Survival rate, abundance, and residency of long-finned pilot whales in the Strait of Gibraltar

Long-finned pilot whales in the Strait of Gibraltar are distributed over the main shipping routes. This exposes them to risks of collisions and probable acoustic and physical disturbance. This species is also the target of whale-watching operations. The aim of this study was to estimate the annual population size, survival rate, and population growth rate of pilot whales occurring in the Strait and their inter-annual variation using photo-identification. A robust design was used to estimate all three parameters. A total of 10,784 individual pilot whale fins were photographed and analyzed. The population size estimation in summer ranged from a low of 147 individuals in 1999 to a high of 265 individuals in 2003. The annual population growth rate was estimated from mark recapture models to be 5.5%. The survival rate of adults was estimated at 0.982 (95% CI: 0.955–0.993). The same individuals have been observed between years. This suggests that this population is resident in the Strait, at least during summer. This study provides baseline knowledge prior to a predicted increase in shipping traffic throughout the main foraging area due to the opening in 2007 of a major shipping harbor along the Moroccan coast of the Strait.     

Long-term social structure of long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas</Emphasis>) in the Strait of Gibraltar

The Strait of Gibraltar is inhabited by around 216 pilot whales, which are present all year round, and nothing is known about their social structure. The aim of this study is to analyse the inter-individual association patterns within this pilot whales community to (1) provide an insight on their long-term social system and (2) to assess the relationship between sexes within this social system. Between 1999 and 2006, 23,004 km was sampled in the Strait of Gibraltar, and 4,887 images of dorsal fins of pilot whales were taken from 226 groups. The sex of 56 of the individuals could be determined genetically. The strength of the behavioural relationships between dyads of individuals was calculated, and the temporal aspects of the social structure were evaluated, showing in a non-random social structure made by constant companions. The preferred associations between individuals consisted in associations of males–females. Eight long-term units could be found with different degrees of association rates. Consequently, we propose that, in the Strait, the pilot whales exhibit a hierarchical social system composed of a population encompassing several clans of pilot whales each containing several pods. Pods will then be formed by several line units, similar to killer whale matrilineal units.     

Discerning between recurrent gene flow and recent divergence under a finite-site mutation model applied to North Atlantic and Mediterranean Sea fin whale (Balaenoptera physalus) populations

Genetic divergence among conspecific subpopulations can be due to either low recurrent gene flow or recent divergence and no gene flow. Here we present a modification of an earlier method developed by Nielsen and Wakeley (2001), which accommodates a finite-site mutation model, to assess which of the two models of divergence is most likely given the observed data. We apply the method to nucleotide sequence data collected from the variable part of the mitochondrial control region in fin whales (Balaenoptera physalus) from the Atlantic coast off Spain and the Mediterranean Sea. Our estimations strongly favor a model of recurrent gene flow over a model of recent divergence and zero gene flow. We estimated the migration rate at two females per generation. While the estimated rate is high by evolutionary standards, exchange rates of this order of magnitude is low from an ecological and conservation perspective and entirely consistent with the current paucity of fin whale sightings in the Strait of Gibraltar today. Intensive commercial shore-based whaling during the 1920s removed substantial numbers of fin whales in the Strait of Gibraltar and this local population has seemingly since failed to recover.     

STOMACH CONTENTS OF LONG-FINNED PILOT WHALES (GLOBICEPHALA MELAS) STRANDED ON THE U.S. MID-ATLANTIC COAST

Ten prey taxa were recorded from the stomach contents of eight long-finned pilot whales ( Globicepbala melas ) independently stranded along the U.S. mid-Atlantic coast. Relative importance of prey species was determined by methods that incorporate prey frequencies of occurrence, proportions of numerical abundance, and proportions of reconstructed mass. Separate analyses of trace (free, durable body parts representing well-digested prey items) and non-trace (relatively intact prey specimens) food material were conducted in order to address biases caused by differential rates of digestion and passage through the gastrointestinal tract. Different measures of prey importance yielded varying results, but the long-finned squid ( Loligo pealei ) was the most important prey species regardless of how prey importance was defined. Fishes were relatively unimportant in the diet. Our results indicate that the diets of western North Atlantic long-finned pilot whales differ substantially from what has been previously reported in the literature and that results from food-habits studies that utilize different techniques may not be comparable.     

Pilot Whales Attracted to Killer Whale Sounds: Acoustically-Mediated Interspecific Interactions in Cetaceans

In cetaceans’ communities, interactions between individuals of different species are often observed in the wild. Yet, due to methodological and technical challenges very little is known about the mediation of these interactions and their effect on cetaceans’ behavior. Killer whales (Orcinus orca) are a highly vocal species and can be both food competitors and potential predators of many other cetaceans. Thus, the interception of their vocalizations by unintended cetacean receivers may be particularly important in mediating interspecific interactions. To address this hypothesis, we conducted playbacks of killer whale vocalizations recorded during herring-feeding activity to free-ranging long-finned pilot whales (Globicephala melas). Using a multi-sensor tag, we were able to track the whales and to monitor changes of their movements and social behavior in response to the playbacks. We demonstrated that the playback of killer whale sounds to pilot whales induced a clear increase in group size and a strong attraction of the animals towards the sound source. These findings provide the first experimental evidence that the interception of heterospecific vocalizations can mediate interactions between different cetacean species in previously unrecognized ways.     

Mortality rate acceleration and post-reproductive lifespan in matrilineal whale species

The strength of selection to increase the span of a life stage is dependent upon individuals at that stage being able to contribute towards individual fitness and the probability of their surviving to that stage. Complete reproductive cessation and a long post-reproductive female lifespan as found in humans are also found in killer whale (Orcinus orca) and short-finned pilot whale (Globicephala macrorhynchus), but not in the long-finned pilot whale (Globicephala melaena). Each species forms kin-based, stable matrilineal groups and exhibits kin-directed behaviours that could increase inclusive fitness. Here, the initial mortality rate and mortality rate-doubling time of females of these three closely related whale species are compared. The initial mortality rate shows little variation among pilot whale species; however mortality rate accelerates almost twice as fast in the long-finned pilot whale as it does in killer whale and short-finned pilot whale. Selection for a long post-reproductive female lifespan in matrilineal whales may therefore be determined by the proportion of females surviving past the point of reproductive cessation.     

Repeated call sequences and behavioural context in long-finned pilot whales off Cape Breton,Nova Scotia,Canada

Repeated calls are part of the vocal repertoire of a diverse array of species, often presented in sequences that take time and effort on the part of the signal producer. Rhythmic repeated call sequences make up a significant portion of long-finned pilot whale (Globicephala melas) vocal production, yet the function of these sequences has not been investigated until now. In this study, we explored the relationship between behavioural context and the presence of these vocal sequences using recordings of a population of pilot whales found off Cape Breton, Nova Scotia, Canada. We used a binomial logit-link generalized linear model to look for possible predictors of the presence of repeated call sequences. They were more common in recordings of socializing whales than in those of whales in other behavioural states, and least common in resting whales. These vocal repetitions were also more common with larger group size. These results suggest that sequences function in maintaining contact and cohesion within this social species, possibly also serving in individual or group identification. The context of repeated call sequences indicate that they are not primarily mother–calf interactions, as they are heard just as commonly from groups without young. Future studies of pilot whale repeated call sequences should include individual-level behaviour and detailed acoustic calling context.     

Local and regional weather patterns influencing post‐breeding migration counts of soaring birds at the Strait of Gibraltar,Spain

Migration is a significant event in the annual cycle of many avian species. During migration birds face many challenges, including unfamiliar foraging and refuge habitats, resulting in a much higher rate of mortality during migration than during other seasons of the year. Weather may significantly affect a bird's decision to initiate migration, the course and pace of migration, and its survival during migration. Each of these influences may impact the counts of migrating birds at geographical convergence zones or bottlenecks. It is important to quantify the effect of short‐term weather on these counts to appropriately interpret and use such counts in other analyses. To this end, we aim to assess the effects of local and regional weather conditions on the migration counts of soaring birds at the Strait of Gibraltar during post‐breeding migration. We used information‐theoretic approaches to analyse the influence of local weather and weather in northern Spain on the migration counts of five soaring bird species from two count sites near the Strait of Gibraltar. Migration counts were higher on days with local northerly and westerly winds, often following a day of easterly winds, on days with local high pressure systems, and often following a day of lower pressure. Weather conditions in northern Spain influenced migration counts at the Strait of Gibraltar, but the effects were much weaker than local weather conditions. We confirm that short‐term weather conditions, locally and regionally, can influence migration counts and should thus be considered when these counts are used in other analyses.     

Small-scale spatial distributions of long-finned pilot whales change over time,but foraging hot spots are consistent: Significance for marine wildlife tourism management

Data collected opportunistically aboard marine wildlife tourism vessels are an inexpensive source of spatial information on the target species. Although these data are often challenging to analyze, they can be used to monitor spatiotemporal changes in species distribution and behavior. Disruptions from whale-watching vessels to behaviors such as foraging can be particularly harmful to cetaceans, but impacts could be reduced if areas essential for these sensitive behaviors are identified. We used data collected onboard whale-watching vessels to explore space-use patterns in long-finned pilot whales (Globicephala melas) off northern Cape Breton Island, Canada, an area where tourism is essential. Encounters with pilot whales between 2011–2016 occurred twice as far offshore than during 2003–2006 and 2008, and foraging activity decreased. Despite the changes in distribution and activity budgets, we identified two hot spots of foraging activity that persisted through time. These identified foraging hot spots comprised only a small proportion (20 km² ) of the range used by whale-watching vessels. Adaptive local management (e.g., voluntary codes of conduct) focused on limiting interactions in these energetically important areas may help reduce any potential impacts from whale-watching and promote the continued viability of the whale population and the tourism industry that relies on it.     

Trends of autumn counts at Iberian migration bottlenecks as a tool for monitoring continental populations of soaring birds in Europe

Migration monitoring may allow us to detect population trends over large geographic areas because the pattern of change in migrant counts may be expected to follow the pattern of change in population size. We analysed recent regional European population trends of migratory soaring birds from rates of change in migration counts over the Strait of Gibraltar (Spain) during the years (1999–2013). An additional bottleneck (Organbidexka, France) within the same migratory route and period was also considered. We estimated count trends by fitting a log-generalized linear model to the time series of each species counts. The counts in Organbidexka were used to test the consistency in the observed trends over the Strait of Gibraltar. Migration counts of White and Black Storks, Black Kites, Short-toed and Booted Eagles as well as Egyptian Vultures showed a linear increase over the Strait of Gibraltar throughout the 15-year period. In contrast, Honey Buzzard numbers remained stable. Trends were highly consistent with those recorded in Organbidexka. We suggest that the larger slopes for the trends in Organbidexka when compared with the Strait reflect an increasing tendency in these species to overwinter in southern Europe. A combination of complementary data sets collected at different bottleneck sites within the European–African flyway system may become a fundamental tool for the investigation of migratory patterns and population trends and changes of soaring migrant birds all over Europe.     

Genetic differentiation in the striped dolphin Stenella coeruleoalba from European waters according to mitochondrial DNA (mtDNA) restriction analysis

We used mitochondrial DNA (mtDNA) restriction analysis to study genetic variation in 98 striped dolphins (Stenella coeruleoalba) stranded on coasts from different European countries and from animals caught by fisheries. A total of 63 different restriction sites was mapped after digestion of mtDNA with 15 restriction endonucleases that yielded a total of 27 haplotypes. No haplotype was shared between Mediterranean and Atlantic areas. All the analyses indicate the existence of two different populations with a very limited gene flow across the Strait of Gibraltar.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

MORBILLIVIRUS INFECTION IN TWO SPECIES OF PILOT WHALE (GLOBICEPHALA SP.) FROM THE WESTERN ATLANTIC

We report evidence of enzootic morbillivirus infection among long-finned, Globicephala melas, and short-finned, G. macrorhynchus, pilot whales in the western Atlantic. A retrospective serologic survey, using five morbilliviruses, was carried out on 99 G. melas from 14 stranding events between 1982 and 1993 and from 25 G. macrorhynchus stranded in 5 events between 1986 and 1994. A blood sample was also obtained from an adult G. melas by-caught in the western North Atlantic. Tissues were collected from 24 G. melas and 15 G. macrorhynchus for histology and immunoperoxidase staining. Neutralizing antibody titers were found in 92 (92%) of 100 G. melas and 16 (64%) of 25 G. macrorynchus, and titers were highest against cetacean morbilliviruses. Seroprevalence was similar between age classes and sexes. The earliest evidence of infection was in a G. melas that stranded in 1982. Stable antibody titers were observed in pilot whales under rehabilitation for up to eight months. Clinical disease consistent with morbillivirus pneumonia was detected in a G. melas calf. Immunoperoxidase staining confirmed that viral antigen was present in the lesions. We propose that enzootic infection in pilot whales is facilitated by population size, social structure, and migration patterns. Furthermore, through mixing with other odontocetes, pilot whales could act as vectors through the Atlantic. Clinical morbillivirus infection may precipitate mass strandings of highly social odontocetes.     

Using a multi-disciplinary approach to identify a critically endangered killer whale management unit

A key goal for wildlife managers is identifying discrete, demographically independent conservation units. Previous genetic work assigned killer whales that occur seasonally in the Strait of Gibraltar (SoG) and killer whales sampled off the Canary Islands (CI) to the same population. Here we present new analyses of photo-identification and individual genotypes to assess the level of contemporary gene flow and migration between study areas, and analyses of biomarkers to assess ecological differences. We identified 47 different individuals from 5 pods in the SoG and 16 individuals in the CI, with no matches found between the areas. Mitochondrial DNA control region haplotype was shared by all individuals sampled within each pod, suggesting that pods have a matrifocal social structure typical of this species, whilst the lack of shared mitogenome haplotypes between the CI and SoG individuals suggests that there was little or no female migration between groups. Kinship analysis detected no close kin between CI and SoG individuals, and low to zero contemporary gene flow. Isotopic values and organochlorine pollutant loads also suggest ecological differences between study areas. We further found that one individual from a pod within the SoG not seen in association with the other four pods and identified as belonging to a potential migrant lineage by genetic analyses, had intermediate isotopic values and contaminant between the two study areas. Overall our results suggest a complex pattern of social and genetic structuring correlated with ecological variation. Consequently at least CI and SoG should be considered as two different management units. Understanding this complexity appears to be an important consideration when monitoring and understanding the viability of these management units. Understand the viability will help the conservation of these threatened management units.     

The fin whale Balaenoptera physalus (L. 1758) in the Mediterranean Sea

1. The ecology and status of fin whales Balaenoptera physalus in the Mediterranean Sea is reviewed. The species’ presence, morphology, distribution, movements, population structure, ecology and behaviour in this semi‐enclosed marine region are summarized, and the review is complemented with original, previously unpublished data. 2. Although the total size of the fin whale population in the Mediterranean is unknown, an estimate for a portion of the western basin, where most of the whales are known to live, was approximately 3500 individuals. High whale densities, comparable to those found in rich oceanic habitats, were found in well‐defined areas of high productivity. Most whales concentrate in the Ligurian‐Corsican‐Provençal Basin, where their presence is particularly noticeable during summer; however, neither their movement patterns throughout the region nor their seasonal cycle are clear. 3. Based on genetic studies, fin whales from the Mediterranean Sea are distinct from North Atlantic conspecifics, and may constitute a resident population, separate from those of the North Atlantic, despite the species’ historical presence in the Strait of Gibraltar. Fin whales are known to calve in the Mediterranean, with births peaking in November but occurring at lower rates throughout the year. They feed primarily on krill Meganyctiphanes norvegica which they capture by diving to depths in excess of 470 m. It is suggested that the extensive vertical migratory behaviour of its main prey may have influenced the social ecology of this population. 4. Known causes of mortality and threats, including collisions with vessels, entanglement in fishing gear, deliberate killing, disturbance, pollution and disease, are listed and discussed in view of the implementation of appropriate conservation measures to ensure the species’ survival in the region.     

Demographic parameters of a free-ranging deep-diving cetacean,the long-finned pilot whale

Demographic parameters provide baselines to estimate future population trajectories which can then be used in management decisions. The aim here was to estimate demographic parameters of long-finned pilot whale (Globicephala melas) from the Strait of Gibraltar by fitting mark-recapture models to photo-identification data of primary and secondary marked individuals. These parameters were used to forecast the future population trajectories in a population viability analysis (PVA) given different scenarios of demographic rates. Survival rate increased with age from 0.629, 95% CI [0.409, 0.805] for calves, 0.869, 95% CI [0.758, 0.934] for juveniles, to 0.972, 95% CI [0.953, 0.983] for adults. A preliminary mean observed interval of viable calves was 4.5 years. The PVA estimated the population would persist over 100 years with a 100% probability for all scenarios except those with lower 95% CI survival values, for which the probability of extinction reached 100%. Population growth rate was negative in all scenarios except those with 95% CI upper survival values. Interbirth interval and juvenile survival were found most influential and depended on the correct identification of secondary marked (e.g., calves and juveniles) individuals on a long-term basis. This population was found in a precarious state prior to a morbillivirus outbreak that might even more endanger its long-term viability.     

Comparing marine mammal acoustic habitats in Atlantic and Pacific sectors of the High Arctic: year-long records from Fram Strait and the Chukchi Plateau

During the International Polar Year (IPY), acoustic recorders were deployed on oceanographic moorings in Fram Strait and on the Chukchi Plateau, representing the first coordinated year-round sampling of underwater acoustic habitats at two sites in the High Arctic. Examination of species-specific marine mammal calls recorded from autumn 2008–2009 revealed distinctly different acoustic habitats at each site. Overall, the Fram Strait site was acoustically complex compared with the Chukchi Plateau site. In Fram Strait, calls from bowhead whales (Balaena mysticetus) and a variety of toothed whales (odontocetes) were recorded year-round, as were airgun pulses from seismic surveys. In addition, calls from blue whales (Balaenoptera musculus) and fin whales (B. physalus) were recorded from June to October and August to March, respectively. Conversely, at the Chukchi Plateau site, beluga (Delphinapterus leucas) and bowhead whale calls were recorded primarily from May to August, with airgun signals detected only in September–October. Ribbon seal (Phoca fasciata) calls were detected in October–November, with no marine mammals calls at all recorded from December to February. Of note, ice-adapted bearded seals (Erignathus barbatus) were recorded at both sites, primarily in spring and summer, corresponding with the mating season for that species. Differences in acoustic habitats between the two sites were related to contrasts in sea ice cover, temperature, patterns of ocean circulation and contributions from anthropogenic noise sources. These data provide a provisional baseline for the comparison of underwater acoustic habitats between Pacific and Atlantic sectors of the High Arctic.     

Biometry and phenology of two sibling Phylloscopus warblers on their circum-Mediterranean migrations

The Mediterranean Sea is known as an ecological barrier for numerous migratory birds flying from European breeding grounds to African wintering sites. Birds generally avoid migration over open sea and fly over land. In the Mediterranean Basin, few land bridges or bottlenecks for migratory birds exist. The narrowest are at the western and eastern extremes: the Strait of Gibraltar and Israel. Comparative studies between these locations are extremely rare to date. Therefore, in order to elucidate the differences between the two flyways, we compared data collected simultaneously for two sister leaf warbler species, the Bonelli’s Warbler complex, Phylloscopus bonelli and Phylloscopus orientalis, at ringing stations in the western Mediterranean Basin Gibraltar, and the eastern Eilat, Israel. Data on biometrics and passage dates of individuals trapped at Gibraltar and Eilat were used, and it was found that mean arrival date of Western Bonelli’s Warblers at Gibraltar was 15 days later than Eastern Bonelli’s Warblers at Eilat. Furthermore, Western Bonelli’s Warblers had shorter wings than Eastern Bonelli’s Warblers. On the other hand, birds in Eilat were in poorer body condition than individuals in Gibraltar. The comparison between geographically distant stop-over sites contributes to furthering our understanding of the development of migration strategies across ecological barriers in sibling species. Our study showed that populations that breed in southwestern Europe migrate through Gibraltar and winter in West Africa are able to accomplish migration in comparatively good body condition. This is in contrast to those that winter in East Africa, migrate through Israel and have to endure the combined challenge of crossing the Sahel, Sahara and Sinai deserts before reaching their breeding grounds across southeast Europe and southwest Asia. Hence, the discrepancies described between the western and the eastern flyway suggest that individuals in the west, in general, migrate shorter distances, have a physiologically less demanding crossing of the North African deserts and appear to stage before their crossing the Strait of Gibraltar, a privilege unavailable to the migrants of the eastern flyway.