Multiple colonizations, in situ speciation, and volcanism-associated stepping-stone dispersals shaped the phylogeography of the Macaronesian red fescues (Festuca L., Gramineae)

Whereas examples of insular speciation within the endemic-rich Macaronesian hotspot flora have been documented, the phylogeography of recently evolved plants in the region has received little attention. The Macaronesian red fescues constitute a narrow and recent radiation of four closely related diploid species distributed in the Canary Islands (F. agustinii), Madeira (F. jubata), and the Azores (F. francoi and F. petraea), with a single extant relative distributed in mainland southwest Europe (F. rivularis). Bayesian structure and priority consensus tree approaches and population spatial correlations between genetic, geographical, and dispersal distances were used to elucidate the phylogeographical patterns of these grasses. Independent versus related origins and dispersal versus isolation by distance (IBD) hypotheses were tested to explain the genetic differentiation of species and populations, respectively. Genetic structure was found to be geographically distributed among the archipelagos and the islands endemics. The high number of shared AFLP fragments in all four species suggests a recent single origin from a continental Pliocene ancestor. However, the strong allelic structure detected among the Canarian, Madeiran, and Azorean endemics and the significant standardized residual values obtained from structured Bayesian analysis for pairwise related origin hypotheses strongly supported the existence of three independent continental-oceanic colonization events. The Canarian F. agustinii, the Madeiran F. jubata, and the two sister F. francoi and F. petraea Azorean species likely evolved from different continental founders in their respective archipelagos. Despite the short span of time elapsed since colonization, the two sympatric Azorean species probably diverged in situ, following ecological adaptation, from a common ancestor that arrived from the near mainland. Simple dispersal hypotheses explained most of the genetic variation at the species level better than IBD models. The optimal dispersal model for F. agustinii was a bidirectional centripetal stepping-stone colonization pattern, an eastern-to-western volcanism-associated dispersion was favored for F. francoi, whereas for the recently derived F. petraea a counterintuitive direction of colonization (west-to-east) was suggested. The population-based phylogeographical trends deduced from our study could be used as predictive models for other Macaronesian plant endemics with similar distribution areas and dispersal abilities.     

Coalescent and biophysical models of stepping‐stone gene flow in neritid snails

Marine species in the Indo‐Pacific have ranges that can span thousands of kilometres, yet studies increasingly suggest that mean larval dispersal distances are less than historically assumed. Gene flow across these ranges must therefore rely to some extent on larval dispersal among intermediate ‘stepping‐stone’ populations in combination with long‐distance dispersal far beyond the mean of the dispersal kernel. We evaluate the strength of stepping‐stone dynamics by employing a spatially explicit biophysical model of larval dispersal in the tropical Pacific to construct hypotheses for dispersal pathways. We evaluate these hypotheses with coalescent models of gene flow among high‐island archipelagos in four neritid gastropod species. Two of the species live in the marine intertidal, while the other two are amphidromous, living in fresh water but retaining pelagic dispersal. Dispersal pathways predicted by the biophysical model were strongly favoured in 16 of 18 tests against alternate hypotheses. In regions where connectivity among high‐island archipelagos was predicted as direct, there was no difference in gene flow between marine and amphidromous species. In regions where connectivity was predicted through stepping‐stone atolls only accessible to marine species, gene flow estimates between high‐island archipelagos were significantly higher in marine species. Moreover, one of the marine species showed a significant pattern of isolation by distance consistent with stepping‐stone dynamics. While our results support stepping‐stone dynamics in Indo‐Pacific species, we also see evidence for nonequilibrium processes such as range expansions or rare long‐distance dispersal events. This study couples population genetic and biophysical models to help to shed light on larval dispersal pathways.     

Geographical patterns of genetic structure in marine species with contrasting life histories

Aim  Phylogeographical breaks may reflect historical or present-day impediments to gene flow, and the congruence of these breaks across multiple species lends insight into evolutionary history and connectivity among populations. In marine systems, examining the concordance of phylogeographical breaks is challenging due to the varied sampling scales in population genetics studies and the diverse life histories of marine organisms. A quantitative approach that considers the effects of sampling scale and species life history is needed.
Location  The south-east and south-west coasts of the United States.
Methods  We quantitatively analysed previously published datasets of marine fauna to look for concordance among phylogeographical breaks. We used a bootstrap approach to determine the regions where phylogeographical breaks are more common than expected by chance among species with planktonic dispersal as well as those with restricted dispersal.
Results  On the south-west coast, breaks were clustered near Point Conception among planktonic dispersers and near Los Angeles among restricted dispersers. On the south-east coast, breaks were most common near the southern tip of Florida for planktonic dispersers and near Cape Canaveral for restricted dispersers.
Main conclusions  Dispersal ability is an important determinant of phylogeographical patterns in marine species. Breaks among planktonic dispersers on both coasts are congruent with present-day flow-mediated barriers to dispersal, suggesting that phylogeographical structure in species with planktonic larvae may reflect contemporary oceanography, while breaks in restricted dispersers reflect historical processes. These results highlight the importance of explicitly considering sampling scale and life history when evaluating phylogeographical patterns.     

Extensive local‐scale gene flow and long‐term population stability in the intertidal mollusc Katharina tunicata (Mollusca: Polyplacophora)

The dispersal capabilities of intertidal organisms may represent a key factor to their survival in the face of global warming, as species that cannot adapt to the various effects of climate change will have to migrate to track suitable habitat. Although species with pelagic larval phases might be expected to have a greater capacity for dispersal than those with benthic larvae, interspecies comparisons have shown that this is not always the case. Consequently, population genetic approaches are being increasingly used to gain insights into dispersal through studying patterns of gene flow. In the present study, we used nuclear single‐nucleotide polymorphisms (SNPs) and mitochondrial DNA (mtDNA) sequencing to elucidate fine‐scale patterns of genetic variation between populations of the Black Katy Chiton, Katharina tunicata, separated by 15–150 km in south‐west Vancouver Island. Both the nuclear and mitochondrial data sets revealed no genetic differentiation between the populations studied, and an isolation‐with‐migration analysis indicated extensive local‐scale gene flow, suggesting an absence of barriers to dispersal. Population demographic analysis also revealed long‐term population stability through previous periods of climate change associated with the Pleistocene glaciations. Together, the findings of the present study suggest that this high potential for dispersal may allow K. tunicata to respond to current global warming by tracking suitable habitat, consistent with its long‐term demographic stability through previous changes in the Earth's climate. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 589–597.     

Long‐term panmixia in a cosmopolitan Indo‐Pacific coral reef fish and a nebulous genetic boundary with its broadly sympatric sister species

Phylogeographical studies have shown that some shallow‐water marine organisms, such as certain coral reef fishes, lack spatial population structure at oceanic scales, despite vast distances of pelagic habitat between reefs and other dispersal barriers. However, whether these dispersive widespread taxa constitute long‐term panmictic populations across their species ranges remains unknown. Conventional phylogeographical inferences frequently fail to distinguish between long‐term panmixia and metapopulations connected by gene flow. Moreover, marine organisms have notoriously large effective population sizes that confound population structure detection. Therefore, at what spatial scale marine populations experience independent evolutionary trajectories and ultimately species divergence is still unclear. Here, we present a phylogeographical study of a cosmopolitan Indo‐Pacific coral reef fish Naso hexacanthus and its sister species Naso caesius, using two mtDNA and two nDNA markers. The purpose of this study was two‐fold: first, to test for broad‐scale panmixia in N. hexacanthus by fitting the data to various phylogeographical models within a Bayesian statistical framework, and second, to explore patterns of genetic divergence between the two broadly sympatric species. We report that N. hexacanthus shows little population structure across the Indo‐Pacific and a range‐wide, long‐term panmictic population model best fit the data. Hence, this species presently comprises a single evolutionary unit across much of the tropical Indian and Pacific Oceans. Naso hexacanthus and N. caesius were not reciprocally monophyletic in the mtDNA markers but showed varying degrees of population level divergence in the two nuclear introns. Overall, patterns are consistent with secondary introgression following a period of isolation, which may be attributed to oceanographic conditions of the mid to late Pleistocene, when these two species appear to have diverged.     

Genetic differentiation of freshwater pond copepods at arctic sites

Freshwater pond copepod species at a low-arctic site show distributional and life history differences which may reflect different dispersal efficiencies. In order to ascertain levels of gene flow among populations, cellulose acetate gel electrophoresis was used to examine allozyme variation in Heterocope septentrionalis, Hesperodiaptomus victoriaensis and Leptodiaptomus tyrrelli near Churchill, Canada. Differentiation of gene frequencies among populations of these species, plus Hesperodiaptomus eiseni and Hesperodiaptomus arcticus at other sites along Hudson Bay was moderate. The variation in gene frequencies was less than that of other passively dispersing organisms from the same habitats, and only slightly greater than that reported for an intertidal copepod with pelagic larvae. The mean number of dispersers exchanged among populations per generation, estimated from Wright's island model, averages 4.1 for the three species. Dispersal efficiency, calculated using population size estimates, revealed differences among the three species. However, these differences were not consistent with that expected from their distributional patterns. This suggests that factors other than dispersal alone determine the distributions of copepod species.     

The evolution of north‐east Atlantic gadfly petrels using statistical phylogeography

Macaronesia (north‐east Atlantic archipelagos) has been host to complex patterns of colonization and differentiation in many groups of organisms including seabirds such as gadfly petrels (genus Pterodroma). Considering the subspecies of widely distributed soft‐plumaged petrel for many years, the taxonomic status of the three gadfly petrel taxa breeding in Macaronesia is not yet settled, some authors advocating the presence of three, two or one species. These birds have already been the subject of genetic studies with only one mtDNA gene and relatively modest sample sizes. In this study, using a total of five genes (two mitochondrial genes and three nuclear introns), we investigated the population and phylogeographical histories of petrel populations breeding on Madeira and Cape Verde archipelagos. Despite confirming complete lineage sorting with mtDNA, analyses with nucDNA failed to reveal any population structuring and Isolation with Migration analysis revealed the absence of gene flow during the differentiation process of these populations. It appears that the three populations diverged in the late Pleistocene in the last 150 000 years, that is 10 times more recently than previous estimates based solely on one mtDNA gene. Finally, our results suggest that the Madeira petrel population is ancestral rather than that from Cape Verde. This study strongly advocates the use of nuclear loci in addition to mtDNA in demographical and phylogeographical history studies.     

High gene flow in oceanic bottlenose dolphins (Tursiops truncatus) of the North Atlantic

Despite the openness of the oceanic environment, limited dispersal and tight social structure often induce genetic structuring in marine organisms, even in large animals such as cetaceans. In the bottlenose dolphin, mitochondrial and nuclear DNA analyses have revealed the existence of genetic differentiation between pelagic (or offshore) and coastal (or nearshore) ecotypes in the western North Atlantic, as well as between coastal populations. Because previous studies concentrated on continental margins, we analysed the population structure of bottlenose dolphins in two of the most isolated archipelagos of the North Atlantic: the Azores and Madeira. We analysed 112 samples collected on live animals in the two archipelagos, and nine samples collected on stranded animals in Madeira and mainland Portugal. Genetic analyses consisted in molecular sexing, sequencing of part of the mitochondrial hyper-variable region, and screening of ten microsatellite loci. We predicted that: (1) there is at least one pelagic and one or more coastal populations in each archipelago; (2) populations are differentiated between and possibly within archipelagos. Contrary to these predictions, results indicated a lack of population structure in the study area. In addition, comparison with published sequences revealed that the samples from the Azores and Madeira were not significantly differentiated from samples of the pelagic population of the western North Atlantic. Thus, bottlenose dolphins occurring in the pelagic waters of the North Atlantic belong to a large oceanic population, which should be regarded as a single conservation unit. Unlike what is known for coastal populations, oceanic bottlenose dolphins are able to maintain high levels of gene flow.     

Population history of Berthelot's pipit: colonization, gene flow and morphological divergence in Macaronesia

The fauna of oceanic islands provide exceptional models with which to examine patterns of dispersal, isolation and diversification, from incipient speciation to species level radiations. Here, we investigate recent differentiation and microevolutionary change in Berthelot's pipit (Anthus berthelotii), an endemic bird species inhabiting three Atlantic archipelagos. Mitochondrial DNA sequence data and microsatellite markers were used to deduce probable colonization pathway, genetic differentiation, and gene flow among the 12 island populations. Phenotypic differentiation was investigated based on eight biologically important morphological traits. We found little mitochondrial DNA variability, with only one and four haplotypes for the control region and cytochrome b, respectively. However, microsatellite data indicated moderate population differentiation (FST=0.069) between the three archipelagos that were identified as genetically distinct units with limited gene flow. Both results, combined with the estimated time of divergence (2.5 millions years ago) from the Anthus campestris (the sister species), suggest that this species has only recently dispersed throughout these islands. The genetic relationships, patterns of allelic richness and exclusive alleles among populations suggest the species originally colonized the Canary Islands and only later spread from there to the Madeiran archipelago and Selvagen Islands. Differentiation has also occurred within archipelagos, although to a lesser degree. Gene flow was observed more among the eastern and central islands of the Canaries than between these and the western islands or the Madeiran Islands. Morphological differences were also more important between than within archipelagos. Concordance between morphological and genetic differentiation provided ambiguous results suggesting that genetic drift alone was not sufficient to explain phenotypic differentiation. The observed genetic and morphological differences may therefore be the result of differing patterns of selection pressures between populations, with Berthelot's pipit undergoing a process of incipient differentiation.     

Introgression versus immigration in hybridizing high-dispersal echinoderms

Phylogeographic studies designed to estimate rates and patterns of genetic differentiation within species often reveal unexpected and graphically striking cases of allele or haplotype sharing between species (introgression) via hybridization and backcrossing. Does introgression between species significantly influence population genetic structure relative to more conventional sources of differentiation (drift) and similarity (dispersal) among populations within species? Here we use mtDNA sequences from four species in two genera of sea urchins and sea stars to quantify the relative magnitude of gene flow across oceans and across species boundaries in the context of the trans-Arctic interchange of marine organisms between the Pacific and Atlantic oceans. In spite of the much smaller distances between sympatric congeners, rates of gene flow between sympatric species via heterospecific gamete interactions were small and significantly lower than gene flow across oceans via dispersal of planktonic larvae. We conclude that, in these cases at least, larvae are more effective than gametes as vectors of gene flow.     

Late Neogene history of the laurel tree (Laurus L., Lauraceae) based on phylogeographical analyses of Mediterranean and Macaronesian populations

Aim The post‐glacial range dynamics of many European plant species have been widely investigated, but information rapidly diminishes as one moves further back in time. Here we infer the historical range shifts of Laurus, a paradigmatic tree of the Tethyan flora that has covered southern Eurasia since the Oligo‐Miocene, by means of phylogenetic and phylogeographical analyses. Location Mediterranean Basin, Black Sea and Macaronesian archipelagos (Azores, Madeira, Canary Islands). Methods We analysed plastid DNA (cpDNA) sequence (trnK–matK, trnD–trnT) variation in 57 populations of Laurus and three Lauraceae genera. Phylogenetic methods (maximum parsimony and Bayesian inference) and statistical parsimony networks were used to reconstruct relationships among haplotypes. These results were contrasted with the fossil record and bioclimatic niche‐based model predictions of past distributions to infer the migration routes and location of refugia. Results The phylogenetic tree revealed monophyly for Laurus. Overall sequence variability was low within Laurus, but six different haplotypes were distinguished and a single network retrieved, portraying three lineages primarily related to geography. A strongly divergent eastern lineage occupied Turkey and the Near East, a second clade was located in the Aegean region and, lastly, a western clade grouped all Macaronesian and central and western Mediterranean populations. A close relationship was observed between the Macaronesian populations of L. azorica and the western populations of L. nobilis. Main conclusions The phylogeographical structure of Laurus preserves the imprints of an ancient contraction and break‐up of the range that resulted in the evolution of separate cpDNA lineages in its western‐ and easternmost extremes. Intense range dynamics in the western Mediterranean and multiple glacial refugia contributed to the generation and long‐term conservation of this phylogeographical pattern, as shown by the fit between the haplotype ranges and past suitable areas inferred from bioclimatic models. Finally, our results challenge the taxonomic separation of Laurus into two distinct species.     

Contrasting demographic history and phylogeographical patterns in two Indo-Pacific gastropods

Marine species with ranges that span the Indo-Australian Archipelago (IAA) exhibit a range of phylogeographical patterns, most of which are interpreted in the context of vicariance between Indian and Pacific Ocean populations during Pliocene and Pleistocene low sea-level stands. However, patterns often vary among ecologically similar taxa, sometimes even within genera. This study compares phylogeographical patterns in two species of highly dispersive neritid gastropod, Nerita albicilla and Nerita plicata, with nearly sympatric ranges that span the Indo-Pacific. Mitochondrial COI sequences from >1000 individuals from 97 sites reveal similar phylogenies in both species (two divergent clades differing by 3.2% and 2.3%, for N. albicilla and N. plicata, respectively). However, despite ecological similarity and congeneric status, the two species exhibit phylogeographical discordance. N. albicilla has maintained reciprocal monophyly of Indian and Pacific Ocean populations, while N. plicata is panmictic between oceans, but displays a genetic cline in the Central Pacific. Although this difference might be explained by qualitatively different demographic histories, parameter estimates from three coalescent models indicate that both species have high levels of gene flow between demes (2Nem>75), and share a common history of population expansion that is likely associated with cyclical flooding of continental shelves and island lagoons following low sea-level stands. Results indicate that ecologically similar, codistributed species may respond very differently to shared environmental processes, suggesting that relatively minor differences in traits such as pelagic larval duration or microhabitat association may profoundly impact phylogeographical structure.     

Phylogeographical structure of the Dover sole Microstomus pacificus: the larval retention hypothesis and genetic divergence along the deep continental slope of the northeastern Pacific Ocean

Population genetic variability and phylogeographical structure of the Dover sole, Microstomus pacificus (Teleostei: Pleuronectidae), were examined along the continental slope of the north-eastern Pacific Ocean using sequences of the left domain of the mitochondrial DNA (mtDNA) control region. Six sites were sampled in three primary biogeographical provinces (based on surface water temperatures), ranging from the subarctic to the warm temperate zones. Many haplotypes were discerned, totalling 90 for 110 individuals. Neighbour joining and parsimony analyses of the entire data set showed that some groups of haplotypes of adult Dover sole were widely distributed, suggesting either panmixia or homoplasy and reversals. However, a substantial number of groups of related haplotypes were geographically circumscribed, and there were significant differences among sites in the distribution of restricted haplotypes (based on a reduced number of characters), supporting geographical structure. Geographical differentiation of populations appeared to be consistent with the member-vagrant hypothesis for retention of the pelagic larvae in currents and recruitment to adult habitats, reducing long-distance dispersal. Results of this study indicate phylogeographical structure among some populations, despite extended pelagic larval periods, and apparent environmental homogeneity in the adult habitats along the continental slope.     

Adult habitat preferences, larval dispersal, and the comparative phylogeography of three Atlantic surgeonfishes (Teleostei: Acanthuridae)

Although many reef fishes of the tropical Atlantic are widely distributed, there are large discontinuities that may strongly influence phylogeographical patterns. The freshwater outflow of the Amazon basin is recognized as a major barrier that produces a break between Brazilian and Caribbean faunas. The vast oceanic distances between Brazil and the mid-Atlantic ridge islands represent another formidable barrier. To assess the relative importance of these barriers, we compared a fragment of the mitochondrial DNA (mtDNA) cytochrome b gene among populations of three species of Atlantic surgeonfishes: Acanthurus bahianus, A. chirurgus and A. coeruleus. These species have similar life histories but different adult habitat preferences. The mtDNA data show no population structure between Brazil and the mid-Atlantic islands, indicating that this oceanic barrier is readily traversed by the pelagic larval stage of all three surgeonfishes, which spend approximately 45-70 days in the pelagic environment. The Amazon is a strong barrier to dispersal of A. bahianus (d = 0.024, phiST = 0.724), a modest barrier for A. coeruleus (phiST = 0.356), and has no discernible effect as a barrier for A. chirurgus. The later species has been collected on soft bottoms with sponge habitats under the Amazon outflow, indicating that relaxed adult habitat requirements enable it to readily cross that barrier. A limited ability to use soft bottom habitats may also explain the low (but significant) population structure in A. coeruleus. In contrast, A. bahianus has not been collected over deep sponge bottoms, and rarely settles outside shallow reefs. Overall, adult habitat preferences seem to be the factor that differentiates phylogeographical patterns in these reef-associated species.     

A comparative study of asymmetric migration events across a marine biogeographic boundary

In many nonclonal, benthic marine species, geographic distribution is mediated by the dispersal of their larvae. The dispersal and recruitment of marine larvae may be limited by temperature gradients that can affect mortality or by ocean currents that can directly affect the movements of pelagic larvae. We focus on Point Conception, a well-known biogeographic boundary between the Californian and Oregonian biogeographic provinces, to investigate whether ocean currents affect patterns of gene flow in intertidal marine invertebrates. The predominance of pelagically dispersing species with northern range limits at Point Conception suggests that ocean currents can affect species distributions by erecting barriers to the dispersal of planktonic larvae. In this paper, we investigate whether the predominantly southward currents have left a recognizable genetic signature in species with pelagically dispersing larvae whose ranges span Point Conception. We use patterns of genetic diversity and a new method for inferring cladistic migration events to test the hypothesis that southward currents increase southward gene flow for species with pelagically dispersing larvae. We collected mitochondrial DNA (mtDNA) sequence data for the barnacles Balanus glandula and Chthamalus fissus and also reanalyzed a previously published mtDNA dataset (Strongylocentrotus purpuratus, Edmands et al. 1996). For all three species, our cladistic approach identified an excess of southward migration events across Point Conception. In data from a fourth species with nondispersing larvae (Nucella emarginata, Marko 1998), our method suggests that ocean currents have not played a role in generating genetic structure.     

What can population genetics tell us about dispersal and biogeographic history of coral-reef fishes?

Abstract Coral-reef fishes, like many other marine organisms, generally possess a benthic adult stage and pelagic larval stage. What can population genetics studies tell us about the demographic, evolutionary and biogeographic consequences of this life cycle? Ten studies of geographical patterns of intraspecific genetic differentiation in reef fishes have been published. These studies have included 2t > species/species complexes (14 in the family Pomacentridae, the remaining 12 in 9 different families) and have been about equally divided between the tropical Pacific and the tropical western Atlantic. A survey of these studies shows the following: (i) the existence of the pelagic larval stage appears to have led to high levels of gene flow even among populations separated by thousands of kilometres of open ocean; (ii) an apparent pattern of increased gene flow among populations connected by intermediate 'stepping stones’; (iii) very tentative evidence for a relationship between length of pelagic larval life and gene flow; (iv) no clear relationship between egg type (pelagic rs non-pelagic) and gene flow; and (v) suggestive evidence that damselfishes (family Pomacentridae) may have more restricted dispersal (less gene flow) than other reef fishes. The application of current and future molecular tools has the strong potential to clarify some of these relationships, particularly by using relatively neutral genetic markers. Additionally, discoveries of DNA markers having very high rates of mutation may allow tracking of demographically relevant levels of larval dispersal. Molecular tools are becoming especially valuable in uncovering the biogeographic and phylogenetic history of reef fishes. The one molecular study to date has suggested that at least some speciation events may have occurred during the climate changes and sea-level regressions associated with Pleistocene glacial episodes. Molecular tools need to be used to further explore the means by which high species diversity can be generated in the face of the apparently high gene flow observed in most coral-reef fishes.     

Geographical patterns of allozymic variation in three species of intertidal sculpins

Synopsis We examined spatial patterns of allozyme variation in three species of intertidal cottids to evaluate how much gene flow occurs between geographically separate populations (most likely via passive dispersal of planktonic larvae by currents). Our results from the analysis of geographical patterns of allele frequencies and, to some extent, from comparison of allele frequencies between newly recruited young and older resident fish are consistent with the notion that sufficient gene flow occurs in these fishes to prevent significant genetic isolation of populations. From these results, we visualize evolutionary changes in populations of these species as occurring most likely over a broad geographic scale, affecting spatially separated but genetically linked populations concurrently, rather than operating independently in individual, genetically isolated populations.     

Disentangling the genetic and morphological structure of Patella candei complex in Macaronesia (NE Atlantic)

The uptake of natural living resources for human consumption has triggered serious changes in the balance of ecosystems. In the archipelagos of Macaronesia (NE Atlantic), limpets have been extensively exploited probably since islands were first colonized. This has led to profound consequences in the dynamics of rocky shore communities. The Patella candei complex includes various subspecies of limpets that are ascribed to a particular archipelago and has been the focus of several taxonomic surveys without much agreement. Under a conservational perspective, we apply morphometric and genetic analyses to test subspecies boundaries in P. candei and to evaluate its current population connectivity throughout Macaronesia (Azores, Madeira, and Canaries). A highly significant genetic break between archipelagos following isolation by distance was detected (F_ST = 0.369, p < .001). Contrastingly, significant genetic differentiation among islands (i.e., Azores) was absent possibly indicating ongoing gene flow via larval exchange between populations. Significant shell‐shape differences among archipelagos were also detected using both distance‐based and geometric morphometric analyses. Adaptive processes associated with niche differentiation and strong barriers to gene flow among archipelagos may be the mechanisms underlying P. candei diversification in Macaronesia. Under the very probable assumption that populations of P. candei from each archipelago are geographically and/or ecologically isolated populations, the various subspecies within the P. candei complex may be best thought of as true species using the denomination: P. candei in Selvagens, Patella gomesii in Azores, Patella ordinaria in Madeira, and Patella crenata for Canaries. This would be in agreement with stock delimitation and units of conservation of P. candei sensu latu along Macaronesia.     

DEVELOPMENTAL MODE AND SPECIES GEOGRAPHIC RANGE IN REGULAR SEA URCHINS (ECHINODERMATA: ECHINOIDEA)

Among marine benthic organisms, the ability to disperse, primarily during the larval stage, is widely thought to influence the extent of species geographic range. Because related species often differ in their modes of larval development (pelagic, feeding larvae; pelagic, nonfeeding larvae; or brooded development), and these can have dramatically different planktonic intervals, the mode of development may influence geographic range. A global survey of 215 regular echinoids shows that species with pelagic, feeding larvae have significantly larger ranges than those with pelagic, nonfeeding larvae, but there is no difference in ranges between species with pelagic, nonfeeding larvae and those with brooded development. These patterns are maintained within the Cidaroida and the Temnopleuroida, which account for the great majority of species with pelagic, nonfeeding development and brooded development. This limited effect of developmental mode on geographic range is found among species occurring predominantly in waters shallower than 100 m. For species occurring deeper than 100 m, there is no significant difference in geographic range related to type of development. The relationship between developmental mode and species range was examined more closely for circa 30 species for which the developmental period was known from laboratory observations. Adjusting the developmental times to a common temperature, 20°C, using realistic values for Q₁₀ from 2.0 to 3.6, showed a highly significant, negative correlation between egg volume and developmental time, indicating the potential for developmental mode to influence the planktonic interval. However, there was no relationship between time in the plankton, estimated from unadjusted developmental times, and extent of species geographic range. These results suggest that developmental mode may influence extent of species geographic ranges indirectly through the consequences of dispersal for gene flow or recovery from disturbance.     

GENETIC STRUCTURE OF GIANT CLAM (TRIDACNA MAXIMA) POPULATIONS IN THE WEST PACIFIC IS NOT CONSISTENT WITH DISPERSAL BY PRESENT-DAY OCEAN CURRENTS

The Pacific marine biota, particularly species with long planktonic larval stages, are thought to disperse widely throughout the Pacific via ocean currents. The little genetic data available to date has supported this view in that little or no significant regional differentiation of populations has been found over large geographical distances. However, recent data from giant clams has demonstrated not only significant regional differentiation of populations, but routes of gene flow that run perpendicular to the main present-day ocean currents. Extensive surveys of genetic variation at eight polymorphic loci in 19 populations of the giant clam Tridacna maxima, sampled throughout the West and Central Pacific, confirmed that the patterns of variation seen so far in T. gigas were not unique to that species, and may reflect a fundamental genetic structuring of shallow-water marine taxa. Populations of T. maxima within highly connected reef systems like the Great Barrier Reef were panmictic (average F_ST < 0.003), but highly significant genetic differences between reef groups on different archipelagos (average F_ST = 0.084) and between West and Central Pacific regions (average F_ST = 0.156) were found. Inferred gene flow was high (N_em usually > 5) between the Philippines and the Great Barrier Reef, between the Philippines and Melanesia (the Solomon Islands and Fiji), and between the Philippines and the Central Pacific island groups (Marshall Islands, Kiribati, Tuvalu and Cook Islands). Gene flow was low between these three sets of island chains (N_em < 2). These routes of gene flow are perpendicular to present-day ocean currents. It is suggested that the spatial patterns of gene frequencies reflect past episodes of dispersal at times of lower sea levels which have not been erased by subsequent dispersal by present-day circulation. The patterns are consistent with extensive dispersal of marine species in the Pacific, and with traditional views of dispersal from the Indo-Malay region. However, they demonstrate that dispersal along present-day ocean surface currents cannot be assumed, that other mechanisms may operate today or that major dispersal events are intermittent (perhaps separated by several thousands of years), and that the nature and timing of dispersal of Pacific marine species is more complex than has been thought.