Defining spatial and temporal patterns of phylogeographic structure in Madagascar's iguanid lizards (genus Oplurus)

Understanding the remarkably high species diversity and levels of endemism found among Madagascar’s flora and fauna has been the focus of many studies. One hypothesis that has received much attention proposes that Quaternary climate fluctuations spurred diversification. While spatial patterns of distribution and phylogenetic relationships can provide support for biogeographic predictions, temporal estimates of divergence are required to determine the fit of these geospatial patterns to climatic or biogeographic mechanisms. We use multilocus DNA sequence data to test whether divergence times among Malagasy iguanid lizards of the subfamily Oplurinae are compatible with a hypotheses of Pliocene–Pleistocene diversification. We estimate the oplurine species tree and associated divergence times under a relaxed‐clock model. In addition, we examine the phylogeographic structure and population divergence times within two sister species of Oplurus primarily distributed in the north‐west and south‐west of Madagascar (Oplurus cuvieri and Oplurus cyclurus, respectively). We find that divergence events among oplurine lineages occurred in the Oligocene and Miocene and are thus far older and incompatible with the hypothesis that recent climate fluctuations are related to current species diversity. However, the timing of intraspecific divergences and spatial patterns of population genetic structure within O. cuvieri and O. cyclurus suggest a role for both intrinsic barriers and recent climate fluctuations at population‐level divergences. Integrating information across spatial and temporal scales allows us to identify and better understand the mechanisms generating patterns diversity.     

Extensive genetic differentiation in Gobiomorphus breviceps from New Zealand

Partial mitochondrial DNA sequences for parts of the cytochrome b gene and control region were obtained for 89 upland bullies Gobiomorphus breviceps from 19 catchments in New Zealand. There were two highly distinctive mtDNA clades: a northern clade corresponding to the North Island, northern South Island and west coast South Island, and a south‐east clade, in the southern and eastern South Island. Within these major clades there were further distinct clades that correlated with geographic sub‐regions and catchments. The marked genetic differentiation has occurred in the absence of obvious morphological divergence. Based on cytochrome b sequence divergences and the molecular clock hypothesis, the northern and southeastern clades correspond with the uplift of the Southern Alps during the Pliocene, while populations in the North Island and northern South Island were estimated to have diverged during the Pleistocene. The widescale geographic divergences were similar to those observed in the galaxiids, Galaxias vulgaris and Galaxias divergens , but biogeographic management boundaries may not be the same, reflecting different evolutionary histories for non‐diadromous species occupying the same areas.     

Biogeography and evolution of European cave salamanders, Hydromantes (Urodela: Plethodontidae), inferred from mtDNA sequences

Aim To infer the phylogenetic relationships and biogeography of Hydromantes, with special emphasis on the European taxa. In particular, we aimed to test: (1) the monophyly of the European species and current views on their interrelationships; and (2) previously proposed timings of the separation of European and American Hydromantes, and of biogeographically important events within Europe. Location California and the Western Mediterranean Basin, specifically south‐east France, Italy, and the island of Sardinia. Methods Partial sequences of mitochondrial genes (cytochrome b and 12S rRNA) were obtained from 45 specimens of Hydromantes, including all European extant species and subspecies, and two species from California. In addition, a fragment of the mitochondrial 16S rRNA gene was amplified for 16 specimens. Data sets were aligned using Clustal X, and well‐supported phylogenetic trees were produced using maximum‐likelihood, Bayesian and maximum‐parsimony methods. Estimates of divergence times were obtained with the program r8s , the molecular clock being calibrated using the opening of the Strait of Gibraltar, the final event in the Messinian Salinity Crisis of 5.3 Ma. Results Separation between the American and European clades occurred approximately 13.5 Ma, most probably before or after westward dispersal across the Bering Land Bridge. In Europe, divergence started in the late Miocene, when Hydromantes (A.) genei separated from other members of the genus 9 Ma and colonized south‐west Sardinia. Movement between the European mainland and Sardinia, by a member of the subgenus Speleomantes, occurred in the Messinian Salinity Crisis, after the Mediterranean Basin desiccated almost completely 5.96 Ma. Subsequent widespread aridification fragmented the geographical ranges of Hydromantes, which live in cool and humid conditions, resulting in the origin of the six species in the subgenus Speleomantes. In contrast, a second period of diversification, in continental Europe 2–1.3 Ma, was probably caused by very cold interludes during the climatic oscillations that characterized the Pleistocene. Main conclusions The molecular clock used here indicates that the separation of Californian and European Hydromantes occurred more recently than previously believed, and the same is true of some subsequent phylogenetic divergences within Europe. Estimated dates for these divergence events are consistent with known geophysical and climatic events that could have caused or facilitated them.     

Multiple fossil calibrations,nuclear loci and mitochondrial genomes provide new insight into biogeography and divergence timing for true seals (Phocidae,Pinnipedia)

Aim To better understand the historical biogeography of the true seals, Phocidae, by combining nuclear DNA (nDNA) and mitochondrial DNA (mtDNA) in a divergence time analysis using multiple fossil calibrations. Location Arctic, Antarctic, Pacific and Atlantic Oceans, Lake Baikal, Caspian Sea. Methods Fifteen nuclear genes totalling 8935 bp plus near‐complete mitochondrial genome sequences were used in a Bayesian divergence time analysis, incorporating eight soft‐bound fossil calibrations across the phylogeny. All species of true seals were included, plus the walrus, three otariids and seven carnivore outgroups. The majority of the nuclear sequences and four phocid mitochondrial genomes (plus three non‐phocid mitochondrial genomes) were newly generated for this study using DNA extracted from tissue samples; other sequences were obtained from GenBank. Results Using multiple nuclear genes and multiple fossil calibrations resulted in most divergence time estimations within Phocidae being much more recent than predicted by other molecular studies incorporating only mtDNA and using a single calibration point. A new phylogenetic hypothesis was recovered for the Antarctic seals. Main conclusions Incorporating multiple nuclear genes and fossil calibrations had a profound effect on the estimated divergence times. Most estimated divergences within Phocinae (Arctic seals) correspond to Arctic oceanic events and all occur within the last 12 Myr, a time when the Arctic and Atlantic oceans were freely exchanging and perennial Arctic sea ice existed, indicating that the Arctic seals may have had a longer association with ice than previously thought. The Monachinae (‘southern’ seals) split from the Phocinae c. 15 Ma on the eastern US coast. Several early trans‐Atlantic dispersals possibly occurred, leaving no living descendants, as divergence estimates suggest that the Monachus (monk seal) species divergences occurred in the western Atlantic c. 6 Ma, with the Mediterranean monk seal ancestor dispersing afterwards. The tribes Lobodontini (Antarctic seals) and Miroungini (elephant seals) are also estimated to have diverged in the eastern Atlantic c. 7 Ma and a single Lobodontini dispersal to Antarctica occurred shortly afterwards. Many of the newly estimated dates are used to infer how extinct lineages/taxa are allied with their living relatives.     

High levels of intraspecific genetic divergences revealed for Antarctic springtails: evidence for small‐scale isolation during Pleistocene glaciation

We examined levels of genetic variability within and among populations of three Antarctic springtail species (Arthropoda: Collembola) and tested the hypothesis that genetic divergences occur among glacially‐isolated habitats. The study was conducted in southern Victoria Land, Ross Dependency, Antarctica, and samples were collected from locations in the vicinity of the Mackay Glacier. We analyzed mtDNA (cytochrome c oxidase subunit I; COI) sequence variability for 97 individuals representing three species (Gomphiocephalus hodgsoni, N = 67; Cryptopygus nivicolus, N = 20; and Antarcticinella monoculata, N = 8). Haplotype diversity and genetic divergences were calculated and used to indicate population variability and also to infer divergence times of isolated populations using molecular clock estimates. Two of the three species showed high levels of genetic divergence. Gomphiocephalus hodgsoni, a widespread and common species, showed 7.6% sequence divergence on opposite sides of the Mackay Glacier. The more range restricted C. nivicolus showed 4.0% divergence among populations. The third species, A. monoculata, was found in only one location. Molecular clock estimates based on sequence divergences suggest that populations separated within the last 4 Mya. We conclude that habitat fragmentation resulting from Pliocene (5 Mya) and Pleistocene (2 Mya to 10 Kya) glaciations has promoted and maintained high levels of diversity among isolated springtail populations on relatively small spatial scales. The region surrounding the Mackay Glacier is likely to have provided refugia for springtail populations during glacial maxima and remains an area of high genetic and species diversity for Collembola within the Ross Sea region.     

Life history characteristics of the estuary eeltail catfish Cnidoglanis macrocephalus (Plotosidae) in eastern Australia

The biology and fisheries of cobbler Cnidoglanis macrocephalus have been examined on the Australian west coast, but not the east coast. This study identified that east coast estuary cobbler was more prevalent over vegetated habitat than bare substrata and that spawning occurred during spring and early summer. Observed longevity was 9⁺ and 20⁺ years for males and females, but there were no discernible differences between sexes or estuaries in mean total length (TL) at age. The von Bertalanffy growth function of combined sexes was L_inf?=?61.2 cm TL, k?=?0.677, t₀?=?0.007. The data demonstrate similarities and differences in life history characteristics of cobbler between coasts.

     

Phylogeographic evidence for two mesic refugia in a biodiversity hotspot

Phylogeographic studies of flora in species-rich south-western Australia point to complex evolutionary histories, reflecting patterns of persistence and resilience to climatic changes during the Pleistocene. We asked whether coastal areas of the mid-west and south, as well as granite outcrops and inland ranges, have acted as major refugia within this region during Pleistocene climatic fluctuations by analysing phylogeographic patterns in the shrub Calothamnus quadrifidus R.Br. (Myrtaceae). We determined variation in chloroplast DNA data for 41 populations across the geographic range. Relationships and major clades were resolved using parsimony and Bayesian analyses. We tested for demographic and spatial expansion of the major clades and estimated clade divergence dates using an uncorrelated, lognormal relaxed clock based on two conservative chloroplast mutation rates. Two distinct phylogeographic clades were identified showing divergence during the Pleistocene, consistent with other phylogeographic studies of south-west Australian flora, emphasising the impact of climatic oscillations in driving divergence in this landscape. The southern clade was more diverse, having higher haplotype diversity and greater genetic structure, while the northern clade showed evidence of fluctuation in population size. Regions of high haplotype diversity with adjacent areas of low diversity observed in each clade indicated the locations of two coastal refugia: one on the south coast and another along the mid-west coast. This is the first evidence for major Pleistocene refugia using chloroplast genetic data in a common, widespread species from this region.     

Impact of Pleistocene aridity oscillations on the population history of a widespread,vagile Australian mammal,Macropus fuliginosus

Aim Climatic fluctuations during the Pleistocene have shaped the population structure of many extant taxa. However, few studies have examined widespread species inhabiting the Australian continent, where periods of increased aridity characterized the Pleistocene. Here we investigate the phylogeography and population history of a widespread and vagile southern Australian marsupial, the western grey kangaroo (Macropus fuliginosus). Location Southern Australia. Methods We examined the variation of the mitochondrial DNA (mtDNA) control region from 511 individuals of M. fuliginosus sampled throughout their transcontinental distribution. Maximum likelihood and Bayesian analyses were used to investigate the phylogeography and coalescence analyses were then used to test hypothesized biogeographical scenarios. Results The combined results of the phylogeographical and coalescence analyses revealed a complex evolutionary history. Macropus fuliginosus originated in the south‐west of the continent, with north‐western and south‐western populations subsequently diverging as a result of vicariance events during the mid‐Pleistocene. Subsequent arid phases affected these populations differently. In the north‐west, the expansion and contraction of the arid zone resulted in repeated vicariance events and multiple divergent north‐western mtDNA subclades. In contrast, the south‐western population was less impacted by climatic oscillations but gave rise to a major transcontinental eastward expansion. Main conclusions Macropus fuliginosus exhibits the genetic signature of divergence due to unidentified barriers in south‐western Western Australia, while previously identified barriers across southern Australia appear to have had little impact despite evidence of a broad‐scale range expansion prior to the Last Glacial Maximum (LGM). This pattern of localized expansion and contraction is comparable to unglaciated regions in both the Northern and Southern Hemispheres. Furthermore, this study indicates that despite the potential similarities between Northern Hemisphere glaciation and the activation of dune systems in the Australian arid zone, both of which rendered large areas inhospitable, the biotic responses and resultant phylogeographical signatures are dissimilar. Whereas a limited number of major geographically concordant refugia are observed in glaciated areas, the Southern Hemisphere arid zone appears to be associated with multiple species‐specific idiosyncratic refugia.     

Implications of acute temperature and salinity tolerance thresholds for the persistence of intertidal invertebrate populations experiencing climate change

To predict whether populations of marine animals will persist in the face of changing climate conditions, it is informative to understand how past climate conditions have shaped present‐day tolerance thresholds. We examined 4 species of intertidal invertebrates (Nucella lamellosa, Littorina scutulata, Littorina sitkana, and Balanus glandula) inhabiting the coasts of Vancouver Island, Canada, where the east coast experiences historically warmer sea surface temperature (SST), warmer low tide (i.e., emersion) rock surface temperature (RST), and lower sea surface salinity (SSS) than the west coast. To determine if east coast populations have higher tolerance thresholds to acute stress than west coast populations, animals from 3 sites per coast were exposed to stressful temperatures and salinities in common garden experiments. Emersion temperature tolerance differed between populations only in N. lamellosa and B. glandula, tolerance thresholds being 1.4–1.5°C higher on the east coast. Water temperature tolerance differed between populations only in B. glandula and L. scutulata but was highest on the west coast. No differences in salinity tolerance were observed within any species. Thus, there is limited evidence of divergence among east and west coast populations in tolerance of acute stress despite the substantial historical differences in extreme temperature and salinity conditions between coasts. However, based on present‐day summertime SST and RST and known rates of change in these parameters, we predict present‐day tolerance thresholds would be sufficient to allow adults of these populations to tolerate extreme temperatures predicted for the next several hundred years, and that even a slow rate of change in acute tolerance thresholds might suffice to keep up with future temperature extremes.     

Trapped in desert springs: phylogeography of Australian desert spring snails

Aim We investigate the phylogeographical history and determine the time‐scale of population divergence of hydrobiid freshwater snails (genus Trochidrobia) inhabiting groundwater springs in the Australian desert. We test the hypothesis that divergence between geographically distinct snail populations occurred simultaneously due to their isolation in hydrologically discrete spring systems, i.e. ‘trapped in desert springs’. Location Groundwater springs of the Great Artesian Basin (GAB) in central Australia. Methods DNA sequence data from the mitochondrial cytochrome c oxidase subunit I gene and the nuclear 28S and internal transcribed spacer rRNA genes were used to reconstruct phylogenetic relationships within and among three species of Trochidrobia (Hydrobiidae): T. punicea (13 spring groups, n = 90), T. smithi (12 spring groups, n = 62) and T. minuta (2 spring groups, n = 4). Bayesian relaxed molecular clock analyses and approximate Bayesian computation were used to date lineage divergence and distinguish between alternative biogeographical scenarios. Results The diversification of the three Trochidrobia species probably occurred between 2.54 and 9.3 Ma, prior to the formation of the springs c. 1 Ma. Intraspecific divergences within the two widespread species occurred after the formation and colonization of the springs. Coalescent modelling and molecular clock analyses supported a simultaneous radiation of five allopatric intraspecific snail lineages within T. punicea (two lineages) and T. smithi (three lineages) across the GAB springs examined. Main conclusions The analyses support the ‘trapped in desert springs’ hypothesis for the diversification of intraspecific lineages within the species T. punicea and T. smithi. This hypothesis suggests that the formation of deserts around Lake Eyre in the early Pleistocene led to the hydrological isolation of spring complexes in the GAB, resulting in significant molecular divergence, but no morphological divergence, of Trochidrobia snail populations.     

The role of vicariance and dispersal on New Zealand's estuarine biodiversity: the case of Paracorophium (Crustacea: Amphipoda)

To investigate the role of vicariance and dispersal on New Zealand's estuarine biodiversity, we examined variability in mitochondrial cytochrome c oxidase subunit I (COI) gene sequences for the amphipod genus Paracorophium. Individuals from the two nominate endemic species (Paracorophium excavatum and Paracorophium lucasi) were collected from sites throughout the North and South Islands. Sequence divergences of 12.8% were detected among the species. However, divergences of up to 11.7% were also observed between well supported clades, suggesting the possibility of cryptic species. Nested clade analyses identified four distinct lineages from within both P. excavatum and P. lucasi, with boundaries between clades corresponding to topographical features (e.g. Cook Straight, North and East Cape). Sequence divergences of 3.7–4.9% were also observed within geographic regions (e.g. east and west coasts of the upper North Island). Genetic structure in Paracorophium appears to represent prolonged isolation and allopatric evolutionary processes dating back to the Upper Miocene and continuing through the Pliocene and early Pleistocene. On the basis of molecular clock estimates from sequence divergences and reconstructions of New Zealand's geological past, we suggest that sea level and landmass changes during the early Pleistocene (2 Mya) resulted in the isolation of previously contiguous populations leading to the present‐day patterns. COI genetic structure was largely congruent with previously observed allozyme patterns and highlights the utility of COI as an appropriate marker for phylogeographic studies of the New Zealand estuarine fauna. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 863–874.     

Population genomic analysis uncovers African and European admixture in Drosophila melanogaster populations from the south‐eastern United States and Caribbean Islands

Drosophila melanogaster is postulated to have colonized North America in the past several 100 years in two waves. Flies from Europe colonized the east coast United States while flies from Africa inhabited the Caribbean, which if true, make the south‐east US and Caribbean Islands a secondary contact zone for African and European D. melanogaster. This scenario has been proposed based on phenotypes and limited genetic data. In our study, we have sequenced individual whole genomes of flies from populations in the south‐east US and Caribbean Islands and examined these populations in conjunction with population sequences from the west coast US, Africa, and Europe. We find that west coast US populations are closely related to the European population, likely reflecting a rapid westward expansion upon first settlements into North America. We also find genomic evidence of African and European admixture in south‐east US and Caribbean populations, with a clinal pattern of decreasing proportions of African ancestry with higher latitude. Our genomic analysis of D. melanogaster populations from the south‐east US and Caribbean Islands provides more evidence for the Caribbean Islands as the source of previously reported novel African alleles found in other east coast US populations. We also find the border between the south‐east US and the Caribbean island to be the admixture hot zone where distinctly African‐like Caribbean flies become genomically more similar to European‐like south‐east US flies. Our findings have important implications for previous studies examining the generation of east coast US clines via selection.     

Tethyan vicariance, relictualism and speciation: evidence from a global molecular phylogeny of the opisthobranch genus Bulla

Aim Our aims were: (1) to reconstruct a molecular phylogeny of the cephalaspidean opisthobranch genus Bulla, an inhabitant of shallow sedimentary environments; (2) to test if divergence times are consistent with Miocene and later vicariance among the four tropical marine biogeographical provinces; (3) to examine the phylogenetic status of possible Tethyan relict species; and (4) to infer the timing and causes of speciation events. Location Tropical and warm‐temperate regions of the Atlantic, Indo‐West Pacific, Australasia and eastern Pacific. Methods Ten of the 12 nominal species of Bulla were sampled, in a total sample of 65 individuals, together with cephalaspidean outgroups. Phylogenetic relationships were inferred by Bayesian analysis of partial sequences of the mitochondrial cytochrome c oxidase I (COI) and 16S rRNA and nuclear 28S rRNA genes. Divergence times and rates of evolution were estimated using uncorrelated relaxed‐clock Bayesian methods with fossil calibrations (based on literature review and examination of fossil specimens), implemented in beast . The geographical pattern of speciation was assessed by estimating the degree of overlap between sister lineages. Results Four clades were supported: Indo‐West Pacific (four species), Australasia (one species), Atlantic plus eastern Pacific (three species) and Atlantic (two species), with estimated mean ages of 35–46 Ma. Nominal species were monophyletic, but deep divergences were found within one Indo‐West Pacific and one West Atlantic species. Species‐level divergences occurred in the Miocene or earlier. The age of a sister relationship across the Isthmus of Panama was estimated at 7.9–32.1 Ma, and the divergence of a pair of sister species on either side of the Atlantic Ocean occurred 20.4–27.2 Ma. Main conclusions Fossils suggest that Bulla originated in the Tethys realm during the Middle Eocene. Average ages of the four main clades fall in the Eocene, and far pre‐date the 18–19 Ma closure of the Tethys Seaway. This discrepancy could indicate earlier vicariant events, selective extinction or errors of calibration. Similarly, the transisthmian divergence estimate far pre‐dates the uplift of the Panamanian Isthmus at about 3 Ma. Speciation events occurred in the Miocene, consistent with tectonic events in the central Indo‐West Pacific, isolation of the Arabian Sea by upwelling and westward trans‐Atlantic dispersal. Differences in habitat between sister species suggest that ecological speciation may also have played a role. The basal position of the Australasian species supports its interpretation as a Tethyan relict.     

Evolution and maintenance of divergent lineages in an endangered freshwater fish, <Emphasis Type="Italic">Macquaria australasica</Emphasis>

Variable hydrological regimes and habitat availability are factors that affect the distribution of freshwater dependent species and are expected to influence their levels of genetic diversity. Although geologically relatively stable, the south eastern region of Australia has experienced significant changes in hydrological conditions during the Quaternary. This area has also been recently affected by anthropogenic activities, resulting in dramatic population declines of Macquarie Perch (Macquaria australasica). We conducted a range-wide phylogeographic study of this endangered fish to assess the relationship between landscape and freshwater fish evolution in south eastern Australia and infer levels of genetic diversity and population structure. Surprisingly, we detected high genetic diversity, with 46 mtDNA control region haplotypes found across 37 sampling locations. Some lineages were remarkably divergent; one represents a putative undescribed species that probably went extinct during the period of this study. Our reconstruction of population history using a combination of coalescent and phylogenetic methods indicates that the species originated on the coast, east of the Great Dividing Range (GDR), with subsequent colonisation of the Murray-Darling basin, west of the GDR. Nested clade and IM analyses inferred a series of range expansions and fragmentations across the species range consistent with the history of climatic oscillations in south eastern Australia during the Pleistocene. We conclude that the unexpected high levels of diversity and divergence observed in M. australasica may be due to specific habitat requirements, localised recruitment, and Pleistocene climate fluctuations. Under expectations of a drier climate and increased sea levels due to global warming, populations of this and other freshwater species may be expected to experience increased habitat fragmentation and loss of genetic diversity. Conservation management should focus on habitat protection, the maintenance of genetic diversity and taxonomic review.     

Shifting nonbreeding distributions of migratory fauna in relation to climatic change

The distributions of eight out of nine common species of waders (Charadrii) overwintering on UK estuaries have changed in association with recent climate change. These birds represent a high proportion of various populations from breeding grounds as far apart as Greenland to the west to high‐arctic Russia to the east. During warmer winters, smaller proportions of seven species wintered in south‐west Britain. The distributions of the smaller species show the greatest temperature dependence. The opposite was found for the largest species and no relationship was found for a particularly site‐faithful species. In north‐west Europe, the winter isotherms have a broadly north to south alignment, with the east being colder than the west. The average minimum winter temperatures across the UK having increased by about 1.5°C since the mid‐1980s, the temperatures on the east coast during recent winters have been similar to those of the west coast during the mid‐1980s. On average, estuaries on the east and south coasts of Britain have muddier sediments than those on the west coast and thus support a higher biomass of the invertebrate prey of waders. We suggest that, with global climatic change, the advantage gained by waders wintering in the milder west to avoid cold weather‐induced mortality is diminished. Consequently, more choose to winter in the east and thus benefit from better foraging opportunities. The implications of these results are considered in terms of a site‐based approach to wildlife protection used in Europe and elsewhere.     

Phylogeographic structure in the gilgie (Decapoda: Parastacidae: Cherax quinquecarinatus): a south‐western Australian freshwater crayfish

The gilgie (Cherax quinquecarinatus) is among the more widespread of the six endemic south‐western Australian freshwater crayfish species. In the present study, the phylogeographic structure of the gilgie was investigated across its distribution to determine whether patterns reflected those identified earlier in a co‐distributed congeneric, the koonac (Cherax preissii). Gilgies were sampled from 20 localities, a 412‐bp fragment of the cytochrome c oxidase subunit I mitochondrial DNA gene was amplified from 75 individuals, and allozyme variation was assayed at nine loci. As in the koonac, three geographically‐restricted lineages were identified: from the north‐western, southern coastal, and intermediate/south‐western regions. Phylogeographic breaks appeared to be congruent with those in the koonac. The extent of genetic differentiation among lineages was comparable to that in the koonac, suggesting temporal congruence of the historical events responsible for the observed structure. A relaxed Bayesian molecular clock suggested that the major clades and lineages in each species diverged in the Late Miocene–Early Pliocene (4.0–9.6 Myr ago), possibly resulting from increasing pulses of aridity. The retrieval of almost‐identical phylogeographic structure in two co‐distributed species suggests that biogeographic regions can be more accurately defined in south‐western Australia. With the geographic fidelity of these lineages, the present data also provide evidence of the translocation of a single individual from the north‐west to the south coast. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 385–402.     

Incipient speciation of Catostylus mosaicus (Scyphozoa, Rhizostomeae, Catostylidae), comparative phylogeography and biogeography in south-east Australia

Aim Phylogeography provides a framework to explain and integrate patterns of marine biodiversity at infra‐ and supra‐specific levels. As originally expounded, the phylogeographic hypotheses are generalities that have limited discriminatory power; the goal of this study is to generate and test specific instances of the hypotheses, thereby better elucidating both local patterns of evolution and the conditions under which the generalities do or do not apply. Location Coastal south‐east Australia (New South Wales, Tasmania and Victoria), and south‐west North America (California and Baja California). Methods Phylogeographic hypotheses specific to coastal south‐east Australia were generated a priori, principally from existing detailed distributional analyses of echinoderms and decapods. The hypotheses are tested using mitochondrial cytochrome c oxidase subunit I (COI) and nuclear internal transcribed spacer 1 (ITS1) DNA sequence data describing population variation in the jellyfish Catostylus mosaicus, integrated with comparable data from the literature. Results Mitochondrial COI distinguished two reciprocally monophyletic clades of C. mosaicus (mean ± SD: 3.61 ± 0.40% pairwise sequence divergence) that were also differentiated by ITS1 haplotype frequency differences; the boundary between the clades was geographically proximate to a provincial zoogeographic boundary in the vicinity of Bass Strait. There was also limited evidence of another genetic inhomogeneity, of considerably smaller magnitude, in close proximity to a second hypothesized zoogeographic discontinuity near Sydney. Other coastal marine species also show genetic divergences in the vicinity of Bass Strait, although they are not closely concordant with each other or with reported biogeographic discontinuities in the region, being up to several hundreds of kilometres apart. None of the species studied to date show a strong phylogeographic discontinuity across the biogeographic transition zone near Sydney. Main conclusions Patterns of evolution in the Bass Strait and coastal New South Wales regions differ fundamentally because of long‐term differences in extrinsic factors. Since the late Pliocene, periods of cold climate and low sea‐level segregated warm temperate organisms east or west of an emergent Bassian Isthmus resulting in population divergence and speciation; during subsequent periods of warmer and higher seas, sister taxa expanded into the Bass Strait region leading to weakly correlated phylogeographic and biogeographic patterns. The Sydney region, by contrast, has been more consistently favourable to shifts in species’ ranges and long‐distance movement, resulting in a lack of intra‐specific and species‐level diversification. Comparisons between the Sydney and Bass Strait regions and prior studies in North America suggest that vicariance plays a key role in generating coastal biodiversity and that dispersal explains many of the deviations from the phylogeographic hypotheses.     

The combination of selection and dispersal helps explain genetic structure in intertidal mussels

Understanding patterns of gene flow, selection and genetic diversity within and among populations is a critical element of predicting how long-term changes in environmental conditions are likely to affect species distribution. The intertidal mussel Perna perna consists of two distinct genetic lineages in South Africa, but the mechanisms maintaining these lineages remains obscure. We used regional oceanography and lineage-specific responses to environmental conditions as proxies for gene flow and local selection, respectively, to test how these mechanisms could shape population genetic structure. Laboratory experiments supported the field findings that mussels on the east coast (eastern lineage) are physiologically more tolerant of sand inundation and high temperatures than those on the south coast (western lineage). Temperature loggers mimicking mussel body temperatures revealed that mussels experience higher body temperatures during aerial exposure on the subtropical east coast than on the temperate south coast. Translocations showed that, on the east coast, the western lineage suffered higher mortality rates than local individuals, while on the south coast, mortality rates did not differ significantly between the lineages. Nearshore drogues showed remarkably little overlap between the trajectories of drifters released off the south coast and those released off the east coast. Physiological tolerances can thus explain the exclusion of western individuals from the east coast, but they cannot explain the exclusion of the eastern lineage from the south coast. In contrast, however, ocean dynamics may limit larval dispersal between the two lineages, helping to explain the absence of eastern individuals from the south coast. We emphasise the importance of a multidisciplinary approach in a macro-ecological context to understand fully the mechanisms promoting evolutionary divergence between genetic entities. Our results suggest that phylogeographic patterns of Perna perna may be maintained by a combination of local conditions and the isolating effect of the Agulhas Current that reduces gene exchange.     

Comparative phylogeography reveals host generalists, specialists and cryptic diversity: Hexabothriid, microbothriid and monocotylid monogeneans from rhinobatid rays in southern Australia

The phylogeography and host specificity of three monogenean species infecting different sites on the southern fiddler ray, Trygonorrhina fasciata (Rhinobatidae) in South Australia (SA) were studied: Branchotenthes octohamatus (Hexabothriidae: gills), Calicotyle australis (Monocotylidae: cloaca) and Pseudoleptobothrium aptychotremae (Microbothriidae: skin). Five rhinobatid species (Aptychotrema vincentiana, T. fasciata, Trygonorrhina sp. A, Aptychotrema rostrata and Rhinobatos typus) with distributions spanning west, south and east Australian coastal waters, were surveyed for monogeneans resembling the three species documented from T. fasciata in SA. The identities of hosts and parasites collected were investigated using the mitochondrial genes ND4 and Cytochrome b (cytb), respectively, in addition to the nuclear marker, Elongation factor 1-alpha (EF1a) for Pseudoleptobothrium. Genetic analyses confirmed that B. octohamatus is geographically widespread and displays little genetic structure, suggesting high levels of gene flow. It was collected from four rhinobatid species throughout its distribution and is not, therefore, host specific. For C. australis, genetic analyses revealed two discrete populations with a genetic divergence of ∼4%, one population occurring west of Bass Strait on two sympatric host species and the other population on the east coast, also occurring on sympatric host species. Similarly, for Pseudoleptobothrium, specimens collected west of Bass Strait were genetically distinct (∼3.5%) from those collected to the east. However, on the east coast, a third Pseudoleptobothrium population was revealed, separated by a genetic distance of >11%, indicating a morphologically cryptic species. Host preferences were indicated for each Pseudoleptobothrium lineage. These genetic discoveries are discussed in relation to life history characteristics of each monogenean species, highlighting the value of phylogeographic analyses to understand the parasite-host relationship.     

Trophic overlap between sexes in the dimorphic African black oystercatcher foraging on an alien mussel

Sex‐specific feeding segregation related to sexual bill dimorphism has been described in several oystercatcher species, including the African black oystercatcher. For the latter, studies concerned only a small number of breeding pairs and were done prior the invasion of the South African rocky shores by the Mediterranean mussel, which is believed to have benefited oystercatchers by increasing overall biomass. Here, we investigated geographic variability in the segregation of diet, biometrics and body condition between sexes in the African species, in relation to changes in foraging habitats along the South African coastline, using stable isotope analyses. Males and females and their potential prey (mussels, limpets, polychaetes and ascidians) were sampled on the southern African west, south‐west and south‐east coasts for stable isotope analyses and biometrics and body conditions of birds were measured. Bill dimorphism occurred throughout the study area and south‐west males had lower body conditions than other males and females in general. Sexes displayed little differences in their δ¹³C ratios and in the relative consumption of the different prey throughout the study area, except on the south‐east coast where males were slightly depleted in ¹³C relative to females and the most abundant prey elsewhere (the Mediterranean mussel) is rare. Females were slightly but significantly enriched in ¹⁵N by 0.3‰ compared to their breeding partners and this did not link clearly to differences in diet. We argue that the combined effect of biogeographic variations in rocky shores diversity and biomass, heterogeneous invasion by the Mediterranean mussel on the South African coastline and bill dimorphism may have altered the sex‐specific feeding behaviour of oystercatchers differently between coastal regions and possibly had an additional cost for male oystercatchers faced with lower prey biomass and diversity on the south‐west coast.