Population size and time since island isolation determine genetic diversity loss in insular frog populations

Understanding the factors that contribute to loss of genetic diversity in fragmented populations is crucial for conservation measurements. Land‐bridge archipelagoes offer ideal model systems for identifying the long‐term effects of these factors on genetic variations in wild populations. In this study, we used nine microsatellite markers to quantify genetic diversity and differentiation of 810 pond frogs (Pelophylax nigromaculatus) from 24 islands of the Zhoushan Archipelago and three sites on nearby mainland China and estimated the effects of the island area, population size, time since island isolation, distance to the mainland and distance to the nearest larger island on reduced genetic diversity of insular populations. The mainland populations displayed higher genetic diversity than insular populations. Genetic differentiations and no obvious gene flow were detected among the frog populations on the islands. Hierarchical partitioning analysis showed that only time since island isolation (square‐root‐transformed) and population size (log‐transformed) significantly contributed to insular genetic diversity. These results suggest that decreased genetic diversity and genetic differentiations among insular populations may have been caused by random genetic drift following isolation by rising sea levels during the Holocene. The results provide strong evidence for a relationship between retained genetic diversity and population size and time since island isolation for pond frogs on the islands, consistent with the prediction of the neutral theory for finite populations. Our study highlights the importance of the size and estimated isolation time of populations in understanding the mechanisms of genetic diversity loss and differentiation in fragmented wild populations.     

Phylogeography of endemic ermine (Mustela erminea) in southeast Alaska

The North Pacific Coast (NPC) of North America is a region of high mammalian endemism, possibly due to its highly fragmented landscape and complex glacial history. For example, four island and one mainland subspecies of ermine, Mustela erminea, have been described as endemic to southeast Alaska alone. To better understand the role of past climatic change in generating diversity in the region, we examined DNA sequence variation in the mitochondrial cytochrome b gene of 210 ermine from across North America, with an emphasis on Alaska and British Columbia. We found three distinct (1.5-3.6% uncorrected 'p') lineages of ermine, all of which occur in southeast Alaska. One lineage includes a southeast Alaska endemic and specimens from Alaska (outside of southeast) and Eurasia. A second lineage includes two southeast Alaskan endemics and ermine from western Canada and the coterminous United States. The close relationships of these purported endemics to ermine outside of southeast Alaska suggest that they colonized the region from Beringian and southern glacial refugia, respectively, following deglaciation of the NPC. The third lineage appears restricted to the Prince of Wales Island complex in southeast Alaska (two subspecies) and Graham Island (Haida Gwaii), British Columbia. This restricted distribution suggests that these populations may be derived from relicts that persisted in a coastal refugium during the Wisconsin glaciation. Studies of nuclear genes and adaptive morphological evolution are necessary to further explore discrepancies between the geographical pattern of differentiation based on mtDNA and the existing subspecific taxonomy based on morphology.     

Large mainland populations of South Island robins retain greater genetic diversity than offshore island refuges

For conservation purposes islands are considered safe refuges for many species, particularly in regions where introduced predators form a major threat to the native fauna, but island populations are also known to possess low levels of genetic diversity. The New Zealand archipelago provides an ideal system to compare genetic diversity of large mainland populations where introduced predators are common, to that of smaller offshore islands, which serve as predator-free refuges. We assessed microsatellite variation in South Island robins (Petroica australis australis), and compared large mainland, small mainland, natural island and translocated island populations. Large mainland populations exhibited more polymorphic loci and higher number of alleles than small mainland and natural island populations. Genetic variation did not differ between natural and translocated island populations, even though one of the translocated populations was established with five individuals. Hatching failure was recorded in a subset of the populations and found to be significantly higher in translocated populations than in a large mainland population. Significant population differentiation was largely based on heterogeneity in allele frequencies (including fixation of alleles), as few unique alleles were observed. This study shows that large mainland populations retain higher levels of genetic diversity than natural and translocated island populations. It highlights the importance of protecting these mainland populations and using them as a source for new translocations. In the future, these populations may become extremely valuable for species conservation if existing island populations become adversely affected by low levels of genetic variation and do not persist.     

Competing hypotheses for the etiology of cryptorchidism in Sitka black-tailed deer: an evaluation of evolutionary alternatives

On the Aliulik Peninsula (AP) of Kodiak Island, Alaska, 70% of male Sitka black-tailed deer (SBTD; Odocoileus hemionus sitkensis ) are bilaterally cryptorchid (both testes fail to descend; male is sterile). Both genetic and environmental factors have been proposed as possible causes of this problem. We investigated the possibility that population genetic processes (isolation, inbreeding and genetic drift) have contributed to an increased frequency of cryptorchidism in this population. Overall, SBTD on major islands throughout Alaska have unusually low levels of genetic diversity, though we identified a likely glacial refugium on Prince of Wales Island in the Alexander Archipelago. Within the Kodiak Archipelago, deer on the AP did not exhibit the patterns of genetic isolation, inbreeding and drift that would be expected if cryptorchidism in this population was the result of a founder mutation(s). Instead, our data favor exposure to environmental contaminants as a likely alternative mechanism causing high prevalence of cryptorchidism on the AP.     

Genetic differentiation within and among island populations of the endangered plant Aster miyagii (Asteraceae), an endemic to the Ryukyu Islands

Genetic diversity was examined at 17 putative allozyme loci in 18 populations of the insular endemic plant Aster miyagii (Asteraceae). This species is geographically restricted to only three islands of the Ryukyu Islands and is on the federal list of threatened plants. Genetic differentiation within an island is small, suggesting that gene flow among populations on the same island is sufficiently large to prevent divergence. By contrast, genetic differentiation among islands is large, especially between Amamioshima Island and the other two islands, suggesting that gene flow between the islands is highly restricted. Two unique alleles are nearly fixed in populations on Amamioshima Island, which is the southernmost island of the three. Comparatively, genetic diversity is the smallest on Amamioshima Island. This genetic paucity on Amamioshima Island is probably a result of a population bottleneck at colonization or the small effective population size on this island. Genetic diversity at the species level of A. miyagii is larger than those of the species with a similar life history and of the congeneric widespread species, suggesting that the species has an old origin as an insular endemic species.     

Is it worth eradicating the invasive pest Rattus norvegicus from Molène archipelago? Genetic structure as a decision-making tool

Genetic variability and structure were estimated by microsatellite analysis at 7 loci in brown rat populations (Rattus norvegicus) from the Iroise insular complex and neighbouring mainland (Brittany, France). Island genetic diversity is lower than on the mainland and a highly significant positive correlation was found between mean heterozygosity and the logarithm of island area, which is consistent with theoretical expectations. Rattus norvegicus populations are substructured at a kilometric scale, both on the islands and on the mainland. Intra-island structuration is extremely high, suggesting that no effective migration occurs between islands or with the mainland. Historical and genetical evidence suggest that R. norvegicus was introduced independently on Ouessant and Molène archipelago, with a low and a high founder effect respectively. These results are discussed in terms of recolonization probability of islands that have been cleared of R. norvegicus, which illustrates the usefulness of genetic markers in determining parameters of interest to the conservation biologist.     

Microsatellite and mitochondrial DNA variation defines island genetic reservoirs for reintroductions of an endangered Australian marsupial, <Emphasis Type="Italic">Perameles bougainville</Emphasis>

Natural populations of the endangered western barred bandicoot (Perameles bougainville) now exist on only two islands in Shark Bay, Western Australia. Our aim was to investigate genetic diversity in natural, reintroduced, and captive populations of the bandicoots and to assess the extent of divergence between the populations. The contemporary isolation of the natural populations has resulted in heterogeneity of allele frequency between the islands, which has acted to maintain a higher combined diversity than would be expected from either population on its own. These findings highlight how remnant island populations can act as genetic reservoirs to maximize diversity for reintroductions into a species former range. Although diversity is high between island populations, diversity within populations, based on six microsatellite loci, are amongst the lowest ever recorded for populations of marsupials. The mtDNA sequence data indicate that the two remaining natural populations show only minor divergence from each other, with the five haplotypes separated by just single base pairs. The reintroduced population and captive colonies show evidence for the loss of diversity related to genetic drift operating on small isolated populations.     

ALLOZYME DIFFERENTIATION AND GENETIC STRUCTURE IN ISLAND AND MAINLAND JAPANESE POPULATIONS OF CAMPANULA PUNCTATA (CAMPANULACEAE)

Allozyme variation was investigated in 17 Japanese populations of Campanula punctata, ten from the Izu Islands and seven in the mainland (Honshu). The data indicate that there are two groups, a mainland group and an island one, and that the systematically problematic Oshima Island (northernmost Izu island) populations are closely related to those of the other islands. Nei's genetic identity values among islands and among mainland populations were 0.95 and 0.97, respectively, while the value between island and mainland populations was 0.84, suggesting that the island populations are an independent species. Total genetic variation was nearly the same among island and mainland populations. However, the apportionment of variation within and among populations was considerably different; 14% of gene diversity exists among mainland populations, while 31% of the diversity exists among island populations. Mean outcrossing rates of self-incompatible mainland and Oshima populations are 0.62–0.79, supporting xenogamy; those in self-compatible island populations are 0.37–0.57 in the northern islands, indicating a mixed mating system, and 0.16–0.25 in southern ones, indicating dominant inbreeding. Total genetic diversity in each island population decreased with distance from the mainland. Genetic and geological data suggest that the ancestors of insular populations were founded on northern islands in a relatively ancient period and that they dispersed progressively to the southern ones. Chromosome number (2n = 34) and isozyme numbers indicate gene duplications in this species, which suggests it is an ancient polyploid.     

Genetic variation and island biogreography: Microsatellite and mitochondrial DNA variation in island populations of the Australian bush rat, Rattus fuscipes greyii

To understand the impact of various factors on the maintenance of genetic variation in natural populations, we need to focus on situations where at least some of these factors are removed or controlled. In this study, we used highly variable, presumably neutral, microsatellite and mtDNA markers to assess the nature of genetic variation in 14 island and two mainland populations of the Australian bush rat, where there is no migration between islands. Thus we are controlling for selection and gene flow. Both marker sets revealed low levels of diversity within the small island populations and extreme differentiation between populations. For six microsatellite loci, all of the small island populations had less genetic variation than the mainland populations; reduction in allelic diversity was more pronounced than loss of heterozygosity. Kangaroo Island, the large island population, had similar levels of diversity to the mainland populations. A 442 base pair (bp) section of the mtDNA control region was screened for variation by outgroup heteroduplex analysis/temperature gradient gel electrophoresis (OHA/TGGE). Only three of the 13 small island populations showed haplotypic diversity: Gambier (2), Waldegrave (2), and Eyere (3). The level of haplotypic diversity in the small island populations was similar to that on the mainland, most likely reflecting a recent population bottleneck on the mainland. In contrast, Kangaroo Island had 9 mtDNA haplotypes. The dominant factor influencing genetic diversity on the islands was island size. No correlation was detected between genetic diversity and the time since isolation or distance form the mainland. The combination of genetic drift within and complete isolation among the small island populations has resulted in rapid and extreme population divergence. Population pair-wise comparisons of allele frequency distributions showed significant differences for all populations for all loci (F _st = 0.11–0.84, R _st = 0.07–0.99). For the mtDNA control region, 92.6% of variation was apportioned between populations; only the Pearson islands shared a haplotype. Mantel tests of pair-wise genetic distance with pair-wise geographic distance showed no significant geographical clustering of haplotypes. However, population substructuring was detected within populations where sampling was conducted over a broader geographical range, as indicated by departures from Hardy-Weinberg equilibrium. Thus substructuring in the ancestral population cannot be ruled out. The dominant evolutionary forces on the islands, after the initial founder event, are stochastic population processes such as genetic drift and mutation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of ecogeographic variables on genetic variation in montane mammals: implications for conservation in a global warming scenario

Aim Evolutionary theory predicts that levels of genetic variation in island populations will be positively correlated with island area and negatively correlated with island isolation. These patterns have been empirically established for oceanic islands, but little is known about the determinants of variation on habitat islands. The goals of this study were twofold. Our first aim was to test whether published patterns of genetic variation in mammals occurring on montane habitat islands in the American Southwest conformed to expectations based on evolutionary theory. The second aim of this research was to develop simple heuristic models to predict changes in genetic variation that may occur in these populations as a result of reductions in available mountaintop habitat in response to global warming. Location Habitat islands of conifer forest on mountaintops in the American Southwest. Methods Relationships between island area and isolation with measures of allozyme variation in four species of small mammal, namely the least chipmunk (Tamias minimus), Colorado chipmunk (Tamias quadrivittatus), red squirrel (Tamiasciurus hudsonicus), and Mexican woodrat (Neotoma mexicana), were determined using correlation and regression techniques. Significant relationships between island area and genetic variation were used to develop three distinct statistical models with which to predict changes in genetic variation following reduction in insular habitat area arising from global warming. Results Patterns of genetic variation in each species conformed to evolutionary predictions. In general, island area was the most important determinant of heterozygosity, while island isolation was the most important determinant of polymorphism and allelic diversity. The heuristic models predicted widespread reductions in genetic variation, the extent of which depended on the population and model considered. Main conclusions The results support a generalized pattern of genetic variation for any species with an insular distribution, with reduced variation in smaller, more isolated populations. We predict widespread reductions in genetic variation in isolated populations of montane small mammals in the American Southwest as a result of global warming. We conclude that climate‐induced reductions in the various dimensions of genetic variation may increase the probability of population extinction in both the short and long term.     

Integrated species delimitation and conservation implications of an endangered weevil Pachyrhynchus sonani (Coleoptera: Curculionidae) in Green and Orchid Islands of Taiwan

Oceanic islands are productive habitats for generating new species and high endemism, which is primarily due to their geographical isolation, smaller population sizes and local adaptation. However, the short divergence times and subtle morphological or ecological divergence of insular organisms may obscure species identity, so the cryptic endemism on islands may be underestimated. The endangered weevil Pachyrhynchus sonani Kôno (Coleoptera: Curculionidae: Entiminae: Pachyrhynchini) is endemic to Green Island and Orchid Island of the Taiwan‐Luzon Archipelago and displays widespread variation in coloration and host range, thus raising questions regarding its species boundaries and degree of cryptic diversity. We tested the species boundaries of P. sonani using an integrated approach that combined morphological (body size and shape, genital shape, coloration and cuticular scale), genetic (four genes and restriction site‐associated DNA sequencing, RAD‐seq) and ecological (host range and distribution) diversity. The results indicated that all the morphological datasets for male P. sonani, except for the colour spectrum, reveal overlapping but statistically significant differences between islands. In contrast, the morphology of the female P. sonani showed minimum divergence between island populations. The populations of P. sonani on the two islands were significantly different in their host ranges, and the genetic clustering and phylogenies of P. sonani established two valid evolutionary species. Integrated species delimitation combining morphological, molecular and ecological characters supported two distinct species of P. sonani from Green Island and Orchid Island. The Green Island population was described as P. jitanasaius sp.n. Chen & Lin, and it is recommended that its threatened conservation status be recognized. Our findings suggest that the inter‐island speciation of endemic organisms inhabiting both islands may be more common than previously thought, and they highlight the possibility that the cryptic diversity of small oceanic islands may still be largely underestimated.     

Demographic history of a recent invasion of house mice on the isolated Island of Gough

Island populations provide natural laboratories for studying key contributors to evolutionary change, including natural selection, population size and the colonization of new environments. The demographic histories of island populations can be reconstructed from patterns of genetic diversity. House mice (Mus musculus) inhabit islands throughout the globe, making them an attractive system for studying island colonization from a genetic perspective. Gough Island, in the central South Atlantic Ocean, is one of the remotest islands in the world. House mice were introduced to Gough Island by sealers during the 19th century and display unusual phenotypes, including exceptionally large body size and carnivorous feeding behaviour. We describe genetic variation in Gough Island mice using mitochondrial sequences, nuclear sequences and microsatellites. Phylogenetic analysis of mitochondrial sequences suggested that Gough Island mice belong to Mus musculus domesticus, with the maternal lineage possibly originating in England or France. Cluster analyses of microsatellites revealed genetic membership for Gough Island mice in multiple coastal populations in Western Europe, suggesting admixed ancestry. Gough Island mice showed substantial reductions in mitochondrial and nuclear sequence variation and weak reductions in microsatellite diversity compared with Western European populations, consistent with a population bottleneck. Approximate Bayesian computation (ABC) estimated that mice recently colonized Gough Island (~100 years ago) and experienced a 98% reduction in population size followed by a rapid expansion. Our results indicate that the unusual phenotypes of Gough Island mice evolved rapidly, positioning these mice as useful models for understanding rapid phenotypic evolution.     

Narrow water barriers prevent multiple colonizations and limit gene flow among California Channel Island wild buckwheats (Eriogonum: Polygonaceae)

The relative roles of chance colonization and subsequent gene flow in the development of insular endemic biotas have been extensively studied in remote oceanic archipelagos, but are less well characterized on nearshore island systems. The current study investigated patterns of colonization and divergence between and within two wild buckwheat species (Polygonaceae), Eriogonum arborescens and E. giganteum, endemic to the California Channel Islands to determine whether geographical isolation is driving diversification. Using plastid and nuclear sequence data and microsatellite allele frequencies, we determined that gene flow in these Eriogonum spp. is restricted by isolation. The data suggest that successful colonization of and gene flow among the islands are infrequent. Colonization appears to have followed a stepping‐stone model that is consistent with a north‐to‐south pattern across the islands. This colonization pattern coupled with relatively little post‐colonization inter‐island gene flow, particularly among southern islands, has generated a pattern of more divergent lineages on the isolated southern islands. These results run counter to the general expectation that all islands close to a continental source should receive a high level of gene flow. Finally, management recommendations focused on protecting the lineages from loss of private alleles and the erosion of the remaining genetic diversity are offered.     

Pathogen richness and abundance predict patterns of adaptive major histocompatibility complex variation in insular amphibians

The identification of the factors responsible for genetic variation and differentiation at adaptive loci can provide important insights into the evolutionary process and is crucial for the effective management of threatened species. We studied the impact of environmental viral richness and abundance on functional diversity and differentiation of the MHC class Ia locus in populations of the black‐spotted pond frog (Pelophylax nigromaculatus), an IUCN‐listed species, on 24 land‐bridge islands of the Zhoushan Archipelago and three nearby mainland sites. We found a high proportion of private MHC alleles in mainland and insular populations, corresponding to 32 distinct functional supertypes, and strong positive selection on MHC antigen‐binding sites in all populations. Viral pathogen diversity and abundance were reduced at island sites relative to the mainland, and islands housed distinctive viral communities. Standardized MHC diversity at island sites exceeded that found at neutral microsatellites, and the representation of key functional supertypes was positively correlated with the abundance of specific viruses in the environment (Frog virus 3 and Ambystoma tigrinum virus). These results indicate that pathogen‐driven diversifying selection can play an important role in maintaining functionally important MHC variation following island isolation, highlighting the importance of considering functionally important genetic variation and host–pathogen associations in conservation planning and management.     

A 9.1-kb insertion/deletion polymorphism suggests a common pattern of genetic diversity in Island Melanesia

Earlier reports suggest a distinct pattern of genetic variation linked to both language and geographic distance in Island Melanesia. Inland Papuan-speaking populations from different islands tend to share one allelic profile, while shore-based or more cosmopolitan populations share another, related to Southeast Asian influence over the past 3000 years. In the present paper, we report the genotypes and allele frequencies of an informative 9.1-thousand-base-pairs (kb) insertion/deletion polymorphism in 19 populations living in Island Melanesia. The populations studied inhabit the islands of New Britain, New Ireland, New Hanover, and Mussau in the Bismarck Archipelago, and speak either Austronesian or Papuan languages. We also include for reference a collection from New Guinea and Bougainville. The data show a marked fluctuation in the allele frequency among the different isolates, with the 9.1-kb(-) allele frequency ranging from 0.67 to 0.98. The deletion allele reaches fixation in some Papuan-speaking interior populations of New Britain, as well as in the interiors of New Guinea and Bougainville. However, certain inland Austronesian-speaking populations also share a similar high frequency of the deletion. Our data suggest that language distinctions are generally, but not invariably, indicative of diverse allelic patterns in this complex region, where inland groups on large islands tend to be often distinctive from shore-based populations.     

Distribution of genetic diversity within and between Western Mediterranean island populations of the black rat Rattus rattus (L. 1758)

Genetic diversity was estimated by allozyme analysis at 26 loci in black rat populations (Rattus rattus) from 15 western Mediterranean islands (Hyéres, Corsica, Sardinia and related islets). Although overall variability levels were low (H = 0.025), the mean heterozygosity values for the islands were similar to those for three reference mainland populations. Within the islands, however, genetic diversity varied in relation to island size and geographic isolation. In particular, most small insular populations were significantly more variable than those on both large and isolated islands. The generic relationships between island populations were established by F_ST analyses indicating possible geographic origins and patterns of colonization. The maintenance of unexpectedly high levels of variability in the small island populations is discussed in relation to changes in the demographic and social structure observed in these populations. These island populations of black rat illustrate how genetic diversity may be efficiently maintained in a series of interconnected spatially fragmented populations.     

The contribution of island populations to in situ genetic conservation

Genetic variation is often lower within island populations, however islands may also harbor divergent genetic variation. The likelihood that insular populations are genetically diverse or divergent should be influenced by island size and isolation. We tested this assumption by comparing patterns of genetic variation across all major island song sparrow populations along the Pacific North American coast. Allelic richness was moderately lowered even on islands which are close to large, potential sources. The most significant differences in allelic richness occurred on very small or highly remote islands. Gene diversity was significantly lower only on remote or very small islands. We found that island populations contribute to regional genetic variation through both the amount of genetic variation and the uniqueness of that variation. The partitioning of this contribution was associated with the size and isolation of the island populations.     

Selection and drift influence genetic differentiation of insular Canada lynx (Lynx canadensis) on Newfoundland and Cape Breton Island

Island populations have long been important for understanding the dynamics and mechanisms of evolution in natural systems. While genetic drift is often strong on islands due to founder events and population bottlenecks, the strength of selection can also be strong enough to counteract the effects of drift. Here, we used several analyses to identify the roles of genetic drift and selection on genetic differentiation and diversity of Canada lynx (Lynx canadensis) across eastern Canada, including the islands of Cape Breton and Newfoundland. Specifically, we assessed whether we could identify a genetic component to the observed morphological differentiation that has been reported across insular and mainland lynx. We used a dinucleotide repeat within the promoter region of a functional gene that has been linked to mammalian body size, insulin‐like growth factor‐1 (IGF‐1). We found high genetic differentiation at neutral molecular markers but convergence of allele frequencies at the IGF‐1 locus. Thus, we showed that while genetic drift has influenced the observed genetic structure of lynx at neutral molecular markers, natural selection has also played a role in the observed patterns of genetic diversity at the IGF‐1 locus of insular lynx.     

The island syndrome and population dynamics of introduced rats

The island syndrome predicts directional changes in the morphology and demography of insular vertebrates, due to changes in trophic complexity and migration rates caused by island size and isolation. However, the high rate of human-mediated species introductions to some islands also increases trophic complexity, and this will reduce the perceived insularity on any such island. We test four hypotheses on the role of increased trophic complexity on the island syndrome, using introduced black rats (Rattus rattus) on two isolated coral atolls in the Mozambique Channel. Europa Island has remained relatively pristine and insular, with few species introductions, whereas Juan de Nova Island has had many species introductions, including predators and competitors of rats, anthropogenically increasing its trophic complexity. In the most insular environments, the island syndrome is expected to generate increases in body size and densities of rodents but decreases in the rates of reproduction and population cycling. Morphology and reproduction were compared using linear regression and canonical discriminant analysis, while density and population cycling were compared using spatially explicit capture–recapture analysis. Results were compared to other insular black rat populations in the Mozambique Channel and were consistent with predictions from the island syndrome. The manifestation of an island syndrome in rodents depends upon the trophic composition of a community, and may not relate to island size alone when many species additions, such as invasions, have occurred. The differing patterns of rodent population dynamics on each island provide information for future rodent eradication operations.     

Phylogeography of Wild Musk Shrew (Suncus Murinus) Populations in Asia Based on Blood Protein/Enzyme Variation

The musk shrew (Suncus murinus) is an insectivore species that inhabits tropical and subtropical Asia widely. To clarify the genetic relationship among wild musk shrew populations, we examined the electrophoretic variants of biparentally inherited genetic markers at 10 loci coding for eight blood proteins/enzymes in a total of 639 animals and compared the results obtained from the mitochondrial DNA data. The principal-component analysis performed using the allele frequency data revealed that the 17 populations could be divided into two major groups, a South Asian group and a Southeast Asian group that includes several island populations bound by Myanmar. The degrees of genetic divergence among populations were higher within the Southeast Asian group than within the South Asian group. This finding was incongruent with the mtDNA diversity. Analysis conducted at the individual level showed that a shrew from the central region in Myanmar that carries a South Asian type of mtDNA showed the electrophoretic variants specific to the Southeast Asian group, suggesting that this region is a contact zone between the two major groups.