Extrinsic and intrinsic influences on the phylogeography of the Austrocochlea constricta species group

Although extrinsic factors, such as oceanic currents and isolation induced by sea level maxima during Plio-Pleistocene glacial cycles, are often suggested as principal determinants of marine phylogeography, they are not always complete explanations. The counterexamples to predominant phylogeographic patterns in southeastern Australia suggest for example, that intrinsic factors such a habitat preference or reproductive mode can have significant influence.We collected DNA sequences from mitochondrial cytochrome c oxidase I (COI) and two nuclear gene introns from the Austrocochlea constricta species group to determine whether its porcata and constricta phenotypes are genetically distinguishable, to examine the phylogeographic effects of the Bassian Isthmus landbridge formed between Tasmania and Victoria during glacial maxima, and to investigate the importance of intrinsic factors in structuring its genetic variability.No fixed genetic differences between the porcata and constricta phenotypes were identified in any gene so the number of species comprised by the group cannot yet be determined. The two major clades recovered in COI analyses were respectively found principally east or west of the Bassian Isthmus. A. constricta is the first lecithotrophic or estuarine invertebrate known to show this pattern. There were no fixed differences, for any of the three genes, between specimens from estuarine and marine populations within either the eastern or western COI clades. Other intrinsic factors such as breeding period (possibly) and larval type (probably) may play roles in modulating phylogeographic patterns.     

Islands increase genetic subdivision and disrupt patterns of connectivity of intertidal snails in a complex archipelago

The view that marine species with planktonic dispersal have highly connected, demographically open populations is giving way to recognition that populations may often be largely self-recruiting, or demographically closed. This raises the question of what local conditions might favor isolation of populations. To test the importance of islands for local isolation in species with planktonic larvae, we examined allozyme variation among 35 populations of the intertidal snail Austrocochlea constricta in the Houtman Abrolhos Islands, Western Australia, spanning 60 km. Heterogeneity of allozyme frequencies among populations was high, with average F(ST) of 0.237, indicating highly localized populations. Increased subdivision was associated with islands at different scales: between island groups, separated by deep water gaps, and between disconnected sets of islands within groups. At short distances, up to two km, subdivision increased fivefold between islands compared with that between populations on the same island. Along 11 km of continuous, sheltered shore, there was isolation by distance but among a linear series of islands over similar distance, there was greater subdivision at short distances but no association with distance. These patterns had been seen previously in the direct-developing snail Bembicium vittatum, but its finding in A. constricta confirms for a planktonic disperser the importance of this complex archipelago for both retention of locally produced larvae and disruption of patterns of connectivity. Taken together, these results indicate that islands can increase both the "open" and the "closed" components of recruitment and that applicable models of genetic connectivity depend substantially on local conditions.     

Cladogenesis as the result of long-distance rafting events in South Pacific topshells (Gastropoda, Trochidae)

We used DNA sequences of lecithotrophic monodontine topshells, belonging to the genera Diloma, Melagraphia, and Austrocochlea, to ascertain how this group became established over a large area of the South Pacific Ocean. The phylogeny of the topshells was estimated using portions of two mitochondrial genes (16S and cytochrome oxidase 1) and one nuclear gene (actin). A range of divergence rates was used to estimate the approximate timing of cladogenetic events within their phylogenetic tree. These estimates allow us to unambiguously reject vicariant explanations for several major divergence events and to infer several dispersal events across wide stretches of ocean. The first were two initial dispersal events from Australia (1) to an area between Samoa and Japan and (2) to New Zealand. Subsequently, at least one, and possibly two, recent eastward dispersals took place from New Zealand to Chile and the Juan Fernandez Islands, and one further dispersal occurred from somewhere in the tropical Pacific to Samoa. Moreover, owing to the short-lived nature of the topshell larvae, transoceanic larval dispersal is unlikely. The apparent paradox of a short larval phase and broad geographic range suggests that dispersal most probably occurred by rafting of adults on a suitable platform such as macroalgae; indeed, naturally buoyant bull kelp is the natural habitat of the most geographically widespread species in this group. Our molecular phylogenies imply that, despite of being an unlikely event, adult rafting in ocean currents has occurred on several occasions throughout the evolutionary history of topshells, resulting in their wide present-day distribution.