Phylogeography of a Sahelian rodent species Mastomys huberti: a Plio-Pleistocene story of emergence and colonization of humid habitats

The multimammate rat Mastomys huberti is a Sahelian species restricted to West Africa. Throughout its distribution area, the species is associated with humid habitats, flood plains and ponds, which make its current distribution highly fragmented. Knowing that humid and dry climatic phases regularly alternated along the Quaternary in West Africa, it can be postulated that the evolutionary history of the species and its genetic variation largely reflect these climatic oscillations. We used mitochondrial cytochrome b sequences to investigate the phylogenetic relationships of M. huberti populations across the totality of the species' geographical range (Mali, Senegal, Guinea and Mauritania). We found that cytochrome b sequence variation is partitioned into four divergent clades (mean Kimura 2-parameter genetic distances varying from 0.57 to 3.08%) corresponding to distinct geographical regions. We dated the separation events of these clades between 0.93 and 0.17 million years ago, suggesting that M. huberti history was strongly influenced by the Quaternary climatic variations and related hydrographic network changes. Relationships between lineages and the partitioning of genetic diversity suggest the occurrence of two refuges along the Atlantic coast during arid periods. Moreover, the species' current range results from a stepwise colonization from west to east. M. huberti colonized recently the Inner Delta of Niger River in Mali, probably during a humid episode some 0.6 million years ago. Demographically stable and highly diversified populations were found in South Senegal and Guinea while populations in North Senegal and in Mali experienced low numbers followed by a demographic expansion during the African Humid Period (c. 14 800-5500 bp). During the last arid period (c. 23 000-18 000 years ago), Malian populations found refuge in the northern parts of the Inner Delta of the Niger River, then expended to the southern parts of the delta and along the course of the Niger River downstream Tombouctou. More recently, M. huberti would have rapidly expanded into irrigated areas along the Senegal River and along the Canal du Sahel, Mali, reflecting the invasive and the pest character of this species.     

Phylogeography of two cryptic species of African desert jerboas (Dipodidae: Jaculus)

The lesser Egyptian jerboa Jaculus jaculus is a desert dwelling rodent that inhabits a broad Arabian–Saharan arid zone. Recently, two distant sympatric lineages were described in North‐West Africa, based on morphometric and molecular data, which may correspond to two cryptic species. In the current study, phylogenetic relationships and phylogeographical structure among those lineages and geographical populations from North Africa and the Middle East were investigated. The phylogeographical patterns and genetic diversity of the cytochrome b gene (1110 bp) were addressed on 111 jerboas from 41 localities. We found that the variation in Africa is partitioned into two divergent mitochondrial clades (10.5% divergence relating to 1.65–4.92 Mya) that corresponds to the two cryptic species: J. jaculus and J. deserti. Diversifications within those cryptic species/clades were dated to 0.23–1.13 Mya, suggesting that the Middle Pleistocene climatic change and its environmental consequences affected the evolutionary history of African jerboas. The third distant clade detected, found in the Middle East region, most likely represents a distinct evolutionary unit, independent of the two African lineages. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Population genetic structure of two ecologically distinct multimammate rats: the commensal Mastomys natalensis and the wild Mastomys erythroleucus in southeastern Senegal

Using the same set of microsatellite markers, we compared the population genetic structure of two Mastomys species, one being exclusively commensal in southeastern Senegal, and the other being continuously distributed outside villages in this region. Both species were sampled in the same landscape context and at the same spatial scale. According to the expectations based on the degree of habitat patchiness (which is higher for commensal populations in this rural area), genetic diversity was lower and genetic differentiation was higher in commensal populations of Mastomys natalensis than in wild populations of Mastomys erythroleucus. Contrasting estimates of effective dispersal and current migration rates corroborates previous data on differences in social structure between the two species. Isolation-by-distance analyses showed that human-mediated dispersal is not a major factor explaining the pattern of genetic differentiation for M. natalensis, and that gene flow is high and random between M. erythroleucus populations at the spatial scale considered.     

The influence of life history and climate driven diversification on the mtDNA phylogeographic structures of two southern African Mastomys species (Rodentia: Muridae: Murinae)

The phylogeographic patterns of small mammals in southern Africa are frequently disjunct. This pattern is predominately attributed to vicariant geographical barriers coupled to climate driven diversification. To gain further insights into this hypothesis, we embarked on a comparative mtDNA phylogeographic study of two common rodent species in southern Africa, Mastomys natalensis and Mastomys coucha. Parsimony haplotype networks and SplitsTrees of mtDNA cytochrome oxidase I data showed a large degree of haplotype sharing throughout the sampling range. Within southern Africa, we found no conclusive evidence to support geographic vicariance as a contributing factor towards Mastomys speciation. We proposed that the regional phylogeographic structures detected for M. natalensis and M. coucha are the result of weak isolation by distance coupled to repeated expansions and contractions of suitable habitat. Both species probably survived in multiple refugia during unfavourable periods and mismatch distributions show signs of population expansion. Mitochondrial DNA nucleotide diversity values (π) show marked differences between the two species (M. natalensis: 0.003 and M. coucha: 0.468), and M. coucha also shows a higher level of population differentiation in AMOVA analyses. These differences are most likely due to life history discrepancies between the two species. Mastomys coucha is regarded to be more of a habitat specialist when compared to M. natalensis, and this probably places a higher constraint on M. coucha dispersal abilities. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 58–68.     

Phylogeographical footprints of the Strait of Gibraltar and Quaternary climatic fluctuations in the western Mediterranean: a case study with the greater white-toothed shrew, Crocidura russula (Mammalia: Soricidae)

We used mitochondrial cyt b sequences to investigate the phylogenetic relationships of Crocidura russula (sensu lato) populations across the Strait of Gibraltar, western Europe, Maghreb, and the Mediterranean and Atlantic islands. This revealed very low genetic divergence between European and Moroccan populations. The application of a molecular clock previously calibrated for shrews suggested that the separation of European from Moroccan lineages occurred less than 60 000 bp, which is at least 5 million years (Myr) after the reopening of the Strait of Gibraltar. This means that an overwater dispersal event was responsible for the observed phylogeographical structure. In contrast, genetic analyses revealed that Moroccan populations were highly distinct from Tunisian ones. According to the molecular clock, these populations separated about 2.2 million years ago (Ma), a time marked by sharp alternations of dry and humid climates in the Maghreb. The populations of the Mediterranean islands Ibiza, Pantelleria, and Sardinia were founded from Tunisian populations by overwater dispersal. In conclusion, overwater dispersal across the Strait of Gibraltar, probably assisted by humans, is possible for small terrestrial vertebrates. Moreover, as in Europe, Quaternary climatic fluctuations had a major effect on the phylogeographical structure of the Maghreb biota.     

Comparative phylogeography of two sibling species of forest‐dwelling rodent (Praomys rostratus and P. tullbergi) in West Africa: different reactions to past forest fragmentation

Two sibling species of the rodent genus Praomys occur in West African forests: P. tullbergi and P. rostratus. By sampling across their geographical ranges (459 individuals from 77 localities), we test the hypothesis that climatic oscillations during the Quaternary made an impact on the observed pattern of cytochrome b sequence variation. We show that, although these two species have parapatric geographical distributions, their phylogeographical histories are dissimilar, which could be related to their distinct ecological requirements. Since the arid phases of the Pleistocene were characterized by isolated forest patches, and intervening wetter periods by forest expansion, these changes in forest cover may be the common mechanism responsible for the observed phylogeographical patterns in both of these species. For example, in both species, most clades had either allopatric or parapatric geographical distributions; however, genetic diversity was much lower in P. tullbergi than in P. rostratus. The genetic pattern of P. tullbergi fits the refuge hypothesis, indicating that a very small number of populations survived in distinct forest blocks during the arid phases, then expanded again with forest recovery. In contrast, a number of populations of P. rostratus appear to have survived during the dry periods in more fragmented forest habitats, with varying levels of gene flow between these patches depending on climatic conditions and forest extent. In addition, historical variations of the West African hydrographic network could also have contributed to the pattern of genetic differentiation observed in both species.     

mtDNA variability in two Bantu‐speaking populations (Shona and Hutu) from Eastern Africa: Implications for peopling and migration patterns in sub‐Saharan Africa

In this study, we report novel data on mitochondrial DNA in two of the largest eastern Bantu‐speaking populations, the Shona from Zimbabwe and the Hutu from Rwanda. The goal is to evaluate the genetic relationships of these two ethnic groups with other Bantu‐speaking populations. Moreover, by comparing our data with those from other Niger‐Congo speaking populations, we aim to clarify some aspects of evolutionary and demographic processes accompanying the spread of Bantu languages in sub‐Saharan Africa and to test if patterns of genetic variation fit with models of population expansion based on linguistic and archeological data. The results indicate that the Shona and Hutu are closely related to the other Bantu‐speaking populations. However, there are some differences in haplogroup composition between the two populations, mainly due to different genetic contributions from neighboring populations. This result is confirmed by estimates of migration rates which show high levels of gene flow not only between pairs of Bantu‐speaking populations, but also between Bantu and non‐Bantu speakers. The observed pattern of genetic variability (high genetic homogeneity and high levels of gene flow) supports a linguistic model suggesting a gradual spread of Bantu‐speakers, with strong interactions between the different lines of Bantu‐speaker descent, and is also in agreement with recent archeological findings. In conclusion, our data emphasize the role that population admixture has played at different times and to varying degrees in the dispersal of Bantu languages. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Phylogeography of the fiscal shrike (Lanius collaris): a novel pattern of genetic structure across the arid zones and savannas of Africa

Aim Savanna occupies a substantial part of Africa, being distributed around the two major tropical rain forest blocks in what is referred to as the Savanna Belt. Our current understanding of the genetic structure within species distributed across the Savanna Belt is primarily derived from mammalian taxa, studies of which have revealed a suture zone or transition between northern and east/southern Africa clades in south‐western Kenya and north‐western Tanzania. We conduct a phylogeographic study of the fiscal shrike (Lanius collaris), a polytypic species distributed across the Savanna Belt of Africa and for which morphological and vocal data are in agreement with the suture zone recovered for mammalian taxa, to test the hypothesis of a spatially congruent genetic break across several taxa, including birds. Location Africa, south of the Sahara. Methods We analysed DNA sequences recovered from four loci (one mitochondrial, two autosomal and one Z‐linked) in 66 individuals, representing all recognized subspecies, as well as putatively closely related species. We make use of a combination of tree‐building and population genetic methods to investigate the phylogeographic structure of the fiscal shrike across Africa. Results The fiscal shrike consists of two primary lineages with a strong geographic component: a northern group distributed from southern Tanzania to Senegal, and a southern group distributed from Botswana/Zambia to South Africa with isolated populations in Tanzania and northern Malawi. Unexpectedly, Souza’s shrike (L. souzae) was nested within L. collaris, as the sister group of the southern group. The positions of Mackinnon’s shrike (L. mackinnoni) and that of the São Tomé shrike (L. newtoni) were variable, being either nested within the fiscal shrike or sister to the L. collaris–L. souzae clade. Our divergence time analyses suggest that the Lanius collaris species complex started to diversify around 2.2 Ma. Main conclusions Our study reveals a distinct biogeographic pattern for a savanna distributed species in Africa, with the transition between the two primary genetic lineages occurring at a latitude of c. 15–16° S, 10° S further south than shown elsewhere for several mammalian species.     

A long‐standing Pleistocene refugium in southern Africa and a mosaic of refugia in East Africa: insights from mtDNA and the common eland antelope

Aim Previous genetic studies of African savanna ungulates have indicated Pleistocene refugial areas in East and southern Africa, and recent palynological, palaeovegetation and fossil studies have suggested the presence of a long‐standing refugium in the south and a mosaic of refugia in the east. Phylogeographic analysis of the common eland antelope, Taurotragus oryx (Bovidae), was used to assess these hypotheses and the existence of genetic signatures of Pleistocene climate change. Location The sub‐Saharan savanna biome of East and southern Africa. Methods Mitochondrial DNA control‐region fragments (414 bp) from 122 individuals of common eland were analysed to elucidate the phylogeography, genetic diversity, spatial population structuring, historical migration and demographic history of the species. The phylogeographic split among major genetic lineages was dated using Bayesian coalescent‐based methods and a calibrated fossil root of 1.6 Ma for the split between the common eland and the giant eland, Taurotragus derbianus. Results Two major phylogeographic lineages comprising East and southern African localities, respectively, were separated by a net nucleotide distance of 4.7%. A third intermediate lineage comprised only three haplotypes, from Zimbabwe in southern Africa. The estimated mutation rate of 0.097 Myr^?1 revealed a more recent common ancestor for the eastern lineage (0.21 Ma; 0.07–0.37) than for the southern lineage (0.35 Ma; 0.10–0.62). Compared with the latter, the eastern lineage showed pronounced geographic structuring, lower overall nucleotide diversity, higher population differentiation, and isolation‐by‐distance among populations. Main conclusions The data support the hypothesis of Pleistocene refugia occurring in East and southern Africa. In agreement with palynological, palaeovegetation and fossil studies, our data strongly support the presence of a longer‐standing population in the south and a mosaic of Pleistocene refugia in the east, verifying the efficacy of genetic tools in addressing such questions. The more recent origin of the common eland inhabiting East Africa could result from colonization following extinction from the region. Only two other dated African ungulate phylogenies have been published, applying different methods, and the similarity of dates obtained from the three distinct approaches indicates a significant event c. 200 ka, which left a strong genetic signature across a range of ungulate taxa.     

Genetic diversity,evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic makeup. Location Sub‐Saharan Africa and India, with special focus on West and Central Africa. Method In this study 126 samples, throughout the lion’s complete geographic range, were subjected to phylogenetic analyses. DNA sequences of a mitochondrial region, containing cytochrome b, tRNAPro, tRNAThr and the left part of the control region, were analysed. Results Bayesian, maximum likelihood and maximum parsimony analyses consistently showed a distinction between lions from West and Central Africa and lions from southern and East Africa. West and Central African lions are more closely related to Asiatic lions than to the southern and East African lions. This can be explained by a Pleistocene extinction and subsequent recolonization of West Africa from refugia in the Middle East. This is further supported by the fact that the West and Central African clade shows relatively little genetic diversity and is therefore thought to be an evolutionarily young clade. Main conclusions The taxonomic division between an African and an Asian subspecies does not fully reflect the overall genetic diversity within lions. In order to conserve genetic diversity within the species, genetically distinct lineages should be prioritized. Understanding the geographic pattern of genetic diversity is key to developing conservation strategies, both for in situ management and for breeding of captive stocks.     

Second Sahelian amphibian endemism suggested by phylogeography of Groove crowned Bullfrog (Hoplobatrachus occipitalis) in western Sahel and hints of polyploid species formation

Several studies have assessed the phylogeographic patterns of vertebrates in North Africa and Sahara–Sahel, but most of the phylogeographic knowledge on amphibians comes from the Mediterranean region while the southern Sahara and Sahel remain poorly studied. Here, we assess the phylogeography of the African Groove crowned frog Hoplobatrachus occipitalis, with a focus on western Sahel in order to better understand the biogeographic patterns of semi-aquatic species in this arid region. Using mitochondrial and nuclear markers, we have assessed the species’ genetic structure, distribution of genetic diversity, and the presence of cryptic diversity. We found evidence of a recent (re-)colonization of the mountains in its northernmost distribution, but also for the role of southern Mauritanian mountains and large rivers as refugia. Two major lineages were detected, one perhaps endemic to Mauritania and the other widespread in Africa. The first lineage possibly constitutes the second Sahelian amphibian endemic; the latter may have originated through an allopolyploidy event, with the Mauritanian lineage being one of the parental ones.     

Phylogeography of the widespread African puff adder (Bitis arietans) reveals multiple Pleistocene refugia in southern Africa

Evidence from numerous Pan‐African savannah mammals indicates that open‐habitat refugia existed in Africa during the Pleistocene, isolated by expanding tropical forests during warm and humid interglacial periods. However, comparative data from other taxonomic groups are currently lacking. We present a phylogeographic investigation of the African puff adder (Bitis arietans), a snake that occurs in open‐habitat formations throughout sub‐Saharan Africa. Multiple parapatric mitochondrial clades occur across the current distribution of B. arietans, including a widespread southern African clade that is subdivided into four separate clades. We investigated the historical processes responsible for generating these phylogeographic patterns in southern Africa using species distribution modelling and genetic approaches. Our results show that interior regions of South Africa became largely inhospitable for B. arietans during glacial maxima, whereas coastal and more northerly areas remained habitable. This corresponds well with the locations of refugia inferred from mitochondrial data using a continuous phylogeographic diffusion model. Analysis of data from five anonymous nuclear loci revealed broadly similar patterns to mtDNA. Secondary admixture was detected between previously isolated refugial populations. In some cases, this is limited to individuals occurring near mitochondrial clade contact zones, but in other cases, more extensive admixture is evident. Overall, our study reveals a complex history of refugial isolation and secondary expansion for puff adders and a mosaic of isolated refugia in southern Africa. We also identify key differences between the processes that drove isolation in B. arietans and those hypothesized for sympatric savannah mammals.     

New polymorphic microsatellite loci for rodents of the genus Mastomys using PCR multiplexing,and cross‐species amplification in Myomys and Praomys

The Praomys complex contains some of the most important agricultural pests of Africa, including Mastomys species. We describe the development of nine supplementary microsatellite markers isolated from Mastomys huberti. We show the potential utility of M. huberti microsatellites as population markers for two other species of Mastomys, and two other species of the Praomys complex, Myomys daltoni and Praomys cf. rostratus.     

Mitochondrial DNA Phylogeography and Comparative Cytogenetics of the Springhare, Pedetes capensis (Mammalia: Rodentia)

Variation in mitochondrial DNA (mtDNA) was used together with comparative cytogenetics to examine the evolutionary history and taxonomic status of an African hystricomorphous rodent, the springhare Pedetes capensis. The mtDNA phylogeographic structure showed that the majority of the southern African populations (P. c. capensis) are characterized by unique but closely related maternal lineages. Based on restriction endonuclease fragment analysis, the east African populations (P. c. surdaster) appear more structured and are distinguished from those in southern Africa by an average sequence divergence of 5.52% (±1.4%). This marked divergence is concordant with results of the cytogenetic study. Specimens from southern Africa have 2n = 38, and those from east Africa 2n = 40. The change in diploid number is due to a single centric fusion. It is suggested that the closure of the Brachystegia or miombo woodland (20,000–10,000 B.P.), which delimits contemporary springhare ranges, may have been too recent to account for the accumulated genetic differences that distinguish these taxa. While rifting and associated habitat changes in east Africa can be invoked to explain genetic structure in this region, the southern African springhare populations, which have a high incidence of locality-specific haplotypes, show a shallow phylogeographic structure, in keeping with a relatively recent range expansion from smaller source populations. Given the magnitude of genetic, morphological, and ethological differences between the two geographic isolates, we believe that there is strong support for the elevation of the east African and southern African springhare populations to full species status, thus supporting earlier taxonomic treatments of this rodent.     

Phylogeography of the Cape velvet worm (Onychophora: Peripatopsis capensis) reveals the impact of Pliocene/Pleistocene climatic oscillations on Afromontane forest in the Western Cape, South Africa

Habitat specialists such as soft-bodied invertebrates characterized by low dispersal capability and sensitivity to dehydration can be employed to examine biome histories. In this study, the Cape velvet worm (Peripatopsis capensis) was used to examine the impacts of climatic oscillations on historical Afromontane forest in the Western Cape, South Africa. Divergence time estimates suggest that the P. capensis species complex diverged during the Pliocene epoch. This period was characterized by dramatic climatic and topographical change. Subsequently, forest expansion and contraction cycles led to diversification within P. capensis. Increased levels of genetic differentiation were observed along a west-to-south-easterly trajectory because the south-eastern parts of the Cape Fold Mountain chain harbour larger, more stable fragments of forest patches, have more pronounced habitat heterogeneity and have historically received higher levels of rainfall. These results suggest the presence of three putative species within P. capensis, which are geographically discreet and genetically distinct.     

Renal physiology of two southern African Mastomys species (Rodentia: Muridae): a salt-loading experiment to assess concentrating ability

Aspects of renal physiology were examined to test the hypothesis that two cryptic species of the genus Mastomys (Mastomys natalensis and Mastomys coucha) are geographically separated by differences in aridity tolerance. Laboratory-bred females of each species were subjected to different levels of salinity in their water source (distilled water, 0.9% NaCl, and 1.5% NaCl; 10 conspecifics in each group) from weaning until sexual maturity. Individuals of the two species exhibited similar rates of water consumption and urine production. The salinity treatments caused sodium diuresis in both species, evident in increased urine volume, decreased osmolality and increased osmotic output. Urine concentration, kidney mass and kidney relative medullary area (RMA) did not differ between species. The results of our study do not support the hypothesis that differences in osmoregulatory ability separate these two cryptic species. Nor do they support the use of salt loading to elicit maximum urine concentrations in mammals.     

Phylogeography of the Alpine salamander, Salamandra atra (Salamandridae) and the influence of the Pleistocene climatic oscillations on population divergence

Fifty individuals of the endemic Alpine salamander, Salamandra atra, representing 13 populations throughout the range of the two currently recognized subspecies, atra and aurorae, were examined for sequence variation in a large portion (1050 bp) of the mitochondrial cytochrome b gene. We revealed a large number of mitochondrial DNA (mtDNA) haplotypes (10). Interpopulation sequence divergence was very low, ranging from 0 to 3.1%. The relationships among haplotypes were poorly resolved. The divergence time estimate between several mtDNA haplotypes suggested a pre-Pleistocene differentiation approximately 3 million years ago. Moreover, the impact of the Pleistocene glaciations on the phylogeographical patterns appears to have been secondary, although a somewhat reduced genetic variability was found in populations living in areas that were directly affected by the glaciation.     

Phylogeography and host association in a pollinating seed parasite Greya politella (Lepidoptera: Prodoxidae)

Inferring the historical context of ecological diversification is an important step in understanding the way that population-level processes result in a diversity of species and interactions in communities. We performed a phylogeographic analysis of mitochondrial DNA haplotypes from the pollinating seed parasite Greya politella (Lepidoptera: Prodoxidae) in order to determine the degree to which populations were structured according to geographical location and host-plant association. Ninety-eight individuals were sampled from 29 locations ranging from southern California to western Idaho. Restriction-site variation in 87 individuals (27 populations) was screened by digestion with 11 endonucleases, followed by Southern blotting; 38 restriction-site positions were mapped by double digests. Haplotypes were further defined by generating fragments 251 bases in length via PCR, screening them for sequence variation using denaturing gel gradient electrophoresis (DGGE), and sequencing the resulting variants. Parsimony analysis of the resulting 12 restriction-site and 15 sequence haplotypes indicated strong geographical structuring of populations: (i) most populations were monomorphic for haplotype; (ii) haplotypes from California and the Pacific Northwest (Oregon, Washington and Idaho) formed robust monophyletic groups. Population structure was significant both within and between the two regions, as reflected by N _ST. Patterns of host-plant association and haplotype phylogeny suggest that populations have recently undergone host-plant shifts in many different parts of the species range, although the direction and number of host shifts cannot be determined at the present level of sampling resolution.     

Phylogeography of Pinus tabulaeformis Carr. (Pinaceae), a dominant species of coniferous forest in northern China

How coniferous trees in northern China changed their distribution ranges in response to Quaternary climatic oscillations remains largely unknown. Here we report a study of the phylogeography of Pinus tabulaeformis, an endemic and dominant species of coniferous forest in northern China. We examined sequence variation of maternally inherited, seed‐dispersed mitochondrial DNA (mtDNA) (nad5 intron 1 and nad4/3–4) and paternally inherited, pollen‐ and seed‐dispersed chloroplast DNA (cpDNA) (rpl16 and trnS‐trnG) within and among 30 natural populations across the entire range of the species. Six mitotypes and five chlorotypes were recovered among 291 trees surveyed. Population divergence was high for mtDNA variation (G_ST = 0.738, N_ST = 0.771) indicating low levels of seed‐based gene flow and significant phylogeographical structure (N_ST > G_ST, P < 0.05). The spatial distribution of mitotypes suggests that five distinct population groups exist in the species: one in the west comprising seven populations, a second with a north–central distribution comprising 15 populations, a third with a southern and easterly distribution comprising five populations, a fourth comprising one central and one western population, and a fifth comprising a single population located in the north‐central part of the species’ range. Each group apart from the fourth group is characterized by a distinct mitotype, with other mitotypes, if present, occurring at low frequency. It is suggested, therefore, that most members of each group apart from Group 4 are derived from ancestors that occupied different isolated refugia in a previous period of range fragmentation of the species, possibly at the time of the Last Glacial Maximum. Possible locations for these refugia are suggested. A comparison of mitotype diversity between northern and southern subgroups within the north‐central group of populations (Group 2) showed much greater uniformity in the northern part of the range both within and between populations. This could indicate a northward migration of the species from a southern refugium in this region during the postglacial period, although alternative explanations cannot be ruled out. Two chlorotypes were distributed across the geographical range of the species, resulting in lower levels of among‐population chlorotype variation. The geographical pattern of variation for all five chlorotypes provided some indication of the species surviving past glaciations in more than one refugium, although differentiation was much less marked, presumably due to the greater dispersal of cpDNA via pollen.