The location,strength, and mechanisms behind marine biogeographic boundaries of the east coast of North America

Classic biogeographic studies emphasized differences in species composition between regions to define biogeographic provinces and delimit biogeographic boundaries. Here we analyze the permeability of biogeographic boundaries to different species to gain mechanistic insight into the processes that maintain species boundaries in the coastal ocean. We identify sites with high frequencies of range boundaries using almost 1800 benthic marine invertebrates along the northwestern Atlantic coast and address whether their magnitude and location vary as a function of species’ taxonomy, pelagic larval duration and depth distribution. We observed clusters of species boundaries at Cape Hatteras, Cape Cod and the Bay of Fundy that are largely independent of taxonomic group. However, the boundaries were permeable and asymmetric, with a higher percentage of species shared across boundaries in the equatorward direction (82%) than in the reverse direction (59%). This pattern was particularly strong for shallow species (median occurrence depth < 20 m). Pelagic larval duration was more important to explain distributions of boundaries for deep species (median occurrence depth > 20 m), where species with long larval dispersal had significantly higher occurrence of boundaries than species with short larval dispersal. When they do exist, species boundaries seem to be set by the interaction of currents, depth distribution and pelagic larval duration. Importantly, species boundaries tend to be pinned to regions of reduced water transport, which might explain why species boundaries are concentrated in narrow geographical areas.     

The oceanic concordance of phylogeography and biogeography: a case study in Notochthamalus

Dispersal and adaptation are the two primary mechanisms that set the range distributions for a population or species. As such, understanding how these mechanisms interact in marine organisms in particular – with capacity for long‐range dispersal and a poor understanding of what selective environments species are responding to – can provide useful insights for the exploration of biogeographic patterns. Previously, the barnacle Notochthamalus scabrosus has revealed two evolutionarily distinct lineages with a joint distribution that suggests an association with one of the two major biogeographic boundaries (~30°S) along the coast of Chile. However, spatial and genomic sampling of this system has been limited until now. We hypothesized that given the strong oceanographic and environmental shifts associated with the other major biogeographic boundary (~42°S) for Chilean coastal invertebrates, the southern mitochondrial lineage would dominate or go to fixation in locations further to the south. We also evaluated nuclear polymorphism data from 130 single nucleotide polymorphisms to evaluate the concordance of the signal from the nuclear genome with that of the mitochondrial sample. Through the application of standard population genetic approaches along with a Lagrangian ocean connectivity model, we describe the codistribution of these lineages through a simultaneous evaluation of coastal lineage frequencies, an approximation of larval behavior, and current‐driven dispersal. Our results show that this pattern could not persist without the two lineages having distinct environmental optima. We suggest that a more thorough integration of larval dynamics, explicit dispersal models, and near‐shore environmental analysis can explain much of the coastal biogeography of Chile.     

Genetic Structuring across Marine Biogeographic Boundaries in Rocky Shore Invertebrates

Biogeography investigates spatial patterns of species distribution. Discontinuities in species distribution are identified as boundaries between biogeographic areas. Do these boundaries affect genetic connectivity? To address this question, a multifactorial hierarchical sampling design, across three of the major marine biogeographic boundaries in the central Mediterranean Sea (Ligurian-Tyrrhenian, Tyrrhenian-Ionian and Ionian-Adriatic) was carried out. Mitochondrial COI sequence polymorphism of seven species of Mediterranean benthic invertebrates was analysed. Two species showed significant genetic structure across the Tyrrhenian-Ionian boundary, as well as two other species across the Ionian Sea, a previously unknown phylogeographic barrier. The hypothesized barrier in the Ligurian-Tyrrhenian cannot be detected in the genetic structure of the investigated species. Connectivity patterns across species at distances up to 800 km apart confirmed that estimates of pelagic larval dispersal were poor predictors of the genetic structure. The detected genetic discontinuities seem more related to the effect of past historical events, though maintained by present day oceanographic processes. Multivariate statistical tools were used to test the consistency of the patterns across species, providing a conceptual framework for across-species barrier locations and strengths. Additional sequences retrieved from public databases supported our findings. Heterogeneity of phylogeographic patterns shown by the 7 investigated species is relevant to the understanding of the genetic diversity, and carry implications for conservation biology.     

Environmental drivers of species composition and functional diversity of dung beetles along the Atlantic Forest–Pampa transition zone

Human activities are causing a rapid loss of biodiversity, which impairs ecosystem functions and services. Therefore, understanding which processes shape how biodiversity is distributed along spatial and environmental gradients is a first step to guide conservation and management efforts. We aimed to determine the relative explanatory importance of biogeographic, environmental, landscape and spatial variables on assemblage dissimilarities and functional diversity of dung beetles along the Atlantic Forest–Pampa (i.e. forest–grassland) transition zone located in Southeast South America. We described each site according to their biogeographic position, environmental conditions, landscape features and spatial patterns. The compositional dissimilarity was partitioned into turnover and nestedness components of β‐diversity. Mantel tests and generalised dissimilarity models were used to relate β‐diversity and its components to biogeographic, environmental, landscape and spatial variables. Variation partitioning analysis was used to estimate the pure and shared variation in species composition and functional diversity explained by the four categories of predictors. Biome domain was the main factor causing dung beetle compositional dissimilarity, with a high species replacement between Atlantic Forest and Pampa. Biogeographic, environmental, landscape and spatial distances also affected the patterns of dung beetle dissimilarity and β‐diversity components. The shared effects of the four sets of predictors explained most of the variation in dung beetle composition. A similar response pattern was found for dung beetle functional diversity, which excluded biogeographic effects. Only the pure effects of environmental and spatial predictors were significant for species composition and functional diversity. Our results indicate that dung beetle species composition and functional diversity are jointly driven by environmental, landscape and spatial predictors with higher pure environmental and spatial effects. The forest–grassland transition zone promotes a strong species and trait replacement highly influenced both by environmental filtering and dispersal limitation.     

Unexpected collective larval dispersal but little support for sweepstakes reproductive success in the highly dispersive brooding mollusc Crepidula fornicata

In many marine invertebrates, long‐distance dispersal is achieved during an extended pelagic larval phase. Although such dispersal should result in high gene flow over broad spatial scales, fine‐scale genetic structure has often been reported, a pattern attributed to interfamilial variance in reproductive success and limited homogenization during dispersal. To examine this hypothesis, the genetic diversity of dispersing larvae must be compared with the postdispersal stages, that is benthic recruits and adults. Such data remain, however, scarce due to the difficulty to sample and analyse larvae of minute size. Here, we carried out such an investigation using the marine gastropod Crepidula fornicata. Field sampling of three to four larval pools was conducted over the reproductive season and repeated over 3 years. The genetic composition of larval pools, obtained with 16 microsatellite loci, was compared with that of recruits and adults sampled from the same site and years. In contrast to samples of juveniles and adults, large genetic temporal variations between larval pools produced at different times of the same reproductive season were observed. In addition, full‐ and half‐sibs were detected in early larvae and postdispersal juveniles, pointing to correlated dispersal paths between several pairs of individuals. Inbred larvae were also identified. Such collective larval dispersal was unexpected given the long larval duration of the study species. Our results suggest that each larval pool is produced by a small effective number of reproducers but that, over a reproductive season, the whole larval pool is produced by large numbers of reproducers across space and time.     

Climate‐driven trends and ecological implications of event‐scale upwelling in the California Current System

Eastern boundary current systems are among the most productive and lucrative ecosystems on Earth because they benefit from upwelling currents. Upwelling currents subsidize the base of the coastal food web by bringing deep, cold and nutrient‐rich water to the surface. As upwelling is driven by large‐scale atmospheric patterns, global climate change has the potential to affect a wide range of significant ecological processes through changes in water chemistry, water temperature, and the transport processes that influence species dispersal and recruitment. We examined long‐term trends in the frequency, duration, and strength of continuous upwelling events for the Oregon and California regions of the California Current System in the eastern Pacific Ocean. We then associated event‐scale upwelling with up to 21 years of barnacle and mussel recruitment, and water temperature data measured at rocky intertidal field sites along the Oregon coast. Our analyses suggest that upwelling events are changing in ways that are consistent with climate change predictions: upwelling events are becoming less frequent, stronger, and longer in duration. In addition, upwelling events have a quasi‐instantaneous and cumulative effect on rocky intertidal water temperatures, with longer events leading to colder temperatures. Longer, more persistent upwelling events were negatively associated with barnacle recruitment but positively associated with mussel recruitment. However, since barnacles facilitate mussel recruitment by providing attachment sites, increased upwelling persistence could have indirect negative impacts on mussel populations. Overall, our results indicate that changes in coastal upwelling that are consistent with climate change predictions are altering the tempo and the mode of environmental forcing in near‐shore ecosystems, with potentially severe and discontinuous ramifications for ecosystem structure and functioning.     

Biogeographic and bathymetric ranges of Atlantic deep-sea echinoderms and ascidians: the role of larval dispersal

Dispersal plays an important role in the establishment and maintenance of biodiversity and, for most deep-sea benthic marine invertebrates, it occurs mainly during the larval stages. Therefore, the mode of reproduction (and thus dispersal ability) will affect greatly the biogeographic and bathymetric distributions of deep-sea organisms. We tested the hypothesis that, for bathyal and abyssal echinoderms and ascidians of the Atlantic Ocean, species with planktotrophic larval development have broader biogeographic and bathymetric ranges than species with lecithotrophic development. In comparing two groups with lecithotrophic development, we found that ascidians, which probably have a shorter larval period and therefore less dispersal potential, were present in fewer geographic regions than elasipod holothurians, which are likely to have longer larval periods. For asteroids and echinoids, both the geographic and bathymetric ranges were greater for lecithotrophic than for planktotrophic species. For these two classes, the relationships of egg diameter with geographic and bathymetric range were either linearly increasing or non-monotonic. We conclude that lecithotrophic development does not necessarily constrain dispersal in the deep sea, probably because species with planktotrophic development may be confined to regions of high detrital input from the sea surface. Our data suggest that more information is necessary on lengths of larval period for different species to accurately assess dispersal in the deep sea.     

51 Characterization of the northwestern gulf of Mexico macroalgae

One of the most commonly predicted effects of global ocean warming on marine communities is a poleward shift in the distribution of species with an associated replacement of cold‐water species by warm‐water species. Such predictions are imprecise and based largely on broad correlations in uncontrolled studies that examine changes in species composition and abundance relative to seawater temperature. Before‐After‐Control‐Impact (BACI) analyses of the effects of a large thermal discharge shows that an induced 3.4 deg. C rise in seawater temperature over 10 years along 2 km of rocky coastline resulted in significant community‐wide changes in 150 species of algae and invertebrates relative to controls. Contrary to predictions from biogeographic models, there was no trend towards warm‐water species with southern geographic affinities replacing cold‐species with northern affinities. Instead, communities were greatly altered in apparently cascading responses to changes in abundance of several habitat‐forming taxa, particularly subtidal kelps (e.g. Pterygophora californica) and intertidal foliose red algae (e.g. Mazzaella flaccida). Many temperature sensitive algae decreased greatly in abundance, whereas many invertebrate grazers increased. The results indicate that the responses of temperate reef communities to ocean warming can be strongly coupled to direct effects on habitat‐forming taxa and indirect effects operating through ecological interactions. Given our understanding of temperate reef ecology and its local variability, the results also suggest that accurate predictions of the effects of global ocean warming will be difficult to make.     

Natural history, biogeography, and endangerment of Hawaiian dry forest trees

We describe the floristic composition of Hawaiian dry forest trees and identify natural history characteristics and biogeographic variables that are associated with risk of endangerment. Hawaiian dry forests are comprised of 109 tree species in 29 families, with 90% of all species endemic, 10% indigenous, and 37% single-island endemics. Forty-five percent of Hawaiian dry forest taxa are at risk of endangerment. Dry forest taxa at risk have a significantly larger range size compared to taxa from other Hawaiian forest types. Dispersal mechanism was a significant predictor of a species occurrence in dry forest compared to other forest types based on logistic regressions clustered by lineage. Among dry forest taxa, hermaphroditic breeding systems, autochorous dispersal mechanisms, conspicuous flowers, and dry fruit were all more likely to be at risk of endangerment. When analyses were clustered by lineage using logistic regressions, only dispersal mechanism and flower size were significant predictors of risk and taxa with autochorous dispersal and conspicuous flowers were more likely to be at risk. The Big Island, Maui, Oahu, and Kauai all have remarkably similar numbers of dry forest taxa (63–65 species) and dry forest taxa at risk of endangerment. However, Big Island and Kauai have the highest number and percentage of single-island endemics. These results demonstrate patterns of endangerment specific to Hawaiian dry forests, the high levels of endangerment in this forest type, and the importance of prioritizing conservation in dry forest regions.     

Inferring speciation history in the Andes with reduced‐representation sequence data: an example in the bay‐backed antpittas (Aves; Grallariidae; Grallaria hypoleuca s. l.)

In the Andes, humid‐forest organisms frequently exhibit pronounced genetic structure and geographic variation in phenotype, often coincident with physical barriers to dispersal. However, phylogenetic relationships of clades have often been difficult to resolve due to short internodes. Consequently, even in taxa with well‐defined genetic structure, the temporal and geographic sequences of dispersal and vicariance events that led to this differentiation have remained opaque, hindering efforts to test the association between diversification and earth history and to understand the assembly of species‐rich communities on Andean slopes. Here, we use mitochondrial DNA and thousands of short‐read sequences generated with genotyping by sequencing (GBS) to examine the geographic history of speciation in a lineage of passerine birds found in the humid forest of the Andes, the ‘bay‐backed’ antpitta complex (Grallaria hypoleuca s. l). Mitochondrial DNA genealogies documented genetic structure among clade but were poorly resolved at nodes relevant for biogeographic inference. By contrast, relationships inferred from GBS loci were highly resolved and suggested a biogeographic history in which the ancestor originated in the northern Andes and dispersed south. Our results are consistent with a scenario of vicariant speciation wherein the range of a widespread ancestor was fragmented as a result of geologic or climatic change, rather than a stepping‐stone series of dispersal events across pre‐existing barriers. However, our study also highlights challenges of distinguishing dispersal‐mediated speciation from static vicariance. Our results further demonstrate the substantial evolutionary timescale over which the diverse biota of the Andes was assembled.     

Role of Upwelling on Larval Dispersal and Productivity of Gooseneck Barnacle Populations in the Cantabrian Sea: Management Implications

The effect of coastal upwelling on the recruitment and connectivity of coastal marine populations has rarely been characterized to a level of detail to be included into sound fishery management strategies. The gooseneck barnacle (Pollicipes pollicipes) fishery at the Cantabrian Coast (Northern Spain) is located at the fringes of the NW Spanish Upwelling system. This fishery is being co-managed through a fine-scale, interspersed set of protected rocks where each rock receives a distinct level of protection. Such interspersion is potentially beneficial, but the extent to which such spacing is consistent with mean larval dispersal distances is as yet unknown. We have simulated the spread of gooseneck barnacle larvae in the Central Cantabrian Coast using a high-resolution time-series of current profiles measured at a nearshore location. During a year of high upwelling activity (2009), theoretical recruitment success was 94% with peak recruitment predicted 56 km west of the emission point. However, for a year of low upwelling activity (2011) theoretical recruitment success dropped to 15.4% and peak recruitment was expected 13 km east of the emission point. This is consistent with a positive correlation between catch rates and the Integrated Upwelling Index, using a 4-year lag to allow recruits to reach commercial size. Furthermore, a net long-term westward larval transport was estimated by means of mitochondrial cytochrome c oxidase subunit I (COI) sequences for five populations in the Cantabrian Sea. Our results call into question the role of long distance dispersal, driven by the mesoscale processes in the area, in gooseneck barnacle populations and point to the prevalent role of small-scale, asymmetric connectivity more consistent with the typical scale of the co-management process in this fishery.     

Soil millipede diversity in tropical forest patches and its relation to landscape structure in northeastern Puerto Rico

Uncontrolled urban development is threatening the survival of many terrestrial species worldwide, especially those with limited dispersal capabilities. Soil invertebrates account for a high proportion of the global biodiversity but there are few studies on how soil biodiversity varies with different parameters of landscape structure, especially in fragmented tropical habitats. Millipedes are among the most abundant detritivore invertebrates in the soil-litter macrofauna. We examined the relationships between soil millipede diversity and landscape structure in 12 forest patches of approximately 30 years of age, in northeastern Puerto Rico. Spatial characteristics of the landscape were determined from aerial color photographs and were digitized into a GIS package for analysis. Millipede species diversity and composition in these forest patches showed correlations with their surroundings (e.g. amount of forest in the matrix) and with the presence of vegetation corridors that connected to other forest patches, rather than forest patch attributes such as patch area and shape. Millipede species richness correlated negatively with the degree of isolation of forest patch (within 600 m radius), while species evenness correlated positively to the amount of forest within a 50 m buffer. Millipede species composition was related with the presence of vegetation corridors and the distance to the Luquillo Experimental Forest reserve. These findings show that a low degree of patch isolation, forested buffers, and presence of vegetation corridors need to be considered for the conservation and management of forest patches surrounded by urban developments, especially to protect terrestrial invertebrate species that require forested habitats for their dispersal.     

Global patterns in marine dispersal estimates: the influence of geography, taxonomic category and life history

We examine estimates of dispersal in a broad range of marine species through an analysis of published values, and evaluate how well these values represent global patterns through a comparison with correlates of dispersal. Our analysis indicates a historical focus in dispersal studies on low-dispersal/low-latitude species, and we hypothesize that these studies are not generally applicable and representative of global patterns. Large-scale patterns in dispersal were examined using a database of correlates of dispersal such as planktonic larval duration (PLD, 318 species) and genetic differentiation (FST, 246 species). We observed significant differences in FST (p<0.001) and PLD (p<0.001) between taxonomic groups (e.g. fishes, cnidarians, etc.). Within marine fishes (more than 50% of datasets), the prevalence of demersal eggs was negatively associated with PLD (R2=0.80, p<0.001) and positively associated with genetic structure (R2=0.74, p<0.001). Furthermore, dispersal within marine fishes (i.e. PLD and FST) increased with latitude, adult body size and water depth. Of these variables, multiple regression identified latitude and body size as persistent predictors across taxonomic levels. These global patterns of dispersal represent a first step towards understanding and predicting species-level and regional differences in dispersal, and will be improved as more comprehensive data become available.     

Connectivity in the cold: the comparative population genetics of vent‐endemic fauna in the Scotia Sea,Southern Ocean

We report the first comparative population genetics study for vent fauna in the Southern Ocean using cytochrome C oxidase I and microsatellite markers. Three species are examined: the kiwaid squat lobster, Kiwa tyleri, the peltospirid gastropod, Gigantopelta chessoia, and a lepetodrilid limpet, Lepetodrilus sp., collected from vent fields 440 km apart on the East Scotia Ridge (ESR) and from the Kemp Caldera on the South Sandwich Island Arc, ~95 km eastwards. We report no differentiation for all species across the ESR, consistent with panmixia or recent range expansions. A lack of differentiation is notable for Kiwa tyleri, which exhibits extremely abbreviated lecithotrophic larval development, suggestive of a very limited dispersal range. Larval lifespans may, however, be extended by low temperature‐induced metabolic rate reduction in the Southern Ocean, muting the impact of dispersal strategy on patterns of population structure. COI diversity patterns suggest all species experienced demographic bottlenecks or selective sweeps in the past million years and possibly at different times. ESR and Kemp limpets are divergent, although with evidence of very recent ESR‐Kemp immigration. Their divergence, possibility indicative of incipient speciation, along with the absence of the other two species at Kemp, may be the consequence of differing dispersal capabilities across a ~1000 m depth range and/or different selective regimes between the two areas. Estimates of historic and recent limpet gene flow between the ESR and Kemp are consistent with predominantly easterly currents and potentially therefore, cross‐axis currents on the ESR, with biogeographic implications for the region.     

Isolation by distance,not rivers,control the distribution of termite species in the Amazonian rain forest

The spatial distribution of species is affected by dispersal barriers, local environmental conditions and climate. However, the effect of species dispersal and their adaptation to the environment across geographic scales is poorly understood. To investigate the distribution of species from local to broad geographic scales, we sampled termites in 198 transects distributed in 13 sampling grids in the Brazilian Amazonian forest. The sampling grids encompassed an area of 271 500 km² and included the five major biogeographic regions delimited by Amazonian rivers. Environmental data for each transect were obtained from local measurements and remote sensing. Similar to previous studies, termite species composition at the local scale was mostly associated with measures of soil texture and chemistry. In contrast, termite species composition at broad geographic scales was associated with soil nutrients, and the geographic position of the transects. Between 17 and 30% of the variance in termite species composition could be attributed exclusively to the geographic position of the transects, but could not be attributed to measured environmental variables or the presence of major rivers. Isolation by distance may have strong effects on termite species composition due to historic processes and the spatially structured environments along distinct geological formations of Amazonia. However, in contrast to many taxa in Amazonia, there is no evidence that major rivers are important barriers to termite dispersal.     

Vertebrate dispersal syndromes along the Atlantic forest: broad-scale patterns and macroecological correlates

Aim To assess the geographical variation in the relative importance of vertebrates, and more specifically of birds and mammals, as seed dispersal agents in forest communities, and to evaluate the influence of geographical and climatic factors on the observed trends. Location One hundred and thirty‐five forest communities in the Brazilian Atlantic forest. Methods We collected data on dispersal modes for 2292 woody species. By combining species × site with species × trait matrices, we obtained the percentages of endozoochory, ornithochory, mastozoochory and the mean fruit diameter for the local forest communities. We used Spearman's correlation to assess bivariate relationships between variables. Subsequently, we performed paired t‐tests to verify if variations in frequency of dispersal modes and mean fruit diameter were influenced by altitude or temperature. Then, we applied multiple linear regressions to evaluate the effect of geographical and climatic variables on variation in the relative frequency of dispersal modes and mean fruit diameter across communities. Results We found no consistent latitudinal or longitudinal trend in the percentage of vertebrate‐dispersed species, neither bird‐ nor mammal‐dispersed species along the Atlantic forest. Endozoochory was affected chiefly by annual mean rainfall, increasing towards moister sites. Forest communities located at higher altitudes had a higher percentage of bird‐dispersed species. Even when sites with identical values of annual mean temperature were compared, altitude had a positive effect on ornithochory. Conversely, we found a higher percentage of mammal‐dispersed species in warmer forests, even when locations at the same altitudinal belts were contrasted. Fruit diameter was clearly related to altitude, decreasing towards higher elevations. Main conclusions This is the first analysis of a large data set on dispersal syndromes in tropical forest communities. Our findings support the hypotheses that: (1) geographical variation in the relative number of fleshy fruit species is mainly driven by moisture conditions and is relatively independent of geographical location, and (2) broad‐scale trends in fruit size correspond to geographical variation in the relative importance of mammals and birds as seed dispersal agents at the community level.     

The relationship between geographic range size and life history traits: is biogeographic history uncovered? A test using the Iberian butterflies

The geographic range of a species is influenced by past phylogenetic and biogeographic patterns. However, other historical interactions, including the interplay between life history and geography, are also likely involved. Therefore, the range size of a species can be explained on the basis of niche‐breadth or dispersal related hypotheses, and previous work on European butterflies suggests that both, under the respective guise of ecological specialisation and colonising ability may apply. In the present study, data from 205 species of butterflies from the Iberian peninsula were processed through multiple regression analyses to test for correlations between geographic range size, life history traits and geographic features of the species distribution types. In addition, the percentage of variance explained by the subsets of variables analyzed in the study, with and without control for phylogenetic effects was tested. Despite a complex pattern of bivariate correlations, we found that larval polyphagy was the single best correlate of range size, followed by dispersal. Models that combined both life history traits and geographic characteristics performed better than models generated independently. The combined variables explained at least 39% of the variance. Bivariate correlations between range size and body size, migratory habits or egg size primarily reflected taxonomic patterning and reciprocal correlations with larval diet breadth and adult phenology. Therefore, aspects of niche breadth i.e. potential larval diet breadth emerged as the most influential determinants of range size. However, the relationships between these types of ecological traits and biogeographic history must still be considered when associations between life history and range size are of interest.     

Predominant east to west colonizations across major oceanic barriers: Insights into the phylogeographic history of the hydroid superfamily Plumularioidea,suggested by a mitochondrial DNA barcoding marker

We provide preliminary insights into the global phylogeographic and evolutionary patterns across species of the hydrozoan superfamily Plumularioidea (Cnidaria: Hydrozoa). We analyzed 1,114 16S sequences of 198 putative species of Plumularioidea collected worldwide. We investigated genetic connections and divergence in relation to present‐day and ancient biogeographic barriers, climate changes and oceanic circulation. Geographical distributions of most species are generally more constrained than previously assumed. Some species able to raft are dispersed widely. Human‐mediated dispersal explains some wide geographical ranges. Trans‐Atlantic genetic connections are presently unlikely for most of the tropical‐temperate species, but were probably more frequent until the Miocene–Pliocene transition, before restriction of the Tethys Sea and the Central American Seaway. Trans‐Atlantic colonizations were predominantly directed westwards through (sub)tropical waters. The Azores were colonized multiple times and through different routes, mainly from the east Atlantic, at least since the Pliocene. Extant geminate clades separated by the Isthmus of Panama have predominantly Atlantic origin. Various ancient colonizations mainly directed from the Indian Ocean to the Atlantic occurred through the Tethys Sea and around South Africa in periods of lower intensity of the Benguela upwelling. Thermal tolerance, population sizes, dispersal strategies, oceanic currents, substrate preference, and land barriers are important factors for dispersal and speciation of marine hydroids.     

Population structure is not a simple function of reproductive mode and larval type: insights from tropical corals

1. For a wide range of organisms, heritable variation in life-history characteristics has been shown to be strongly subject to selection, reflecting the impact that variation in characters such as genotypic diversity, duration of larval development and adaptations for dispersal can have on the fitness of offspring and the make-up of populations. Indeed, variation in life-history characteristics, especially reproduction and larval type, have often been used to predict patterns of dispersal and resultant population structures in marine invertebrates. 2. Scleractinian corals are excellent models with which to test this relationship, as they exhibit almost every possible combination of reproductive mode and larval type. Some general patterns are emerging but, contrary to expectations, genetic data suggest that while populations of broadcast spawning species may be genotypically diverse they may be heavily reliant on localized recruitment rather than widespread dispersal of larvae. 3. Here we use microsatellites to test the importance of localized recruitment by comparing the genetic structure of populations of two broadcast spawning corals with contrasting modes of reproduction and larval development; Goniastrea favulus is self-compatible, has sticky, negatively buoyant eggs and larvae and is expected to have restricted dispersal of gametes and larvae. In contrast, Platygyra daedalea is self-incompatibile, spawns positively buoyant egg-sperm bundles and has planktonic development. 4. Surprisingly, spatial-autocorrelation revealed no fine-scale clustering of similar genotypes within sites for G. favulus, but showed a non-random distribution of genotypes in P. daedalea. Both species showed similar levels of genetic subdivision among sites separated by 50-100 m (F(ST) = 0.03), suggesting that larval dispersal may be equivalent in both species. 5. Interestingly, as fragmentation has been considered rare in massive corals, our sample of 284 P. daedalea colonies included 28 replicated genotypes that were each unlikely (P < 0.05) to have been derived independently from sexual reproduction. 6. We conclude that the extreme life history of G. favulus does not produce unusually fine-scale genetic structure and subsequently, that reproductive mode and larval type may not be not good predictors of population structure or dispersal ability.     

Larval dispersal dampens population fluctuation and shapes the interspecific spatial distribution patterns of rocky intertidal gastropods

Many marine benthic invertebrates pass through a planktonic larval stage whereas others spend their entire lifetimes in benthic habitats. Recent studies indicate that non‐planktonic species show relatively greater fine‐scale patchiness than do planktonic species, but the underlying mechanisms remain unknown. One hypothesis for such a difference is that larval dispersal enhances the connectivity of populations and buffers population fluctuations and reduces local extinction risk, consequently increasing patch occupancy rate and decreasing spatial patchiness. If this mechanism does indeed play a significant role, then the distribution of non‐planktonic species should be more aggregated – both temporally and spatially – than the distribution of species with a planktonic larval stage. To test this prediction, we compared 1) both the spatial and the temporal abundance–occupancy relationships and 2) both the spatial and the temporal mean–variance relationships of population size across species of rocky intertidal gastropods with differing dispersive traits from the Pacific coast of Japan. We found that, compared to planktonic species, non‐planktonic species exhibited 1) a smaller occupancy rate for any given level of mean population size and 2) greater variations in population size, both spatially and temporally. This suggests that the macroecological patterns observed in this study (i.e. the abundance–occupancy relationships and mean–variance relationships of population size across species) were shaped by the effect of larval dispersal dampening population fluctuation, which works over both space and time. While it has been widely assumed that larval dispersal enhances population fluctuations, larval dispersal may in fact enhance the connectively of populations and buffer population fluctuations and reduce local extinction risks.