or directional selection. Interpopulation differentiation within units was significant in the absence of intervening forest habitat and was substantially less in its presence, indicating that gene flow is likely only when forest corridors exist. To foster preservation of the existing gene pool and enhance participation in it, management of the units of At. riversi should focus on maintaining as many populations in situ as possible and facilitating connections between them, while also creating or restoring habitat for potential colonization.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 27–37.     

Allozymic differentiation of Tunisian populations of Androctonus species and Buthus occitanus (Scorpiones: Buthidae)

We examined the population structure and phylogeny of Buthus occitanus (Amoreux, 1789), Androctonus amoreuxi (Audouin & Savigny 1812) and Androctonus australis (Linnaeus, 1758) in Tunisia, using horizontal starch gel and polyacrylamide gel electrophoresis to resolve allozymes at 18 loci. Populations of B. occitanus and A. amoreuxi showed little genetic variation, whereas A. australis populations showed moderate genetic variation. Moreover, despite the restricted dispersal abilities of these species, the allozyme analysis indicated very low levels of intraspecific population differentiation. The Island model of population differentiation best fitted the intraspecific variation, both for mainland populations and for the slightly more differentiated populations from two Mediterranean islands. No diagnostic, fixed alleles were observed between the species of Androctonus , which concurs with the results of nuclear ribosomal DNA analysis. Two subspecies of A. australis , A. a. garzonii Goyffon & Lamy, 1973 and A. a. hector Vachon, 1948, were not resolved by a phylogenetic analysis of the allozyme data. The most likely reasons for the slight population differentiation are either low mutation rates or recent range extensions promoted by human activities. The possibility of indirect gene flow between Androctonus species should also be explored.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 255–265.     

Molecular and morphometric data demonstrate the presence of cryptic lineages among freshwater crabs (Decapoda: Potamonautidae: Potamonautes) from the Drakensberg Mountains, South Africa

The evolutionary importance of cryptic taxa is well documented, yet few studies have examined them with a view to conservation. In the present study, the significance of cryptic speciation in freshwater crabs is examined. Allozyme and 16S rRNA sequence data were used to explore the degree of population differentiation between mountain stream populations of two distinct freshwater crab species. Marked patterns of differentiation were evident among populations; those in close geographical proximity were characterized by the fixation of alternate alleles at certain loci, indicating that currently there is no gene flow among populations. Both allozyme and sequence data provide evidence for the recognition of at least five distinct evolutionary lineages with pronounced levels of genetic differentiation. Morphometric data failed to detect any distinct geographically intermediate population groupings. Our findings indicate the presence of five phylogeographical units, all worthy of conservation, three of which are evolutionarily significant.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 129–147.     

Molecular and morphological data reveal cryptic taxonomic diversity in the terrestrial slug complex Arion subfuscus/fuscus (Mollusca, Pulmonata, Arionidae) in continental north-west Europe

The importance and abundance of cryptic species among invertebrate taxa is well documented. Nowadays, taxonomic, phylogenetic and conservation biological studies frequently use molecular markers to delineate cryptic taxa. Such studies, however, often face the problem of the differential resolution of the molecular markers and techniques involved. This issue is explored in the present study of cryptic taxa within the terrestrial slug complex Arion subfuscus/fuscus in continental north-west Europe. To this end, morphological, allozyme and mitochondrial 16S rDNA sequence data have been jointly evaluated. Using allozyme data and gonad type, two distinct groups were consistently delineated, even under sympatric conditions. The 16S rDNA data strongly supported both those groups and even suggested the presence of three distinct taxa within one of them. However, in view of: (1) the allopatric distribution of three OTUs, (2) the lack of allozyme or morphological differentiation, and (3) the extremely high degree of intraspecific mtDNA variation reported in pulmonate gastropods, they are, for the time being, not regarded as valid species under the biological species concept. By means of 16S rDNA and allozyme data, the position of type and topotype material of A. subfuscus s.s. and A. fuscus relative to the newly defined OTUs was determined, thus clarifying the nomenclature of this species complex. Additionally, gonad type proved to be a useful character for distinguishing the two species in north-west Europe.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 23–38.     

Species diversity and endemism in the Daphnia of Argentina: a genetic investigation

Although the temperate regions of South America are known to have a diverse daphniid fauna, there has been no genetic evaluation of the existing taxonomic system or of the affinities between the North and South American faunas. The present study analyses mitochondrial DNA sequences and allozyme variation to investigate species diversity in 176 Daphnia populations from Argentina. This work established the presence of at least 15 species in Argentina, six of which are either undescribed or are currently misidentified and two of which represent range extensions of North American taxa. Eleven of the Argentine species appear endemic to South America, while the remaining four also occur in North America. In the latter cases, the close genetic similarity between populations from North and South America indicates the recent exchange of propagules between the continents. While biological interactions and habitat availability have undoubtedly contributed to the observed species distributions, chance dispersal has apparently played a dominant role in structuring large-scale biogeographical patterns in this genus and probably in other passively-dispersed organisms.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 171−205.     

Life history and allozyme diversification in regional ecomorphs of the Hyalella azteca (Crustacea: Amphipoda) species complex

In North America, several species in the freshwater amphipod genus Hyalella occur as one of two alternative phenotypic types, or 'ecomorphs', each possessing life history traits that allow success in alternative habitats that differ in predation regime. This study documents life history diversification, reproductive isolation and allozyme differentiation of Hyalella ecomorphs in Oklahoma, and compares these results to previously reported patterns of phenotypic and systematic diversification in Michigan. As in Michigan, two ecomorphs are common in Oklahoma, with an early maturing, small sized ecomorph found in habitats containing Lepomis sunfish, which prey on Hyalella , and a late reproducing, large sized ecomorph found in fishless habitats and in very shallow margins of large reservoirs. Allozyme analysis and laboratory interbreeding trials demonstrated that ecomorphs in Oklahoma are reproductively isolated species. Phenotypically, these species are very similar to species of the same ecomorph in Michigan. Large ecomorph species in the two regions differ substantially in allozyme allele composition in a pattern consistent with reproductive isolation, yet these species did not differ in a comparison of phenotype. The small ecomorph in Oklahoma is similar in phenotype to two of three small ecomorph species in Michigan. Overall, this study supports the hypothesis that Hyalella diversification in North America is characterized by the evolution of similar phenotypic solutions to comparable ecological challenges.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 161–175.     

Geographical variation and diversification in the flightless leaf beetles of the Chrysolina angusticollis species complex (Chrysomelidae, Coleoptera) in northern Japan

Geographical variation and differentiation of flightless leaf beetles of the Chrysolina angusticollis species complex were analysed. Nine morphological groups were recognized on the basis of metric characters relating to body size, shape of hind wing and shape of male genitalia, and binary characters relating to females' elytral surface and larval pubescence. The nine groups were essentially parapatric or allopatric to each other. Five groups occupied a continuous geographical range each, whereas another four groups were disjunctively distributed. Morphometric distance and geographical distance between populations were significantly correlated. Analyses based on 13 allozyme loci demonstrated that six out of the nine morphological groups were further divided into more than one genetically distinct group each. Genetic similarity between populations was significantly correlated with morphometric similarity and geographical closeness. Disjunctively distributed populations of a single morphological group more often resembled geographically adjacent populations of different morphological groups than geographically remote populations of the same group. In a morphological group that occupied a wide continuous range, remote populations substantially differed genetically from each other. The results suggest that there exist at least 19 morphologically and/or genetically differentiated units in the C. angusticollis complex in northern Japan. The C. angusticollis complex is likely a superspecies composed of a number of semispecies. We postulate that the diversification of this species complex might have resulted from the repeated range contraction and expansion during the ice ages. Movement of tension zones that separate two adjacent forms might contribute to enhance their geographical differentiation.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 557–578.     

Molecular differentiation between coexisting species of Taraxacum sect. Erythrosperma (Asteraceae) from populations in south-east and west Germany

The present study focuses on the genetic differentiation between six coexisting species of Taraxacum sect. Erythrosperma . Twenty-six individuals of T. rubicundum , T. brachyglossum , T. tortilobum , T. lacistophyllum , T. parnassicum and T. scanicum were sampled at 14 locations in south-east (Bavaria) and west Germany (Eifel). Using random amplified polymorphic DNA (RAPD) analysis 139 fragments were amplified. Nineteen of these were found to be species specific. The level of polymorphic bands within the different Taraxacum species varied between 0% and 15.8%. An analysis of molecular variance (AMOVA) revealed strong differentiation between the investigated species: 14.3% of the molecular variance were found within the species, while 85.7% of the genetic variability was observed between the species. However, a geographical differentiation between the different regions could not be observed. All analysed species were clearly separated from each other in a cluster analysis using the neighbour joining technique. The observed strong genetic differentiation between the Taraxacum species clearly supports the agamospecies concept of Taraxacum sect. Erythrosperma , which is at present mainly based on morphological and cytological criteria  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 109–117.     

Falling apart and merging: diversification of slender salamanders (Plethodontidae: Batrachoseps) in the American West

The plethodontid genus Batrachoseps , the slender salamanders, is the most diverse clade of salamanders in western North America, but it has posed taxonomic difficulties because it contains many morphologically cryptic species. A segment of the mitochondrial DNA gene cytochrome b was studied for 278 individuals densely sampled from throughout the range of all 18 described species and several undescribed species. Phylogenetic analyses of the mtDNA data identify six major clades, one corresponding to the subgenus Plethopsis and five within a monophyletic subgenus Batrachoseps. All major clades and most species within these clades display strong phylogeographic structuring. Comparisons of mtDNA and allozyme data show that several allozymically cohesive groups are not monophyletic with respect to mtDNA. We suggest that this phenomenon results from fragmentation of populations, divergence in allopatry, and then recontact and gradual merging of units caused predominantly by male-mediated gene flow. The mtDNA offers evidence that populations were once more isolated than they are now, while the patterns of allozyme variation reflect recent and current interactions among populations. The complex patterns of morphological, allozymic and mtDNA variation associated with the constantly changing geological landscape give insight into the nature of processes responsible for species formation in Batrachoseps .  © 2002 The Linnean Society of London. Biological Journal of the Linnean Society , 2002, 76 , 361–391.     

Natural hybridization does not dissolve species boundaries in commercially important sea cucumbers

The study of species boundaries in areas of sympatry provides important insight into speciation processes. We investigated whether (i) two sympatric holothurians, Holothuria scabra and H. s. var. versicolor constituted species, and (ii) specimens of intermediate phenotype hybrids. Results from allozyme and 16S mtDNA sequence analyses indicated these two sea cucumbers to be distinct but young biological and phylogenetic species. Several private allozyme alleles existed and a Bayesian analysis grouped varieties into separate clusters. MtDNA sequences hardly varied within each taxon, and nine single bp changes were diagnostic between these two taxa. Allozyme allele frequencies in individuals of intermediate phenotype were intermediate to those of H. scabra and H. s.  var.  versicolor , most private alleles were present and heterozygote frequencies were higher than in either species. Ancestry coefficients modelled for these individuals were close to 0.5, indicating that the two taxa contributed equally to their genome. MtDNA sequences were identical to those of either species. We conclude that individuals of intermediate phenotype represent F1 hybrids. The presence of hybrids demonstrates that the opportunity for introgression exists, but is not realized, as backcrossing and introgression were not supported by the data. Thus, the genetic integrity of either holothurian species remains intact through an unknown postzygotic mechanism, possibly hybrid sterility.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 261–270.     

Morphological and genetic evidence supports the existence of two species in the genus Ophelia (Annelida, Polychaeta) from the Western Mediterranean

About 900 individuals of the intertidal polychaete Ophelia cf. bicornis were collected at four sandy beaches in Corsica, ten in Sardinia and one in Tuscany (Western Mediterranean). They were analysed morphologically and genetically to clarify the taxonomic status of the littoral species of Ophelia in the study area and to assess the population genetic structuring. Two morphotypes were distinguished on the basis of the number of nephridiopore pairs (five or six). Genetic distances, F -statistics and analysis of molecular variance based on eight allozyme genetic markers were consistent in showing the two morphotypes to separate at species level. Thus, the two morphotypes which most authors have included in O. bicornis s.l. are not one morphologically variable taxon, but instead constitute two valid species, namely O. bicornis Savigny, 1818, with six pairs of nephridiopores, and O. barquii Fauvel, 1927, with five pairs of nephridiopores. Mechanisms of ecological speciation in a background of gene flow can be taken into account to explain the origin of the two species. Moderate genetic differentiation was found among populations within each species, suggesting that dispersal may not be effective over large distances, or that local selection plays an important role in population differentiation.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 101–113.     

Genetic variation and its evolutionary implications in a Mediterranean island endemic lizard

The peculiar bioclimatic and geographic features of Corso–Sardinian islands may provide an ideal scenario for investigating microevolutionary processes, given their large heterogeneity of environments, which could affect dispersal and gene flow among populations, as well as processes of local adaptation. The genetic variation and differentiation among populations of the endemic lizard Archaeolacerta bedriagae were studied by allozyme electrophoresis at 20 presumptive loci. The genetic structure of this species is characterized by relatively high levels of polymorphism and low differentiation among populations. The pattern of genetic differentiation cannot be explained by genetic drift as a function of geographic distance. Genetic distance data show that genetic variation is distributed into three geographically coherent population groups and suggest a recent (Late Pleistocene) origin for the observed geographic fragmentation. The analysis of environmental correlates of allozymic variation indicates a strong correlation of the Idh-1 locus with climatic variables. The frequency of the Idh-1¹⁰⁶ allele is negatively correlated with annual temperature, and positively correlated with annual precipitation. In addition, the observed heterozygosity at this locus decreases towards more arid climatic regimes. The results obtained support the assumption of differential selection acting on Idh-1 allozymes under diverse climates. An association between Idh-1 allozymes and local bioclimatic regimes was also observed for the sympatric lizard Podarcis tiliguerta , suggesting a key role for such selective agents on Idh-1 polymorphism in these two Corso–Sardinian lacertids.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 661–676.     

Genetic differentiation between populations of the European rose hip fly Rhagoletis alternata

We tested for genetic differentiation between populations of Rhagoletis alternata Fall. (Diptera: Tephritidae) on three different host species. We collected larvae from three rose species of the section Caninae ( Rosa canina L., Rosa corymbifera Borkh . , and Rosa rubiginosa L.) from 15 sites across Germany, where the three roses occurred together. Additionally, we sampled three sites in Switzerland. Roses differ in morphology (e.g. leaf glands) as well as phenology. We were able to score nine allozyme loci (five polymorphic). Populations from the three hosts did not differ in genetic variability. We found significant genetic differentiation between populations from different host species. However, the differentiation was very low (0.9%). Hence, we found no indication for host races. Furthermore, surprisingly little geographical structure of genetic differentiation was found between populations of this fruit fly across central Europe. We offer three mutually non-exclusive explanations for these findings. First, gene flow between populations of Rh. alternata is high. Second, the pattern of genetic differentiation is based on a recent expansion of the distributional range . Third, the ongoing gene flow between roses of the section Caninae acts as a hybrid bridge.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 619–625.     

Immigration history and gene dispersal: allozyme variation in Nordic populations of the red campion, Silene dioica (Caryophyllaceae)

Most of the Nordic region was ice-covered during the last (Weichselian) glaciation. During the postglacial period, plant and animal species recolonized the region from several directions and the geographic structuring of genetic variation within Nordic species may still contain a historic component that reflects patterns of postglacial immigration. The present investigation of 69 populations of Silene dioica represents the first large-scale allozyme study of a widespread herbaceous plant in the Nordic region. Although the frequencies of individual alleles showed a range of different geographic patterns, mapping of the axis scores from an ordination of variation at eight polymorphic loci revealed a division into two main geographic groups of populations. The broadly south-western and north-eastern distributions of these two groups of populations suggest that immigration into the region may have involved both eastern and southern geographic sources. However, the geographic boundaries between the two groups of populations are diffuse, and the relatively low between-population component of genetic diversity (G_ST = 16.4%) suggests a history of extensive gene dispersal by pollen.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 23–34.     

Extremely low levels of genetic diversity in the terrestrial orchid Epipactis thunbergii (Orchidaceae) in South Korea: implications for conservation

Levels of allozyme variation, population genetic structure, and fine-scale genetic structure (FSGS) of the rare, both sexually and clonally reproducing terrestrial orchid Epipactis thunbergii were examined for eight ( N  = 734) populations in a 20 × 20-km area in South Korea. Twenty-three putative allozyme loci resolved from 15 enzyme systems were used. Extremely low levels of allozyme variation were found within populations: the mean frequency of polymorphic loci was 3.8% [isocitrate dehydrogenase ( Idh-2 ) with two alleles was polymorphic across populations], the mean number of alleles per locus was 1.04, and the mean expected heterozygosity was 0.013. The overall fixation index was not significantly different from zero ( F _IS = 0.069), although the species is self-compatible. However, a significantly high degree of population differentiation was found between populations at Idh-2 ( F _ST = 0.388) in the studied area. Furthermore, spatial autocorrelation analyses revealed a significant FSGS (up to 3 m) within populations. These observations suggest that the main explanatory factors for the extremely low levels of genetic diversity and the shaping of the population genetic structure of E. thunbergii are genetic drift as a result of a small effective population size, a restricted gene flow, and the isolation of populations. Considering the current genetic structure of E. thunbergii , three guidelines are suggested for the development of conservation strategies for the species in South Korea: (1) protection of habitats of standing populations; (2) prohibition by law of any collection of E. thunbergii ; and (3) protection of nearby pollinator populations, given the fact that fruit set in natural habitats is very low.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 161–169.     

Systematics and evolutionary relationships of the mountain lizard Liolaemus monticola (Liolaemini): how morphological and molecular evidence contributes to reveal hidden species diversity

The delimitation of species is a major issue in systematic biology and has been a re-emerging discipline in the last decade. A number of studies have shown that the use of multiple data sets is critical for the identification of cryptic species, particularly in groups with complex evolutionary histories. Liolaemus monticola is a montane lizard species distributed in central Chile (32°–42°S), with four described subspecies in a latitudinal gradient from north to south: L. m. monticola , L. m. chillanensis , L. monticola ssp. and L. m. villaricensis . In order to test the systematic status and phylogenetic relationships of the taxa included in the L. monticola group, we analysed morphological (morphometric and meristic) and molecular (allozyme and mitochondrial DNA) data sets. The results of the morphological analyses showed that meristic variables correctly assigned individuals with higher accuracy than did morphometric characters. The results of the analyses of allozyme data revealed eight diagnostic loci that are evidence for significant differences among the four L. monticola subspecies. Phylogenetic analyses with mitochondrial DNA data, including additional species, showed that the L. monticola group is polyphyletic. We postulate that the four current subspecies represent independent evolutionary lineages and must be raised to the specific level as L. monticola , L. chillanensis and L. villaricensis . The taxonomic status of the unnamed L. monticola ssp. remains unresolved, although we provide a preliminary proposal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 635–650.     

Speciation in Darwin's darklings: taxonomy and evolution of Stomion beetles in the Galápagos Islands, Ecuador (Insecta: Coleoptera: Tenebrionidae)

Stomion is one of three speciose and taxonomically difficult genera of flightless Darwin's darkling beetles of the Galápagos Islands. The previously named species and subspecies of Stomion are analysed by morphometric and cladistic methods. Eleven species are here recognized, which probably descended from colonization by a single ancestral species. Cladistic analysis yielded two weakly supported clades that are different from previously published morphological and allozyme genotype groups. Phenotypic and genotypic characters may be evolving independently. There is much within-species variation, but there is no compelling evidence of adaptive radiation in Stomion , in contrast to that in the famous Darwin's finches. The analyses show the need for changes in morpho-species taxonomy: S. punctipennis Van Dyke ( syn. nov. ) is elevated to full species from a subspecies of S. galapagoensis Waterhouse. There is no evidence to support the validity of: S. laevigatum santacruzensi Franz ( syn. nov. ), synonomized here under S. l. laevigatum Waterhouse; S. galapagoensis leleupi Kaszab ( syn. nov. ), synonomized here under S. linelli Blair; S. galapagoensis pinzoni Franz ( syn. nov. ), synonomized here under S. obesum Van Dyke. S. genovesa sp. nov. is described from Genovesa Island. Lectotypes are designated for S. galapagoensis Waterhouse, S. laevigatum Waterhouse and S. helopoides Waterhouse .   © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 135–152.     

Phylogeography of the freshwater crayfish Cherax destructor Clark (Parastacidae) in inland Australia: historical fragmentation and recent range expansion

The yabby, Cherax destructor Clark, is the most widespread species in the most widespread genus of Australian freshwater crayfish. It has a distribution that spans several distinct drainage basins and biogeographical regions within semiarid and arid inland Australia. Here we report a study designed to investigate patterns of genetic variation within the species and hypotheses put forward to account for its extensive distribution using DNA sequences from the mitochondrial 16S rRNA gene region. Results of phylogenetic analyses contradicted previous allozyme data and revealed relatively deep phylogenetic structure in the form of three geographically correlated clades. The degree of genetic divergences between clades (8–15 bp) contrasted with the relatively limited haplotype diversity within clades (1–3 bp). Network-based analyses confirmed these results and revealed genetic structure on both larger and more restricted geographical scales. Nevertheless some haplotypes and 1-step clades had large distributions, some of which crossed boundaries between river basins and aquatic biogeographical regions. Thus both older and more recent historical processes, including fragmentation on a larger geographical scale and more recent range expansion on a local scale, appear to be responsible for the observed pattern of genetic variation within C. destructor . These results support elements of alternative hypotheses previously put forward to account for the evolutionary history of C. destructor and the origin of its large distribution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 539–550.