Diane Gendron

Insights into the Underwater Diving, Feeding, and Calling Behavior of Blue Whales from a Suction-Cup-Attached Video-Imaging Tag (CRITTERCAM)

Abstract : We examined the underwater behavior of blue whales using a suction-cup-attached video-imaging instrument (CRITTERCAM). We made 13 successful deployments (defined as tag duration of 15 min and successful recovery of the tag and data) totaling 19 hours of CRITTERCAMs on blue whales off California and in the Sea of Cortez from spring through fall (26 February to 30 September) between 1999 and 2003 . Whale diving depth and behavior varied widely by region and period, although deployments on different individuals in the same area and period often showed very similar feeding behavior. One deployment extending into night showed a diurnal shift in diving behavior with progressively shallower feeding dives as it became dark, with shift to shallow, apparently non-feeding dives during the night. Data and video from tags demonstrated that the characteristic series of vertical movements blue whales make at depth are lunges into dense aggregations of krill. These krill were visible streaming by the camera immediately before these lunges and more clearly when the whales? forward motion stopped as a result of the lunge. The progression of events leading up to and during the lunge could be documented from the head movement of whales and occasional views of the expanding throat pleats or lower jaw, and by changes in flow noise past the tag, indicating a rapid deceleration. One set of deployments in the Southern California Bight revealed consistent feeding at depths of 250-300 m, deeper than has been previously reported for blue whales. A loud blue whale vocalization was heard on only one deployment on a male blue whale in an interacting trio of animals.

Stable isotopes on jumbo squid (Dosidicus gigas) beaks to esitmate its trophic positon: comparison between stomach contents and stable isotopes

Stomach contents and carbon (C) and nitrogen (N) stable isotope analysis were used to evaluate trophic relationships of jumbo squid, Dosidicus gigas . Buccal masses, beaks and stomach contents of large and medium maturing-sized jumbo squid and muscle from its main prey, the myctophid Benthosema panamense , were collected in the Gulf of California, Mexico during 1996, 1997 and 1999. Both the quantified C and N-isotope ratios in muscle, and stomach content analysis revealed that larger-sized maturing squid showed a higher trophic position than medium-sized individuals. However, a discrepancy between stomach contents versus stable isotope analyses was found in evaluating trophic relationships. Simple dilution models as a function of growth were used to estimate the C and N renewal dietary shift for jumbo squid. Estimates of the initial C and N pools in D. gigas with an initial age of 70 days and 210 days indicated isotopic shifts of 32% after a threefold biomass increase and 25% after a fourfold biomass increase, respectively. Additionally, beak samples of jumbo squid were evaluated as an alternative tissue to estimate squid trophic position using stable isotopes. The results showed a significant correlation between stable isotope ratios from muscle and beak samples. Muscle isotope values were higher than beak by 1% and 4% for δ 13 C and δ 15 N respectively. A test with jumbo squid beaks collected from a stomach of a stranded sperm whale confirmed the viability of this method.

Acute sun damage and photoprotective responses in whales

Rising levels of ultraviolet radiation (UVR) secondary to ozone depletion are an issue of concern for public health. Skin cancers and intraepidermal dysplasia are increasingly observed in individuals that undergo chronic or excessive sun exposure. Such alterations of skin integrity and function are well established for humans and laboratory animals, but remain unexplored for mammalian wildlife. However, effects are unlikely to be negligible, particularly for species such as whales, whose anatomical or life-history traits force them to experience continuous sun exposure. We conducted photographic and histological surveys of three seasonally sympatric whale species to investigate sunburn and photoprotection. We find that lesions commonly associated with acute severe sun damage in humans are widespread and that individuals with fewer melanocytes have more lesions and less apoptotic cells. This suggests that the pathways used to limit and resolve UVR-induced damage in humans are shared by whales and that darker pigmentation is advantageous to them. Furthermore, lesions increased significantly in time, as would be expected under increasing UV irradiance. Apoptosis and melanocyte proliferation mirror this trend, suggesting that whales are capable of quick photoprotective responses. We conclude that the thinning ozone layer may pose a risk to the health of whales and other vulnerable wildlife.

Whales Use Distinct Strategies to Counteract Solar Ultraviolet Radiation

A current threat to the marine ecosystem is the high level of solar ultraviolet radiation (UV). Large whales have recently been shown to suffer sun-induced skin damage from continuous UV exposure. Genotoxic consequences of such exposure remain unknown for these long-lived marine species, as does their capacity to counteract UV-induced insults. We show that UV exposure induces mitochondrial DNA damage in the skin of seasonally sympatric fin, sperm, and blue whales and that this damage accumulates with age. However, counteractive molecular mechanisms are markedly different between species. For example, sperm whales, a species that remains for long periods at the sea surface, activate genotoxic stress pathways in response to UV exposure whereas the paler blue whale relies on increased pigmentation as the season progresses. Our study also shows that whales can modulate their responses to fluctuating levels of UV, and that different evolutionary constraints may have shaped their response strategies.

Fine-Scale Population Structure of Blue Whale Wintering Aggregations in the Gulf of California

Population differentiation in environments without well-defined geographical barriers represents a challenge for wildlife management. Based on a comprehensive database of individual sighting records (1988–2009) of blue whales from the winter/calving Gulf of California, we assessed the fine-scale genetic and spatial structure of the population using individual-based approaches. Skin samples of 187 individuals were analyzed for nine microsatellite loci. A single population with no divergence among years and months and no isolation by distance (Rxy = 0.1–0.001, p>0.05) were found. We ran two Bayesian clustering methods using Structure and Geneland softwares in two different ways: 1) a general analysis including all individuals in which a single cluster was identified with both softwares; 2) a specific analysis of females only in which two main clusters (Loreto Bay and northern areas, and San Jose-La Paz Bay area) were revealed by Geneland program. This study provides information indicating that blue whales wintering in the Gulf of California are part of a single population unit and showed a fine-scale structure among females, possibly associated with their high site fidelity, particularly when attending calves. It is likely that the loss of genetic variation is minimized by male mediated gene flow, which may reduce the genetic drift effect. Opportunities for kin selection may also influence calf survival and, in consequence, have a positive impact on population demography in this small and endangered population.

The social organization of sperm whales in the Gulf of California and comparisons with other populations

Intra-specific variation in social organization provides valuable insights into the selective forces driving social evolution. Sperm whales are distributed globally and live far from shore, thus obtaining large sample sizes on social organization in multiple areas is logistically challenging and few comparative studies exist. In order to address how ecological factors influence sociality, we investigated the sacial organization of sperm whales in the Gulf of California (GoC) using a long-term study (1998―2004) and compare our results to other published studies. Standard photo-identification and behavioural observation techniques were used. Group size was calculated from photographic mark―recaptures using a Petersen estimator. Social organization was investigated using SocProg 2.3. Mean typical group sizes in the GoC were similar to those in the Galapagos Islands, Chile and Seychelles (24.7, 24.8, 30.4 and 18 individuals respectively), but substantially larger than in the Sargasso Sea, Caribbean and northern Gulf of Mexico (12.0, 6.4 and 6.9 individuals respectively). Sperm whale social organization in the GoC best fitted a constant companion/casual acquaintance model, where permanent units sizes were 12.5 individuals and two units usually associated together to form a group. This structure is similar to the situation in the Galapagos Islands and Chile areas. However, groups were more stable in the GoC than in the South Pacific, as groups stayed together for periods of about 80 days versus about ten days in the Galapagos Islands and Chile. It is likely that differences in the social organization between the study areas in the Pacific and Atlantic Oceans were due to differences in predation pressure and/or food resources. We suggest that, site-specific ecological factors are likely to infiuence fundamental aspects of sperm whale social organization.

Inferring cetacean population densities from the absolute dynamic topography of the ocean in a hierarchical Bayesian framework

We inferred the population densities of blue whales (Balaenoptera musculus) and short-beaked common dolphins (Delphinus delphis) in the Northeast Pacific Ocean as functions of the water-column’s physical structure by implementing hierarchical models in a Bayesian framework. This approach allowed us to propagate the uncertainty of the field observations into the inference of species-habitat relationships and to generate spatially explicit population density predictions with reduced effects of sampling heterogeneity. Our hypothesis was that the large-scale spatial distributions of these two cetacean species respond primarily to ecological processes resulting from shoaling and outcropping of the pycnocline in regions of wind-forced upwelling and eddy-like circulation. Physically, these processes affect the thermodynamic balance of the water column, decreasing its volume and thus the height of the absolute dynamic topography (ADT). Biologically, they lead to elevated primary productivity and persistent aggregation of low-trophic-level prey. Unlike other remotely sensed variables, ADT provides information about the structure of the entire water column and it is also routinely measured at high spatial-temporal resolution by satellite altimeters with uniform global coverage. Our models provide spatially explicit population density predictions for both species, even in areas where the pycnocline shoals but does not outcrop (e.g. the Costa Rica Dome and the North Equatorial Countercurrent thermocline ridge). Interannual variations in distribution during El Niño anomalies suggest that the population density of both species decreases dramatically in the Equatorial Cold Tongue and the Costa Rica Dome, and that their distributions retract to particular areas that remain productive, such as the more oceanic waters in the central California Current System, the northern Gulf of California, the North Equatorial Countercurrent thermocline ridge, and the more southern portion of the Humboldt Current System. We posit that such reductions in available foraging habitats during climatic disturbances could incur high energetic costs on these populations, ultimately affecting individual fitness and survival.

The simultaneous detection of mitochondrial DNA damage from sun-exposed skin of three whale species and its association with UV-induced microscopic lesions and apoptosis

Due to life history and physiological constraints, cetaceans (whales) are unable to avoid prolonged exposure to external environmental insults, such as solar ultraviolet radiation (UV). The majority of studies on the effects of UV on skin are restricted to humans and laboratory animals, but it is important to develop tools to understand the effects of UV damage on large mammals such as whales, as these animals are long-lived and widely distributed, and can reflect the effects of UV across a large geographical range. We and others have used mitochondrial DNA (mtDNA) as a reliable marker of UV-induced damage particularly in human skin. UV-induced mtDNA strand breaks or lesions accumulate throughout the lifespan of an individual, thus constituting an excellent biomarker for cumulative exposure. Based on our previous studies in human skin, we have developed for the first time in the literature a quantitative real-time PCR methodology to detect and quantify mtDNA lesions in skin from sun-blistered whales. Furthermore the methodology allows for simultaneous detection of mtDNA damage in different species. Therefore using 44 epidermal mtDNA samples collected from 15 blue whales, 10 fin whales, and 19 sperm whales from the Gulf of California, Mexico, we quantified damage across 4.3 kilobases, a large region of the ~16,400 base pair whale mitochondrial genome. The results show a range of mtDNA damage in the skin of the three different whale species. This previously unreported observation was correlated with apoptotic damage and microscopic lesions, both of which are markers of UV-induced damage. As is the case in human studies, this suggests the potential use of mtDNA as a biomarker for measuring the effect of cumulative UV exposure in whales and may provide a platform to help understand the effects of changing global environmental conditions.

Estimating blue whale skin isotopic incorporation rates and baleen growth rates: Implications for assessing diet and movement patterns in mysticetes

Stable isotope analysis in mysticete skin and baleen plates has been repeatedly used to assess diet and movement patterns. Accurate interpretation of isotope data depends on understanding isotopic incorporation rates for metabolically active tissues and growth rates for metabolically inert tissues. The aim of this research was to estimate isotopic incorporation rates in blue whale skin and baleen growth rates by using natural gradients in baseline isotope values between oceanic regions. Nitrogen (δ15N) and carbon (δ13C) isotope values of blue whale skin and potential prey were analyzed from three foraging zones (Gulf of California, California Current System, and Costa Rica Dome) in the northeast Pacific from 1996–2015. We also measured δ15N and δ13C values along the lengths of baleen plates collected from six blue whales stranded in the 1980s and 2000s. Skin was separated into three strata: basale, externum, and sloughed skin. A mean (±SD) skin isotopic incorporation rate of 163±91 days was estimated by fitting a generalized additive model of the seasonal trend in δ15N values of skin strata collected in the Gulf of California and the California Current System. A mean (±SD) baleen growth rate of 15.5±2.2 cm y-1 was estimated by using seasonal oscillations in δ15N values from three whales. These oscillations also showed that individual whales have a high fidelity to distinct foraging zones in the northeast Pacific across years. The absence of oscillations in δ15N values of baleen sub-samples from three male whales suggests these individuals remained within a specific zone for several years prior to death. δ13C values of both whale tissues (skin and baleen) and potential prey were not distinct among foraging zones. Our results highlight the importance of considering tissue isotopic incorporation and growth rates when studying migratory mysticetes and provide new insights into the individual movement strategies of blue whales.

Genetic Variation at Exon 2 of the MHC Class II DQB Locus in Blue Whale (Balaenoptera musculus) from the Gulf of California.

The genes of the Major Histocompatibility Complex (MHC) play an important role in the vertebrate immune response and are among the most polymorphic genes known in vertebrates. In some marine mammals, MHC genes have been shown to be characterized by low levels of polymorphism compared to terrestrial taxa; this reduction in variation is often explained as a result of lower pathogen pressures in marine habitats. To determine if this same reduction in variation applies to the migratory population of blue whales (Balaenoptera musculus) that occurs in the Gulf of California, we genotyped a 172 bp fragment of exon 2 of the MHC Class II DQB locus for 80 members of this population. Twenty-two putatively functional DQB allotypes were identified, all of which were homologous with DQB sequences from other cetacean species. Up to 5 putative alleles per individual were identified, suggesting that gene duplication has occurred at this locus. Rates of non-synonymous to synonymous substitutions (ω) and maximum likelihood analyses of models of nucleotide variation provided potential evidence of ongoing positive selection at this exon. Phylogenetic analyses of DQB alleles from B. musculus and 16 other species of cetaceans revealed trans-specific conservation of MHC variants, suggesting that selection has acted on this locus over prolonged periods of time. Collectively our findings reveal that immunogenic variation in blue whales is comparable to that in terrestrial mammals, thereby providing no evidence that marine taxa are subject to reduced pathogen-induced selective pressures.