Marine biogeographical structure in two highly dispersive gastropods: implications for trans-Tasman dispersal

Aim  Recent genetic and ecological studies of marine invertebrate species with planktotrophic larvae have inferred high rates of gene flow across wide oceanic barriers. We therefore aim to test for the genetic signature of long-distance dispersal in two widespread and abundant marine gastropod taxa.
Location  The intertidal and shallow subtidal zones of southern Australia and New Zealand (NZ), which house similar marine invertebrate assemblages despite being separated by the 2000-km-wide Tasman Sea.
Methods  We used mtDNA cytochrome oxidase I gene sequence analysis of two gastropod genera exhibiting trans-Tasman distributions, namely Austrolittorina (Littorinidae) (139 specimens; 28 localities) and Scutus (Fissurellidae) (154 specimens; 32 localities). The cool-temperate Australian ( A. unifasciata ; S. antipodes ) and NZ ( A. antipodum ; S. breviculus ) taxa within each genus are morphologically similar but of uncertain taxonomic status.
Results  The mtDNA analyses indicate major trans-Tasman genetic discontinuities for both gastropod genera, with no evidence of recent or ongoing intercontinental gene flow. Although both Scutus and Austrolittorina show significant east–west structure within southern Australia – consistent with recent studies of regional marine phylogeography – neither taxon exhibits significant differentiation within NZ.
Main conclusions  Morphologically conserved but biogeographically disjunct gastropod populations may exhibit striking phylogeographic discontinuities, even when dispersal abilities appear to be high. On the basis of these data we reject recent calls for the synonymy of NZ and Australian lineages.     

The Influence of Interspecific Competition and Host Preference on the Phylogeography of Two African Ixodid Tick Species

A comparative phylogeographic study on two economically important African tick species, Amblyomma hebraeum and Hyalomma rufipes was performed to test the influence of host specificity and host movement on dispersion. Pairwise AMOVA analyses of 277 mtDNA COI sequences supported significant population differentiation among the majority of sampling sites. The geographic mitochondrial structure was not supported by nuclear ITS-2 sequencing, probably attributed to a recent divergence. The three-host generalist, A. hebraeum, showed less mtDNA geographic structure, and a lower level of genetic diversity, while the more host-specific H. rufipes displayed higher levels of population differentiation and two distinct mtDNA assemblages (one predominantly confined to South Africa/Namibia and the other to Mozambique and East Africa). A zone of overlap is present in southern Mozambique. A mechanistic climate model suggests that climate alone cannot be responsible for the disruption in female gene flow. Our findings furthermore suggest that female gene dispersal of ticks is more dependent on the presence of juvenile hosts in the environment than on the ability of adult hosts to disperse across the landscape. Documented interspecific competition between the juvenile stages of H. rufipes and H. truncatum is implicated as a contributing factor towards disrupting gene flow between the two southern African H. rufipes genetic assemblages.     

Significant population structure and asymmetric gene flow patterns amidst expanding populations of Clinus cottoides (Perciformes,Clinidae): application of molecular data to marine conservation planning in South Africa

Clinus cottoides is a fish endemic to the coast of South Africa, predominantly inhabiting rock pools. All South African clinids are viviparous, but probably breed throughout the year; as such, their dispersal may be limited, unlike species with pelagic larval stages. We analysed 343 fish from 14 localities on the west, south and east coasts using two mitochondrial genes and the second intron of the S7 ribosomal gene. Mitochondrial DNA analyses recovered significant genetic differentiation between fish populations from the east coast and other sampling locations, with a second break found between Gansbaai and Cape Agulhas on the south coast. Nuclear DNA recovered shallower, but significant, levels of population structure. Coalescent analyses suggested remarkably asymmetrical gene flow between sampling locations, suggesting that the cold Atlantic Benguela Current and Indian Ocean Agulhas counter‐current play important roles in facilitating dispersal. There was no gene flow between the east coast and the other sites, suggesting that these populations are effectively isolated. Divergence times between them were estimated to at least 68 000 years. Neutrality tests and mismatch distributions suggest recent population expansions, with the exception of peripheral western and eastern populations (possibly a consequence of environmental extremes at the edge of the species distribution). Analyses of the current South African marine protected areas network show that it is not connected and that De Hoop, one of South Africa's largest marine reserves, appears to be an important source population of recruits to both the south and southwest coasts.     

Contemporary population structure and post‐glacial genetic demography in a migratory marine species,the blacknose shark,Carcharhinus acronotus

Patterns of population structure and historical genetic demography of blacknose sharks in the western North Atlantic Ocean were assessed using variation in nuclear‐encoded microsatellites and sequences of mitochondrial (mt)DNA. Significant heterogeneity and/or inferred barriers to gene flow, based on microsatellites and/or mtDNA, revealed the occurrence of five genetic populations localized to five geographic regions: the southeastern U.S Atlantic coast, the eastern Gulf of Mexico, the western Gulf of Mexico, Bay of Campeche in the southern Gulf of Mexico and the Bahamas. Pairwise estimates of genetic divergence between sharks in the Bahamas and those in all other localities were more than an order of magnitude higher than between pairwise comparisons involving the other localities. Demographic modelling indicated that sharks in all five regions diverged after the last glacial maximum and, except for the Bahamas, experienced post‐glacial, population expansion. The patterns of genetic variation also suggest that the southern Gulf of Mexico may have served as a glacial refuge and source for the expansion. Results of the study demonstrate that barriers to gene flow and historical genetic demography contributed to contemporary patterns of population structure in a coastal migratory species living in an otherwise continuous marine habitat. The results also indicate that for many marine species, failure to properly characterize barriers in terms of levels of contemporary gene flow could in part be due to inferences based solely on equilibrium assumptions. This could lead to erroneous conclusions regarding levels of connectivity in species of conservation concern.     

Phylogeography and ecological niche modeling unravel the evolutionary history of the African green toad,Bufotes boulengeri boulengeri (Amphibia: Bufonidae), through the Quaternary

Recent integration of ecological niche models in phylogeographic studies is improving our understanding of the processes structuring genetic variation across landscapes. Previous studies on the amphibian Bufotes boulengeri boulengeri uncovered a surprisingly weak intraspecific differentiation across the Maghreb region. We widely sampled this species from Morocco to Egypt and sequenced one nuclear and three mitochondrial (mtDNA) genes to determine the level of genetic variability across its geographic range. We evaluated these data with ecological niche modeling to reveal its evolutionary history in response to climate change during the Quaternary. Our results highlight some mtDNA phylogeographic structure within this species, with one haplogroup endemic to coastal Morocco, and one haplogroup widely distributed throughout North Africa. No or little genetic differentiation is observed between isolated populations from the Hoggar Mountains, the Sabha district and the islands of Kerkennah and Lampedusa, compared to others populations. This can be explained by the expansion of the distribution range of B. b. boulengeri during glacial periods. This might have facilitated the species’ dispersal and subsequent gene flow between most North African localities.     

Genetic structure of sigmodontine rodents (Cricetidae) along an altitudinal gradient of the Atlantic Rain Forest in southern Brazil

The population genetic structure of two sympatric species of sigmodontine rodents (Oligoryzomys nigripes and Euryoryzomys russatus) was examined for mitochondrial DNA (mtDNA) sequence haplotypes of the control region. Samples were taken from three localities in the Atlantic Rain Forest in southern Brazil, along an altitudinal gradient with different types of habitat. In both species there was no genetic structure throughout their distribution, although levels of genetic variability and gene flow were high.     

Genetic Variation and Geographic Differentiation in Mitochondrial DNA of the Horseshoe Crab, LIMULUS POLYPHEMUS

Restriction site variation in mitochondrial DNA (mtDNA) of the horseshoe crab (Limulus polyphemus) was surveyed in populations ranging from New Hampshire to the Gulf Coast of Florida. MtDNA clonal diversity was moderately high, particularly in southern samples, and a major genetic "break" (nucleotide sequence divergence approximately 2%) distinguished all sampled individuals which were north vs. south of a region in northeastern Florida. The area of genotypic divergence in Limulus corresponds to a long-recognized zoogeographic boundary between warm-temperate and tropical marine faunas, and it suggests that selection pressures and/or gene flow barriers associated with water mass differences may also influence the evolution of species widely distributed across such transition zones. On the other hand, a comparison of the mtDNA divergence patterns in Limulus with computer models involving stochastic lineage extinction in species with limited gene flow demonstrates that deterministic explanations need not necessarily be invoked to account for the observations. Experiments to distinguish stochastic from deterministic possibilities are suggested. Overall, the pattern and magnitude of mtDNA differentiation in horseshoe crabs is very similar to that typically reported for freshwater and terrestrial species assayed over a comparable geographic range. Results demonstrate for the first time that, geographically, at least some continuously distributed marine organisms can show considerable mtDNA genetic differentiation.     

Regional differences in the distribution of the sub-Saharan, West Eurasian, and South Asian mtDNA lineages in Yemen

Despite its key location for population movements out of and back into Africa, Yemen has not yet been sampled on a regional level for an investigation of sub-Saharan, West Eurasian, and South Asian genetic contributions. In this study, we present mitochondrial DNA (mtDNA) data for regionally distinct Yemeni populations that reveal different distributions of mtDNA lineages. An extensive database of mtDNA sequences from North and East African, Middle Eastern and Indian populations was analyzed to provide a context for the regional Yemeni mtDNA datasets. The groups of western Yemen appear to be most closely related to Middle Eastern and North African populations, while the eastern Yemeni population from Hadramawt is most closely related to East Africa. Furthermore, haplotype matches with Africa are almost exclusively confined to West Eurasian R0a haplogroup in southwestern Yemen, although more sub-Saharan L-type matches appear in more northern Yemeni populations. In fact, Yemeni populations have the highest frequency of R0a haplotypes detected to date, thus Yemen or southern Arabia may be the site of the initial expansion of this haplogroup. Whereas two variants of the sub-Saharan haplogroup M1 were detected only in southwestern Yemen close to the Bab el-Mandeb Strait, different non-African M haplotypes were detected at low frequencies (approximately 2%) in western parts of the country and at a higher frequency (7.5%) in the Hadramawt. We conclude that the Yemeni gene pool is highly stratified both regionally and temporally and that it has received West Eurasian, Northeast African, and South Asian gene flow.     

Population structure of the African savannah elephant inferred from mitochondrial control region sequences and nuclear microsatellite loci

Two hundred and thirty-six mitochondrial DNA nucleotide sequences were used in combination with polymorphism at four nuclear microsatellite loci to assess the amount and distribution of genetic variation within and between African savannah elephants. They were sampled from 11 localities in eastern, western and southern Africa. In the total sample, 43 haplotypes were identified and an overall nucleotide diversity of 2.0% was observed. High levels of polymorphism were also observed at the microsatellite loci both at the level of number of alleles and gene diversity. Nine to 14 alleles per locus across populations and 44 alleles in the total sample were found. The gene diversity ranged from 0.51 to 0.72 in the localities studied. An analysis of molecular variance showed significant genetic differentiation between populations within regions and also between regions. The extent of subdivision between populations at the mtDNA control region was approximately twice as high as shown by the microsatellite loci (mtDNA F(ST) = 0.59; microsatellite R(ST) = 0.31). We discuss our results in the light of Pleistocene refugia and attribute the observed pattern to population divergence in allopatry accompanied by a recent population admixture following a recent population expansion.     

Genetic differentiation in the mud crab Scylla serrata (Decapoda: Portunidae) within the Indian Ocean

Scylla serrata (Decapoda: Portunidae) is a swimming crab that is widespread in the Indo-Pacific region and commonly found in estuarine and mangrove waters. An extended planktonic larval phase suggests high dispersal potential and the possibility of extensive gene flow between conspecific populations at least on a geographic mesoscale (tens to hundreds of kilometres).Intraspecific variation of the mitochondrial DNA cytochrome oxidase subunit I (mtDNA COI) gene was investigated in 77 individuals from four representative mangrove swamps of the African tropics (Kenya and Zanzibar) by means of DNA sequencing. We examined 535 base pairs (bp) and identified 24 different haplotypes. Each population sample is characterised by a single most frequent haplotype, shared among all four populations, and a small number of rare ones, typically present in only one or two individuals and representative of a specific population.Analysis of molecular variance (AMOVA), F_ST statistics and χ² contingency analysis of spatial distribution of mtDNA haplotype frequencies revealed in toto a significant genetic differentiation among populations. These results could indicate that gene flow might be reduced, even between geographically close sites, despite the high potential for dispersal; anyway, at the recorded level of divergence and owing to the abundance of rare haplotypes and singletons in our data set, repeated sampling over time is necessary to establish whether the recorded pattern of genetic differentiation is stable and biologically significant.Finally, integration of our data with those reported by Gopurenko et al. [Mar. Biol. 134 (1999) 227] on S. serrata from South Africa, Red Sea and Mauritius Islands allowed to infer S. serrata population structure within a larger area of the Indian Ocean region.     

A re-evaluation of management units based on gene flow of a rare waterbird in the Americas

The maintenance of gene flow in species that have experienced population contractions and are geographically fragmented is important to the maintenance of genetic variation and evolutionary potential; thus, gene flow is also important to conservation and management of these species. For example, the Reddish Egret (Egretta rufescens) has recovered after severe population reductions during the 19th and 20th centuries, but population numbers remain below historical levels. In this study, we characterized gene flow among management units of the Reddish Egret by using ten nuclear microsatellite markers and part of the mitochondrial (mtDNA) control region from 176 nestlings captured at eight localities in Mexico (Baja California, Chiapas, Tamaulipas, and Yucatan), the USA (Texas, Louisiana, and Florida), and the Bahamas. We found evidence of population structure and that males disperse more often and across longer distances compared with females, which is congruent with previous banding and telemetry data. The maternally inherited mtDNA and biparentally inherited microsatellite data supported slightly different MU models; however, when interpreted together, a four MU model that considered population structure and geographic proximity was most optimal. Namely, MU 1 (Baja California); MU 2 (Chiapas); MU 3 (Yucatan, Tamaulipas, Texas, and Louisiana); and MU 4 (Florida and the Bahamas). Regions outside our sampled localities (e.g., the Greater Antilles and South America) require additional sampling to fully understand gene flow and movement of individuals across the species’ entire range. However, the four MUs we have defined group nesting localities into genetically similar subpopulations, which can guide future management plans.     

Mitochondrial DNA genealogy of chimpanzees in the Nimba mountains and Bossou, West Africa

The chimpanzee populations of the Bossou and Nimba regions in West Africa were genetically surveyed to 1) reveal the genetic relationship between the Bossou and Nimba populations, and 2) elucidate the evolutionary relationship between the Bossou-Nimba and other West African populations. The chimpanzee group at Bossou is characterized by its small population size, no evidence of contact with neighboring populations, and no female immigration. It is believed that most females and adolescent males emigrate from this population. To reveal the genetic signature of these characteristics, we examined the genetic diversity of Bossou and two neighboring populations (Seringbara and Yealé) in the Nimba Mountains by sequencing approximately 605 bp of the mitochondrial DNA (mtDNA) control region. A total of 20 distinct mtDNA variants were observed from 56 sequences of noninvasively collected, anonymous samples. Nucleotide diversity in the Nimba Mountain populations was 0.03-0.04, and did not differ significantly from that in the Bossou population. Very few mitochondrial variants are shared among the sites sampled, which suggests that there is little gene flow involving mtDNA. Nevertheless, no clear population structures were revealed in either population. A comparison with published sequences from West African chimpanzees (Pan troglodytes verus) indicates that the variants observed in the Bossou and Nimba regions are scattered throughout the subspecies, rather than clustered according to geographic region. This suggests that the Bossou-Nimba populations derived only recently from the common ancestral population of the West African chimpanzees, and did not pass through a bottleneck.     

Population genetic structure and male-biased dispersal in the queenless ant Diacamma cyaneiventre

In this study we investigated the population genetic structure of the queenless ant Diacamma cyaneiventre. This species, lacking winged queens, is likely to have a restricted female dispersal. We used both mitochondrial and microsatellite markers to assess the consequence of such restricted female dispersal at three geographical scales: within a given locality (< 1 km), between localities within a given region (< 10 km) and between regions (> 36 km). Within a locality, a strong population structure was observed for mitochondrial DNA (mtDNA) whereas weak or nonexistent population genetic structure was observed for the microsatellites (around 5% of the value for mtDNA). Male gene flow was estimated to be about 20-30 times higher than female gene flow at this scale. At a larger spatial scale, very strong genetic differentiation for both markers was observed between localities - even within a single region. Female dispersal is nonexistent at these scales and male dispersal is very restricted, especially between regions. The phylogeographical structure of the mtDNA haplotypes as well as the very low genetic diversity of mtDNA within localities indicate that new sites are colonized by a single migration event from adjacent localities, followed by successive colony fissions. These patterns of genetic variability and differentiation agree with what is theoretically expected when colonization events are kin-structured and when, following colonization, dispersion is mainly performed by males.     

Demographic inference and genetic diversity of <Emphasis Type="Italic">Octopus mimus</Emphasis> (Cephalopoda: Octopodidae) throughout the Humboldt Current System

Climatic and oceanographic events occurring during the last glacial cycle in the Humboldt Current System (HCS) have left genetic footprints in marine invertebrate populations. The objective of this study was to evaluate the effect of the glacial period on Octopus mimus populations found throughout the HCS. This species lays a large number of small eggs which hatch into planktonic paralarvae with the potential to undergo wide dispersal. We sequenced the COIII gene to perform phylogeographic analyses of 197 octopuses sampled from seven localities. The genetic diversity of Octopus mimus was low and decreased towards the southern end of the distribution range, which comprises a single population. The haplotype genealogy and Bayesian Skyride plot suggest that O. mimus underwent a demographic expansion after the last glacial maximum (LGM). This would imply a contraction of the range of this organism toward northern latitudes during the LGM followed by southward expansion and recolonization once the contemporary interglacial period began.     

Revisiting species delimitation within the genus Oxystele using DNA barcoding approach

The genus Oxystele, a member of the highly diverse marine gastropod superfamily Trochoidea, is endemic to southern Africa. Members of the genus include some of the most abundant molluscs on southern African shores and are important components of littoral biodiversity in rocky intertidal habitats. Species delimitation within the genus is still controversial, especially regarding the complex O. impervia / O. variegata. Here, we assessed species boundaries within the genus using DNA barcoding and phylogenetic tree reconstruction. We analysed 56 specimens using the mitochondrial gene COI. Our analysis delimits five molecular operational taxonomic units (MOTUs), and distinguishes O. impervia from O. variegata. However, we reveal important discrepancies between MOTUs and morphology-based species identification and discuss alternative hypotheses that can account for this. Finally, we indicate the need for future study that includes additional genes, and the combination of both morphology and genetic techniques (e.g. AFLP or microsatellites) to get deeper insight into species delimitation within the genus.     

Is Gene Flow Promoting the Reversal of Pleistocene Divergence in the Mountain Chickadee (Poecile gambeli)?

The Pleistocene glacial cycles left a genetic legacy on taxa throughout the world; however, the persistence of genetic lineages that diverged during these cycles is dependent upon levels of gene flow and introgression. The consequences of secondary contact among taxa may reveal new insights into the history of the Pleistocene’s genetic legacy. Here, we use phylogeographic methods, using 20 nuclear loci from regional populations, to infer the consequences of secondary contact following divergence in the Mountain Chickadee (Poecile gambeli). Analysis of nuclear data identified two geographically-structured genetic groups, largely concordant with results from a previous mitochondrial DNA (mtDNA) study. Additionally, the estimated multilocus divergence times indicate a Pleistocene divergence, and are highly concordant with mtDNA. The previous mtDNA study showed a paucity of sympatry between clades, while nuclear patterns of gene flow show highly varied patterns between populations. The observed pattern of gene flow, from coalescent-based analyses, indicates southern populations in both clades exhibit little gene flow within or between clades, while northern populations are experiencing higher gene flow within and between clades. If this pattern were to persist, it is possible the historical legacy of Pleistocene divergence may be preserved in the southern populations only, and the northern populations would become a genetically diverse hybrid species.     

Barriers to Gene Flow in the Marine Environment: Insights from Two Common Intertidal Limpet Species of the Atlantic and Mediterranean

Knowledge of the scale of dispersal and the mechanisms governing gene flow in marine environments remains fragmentary despite being essential for understanding evolution of marine biota and to design management plans. We use the limpets Patella ulyssiponensis and Patella rustica as models for identifying factors affecting gene flow in marine organisms across the North-East Atlantic and the Mediterranean Sea. A set of allozyme loci and a fragment of the mitochondrial gene cytochrome C oxidase subunit I were screened for genetic variation through starch gel electrophoresis and DNA sequencing, respectively. An approach combining clustering algorithms with clinal analyses was used to test for the existence of barriers to gene flow and estimate their geographic location and abruptness. Sharp breaks in the genetic composition of individuals were observed in the transitions between the Atlantic and the Mediterranean and across southern Italian shores. An additional break within the Atlantic cluster separates samples from the Alboran Sea and Atlantic African shores from those of the Iberian Atlantic shores. The geographic congruence of the genetic breaks detected in these two limpet species strongly supports the existence of transpecific barriers to gene flow in the Mediterranean Sea and Northeastern Atlantic. This leads to testable hypotheses regarding factors restricting gene flow across the study area.     

Shifting clinal patterns and microsatellite variation in Drosophila serrata populations: a comparison of populations near the southern border of the species range

Geographical patterns for quantitative traits in Drosophila and other insects are commonly used to investigate climatic selection. They are usually determined from comparisons of populations over extensive areas and based on one collection per population. Here we consider patterns in the Australian endemic species Drosophila serrata established over a shorter transect with repeated sampling. Summer (prewinter) and spring (post-winter) collections were made from 10 to 14 localities, incorporating the southern border of D.serrata and extending approximately 1000 km northwards along the eastern coast of Australia. Linear or curvilinear associations with latitude were evident for development time, viability and cold resistance but patterns differed between collections. Some geographical (population) and genetic associations between traits were found and these also tended to differ between collections. Results confirm the importance of cold stress resistance over winter to the southern border of this species. Microsatellite markers were developed for D.serrata. These indicated a low level of genetic differentiation between populations, high levels of gene flow and no evidence that the most southerly populations were isolated. The results suggest that selection generated geographical patterns in cold resistance, development time and viability, and that substantial gene flow may prevent adaptation at the border to conditions beyond the current distribution of D. serrata.     

Phylogeography and historical demography of Sicydium salvini in the eastern Pacific

Amphidromy, characterized by freshwater adult and marine larval stages, has been shown to be an important influence on the genetic structure of aquatic populations. Sicydium salvini is a widespread goby species (Teleostei: Gobiidae: Sicydiinae) in the eastern Pacific with a continuous distribution from Mexico to Panama. Here, we use mitochondrial data to infer population genetic and historical demography of this species. Sequences were collected for the mitochondrial gene cytochrome b for 162 specimens sampled across the range of S. salvini with a concentration on rivers in Costa Rica. No genetic structure was detected between regions or rivers in the AMOVA analysis; the phylogeny for this species showed no geographic affinities and very little resolution. Historical demographic analyses indicated a population expansion during the late Pleistocene. These results are consistent with a panmictic population with expansion influenced strongly by Pleistocene glacial cycles and geologic uplift.     

Environmental variables,habitat discontinuity and life history shaping the genetic structure of Pomatoschistus marmoratus

Coastal lagoons are semi-isolated ecosystems exposed to wide fluctuations of environmental conditions and showing habitat fragmentation. These features may play an important role in separating species into different populations, even at small spatial scales. In this study, we evaluate the concordance between mitochondrial (previous published data) and nuclear data analyzing the genetic variability of Pomatoschistus marmoratus in five localities, inside and outside the Mar Menor coastal lagoon (SE Spain) using eight microsatellites. High genetic diversity and similar levels of allele richness were observed across all loci and localities, although significant genic and genotypic differentiation was found between populations inside and outside the lagoon. In contrast to the F _ST values obtained from previous mitochondrial DNA analyses (control region), the microsatellite data exhibited significant differentiation among samples inside the Mar Menor and between lagoonal and marine samples. This pattern was corroborated using Cavalli-Sforza genetic distances. The habitat fragmentation inside the coastal lagoon and among lagoon and marine localities could be acting as a barrier to gene flow and contributing to the observed genetic structure. Our results from generalized additive models point a significant link between extreme lagoonal environmental conditions (mainly maximum salinity) and P. marmoratus genetic composition. Thereby, these environmental features could be also acting on genetic structure of coastal lagoon populations of P. marmoratus favoring their genetic divergence. The mating strategy of P. marmoratus could be also influencing our results obtained from mitochondrial and nuclear DNA. Therefore, a special consideration must be done in the selection of the DNA markers depending on the reproductive strategy of the species.