Dive characteristics can predict foraging success in Australian fur seals (arctocephalus pusillus doriferus) as validated by animal-borne video

Dive characteristics and dive shape are often used to infer foraging success in pinnipeds. However, these inferences have not been directly validated in the field with video, and it remains unclear if this method can be applied to benthic foraging animals. This study assessed the ability of dive characteristics from time-depth recorders (TDR) to predict attempted prey capture events (APC) that were directly observed on animal-borne video in Australian fur seals (Arctocephalus pusillus doriferus, n=11). The most parsimonious model predicting the probability of a dive with ≥1 APC on video included only descent rate as a predictor variable. The majority (94%) of the 389 total APC were successful, and the majority of the dives (68%) contained at least one successful APC. The best model predicting these successful dives included descent rate as a predictor. Comparisons of the TDR model predictions to video yielded a maximum accuracy of 77.5% in classifying dives as either APC or non-APC or 77.1% in classifying dives as successful verses unsuccessful. Foraging intensity, measured as either total APC per dive or total successful APC per dive, was best predicted by bottom duration and ascent rate. The accuracy in predicting total APC per dive varied based on the number of APC per dive with maximum accuracy occurring at 1 APC for both total (54%) and only successful APC (52%). Results from this study linking verified foraging dives to dive characteristics potentially opens the door to decades of historical TDR datasets across several otariid species.

Flipper strokes can predict energy expenditure and locomotion costs in free-ranging northern and Antarctic fur seals

Flipper strokes have been proposed as proxies to estimate the energy expended by marine vertebrates while foraging at sea, but this has never been validated on free-ranging otariids (fur seals and sea lions). Our goal was to investigate how well flipper strokes correlate with energy expenditure in 33 foraging northern and Antarctic fur seals equipped with accelerometers, GPS, and time-depth recorders. We concomitantly measured field metabolic rates with the doubly-labelled water method and derived activity-specific energy expenditures using fine-scale time-activity budgets for each seal. Flipper strokes were detected while diving or surface transiting using dynamic acceleration. Despite some inter-species differences in flipper stroke dynamics or frequencies, both species of fur seals spent 3.79 ± 0.39 J/kg per stroke and had a cost of transport of ~1.6-1.9 J/kg/m while diving. Also, flipper stroke counts were good predictors of energy spent while diving (R(2) = 0.76) and to a lesser extent while transiting (R(2) = 0.63). However, flipper stroke count was a poor predictor overall of total energy spent during a full foraging trip (R(2) = 0.50). Amplitude of flipper strokes (i.e., acceleration amplitude × number of strokes) predicted total energy expenditure (R(2) = 0.63) better than flipper stroke counts, but was not as accurate as other acceleration-based proxies, i.e. Overall Dynamic Body Acceleration.

Estimates of Marine Mammal, Sea Turtle, and Seabird Mortality in the California Drift Gillnet Fishery for Swordfish and Thresher Shark, 1996–2002

Estimates of incidental marine mammal, sea turtle, and seabird mortality in the California drift gillnet fishery for broadbill swordfish, Xiphias gladius, and common thresher shark, Alopias vulpinus, are summarized for the 7-year period, 1996 to 2002. Fishery observer coverage was 19% over the period (3,369 days observed/17,649 days fished). An experiment to test the effectiveness of acoustic pingers on reducing marine mammal entanglements in this fishery began in 1996 and resulted in statistically significant reductions in marine mammal bycatch. The most commonly entangled marine mammal species were the short-beaked common dolphin, Delphinus delphis; California sea lion, Zalophus californianus; and northern right whale dolphin, Lissodelphis borealis. Estimated mortality by species (CV and observed mortality in parentheses) from 1996 to 2002 is 861 (0.11, 133) short-beaked common dolphins; 553 (0.16, 103) California sea lions; 151 (0.25, 31) northern right whale dolphins; 150 (0.21, 27) northern elephant seals, Mirounga angustirostris; 54 (0.41, 10) long-beaked common dolphins, Delphinus capensis; 44 (0.53, 6) Dall’s porpoise, Phocoenoides dalli; 19 (0.60, 5) Risso’s dolphins, Grampus griseus; 11 (0.71, 2) gray whales, Eschrichtius robustus; 7 (0.83, 2) sperm whales, Physeter macrocephalus; 7 (0.96, 1) short-finned pilot whales, Globicephala macrorhychus; 12 (1.06, 1) minke whales, Balaenoptera acutorostrata; 5 (1.05, 1) fin whales, Balaenoptera physalus; 11 (0.68, 2) unidentified pinnipeds; 33 (0.52, 4) leatherback turtles, Dermochelys coriacea; 18 (0.57, 3) loggerhead turtles, Caretta caretta; 13 (0.73, 3) northern fulmars, Fulmarus glacialis; and 6 (0.86, 2) unidentified birds.

Distribution and abundance of humpback whales (Megaptera novaeangliae) and other marine mammals off the northern Washington coast

We examined the summer distribution of marine mammals off the northern Washington coast based on six ship transect surveys conducted between 1995 and 2002, primarily from the NOAA ship McArthur. Additionally, small boat surveys were conducted in the same region between 1989 and 2002 to gather photographic identification data on humpback whales (Megaptera novaeangliae) and killer whales (Orcinus orca) to examine movements and population structure. In the six years of ship survey effort, 706 sightings of 15 marine mammal species were made. Humpback whales were the most common large cetacean species and were seen every year and a total of 232 sightings of 402 animals were recorded during ship surveys. Highest numbers were observed in 2002, when there were 79 sightings of 139 whales. Line-transect estimates for humpback whales indicated that about 100 humpback whales inhabited these waters each year between 1995 and 2000; in 2002, however, the estimate was 562 (CV= 0.21) whales. A total of 191 unique individuals were identified photographically and mark recapture estimates also indicated that the number of animals increased from under 100 to over 200 from 1995 to 2002. There was only limited interchange of humpback whales between this area and feeding areas off Oregon and California. Killer whales were also seen on every ship survey and represented all known ecotypes of the Pacific Northwest, including southern and northern residents, transients, and offshore-type killer whales. Dall’s porpoise (Phocoenoides dalli) were the most frequently sighted small cetacean; abundance was estimated at 181−291 individuals, except for 2002 when we observed dramatically higher numbers (876, CV= 0.30). Northern fur seals (Callorhinus ursinus) and elephant seals (Mirounga angustirostris) were the most common pinnipeds observed. There were clear habitat differences related to distance offshore and water depth for different species.

Variance in male reproductive success and sexual size dimorphism in pinnipeds: testing an assumption of sexual selection theory

The theory of evolution by sexual selection for sexual size dimorphism (SSD) postulates that SSD primarily reflects the adaptation of males and females to their different reproductive roles. For example, competition among males for access to females increases male body size because larger males are better able to maintain dominant status than smaller males. Larger dominant males sire most offspring while smaller subordinate males are unsuccessful, leading to skew in reproductive success. Therefore, species with male-biased SSD are predicted to have greater variance in male reproductive success than those in which both sexes are similar in size. We tested this prediction among the Pinnipedia, a mammalian group with a great variation in SSD. From a literature review, we identified genetic estimates of male reproductive success for 10 pinniped taxa (eight unique species and two subspecies of a ninth species) that range from seals with similarly sized males and females to species in which males are more than four times as large as females. We found no support for a positive relationship between variance in reproductive success and SSD among pinnipeds after excluding the elephant seals Mirounga leonina and Mirounga angustirostris, which we discuss as distinctive cases. Several explanations for these results are presented, including the revival of one of Darwin's original ideas. Darwin proposed that natural selection may explain SSD based on differences in energetic requirements between sexes and the potential for sexual niche segregation. Males may develop larger bodies to exploit resources that remain unavailable to females due to the energetic constraints imposed on female mammals by gestation and lactation. The importance of this alternative explanation remains to be tested.

Terrestrial apnoeas and the development of cardiac control in Australian fur seal (Arctocephalus pusillus doriferus) pups

The development of cardiac control in association with terrestrial respiration patterns was examined throughout the period of maternal dependence in Australian fur seal pups. Resting eupnoic heart rate and respiration rate were significantly correlated (r² = 0.49) and both decreased with age (P < 0.05 in both cases). From an early age (1 month), pups displayed terrestrial apnoeas (18.1 ± 0.5 s) accompanied by substantial bradycardia (127 beats min^-1, a 13% decrease from eupnoic HR). Terrestrial apnoea duration increased significantly with age reaching a mean of 41 s just prior to weaning, slightly lower than the mean dive duration (52 s) previously recorded for pups of the same age. Correspondingly, mean apnoic heart rate decreased with age to 74 beats min^-1 just prior to weaning, representing a 25% decrease on eupnoic heart rate. Importantly, concomitant with the decrease in mean apnoic heart rate with age, an increase in the control
of bradycardia was evident with the variability in instantaneous apnoic heart decreasing such that older pups were able to maintain a low steady heart rate for the duration of the apnoea. The changes seen in these parameters are similar to those reported during postnatal development in elephant seals (Mirounga spp.) and harbour seals (Phoca vitulina), and are considered indicative of the development of cardiac control. These findings suggest a common strategy for the development of bradycardia control in both otariid and phocid seals.

Accuracy of ARGOS locations of pinnipeds at-sea estimated using Fastloc GPS

Background: ARGOS satellite telemetry is one of the most widely used methods to track the movements of free-ranging marine and terrestrial animals and is fundamental to studies of foraging ecology, migratory behavior and habitat-use. ARGOS location estimates do not include complete error estimations, and for many marine organisms, the most commonly acquired locations (Location Class 0, A, B, or Z) are provided with no declared error estimate.
Methodology/Principal Findings: We compared the accuracy of ARGOS locations to those obtained using Fastloc GPS from the same electronic tags on five species of pinnipeds: 9 California sea lions (Zalophus californianus), 4 Galapagos sea lions (Zalophus wollebaeki), 6 Cape fur seals (Arctocephalus pusillus pusillus), 3 Australian fur seals (A. p. doriferus) and 5 northern elephant seals (Mirounga angustirostris). These species encompass a range of marine habitats (highly pelagic vs coastal), diving behaviors (mean dive durations 2–21 min) and range of latitudes (equator to temperate). A total of 7,318 ARGOS positions and 27,046 GPS positions were collected. Of these, 1,105 ARGOS positions were obtained within five minutes of a GPS position and were used for comparison. The 68^th percentile ARGOS location errors as measured in this study were LC-3
0.49 km, LC-2 1.01 km, LC-1 1.20 km, LC-0 4.18 km, LC-A 6.19 km, LC-B 10.28 km.
Conclusions/Significance: The ARGOS errors measured here are greater than those provided by ARGOS, but within the range of other studies. The error was non-normally distributed with each LC highly right-skewed. Locations of species that make short duration dives and spend extended periods on the surface (sea lions and fur seals) had less error than species like elephant seals that spend more time underwater and have shorter surface intervals. Supplemental data (S1) are provided allowing the creation of density distributions that can be used in a variety of filtering algorithms to improve the quality of ARGOS tracking data.

Global compilation of chlorophyll fluorescence profiles from several data bases, and from published and unpublished individual sources

The present data set includes 268,127 vertical in situ fluorescence profiles obtained from several available online databases and from published and unpublished individual sources. Metadata about each profiles are given in the file provided here in further details. The majority of profiles comes from the National Oceanographic Data Center (NODC) and the fluorescence profiles acquired by Bio-Argo floats available on the Oceanographic Autonomous Observations (OAO) platform (63.7% and 12.5% respectively). Different modes of acquisition were used to collect the data presented in this study: (1) CTD profiles are acquired using a fluorometer mounted on a CTD-rosette; (2) OSD (Ocean Station Data) profiles are derived from water samples and are defined as low resolution profiles; (3) the UOR (Undulating Oceanographic Recorder) profiles are acquired by a equipped with a fluorometer and towed by a research vessel; (4) PA profiles are acquired by autonomous platforms (here profiling floats or elephant seals equipped with a fluorometer). Data acquired from gliders are not included in the compilation.

Biotelemetry: a mechanistic approach to ecology

Remote measurement of the physiology, behaviour and energetic status of free-living animals is made possible by a variety of techniques that we refer to collectively as 'biotelemetry'. This set of tools ranges from transmitters that send their signals to receivers up to a few kilometers away to those that send data to orbiting satellites and, more frequently, to devices that log data. They enable researchers to document, for long uninterrupted periods, how undisturbed organisms interact with each other and their environment in real time. In spite of advances enabling the monitoring of many physiological and behavioural variables across a range of taxa of various sizes, these devices have yet to be embraced widely by the ecological community. Our review suggests that this technology has immense potential for research in basic and applied animal ecology. Efforts to incorporate biotelemetry into broader ecological research programs should yield novel information that has been challenging to collect historically from free-ranging animals in their natural environments. Examples of research that would benefit from biotelemetry include the assessment of animal responses to different anthropogenic perturbations and the development of life-time energy budgets.

Male reproductive tactics and female choice in the solitary, promiscuous bridled nailtail wallaby (Onychogalea fraenata)

Several behavioral studies of large, gregarious, and sexually dimorphic macropods have shown that males form dominance hierarchies and large males have the highest reproductive success. The bridled nailtail wallaby (Onychogalea fraenata) is a smaller and strongly sexually dimorphic macropod, but is also highly solitary and males do not form dominance hierarchies that are maintained temporally or spatially. Genetic studies of paternity have shown that large males are the most reproductively successful and only one-quarter of males sire offspring at any one time. The aim of this study was to investigate the tactics that males adopt to secure access to females at the time of estrus and to investigate whether females can influence which males have access to them. This study was conducted using 2 wild, free-ranging populations of bridled nailtail wallabies. Females in estrus were located and observed. and the total number of males present, the relative weight rank of each mate, and interactions between individuals were recorded. Females showed a preference for large males and incited male-male competition when the group of males present was large. Unlike other dimorphic macropods, fights among males were rare and were restricted to males of similar size. Large males gained access to females by guarding and following them closely and threatening other males who attempted to gain access. Smaller males spent less time with females, suggesting that small males may leave multimale groups in an attempt to locate unguarded females. Given the solitary nature of this species and the lack of a stable dominance hierarchy to influence male reproductive success. mate searching and mate guarding may be important male reproductive tactics in this species.

Early-life sexual segregation: ontogeny of isotopic niche differentiation in the Antarctic fur seal

Investigating the ontogeny of niche differentiation enables to determine at which life-stages sexual segregation arises, providing insights into the main factors driving resource partitioning. We investigated the ontogeny of foraging ecology in Antarctic fur seals (Arctocephalus gazella), a highly dimorphic species with contrasting breeding strategies between sexes. Sequential δ(13)C and δ(15)N values of whiskers provided a longitudinal proxy of the foraging niche throughout the whole life of seals, from weaning, when size dimorphism is minimal to the age of 5. Females exhibited an early-life ontogenetic shift, from a total segregation during their first year at-sea, to a similar isotopic niche as breeding females as early as age 2. In contrast, males showed a progressive change in isotopic niche throughout their development such that 5-year-old males did not share the same niche as territorial bulls. Interestingly, males and females segregated straight after weaning with males appearing to feed in more southerly habitats than females. This spatial segregation was of similar amplitude as observed in breeding adults and was maintained throughout development. Such early-life niche differentiation is an unusual pattern and indicates size dimorphism and breeding constraints do not directly drive sexual segregation contrary to what has been assumed in otariid seals.

First molecular detection and characterization of herpesvirus and poxvirus in a Pacific walrus (Odobenus rosmarus divergens)

BACKGROUND Herpesvirus and poxvirus can infect a wide range of species: herpesvirus genetic material has been detected and amplified in five species of the superfamily Pinnipedia; poxvirus genetic material, in eight species of Pinnipedia. To date, however, genetic material of these viruses has not been detected in walrus (Odobenus rosmarus), another marine mammal of the Pinnipedia clade, even though anti-herpesvirus antibodies have been detected in these animals. CASE PRESENTATION In February 2013, a 9-year-old healthy captive female Pacific walrus died unexpectedly at L'Oceanografic (Valencia, Spain). Herpesvirus was detected in pharyngeal tonsil tissue by PCR. Phylogenetic analysis revealed that the virus belongs to the subfamily Gammaherpesvirinae. Poxvirus was also detected by PCR in skin, pre-scapular and tracheobronchial lymph nodes and tonsils. Gross lesions were not detected in any tissue, but histopathological analyses of pharyngeal tonsils and lymph nodes revealed remarkable lymphoid depletion and lymphocytolysis. Similar histopathological lesions have been previously described in bovine calves infected with an alphaherpesvirus, and in northern elephant seals infected with a gammaherpesvirus that is closely related to the herpesvirus found in this case. Intracytoplasmic eosinophilic inclusion bodies, consistent with poxviral infection, were also observed in the epithelium of the tonsilar mucosa. CONCLUSION To our knowledge, this is the first molecular identification of herpesvirus and poxvirus in a walrus. Neither virus was likely to have contributed directly to the death of our animal.

Population Growth of Antarctic Fur Seals: Limitation by a Top Predator, The Leopard Seal?

Antarctic fur seals (Arctocephalus gazella) in the South Shetland Islands are recovering from 19th-century exploitation more slowly than the main population at South Georgia. To document demographic changes associated with the recovery in the South Shetlands, we monitored fur seal abundance and reproduction in the vicinity of Elephant Island during austral summers from 1986/1987 through 1994/1995. Total births, mean and variance of birth dates, and average daily mortality rates were estimated from daily live pup counts at North Cove (NC) and North Annex (NA) colonies on Seal Island. Sightings of leopard seals (Hydrurga leptonyx) and incidents of leopard seal predation on fur seal pups were recorded opportunistically during daily fur seal research at both sites. High mortality of fur seal pups, attributed to predation by leopard seals frequently observed at NC, caused pup numbers to decline rapidly between January and March (i.e., prior to weaning) each year and probably caused a long-term decline in the size of that colony. The NA colony, where leopard seals were never observed, increased in size during the study. Pup mortality from causes other than leopard seal predation appeared to be similar at the two sites. The number of pups counted at four locations in the Elephant Island vicinity increased slowly, at an annual rate of 3.8%, compared to rates as high as 11% at other locations in the South Shetland Islands. Several lines of circumstantial evidence are consistent with the hypothesis that leopard seal predators limit the growth of the fur seal population in the Elephant Island area and perhaps in the broader population in the South Shetland Islands. The sustained growth of this fur seal population over many decades rules out certain predator–prey models, allowing inference about the interaction between leopard seals and fur seals even though it is less thoroughly studied than predator–prey systems of terrestrial vertebrates of the northern hemisphere. Top-down forces should be included in hypotheses for future research on the factors shaping the recovery of the fur seal population in the South Shetland Islands.