Distribution and prevalence of the predatory planarian Artioposthia triangulata (Dendy) (Trieladida: Terricola) in Scotland

The distribution of the New Zealand flatworm (Artioposthia triangulata) in Scotland was surveyed between July 1991 and February 1993. There were 348 records from domestic gardens, 56 from botanic gardens, garden centres and nurseries, with only 13 from farms. Although most of the records came from around the major cities the flatworm was found to have become established throughout the Scottish mainland and some of the Islands, e.g. Bute, Gigha, Orkney and Skye. The impact of the flatworm on earthworm populations in agricultural land in Scotland was, as yet, found to be minimal but the fact that seven adjacent farms near Dunoon were infected suggested it could be spread from farm to farm and that in the West of Scotland it could become widespread in agricultural land.     

Molecular evidence for hybridization ofIlex x wandoensis (Aquifoliaceae) by RAPD analysis

Based on the presence of intermediate morphological characters, such as serrated leaf margins and flower structures,Ilex x wandoensis was initially described as a putative natural hybrid betweenI. cornuta andI. Integra, and was formally described as a new hybrid species,I. x wandoensis C. F. Mill., and M. Kim. However, using molecular markers generated via random amplified polymorphic DNA (RAPD), we have now discovered hybridization in populations of theI. x wandoensis complex collected from Wando and Jeju Islands, Korea. Marker bands of the putative parent taxa also were found in some populations ofI. x wandoensis, confirming its hybrid origin. Morphological variability within and among those populations was confirmed by model-based clustering methods, using multilocus genotype data. Phenograms generated from RAPD bands indicated that some accessions ofI. x wandoensis clustered with one of the parental species. This implied the occurrence of hybridization and recurring backcrosses of the hybrid to both parents, resulting in various hybrid derivatives because of the segregation and recombination of traits.Ilex x wandoensis was more closely related toI. cornuta than toI. integra suggesting that it backcrossed more with the former than with the latter.     

Multiple diploid hybrid speciation of the Canary Island endemicArgyranthemum sundingii (Asteraceae)

There are several well-documented examples of multiple hybrid origins of polyploid species. Herein we report the first, to our knowledge, explicit example of a species that most probably has originated recurrently by diploid hybrid speciation. Genetic relationships and stabilization of twoArgyranthemum populations of putative hybrid origin on Tenerife, the Canary Islands, were investigated using chromosomal, morphometric, and fertility analyses of cultivated progeny families and artificial F₁ and F₂ hybrids. These data were compared to a recently published chloroplast DNA phylogeny of the genus, in which the same populations were included. The results suggest that the two populations must be referred to a single species,A. sundingii, which is diploid, fully fertile, genetically stabilized, and occurs in an ecologically intermediate habitat opened by deforestation, and that this species has originated at least twice following local hybridization in two valleys. The same parental species were involved in each origin; the montaneA. broussonetii and the coastalA. frutescens. The montane species was the chloroplast donor in one of the valleys and the coastal species in the other.     

New chromosome counts inStreptocarpus (Gesneriaceae) from Madagascar and the Comoro Islands and their taxonomic significance

This study records the chromosome numbers of 10 species ofStreptocarpus; nine of the counts are new. With the exception ofS. buchananii of mainland Africa, all the results are for plants endemic to Madagascar and the Comoro Islands. While there is a strong correlation between basic number and growth form in the two subgenera of the genus on the African mainland (x = 15 among caulescent species in subgenusStreptocarpella; x = 16 among acaulescent species in subgenusStreptocarpus), the situation appears more complex among Madagascan and Comoro Island species. One notable example of deviation from this correlation is shown byS. papangae, a shrubby caulescent species, with 2n = 32 (x = 16). Polyploidy in the genus appears to be absent on mainland Africa, but is present in Madagascar and the Comoro Islands, ranging from tetraploidy to octoploidy. Evolutionary implications of the cytological observations are considered.     

Impact of geographical isolation on genetic differentiation in insular and mainland populations of Weigela coraeensis (Caprifoliaceae) on Honshu and the Izu Islands

Aim To provide insights into genetic differentiation between insular endemic Weigela coraeensis var. fragrans and its progenitor variety W. coraeensis var. coraeensis, the population genetic structure of both varieties was examined, and factors promoting genetic differentiation between the two taxa were explored. Location The natural range of W. coraeensis (sensu lato) throughout mainland Japan (Honshu) and the Izu Islands. Methods The analysis included 349 and 504 individuals across the mainland (Honshu) and the Izu Islands, respectively, using 10 allozyme and 10 microsatellite loci. The population genetic structure of W. coraeensis was assessed by analysing genetic diversity indices for each population, genetic differentiation among populations, model‐based Bayesian clustering or distance‐based clustering, and bottleneck tests. Results The level of genetic diversity in each of the populations on the Izu Islands was negatively correlated with geographical distance between each island and the mainland. The populations on the mainland and on the Izu Islands were genetically differentiated to a certain extent; however, the microsatellite analyses suggested that gene flow also occurred between the mainland and the islands, and among individual islands. These microsatellite analyses also suggested recent bottlenecks in several populations in both areas. Main conclusions The decrease in genetic diversity throughout the Izu Islands, which correlated with distance to the mainland, Honshu, may be the result of a repeated founder effect occurring at a series of inter‐island colonizations from north to south. The stepping stone‐like configuration of the islands may have played a role in the dispersal of the species. Geographical isolation by sea would effectively result in genetic differentiation of W. coraeensis between mainland Honshu and the Izu Islands, although some gene flow may still occur between Honshu and the northern Izu Islands. The differentiation process of the endemic plants on the Izu Islands is anagenetic but not completed, and the study of these plants will provide insightful knowledge concerning the evolution of insular endemics.     

Scrophularia arguta,a widespread annual plant in the Canary Islands: a single recent colonization event or a more complex phylogeographic pattern?

Many studies have addressed evolution and phylogeography of plant taxa in oceanic islands, but have primarily focused on endemics because of the assumption that in widespread taxa the absence of morphological differentiation between island and mainland populations is due to recent colonization. In this paper, we studied the phylogeography of Scrophularia arguta, a widespread annual species, in an attempt to determine the number and spatiotemporal origins of dispersal events to Canary Islands. Four different regions, ITS and ETS from nDNA and psbA‐trnH and psbJ‐petA from cpDNA, were used to date divergence events within S. arguta lineages and determine the phylogenetic relationships among populations. A haplotype network was obtained to elucidate the phylogenetic relationships among haplotypes. Our results support an ancient origin of S. arguta (Miocene) with expansion and genetic differentiation in the Pliocene coinciding with the aridification of northern Africa and the formation of the Mediterranean climate. Indeed, results indicate for Canary Islands three different events of colonization, including two ancient events that probably happened in the Pliocene and have originated the genetically most divergent populations into this species and, interestingly, a recent third event of colonization of Gran Canaria from mainland instead from the closest islands (Tenerife or Fuerteventura). In spite of the great genetic divergence among populations, it has not implied any morphological variation. Our work highlights the importance of nonendemic species to the genetic richness and conservation of island flora and the significance of the island populations of widespread taxa in the global biodiversity.     

Genetic diversity in the Eurasian otter, Lutra lutra, in Scotland. Evidence from microsatellite polymorphism

The relationship between microsatellite diversity and geographical fragmentation and isolation was studied in Scottish populations of the Eurasian otter, Lutra lutra. The geographic range of the study encompassed isolated archipelagos, islands adjacent to the Scottish mainland and both fragmented and continuous mainland populations. Tissue samples of 496 individuals from across Scotland were assayed for polymorphism at ten microsatellites. The isolation of populations on Shetland, and to a lesser degree on Orkney, was associated with reduced levels of microsatellite diversity. Most of the remaining island and fragmented mainland populations contained levels of microsatellite diversity similar to the high levels observed in die continuous mainland populations. Unexpectedly, both island and continuous mainland populations showed similar rates of departures from mutation-drift equilibrium. Such departures could have arisen from a variety of local demographic processes besides population bottlenecks. Gene flow appeared to be a major factor maintaining microsatellite diversity in all of these populations except the one on Shedand.     

Evolution and phylogeny of the New Zealand cicada genus Kikihia Dugdale (Homoptera: Auchenorrhyncha: Cicadidae) with special reference to the origin of the Kermadec and Norfolk Islands' species

Aim Determine the phylogeny and dispersal patterns of the cicada genus Kikihia in New Zealand and the origin of the Norfolk, Kermadec, and Chatham Island cicadas. Location New Zealand, Norfolk Island, Kermadec Islands and Chatham Island. Methods DNA sequences from 16 species and four soon to be described species of cicadas from New Zealand and Norfolk Island (Australia) were examined. A total of 1401 base pairs were analysed from whole genome extraction of three mitochondrial genes (cytochrome oxidase subunit II, ATPase6 and ATPase8). These DNA sequences were aligned and analysed using standard likelihood approaches to phylogenetic analysis. Dates of divergences between clades were determined using a molecular clock based on Bayesian statistics. Results Most species in the genus Kikihia diverged between 3 and 5 million years ago (Ma) coincident with a period of rapid mountain building in New Zealand. Cicada species on the Kermadec and Norfolk Islands invaded recently from New Zealand and are closely related to the New Zealand North Island species Kikihia cutora. Main conclusions Speciation in the genus Kikihia was likely due in large part to the appearance of new habitats associated with the rise of the Southern Alps, starting c. 5 Ma. Dispersal of Kikihia species within mainland New Zealand probably occurred gradually rather than through long‐distance jumps. However, invasion of Norfolk, the Kermadecs and Chatham Islands had to have occurred through long‐distance dispersal.     

DNA bar-coding reveals source and patterns of <Emphasis Type="Italic">Thaumastocoris peregrinus</Emphasis> invasions in South Africa and South America

Thaumastocoris peregrinus is a recently introduced invertebrate pest of non-native Eucalyptus plantations in the Southern Hemisphere. It was first reported from South Africa in 2003 and in Argentina in 2005. Since then, populations have grown explosively and it has attained an almost ubiquitous distribution over several regions in South Africa on 26 Eucalyptus species. Here we address three key questions regarding this invasion, namely whether only one species has been introduced, whether there were single or multiple introductions into South Africa and South America and what the source of the introduction might have been. To answer these questions, bar-coding using mitochondrial DNA (COI) sequence diversity was used to characterise the populations of this insect from Australia, Argentina, Brazil, South Africa and Uruguay. Analyses revealed three cryptic species in Australia, of which only T. peregrinus is represented in South Africa and South America. Thaumastocoris peregrinus populations contained eight haplotypes, with a pairwise nucleotide distance of 0.2–0.9% from seventeen locations in Australia. Three of these haplotypes are shared with populations in South America and South Africa, but the latter regions do not share haplotypes. These data, together with the current distribution of the haplotypes and the known direction of original spread in these regions, suggest that at least three distinct introductions of the insect occurred in South Africa and South America before 2005. The two most common haplotypes in Sydney, one of which was also found in Brisbane, are shared with the non-native regions. Sydney populations of T. peregrinus, which have regularly reached outbreak levels in recent years, might thus have served as source of these three distinct introductions into other regions of the Southern Hemisphere.     

Diploicia canescens (Dicks.) Massal. in Shetland

Summary

The lichenized fungus Diploicia canescens (Dicks.) Massal. has recently been discovered in Shetland. Its world distribution is centred in Europe where it is essentially a western and southern species. Its habitats in Shetland are discussed in comparison with its stations in Orkney and in mainland Britain. It is concluded that the species does not occupy significantly different habitats at the extreme north of its world range, although the precise environmental controls for this species must remain partly unidentified.     

Polyploidy and microsatellite variation in the relict tree Prunus lusitanica L.: how effective are refugia in preserving genotypic diversity of clonal taxa?

Refugia are expected to preserve genetic variation of relict taxa, especially in polyploids, because high gene dosages could prevent genetic erosion in small isolated populations. However, other attributes linked to polyploidy, such as asexual reproduction, may strongly limit the levels of genetic variability in relict populations. Here, ploidy levels and patterns of genetic variation at nuclear microsatellite loci were analysed in Prunus lusitanica, a polyploid species with clonal reproduction that is considered a paradigmatic example of a Tertiary relict. Sampling in this study considered a total of 20 populations of three subspecies: mainland lusitanica (Iberian Peninsula and Morocco), and island azorica (Azores) and hixa (Canary Islands and Madeira). Flow cytometry results supported an octoploid genome for lusitanica and hixa, whereas a 16‐ploid level was inferred for azorica. Fixed heterozygosity of a few allele variants at most microsatellite loci resulted in levels of allelic diversity much lower than those expected for a high‐order polyploid. Islands as a whole did not contain higher levels of genetic variation (allelic or genotypic) than mainland refuges, but island populations displayed more private alleles and higher genotypic diversity in old volcanic areas. Patterns of microsatellite variation were compatible with the occurrence of clonal individuals in all but two island populations, and the incidence of clonality within populations negatively correlated with the estimated timing of colonization. Our results also suggest that gene flow has been very rare among populations, and thus population growth following founder events was apparently mediated by clonality rather than seed recruitment, especially in mainland areas. This study extends to clonal taxa the idea of oceanic islands as important refugia for biodiversity, since the conditions for generation and maintenance of clonal diversity (i.e. occasional events of sexual reproduction, mutation and/or seed immigration) appear to have been more frequent in these enclaves than in mainland areas.     

The genome sequence of the biocontrol fungus Metarhizium anisopliae and comparative genomics of Metarhizium species

Background

Metarhizium anisopliae is an important fungal biocontrol agent of insect pests of agricultural crops. Genomics can aid the successful commercialization of biopesticides by identification of key genes differentiating closely related species, selection of virulent microbial isolates which are amenable to industrial scale production and formulation and through the reduction of phenotypic variability. The genome of Metarhizium isolate ARSEF23 was recently published as a model for M. anisopliae, however phylogenetic analysis has since re-classified this isolate as M. robertsii. We present a new annotated genome sequence of M. anisopliae (isolate Ma69) and whole genome comparison to M. robertsii (ARSEF23) and M. acridum (CQMa 102).

Results

Whole genome analysis of M. anisopliae indicates significant macrosynteny with M. robertsii but with some large genomic inversions. In comparison to M. acridum, the genome of M. anisopliae shares lower sequence homology. While alignments overall are co-linear, the genome of M. acridum is not contiguous enough to conclusively observe macrosynteny. Mating type gene analysis revealed both MAT1-1 and MAT1-2 genes present in M. anisopliae suggesting putative homothallism, despite having no known teleomorph, in contrast with the putatively heterothallic M. acridum isolate CQMa 102 (MAT1-2) and M. robertsii isolate ARSEF23 (altered MAT1-1). Repetitive DNA and RIP analysis revealed M. acridum to have twice the repetitive content of the other two species and M. anisopliae to be five times more RIP affected than M. robertsii. We also present an initial bioinformatic survey of candidate pathogenicity genes in M. anisopliae.

Conclusions

The annotated genome of M. anisopliae is an important resource for the identification of virulence genes specific to M. anisopliae and development of species- and strain- specific assays. New insight into the possibility of homothallism and RIP affectedness has important implications for the development of M. anisopliae as a biopesticide as it may indicate the potential for greater inherent diversity in this species than the other species. This could present opportunities to select isolates with unique combinations of pathogenicity factors, or it may point to instability in the species, a negative attribute in a biopesticide.

Electronic supplementary material

The online version of this article (doi:10.1186/1471-2164-15-660) contains supplementary material, which is available to authorized users.     

Committee for the study of Scottish Flora

Summary

Evidence is presented for and against the proposal that two new plant species have originated in Scotland in the very recent past as a result of interspecific hybridization between plants that have colonized ‘man-made’ habitats. Isozyme and chloroplast DNA evidence has confirmed a recent origin of the new allopolyploid species Welsh groundsel, Senecio cambrensis, in Leith, Edinburgh. Scottish and Welsh forms of the species differ in a number of morphological and life history features, but overall are very similar in morphology. A survey of isozyme variation in what has been considered as another new hybrid species, Epipactis youngiana (Young's helleborine), and its putative parent species, has cast doubt on its supposed origins, i.e. by hybridization between E. helleborine x E. leptochila or E. helleborine x E. phyllanthes and the stabilization of a hybrid product via autogamy. Contrary to expectation, E. youngiana was found to contain a high level of genetic diversity, and a genotypic structure that is indicative of random mating. Moreover, at a Glasgow site it appears to be interbreeding with two of its putative parents, E. helleborine and E. leptochila, to form part of a hybrid swarm. From the isozyme data, it is possible to dismiss the idea that E. phyllanthes may have acted as a parent of E. youngiana; however, it can not be ruled out that the ‘new species’ is merely a variant form of E. helleborine or an introduction rather than a product of hybridization.     

Evolution and history of hummingbirds (Aves: Trochilidae) from the Juan Fernandez Islands, Chile

The Juan Fernandez Firecrown Sephanoides fernandensis is an endangered endemic hummingbird that inhabits the Juan Fernandez Islands, 667 km off the coast of Chile. Its population has decreased from several thousand in the early part of this century to approximately 250–400 individuals at present. The reasons for its decline include habitat degradation by anthropogenic forest clearance and the introduction of grazing mammals and rodents. Another hummingbird, the Green-backed Firecrown Sephanoides sephaniodes, inhabits the Juan Fernandez Islands but is also found on the Chilean mainland. It currently numbers several thousand on the Juan Fernandez Islands but was considered rare during the late 19th and early 20th centuries. The sister relationship between the two species has not been critically tested, and so their evolutionary histories on the Juan Fernandez Islands remain uncertain. With the use of mtDNA cytochrome b and ND2 phylogenetic reconstructions, our study supports the two species as sister taxa. Moreover, the molecular data suggest that the genus Sephanoides is closely related to the higher altitude Andean hummingbirds typical of the paramo and puna habitats. The molecular divergence between the two species of Sephanoides indicates they may have become isolated from each other less than 1 million years ago, suggesting that S. fernandensis evolved in situ on the Juan Fernandez Islands. We find no evidence of genetic subdivision between populations of S. sephaniodes from the Juan Fernandez Islands and the mainland. In addition, high genetic variation of the Juan Fernandez Islands population does not indicate a long period of isolation of a limited number of S. sephaniodes but instead suggests a recent colonization event, perhaps from several mainland populations. As a result of molecular, morphological and apparent ecological similarities, we suggest that competition by S. sephaniodes may be an additional factor stifling the recovery of S. fernandensis. Possible conservation strategies include habitat restoration and the removal of introduced mammals; immediate implementation of such conservation management plans are necessary to save this species from extinction.     

VII. A note on the Institute of Seaweed Research

A new genus and two new species are described and compared with related algae. All were isolated from terrestrial habitats on Signy Island, South Orkney Islands, Antarctica. They are Signiosphaera multinucleata gen. et sp. nov. (Tetrasporales, Chlorophyceae), Chlamydocapsa lobata sp. nov. (Tetrasporales, Chlorophyceae) and Chloromonas palmelloides sp. nov. (Volvocales, Chlorophyceae).     

Inversion Polymorphisms in Populations of Drosophila Melanogaster in the South-West Islands of Japan: Comparisons with the Mainland Populations

Natural populations of D. melanogaster newly collected from the South-West Islands of Japan were examined for the frequencies of inversions, and the results, together with previous data, were compared with those of the mainland populations. The four Common cosmopolitan inversions (2Lt, 2RNS, 3LP, 3RP) were detected from most of the populations with higher frequencies. The three Rare cosmopolitan inversions (3LM, 3RC, 3RMo), three Quasi cosmopolitan inversions (2LA, 3LY, 3RK) and two new Endemic inversions were also found but with lower frequencies. Some of the South-West Island populations still maintained higher levels of inversion polymorphisms compared to the mainland populations, while others lost the polymorphisms to various degrees. The South-West Island populations were characterized by the higher frequency of In(3L)P. The ratios of In(3L)P/In(3R)P in the South-West Islands are significantly higher than those of the mainland populations, suggesting that the In(3L)P is relatively adaptive in the South-West Islands.     

Parasites and the island syndrome: the colonization of the western Mediterranean islands by Heligmosomoides polygyrus (Dujardin, 1845)

Aim Populations of free‐living vertebrates on islands frequently differ from their mainland counterparts by a series of changes in morphometric, life‐history, behavioural, physiological and genetic traits, collectively referred to as the ‘island syndrome’. It is not known, however, whether the ‘island syndrome’ also affects parasitic organisms. The present study establishes the colonization pattern of the Mediterranean islands by the nematode Heligmosomoides polygyrus, a direct and specific parasite of rodent hosts of the Apodemus genus, and evaluates the effects of island colonization by this species on two components of the island syndrome: the loss of genetic diversity and the enlargement of the ecological niche. Location Heligmosomoides polygyrus was sampled on seven western Mediterranean islands ? Corsica, Crete, Elba, Majorca, Minorca, Sardinia and Sicily ? as well as in 20 continental locations covering the Mediterranean basin. Methods The mitochondrial cytochrome b gene (690 base pairs) was sequenced in 166 adult H. polygyrus individuals sampled in the 27 continental and island locations. Phylogenetic reconstructions in distance, parsimony, maximum likelihood and Bayesian posterior probabilities were carried out on the whole cytochrome b gene data set. The levels of nucleotide, haplotype and genetic divergence (Kimura two‐parameter distance estimator) diversities were estimated in each island population and in the various continental lineages. Results Phylogenetic reconstructions show that the mainland origins of H. polygyrus were continental Spain for the Balearic Islands (Majorca, Minorca), northern Italy for the Tyrrhenian Islands (Corsica, Sardinia, Elba), southern Italy for Sicily, and the Balkan region for Crete. A comparison of island H. polygyrus populations with their mainland source populations revealed two characteristic components of the island syndrome in this parasite. First, island H. polygyrus populations display a significant loss of genetic diversity, which is related (r² = 0.73) to the distance separating the island from the mainland source region. Second, H. polygyrus exhibits a niche enlargement following insularization. Indeed, H. polygyrus in Corsica is present in both A. sylvaticus and Mus musculus domesticus, while mainland H. polygyrus populations are present exclusively in Apodemus hosts. Main conclusions Our results show that H. polygyrus has undergone a loss of genetic diversity and a niche (host) enlargement following colonization of the western Mediterranean islands. To our knowledge, this study provides the first evidence for components of the ‘island syndrome’ in a parasitic nematode species.     

A genomic approach to identify hybrid incompatibility genes

Uncovering the genetic and molecular basis of barriers to gene flow between populations is key to understanding how new species are born. Intrinsic postzygotic reproductive barriers such as hybrid sterility and hybrid inviability are caused by deleterious genetic interactions known as hybrid incompatibilities. The difficulty in identifying these hybrid incompatibility genes remains a rate-limiting step in our understanding of the molecular basis of speciation. We recently described how whole genome sequencing can be applied to identify hybrid incompatibility genes, even from genetically terminal hybrids. Using this approach, we discovered a new hybrid incompatibility gene, gfzf, between Drosophila melanogaster and Drosophila simulans, and found that it plays an essential role in cell cycle regulation. Here, we discuss the history of the hunt for incompatibility genes between these species, discuss the molecular roles of gfzf in cell cycle regulation, and explore how intragenomic conflict drives the evolution of fundamental cellular mechanisms that lead to the developmental arrest of hybrids.     

Variation in coat colour gene frequencies and selection in the cats of Scotland

Estimates of Felis catus coat colour gene frequencies were obtained from samples in 27 Scottish localities. These areas ranged from island samples in Shetland, Orkney and the Outer Hebrides to remote highland areas on the Scottish mainland. Animals from several towns and cities were also sampled. The results reveal considerable heterogeneity in the frequencies of the O, a, t ^b, l, S and W alleles. The frequencies of d were relatively homogeneous throughout Scotland. There were indications that island and mainland cat populations may have been derived from separate ancestral gene pools. Much of the observed gene frequency heterogeneity in rural areas could be attributed to differential migration or lack of penetrance. Geographical isolation and possible hybridization with F. sylvestris may explain the high frequency of wild-type alleles in mainland rural areas. A correlation between cat population coefficient of darkness and human population was observed. Such an observation supports the concept of cumulative urban pressure and indicates that a process analogous to industrial melanism is occurring in urban cat populations. The darkening of phenotype appears to be acting through the t, a and O loci and may be the result of pleiotropic effects of alleles at these loci.     

ANALYSIS OF MITOCHONDRIAL DNA POLYMORPHISMS AMONG CHANNEL ISLAND DEER MICE

Mitochondrial DNA (mtDNA) from 131 deer mice, Peromyscus maniculatus, collected on the eight California Channel Islands and from seven southern California mainland locations, was isolated and analyzed for restriction endonuclease fragment polymorphisms. A total of 26 mtDNA genotypes were distinguishable among the deer mice sampled. All of the island samples had mtDNA restriction-fragment patterns not found among the mainland samples. Distributions of specific restriction-fragment patterns provide evidence for at least four separate colonization events to the Channel Islands. The estimated percentage of sequence divergence between all mtDNA's in this study was less than 1%, suggesting that colonization of the islands occurred fairly recently, probably within the last 500,000 years. Levels of mtDNA heterogeneity were much lower within island populations than within mainland populations.