Trace element concentrations in common dolphins (Delphinus delphis) in the Celtic Seas ecoregion: Interelement relationships and effects of life history and health status.

Given the increased extraction of trace elements for use by new and emerging technologies, monitoring the environmental fate and potential effects of these compounds within the aquatic environment has never been more critical. Here, hepatic trace element concentrations were assessed in a key sentinel predator, the common dolphin (Delphinus delphis), using a long-term dataset. Variation in concentrations were assessed in relation to other elements, time period, decomposition state, sex, age, total body length, sexual maturity and nutritional status, and cause of death. Additionally, mercury toxicity thresholds for evaluating risk were reviewed and employed. Concentrations of elements which bioaccumulate, THg, MeHg, Cd, and Pb, in addition to Se and V, were strongly correlated with age, and/or body length. An association was observed between Zn concentrations and disease status, with significantly higher concentrations measured in individuals that died from infectious disease, compared to other causes. Strong inter-elemental relationships were detected, namely between Hg and Se, MeHg and Se, Cd and Se, and Cu and Zn. While THg:Se molar ratio values were observed to increase with age and body length, approaching equimolarity. THg was largely comprised of inorganic Hg in older individuals, potentially bound to Se, therefore the effects from THg toxicity may possibly be less important than originally assumed. In contrast, higher MeHg:Hg ratio values were reported in juveniles, suggesting a poorer efficiency in demethylation and a higher sensitivity. The generation of data on proportions of hepatic MeHg and inorganic Hg is highly informative to both future toxicity threshold assessments within pollutant indicator assessments, and to understanding the ultimate fate of mercury in the marine web.

Population genomics of the white-beaked dolphin (Lagenorhynchus albirostris): Implications for conservation amid climate-driven range shifts.

Climate change is rapidly affecting species distributions across the globe, particularly in the North Atlantic. For highly mobile and elusive cetaceans, the genetic data needed to understand population dynamics are often scarce. Cold-water obligate species such as the white-beaked dolphin (Lagenorhynchus albirostris) face pressures from habitat shifts due to rising sea surface temperatures in addition to other direct anthropogenic threats. Unravelling the genetic connectivity between white-beaked dolphins across their range is needed to understand the extent to which climate change and anthropogenic pressures may impact species-wide genetic diversity and identify ways to protect remaining habitat. We address this by performing a population genomic assessment of white-beaked dolphins using samples from much of their contemporary range. We show that the species displays significant population structure across the North Atlantic at multiple scales. Analysis of contemporary migration rates suggests a remarkably high connectivity between populations in the western North Atlantic, Iceland and the Barents Sea, while two regional populations in the North Sea and adjacent UK and Irish waters are highly differentiated from all other clades. Our results have important implications for the conservation of white-beaked dolphins by providing guidance for the delineation of more appropriate management units and highlighting the risk that local extirpation may have on species-wide genetic diversity. In a broader context, this study highlights the importance of understanding genetic structure of all species threatened with climate change-driven range shifts to assess the risk of loss of species-wide genetic diversity.

Ancient dolphin genomes reveal rapid repeated adaptation to coastal waters.

Parallel evolution provides strong evidence of adaptation by natural selection due to local environmental variation. Yet, the chronology, and mode of the process of parallel evolution remains debated. Here, we harness the temporal resolution of paleogenomics to address these long-standing questions, by comparing genomes originating from the mid-Holocene (8610-5626 years before present, BP) to contemporary pairs of coastal-pelagic ecotypes of bottlenose dolphin. We find that the affinity of ancient samples to coastal populations increases as the age of the samples decreases. We assess the youngest genome (5626 years BP) at sites previously inferred to be under parallel selection to coastal habitats and find it contained coastal-associated genotypes. Thus, coastal-associated variants rose to detectable frequencies close to the emergence of coastal habitat. Admixture graph analyses reveal a reticulate evolutionary history between pelagic and coastal populations, sharing standing genetic variation that facilitated rapid adaptation to newly emerged coastal habitats.

Estimated mortality of the highly pathogenic avian influenza pandemic on northern gannets (Morus bassanus) in southwest Ireland.

The 2022 highly pathogenic avian influenza (HPAI) outbreak that occurred in many European countries affected several seabird species. Among them, northern gannets (Morus bassanus) were particularly impacted. We conducted aerial surveys in waters around the two largest gannet colonies in southwest Ireland (Little Skellig and Bull Rock, together representing 87% of the national population) in September 2022. During surveys dead and alive northern gannets were counted on survey effort. A total of 184 dead gannets were recorded on survey effort, representing 3.74% of the total number of gannets recorded. We estimated the abundance of dead gannets in the surveyed area at 1526 (95% confidence intervals (CIs) 1450-1605) individuals. The percentage of dead gannets observed was used to estimate a minimum local population mortality of 3126 (95% CIs 2993-3260) individuals across both colonies. Aerial surveys provided key information on gannet mortality from HPAI at sea. The study provides the first estimate of gannet mortality in the two largest gannetries in Ireland.

What the F-POD? Comparing the F-POD and C-POD for monitoring of harbor porpoise (Phocoena phocoena).

Passive acoustic monitoring (PAM) is a cost-effective method for monitoring cetacean populations compared with techniques such as aerial and ship-based surveys. The Cetacean POrpoise Detector (C-POD) has become an integral tool in monitoring programs globally for over a decade, providing standardized metrics of occurrence that can be compared across time and space. However, the phasing out of C-PODs following the development of the new Full waveform capture POD (F-POD) with increased sensitivity, improved train detection, and reduced false-positive rates represents an important methodological change in data collection, particularly when being introduced into existing monitoring programs. Here, we compare the performance of the C-POD with that of its successor, the F-POD, co-deployed in a field setting for 15 months, to monitor harbor porpoise (Phocoena phocoena). While similar temporal trends in detections were found for both devices, the C-POD detected only 58% of the detection-positive minutes (DPM), recorded by the F-POD. Differences in detection rates were not consistent through time making it difficult to apply a correction factor or directly compare results obtained from the two PODs. Generalized additive models (GAMs) were used to test whether these differences in detection rates would have an effect on analyses of temporal patterns and environmental drivers of occurrence. No differences were found in seasonal patterns or the environmental correlates of porpoise occurrence (month, diel period, temperature, environmental noise, and tide). However, the C-POD failed to detect sufficient foraging rates to identify temporal patterns in foraging behavior, which were shown by the F-POD. Our results suggest that the switch to F-PODs will have little effect on determining broad-scale seasonal patterns of occurrence but may improve our understanding of fine-scale behaviors such as foraging. We highlight how care must be taken interpreting F-POD results as indicative of increased occurrence when used in time-series analysis.

Speciation in the deep: genomics and morphology reveal a new species of beaked whale Mesoplodon eueu.

The deep sea has been described as the last major ecological frontier, as much of its biodiversity is yet to be discovered and described. Beaked whales (ziphiids) are among the most visible inhabitants of the deep sea, due to their large size and worldwide distribution, and their taxonomic diversity and much about their natural history remain poorly understood. We combine genomic and morphometric analyses to reveal a new Southern Hemisphere ziphiid species, Ramari's beaked whale, Mesoplodon eueu, whose name is linked to the Indigenous peoples of the lands from which the species holotype and paratypes were recovered. Mitogenome and ddRAD-derived phylogenies demonstrate reciprocally monophyletic divergence between M. eueu and True's beaked whale (M. mirus) from the North Atlantic, with which it was previously subsumed. Morphometric analyses of skulls also distinguish the two species. A time-calibrated mitogenome phylogeny and analysis of two nuclear genomes indicate divergence began circa 2 million years ago (Ma), with geneflow ceasing 0.35-0.55 Ma. This is an example of how deep sea biodiversity can be unravelled through increasing international collaboration and genome sequencing of archival specimens. Our consultation and involvement with Indigenous peoples offers a model for broadening the cultural scope of the scientific naming process.

Modelling ecosystem dynamics to assess the effect of coastal fisheries on cetacean species.

The expansion of fisheries and its increased efficiency are causing severe detrimental impacts on marine species and ecosystems, that can be categorised into operational and ecological effects. While impacts directly caused by fishing activities have been extensively documented, it is difficult to set an empirical link between fisheries and changes in predator biomass and abundance. Therefore, exploring the functioning of ecosystems as a whole, the interactions between the different species within them and the impact of human activities, is key to understanding the ecological effects of fisheries on top predators and ecosystems, and to develop effective conservation measures, while ensuring a more sustainable exploitation of fishing resources. For instance, mass balance models, such as Ecopath with Ecosim, have proven to be a useful tool to develop more holistic fisheries management and conservation strategies. In this study, Ecopath with Ecosim was used to investigate the temporal dynamics of the Rías Baixas shelf ecosystem (North-West Spain) between 2005 and 2017. Additionally, nine 30-year forward projecting simulations covering the period 2018-2047 were developed to examine the effects of differing fisheries management strategies on common dolphins (Delphinus delphis), bottlenose dolphins (Tursiops truncatus) and harbour porpoises (Phocoena phocoena). Results from these models suggest that when intense fishing increases it poses a major threat to the conservation of these top predators in the area, by reducing the variety of their available prey and potentially enhancing competition amongst them. The study highlights the applicability of Ecopath with Ecosim to develop cetacean conservation measures and despite its small spatial scale, it provides a general framework that can be used to assess cetacean conservation in larger and impacted areas.

Food-web interactions in a coastal ecosystem influenced by upwelling and terrestrial runoff off North-West Spain.

Ecopath with Ecosim has been used to create mass-balance models of different type of ecosystems around the world to explore and analyse their functioning and structure. This modelling framework has become a key tool in the ecosystem approach to fisheries management, by providing a more comprehensive and holistic understanding of the interactions between the different species. Additionally, Ecopath with Ecosim has provided a useful framework to study ecosystem maturity, changes in the ecosystem functioning over time and the impact of fisheries and aquaculture on the ecosystem, among other aspects. The present work explores the ecosystem functioning and structure in an anthropogenically impacted coastal area, influenced by seasonal coastal upwelling and high input of nutrients from rias (ancient drowned tectonic valleys) off North-West Spain. A mass-balance model with 23 functional groups was created using Ecopath to study the trophic interactions in the ecosystem during the post-upwelling period (August to October) in 2017. The model described an immature, wasp-waist ecosystem, that shared characteristics of ecosystems found in upwelling areas and ecosystems found in fjords or coastal embayments. Deeper analyses highlighted the importance of small planktivorous pelagic fish as a keystone functional group, and of zooplankton, blue whiting (Micromesistius poutassou) and phytoplankton as structuring groups in the ecosystem. Additionally, the study revealed that the existing fishing pressure on species of intermediate-high trophic levels could alter ecosystem functioning and structure, and ultimately affect top predators in the area. Findings of this study provide baseline information in ecosystem functioning and structure in the area and highlight the need to deeper study the effects of fisheries and their potential impacts on top predators.

Postglacial Colonization of Northern Coastal Habitat by Bottlenose Dolphins: A Marine Leading-Edge Expansion?

Oscillations in the Earth's temperature and the subsequent retreating and advancing of ice-sheets around the polar regions are thought to have played an important role in shaping the distribution and genetic structuring of contemporary high-latitude populations. After the Last Glacial Maximum (LGM), retreating of the ice-sheets would have enabled early colonizers to rapidly occupy suitable niches to the exclusion of other conspecifics, thereby reducing genetic diversity at the leading-edge. Bottlenose dolphins (genus Tursiops) form distinct coastal and pelagic ecotypes, with finer-scale genetic structuring observed within each ecotype. We reconstruct the postglacial colonization of the Northeast Atlantic (NEA) by bottlenose dolphins using habitat modeling and phylogenetics. The AquaMaps model hindcasted suitable habitat for the LGM in the Atlantic lower latitude waters and parts of the Mediterranean Sea. The time-calibrated phylogeny, constructed with 86 complete mitochondrial genomes including 30 generated for this study and created using a multispecies coalescent model, suggests that the expansion to the available coastal habitat in the NEA happened via founder events starting ~15 000 years ago (95% highest posterior density interval: 4 900-26 400). The founders of the 2 distinct coastal NEA populations comprised as few as 2 maternal lineages that originated from the pelagic population. The low effective population size and genetic diversity estimated for the shared ancestral coastal population subsequent to divergence from the pelagic source population are consistent with leading-edge expansion. These findings highlight the legacy of the Late Pleistocene glacial cycles on the genetic structuring and diversity of contemporary populations.

Spatial variation in a top marine predator's diet at two regionally distinct sites.

In ecological studies it is often assumed that predator foraging strategies and resource use are geographically and seasonally homogeneous, resulting in relatively static trophic relationships. However, certain centrally placed foragers (e.g. seals) often have terrestrial sites for breeding, resting, and moulting that are geographically distinct, and associated with different habitat types. Therefore, accurate estimations of predator diet at relevant spatial and temporal scales are key to understanding energetic requirements, predator-prey interactions and ecosystem structure. We investigate geographic variation in the diet of grey seals (Halichoerus grypus), a relatively abundant and widely distributed central place forager, to provide insights into geographic variation in resource use. Prey composition was identified using scat samples collected over concurrent timescales and a multivariate approach was used to analyse diet from two contrasting habitats. Regional differences in prey assemblages occurred within all years (2011-2013) and all seasons (ANOSIM, all p<0.05), apart from in winter. Telemetry data were used to identify core foraging areas and habitats most likely associated with scat samples collected at the two haul-out sites. Regional differences in the diet appear to reflect regional differences in the physical habitat features, with seals foraging in deeper waters over sandy substrates showing a higher prevalence of pelagic and bentho-pelagic prey species such as blue whiting and sandeels. Conversely, seals foraging in comparatively shallow waters had a greater contribution of demersal and groundfish species such as cephalopods and flatfish in their diet. We suggest that shallower waters enable seals to spend more time foraging along the benthos while remaining within aerobic dive limits, resulting in more benthic species in the diet. In contrast, the diet of seals hauled-out in areas adjacent to deeper waters indicates that either seals engage in a more pelagic foraging strategy, or that seals can spend less time at the benthos, resulting in comparatively more pelagic prey recovered in the diet. The substantial differences in prey assemblages over a small spatial scale (<300 km) demonstrates the importance of using regionally-specific diet information in ecosystem-based models to better account for different trophic interactions.

Quantifying dispersal between marine protected areas by a highly mobile species, the bottlenose dolphin, Tursiops truncatus.

The functioning of marine protected areas (MPAs) designated for marine megafauna has been criticized due to the high mobility and dispersal potential of these taxa. However, dispersal within a network of small MPAs can be beneficial as connectivity can result in increased effective population size, maintain genetic diversity, and increase robustness to ecological and environmental changes making populations less susceptible to stochastic genetic and demographic effects (i.e., Allee effect). Here, we use both genetic and photo-identification methods to quantify gene flow and demographic dispersal between MPAs of a highly mobile marine mammal, the bottlenose dolphin Tursiops truncatus. We identify three populations in the waters of western Ireland, two of which have largely nonoverlapping core coastal home ranges and are each strongly spatially associated with specific MPAs. We find high site fidelity of individuals within each of these two coastal populations to their respective MPA. We also find low levels of demographic dispersal between the populations, but it remains unclear whether any new gametes are exchanged between populations through these migrants (genetic dispersal). The population sampled in the Shannon Estuary has a low estimated effective population size and appears to be genetically isolated. The second coastal population, sampled outside of the Shannon, may be demographically and genetically connected to other coastal subpopulations around the coastal waters of the UK. We therefore recommend that the methods applied here should be used on a broader geographically sampled dataset to better assess this connectivity.

Using tagging data and aerial surveys to incorporate availability bias in the abundance estimation of blue sharks (Prionace glauca).

There is worldwide concern about the status of elasmobranchs, primarily as a result of overfishing and bycatch with subsequent ecosystem effects following the removal of top predators. Whilst abundant and wide-ranging, blue sharks (Prionace glauca) are the most heavily exploited shark species having suffered marked declines over the past decades, and there is a call for robust abundance estimates. In this study, we utilized depth data collected from two blue sharks using pop-up satellite archival tags, and modelled the proportion of time the sharks were swimming in the top 1-meter layer and could therefore be detected by observers conducting aerial surveys. The availability models indicated that the tagged sharks preferred surface waters whilst swimming over the continental shelf and during daytime, with a model-predicted average proportion of time spent at the surface of 0.633 (SD = 0.094) for on-shelf, and 0.136 (SD = 0.075) for off-shelf. These predicted values were then used to account for availability bias in abundance estimates for the species over a large area in the Northeast Atlantic, derived through distance sampling using aerial survey data collected in 2015 and 2016 and modelled with density surface models. Further, we compared abundance estimates corrected with model-predicted availability to uncorrected estimates and to estimates that incorporated the average time the sharks were available for detection. The mean abundance (number of individuals) corrected with modelled availability was 15,320 (CV = 0.28) in 2015 and 11,001 (CV = 0.27) in 2016. Depending on the year, these estimates were ~7 times higher compared to estimates without the bias correction, and ~3 times higher compared to the abundances corrected with average availability. When the survey area contains habitat heterogeneity that may affect surfacing patterns of animals, modelling animals' availability provides a robust alternative to correcting for availability bias and highlights the need for caution when applying "average" correction factors.

Estimating common dolphin bycatch in the pole-and-line tuna fishery in the Azores.

Small-scale artisanal fisheries can have a significant negative impact in cetacean populations. Cetacean bycatch has been documented in the pole-and-line tuna fishery in the Azores with common dolphins being the species more frequently taken. Based on data collected by observers on ∼50% of vessels operating from 1998 to 2012, we investigate the influence of various environmental and fisheries-related factors in common dolphin bycatch and calculate fleet-wide estimates of total bycatch using design-based and model-based methods. Over the 15-year study dolphin bycatch occurred in less than 0.4% of the observed fishing events. Generalized additive modelling results suggest a significant relationship between common dolphin bycatch and duration of fishing events, sea surface temperature and location. Total bycatch calculated from the traditional stratified ratio estimation approach was 196 (95% CI: 186-205), while the negative binomial GAM estimated 262 (95% CI: 249-274) dolphins. Bycatch estimates of common dolphin were similar using statistical approaches suggesting that either of these methods may be used in future bycatch assessments for this fishery. Our work shows that rates of common dolphin bycatch in the pole-and-line tuna fishery in the Azores are low, despite considerable variations between years. Dolphins caught were released alive although the fate of these individuals is unknown. Continued monitoring will provide a better understanding of dolphin bycatch and more accurate estimates essential in the development of potential mitigation measures.

Incidence of marine debris in cetaceans stranded and bycaught in Ireland: Recent findings and a review of historical knowledge.

Interactions between marine mammals and plastic debris have been the focus of studies for many years. Examples of interactions include entanglement in discarded fishing items or the presence of ingested debris in digestive tracts. Plastics, including microplastics, are a form of marine debris globally distributed in coastal areas, oceanic waters and deep seas. Cetaceans which strand along the coast present a unique opportunity to study interactions between animals with macro- and microplastics. A combination of novel techniques and a review of historical data was used to complete an extensive study of cetaceans interacting with marine debris within Irish waters. Of the 25 species of marine mammals reported in Irish waters, at least 19 species were reported stranded between 1990 and 2015 (n = 2934). Two hundred and forty-one of the stranded cetaceans presented signs of possible entanglement or interactions with fisheries. Of this number, 52.7% were positively identified as bycatch or as entangled in fisheries items, 26.6% were classified as mutilated and 20.7% could not be related to fisheries but showed signs of entanglement. In addition, 274 cetaceans were recorded as by-catch during observer programmes targeting albacore tuna. Post-mortem examinations were carried out on a total of 528 stranded and bycaught individuals and 45 (8.5%) had marine debris in their digestive tracts: 21 contained macrodebris, 21 contained microdebris and three had both macro- and microdebris. Forty percent of the ingested debris were fisheries related items. All 21 individuals investigated with the novel method for microplastics contained microplastics, composed of fibres (83.6%) and fragments (16.4%). Deep diving species presented more incidences of macrodebris ingestion but it was not possible to investigate this relationship to ecological habitat. More research on the plastic implications to higher trophic level organisms is required to understand the effects of these pollutants.

New insights into ocean sunfish (Mola mola) abundance and seasonal distribution in the northeast Atlantic.

The ocean sunfish, Mola mola, is the largest teleost fish in the world. Despite being found in all oceans of the world, little is known about its abundance and factors driving its distribution. In this study we provide the first abundance estimates for sunfish in offshore waters in the northeast Atlantic and the first record of extensive sunfish presence in these waters year-round. Abundance estimates and predictive distributions for sunfish in approximately 300,000 km² of the northeast Atlantic were derived from large scale offshore aerial surveys in 2015-2016 using distance sampling techniques. Generalized additive models of sunfish density were fitted to survey data from 17,360 km of line transect effort resulting in minimum abundance estimates of 12,702 (CI: 9,864-16,357) in the summer (Density = 0.043 ind/km²) and 8,223 individuals (CI: 6,178-10,946) (Density = 0.028 ind/km²) in the winter. Density surface models predicted seasonal shifts in distribution and highlighted the importance of the mixed layer depth, possibly related to thermoregulation following deep foraging dives. The abundance estimate and estimated daily consumption of 2,600 tonnes of jellyfish in the northeast Atlantic highlights the need to re-assess the importance of this species in the pelagic ecosystem, and its role in top-down control of jellyfish blooms.

Evidence for distinct coastal and offshore communities of bottlenose dolphins in the north east Atlantic.

Bottlenose dolphin stock structure in the northeast Atlantic remains poorly understood. However, fine scale photo-id data have shown that populations can comprise multiple overlapping social communities. These social communities form structural elements of bottlenose dolphin (Tursiops truncatus) [corrected] populations, reflecting specific ecological and behavioural adaptations to local habitats. We investigated the social structure of bottlenose dolphins in the waters of northwest Ireland and present evidence for distinct inshore and offshore social communities. Individuals of the inshore community had a coastal distribution restricted to waters within 3 km from shore. These animals exhibited a cohesive, fission-fusion social organisation, with repeated resightings within the research area, within a larger coastal home range. The offshore community comprised one or more distinct groups, found significantly further offshore (>4 km) than the inshore animals. In addition, dorsal fin scarring patterns differed significantly between inshore and offshore communities with individuals of the offshore community having more distinctly marked dorsal fins. Specifically, almost half of the individuals in the offshore community (48%) had characteristic stereotyped damage to the tip of the dorsal fin, rarely recorded in the inshore community (7%). We propose that this characteristic is likely due to interactions with pelagic fisheries. Social segregation and scarring differences found here indicate that the distinct communities are likely to be spatially and behaviourally segregated. Together with recent genetic evidence of distinct offshore and coastal population structures, this provides evidence for bottlenose dolphin inshore/offshore community differentiation in the northeast Atlantic. We recommend that social communities should be considered as fundamental units for the management and conservation of bottlenose dolphins and their habitat specialisations.

Characteristics of fishing operations, environment and life history contributing to small cetacean bycatch in the northeast Atlantic.

Fisheries bycatch is a key threat to cetacean species globally. Managing the impact requires an understanding of the conditions under which animals are caught and the sections of the population affected. We used observer data collected on an albacore tuna gillnet fishery in the northeast Atlantic, to assess operational and environmental factors contributing to bycatch of common and striped dolphins, using generalised linear models and model averaging. Life history demographics of the captured animals were also investigated. In both species, young males dominated the catch. The age ratio of common dolphins was significantly different from that estimated for the population in the region, based on life tables (G = 17.1, d.f. = 2, p = 0.002). Skewed age and sex ratios may reflect varying vulnerability to capture, through differences in behaviour or segregation in populations. Adult females constituted the second largest portion of the bycatch for both species, with potential consequences for population sustainability. Depth was the most important parameter influencing bycatch of both species and reflected what is known about common and striped dolphin habitat use in the region as the probability of catching common dolphins decreased, and striped dolphins increased, with increasing depth. Striped dolphin capture was similarly influenced by the extent to which operations were conducted in daylight, with the probability of capture increasing with increased operations in the pre-sunset and post-sunrise period, potentially driven by increased ability of observers to record animals during daylight operations, or by diurnal movements increasing contact with the fishery. Effort, based on net length and soak time, had little influence on the probability of capturing either species. Our results illustrate the importance of assessing the demographic of the animals captured during observer programmes and, perhaps more importantly, suggest that effort restrictions alone may not be sufficient to eradicate bycatch in areas where driftnets and small cetaceans co-occur.

Postglacial climate changes and rise of three ecotypes of harbour porpoises, Phocoena phocoena, in western Palearctic waters.

Despite no obvious barriers to gene flow in the marine realm, environmental variation and ecological specializations can lead to genetic differentiation in highly mobile predators. Here, we investigated the genetic structure of the harbour porpoise over the entire species distribution range in western Palearctic waters. Combined analyses of 10 microsatellite loci and a 5085 base-pair portion of the mitochondrial genome revealed the existence of three ecotypes, equally divergent at the mitochondrial genome, distributed in the Black Sea (BS), the European continental shelf waters, and a previously overlooked ecotype in the upwelling zones of Iberia and Mauritania. Historical demographic inferences using approximate Bayesian computation (ABC) suggest that these ecotypes diverged during the last glacial maximum (c. 23-19 kilo-years ago, kyrbp). ABC supports the hypothesis that the BS and upwelling ecotypes share a more recent common ancestor (c. 14 kyrbp) than either does with the European continental shelf ecotype (c. 28 kyrbp), suggesting they probably descended from the extinct populations that once inhabited the Mediterranean during the glacial and post-glacial period. We showed that the two Atlantic ecotypes established a narrow admixture zone in the Bay of Biscay during the last millennium, with highly asymmetric gene flow. This study highlights the impacts that climate change may have on the distribution and speciation process in pelagic predators and shows that allopatric divergence can occur in these highly mobile species and be a source of genetic diversity.

Structure, material characteristics and function of the upper respiratory tract of the pygmy sperm whale.

Cetaceans are neckless, so the trachea is very short. The upper respiratory tract is separate from the mouth and pharynx, and the dorsal blowhole connects, via the vestibular and nasopalatine cavities, directly to the larynx. Toothed cetaceans (Odontoceti) are capable of producing sounds at depth, either for locating prey or for communication. It has been suggested that during dives, air from the lungs and upper respiratory tract can be moved to the vestibular and nasal cavities to permit sound generation to continue when air volume within these cavities decreases as ambient pressure rises. The pygmy sperm whale, Kogia breviceps, is a deep diver (500-1000 m) that is known to produce hunting clicks. Our study of an immature female shows that the upper respiratory tract is highly asymmetrical: the trachea and bronchi are extremely compressible, whereas the larynx is much more rigid. Laryngeal and tracheal volumes were established. Calculations based on Boyle's Law imply that all air from the lungs and bronchi would be transferred to the larynx and trachea by a depth of 270 m and that the larynx itself could not accommodate all respiratory air mass at a depth of 1000 m. This suggests that no respiratory air would be available for vocalisation. However, the bronchi, trachea and part of the larynx have a thick vascular lining featuring large, thin-walled vessels. We propose that these vessels may become dilated during dives to reduce the volume of the upper respiratory tract, permitting forward transfer of air through the larynx.

Genetic differentiation among North Atlantic killer whale populations.

Population genetic structure of North Atlantic killer whale samples was resolved from differences in allele frequencies of 17 microsatellite loci, mtDNA control region haplotype frequencies and for a subset of samples, using complete mitogenome sequences. Three significantly differentiated populations were identified. Differentiation based on microsatellite allele frequencies was greater between the two allopatric populations than between the two pairs of partially sympatric populations. Spatial clustering of individuals within each of these populations overlaps with the distribution of particular prey resources: herring, mackerel and tuna, which each population has been seen predating. Phylogenetic analyses using complete mitogenomes suggested two populations could have resulted from single founding events and subsequent matrilineal expansion. The third population, which was sampled at lower latitudes and lower density, consisted of maternal lineages from three highly divergent clades. Pairwise population differentiation was greater for estimates based on mtDNA control region haplotype frequencies than for estimates based on microsatellite allele frequencies, and there were no mitogenome haplotypes shared among populations. This suggests low or no female migration and that gene flow was primarily male mediated when populations spatially and temporally overlap. These results demonstrate that genetic differentiation can arise through resource specialization in the absence of physical barriers to gene flow.