Multiple Founder Events Research Articles

Population genetic and phylogeographic relationships of the dengue vector <em>Aedes aegypti</em> inferred from three localities of Sri Lanka

Aedes aegypti is the major epidemic vector of dengue outbreaks in the world. Here, a 359 bp region of the mitochondrial NADH hydrogenase subunit 4 (ND4) gene was taken as the candidate locus for study purposes from 27 Aedes aegypti mosquito larvae collected from an inland mountain area (Badulla), and two distantly located coastal areas (Hambantota and Trincomalee) in Sri Lanka. Eleven parsimonious informative sites were observed, and 7 haplotypes were identified from the 27 samples. The coastal populations displayed higher genetic diversity compared to those from the inland mountain-contained region. The phylogenetic analyses of Ae. aegypti from Badulla, Hambantota and Trincomalee including published Sri Lankan and global samples (a total of 106 sequences) revealed two separate clades: a basal clade and a derived clade. Eighty percent of the samples found from Badulla, were of the basal haplotype (H1) clustered together with West African (Senegal) samples in close proximity to the outgroup. On the contrary, Hambantota and Trincomalee samples were found dispersed in both the basal and derived clades clustering closely with samples from Mexico, Brazil, Kenya, USA and Myanmar indicating their close genetic relationship to many New World, East African and Asian varieties. Taken together, our study proposes that colonization of Ae aegypti in Sri Lanka occurred through multiple founder events (ancient multiple invasions in coastal sites and a more recent West African invasion to the inland mountain-demarcated Badulla region) and continue to exist in mutational drift equilibrium following the settlement event.

An island-hopping bird reveals how founder events shape genome-wide divergence.

When populations colonize new areas, both strong selection and strong drift can be experienced due to novel environments and small founding populations, respectively. Empirical studies have predominantly focused on the phenotype when assessing the role of selection, and limited neutral-loci when assessing founder-induced loss of diversity. Consequently, the extent to which processes interact to influence evolutionary trajectories is difficult to assess. Genomic-level approaches provide the opportunity to simultaneously consider these processes. Here, we examine the roles of selection and drift in shaping genomic diversity and divergence in historically documented sequential island colonizations by the silvereye (Zosterops lateralis). We provide the first empirical demonstration of the rapid appearance of highly diverged genomic regions following population founding, the position of which are highly idiosyncratic. As these regions rarely contained loci putatively under selection, it is most likely that these differences arise via the stochastic nature of the founding process. However, selection is required to explain rapid evolution of larger body size in insular silvereyes. Reconciling our genomic data with these phenotypic patterns suggests there may be many genomic routes to the island phenotype, which vary across populations. Finally, we show that accelerated divergence associated with multiple founding steps is the product of genome-wide rather than localized differences, and that diversity erodes due to loss of rare alleles. However, even multiple founder events do not result in divergence and diversity levels seen in evolutionary older subspecies, and therefore do not provide a shortcut to speciation as proposed by founder-effect speciation models.

The Counteracting Effects of Demography on Functional Genomic Variation: The Roma Paradigm.

Demographic history plays a major role in shaping the distribution of genomic variation. Yet the interaction between different demographic forces and their effects in the genomes is not fully resolved in human populations. Here, we focus on the Roma population, the largest transnational ethnic minority in Europe. They have a South Asian origin and their demographic history is characterized by recent dispersals, multiple founder events, and extensive gene flow from non-Roma groups. Through the analyses of new high-coverage whole exome sequences and genome-wide array data for 89 Iberian Roma individuals together with forward simulations, we show that founder effects have reduced their genetic diversity and proportion of rare variants, gene flow has counteracted the increase in mutational load, runs of homozygosity show ancestry-specific patterns of accumulation of deleterious homozygotes, and selection signals primarily derive from preadmixture adaptation in the Roma population sources. The present study shows how two demographic forces, bottlenecks and admixture, act in opposite directions and have long-term balancing effects on the Roma genomes. Understanding how demography and gene flow shape the genome of an admixed population provides an opportunity to elucidate how genomic variation is modeled in human populations.

Low genetic differentiation despite high fragmentation in the endemic serpentinophyte Minuartia smejkalii (M. verna agg., Caryophyllaceae) revealed by RADSeq SNP markers

Minuartia smejkalii is an obligate serpentinophyte plant endemic to the Czech Republic. Since the 1960s, the species’ habitat has undergone strong human-mediated fragmentation, resulting in extinction of some populations and dramatic size reduction of the remaining populations. Thus, contrary to the typically stable serpentine habitats, M. smejkalii habitats underwent a recent and severe decline, which can exacerbate the effects of fragmentation on population genetic structure. We examined the genetic structure of all known M. smejkalii populations and two populations of M. corcontica and M. caespitosa, which are closely related, using RADSeq. The results indicate low, but clear differentiation among the three species, thus supporting the status of M. smejkalii as an independent taxon, though more extensive analysis of the whole group is needed. We further show high genetic diversity within M. smejkalii populations, low to moderate among-populations differentiation, and moderate regional differentiation. This could be due to the outcrossing mating system of M. smejkalii promoting high levels of gene flow and historical factors (multiple founder events, a recent bottleneck and/or a genetic time lag). We finally demonstrate that 2–3% of the markers show differentiation patterns consistent with divergent selection, suggesting that some local adaptation might have occured in M. smejkalii. Based on our observations, but without any experimental testing for local adaptation, if a conservation action is to be carried out, we recommend strictly separating the material from the two regions, and if possible, separating the populations within a region.

Multiple founder events explain the genetic diversity and structure of the model allopolyploid grass Brachypodium hybridum in the Iberian Peninsula hotspot.

It is accepted that contemporary allopolyploid species have originated recurrently, but very few cases have been documented using multiple natural formations of the same species. To extend our knowledge, we have investigated the multiple origins, genetic variation and structure of the allotetraploid grass Brachypodium hybridum with respect to its progenitor diploid species B. distachyon (D genome) and B. stacei (S genome). For this, our primary focus is the Iberian Peninsula, an evolutionary hotspot for the genus Brachypodium. We analysed 342 B. hybridum individuals from 36 populations using ten nuclear SSR loci and two plastid loci. The B. hybridum genetic profiles were compared with those previously reported for B. stacei and B. distachyon. In addition, phylogenetic analysis of the plastid data was performed for a reduced subset of individuals. The nuclear simple sequence repeat (SSR) genetic analysis detected medium to high genetic diversity, with a strong south-to-north genetic structure cline, and a high selfing rate in B. hybridum. Comparative genetic analysis showed a close relatedness of current B. hybridum D allelic profiles with those of B. distachyon, but a lack of similarity with those of B. stacei, suggesting another B. stacei source for the B. hybridum S alleles. Plastid analysis detected three different bidirectional allopolyploidization events: two involved distinct B. distachyon-like ancestors and one involved a B. stacei-like ancestor. The south-eastern Iberian Peninsula B. hybridum populations were more genetically diverse and could have originated from at least two hybridization events whereas north-eastern/north-western Iberian Peninsula B. hybridum populations were less diverse and may have derived from at least one hybridization event. The genetic and evolutionary evidence supports the plausible in situ origin of the south-eastern and northern Iberian Peninsula B. hybridum allopolyploids from their respective local B. distachyon and unknown B. stacei ancestors. The untapped multiple origins and genetic variation detected in these B. hybridum populations opens the way to future evolutionary analysis of allopolyploid formation and genomic dominance and expression in the B. hybridum-B. distachyon-B. stacei grass model complex.

M77 - COMPLEX SPATIO-TEMPORAL DISTRIBUTION AND GENOGEOGRAPHIC AFFINITY OF MITOCHONDRIAL DNA HAPLOGROUPS IN 24,216 DANES

Background Mitochondrial DNA (mtDNA) haplogroups (hgs) are evolutionarily conserved sets of mtDNA SNPs. Associations of hgs with geographical origin, disease and physiological characteristics have been reported, but have frequently not been reproducible. We assessed, using 418 mtDNA SNPs on the PsychChip (Illumina), the spatio-temporal distribution of mtDNA hgs in DNA isolated from geographically un-biased dried blood spots (DBS), collected from 1981 to 2005 through the Danish National Neonatal Screening program. Methods As part of the iPSYCH (www.iPSYCH.au.dk) recruitment protocol, 24,651 singletons (47.1% female), born between May 1 1981 and Dec 31 2005 were selected at random from the Danish Central Person Registry. The Samples were extracted from the Danish National Biobank and Genotyped at the Broad Institute as part of iPSYCH and PGC. Haplotyping of mtDNA was performed using the defining SNPs reported in www.phylotree.org. First the affiliation to macro-hg i.e. L0 – L6, M, N, R, and then to hgs – units more distal in the cladogram was established. Ancestry estimation was done using ADMIXTURE 1.3.052. Briefly, a reference population consisting of Human Genome Diversity Project (HGDP) ( http://www.hagsc.org/hgdp/ ) genotyping SNP data set, supplemented with representative samples of danes (716 individuals) and greenlanders (592 individuals) available at SSI, was used. Results The hg distribution was typically Northern European, and hgs were highly variable based on median-joining analysis, suggesting multiple founder events. Considerable heterogeneity and variation in autosomal geno-geographic affinity (ancestry) was observed. Thus, individuals with hg H exhibited 95%, and U hgs 38.2% - 92.5%, Danish ancestry. Significant clines between geographical regions and rural and metropolitan populations were found. Over 25 years, macro-hg L increased from 0.2% to 1.2% (p=1.1⁎E-10), and M from 1% to 2.4% (p=3.7⁎E-8). Hg U increased among the R macro-hg from 14.1% to 16.5% (p=1.9⁎E-3). Geno-geographic affinity, geographical skewedness, and sub-hg distribution suggested that the L, M and U increases are due to immigration. Discussion This study shows that the distribution of mtDNA hgs in Denmark is highly dynamic and complex. It comprises 1.6% of the Danish population over a 25-year period, and is by far the largest ever performed of the distribution of mtDNA hgs in any country. The method of collecting stored DBS from the PKU biobank, where the coverage is ~99%, enabled us to survey a true population based sample, where the time and place of birth was known from national electronic registries. This is in contrast to most other population genetic studies of adults sampled in a specific bias-prone context, e.g. hospitalized patients or localized samplings. The mtDNA haplotyping was based on array data with only 418 mtDNA SNPs and as a consequence not all sub-hgs could be called. The stringent adherence to specific SNPs left a number of persons, normally in the order of 1–2%, not haplotypable. An advantage of only using a limited set of markers is that confounding due to private variants is avoided.

Complex spatio-temporal distribution and genomic ancestry of mitochondrial DNA haplogroups in 24,216 Danes.

Mitochondrial DNA (mtDNA) haplogroups (hgs) are evolutionarily conserved sets of mtDNA SNP-haplotypes with characteristic geographical distribution. Associations of hgs with disease and physiological characteristics have been reported, but have frequently not been reproducible. Using 418 mtDNA SNPs on the PsychChip (Illumina), we assessed the spatio-temporal distribution of mtDNA hgs in Denmark from DNA isolated from 24,642 geographically un-biased dried blood spots (DBS), collected from 1981 to 2005 through the Danish National Neonatal Screening program. ADMIXTURE was used to establish the genomic ancestry of all samples using a reference of 100K+ autosomal SNPs in 2,248 individuals from nine populations. Median-joining analysis determined that the hgs were highly variable, despite being typically Northern European in origin, suggesting multiple founder events. Furthermore, considerable heterogeneity and variation in nuclear genomic ancestry was observed. Thus, individuals with hg H exhibited 95%, and U hgs 38.2% - 92.5%, Danish ancestry. Significant clines between geographical regions and rural and metropolitan populations were found. Over 25 years, macro-hg L increased from 0.2% to 1.2% (p = 1.1*E-10), and M from 1% to 2.4% (p = 3.7*E-8). Hg U increased among the R macro-hg from 14.1% to 16.5% (p = 1.9*E-3). Genomic ancestry, geographical skewedness, and sub-hg distribution suggested that the L, M and U increases are due to immigration. The complex spatio-temporal dynamics and genomic ancestry of mtDNA in the Danish population reflect repeated migratory events and, in later years, net immigration. Such complexity may explain the often contradictory and population-specific reports of mito-genomic association with disease.

Analysis of genetic diversity and population structure of the endangered Origanum compactum from Morocco, using SSR markers: Implication for conservation

Origanum compactum L. (Lamiaceae) is one of the most popular medicinal and culinary herbs with a long history of traditional use. This species, endemic to Morocco and Southern Spain, is considered as threatened, due mainly to over-harvesting and habitat destruction. Assessment of genetic polymorphism of threatened species constitutes a prerequisite for germplasm conservation. To support and monitor future conservation and management measures, we used 15 microsatellite loci to genotype 670 individuals of O. compactum distributed over 59 localities covering the entire geographic distribution of the species in Morocco. In contrast to what was argued in most previous studies, the genetic diversity of the Moroccan oregano was greater mostly on isolated and declined populations compared to less fragmented accessions. Accordingly, our study identifies the most relevant management units for conservation purposes on the basis of their genetic diversity level. Factors such as the perennity of the species, somatic mutations, or multiple founder events could explain this unexpected level of genetic diversity at the species level in this endangered plant. The population genetic results reflect a strong genetic structure and identify three major clusters among O. compactum populations: a Northern cluster composed of populations near the strait of Gibraltar (Tangier-Tetouan and Chefchaouen regions as well as some adjacent populations from Taounate), and two other clusters with significant mixed lineage. This result supports the hypothesis that the strait of Gibraltar could be a migration route for plants between Southern Spain and Northern Morocco. The high differentiation of populations (Fst=0.22) and the low rate of gene flow between them (Nm=0.88) might been caused by recent isolation of the populations through habitat disturbances. Historical demographic patterns inferred with a generalized stepwise mutation model (SSM) and two-phase model (TPM) using maximum likelihood showed significant past demographic changes in 39 of the 59 sampled populations. Implications of these results to develop appropriate conservation strategies of this endangered species are proposed.

A three-dimensional geometric morphometrics view of the cranial shape variation and population history in the New World.

Craniofacial variation in past and present Amerindians has been attributed to the effect of multiple founder events, or to one major migration followed by in situ differentiation and possibly recurrent contacts among Circum-Arctic groups. Our study aims to: (i) detect morphological differences that may indicate several migrations; (ii) test for the presence of genetic isolation; and (iii) test the correlation between shape data and competing settlement hypotheses by taking into account geography, chronology, climate effects, the presence of genetic isolation and recurrent gene flow. We analyzed a large sample of three-dimensional (3D) cranial surface scans (803 specimens) including past and modern groups from America and Australasia. Shape variation was investigated using geometric morphometrics. Differential external gene flow was evaluated by applying genetic concepts to morphometric data (Relethford-Blangero approach). Settlement hypotheses were tested using a matrix correlation approach (Mantel tests). Our results highlight the strong dichotomy between Circum-Arctic and continental Amerindians as well as the impact of climate adaptation, and possibly recurrent gene flow in the Circum-Arctic area. There is also evidence for the impact of genetic isolation on phenetic variation in Baja California. Several settlement hypotheses are correlated with our data. The three approaches used in this study highlight the importance of local processes especially in Baja California, and caution against the use of overly simplistic models when searching for the number of migration events. The results stress the complexity of the settlement of the Americas as well as the mosaic nature of the processes involved in this process. Am. J. Hum. Biol. 28:646-661, 2016. © 2016 Wiley Periodicals, Inc.

Evidence of rapid change in genetic structure and diversity during range expansion in a recovering large terrestrial carnivore.

Recovery of natural populations occurs often with simultaneous or subsequent range expansions. According to population genetic theory, genetic structuring emerges at the expansion front together with decreasing genetic diversity, owing to multiple founder events. Thereupon, as the expansion proceeds and connectivity among populations is established, homogenization and a resurgence of genetic diversity are to be expected. Few studies have used a fine temporal scale combined with genetic sampling to track range expansions as they proceed in wild animal populations. As a natural experiment, the historical eradication of large terrestrial carnivores followed by their recovery and recolonization may facilitate empirical tests of these ideas. Here, using brown bear (Ursus arctos) as model species, we tested predictions from genetic theory of range expansion. Individuals from all over Finland were genotyped for every year between 1996 and 2010 using 12 validated autosomal microsatellite markers. A latitudinal shift of about 110 km was observed in the distribution and delineation of genetic clusters during this period. As the range expansion proceeded, we found, as theory predicts, that the degree of genetic structure decreased, and that both genetic variation and admixture increased. The genetic consequences of range expansions may first be detected after multiple generations, but we found major changes in genetic composition after just 1.5 generations, accompanied by population growth and increased migration. These rapid genetic changes suggest an ongoing concerted action of geographical and demographic expansion combined with substantial immigration of bears from Russia during the recovery of brown bears within the large ecosystem of northern Europe.

Rethinking refugia: Tree topology, divergence dates, and demographic history trace the distribution of the endangered Plymouth gentian (Sabatia kennedyana) from the Pleistocene glaciation to present day.

Molecular population genetics is a powerful tool to infer how species responded to past environmental change. In the northern hemisphere, interest is increasing in how species responded to changes in ice coverage and temperature during the last glaciation maximum (LGM, between 18000-21000 yr ago) with a common assumption that glacial refugia were located at the southern edge of a species range. We reconstructed the glacial and postglacial phylogeography of Sabatia kennedyana, a member of the Atlantic Coastal Plains Flora with a current distribution from Nova Scotia (NS) to South Carolina, using both cpDNA and nuclear markers. We also examined clinal variation in morphological traits, in particular relative investment in asexual vs sexual growth. We find strong evidence that the species did not reside in southern glacial refugia, but rather in primary glacial refugia off the exposed continental shelf extending from Cape Cod and that this area was responsible for the founding of modern populations across the range from Nova Scotia (NS) to the United States. Additionally, based on the finding of higher cpDNA diversity and older cpDNA lineages in NS, we propose that multiple founder events occurred in NS, while only a single lineage gave rise to current populations in the United States. By understanding how S. kennedyana responded to past shifts in climate and by identifying areas of high genetic diversity in the northern range edge, we discuss the potential response of the species to future climate change scenarios.

Strong genetic differentiation due to multiple founder events during a recent range expansion of an introduced wall lizard population

Biological invasions represent ideal sys- tems for the study of evolutionary processes associated with colonization events. It has been hypothesized that the genetic diversity is generally decreasing from the centre of the range to the margins due to multiple founder events. Invasive populations offer the oppor- tunity to test this hypothesis at a fine spatial and temporal scale. We analysed the genetic structure of a large expanding non-native population of the Com- mon Wall Lizard (Podarcis muralis) in Passau (Ger- many) using thirteen microsatellite loci. We analyzed the genetic structure and levels of admixture across a transect reflecting the expansion process and tested for a loss of genetic diversity and an increase of genetic differentiation from the centre to the invasion front. Our results demonstrate that significant genetic pop- ulation structure can emerge rapidly at a small spatial scale. We found a trend for an increase in genetic differentiation and a decrease in genetic diversity from the invasion centre to the expanding range margin, suggesting that genetic drift is the major factor causing this pattern. The correlation between genetic diversity and average genetic differentiation was significant among sites. We hypothesize that the territoriality of P. muralis generates sufficient rates of noncontiguous and stratified dispersal from longer established sites to maintain significant genetic diversity at the invasion front. Simultaneously, territoriality might restrict the colonization success of migrants at established sites, so that in combination with founder events a strong differentiation arises.

Low genetic and morphological differentiation between an introduced population of dunnocks in New Zealand and an ancestral population in England

Species invasions and exotic species introductions can be considered as ‘unplanned experiments’, which help us to understand the evolution of organisms. In this study, we investigated whether an exotic bird species, the dunnock (Prunella modularis), has diverged genetically and morphologically from its native source population (Cambridge, England) after introduction into a new environment (Dunedin, South Island of New Zealand; exotic population). We used a set of microsatellite markers and three morphological traits to quantify the divergence between these two populations. We quantified neutral genotypic differentiation between the populations, and also used an individual-based Bayesian clustering method to assess genetic structure. We compared morphological divergence using univariate and principal components analyses. We found that individuals from the Dunedin population are genetically distinct from the Cambridge population, but levels of differentiation are very low. Overall within-population levels of genetic diversity are low compared to other bird species, and effective population sizes are small; indicating that the native population probably has a historically low level of genetic diversity, and that the introduced population retained most of that diversity after its introduction into New Zealand. We found little evidence of morphological divergence, and the evolutionary rate of change in these traits is below the average for other taxa. Our study adds support to the growing literature showing that invasive species maintain most of their initial genetic diversity after multiple founder events, even when population size is severely reduced. Moreover, our morphological data indicate slow evolutionary rates in species introduced to similar habitats.

Strong gene flow and lack of stable population structure in the face of rapid adaptation to local temperature in a spring-spawning salmonid, the European grayling (Thymallus thymallus).

Gene flow has the potential to both constrain and facilitate adaptation to local environmental conditions. The early stages of population divergence can be unstable because of fluctuating levels of gene flow. Investigating temporal variation in gene flow during the initial stages of population divergence can therefore provide insights to the role of gene flow in adaptive evolution. Since the recent colonization of Lake Lesjaskogsvatnet in Norway by European grayling (Thymallus thymallus), local populations have been established in over 20 tributaries. Multiple founder events appear to have resulted in reduced neutral variation. Nevertheless, there is evidence for local adaptation in early life-history traits to different temperature regimes. In this study, microsatellite data from almost a decade of sampling were assessed to infer population structuring and its temporal stability. Several alternative analyses indicated that spatial variation explained 2-3 times more of the divergence in the system than temporal variation. Over all samples and years, there was a significant correlation between genetic and geographic distance. However, decomposed pairwise regression analysis revealed differing patterns of genetic structure among local populations and indicated that migration outweighs genetic drift in the majority of populations. In addition, isolation by distance was observable in only three of the six years, and signals of population bottlenecks were observed in the majority of samples. Combined, the results suggest that habitat-specific adaptation in this system has preceded the development of consistent population substructuring in the face of high levels of gene flow from divergent environments.

Population genetic differences along a latitudinal cline between original and recently colonized habitat in a butterfly.

BackgroundPast and current range or spatial expansions have important consequences on population genetic structure. Habitat-use expansion, i.e. changing habitat associations, may also influence genetic population parameters, but has been less studied. Here we examined the genetic population structure of a Palaeartic woodland butterfly Pararge aegeria (Nymphalidae) which has recently colonized agricultural landscapes in NW-Europe. Butterflies from woodland and agricultural landscapes differ in several phenotypic traits (including morphology, behavior and life history). We investigated whether phenotypic divergence is accompanied by genetic divergence between populations of different landscapes along a 700 km latitudinal gradient.Methodology/Principal FindingsPopulations (23) along the latitudinal gradient in both landscape types were analyzed using microsatellite and allozyme markers. A general decrease in genetic diversity with latitude was detected, likely due to post-glacial colonization effects. Contrary to expectations, agricultural landscapes were not less diverse and no significant bottlenecks were detected. Nonetheless, a genetic signature of recent colonization is reflected in the absence of clinal genetic differentiation within the agricultural landscape, significantly lower gene flow between agricultural populations (3.494) than between woodland populations (4.183), and significantly higher genetic differentiation between agricultural (0.050) than woodland (0.034) pairwise comparisons, likely due to multiple founder events. Globally, the genetic data suggest multiple long distance dispersal/colonization events and subsequent high intra- and inter-landscape gene flow in this species. Phosphoglucomutase deviated from other enzymes and microsatellite markers, and hence may be under selection along the latitudinal gradient but not between landscape types. Phenotypic divergence was greater than genetic divergence, indicating directional selection on some flight morphology traits.Main Conclusions/SignificanceClinal differentiation characterizes the population structure within the original woodland habitat. Genetic signatures of recent habitat expansion remain, notwithstanding high gene flow. After differentiation through drift was excluded, both latitude and landscape were significant factors inducing spatially variable phenotypic variation.

A geographic mosaic of passive dispersal: population structure in the endemic Hawaiian amber snail Succinea caduca (Mighels, 1845)

We used 276 cytochrome c oxidase subunit I (COI, 645 bp) and a subset of 84 16S large ribosomal subunit (16S, 451 bp) sequences to evaluate geographic patterns of genetic variation in 24 populations of the endemic Hawaiian land snail Succinea caduca spanning its range on six islands. Haplotype networks, gene tree topologies, pairwise molecular divergence and F(ST) matrices suggest substantial geographic genetic structuring and complex dispersal patterns. Low nucleotide diversity and low pairwise molecular divergence values within populations coupled with higher between population values suggest multiple founder events. High overall haplotype diversity suggests diversification involving rare interpopulation dispersal, fragmentation by historical lava flows and variation in habitat structure. Within-island rather than between-island population comparisons accounted for the majority of molecular variance. Although 98% of 153 COI haplotypes were private by population, a Mantel test showed no evidence for isolation by distance. Mismatch distributions and population partitioning patterns suggest that genetic fragmentation has been driven by punctuated, passive dispersal of groups of closely related haplotypes that subsequently expanded and persisted in isolation for long periods (average > 2 million years ago), and that Pleistocene island connections may have been important in enhancing gene flow. Historical availability of mesic coastal habitat, together with effective dispersal may explain the long-term persistence and unusual multi-island distribution of this species, contrasting with the single-island endemism of much of the Hawaiian biota.

Molecular systematics and phylogeography in the fairy shrimp Tanymastix stagnalis based on mitochondrial DNA

AbstractPatterns of sequence divergence in about 1 kb of mitochondrial DNA coding for two genes (16s rRNA and cytochrome oxidase I, COI) were analysed in 13 populations of the fairy shrimp Tanymastix stagnalis from Norway, Germany, France, Italy (northern and central Italy plus insular populations from Sardinia and the Tuscan Archipelago) and Spain, and in one presumed population of Tanymastix stellae from Corsica. The latter species was originally known only from a single locality in Sardinia, which has been destroyed by urbanization; the Corsican population was referred to T. stellae by some French authors on the basis of the collection of several cysts from mud. mtDNA data revealed a very low level of genetic divergence between the presumed population of T. stellae and the other T. stagnalis populations included in the study. Our genetic findings do not support the presence of T. stellae in Corsica and are in line with previous SEM studies revealing that all species belonging to the genus Tanymastix produce cysts with identical morphology. The results indicate complex phylogeographic relationships and pronounced genetic differentiation among T. stagnalis populations. The islands of Corsica and Sardinia on the one hand and the island of Capraia (Tuscan Archipelago) on the other were probably colonized independently at different times. Genetic relationships among continental populations do not follow a clear geographical trend, indicating that geographical distance is not the main force shaping the pattern of genetic structuring of the species. Stochastic factors such as multiple and independent founder events probably contributed to the striking pattern of genetic differentiation along with subsequent local adaptation. These results agree with previously published molecular work on several groups of aquatic organisms and further support the high potential for dispersal–low gene flow paradox shown by a large array of animals living in lentic habitats.

Genetic differentiation of three species of Matthiola (Brassicaceae) in the Sicilian insular system

We examined the genetic variation of 12 isozyme loci in 14 populations of Matthiola (Brassi- caceae) representing the geographic distribution of the species M. incana, M. fruticulosa ssp. fruticulosa and M. tricuspidata in the Sicilian insular system and the adjacent mainland areas to estimate the levels and distribution of genetic variation in the insular populations and to understand their popu- lation dynamics. The disparity in the distribution of polymorphism in populations of M. incana ssp. incana (low within populations but with high values of FST and GST) contrasts with the homogeneity in the inter-population distribution of the high genetic variation detected in M. tricuspidata and M. fruticulosa ssp. fruticulosa. While the low polymorphism found in M. incana ssp. incana is consistent with its origin through cultivation and the associated lack of gene flow, the Sicilian populations of the other two taxa probably derived from multiple founder events from nearby conti- nental areas and, according to our estimates, have maintained high interpopulational gene flow. Unlike M. incana, the Sicilian populations of M. tricuspidata and M. fruticulosa ssp. fruticulosa could have survived the glaciations in refugia. This higher antiquity, together with the maintenance of abundant gene flow, largely explains their high values of genetic variation. In contrast, M. incana ssp. pulchella and M. incana ssp. rupestris have low indices of polymorphism and they are probably neo-endemics, as their distribution areas were severely affected by the Plio-Pleistocene glaciations.

Evidence for multiple genetic forms with similar eyeless phenotypes in the blind cavefish, Astyanax mexicanus.

A diverse group of animals has adapted to caves and lost their eyes and pigmentation, but little is known about how these animals and their striking phenotypes have evolved. The teleost Astyanax mexicanus consists of an eyed epigean form (surface fish) and at least 29 different populations of eyeless hypogean forms (cavefish). Current alternative hypotheses suggest that adaptation to cave environments may have occurred either once or multiple times during the evolutionary history of this species. If the latter is true, the unique phenotypes of different cave-dwelling populations may result from convergence of form, and different genetic changes and developmental processes may have similar morphological consequences. Here we report an analysis of variation in the mitochondrial NADH dehydrogenase 2 (ND2) gene among different surface fish and cavefish populations. The results identify a minimum of two genetically distinctive cavefish lineages with similar eyeless phenotypes. The distinction between these divergent forms is supported by differences in the number of rib-bearing thoracic vertebrae in their axial skeletons. The geographic distribution of ND2 haplotypes is consistent with roles for multiple founder events and introgressive hybridization in the evolution of cave-related phenotypes. The existence of multiple genetic lineages makes A. mexicanus an excellent model to study convergence and the genes and developmental pathways involved in the evolution of the eye and pigment degeneration.

The Genetic Basis of Photoperiodism and Its Evolutionary Divergence Among Populations of the Pitcher-Plant Mosquito, Wyeomyia smithii

We measured the additive genetic variance within populations and the composite additive, dominance, and epistatic effects contributing to differentiation of photoperiodic response between two southern (ancestral) and each of four progressively more northern (derived) populations of the pitcher-plant mosquito, Wyeomyia smithii. Critical photoperiod and its additive genetic variance but not its heritability increased with latitude. Directional selection on critical photoperiod during the northward divergence of W. smithii has therefore not eroded the additive genetic variance underlying this trait. Joint scaling tests of crosses between populations showed that epistatic effects, especially additive x additive and dominance x dominance interactions, overwhelm composite additive and dominance effects on critical photoperiod. The presence of substantial epistasis suggests that multiple founder events during the northward divergence of W. smithii may have been responsible for the release of progressively greater additive genetic variance in derived populations, despite directional and stabilizing selection to reduce it. If epistasis makes a similar contribution to the genetic differentiation of populations in other species, then current models of adaptive evolution that consider only additive genetic variation and covariation within populations may be of limited value in predicting how natural populations differentiate in life history.