The hidden diversity of Coleodactylus amazonicus (Sphaerodactylinae,Gekkota) revealed by molecular data

Coleodactylus amazonicus, a small leaf-litter diurnal gecko widely distributed in Amazon Basin, has been considered a single species with no significant morphological differences between populations along its range. A recent molecular study, however, detected large genetic differences between populations of central Amazonia and those in the easternmost part of the Amazon Basin, suggesting the presence of taxonomically unrecognised diversity. In this study, DNA sequences of three mitochondrial (16S, cytb, and ND4) and two nuclear genes (RAG-1, c-mos) were used to investigate whether the species currently identified as C.amazonicus contains morphologically cryptic species lineages. The present phylogenetic analysis reveals further genetic subdivision including at least five potential species lineages, restricted to northeastern (lineage A), southeastern (lineage B), central-northern (lineage E) and central-southern (lineages C and D) parts of Amazon Basin. All clades are characterized by exclusive groups of alleles for both nuclear genes and highly divergent mitochondrial haplotype clades, with corrected pairwise net sequence divergence between sister lineages ranging from 9.1% to 20.7% for the entire mtDNA dataset. Results of this study suggest that the real diversity of “C. amazonicus” has been underestimated due to its apparent cryptic diversification.     

Pronounced phylogeographic structure on a small spatial scale: Geomorphological evolution and lineage history in the salamander ring species Ensatina eschscholtzii in central coastal California

The salamander Ensatina eschscholtzii is a classic example of a ring species, and has an intricate biogeographic history. Within a part of the ring distribution, earlier work using allozymes disclosed high levels of genetic structure in central coastal California, where the subspecies oregonensis, xanthoptica, and eschscholtzii meet. We used mitochondrial cytochrome b sequences to further examine patterns of divergence in this area, including data from 155 localities (309 individuals). Our focus is on the documentation of population-level haplotype lineages. We show that oregonensis is represented by two unrelated, phenotypically similar clades, both of which possess substantial substructure of their own. The subspecies xanthoptica includes two lineages that differ in phenotype, one of which has colonized the foothills of the Sierra Nevada. The subspecies eschscholtzii occurs mainly to the south, but some populations from a northern lineage extend into the Monterey Bay region, where they approach xanthoptica geographically. In sum, populations in the central coastal California region form a distributional patchwork, including three subspecies, three clades (which differ from the three subspecies), and ten haplotype lineages. We conclude that such striking levels of phylogeographic structure reflect interspersed episodes of spatial fragmentation, in part driven by the complex geomorphological evolution of the California Coast Range system.     

Vocal Diversity and Taxonomy of <Emphasis Type="Italic">Nomascus</Emphasis> in Central Vietnam and Southern Laos

Previous researchers suggested that gibbon song repertoire is genetically determined and song characteristics are useful for assessing systematic relationships. The southern white-cheeked crested gibbon is regarded as either a subspecies of Nomascus leucogenys or its own species (Nomascus siki). I studied vocal diversity among different wild populations of Nomascus in central Vietnam and southern Laos to assess their taxonomic relationships and to examine whether their vocal patterns correspond to forms previously described for Nomascus siki. I examined the songs of 7 Nomascus populations in Vietnam and Laos. I analyzed 192 song bouts from different gibbon groups including 173 phrases of 42 females and 192 phrases of 42 males. Linear discriminant analysis, classification trees, and multidimensional scaling revealed marked separation of groups in the northern and southern populations. Within the 2 geographic populations, there is little variability and the vocal characteristics exhibited no apparent cline. I conclude that the northern and southern geographic populations may represent 2 distinct taxa. I postulate that a taxonal boundary such as large rivers existing between southern Quang Binh province and northern Thua-Thien Hue province in Vietnam and northern Phou Xang He NBCA and southern Dong Phou Vieng NBCA in Laos has limited gene flow between the populations. Differing topographic features could also serve as a selective force for improved sound transmission in a highly territorial species, driving the divergence between the 2 populations.     

Population genetic structure and demography of Magnolia kobus: variety borealis is not supported genetically

Species delimitations by morphological and by genetic markers are not always congruent. Magnolia kobus consists of two morphologically different varieties, kobus and borealis. The latter variety is characterized by larger leaves than the former. For the conservation of M. kobus genetic resources in natural forests, the relationships between morphological and genetic variation should be clarified. We investigated variations in nuclear microsatellites, chloroplast DNA (cpDNA) sequences and leaf morphological traits in 23 populations of M. kobus over the range of species. Two genetically divergent lineages, northern and southern were detected and their geographical boundary was estimated to be at 39°N. The northern lineage consisted of two genetic clusters and a single cpDNA haplotype, while the southern one had multiple genetic clusters and cpDNA haplotypes. The northern lineage showed significantly lower genetic diversity than the southern. Approximate Bayesian computation indicated that the northern and southern lineages had experienced, respectively, population expansion and long-term stable population size. The divergence time between the two lineages was estimated to be 565,000 years ago and no signature of migration between the two lineages after divergence was detected. Ecological niche modeling showed that the potential distribution area in northern Japan at the last glacial maximum was very small. It is thus considered that the two lineages have experienced different population histories over several glacial-inter-glacial cycles. Individuals of populations in the central to northern part of Honshu on the Sea of Japan side and in Hokkaido had large leaf width and area. These leaf characteristics corresponded with those of variety borealis. However, the delimitation of the northern and southern lineages detected by genetic markers (39°N) was not congruent with that detected by leaf morphologies (36°N). It is therefore suggested that variety borealis is not supported genetically and the northern and southern lineages should be considered separately when identifying conservation units based not on morphology but on genetic markers.

     

Mitochondrial evidence uncovers a refugium for the fat dormouse (Glis glis Linnaeus, 1766) in Hyrcanian forests of northern Iran

Fat dormouse is a squirrel-like rodent which is closely tied to deciduous forest ecosystem in southwestern Eurasia. As such it is a valuable indicator of forest survival in refugia during glacial-interglacial periods. Previous phylogeographic analyses uncovered divergent fat dormouse lineages in southern refugia in Italy and the Balkans, but retrieved a surprisingly low overall genetic diversity across the majority of the species’ range. We explored 812 bp long fragment of a cytochrome b (cyt b) gene in ten fat dormice from refugial Hyrcanian forests in northern Iran. We identified 10 new cyt b haplotypes, which generated a total dataset of 28 fat dormouse haplotypes. The phylogenetic reconstruction clustered the new haplotypes into the Iranian lineage which hold a sister position against all other fat dormouse haplotypes from Europe and Asia Minor. The divergence between these lineages suggests a fragmentation event of an ancestral population at 5.76 mya (95% HPD = 3.21–8.92). This early evolutionary divergence was possibly triggered in the Middle East by dramatically divergent environmental conditions at the Messinian Salinity Crisis. The divergence clearly exceeds the intraspecific divergence, and is well within the range between congeneric rodent species. We suggest a long-term persistence of the Iranian lineage in the Hyrcanian refugium which is consitent with a high number of endemics along the southern Caspian coastal areas.     

A new cytochrome b phylogroup of the common vole (Microtus arvalis) endemic to the Balkans and its implications for the evolutionary history of the species

The phylogeographic architecture of the common vole, Microtus arvalis, has been well‐studied using mitochondrial DNA and used to test hypotheses relating to glacial refugia. The distribution of the five described cytochrome b (cyt b) lineages in Europe west of Russia has been interpreted as a consequence of postglacial expansion from both southern and central European refugia. A recently proposed competing model suggests that the ‘cradle’ of the M. arvalis lineages is in western central Europe from where they dispersed in different directions after the Last Glacial Maximum. In the present study, we report a new cyt b lineage of the common vole from the Balkans that is not closely related to any other lineage and whose presence might help resolve these issues of glacial refugia. The Balkan phylogroup occurs along the southern distributional border of M. arvalis in central and eastern Bosnia and Herzegovina, Montenegro, and eastern Serbia. Further north and west in Slovenia, Bosnia and Herzegovina, and Serbia, common voles belong to the previously‐described Eastern lineage, whereas both lineages are sympatric in one site in Bosnia and Herzegovina. The Balkan phylogroup most reasonably occupied a glacial refugium already known for various Balkan endemic species, in contrast to the recently proposed model. South‐east Europe is an absolutely crucial area for understanding the postglacial colonization history of small mammals in Europe and the present study adds to the very few previous detailed phylogeographic studies of this region. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 788–796.     

Definition and review of the lancangjiang species group of the termitophilous genus Orthogonius Macleay, 1825 (Coleoptera,Carabidae, Orthogoniini)

The lancanjiang species group of the termitophilous ground beetle genus Orthogonius Macleay, 1825, is defined and reviewed. This group is characterized by the black and rather elongate body, dense punctation on head and elytra, long and slender appendages, thin fore tibiae, and elytral interval 3 without a subapical setiferous pore. To date, the lancanjiang species group is composed of four species and one subspecies, including three new species and one new subspecies which are described in the present paper: O. macrophthalmus sp. n. (northern Vietnam), O. euthyphallus sp. n. (southern Vietnam), O. euthyphallus bolavenensis ssp. n. (southern Laos) and O. carinatus sp. n. (northern Laos). A distribution map and a key to all species of this group are also provided.     

Speciation on the Rocks: Integrated Systematics of the Heteronotia spelea Species Complex (Gekkota; Reptilia) from Western and Central Australia

The isolated uplands of the Australian arid zone are known to provide mesic refuges in an otherwise xeric landscape, and divergent lineages of largely arid zone taxa have persisted in these regions following the onset of Miocene aridification. Geckos of the genus Heteronotia are one such group, and have been the subject of many genetic studies, including H. spelea, a strongly banded form that occurs in the uplands of the Pilbara and Central Ranges regions of the Australian arid zone. Here we assess the systematics of these geckos based on detailed examination of morphological and genetic variation. The H. spelea species complex is a monophyletic lineage to the exclusion of the H. binoei and H. planiceps species complexes. Within the H. spelea complex, our previous studies based on mtDNA and nine nDNA loci found populations from the Central Ranges to be genetically divergent from Pilbara populations. Here we supplement our published molecular data with additional data gathered from central Australian samples. In the spirit of integrative species delimitation, we combine multi-locus, coalescent-based lineage delimitation with extensive morphological analyses to test species boundaries, and we describe the central populations as a new species, H. fasciolatus sp. nov. In addition, within the Pilbara there is strong genetic evidence for three lineages corresponding to northeastern (type), southern, and a large-bodied melanic population isolated in the northwest. Due to its genetic distinctiveness and extreme morphological divergence from all other Heteronotia, we describe the melanic form as a new species, H. atra sp. nov. The northeastern and southern Pilbara populations are morphologically indistinguishable with the exception of a morpho-type in the southeast that has a banding pattern resembling H. planiceps from the northern monsoonal tropics. Pending more extensive analyses, we therefore treat Pilbara H. spelea as a single species with phylogenetic structure and morphological heterogeneity.     

New species of Peliosanthes,Rohdea and Tupistra (Asparagaceae) from Laos and Vietnam

We describe and illustrate four new species, Peliosanthes choriandra, P. tatianae and Tupistra orlovii from central to northern Laos, and Rohdea filosa from northern Vietnam. These are all very local in distribution and endemic to the respective countries. We also report new localities and their ecological conditions for five other species of Peliosanthes (P. argenteostriata, P. hirsuta, P. irinae, P. micrantha and P. nivea) recently described from Laos and/or Vietnam. Furthermore, Peliosanthes nivea is recorded as new to the flora of Laos.     

Genomic divergence in sympatry indicates strong reproductive barriers and cryptic species within Eucalyptus salubris

Genetic studies are increasingly detecting cryptic taxa that likely represent a significant component of global biodiversity. However, cryptic taxa are often criticized because they are typically detected serendipitously and may not receive the follow‐up study required to verify their geographic or evolutionary limits. Here, we follow‐up a study of Eucalyptus salubris that unexpectedly detected two divergent lineages but was not sampled sufficiently to make clear interpretations. We undertook comprehensive sampling for an independent genomic analysis (3,605 SNPs) to investigate whether the two purported lineages remain discrete genetic entities or if they intergrade throughout the species’ range. We also assessed morphological and ecological traits, and sequenced chloroplast DNA. SNP results showed strong genome‐wide divergence (F _ST = 0.252) between two discrete lineages: one dominated the north and one the southern regions of the species’ range. Within lineages, gene flow was high, with low differentiation (mean F _ST = 0.056) spanning hundreds of kilometers. In the central region, the lineages were interspersed but maintained their genomic distinctiveness: an indirect demonstration of reproductive isolation. Populations of the southern lineage exhibited significantly lower specific leaf area and occurred on soils with lower phosphorus relative to the northern lineage. Finally, two major chloroplast haplotypes were associated with each lineage but were shared between lineages in the central distribution. Together, these results suggest that these lineages have non‐contemporary origins and that ecotypic adaptive processes strengthened their divergence more recently. We conclude that these lineages warrant taxonomic recognition as separate species and provide fascinating insight into eucalypt speciation.     

Coalescent species delimitation of a Sumatran parachuting frog

We conduct species delimitation of the widespread parachuting frog species Rhacophorus catamitus using samples from across the island of Sumatra, Indonesia. We use mitochondrial, genomic and morphological data, and find that R. catamitus is composed of three lineages corresponding to northern, central and southern lineages. Mitochondrial and genomic data show admixture or incomplete lineage sorting between the central and southern lineages, but deep divergence from the northern lineage. Coalescent species delimitation supports a three species model for this complex, and we recommend that the northern lineage be described as a new species. Our study highlights the power of coalescent species delimitation in an integrative framework for identifying unrecognised diversity in understudied tropical species complexes. We also emphasise the evolutionary importance of northern Sumatra, a region that harboured montane refugia during Pliocene–Pleistocene climate change, but has also been heavily affected by volcanic activity.     

CHLOROPLAST-DNA VARIATION IN TELLIMA GRANDIFLORA (SAXIFRAGACEAE)

Tellima grandiflora, a herbaceous, diploid (2n = 14) perennial, is distributed from the peninsula and panhandle of Alaska to central California. Restriction site variation of chloroplast DNA was surveyed in 51 populations representing the geographic range of T. grandiflora using 20 endonucleases. Two well-differentiated clades of populations differing by 19 restriction site mutations and several length mutations are geographically structured. A northern group comprises populations from Alaska to central Oregon; populations from central Oregon to San Francisco, California, form a southern group. The southern lineage of the monotypic Tellima appears to have obtained its chloroplast genome via ancient hybridization with a species of Mitella. Although northern and southern lineages have well-differentiated chloroplast genomes, populations of T. grandiflora show a high degree of genetic similarity of nuclear-encoded allozymes; furthermore, no apparent morphological differences characterize the lineages. Significantly, several populations of T. grandiflora that possess the typical southern chloroplast genome also occur disjunctly on Prince of Wales Island, Alaska, and the Olympic Peninsula, Washington. Because both areas are proposed glacial refugia, we suggest that past glaciation may have created discontinuities in the geographic distribution of T. grandiflora. Following glaciation, migration of once-isolated populations possessing different chloroplast genomes resulted in the formation of a continuous geographic distribution with a major organellar discontinuity. Additional support for this hypothesis is provided by the presence of well-differentiated northern and southern chloroplast DNA lineages in Tolmiea menziesii, a species having a geographic distribution and life history traits similar to those of Tellima.     

Playing hide-and-seek with the tiny dragonfly: DNA barcoding discriminates multiple lineages of <Emphasis Type="Italic">Nannophya pygmaea</Emphasis> in Asia

We examined the utility of DNA barcode data for assessing genetic diversity of the tiny dragonfly Nannophya pygmaea Rambur in Asia. Data analyses inferred from the barcode region of cytochrome oxidase subunit I (COI) were performed with Malaysian N. pygmaea, along with the existing COI haplotypes distributed in Asia. We applied four species delimitation analyses [automatic barcode gap discovery (ABGD), generalised mixed yule coalescent (GMYC), poisson tree processes maximum likelihood (PTP_ML) and poisson tree processes simple heuristic solutions (PTP_sh)] to investigate potential lineages in this geographically widespread species. Based on our dataset, we provisionally recognize four distinct lineages or operational taxonomic units of N. pygmaea, which were represented by the taxa from Japan/Korea, China/Laos/Taiwan, Malaysia and Vietnam, respectively. Phylogenetic analyses showed two well-supported assemblages of N. pygmaea: one restricted to the taxa from Malaysia and Vietnam; and the other covering all populations further north (i.e., China, Japan, Korea, Laos and Taiwan). An extraordinarily high degree of genetic distance (up to >12 %) was detected between these two assemblages—suggesting they represent two separate species.     

New data on the distribution and intraspecific variation of the Millard’s giant rat <Emphasis Type="Italic">Dacnomys millardi</Emphasis> (Mammalia,Rodentia) from Vietnam

The paper presents an analysis of a genetic and morphological variation of Dacnomys millardi from seven Vietnamese localities, including a new finding from southern Vietnam which lies some 1000 km to the south of the currently known limit of the species range. Samples from northern and southern Vietnam do not differ significantly in their cranial and oversize characteristics. Surprisingly, they are quite similar genetically as well. The genetic distances between the samples are very low and do not exceed 0.0232 for cyt b, 0.0058 for subunit I of the cytochrome oxidase gene, 0.0019 for interphotoreceptor retinoid-binding protein, and 0.0017 for recombination activation factor gene. A low overall genetic diversity in the D. millardi could reflect its ecological specialization, as well as a population bottleneck or the colonization history of eastern Indochina.     

The genetic architecture of hybridisation between two lineages of greenshell mussels

A multidisciplinary approach has identified sigmoidal genetic clines on the east and west coasts in central New Zealand where low-density ecological interactions occur between northern and southern lineages of the endemic greenshell mussel, Perna canaliculus. The sigmoidal clines indicate the existence of a mussel hybrid zone in a region of genetic discontinuities for many continuously distributed coastal taxa, in particular marine invertebrates. Examination of the genetic architecture of the hybrid zone revealed the differential contribution of individual microsatellite loci and/or alleles to defining the zone of interaction and no evidence of increased allelic richness or heterozygosity inside versus outside the hybrid zone. Genomics cline analysis identified one locus in particular (Pcan1–27) as being different from neutral expectations, thereby contributing to lineage differentiation. Estimates of contemporary gene flow revealed very high levels of within-lineage self-recruitment and a hybrid zone composed mostly (~85%) of northern immigrants. Broad scale interpretation of these results is consistent with a zone of genetic interaction that was generated between 0.3 and 1.3 million years before present at a time of pronounced global sea-level change. At that time, the continuous distribution of the greenshell mussel was split into northern and southern groups, which differentiated to become distinct lineages, and which have subsequently been reunited (secondary contact) resulting in the generation of the hybrid zone at ~42°S.     

The potential key seed-dispersing role of the arboreal marsupial Dromiciops gliroides

Marsupial seed dispersal is a rare phenomenon, although it may be ecologically significant in southern South America. The marsupial Dromiciops gliroides is endemic to the northern part of the temperate forest of South America. Here we describe the food habits and examine the potential role of D. gliroides as a seed disperser. We evaluated the diet of this marsupial in its natural habitat and in captivity. Dromiciops gliroides is omnivorous showing high consumption of a diversity of fruits. In captivity, D. gliroides consumed fruits from 80% of 22 native plant species we examined. Experiments conducted with fruits from two common understory shrubs show that seed passage through the digestive tract of D. gliroides enhances germination. Our results suggest that this species may have an important role as a seed disperser in the temperate forest of South America, which might offset a scarcity of frugivorous bird species.     

Identity of Pelodiscus sinensis revealed by DNA sequences of an approximately 180‐year‐old type specimen and a taxonomic reappraisal of Pelodiscus species (Testudines: Trionychidae)

Recent studies identified several distinct genetic lineages within the softshell turtle genus Pelodiscus that could represent valid species. Traditionally, Pelodiscus was regarded to comprise only a single species (P. sinensis). These softshell turtles are economically the most important chelonians of the world, with hundreds of millions of specimens traded as food every year. Moreover, Pelodiscus is used as a model organism for embryological and physiological studies, making correct species identification of paramount interest for disciplines beyond taxonomy. However, the understanding of the diversity of Pelodiscus was seriously hampered by the unclear taxonomic allocation of the oldest available species name, Trionyx (Aspidonectes) sinensis Wiegmann, 1834 . To clarify its identity, we reconstructed two mitochondrial DNA fragments of 1013 bp (cytb) and 468 bp (ND4) length of one of the two surviving syntypes and designate this specimen as lectotype (ZMB 38, Museum für Naturkunde Berlin). The sequences obtained from the lectotype represent a previously unknown lineage. Using the phylogenetic placement of all lineages and uncorrected p distances of the mitochondrial cytb gene as a yardstick, we suggest that the observed sequence variation is consistent with the existence of at least four distinct species within Pelodiscus. The name P. sinensis should be restricted to turtles harbouring the mitochondrial lineages B, C, D and the lineage of the lectotype. More divergent lineages are to be identified with P. axenaria, P. maackii and P. parviformis, which are recognized as valid species.     

Phylogeography and cryptic variation within the Lacerta viridis complex (Lacertidae, Reptilia)

It is well known that the current genetic pattern of many European species has been highly influenced by climatic changes during the Pleistocene. While there are many well known vertebrate examples, knowledge about squamate reptiles is sparse. To obtain more data, a range‐wide sampling of Lacerta viridis was conducted and phylogenetic relations within the L. viridis complex were analysed using an mtDNA fragment encompassing part of cytochrome b, the adjacent tRNA genes and the noncoding control region. Most genetic divergence was found in the south of the distribution range. The Carpathian Basin and the regions north of the Carpathians and Alps are inhabited by the same mitochondrial lineage, corresponding to Lacerta viridis viridis. Three distinct lineages occurred in the south‐eastern Balkans — corresponding to L. v. viridis, L. v. meridionalis, L. v. guentherpetersi— as well as a fourth lineage for which no subspecies name is available. This distribution pattern suggests a rapid range expansion of L. v. viridis after the Holocene warming, leading to a colonization of the northern part of the species range. An unexpected finding was that a highly distinct genetic lineage occurs along the western Balkan coast. Phylogenetic analyses (Bayesian, maximum likelihood, maximum parsimony) suggested that this west Balkan lineage could represent the sister taxon of Lacerta bilineata. Due to the morphological similarity of taxa within the L. viridis complex this cryptic taxon was previously assigned to L. v. viridis. The distribution pattern of several parapatric, in part highly, distinct genetic lineages suggested the existence of several refuges in close proximity on the southern Balkans. Within L. bilineata sensu stricto a generally similar pattern emerged, with a high genetic diversity on the Apennine peninsula, arguing for two distinct refuges there, and a low genetic diversity in the northern part of the range. Close to the south‐eastern Alps, three distinct lineages (L. b. bilineata, L. v. viridis, west Balkan taxon) occurred within close proximity. We suggest that the west Balkan lineage represents an early offshoot of L. bilineata that was isolated during a previous Pleistocene glacial from the more western L. bilineata populations, which survived in refuges on the Apennine peninsula.     

Phylogeographic structure of the dwarf snakehead (Channa gachua) around Gulf of Tonkin: Historical biogeography and pronounced effects of sea‐level changes

Geological events, landscape features, and climate fluctuations have shaped the distribution of genetic diversity and evolutionary history in freshwater fish, but little attention has been paid to that around the Gulf of Tonkin; therefore, we investigated the phylogeographic structure of the dwarf snakehead (Channa gachua) on Hainan Island and mainland China, as well as two populations in Vietnam. We attempted to elucidate the origins of freshwater fish in South Hainan by incorporating genetic data from DNA markers on both the mitochondrial cytochrome b gene (cyt b) and the nuclear recombination‐activating gene 1 (RAG‐1). Mitochondrial phylogenetic analysis identified two major lineages (lineages A and B), which may represent separate species. Divergence data suggested that C. gachua populations diverged between 0.516 and 2.376 myr. The divergence of the two cryptic species is congruent with sea‐level rise, which subsequently isolated Hainan from the mainland. During the Pleistocene glaciations, the entire region of the Gulf of Tonkin and the Qiongzhou Strait became part of the coastal plain of the Asian continent, which might have resulted in the current distribution patterns and dispersal routes of C. gachua populations. The formation of three sublineages in lineage A indicated that the Gulf of Tonkin was a geographical barrier between Hainan Island and mainland China but not between Vietnam and Hainan Island. The results of this study may help to elucidate the origins of freshwater fish in South Hainan and the phylogeographic structure of C. gachua.     

Evolutionary history of the Dactylorhiza maculata polyploid complex (Orchidaceae)

Taxonomic complexity may be associated with migration history and polyploidy. We used plastid and nuclear DNA markers to investigate the evolutionary history of the systematically challenging Dactylorhiza maculata polyploid complex. A total of 1833 individuals from 298 populations from throughout Europe were analysed. We found that gene flow was limited between the two major taxa, diploid ssp. fuchsii (including ssp. saccifera) and tetraploid ssp. maculata. A minimum of three autotetraploid lineages were discerned: (1) southern/western ssp. maculata; (2) northern/eastern ssp. maculata; and (3) Central European ssp. fuchsii. The two ssp. maculata lineages, which probably pre‐date the last glaciation, form a contact zone with high genetic diversity in central Scandinavia. Intermediate plastid haplotypes in the contact zone hint at recombination. Central Europe may have been a source area for the postglacial migration for the southern/western lineage of ssp. maculata, as well as for ssp. fuchsii. The northern/eastern lineage of ssp. maculata may have survived the LGM in central Russia west of the Urals. The tetraploid lineage of ssp. fuchsii is indistinguishable from diploid ssp. fuchsii, and is probably of postglacial origin. The Mediterranean region and the Caucasus have not contributed to the northward migration of either ssp. fuchsii or ssp. maculata. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 503–525.