Biogeography and diversification of the Pacific ant genus Lordomyrma Emery

Aim This study addresses the origins of terrestrial biodiversity of the Fijian islands using the ant genus Lordomyrma (Hymenoptera: Formicidae: Myrmicinae) as a model system. We derive the evolution of the genus and determine its closest extra-Fijian relatives from geological data, molecular phylogenetic reconstruction and divergence estimates. Location Ant taxa were sampled in the Southwest Pacific, Melanesia, Southeast Asia, Australia and mainland China. Methods Phylogeny and divergence estimates of the ant genus Lordomyrma based on four nuclear genes (28S, ArgK, LW Rh, CAD) plus data on Indo-Pacific geological history are used to address current hypotheses regarding the origins of the Fijian biota. Results The genus Lordomyrma probably originated in mainland Asia, with subsequent colonization of Australia and the Pacific. The Fijian Lordomyrma clade is monophyletic, and originated c. 8.8 Ma, when it diverged from a sister group in Papua New Guinea. Main conclusions The colonization of Fiji by Lordomyrma is probably a result of long-distance dispersal from New Guinea, possibly aided by island hopping across the Vitiaz Arc. The timeline of diversification in Lordomyrma is broadly congruent with the Miocene fragmentation of the Vitiaz Arc and the Pliocene emergence of Vanua Levu. The biotic shuttle hypothesis, which posits ‘Eua Island as the source of Fijian endemics, is rejected based on the sister relationship of Fiji and New Guinea lineages, as well as on the Miocene submergence of the terrane below sea level. The diversity of Fijian Lordomyrma results from the radiation of a single lineage, which diverged from a New Guinea sister group. The genus appears to have originated in Asia rather than in Australia.     

Genetic variability,local selection and demographic history: genomic evidence of evolving towards allopatric speciation in Asian seabass

Genomewide analysis of genetic divergence is critically important in understanding the genetic processes of allopatric speciation. We sequenced RAD tags of 131 Asian seabass individuals of six populations from South‐East Asia and Australia/Papua New Guinea. Using 32 433 SNPs, we examined the genetic diversity and patterns of population differentiation across all the populations. We found significant evidence of genetic heterogeneity between South‐East Asian and Australian/Papua New Guinean populations. The Australian/Papua New Guinean populations showed a rather lower level of genetic diversity. F_ST and principal components analysis revealed striking divergence between South‐East Asian and Australian/Papua New Guinean populations. Interestingly, no evidence of contemporary gene flow was observed. The demographic history was further tested based on the folded joint site frequency spectrum. The scenario of ancient migration with historical population size changes was suggested to be the best fit model to explain the genetic divergence of Asian seabass between South‐East Asia and Australia/Papua New Guinea. This scenario also revealed that Australian/Papua New Guinean populations were founded by ancestors from South‐East Asia during mid‐Pleistocene and were completely isolated from the ancestral population after the last glacial retreat. We also detected footprints of local selection, which might be related to differential ecological adaptation. The ancient gene flow was examined and deemed likely insufficient to counteract the genetic differentiation caused by genetic drift. The observed genomic pattern of divergence conflicted with the ‘genomic islands’ scenario. Altogether, Asian seabass have likely been evolving towards allopatric speciation since the split from the ancestral population during mid‐Pleistocene.     

Mosquito seasonality and arboviral disease incidence in Murray Valley, southeast Australia

Abstract. Adult female mosquito populations were monitored at weekly intervals during spring-autumn (November-March) for 4 years (1991-95) using dry-ice-baited light traps at forty sites in the Murray Valley of Victoria, Australia. Among twenty species of mosquitoes collected, Culex annulirostriswas the most abundant (66.6 ± 9.3%) followed by Cx australicus(15.3 ± 7.7%). From a total of 476, 682 mosquitoes collected, nearly all were females and only 1295 (0.27%) were males. Mosquito population densities were generally higher in 1992-93 and 1993-94 seasons than in 1991-92 and 1994-95 seasons. Greatest densities of Cx annulirostrisand Cx australicusoccurred in 1992-93, coinciding with outbreaks of Ross River (RR) and Barman Forest (BF) arboviruses causing human polyarthritis. In the majority of shires, Cx australicuswas the predominant species from spring to early summer (November and December), then was replaced by Cx annulirostrisfrom mid-summer to autumn (January-April). In three shires, Aedes bancroftianusand Ae.sagaxpredominated during the early part of the season. Densities of both Cx annulirostrisand Cx australicuswere related to temperature. Cx australicusadults were found to be trapped when the mean ambient temperature exceeded 6d?C, with peak population recorded at 20d?C. Cx annulirostrisadult density increased when the mean temperature rose above 12d?C, reaching a peak during February and March when temperature exceeded 25d?C. Cx annulirostrisdeclined rapidly from April onwards, with no adult activity evident from May to November. Population densities of Aedesspp. were generally less than reported from earlier studies, possibly due to lower rainfall in spring and summer as well as reduced flood irrigation practices. In each year, a significant correlation was detected between Cx annulirostrisdensity and RR virus incidence in humans. As Cx annulirostrisis the predominant local mosquito species and feeds on a wide spectrum of hosts including man, it seems likely that Cx annulirostrisis the major vector of RR in the inland Murray Valley region.     

Characterization of population structure from the mitochondrial DNA vis‐à‐vis language and geography in Papua New Guinea

Situated along a corridor linking the Asian continent with the outer islands of the Pacific, Papua New Guinea has long played a key role in understanding the initial peopling of Oceania. The vast diversity in languages and unique geographical environments in the region have been central to the debates on human migration and the degree of interaction between the Pleistocene settlers and newer migrants. To better understand the role of Papua New Guinea in shaping the region's prehistory, we sequenced the mitochondrial DNA (mtDNA) control region of three populations, a total of 94 individuals, located in the East Sepik Province of Papua New Guinea. We analyzed these samples with a large data set of Oceania populations to examine the role of geography and language in shaping population structure within New Guinea and between the region and Island Melanesia. Our results from median‐joining networks, star‐cluster age estimates, and population genetic analyses show that while highland New Guinea populations seem to be the oldest settlers, there has been significant gene flow within New Guinea with little influence from geography or language. The highest genetic division is between Papuan speakers of New Guinea versus East Papuan speakers located outside of mainland New Guinea. Our study supports the weak language barriers to genetic structuring among populations in close contact and highlights the complexity of understanding the genetic histories of Papua New Guinea in association with language and geography. Am J Phys Anthropol 142:613–624, 2010. © 2010 Wiley‐Liss, Inc.     

Population genetics of Anopheles koliensis through Papua New Guinea: New cryptic species and landscape topography effects on genetic connectivity

New Guinea is a topographically and biogeographically complex region that supports unique endemic fauna. Studies describing the population connectivity of species through this region are scarce. We present a population and landscape genetic study on the endemic malaria‐transmitting mosquito, Anopheles koliensis (Owen). Using mitochondrial and nuclear sequence data, as well as microsatellites, we show the evidence of geographically discrete population structure within Papua New Guinea (PNG). We also confirm the existence of three rDNA ITS2 genotypes within this mosquito and assess reproductive isolation between individuals carrying different genotypes. Microsatellites reveal the clearest population structure and show four clear population units. Microsatellite markers also reveal probable reproductive isolation between sympatric populations in northern PNG with different ITS2 genotypes, suggesting that these populations may represent distinct cryptic species. Excluding individuals belonging to the newly identified putative cryptic species (ITS2 genotype 3), we modeled the genetic differences between A. koliensis populations through PNG as a function of terrain and find that dispersal is most likely along routes with low topographic relief. Overall, these results show that A. koliensis is made up of geographically and genetically discrete populations in Papua New Guinea with landscape topography being important in restricting dispersal.     

Evolutionary origins of taro (Colocasia esculenta) in Southeast Asia

As an ancient clonal root and leaf crop, taro (Colocasia esculenta, Araceae) is highly polymorphic with uncertain genetic and geographic origins. We explored chloroplast DNA diversity in cultivated and wild taros, and closely related wild taxa, and found cultivated taro to be polyphyletic, with tropical and temperate clades that appear to originate in Southeast Asia sensu lato. A third clade was found exclusively in wild populations from Southeast Asia to Australia and Papua New Guinea. Our findings do not support the hypothesis of taro domestication in Papua New Guinea, despite archaeological evidence for early use or cultivation there, and the presence of apparently natural wild populations in the region (Australia and Papua New Guinea).     

Mosquito host-feeding patterns and implications for Japanese encephalitis virus transmission in northern Australia and Papua New Guinea

Japanese encephalitis (JE) virus spread to northern Australia during the 1990s, transmitted by Culex annulirostris Skuse and other mosquitoes (Diptera: Culicidae). To determine the relative importance of various hosts for potential vectors of JE virus, we investigated the host-feeding patterns of mosquitoes in northern Australia and Western Province of Papua New Guinea, with particular attention to pigs, Sus scrofa L. - the main amplifying host of JE virus in South-east Asia. Mosquitoes were collected by CDC light traps baited with dry ice and 1-octen-3-ol, run 16.00-08.00 hours, mostly set away from human habitations, if possible in places frequented by feral pigs. Bloodmeals of 2569 mosquitoes, representing 15 species, were identified by gel diffusion assay. All species had fed mostly on mammals: only <10% of bloodmeals were from birds. The predominant species was Cx. annulirostris (88%), with relatively few (4.4%) bloodmeals obtained from humans. From all 12 locations sampled, the mean proportion of Cx. annulirostris fed on pigs (9.1%) was considerably lower than fed on other animals (90.9%). Highest rates of pig-fed mosquitoes (>30%) were trapped where domestic pigs were kept close to human habitation. From seven of eight locations on the Australian mainland, the majority of Cx. annulirostris had obtained their bloodmeals from marsupials, probably the Agile wallaby Macropus agilis (Gould). Overall proportions of mosquito bloodmeals identified as marsupial were 60% from the Gulf Plains region of Australia, 78% from the Cape York Peninsula and 64% from the Daru area of Papua New Guinea. Thus, despite the abundance of feral pigs in northern Australia, our findings suggest that marsupials divert host-seeking Cx. annulirostris away from pigs. As marsupials are poor JE virus hosts, the prevalence of marsupials may impede the establishment of JE virus in Australia.     

Bats Without Borders: Long-Distance Movements and Implications for Disease Risk Management

Fruit bats of the genus Pteropus (commonly known as flying-foxes) are the natural hosts of several recently emerged zoonotic viruses of animal and human health significance in Australia and Asia, including Hendra and Nipah viruses. Satellite telemetry was used on nine flying-foxes of three species (Pteropus alecto n = 5, P. vampyrus n = 2, and P. neohibernicus n = 2) to determine the scale and pattern of their long-distance movements and their potential to transfer these viruses between countries in the region. The animals were captured and released from six different locations in Australia, Papua New Guinea, Indonesia, and Timor-Leste. Their movements were recorded for a median of 120 (range, 47–342) days with a median total distance travelled of 393 (range, 76–3011) km per individual. Pteropus alecto individuals were observed to move between Australia and Papua New Guinea (Western Province) on four occasions, between Papua New Guinea (Western Province) and Indonesia (Papua) on ten occasions, and to traverse Torres Strait on two occasions. Pteropus vampyrus was observed to move between Timor-Leste and Indonesia (West Timor) on one occasion. These findings expand upon the current literature on the potential for transfer of zoonotic viruses by flying-foxes between countries and have implications for disease risk management and for the conservation management of flying-fox populations in Australia, New Guinea, and the Lesser Sunda Islands.     

Genetic diversity and colony structure of Tapinoma melanocephalum on the islands and mainland of South China

Aim

Tapinoma melanocephalum is listed as one of the most important invasive pest species in China. Information regarding the patterns of invasion and effects of geographic isolation on the population genetics of this species is largely lacking.

Location

South China.

Methods

To address this problem, we genotyped 39 colonies (two colonies were collapsed due to genetic similarity) using microsatellite markers and mitochondrial DNA sequencing to compare colony genetic structure of T. melanocephalum on the mainland and islands of South China.

Results

An analysis of the colony genotypes showed that the genetic diversity of the mainland population was slightly higher than that of the island populations but not significantly so. However, the observed heterozygosity on Shangchuan Island (SCD) was significantly lower than that of the other colonies. We also found six haplotypes in 111 mitochondrial DNA COI sequences. The relatedness (r) value between colonies of SCD was 0.410, higher than that of the other populations. The genetic clusters among colonies were not related to geographic locations and exhibited admixture likely due to frequent human‐mediated dispersal associated with trade between the mainland population and the islands. Pairwise F_STs between populations showed differentiation among mainland populations, while SCD displayed high levels of divergence (F_ST > 0.15) from most mainland populations. There was no significant isolation by distance among colonies. Most populations showed signs of a bottleneck effect.

Main conclusions

Our study suggests that there was no significant difference in the genetic diversity among the islands and the mainland; however, the lower genetic diversity, the higher degree of genetic divergence from other colonies, and the higher relatedness among nestmates made the SCD population stand out from all the others.     

Contributions to the Cladocera fauna from Papua New Guinea

Twenty-eight taxa of the Cladocera are identified in collections from Papua New Guinea, 17 being new records for New Guinea, bringing the total number of Cladocera taxa reported for this region to 39. Most of the taxa are circumtropical. One species (Sarsilatona papuana) is endemic to Papua New Guinea and northern Australia. The species list includes two species that are normally listed as Holarctic:Alonella nana andAlona rustica. Widespread genera such asDaphnia, Pleuroxus, Disparalona, Acroperus were strikingly absent from the Papua New Guinean material.     

High species richness and genetic diversity of the genus Caloglossa (Delesseriaceae,Rhodophyta) in New Ireland,Papua New Guinea

Distribution patterns and genetic diversity of Caloglossa species have been studied in various regions but are still poorly understood around the tropical western Pacific where many marine organisms show high species richness. We explored the diversity of Caloglossa species in New Ireland, Papua New Guinea using morphological and genetic analyses. Seven species of Caloglossa (C. adhaerens, C. leprieurii, C. ogasawaraensis, C. bengalensis, C. postiae, C. saigonensis and C. vieillardii) were collected from eight sites; the latter four species are new records in Papua New Guinea. All specimens collected were cultured to compare their morphology under the same culture condition. Reproductive structures of gametophytes and/or sporophytes were described for the first time in some species from these collections. In C. leprieurii, gall‐like structure appeared in most gametophyte strains but not in tetrasporophytes. Such galls were produced from wing cells, and spermatangia and/or procarps were formed on many galls. The LSU rDNA and rbcL analyses revealed high intraspecific diversity in many species although the number of samples was limited and the sampling sites were within 200 km. We revealed high species richness and intraspecific genetic variation in Papua New Guinea, suggesting a hot spot for the diversity of Caloglossa.     

The mosquito and predatory insect fauna inhabiting fresh-water ponds,with particular reference to Culex annulirostris Skuse (Diptera: Culicidae)

Colonization and abundance of mosquitoes and other insects were studied during four trials in man-made ponds at Mildura, Victoria. In December and February, respectively, Culex australicus and Cx annulirostris colonized the ponds within 1 day of formation. Maximum larval densities were attained within 8 days and, thereafter, densities gradually declined. The ponds were also colonized initially by predatory Coleoptera and Hemiptera, then Odonata. In three of the four trials, a significant inverse relationship existed between mosquito and predator densities. From life tables compiled during one of these trials, a survival rate of Cx annulirostris, from egg hatch to eclosion, was estimated to be 11%. Predation was considered to be largely responsible for the low survival. The results are discussed in relation to control of Cx annulirostris in temporary and permanent grassy ponds.     

Island biogeography theory explains the genetic diversity of a fragmented rock ptarmigan (Lagopus muta) population

The island biogeography theory is one of the major theories in ecology, and its applicability to natural systems is well documented. The core model of the theory, the equilibrium model of island biogeography, predicts that species diversity on an island is positively related to the size of the island, but negatively related by the island's distance to the mainland. In recent years, ecologists have begun to apply this model when investigating genetic diversity, arguing that genetic and species diversity might be influenced by similar ecological processes. However, most studies have focused on oceanic islands, but knowledge on how the theory applies to islands located on the mainland (e.g., mountain islands, forest islands) is scarce. In this study, we examined how the size and degree of isolation of mountain islands would affect the genetic diversity of an alpine bird, the rock ptarmigan (Lagopus muta). Within our study area, we defined the largest contiguous mountain area as the mainland, while smaller mountains surrounding the mainland were defined as islands. We found that the observed heterozygosity (H_o) was significantly higher, and the inbreeding coefficient (F_is) significantly lower, on the mainland compared to islands. There was a positive significant relationship between the unbiased expected heterozygosity (H_n.b.) and island size (log km²), but a negative significant relationship between H_o and the cost distance to the mainland. Our results are consistent with the equilibrium model of island biogeography and show that the model is well suited for investigating genetic diversity among islands, but also on islands located on the mainland.     

Conservation genetics of Boelen’s python (Morelia boeleni) from New Guinea: reduced genetic diversity and divergence of captive and wild animals

Boelen’s python (Morelia boeleni) is a montane New Guinea endemic found in highlands above 1000 m and below the tree line. The ecology, natural history, distribution, population size, and conservation status of this species are largely unknown. It has a protected status in Papua New Guinea but not in Indonesian Papua and several US and European zoos have active captive breeding programs that have been largely unsuccessful. To understand the degree of genetic diversity in wild and captive animals we undertook a genetic analysis of 90 M. boeleni for which we sequenced two mtDNA loci and one nuclear locus for a total of 1,418 bp of sequence data per individual. All 16 wild-caught M. boeleni from Indonesia and all captive M. boeleni are genetically uniform for all three loci. The single wild-caught animal from Papua New Guinea showed extremely low levels of genetic divergence and diversity from the Indonesian and captive samples. Data from two congeners, M. amethistina and M. viridis, suggests that M. boeleni have reduced genetic variation with a small effective population size possibly due to historical bottlenecks. These data demonstrate the need for further studies of genetic diversity of M. boeleni from across its range and raise particular concern for the limited genetic diversity of M. boeleni used captive breeding programs in zoological parks.     

Genetic evidence for the origins of range disjunctions in the Australian dry sclerophyll plant Hardenbergia violacea

Aim The distribution of genetic variation in the Australian dry sclerophyll plant Hardenbergia violacea (Fabaceae) is examined in the context of Pleistocene climate change in order to identify likely refugia. Particular consideration is given to the origin of range disjunctions in South Australia and Tasmania, and to determining whether the Tasmanian population is indigenous or recently introduced from mainland Australia. Location Southeastern Australian mainland and Tasmania. Methods A combination of chloroplast polymerase chain reaction–restriction fragment length polymorphism and genomic amplified fragment length polymorphism (AFLP) marker systems was used to examine the genetic structure of 292 individuals from 13 populations across the range of H. violacea in southeastern Australia. Results Hardenbergia violacea populations in Tasmania and southern Victoria were characterized by low, almost monotypic chloroplast diversity. New South Wales showed higher haplotype diversity and haplotype sharing among widely distributed populations. Principal coordinates analysis (PCoA) of the AFLP data found a strong latitudinal cline in AFLP variation from northern New South Wales south to Tasmania. The Tasmanian population formed an isolated and somewhat disjunct genetic cluster at one end of this cline. However, the South Australian population was an exception to the clinal variation shown by all other populations, forming a highly disjunct cluster in the PCoA. Within‐population genetic diversity was low in both disjunct populations. Main conclusions The genetic evidence indicates that the Tasmanian population is likely to be indigenous and probably the product of vicariance, which was followed by range contraction at the Last Glacial Maximum or an earlier glacial event. The deep phylogenetic disjunction in South Australia is evidence of a much earlier separation on mainland Australia. The chloroplast structure indicates that, during the Pleistocene, H. violacea underwent broad‐scale recolonization in southern Victoria and Tasmania, possibly from a large continental refugium in eastern New South Wales. We conclude that H. violacea, and presumably the sclerophyll communities in which it occurs, have undergone multiple range contractions to large continental refugia during different Pleistocene glaciations in southeastern Australia.     

Genetic diversity and population structure of Culex modestus across Europe: does recent appearance in the United Kingdom reveal a tendency for geographical spread?

In mainland Europe, the mosquito species Culex modestus Ficalbi (1890) is a bridge vector for West Nile virus (WNV) from its natural bird-mosquito cycle to mammals. The present study assessed the genetic diversity of Cx. modestus, as well as related Culex species, using the mitochondrial COI DNA barcoding region and compared this with the population structure across Europe. A haplotype network was mapped to determine genealogical relationships among specimens. The intraspecific genetic diversity within individual Culex species was below 2%, whereas the interspecific genetic divergence varied from 2.99% to 13.74%. In total, 76 haplotypes were identified among 198 sequences. A median-joining network determined from 198 COI sequences identified two major lineages that were separated by at least four mutation steps. A high level of intraspecific genetic diversity was not detected in Cx. modestus in samples submitted from different European populations, which indicates that morphologically identified specimens represent a single species and not a species complex. Therefore, it is deduced that different populations of Cx. modestus will show a similar potential to transmit WNV, lending support to concerns that the population present in southeast England represents a risk of transmission to humans.     

Dry-season Mosquito Breeding Associated with Irrigation in the Northeast Kimberley Region of Western Australia: Potential Impact on Mosquito-borne Disease Transmission

Recent Ross River virus activity prior to the onset of the wet season in the Kimberley region of Western Australia points to an increased dry season mosquito-borne disease risk at Kununurra, compared to other Kimberley towns. This study describes a preliminary investigation into the role of the Ord River Irrigation Area at Kununurra in mosquito production during the dry season. Specifically, the study sought to determine whether the irrigation area has provided the opportunity for year-round breeding of arbovirus vector mosquitoes. A 10-day adult and larval mosquito sampling survey, in August 2003, revealed that mosquito breeding was occurring during the driest month of the year at Kununurra, supporting the hypothesis that mosquitoes can breed year-round. Importantly, significantly larger numbers and percentage of adult Culex annulirostris, an important disease vector in Australia, were collected within the irrigation area (44.6% of total catch) compared to nearby reference (nonirrigated) breeding sites (9.8%) (mean difference 76.2 per trap per night; 95% CI 38.6, 113.7; P <0.001). Larval Cx. annulirostris were also collected at several sites within the irrigation area, whereas none were collected at reference sites. These results indicate that mosquito breeding associated with anthropogenic environmental changes may be responsible for an increased health risk at the end of the dry season. Mosquito management needs to be given a high priority to ensure this potential health risk is not further exacerbated. Several control strategies to reduce breeding of disease vector mosquitoes are identified, primarily focusing on modification of breeding habitats and alteration of irrigation protocols.     

Molecular phylogeny of Burkholderia pseudomallei from a remote region of Papua New Guinea

Background

The island of New Guinea is located midway between the world''s two major melioidosis endemic regions of Australia and Southeast Asia. Previous studies in Papua New Guinea have demonstrated autochthonous melioidosis in Balimo, Western province. In contrast to other regions of endemicity, isolates recovered from both environmental and clinical sources demonstrate narrow genetic diversity over large spatial and temporal scales.

Methodology/Principal Findings

We employed molecular typing techniques to determine the phylogenetic relationships of these isolates to each other and to others worldwide to aid in understanding the origins of the Papua New Guinean isolates. Multi-locus sequence typing of the 39 isolates resolved three unique sequence types. Phylogenetic reconstruction and Structure analysis determined that all isolates were genetically closer to those from Australia than those from Southeast Asia. Gene cluster analysis however, identified a Yersinia-like fimbrial gene cluster predominantly found among Burkholderia pseudomallei derived from Southeast Asia. Higher resolution VNTR typing and phylogenetic reconstruction of the Balimo isolates resolved 24 genotypes with long branch lengths. These findings are congruent with long term persistence in the region and a high level of environmental stability.

Conclusions/Significance

Given that anthropogenic influence has been hypothesized as a mechanism for the dispersal of B. pseudomallei, these findings correlate with limited movement of the indigenous people in the region. The palaeogeographical and anthropogenic history of Australasia and the results from this study indicate that New Guinea is an important region for the further study of B. pseudomallei origins and dissemination.     

New Insights into the Population Structure of Anopheles gambiae s.s. in the Gulf of Guinea Islands Revealed by Herves Transposable Elements

Transposable elements (TEs) are mobile portions of DNA that are able to replicate and spread in the genome of many organisms. TEs can be used as a means to insert transgenes in insects, being stably inherited throughout generations. Anopheles gambiae is the main vector of human malaria in Sub-Saharan Africa. Given the extraordinary burden this disease imposes, the mosquito became a choice target for genetic control approaches with the purpose of reducing malaria transmission. In this study, we investigated the abundance and distribution of Herves TE in An. gambiae s.s. from Cameroon and four islands in the Gulf of Guinea, in order to determine their genetic structure. We have detected a population subdivision between Equatorial Guinea islands and the islands of São Tomé, Príncipe and mainland. This partitioning associates more with political rather than geographic boundaries, possibly reflecting different mainland source populations colonizing the islands.