Phylogeographic analysis of the brooding brittle star Amphipholis squamata (Echinodermata) along the coast of New Zealand reveals high cryptic genetic variation and cryptic dispersal potential.

Direct development in benthic marine invertebrates is usually associated with narrow geographical range, low rates of colonization, and low levels of gene flow. Paradoxically, the small brittle star Amphipholis squamata broods its larvae to a crawl-away juvenile stage, yet has a cosmopolitan distribution. Using sequence and restriction-fragment-length-polymorphisms (RFLP) analyses of nuclear and mitochondrial DNA from 16 coastal populations throughout New Zealand, we tested whether the species is indeed a poor disperser, as may be expected from its brooding habit. We predicted that local and regional populations would be genetically structured according to isolation by distance. We also suspected that this ubiquitous "species" is composed of a variety of cryptic taxa in different geographic areas, as has been discovered in an increasing number of marine invertebrates. We found evidence of four genetically divergent and reproductively isolated lineages that can exist in syntopy. Lineages vary in abundance, haplotype diversity, and geographic distribution. The partitioning of genetic variation within the most common lineage, as well as the geographic distribution of the four lineages, suggest a north/south split. This pattern is consistent with known New Zealand marine biogeographic zones and appears to be linked to the regime of oceanic circulation, which is characterized by subtropical, southward-moving water masses in the north, and sub-Antarctic, northward-moving water in the south. We conclude that the dispersal ability of A. squamata is regionally restricted but with sporadic long-distance dispersal, which serves to increase local genetic variation. Our results support the idea that dispersal occurs through passive transport by drifting or rafting on macroalgae, which A. squamata commonly inhabits, and emphasize that poor dispersal ability is not necessarily a corollary of direct development.

Out of Africa: the slow train to australasia.

We used mitochondrial DNA (mtDNA) sequences to test biogeographic hypotheses for Patiriella exigua (Asterinidae), one of the world's most widespread coastal sea stars. This small intertidal species has an entirely benthic life history and yet occurs in southern temperate waters of the Atlantic, Indian, and Pacific oceans. Despite its abundance around southern Africa, southeastern Australia, and several oceanic islands, P. exigua is absent from the shores of Western Australia, New Zealand, and South America. Phylogenetic analysis of mtDNA sequences (cytochrome oxidase I, control region) indicates that South Africa houses an assemblage of P. exigua that is not monophyletic (P = 0.04), whereas Australian and Lord Howe Island specimens form an interior monophyletic group. The placement of the root in Africa and small genetic divergences between eastern African and Australian haplotypes strongly suggest Pleistocene dispersal eastward across the Indian Ocean. Dispersal was probably achieved by rafting on wood or macroalgae, which was facilitated by the West Wind Drift. Genetic data also support Pleistocene colonization of oceanic islands (Lord Howe Island, Amsterdam Island, St. Helena). Although many biogeographers have speculated about the role of long-distance rafting, this study is one of the first to provide convincing evidence. The marked phylogeographic structure evident across small geographic scales in Australia and South Africa indicates that gene flow among populations may be generally insufficient to prevent the local evolution of monophyly. We suggest that P. exigua may rely on passive mechanisms of dispersal.

Effects of hydrographic barriers on population genetic structure of the sea star Coscinasterias muricata (Echinodermata, Asteroidea) in the New Zealand fiords.

New Zealand's 14 deep-water fiords possess persistent salinity stratification and mean estuarine circulation that may serve to isolate populations of marine organisms that have a dispersal larval phase. In order to investigate this idea, we analysed the population structure of the sea star Coscinasterias muricata using a mitochondrial DNA marker. Genetic differentiation among populations of C. muricata was analysed using 366 base pairs of mtDNA D-loop. We compared populations from the fiords with several others sampled from around New Zealand. At a macro-geographical scale (> 1000 km), restricted gene flow between the North and South Islands was observed. At a meso-geographical scale (10-200 km), significant population structure was found among fiords and between fiords and open coast. The pattern of population genetic structure among the fiords suggests a secondary contact between a northern population and a southern one, separated by a contact or mixing zone. These populations may have diverged by the effects of random genetic drift and population isolation as a consequence of the influence of estuarine circulation on dispersal. In northern Fiordland, genetic structure approximated an isolation by distance model. However, the pattern in genetic differences suggests that distance alone cannot explain the most divergent populations and that fiord hydrography may increase the effect of genetic drift within populations in the fiords. Finally, our study indicates that populations within the fiords underwent recent rapid expansion, followed most probably by genetic drift due to a lack of gene flow among the fiords.

Cladogenesis in a starfish species complex from southern Australia: evidence for vicariant speciation?

DNA sequencing (cytochrome oxidase I; 82 sequences; 25 locations) of a species complex of Australian six-rayed sea-stars (genus Patiriella) reveals four well-supported mtDNA clades, corresponding to P. oriens, P. occidens, P. medius, and P. gunnii. These clades have non-random geographic distributions along an east to west axis that are broadly consistent with the biogeographic provinces of southern Australia proposed by. The taxa are deeply divergent (minimum 7.5%) and are estimated to have originated during the late Pliocene. By contrast, intra-clade divergences are small, typically less than 1.0%. Phylogenetic analysis of mtDNA provides strong support for the combined monophyly of multicoloured forms (P. oriens, P. occidens, and P. medius; 100% bootstrap support) and suggests that P. medius (central) and P. occidens (western) may be sister taxa (up to 76% bootstrap support). Maximum likelihood analysis of nuclear DNA sequences (actin; 1437 bp) yields an optimal tree largely consistent with mtDNA groupings, but with little bootstrap support. The biogeographic distribution of P. oriens (eastern) and P. occidens (western) is roughly consistent with a vicariant model involving allopatric divergence during glaciation. In addition, we propose that the Great Australian Bight may also have retained isolated populations during glacial periods, perhaps explaining the "central" distributions of P. gunnii and P. medius.