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.     

Genetic diversity and population size: island populations of the common shrew, Sorex araneus

Populations of many species are currently being fragmented and reduced by human interactions. These processes will tend to reduce genetic diversity within populations and reduce individual heterozygosities because of genetic drift, inbreeding and reduced migration. Conservation biologists need to know the effect of population size on genetic diversity, as this is likely to influence a population's ability to persist. Island populations represent an ideal natural experiment with which to study this problem. In a study of common shrews (Sorex araneus) on offshore Scottish islands, 497 individuals from 13 islands of different sizes and 6 regions on the mainland were trapped and genotyped at eight microsatellite loci. Previous genetic work had revealed that most of the islands in this study were highly genetically divergent from one another and the mainland. We found that most of the islands exhibited lower genetic diversity than the mainland populations. In the island populations, mean expected heterozygosity, mean observed heterozygosity and mean allelic richness were significantly positively correlated with log island size and log population size, which were estimated using habitat population density data and application of a Geographic Information System.     

Population genetic structure of Sorex unguiculatus and Sorex caecutiens (Soricidae, Mammalia) in Hokkaido, based on microsatellite DNA polymorphism

We investigated the genetic structure of Sorex unguiculatus and Sorex caecutiens populations in Hokkaido, Japan, using hypervariable microsatellite DNA markers. We used five microsatellite loci to type 475 S. unguiculatus individuals from 20 localities on the Hokkaido mainland and four localities from each of four offshore islands (and 11 shrews from one locality in southern Sakhalin for a particular analysis). We used six microsatellite loci to type 240 S. caecutiens individuals from 13 localities on the Hokkaido mainland. Genetic variation was high in mainland populations of both species and low in the island populations of S. unguiculatus. Allelic richness and island size were positively correlated for S. unguiculatus, suggesting that genetic drift occurred on those islands due to small population size. In addition, four insular populations of S. unguiculatus were genetically differentiated from the mainland populations, although clear phylogeographic clustering was not confirmed among populations on the Hokkaido mainland for either S. unguiculatus or S. caecutiens. Heterozygosity excess was observed in more than half of the populations including the mainland populations of the two species, suggesting recent bottleneck events in these populations. Population dynamics of the shrews might be explained by a metapopulation scheme. According to autocorrelation analysis, the extent of non-random spatial genetic structure was approximately 100 km. Isolation by distance was observed in S. unguiculatus, but not in S. caecutiens although there is a positive trend. The lack of correlation for S. caecutiens might have been due to small sample size. Thus, no obvious differences in population genetic structure were found between the two species on the Hokkaido mainland in the present study, while previous investigations using mitochondrial DNA sequences inferred that these two species might have rather different biogeographic histories.     

Low outcrossing rates and shift in pollinators in New Zealand pohutukawa (Metrosideros excelsa; Myrtaceae)

New Zealand pohutukawa (Metrosideros excelsa), a member of the Myrtaceae, is a large, mass-flowering tree endemic to northern New Zealand coastlines. Mainland populations have been reduced to fragmented stands, and the original suite of bird pollinators has been largely replaced by introduced species. The native pollinator fauna on several offshore islands is largely intact and includes three species of the New Zealand honeyeaters (Meliphagidae) and native, solitary bees. We estimated multilocus outcrossing rates for three mainland and two island populations and found that they were among the lowest in the Myrtaceae (t(m) = 0.22-0.53). The shift in pollinators had no measurable effect on the mating system. Mass-flowering facilitates geitonogamous selfing, and inbreeding depression in seedling height was detectable at 6 mo of growth. F(s) [Wright's (1965) Fixation Index] was consistently higher than F(m) in all populations, indicating that selection may eliminate selfed offspring from populations prior to achieving reproductive maturity. Results suggest that increased selfing in mainland populations due to pollinator changes is not responsible for current patterns of poor regeneration of this species.     

Species limits and population differentiation in New Zealand snipes (Scolopacidae: <Emphasis Type="Italic">Coenocorypha</Emphasis>)

At least four species of New Zealand snipes (Coenocorypha) became extinct following the introduction of predatory mammals, and another two species suffered massive range reductions. To investigate species limits and population differentiation in six of the seven remaining offshore populations, we assayed variation in nine microsatellite loci and 1,980 base pairs of four mitochondrial DNA (mtDNA) genes. Genetic diversity in all populations except the largest one on Adams Island in the Auckland Islands was very low in both genomes. Alleles were fixed at many microsatellite loci and for single mtDNA haplotypes, particularly in the populations in the Chathams, Snares, Antipodes and Campbell Islands. Strong population structure has developed, and Chathams and Snares Islands populations are effectively genetically isolated from one another and from the more southern island populations. Based on reciprocal monophyly of lineages and their morphological distinctiveness we recommend that three phylogenetic species should be recognized, C. pusilla in the Chatham Islands, C. huegeli in the Snares Islands and C. aucklandica in the southern islands. The populations of C. aucklandica in the Auckland Islands, Antipodes Island and Campbell Island may warrant recognition as subspecies, and all should be managed as separate conservation units.     

Genetic structure of an introduced paper wasp,Polistes chinensis antennalis (Hymenoptera,Vespidae) in New Zealand

Several eusocial wasps are prominent invaders to remote islands. The paper wasp Polistes chinensis antennalis is native to East Asia, was introduced to New Zealand in 1979 and has expanded its distribution there. This provides an excellent opportunity to examine the impacts of an initial bottleneck and subsequent expansion on genetic structure. We analysed and compared the genetic population structures of the native (Japan and South Korea) and invasive New Zealand populations. Although 94% of individuals had shared haplotypes detected across both populations, the remaining 6% had private haplotypes identified in only one of the three countries. The genetic variation at microsatellite loci was lower in New Zealand than in native countries, and the genetic structure in New Zealand was clearly distinct from that in its native range. Higher frequencies of diploid‐male‐ and triploid‐female‐producing colonies were detected in New Zealand than in the native countries, showing the reduction in genetic variation via a genetic bottleneck. At least two independent introductions were suggested, and the putative source regions for New Zealand were assigned as Kanto (central island) and Kyushu (south island) in Japan. Serial founder events following the initial introduction were also indicated. The estimated dispersal distance between mother and daughter in New Zealand was twice that in Japan. Thus, the introduction history of P. chinensis antennalis in New Zealand is probably the result of at least two independent introductions, passing through a bottleneck during introduction, followed by population expansion from the point of introduction.     

Closing the gap: Avian lineage splits at a young,narrow seaway imply a protracted history of mixed population response

The evolutionary significance of spatial habitat gaps has been well recognized since Alfred Russel Wallace compared the faunas of Bali and Lombok. Gaps between islands influence population structuring of some species, and flightless birds are expected to show strong partitioning even where habitat gaps are narrow. We examined the population structure of the most numerous living flightless land bird in New Zealand, Weka (Gallirallus australis). We surveyed Weka and their feather lice in native and introduced populations using genetic data gathered from DNA sequences of mitochondrial genes and nuclear β‐fibrinogen and five microsatellite loci. We found low genetic diversity among extant Weka population samples. Two genetic clusters were evident in the mtDNA from Weka and their lice, but partitioning at nuclear loci was less abrupt. Many formerly recognized subspecies/species were not supported; instead, we infer one subspecies for each of the two main New Zealand islands. Although currently range restricted, North Island Weka have higher mtDNA diversity than the more wide‐ranging southern Weka. Mismatch and neutrality statistics indicate North Island Weka experienced rapid and recent population reduction, while South Island Weka display the signature of recent expansion. Similar haplotype data from a widespread flying relative of Weka and other New Zealand birds revealed instances of North Island—South Island partitioning associated with a narrow habitat gap (Cook Strait). However, contrasting patterns indicate priority effects and other ecological factors have a strong influence on spatial exchange at this scale.     

Systematic and conservation implications of geographic variation in pipits (Anthus: Motacillidae) in New Zealand and some offshore islands

The New Zealand Pipit Anthus novaeseelandiae occurs as a single species in Europe (as Richard's Pipit), Asia, Africa, New Guinea, Australia and New Zealand. In the New Zealand region, subspecific status has been accorded to allopatric populations on the mainland (A. n. novaeseelandiae) , on the Auckland and Campbell Islands (A. n. aucklandicus) , on the Antipodes Islands (A. n. steindachneri) and on the Chatham Islands (A. n. chathamensis) . Analyses of 23 allozyme loci and morphometric variation of populations on mainland New Zealand and on the Auckland, Campbell, Antipodes and Chatham Islands showed an appreciable and significant divergence between mainland and island populations. The magnitude of the allozyme difference is sufficient to suggest full species designation for the island birds, a conclusion supported by morphometric analyses and in sharp contrast to current taxonomy. However, formal taxonomic change will not be undertaken until the two New Zealand groups have been compared with more distant populations of this wideranging genus. The cryptic variation found in this species has conservation management implications for the small offshore populations.     

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.     

Genetic isolation of insular populations of the Maghrebian bat,Myotis punicus,in the Mediterranean Basin

Aim We investigate the population genetic structure of the Maghrebian bat, Myotis punicus, between the mainland and islands to assess the island colonization pattern and current gene flow between nearby islands and within the mainland. Location North Africa and the Mediterranean islands of Corsica and Sardinia. Methods We sequenced part of the control region (HVII) of 79 bats across 11 colonies. The phylogeographical pattern was assessed by analysing molecular diversity indices, examining differentiation among populations and estimating divergence time. In addition, we genotyped 182 bats across 10 colonies at seven microsatellite loci. We used analysis of molecular variance and a Bayesian approach to infer nuclear population structure. Finally, we estimated sex‐specific dispersal between Corsica and Sardinia. Results Mitochondrial analyses indicated that colonies between Corsica, Sardinia and North Africa are highly differentiated. Within islands there was no difference between colonies, while at the continental level Moroccan and Tunisian populations were highly differentiated. Analyses with seven microsatellite loci showed a similar pattern. The sole difference was the lack of nuclear differentiation between populations in North Africa, suggesting a male‐biased dispersal over the continental area. The divergence time of Sardinian and Corsican populations was estimated to date back to the early and mid‐Pleistocene. Main conclusions Island colonization by the Maghrebian bats seems to have occurred in a stepping‐stone manner and certainly pre‐dated human colonization. Currently, open water seems to prevent exchange of bats between the two islands, despite their ability to fly and the narrowness of the strait of Bonifacio. Corsican and Sardinian populations are thus currently isolated from any continental gene pool and must therefore be considered as different evolutionarily significant units (ESU).     

Genetic structure of kestrel populations and colonization of the Cape Verde archipelago

Genetic diversity and population structure were studied in eight populations of the kestrel Falco tinnunculus to identify the genetic consequences of spatial distribution and to infer the colonization patterns of the Cape Verde archipelago. We studied genetic differentiation and gene flow among seven island populations and one mainland population using nine microsatellite loci. Within the archipelago, differentiation was strong and genetic diversity and heterozygosity were low but variable among populations. Two subspecies F. tinnunculus neglectus on the northwestern islands and F. tinnunculus alexandri on all the other islands were identified as genetically distinct units. F. t. alexandri could be further separated into two groups on eastern and southern islands. Populations are probably founded by birds originating from the mainland. Immigration is more likely to the eastern and southern populations, whereas the northwestern islands with the lowest genetic diversity and highest differentiation are likely to exhibit fewer founding events by immigrants. The number of founding events on each island may depend not only on geographical distance to neighbouring populations, but also on directional immigration due to the northeastern trade winds. This may explain differences in genetic differentiation and diversity between populations and subspecies and may enable allopatric speciation.     

Evolutionary history of Scinax treefrogs on land‐bridge islands in south‐eastern Brazil

Aim We investigated how Pleistocene refugia and recent (c. 12,000 years ago) sea level incursions shaped genetic differentiation in mainland and island populations of the Scinax perpusillus treefrog group. Location Brazilian Atlantic Forest, São Paulo state, south‐eastern Brazil. Methods Using mitochondrial and microsatellite loci, we examined population structure and genetic diversity in three species from the S. perpusillus group, sampled from three land‐bridge islands and five mainland populations, in order to understand the roles of Pleistocene forest fragmentation and sea level incursions on genetic differentiation. We calculated metrics of relatedness and genetic diversity to assess whether island populations exhibit signatures of genetic drift and isolation. Two of the three island populations in this study have previously been described as new species based on a combination of distinct morphological and behavioural characters, thus we used the molecular datasets to determine whether phenotypic change is consistent with genetic differentiation. Results Our analyses recovered three distinct lineages or demes composed of northern mainland São Paulo populations, southern mainland São Paulo populations, and one divergent island population. The two remaining island populations clustered with samples from adjacent mainland populations. Estimates of allelic richness were significantly lower, and estimates of relatedness were significantly higher, in island populations relative to their mainland counterparts. Main conclusions Fine‐scale genetic structure across mainland populations indicates the possible existence of local refugia within São Paulo state, underscoring the small geographic scale at which populations diverge in this species‐rich region of the Atlantic Coastal Forest. Variation in genetic signatures across the three islands indicates that the populations experienced different demographic processes after marine incursions fragmented the distribution of the S. perpusillus group. Genetic signatures of inbreeding and drift in some island populations indicate that small population sizes, coupled with strong ecological selection, may be important evolutionary forces driving speciation on land‐bridge islands.     

Factors determining the number of land bird species on Islands around South-Western Australia

Summary New evidence from the passerine faunas of islands off Southwestern Australia agrees with the hypothesis that the passerine faunas of Australian and New Zealand islands are impoverished because most passerine species are poor colonizers. Dispersal of landbirds onto Carnac Island near Perth was infrequent, and many of those species that arrived were represented by single birds. Comparison of similarly structured island and mainland habitats showed that island habitats still have fewer passerine bird species than mainland habitats. Island bird faunas are more stable over short periods of time than over long periods; this is contrary to island avifaunas in the Northern Hemisphere.The following features typify the avifaunas of Australian islands: immigration of species of land birds occurs infrequently; (natural) extinction is rare; and the degree of saturation of the avifaunas is low. Without more direct evidence, competitive interactions should not be invoked to account for the species poverty of these insular avifaunas.     

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.     

Microsatellites reveal a lack of structure in Australian populations of the diamondback moth, Plutella xylostella (L.)

The diamondback moth, Plutella xylostella, is renowned for developing resistance to insecticides and causing significant economic damage to Brassica vegetable crops throughout the world. Yet despite its economic importance, little is known about the population structure and movement patterns of this pest both at local and regional scales. In Australia, the movement patterns and insecticide resistance status of P. xylostella infesting canola, vegetables, forage brassicas and weeds have fundamental implications for the management of this pest. Here we use six polymorphic microsatellite loci to investigate population structure and gene flow in Australian populations of P. xylostella. Samples of P. xylostella from New Zealand, Malaysia, Indonesia and Kenya were also scored at these loci. We found no evidence of population structure within Australia, with most populations having low inbreeding coefficients and in Hardy-Weinberg equilibrium. In addition, a sample from the North Island of New Zealand was indistinguishable from the Australian samples. However, large genetic differences were found between the Australia/New Zealand samples and samples from Kenya, Malaysia and Indonesia. There was no relationship between genetic distance and geographic distance among Australian and New Zealand samples. Two of the loci were found to have null alleles, the frequency of which was increased in the populations outside the Australia/New Zealand region. We discuss these results with reference to insecticide resistance management strategies for P. xylostella in Australia.     

Distribution and phylogenetic analyses of an endangered tick,Amblyomma sphenodonti

Abstract

In this study we investigate the geographic distribution, genetic diversity, and phylogenetic relationships of an endangered tick, Amblyomma sphenodonti (Family Ixodidae). Amblyomma sphenodonti and its host, the tuatara (Sphenodon), are found only on small offshore islands around New Zealand. Our results show that Amblyomma sphenodonti has a more severely restricted geographic distribution than its host, as it was found on only eight of 28 islands (four out of 12 island groups) where tuatara still live. The prevalence of A. sphenodonti is likely to have been affected by low host density and fluctuations in host population size as tuatara populations became isolated on offshore islands. Analysis of A. sphenodonti cytochrome oxidase 1 (CO1) sequences indicated a lack of gene flow between islands, with fixed differences in CO1 sequences between islands, but almost no genetic diversity within island populations. A similar phylogenetic pattern to that observed in tuatara mtDNA was observed, indicating co‐evolution of two species, at least since the Pleistocene. Phylogenetic analysis using 18S rRNA sequences suggest that A. sphenodonti is not closely related to other Amblyomma species, and that a separate genus for this species may be warranted. However, data from other ixodid ticks are required before the distinctiveness of A. sphenodonti can be confirmed and the phylogenetic relationships among ixodid ticks fully understood.     

The invasion biology of the invasive earwig, <Emphasis Type="Italic">Forficula auricularia</Emphasis> in Australasian ecosystems

The European earwig, Forficula auricularia, is a cosmopolitan insect endemic to Europe, West Asia and North Africa, which has invaded many temperate regions of the world including Australia and New Zealand. F. auricularia has been shown to be a complex of morphologically identical, reproductively isolated lineages that possess two distinct clades of mitochondrial DNA. Entomological collection data, historical literature and further field collections were used to develop a greater understanding of Australian F. auricularia’s invasion biology and its current distribution. Genetic analysis of F. auricularia collected from Australia and New Zealand using two mitochondrial genes (COI and a fragment overlapping parts of the COI -COII genes) was also undertaken. To identify the possible source populations of the Australasian invasion these sequences were compared to those from 16 locations within Britain and continental Europe. All Australasian populations were shown to be of the clade B lineage. Tasmanian and New Zealand populations consist of a single subclade comprised of only 4 and 1 haplotypes respectively. The Australian mainland populations also contained a second subclade consisting of up to 11 haplotypes indicating that multiple introductions possibly occurred on the Australian mainland. Comparison of mitochondrial genomes from Australasian and European populations showed the Australian mainland subclade to be most closely related to Portuguese haplotypes, and the Tasmanian and New Zealand clade closely related to those in Brittany, France. No European haplotypes perfectly matched the Australasian sequences. Therefore, the original source populations are still to be identified with harbours on the Iberian Peninsula’s western coast and those on the English Channel likely candidate areas.     

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.     

Identifying eradication units in an invasive mammalian pest species

Eradication of invasive species is an important component for species conservation and ecosystem restoration. Success of eradications is dependent on knowledge of population connectivity in order to determine reinvasion pathways, and hence populations requiring simultaneous eradication (eradication units). The common brushtail possum (Trichosurus vulpecula) was introduced into New Zealand from Australia and Tasmania, and now occupies a wide range of habitats across the majority of New Zealand. Possums are one of the most destructive invaders within New Zealand, with extensive control operations occurring throughout the country. Understanding the population connectivity of possums on mainland New Zealand (North and South Islands) will enhance the success of planned eradications. We examined the genetic population structure of invasive possums to identify gene flow, thus reinvasion pathways, between seven populations around Dunedin and on the Otago Peninsula where eradication of possums is occurring. Genetic variation at 12 microsatellite loci was comparable between all sampled populations and exhibited a significant isolation by distance pattern. Bayesian clustering methods supported the existence of two population clusters, indicating the presence of a reinvasion pathway onto the Otago Peninsula from urban areas at the Southern end of the Peninsula. To avoid recolonisation, all possums on the Otago Peninsula should be eradicated simultaneously, with the implementation and ongoing maintenance and monitoring of an urban buffer zone. We recommend pre-eradication genetic analyses be adopted by all pest managers to define appropriate eradication units, thereby maximising eradication success and avoiding costly failures.