Incorporating non-equilibrium dynamics into demographic history inferences of a migratory marine species.

Understanding how dispersal and gene flow link geographically separated the populations over evolutionary history is challenging, particularly in migratory marine species. In southern right whales (SRWs, Eubalaena australis), patterns of genetic diversity are likely influenced by the glacial climate cycle and recent history of whaling. Here we use a dataset of mitochondrial DNA (mtDNA) sequences (n = 1327) and nuclear markers (17 microsatellite loci, n = 222) from major wintering grounds to investigate circumpolar population structure, historical demography and effective population size. Analyses of nuclear genetic variation identify two population clusters that correspond to the South Atlantic and Indo-Pacific ocean basins that have similar effective breeder estimates. In contrast, all wintering grounds show significant differentiation for mtDNA, but no sex-biased dispersal was detected using the microsatellite genotypes. An approximate Bayesian computation (ABC) approach with microsatellite markers compared the scenarios with gene flow through time, or isolation and secondary contact between ocean basins, while modelling declines in abundance linked to whaling. Secondary-contact scenarios yield the highest posterior probabilities, implying that populations in different ocean basins were largely isolated and came into secondary contact within the last 25,000 years, but the role of whaling in changes in genetic diversity and gene flow over recent generations could not be resolved. We hypothesise that these findings are driven by factors that promote isolation, such as female philopatry, and factors that could promote dispersal, such as oceanographic changes. These findings highlight the application of ABC approaches to infer the connectivity in mobile species with complex population histories and, currently, low levels of differentiation.

The phylogeography of dusky dolphins (Lagenorhynchus obscurus): a critical examination of network methods and rooting procedures.

We investigated the phylogeography and evolutionary history of dusky dolphins (Lagenorhynchus obscurus) using DNA sequences of the full mitochondrial cytochrome b gene in 124 individuals from the putative stocks off Peru, Argentina and Southwest Africa. While genetic differentiation within oceans is surprisingly low, there is no evidence for recent female gene flow between Atlantic and Pacific waters. Highest genetic variability in terms of sequence divergence and number of haplotypes is found in the Atlantic. Our analyses also indicate that the eastern South Pacific dusky dolphins stock should be considered a separate management unit. Given the high level of mortality experienced by the Peruvian dusky dolphin in local fishery activities, these findings have important implications for an objective management of the species. Furthermore, we analysed our mitochondrial sequence data with several widely used network estimation and rooting methods. The resulting intraspecific gene genealogies and rooting inferences exhibited substantial differences, underlying the limitations of some algorithms. Given that scientific hypotheses and management decisions depend strongly on inferred tree or network topologies, there is a clear need for a systematic comparative analysis of available methods. Finally, the present study indicates that (i) the dusky and the Pacific white-sided dolphins are sister species and (ii) not only the Westwind Drift hypothesis but also other models of dispersion are compatible with the current geographical distribution of dusky dolphins.

World-wide genetic differentiation of Eubalaena: questioning the number of right whale species.

Few studies have examined systematic relationships of right whales (Eubalaena spp.) since the original species descriptions, even though they are one of the most endangered large whales. Little morphological evidence exists to support the current species designations for Eubalaena glacialis in the northern hemisphere and E. australis in the southern hemisphere. Differences in migratory behaviour or antitropical distribution between right whales in each hemisphere are considered a barrier to gene flow and maintain the current species distinctions and geographical populations. However, these distinctions between populations have remained controversial and no study has included an analysis of all right whales from the three major ocean basins. To address issues of genetic differentiation and relationships among right whales, we have compiled a database of mitochondrial DNA control region sequences from right whales representing populations in all three ocean basins that consist of: western North Atlantic E. glacialis, multiple geographically distributed populations of E. australis and the first molecular analysis of historical and recent samples of E. glacialis from the western and eastern North Pacific Ocean. Diagnostic characters, as well as phylogenetic and phylogeographic analyses, support the possibility that three distinct maternal lineages exist in right whales, with North Pacific E. glacialis being more closely related to E. australis than to North Atlantic E. glacialis. Our genetic results provide unequivocal character support for the two usually recognized species and a third distinct genetic lineage in the North Pacific under the Phylogenetic Species Concept, as well as levels of genetic diversity among right whales world-wide.

Anatomy of the laryngeal apparatus of the pygmy right whale, Caperea marginata (Gray 1846).

The laryngeal apparatus of Caperea marginata is described for the first time and proves to be significantly different from that of any mysticete previously described. This difference is especially noticeable in the position of the laryngeal sac, which is separate from the tracheal rings and lies to the right of the animal. Massive ontogenetic development of the sac in adult males is demonstrated, far greater than that seen in most other mysticetes. Histological analysis of the laryngeal sac shows the walls to be very muscular, abundantly innervated, and vascularized, indicating an active organ. Coiled blood vessels and nerves support the hypothesis that the sac undergoes extensive expansion and contraction. A possible association with the unique thoracic development of the species is suggested. The roles of the laryngeal sac and arytenoid cartilages in mysticete sound production are also discussed.

Dorsal skin color patterns among southern right whales (Eubalaena australis): genetic basis and evolutionary significance.

Distribution and inheritance of dorsal skin color markings among two populations of southern right whales (Eubalaena australis) suggest that two genes influence dorsal skin color. The grey-morph and partial-grey-morph phenotypes (previously known as partial albino and grey-blaze, respectively) appear to be controlled by an X-linked gene, whereas the white blaze appears controlled by an autosomal gene (recessive phenotype). Calving intervals, calf size, and length of sighting history data suggest that partial-grey-morph, white-blaze, and black cows experience similar levels of reproductive success. Grey-morph cows (XgXg) are rare or absent in the two populations, but this was not unexpected given observed population frequencies of grey-morph males (XgY) and partial-grey-morph females (XGXg). The proportion of partial-grey-morph calves produced by black cows (XGXG) suggests that the reproductive success of grey-morph males was equal to that of black males, however, larger sample sizes are required to determine whether grey-morph males tend to have shorter sighting histories. The reproductive success of white-blaze males appeared similar to that of black males among whales off Argentina. There were significantly fewer white-blaze calves than expected off South Africa, which could be due to white-blaze males experiencing reduced reproductive success or to sighting blases that result in white-marked calves being misidentified as black calves. The relative frequencies of both types of dorsal color markings varied between the South African and Argentinian right whale populations, suggesting limited nuclear gene flow between these populations; analyses using other nuclear markers are under way to confirm the extent of gene flow.

Genetic differentiation between parapatric 'nearshore' and 'offshore' populations of the bottlenose dolphin.

The existence of nearshore and offshore populations of the bottlenose dolphin has been documented throughout its range. In several cases the two regional forms have been shown to be morphologically distinct, although there is considerable overlap for most characters. The populations off the eastern coast of North America have been the subject of a long-term programme of research on their distribution and movements. In this study, we compare mitochondrial and nuclear genetic markers between dolphins classified as either nearshore or offshore type. These putative populations were found to be distinct at both nuclear and mitochondrial genetic markers. Further, the level of variation among the nearshore dolphins was reduced compared with the offshore population. A broader geographical comparison suggests a shared lineage between offshore dolphins from the western North Atlantic and both offshore and nearshore dolphins from the eastern Atlantic. These results are consistent with local differentiation based on habitat or resource specialization in the western North Atlantic, and suggest differences in the character of the nearshore/offshore distinction in different parts of the world.

Systematics of cetaceans using restriction site mapping of mitochondrial DNA.

Phylogenetic analysis of 14 cetacean species, including members from two baleen whale families and three toothed whale families, was undertaken using restriction site mapping of mitochondrial DNA and using cladistic and distance measures to infer phylogenies. The amount of between-taxa sequence divergence inferred from the data was lower than expected from the standard interpretation of the fossil record, but more consistent with some recent estimates of sequence divergence in cetacean mitochondrial DNA or nuclear DNA. This implies either that the rate of molecular evolution of cetacean DNA is much lower than that of other mammalian orders or that the fossil record of cetaceans requires reinterpretation. The incompleteness of the cetacean fossil record precludes resolution of the paradox at the present time. However, this discrepancy could in part be attributed to the sampling error inherent in the restriction site mapping technique, as comparative studies using the complete mtDNA genome and restriction site data of the blue and fin whales (genus Balaenoptera) indicate that the restriction site maps underestimate sequence divergence by about 40%. In contrast to a recent study suggesting that toothed whales were paraphyletic, with the sperm whales being more closely related to the rorquals than to the other toothed whales, the restriction data tend to support the monophyly of the baleen and the toothed whales, a finding which is consistent with a recent molecular-based study and with morphological and paleontological data. Topologies of the subfamily and generic levels are generally consistent with morphologically based schemes.

Persistent organochlorine residues in small cetaceans from the east and west coasts of southern Africa.

Organochlorine pollutant levels in the blubber of small cetaceans and ratios between concentrations of compounds, such as DDE/tDDT and tDDT/PCB, were used to identify the chronology of input of DDT and PCB into the coastal waters along the west and east coasts of southern Africa. Although regression analyses, with time as the only independent variable, do not show a statistically significant decline in tDDT from 1980 to 1987 in the common dolphin from the east coast of South Africa, there is a fairly strong indication that tDDT concentrations did not increase or even stay constant with time. tDDT concentrations in the bottlenose dolphin declined significantly from 1980 to 1987 (P < 0.05). The regression analyses for PCB in the bottlenose dolphin from the east coast show no decline (P > 0.05). The data from this study illustrate that it would take a long time in the coastal waters and open ocean before the restriction and prohibition on the production and use of organochlorines will take effect.