Flying shells: historical dispersal of marine snails across Central America

The geological rise of the Central American Isthmus separated the Pacific and the Atlantic oceans about 3 Ma, creating a formidable barrier to dispersal for marine species. However, similar to Simpson's proposal that terrestrial species can 'win sweepstakes routes'-whereby highly improbable dispersal events result in colonization across geographical barriers-marine species may also breach land barriers given enough time. To test this hypothesis, we asked whether intertidal marine snails have crossed Central America to successfully establish in new ocean basins. We used a mitochondrial DNA genetic comparison of sister snails (Cerithideopsis spp.) separated by the rise of the Isthmus. Genetic variation in these snails revealed evidence of at least two successful dispersal events between the Pacific and the Atlantic after the final closure of the Isthmus. A combination of ancestral area analyses and molecular dating techniques indicated that dispersal from the Pacific to the Atlantic occurred about 750 000 years ago and that dispersal in the opposite direction occurred about 72 000 years ago. The geographical distribution of haplotypes and published field evidence further suggest that migratory shorebirds transported the snails across Central America at the Isthmus of Tehuantepec in southern Mexico. Migratory birds could disperse other intertidal invertebrates this way, suggesting the Central American Isthmus may not be as impassable for marine species as previously assumed.     

Phylogeography of Middle American gophersnakes: mixed responses to biogeographical barriers across the Mexican Transition Zone

Aim We used inferences of phylogeographical structure and estimates of divergence times for three species of gophersnakes (Colubridae: Pituophis) distributed across the Mexican Transition Zone (MTZ) to evaluate the postulated association of three Neogene geological events (marine seaway inundation of the Isthmus of Tehuantepec, formation of the Transvolcanic Belt across central Mexico, and secondary uplifting of the Sierra Madre Occidental) and of Pleistocene climate change with inter‐ and intraspecific diversification. Location Mexico, Guatemala, and the western United States. Methods We combined range‐wide sampling (67 individuals representing three putative species distributed across northern Middle America and western North America) and phylogenetic analyses of 1637 base pairs of mitochondrial DNA to estimate genealogical relationships and divergence times. The hypothesized concordance of inferred gene trees with geological histories was assessed using topology tests. Results We identified three major lineages of Middle American gophersnakes, and strong phylogeographical structure within each lineage. Gene trees were statistically congruent with hypothesized geological histories for two of the three postulated geological events. Estimated divergence dates and the geographical distribution of genetic variation further support mixed responses to these geological events. Considerable phylogeographical structure appears to have been generated during the Pleistocene. Main conclusions Phylogenetic and phylogeographical structure in gophersnakes distributed across northern Middle America and western North America highlights the influence of both Neogene vicariance events and Pleistocene climate change in shaping genetic diversity in this region. Despite the presence of two major geographical barriers in southern Mexico, extreme geological and environmental heterogeneity in this area may have differentially structured genetic diversity in highland taxa. To the north, co‐distributed taxa may display a more predictable pattern of diversification across the warm desert regions. Future studies should incorporate nuclear data to disentangle inferred lineage boundaries and further elucidate patterns of mitochondrial introgression.     

Patterns of morphological and ecological similarities of small-eared shrews (Soricidae,Cryptotis) in tropical montane cloud forests from Mesoamerica

It has been proposed that high morphological similarity between closely related species of small-eared shrews resulted from a recent divergence and intermittent population connectivity, presumably due to Pleistocene climatic fluctuations and associated changes in forest habitat distribution. Here we examined the morphological variation of two sister species of small-eared shrews inhabiting cloud forests from Mexico, Cryptotis obscurus and C. mexicanus. We then used ecological niche modelling to provide compelling evidence for current environmental barriers for population connectivity, and for detecting divergent ecological niches between candidate species. Our results indicated that the species boundaries in this clade should be subject to change. High morphological similarity suggested that populations of C. obscurus and C. mexicanus located west of the Isthmus of Tehuantepec, a major geographic barrier for montane species, are conspecific. Niche divergence between these two putative species was not supported indicating niche conservatism across the evolutionary history of these small-eared shrews. In addition, several barriers seem to play a main role for current lineage divergence between populations within this clade. The population located east of the Isthmus, previously referred to C. mexicanus, might prove to represent a new species based on morphological distinction and current geographic isolation. We have highlighted that estimating species’ potential distributions provides insights to evaluating the effect of geographic barriers on lineage divergence and making stronger inferences when delimiting species.     

Complex phylogeographic patterns indicate Central American origin of two widespread Mesoamerican <Emphasis Type="Italic">Quercus</Emphasis> (Fagaceae) species

The northern Neotropical region is characterized by a heterogeneous geological and climatic history. Recent studies have shown contrasting patterns regarding the role of geographic elements as barriers that could have determined phylogeographic structure in various species. Recently, the phylogeography and biogeography of Quercus species have been studied intensively, and the patterns observed so far suggest contrasting evolutionary histories for Neotropical species in comparison with their Holarctic relatives. The goal of this study was to describe the phylogeographic structure of two Neotropical oak species (Quercus insignis and Quercus sapotifolia) in the context of the geological and palaeoclimatic history of the northern Neotropics. Populations through the distribution range of both species were collected and characterized using nine chloroplast DNA microsatellite loci. Both oak species showed high levels of genetic diversity and strong phylogeographic structure. The distribution of genetic variation in Q. insignis suggested an influence of two major barriers, the Isthmus of Tehuantepec and the Nicaraguan Depression, while Q. sapotifolia exhibited a genetic structure defined by the heterogeneity of the Chortis highlands. The haplotype networks of both species indicated complex histories, suggesting that colonization from the Sierra Madre de Chiapas to central Mexico and from the north of the Nicaraguan Depression to the Costa Rican mountains may have occurred during different stages, and apparently more than one time. In conclusion, the phylogeographic structure of Neotropical oak species seems to be defined by a combination of geological and climatic events.     

Eukaryotic microbes, species recognition and the geographic limits of species: examples from the kingdom Fungi

The claim that eukaryotic micro-organisms have global geographic ranges, constituting a significant departure from the situation with macro-organisms, has been supported by studies of morphological species from protistan kingdoms. Here, we examine this claim by reviewing examples from another kingdom of eukaryotic microbes, the Fungi. We show that inferred geographic range of a fungal species depends upon the method of species recognition. While some fungal species defined by morphology show global geographic ranges, when fungal species are defined by phylogenetic species recognition they are typically shown to harbour several to many endemic species. We advance two non-exclusive reasons to explain the perceived difference between the size of geographic ranges of microscopic and macroscopic eukaryotic species when morphological methods of species recognition are used. These reasons are that microbial organisms generally have fewer morphological characters, and that the rate of morphological change will be slower for organisms with less elaborate development and fewer cells. Both of these reasons result in fewer discriminatory morphological differences between recently diverged lineages. The rate of genetic change, moreover, is similar for both large and small organisms, which helps to explain why phylogenetic species of large and small organisms show a more similar distribution of geographic ranges. As a consequence of the different rates in fungi of genetic and morphological changes, genetic isolation precedes a recognizable morphological change. The final step in speciation, reproductive isolation, also follows genetic isolation and may precede morphological change.     

Recovering cryptic diversity and ancient drainage patterns in eastern North America: historical biogeography of the Notropis rubellus species group (Teleostei: Cypriniformes)

The Central Highlands of North America contain a strikingly diverse assemblage of temperate freshwater fishes and have long been a focus of biogeographic studies. The rosyface shiner complex, Notropis rubellus and related species, is a member of this fauna exhibiting a disjunct highlands distribution occurring in the unglaciated regions of the Central Highlands and glaciated regions of the Central Lowlands. Until recently, N. rubellus was considered a single, widespread species exhibiting geographic variation in morphological characters. However, several studies have revealed that N. rubellus is a multi-species complex with closely related species endemic to drainages within each highland region. We examined genetic variation of the N. rubellus complex using a complete mtDNA cytochrome b gene sequence data set and combined mtDNA and published allozyme data sets. Parsimony and Bayesian analyses of the mitochondrial data set and parsimony analyses of a combined mitochondrial and allozyme data sets were largely consistent. Results of these analyses revealed ancient cryptic diversity within the N. rubellus complex that existed prior to the onset of Pleistocene glaciations. We identified seven strongly supported clades within the N. rubellus complex. Four clades are diagnosed as separate species (N. percobromus, N. rubellus, N. micropteryx and N. suttkusi) and three clades may represent undescribed forms. Relationships among these groups and their biogeographical patterns provided significant inferences on ichthyofaunal distributions in southeastern North America. These include the timing of the origin of the diversity, ancient drainage patterns and barriers to dispersal in the Central Highlands. The observation of increased diversity in N. rubellus suggests there may be greater diversity within other taxa with a similar distribution.     

The Necessity of DNA Taxonomy to Reveal Cryptic Diversity and Spatial Distribution of Meiofauna,with a Focus on Nemertea

Meiofauna represent one of the most abundant and diverse communities in marine benthic ecosystems. However, an accurate assessment of diversity at the level of species has been and remains challenging for these microscopic organisms. Therefore, for many taxa, especially the soft body forms such as nemerteans, which often lack clear diagnostic morphological traits, DNA taxonomy is an effective means to assess species diversity. Morphological taxonomy of Nemertea is well documented as complicated by scarcity of unambiguous character states and compromised by diagnoses of a majority of species (and higher clades) being inadequate or based on ambiguous characters and character states. Therefore, recent studies have advocated for the primacy of molecular tools to solve the taxonomy of this group. DNA taxonomy uncovers possible hidden cryptic species, provides a coherent means to systematize taxa in definite clades, and also reveals possible biogeographic patterns. Here, we analyze diversity of nemertean species by considering the barcode region of the mitochondrial gene Cytochrome Oxidase subunit I (COI) and different species delineation approaches in order to infer evolutionarily significant units. In the aim to uncover actual diversity of meiofaunal nemerteans across different sites in Central America, COI sequences were obtained for specimens assigned here to the genera Cephalothrix, Ototyphlonemertes, and Tetrastemma-like worms, each commonly encountered in our sampling. Additional genetic, taxonomic, and geographic data of other specimens belonging to these genera were added from GenBank. Results are consistent across different DNA taxonomy approaches, and revealed (i) the presence of several hidden cryptic species and (ii) numerous potential misidentifications due to traditional taxonomy. (iii) We additionally test a possible biogeographic pattern of taxonomic units revealed by this study, and, except for a few cases, the putative species seem not to be widely distributed, in contrast to what traditional taxonomy would suggest for the recognized morphotypes.     

Evaluating connectivity in the brooding brittle star Astrotoma agassizii across the drake passage in the Southern Ocean

Studies examining population structure and genetic diversity of benthic marine invertebrates in the Southern Ocean have emerged in recent years. However, many taxonomic groups remain largely unstudied, echinoderms being one conspicuous example. The brittle star Astrotoma agassizii is distributed widely throughout Antarctica and southern South America. This species is a brooding echinoderm and therefore may have limited dispersal capacity. In order to determine the effect of hypothesized isolating barriers in the Southern Ocean, such as depth, geographic distance, and the polar front, 2 mitochondrial DNA markers were used to compare populations from the South American and Antarctic continental shelves. Astrotoma agassizii was shown to be genetically discontinuous across the polar front. In fact, populations previously assumed to be panmictic instead represent 3 separate lineages that lack morphological distinction. However, within lineages, genetic continuity was displayed across a large geographic range (>500 km). Therefore, despite lacking a pelagic larval stage, A. agassizii can disperse across substantial geographic distance within continental shelf regions. These results indicate that geographic distance alone may not be a barrier to dispersal, but rather the combined effects of distance, depth, and the polar front act to prevent gene flow between A. agassizii populations in the Southern Ocean.     

Cryptic species in tropic sands--interactive 3D anatomy, molecular phylogeny and evolution of meiofaunal Pseudunelidae (Gastropoda, Acochlidia)

Background

Towards realistic estimations of the diversity of marine animals, tiny meiofaunal species usually are underrepresented. Since the biological species concept is hardly applicable on exotic and elusive animals, it is even more important to apply a morphospecies concept on the best level of information possible, using accurate and efficient methodology such as 3D modelling from histological sections. Molecular approaches such as sequence analyses may reveal further, cryptic species. This is the first case study on meiofaunal gastropods to test diversity estimations from traditional taxonomy against results from modern microanatomical methodology and molecular systematics.

Results

The examined meiofaunal Pseudunela specimens from several Indo-Pacific islands cannot be distinguished by external features. Their 3D microanatomy shows differences in the organ systems and allows for taxonomic separation in some cases. Additional molecular analyses based on partial mitochondrial cytochrome c oxidase subunit I (COI) and 16S rRNA markers revealed considerable genetic structure that is largely congruent with anatomical or geographical patterns. Two new species (Pseudunela viatoris and P. marteli spp. nov.) are formally described integrating morphological and genetic analyses. Phylogenetic analysis using partial 16S rRNA, COI and the nuclear 18S rRNA markers shows a clade of Pseudunelidae species as the sister group to limnic Acochlidiidae. Within Pseudunela, two subtypes of complex excretory systems occur. A complex kidney already evolved in the ancestor of Hedylopsacea. Several habitat shifts occurred during hedylopsacean evolution.

Conclusions

Cryptic species occur in tropical meiofaunal Pseudunela gastropods, and likely in other meiofaunal groups with poor dispersal abilities, boosting current diversity estimations. Only a combined 3D microanatomical and molecular approach revealed actual species diversity within Pseudunela reliably. Such integrative methods are recommended for all taxonomic approaches and biodiversity surveys on soft-bodied and small-sized invertebrates. With increasing taxon sampling and details studied, the evolution of acochlidian panpulmonates is even more complex than expected.     

The genetic structure of fish: differences in the intraspecific distribution of biochemical genetic variation between marine, anadromous, and freshwater species

Biochemical genetic markers have been increasingly used for inferences on the population genetic structure of various fish, but little attention has been paid to relative differences in the distribution of variability within species in different groups of fish. Nested gene diversity analyses of multiple locus electrophoretic data from the literature for 19 species (6 marine species, 4 anadromous salmonids, and 9 freshwater species (5 salmonids)), mainly from Scandinavia and North America, show that (i) average heterozygosity is higher in marine species than in freshwater species, (ii) the average fraction of the total gene diversity allocated between localities increases in the order marine (1.6%), anadromous (3.7%), and freshwater species (29.4%), and (iii) the fraction of the total gene diversity associated with differences between geographic clusters of localities is generally smaller than that between localities within clusters. This finding is consistent with that the differentiation observed in most species is of postglacial origin, and that the timespan available since deglaciation has been too short to result in the accumulation of substantial differences between clusters.     

Phylogeography of a morphologically diverse Neotropical montane species, the Common Bush-Tanager (Chlorospingus ophthalmicus)

The Common Bush-Tanager (Chlorospingus ophthalmicus) is distributed in Neotropical cloud-forests from Mexico to Argentina and contains 25 subspecies divided into eight subspecies groups based on biogeography, eye coloration, presence of a postocular spot and chest band. All of Central America is occupied by a single subspecies group; whereas the Andes are believed to be occupied by seven additional subspecies groups. We used five mitochondrial genes to investigate the phylogeography and possible species limits of the ophthalmicus complex. A total of 14 monophyletic lineages were uncovered within the ophthalmicus complex, including three clades currently classified as separate species (C. semifuscus, inornatus and tacarcunae). Divergence estimates for these clades date between 0.8 and 5.2 million years ago (Ma). Contrary to expectations based on morphological diversity, phylogeographic structure was greatest in Mexico and Central America and weakest in the Andes. Morphological and genetic divergences were not significantly correlated and most morphologically defined subspecies groups were not supported. Our evidence suggests the ophthalmicus complex originated in Mexico ca. 6.0 Ma (million years ago) and spread south into the Andes ca. 4.7 Ma before the completion of the Isthmus of Panama. Three genetically divergent lineages of ophthalmicus that formed in the Andes possess a complex checkerboard distribution, with a single lineage represented by disjunct populations from Venezuela and the southern Andes, while intervening populations in Ecuador and Central Peru form two genetically and morphologically divergent lineages.     

Seaweeds, latitudinal diversity patterns, and Rapoport's Rule

We analysed the geographic distribution of 645 species of marine benthic algae along the Atlantic coast of Europe and Pacific coast of temperate South America to test for the existence of an association between geographic range size and latitude (Rapoport's Rule) and for three key components of the explanations offered for it. We found that species in high diversity areas are characterized by small geographic ranges and by low specific growth rates as compared to species with large geographic ranges, thus supporting the Rapoport-rescue hypothesis. However, the pattern is not related to species' tolerances, to abiotic conditions or to climatic variability. Further, the inverse latitudinal diversity pattern shown by the marine algal flora of temperate Pacific South America, and the opposite patterns shown by tropical and subantarctic species within this flora, stressed that small geographic ranges are linked to high diversity areas in general, and not only in relation to the pole to tropic species diversity gradient.     

Evolutionary history of the northern leopard frog: reconstruction of phylogeny, phylogeography, and historical changes in population demography from mitochondrial DNA

This study uses a combined methodological approach including phylogenetic, phylogeographic, and demographic analyses to understand the evolutionary history of the northern leopard frog, Rana pipiens. We tested hypotheses concerning how (or if) known geological events and key features of the species biology influenced the contemporary geographic and genetic distribution of R. pipiens. We assayed mitochondrial DNA variation from 389 individuals within 35 populations located throughout the species range. Our a priori expectations for patterns and processes influencing the current genetic structure of R. pipiens were supported by the data. However, our analyses revealed specific aspects of R. pipiens evolutionary history that were unexpected. The phylogenetic analysis indicated that R. pipiens is split into populations containing discrete eastern or western haplotypes, with the Mississippi River and Great Lakes region dividing the geographic ranges. Nested clade analysis indicated that the biological process most often invoked to explain the pattern of haplotype position is restricted gene flow with isolation by distance. Demographic analyses showed evidence of both historical bottlenecks and population expansions. Surprisingly, the genetic evidence indicated that the western haplotypes had significantly reduced levels of genetic diversity relative to the eastern haplotypes and that major range expansions occurred in both regions well before the most recent glacial retreat. This study provides a detailed history of how a widespread terrestrial vertebrate responded to episodic Pleistocene glacial events in North America. Moreover, this study illustrates how complementary methods of data analysis can be used to disentangle recent and ancient effects on the genetic structure of a species.     

Population genetics and geometric morphometrics of the <Emphasis Type="Italic">Bombus ephippiatus</Emphasis> species complex with implications for its use as a commercial pollinator

Mexico and Central America are among the most biodiverse regions on Earth, harboring many species with high levels of interpopulation morphological and genetic diversity. The mountainous topography of this region contains isolated sky island habitats that have the potential to promote speciation. This has been studied in vertebrates, yet few studies have examined the phylogeographic and genetic structure of insect species encompassing this region. Here we investigate geographic patterns of genetic and morphological divergence and speciation among widespread populations of the highly polymorphic bumble bee Bombus ephippiatus and its closest relative B. wilmattae. We used DNA sequences from a fragment of cytochrome oxidase I (COI), genotypes for twelve microsatellite markers, and morphometric data from wings to construct a well-supported inference of the divergences among these taxa. We have found complex patterns of genetic isolation and morphological divergence within B. ephippiatus across its geographic range and present evidence that B. ephippiatus comprises multiple independent evolutionary lineages. The pattern of their diversification corresponds to geographic and environmental isolating mechanisms, including the Mexican highlands, the lowlands of the Isthmus of Tehuantepec in southern Mexico, the Nicaraguan Depression, the patchily distributed volcanic ranges in Nuclear Central America and Pleistocene glacial cycles. These results have important implications for the development and distribution of B. ephippiatus as a commercial pollinator in Mexico and Central America.     

Genetic and phenotypic variation in a colourful treefrog across five geographic barriers

Aim Understanding the patterns and processes underlying phenotype in a polytypic species provides key insights into microevolutionary mechanisms of diversification. The red‐eyed treefrog, Agalychnis callidryas, exhibits strong regional differentiation in colour pattern, corresponding to five admixed mitochondrial DNA clades. We evaluated spatial diversity patterns across multiple, putative barriers to examine the fine‐scale processes that mediate phenotypic divergence between some regions while maintaining homogeneity between others. Location We examined patterns of phenotypic diversification among 17 sites that span five putative biogeographic barriers in lower Central America (Costa Rica and Panama). Methods We tested the extent to which genetic distance (F_ST) derived from six multilocus nuclear genotypes covaried with measures of phenotypic distance (leg coloration) within and between biogeographic regions. We used linear regression analyses to determine the role of geographic and genetic factors in structuring spatial patterns of phenotypic diversity. Results The factors that best explained patterns of phenotypic diversity varied among biogeographic regions. We identified one geographic barrier that impeded gene exchange and resulted in concordant phenotypic divergence across the Continental Divide, isolating Caribbean and Pacific populations. Across Caribbean Costa Rican populations, one barrier structured phenotypic but not genetic diversity patterns, indicating a role for selection. In other regions, the putative barriers had no determining effect on either genetic or leg colour structure. Main conclusions The processes mediating the distribution and diversification of colour pattern in this polytypic, wide‐ranging treefrog varied among biogeographic regions. Spatially varying selection combined with the isolating effects of geographic factors probably resulted in the patchy distribution of colour diversity across Costa Rican and Panamanian populations.     

Avian host composition,local speciation and dispersal drive the regional assembly of avian malaria parasites in South American birds

Identifying the ecological factors that shape parasite distributions remains a central goal in disease ecology. These factors include dispersal capability, environmental filters and geographic distance. Using 520 haemosporidian parasite genetic lineages recovered from 7,534 birds sampled across tropical and temperate South America, we tested (a) the latitudinal diversity gradient hypothesis and (b) the distance–decay relationship (decreasing proportion of shared species between communities with increasing geographic distance) for this host–parasite system. We then inferred the biogeographic processes influencing the diversity and distributions of this cosmopolitan group of parasites across South America. We found support for a latitudinal gradient in diversity for avian haemosporidian parasites, potentially mediated through higher avian host diversity towards the equator. Parasite similarity was correlated with climate similarity, geographic distance and host composition. Local diversification in Amazonian lineages followed by dispersal was the most frequent biogeographic events reconstructed for haemosporidian parasites. Combining macroecological patterns and biogeographic processes, our study reveals that haemosporidian parasites are capable of circumventing geographic barriers and dispersing across biomes, although constrained by environmental filtering. The contemporary diversity and distributions of haemosporidian parasites are mainly driven by historical (speciation) and ecological (dispersal) processes, whereas the parasite community assembly is largely governed by host composition and to a lesser extent by environmental conditions.     

The description of a new species of the Neotropical land crab genus Gecarcinus Leach, 1814 (Crustacea,Decapoda, Brachyura,Gecarcinidae)

In this contribution a new species of the land crab genus Gecarcinus Leach, 1814, from the Neotropical Pacific coast of South America is described and illustrated. In addition to its unique body color, Gecarcinus nobilii sp. n. is distinguished from congeners by a distinctly wider carapace front and differences in the shape of the infraorbital margin. The new species is not isolated from Gecarcinus populations from the Pacific coast of Central America by an insurmountable geographic barrier. Considering the closure of the Panamanian Isthmus as a calibration point for morphological divergence between the trans-isthmian mainland populations of Gecarcinus, the virtual lack of morphological differentiation (other than color) between them and the distinctness of G. nobilii sp. n. suggests that G. nobilii sp. n. evolved from a common ancestor before the Isthmus closed.     

Placing the Floridian marine genetic disjunction into a regional evolutionary context using the scorched mussel, Brachidontes exustus, species complex

The well-documented Floridian Gulf/Atlantic marine genetic disjunction provides an influential example of presumed vicariant cladogenesis along a continental coastline for major elements of a diverse nearshore fauna. However, it is unclear if this disjunction represents a local anomaly for regionally distributed morphospecies, or if it is merely one of many such cryptic phylogenetic splits that underlay their assumed genetic cohesiveness. We aimed to place the previously characterized scorched mussel Gulf/Atlantic genetic disjunction into a regional phylogenetic perspective by incorporating genotypes of nominal conspecifics sampled throughout the Caribbean Basin as well as those of eastern Pacific potential geminate species. Our results show it to be one of multiple latent regional genetic disjunctions, involving five cryptic Caribbean species, that appear to be the product of a long history of regional cladogenesis. Disjunctions involving three stem lineages clearly predate formation of the Isthmus of Panama and of the Caribbean Sea, although four of the five cryptic species have within-basin sister relationships. Surprisingly, the Atlantic clade was also found to be widespread in the southern Caribbean, and ancestral demography calculations through time for Atlantic coast-specific genotypes are consistent with a northward range extension after the last glacial maximum. Our new data seriously undermine the hypothesis of a Floridian vicariant genesis and imply that the scorched mussel Gulf/Atlantic disjunction represents a case of geographic and temporal pseudocongruence. All five Caribbean Basin cryptic species exhibited an intriguing pattern of predominantly allopatric distribution characterized by distinct geographic areas of ecological dominance, often adjoining those of sister taxa. This pattern of distribution is consistent with allopatric speciation origins, coupled with restricted postspeciation range extensions. Several lines of indirect evidence favor the hypothesis that the predominantly allopatric distributions are maintained over evolutionary time scales, primarily by postrecruitment ecological filters rather than by oceanographic barriers to larval-mediated gene flow.     

Polymorphism hides cryptic species in Oerstedia dorsalis (Nemertea, Hoplonemertea)

The marine hoplonemertean Oerstedia dorsalis is considered to be a highly polymorphic species with extensive geographic distribution. We show, based on mitochondrial cytochrome c oxidase subunit I, 16S and nuclear internal transcribed spacer sequences, that there are genetic subsets withing this species. Seventy-one specimens of various colours from different geographic localities (in Europe) were sequenced and analysed using statistical parsimony and Bayesian analysis. Both analyses supported nine major clades. We conclude that O. dorsalis hides different species with geographic resolution. These species, however, appear to be polymorphic as well, and we find no diagnostic features in pigmentation or external characters to separate species within this complex.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 556–567.