Ecotypic variation and predatory behavior among killer whales (Orcinus orca) off the eastern Aleutian Islands, Alaska

From 2001 to 2004 in the eastern Aleutian Islands, Alaska, killer whales (Orcinus orca) were encountered 250 times during 421 days of surveys that covered a total of 22,491 miles. Three killer whale groups (resident, transient, and offshore) were identified acoustically and genetically. Resident killer whales were found 12 times more frequently than transient killer whales, and offshore killer whales were encountered only once. A minimum of 901 photographically identified resident whales used the region during our study. A total of 165 mammal-eating transient killer whales were identified, and the majority (70%) were encountered during spring (May and June). The diet of transient killer whales in spring was primarily gray whales (Eschrichtius robustus), and in summer primarily northern fur seals (Callorhinus ursinus). Steller sea lions (Eumetopias jubatus) did not appear to be a preferred prey or major prey item during spring and summer. The majority of killer whales in the eastern Aleutian Islands are the resident ecotype, which does not consume marine mammals.

Trends and Potential Interactions Between Pinnipeds and Fisheries of New England and the U.S. West Coast

Long-term trends in the abundance and distribution of several pinniped species and commercially important fisheries of New England and the contiguous U.S. west coast are reviewed, and their actual and potential interactions discussed. Emphasis is on biological interactions or competition. The pinnipeds include the western North Atlantic stock of harbor seals, Phoca vitulina concolor; western North Atlantic gray seals, Halochoerus grypus; the U.S. stock of California sea lions, Zalophus californianus californianus; the eastern stock of Steller sea lions, Eumetopias jubatus; and Pacific harbor seals, Phoca vitulina richardii. Fisheries included are those for Atlantic cod, Gadus morhua; silver hake, Merluccius bilinearis; Atlantic herring, Clupea harengus; the coastal stock of Pacific whiting, Merluccius productus; market squid, Loligo opalescens; northern anchovy, Engraulis mordax; Pacific her-ring, Clupea pallasi; and Pacific sardine, Sardinops sagax. Most of these pinniped populations have grown exponentially since passage of the U.S. Marine Mammal Protection Act in 1972. They exploit a broad prey assemblage that includes several commercially valuable species. Direct competition with fisheries is therefore possible, as is competition for the prey of commercially valuable fish. The expanding pinniped populations, fluctuations in commercial fish biomass, and level of exploitation by the fisheries may affect this potential for competition. Concerns over pinnipeds impacting fisheries (especially those with localized spawning stocks or at low biomass levels) are more prevalent than concerns over fisheries’ impacts on pinnipeds. This review provides a framework to further evaluate potential biological interactions between these pinniped populations and the commercial fisheries with which they occur.

Domoic acid contamination within eight representative species from the benthic food web of Monterey Bay, California, USA

Benthic food webs often derive a significant fraction of their nutrient inputs from phytoplankton in the overlying waters. If the phytoplankton include harmful algal species like Pseudo-nitzschia australis, a diatom capable of producing the neurotoxin domoic acid (DA), the benthic food web can become a depository for phycotoxins. We tested the general hypothesis that DA contaminates benthic organisms during local blooms of P. australis, a widespread toxin producer along the US west coast. To test for trophic transfer and uptake of DA into the benthic food web, we sampled 8 benthic species comprising 4 feeding groups: filter feeders (Emerita analoga and Urechis caupo); a predator (Citharichthys sordidus); scavengers (Nassarius fossatus and Pagurus samuelis) and deposit feeders (Neotrypaea californiensis, Dendraster excentricus and Olivella biplicata). Sampling occurred before, during and after blooms of P. australis in Monterey Bay, CA, USA during 2000 and 2001. DA was detected in all 8 species, with contamination persisting over variable time scales. Maximum DA levels in N. fossatus (674 ppm), E. analoga (278 ppm), C. sordidus (515 ppm), N. californiensis (145 ppm), P. samuelis (56 ppm), D. excentricus (15 ppm) and O. biplicata (3 ppm) coincided with P. australis blooms, while DA levels in U. caupo remained above 200 ppm (max. = 751 ppm) throughout the study period. DA in 6 species exceeded levels thought to be safe for higher level consumers (i.e. ≥20 ppm) and thus is likely to have deleterious effects on marine birds, sea lions and the endangered California sea otter, known to prey upon these benthic species.

A method to improve size estimates of walleye pollock (Theragra chalcogramma) Atka mackerel (Pleurogrammus monopterygius) consumed by pinnipeds: digestion correction factors applied to bones and otoliths recovered in scats

The lengths of otoliths and other skeletal structures recovered from the scats of pinnipeds, such as Steller sea lions (Eumetopias jubatus), correlate with body size and can be used to estimate the length of prey consumed. Unfortunately, otoliths are often found in too few scats or are too digested to usefully estimate prey size. Alternative diagnostic bones are frequently recovered, but few bone-size to prey-size correlations exist and bones are also reduced in size by various degrees owing to digestion. To prevent underestimates in prey sizes consumed techniques are required to account for the degree of digestion of alternative bones prior to estimating prey size. We developed a method (using defined criteria and photo-reference material) to assign the degree of digestion for key cranial structures of two prey species: walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). The method grades each structure into one of three condition categories; good, fair or poor. We also conducted feeding trials with captive Steller sea lions, feeding both fish species to determine the extent of erosion of each structure and to derive condition-specific digestion correction factors to reconstruct the original sizes of the structures consumed. In general, larger structures were relatively more digested than smaller ones. Mean size reduction varied between different types of structures (3.3−26.3%), but was not influenced by the size of the prey consumed. Results from the observations and experiments were combined to be able to reconstruct the size of prey consumed by sea lions and other pinnipeds. The proposed method has four steps: 1) measure the recovered structures and grade the extent of digestion by using defined criteria and photo-reference collection; 2) exclude structures graded in poor condition; 3) multiply measurements of structures in good and fair condition by their appropriate digestion correction factors to derive their original size; and 4) calculate the size of prey from allometric regressions relating corrected structure measurements to body lengths. This technique can be readily applied to piscivore dietary studies that use hard remains of fish.

Changes over time in the spatial distribution of walleye pollock (Theragra chalcogramma) in the Gulf of Alaska, 1984-1996

Triennial bottom trawl survey data from 1984 to 1996 were used to evaluate changes in the summer distribution of walleye pollock in the western and central Gulf of Alaska. Differences between several age groups of pollock were evaluated. Distribution was examined in relation to several physical characteristics, including bottom depth and distance from land. Interspecies associations were also analyzed with the Bray-Curtis clustering technique to better understand community structure. Our results indicated that although the population numbers decreased, high concentrations of pollock remained in the same areas during 1984–96. However, there was an increase in the number of stations where low-density pollock concentrations of all ages were observed, which resulted in a decrease in mean population density of pollock within the GOA region. Patterns emerging from our data suggested an alternative to Mac-Call’s “basin hypothesis” which states that as population numbers decrease, there should be a contraction of the population range to optimal habitats. During 1984–96 there was a concurrent precipitous decline in Steller sea lions in the Gulf of Alaska. The results of our study suggest that decreases in the mean density of adult pollock, the main food in the Steller sea lion diet, combined with slight changes in the distribution of pollock (age-1 pollock in particular) in the mid-1980s, may have contributed to decreased foraging efficiency in Steller sea lions. Our results support the prevailing conceptual model for pollock ontogeny, although there is evidence that substantial spawning may also occur outside of Shelikof Strait.

Rensch's rule in large herbivorous mammals derived from metabolic scaling

Rensch’s rule, which states that the magnitude of sexual size dimorphism tends to increase with increasing body size, has evolved independently in three lineages of large herbivorous mammals: bovids (antelopes), cervids (deer), and macropodids (kangaroos). This pattern can be explained by a model that combines allometry,life-history theory, and energetics. The key features are thatfemale group size increases with increasing body size and that males have evolved under sexual selection to grow large enough to control these groups of females. The model predicts relationships among body size and female group size, male and female age at first breeding,death and growth rates, and energy allocation of males to produce body mass and weapons. Model predictions are well supported by data for these megaherbivores. The model suggests hypotheses for why some other sexually dimorphic taxa, such as primates and pinnipeds(seals and sea lions), do or do not conform to Rensh’s rule.

Variation in the fatty acid composition of the blubber in Cape fur seals (Arctocephalus pusillus pusillus) and the implications for dietary interpretation

Analysis of the fatty acid (FA) composition of blubber is a valuable tool in interpreting the diet of marine mammals. This technique is based on the principle that particular FA present in prey can be incorporated largely untransformed into predator adipose tissue stores, thereby providing biochemical signatures with which to identify prey species. Several studies of phocid seals and cetaceans have documented vertical stratification in the FA composition of blubber such that inferences about diet may vary greatly depending on the layer of the blubber that is analysed. It is not known whether blubber in otariid seals (fur seals and sea lions) also displays vertical stratification in FA composition. Furthermore, it is not known whether the FA composition of blubber is uniform in these species. In the present study, the vertical and regional variation in FA composition of blubber was investigated in seven adult female Cape fur seals (Arctocephalus pusillus pusillus). The proportion of monounsaturated fatty acids (MUFA) was greater in the outer (43.6±1.3%) than inner portion (40.9±1.2%; t20=5.59, P<0.001) whereas the proportions were greater in the inner than outer portions for saturated fatty acids (23.6±0.5% and 21.9±0.6%, respectively, t20 = 5.31, P<0.001) and polyunsaturated fatty acids (PUFA, 35.5±0.7% and 34.5±0.7%, respectively, t20 = 3.81, P < 0.001). There was an inverse relationship between MUFA and PUFA in the blubber, independent of sampling location. In addition, with the exception of the inner portion from non-lactating females, blubber from the mammary area had the highest proportions of 18:1ω9c and total MUFA, followed by blubber from the rump and neck, suggesting that the deposition and mobilisation of blubber lipids may not be uniform around the body in otariid seals. These results support the need for blubber tissue to be sampled from the same site on animals, and to the full depth of the blubber layer, to minimise variation in FA profiles that could occur if different sites and depths were sampled. Such standardisation of sampling will further aid in interpreting diet in otariid seals using the FA Signature Analysis approach.

Evolution of Pinnipedia lactation strategies: a potential role for α-lactalbumin?

Despite the considerable variation in milk composition found among mammals, a constituent common across all groups is lactose, the main sugar and osmole in most eutherians milk. Exceptions to this are the families Otariidae (fur seals and sea lions) and Odobenidae (walruses), where lactose has not been detected. We investigated the molecular basis for this by cloning α-lactalbumin, the modifier protein of the lactose synthase complex. A mutation was observed which, in addition to preventing lactose production, may enable otariids to maintain lactation despite the extremely long inter-suckling intervals during the mother's time at sea foraging (more than 23 days in some species).

Effect of proximity to the shelf edge on the diet of female Australian fur seals

The Australian fur seal Arctocephalus pusillus doriferus is a temperate latitude species with a breeding distribution restricted to Bass Strait, Australia. Recent studies of the foraging behaviour of female Australian fur seals indicated that they feed demersally in the shallow continental shelf waters, a behaviour that is in contrast to the epipelagic foraging of females of most other arctocephaline (Southern Hemisphere fur seals) species and akin to that observed in sea lions. These studies, however, were conducted at one colony (Kanowna Island) located in central northern Bass Strait, and it was suggested that the observed foraging behaviour may have been due to the distance of this colony from the continental shelf edge (180 km), making it inefficient to forage beyond it. Here, the diet of lactating Australian fur seals was compared between 2 colonies to test if differing proximity to the continental shelf edge resulted in differences in foraging behaviour. The 2 breeding colonies studied, Kanowna Island and The Skerries, were 180 and 25 km from the nearest shelf edge, respectively. We analysed a total of 917 scat samples collected at the 2 colonies between 1997 and 2001. From faecal analysis, 45 primarily demersal on-shelf species of fishes and cephalopods were identified. Only 4 species had a frequency of occurrence greater than 10%: redbait Emmelichthys nitidus, jack mackerel Trachurus sp., red rock cod Pseudophycis bachus, and Gould’s squid Nototodarus gouldi. No seasonal, annual or spatial differences were found between the 2 colonies, indicating that proximity to the shelf edge does not influence diet.

Individual variation in the pup attraction call produced by female Australian fur seals during early lactation

Otariid seals (fur seals and sea lions) are colonial breeders with large numbers of females giving birth on land during a synchronous breeding period. Once pups are born, females alternate between feeding their young ashore and foraging at sea. Upon return, both mother and pup must relocate each other and it is thought to be primarily facilitated by vocal recognition. Vocalizations of thirteen female Australian fur seals (Arctocephalus pusillus doriferus) were recorded during the breeding seasons of December 2000 and 2001, when pups are aged from newborns to one month. The pup attraction call was examined to determine whether females produce individually distinct calls which could be used by pups as a basis for vocal recognition. Potential for individual coding, discriminant function analysis (DFA), and classification and regression tree analysis were used to determine which call features were important in separating individuals. Using the results from all three analyses: F0, MIN F and DUR were considered important in separating individuals. In 76% of cases, the PAC was classified to the correct caller, using DFA, suggesting that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition by pups of this species.

When does physiology limit the foraging behaviour of freely diving mammals?

Diving animals offer a unique opportunity to study the importance of physiological constraint and the limitation it can impose on animal's behaviour in nature. This paper examines the interaction between physiology and behaviour and its impact on the diving capability of five eared seal species (Family Otariidae; three sea lions and two fur seals). An important physiological component of diving marine mammals is the aerobic dive limit (ADL). The ADL of these five seal species was estimated from measurements of their total body oxygen stores, coupled with estimates of their metabolic rate while diving. The tendency of each species to exceed its calculated ADL was compared relative to its diving behaviour. Overall, our analyses reveal that seals which forage benthically (i.e. on the sea floor) have a greater tendency to approach or exceed their ADL compared to seals that forage epipelagically (i.e. near the sea surface). Furthermore, the marked differences in foraging behaviour and physiology appear to be coupled with a species demography. For example, benthic foraging species have smaller populations and lower growth rates compared to seal species that forage epipelagically. These patterns are relevant to the conservation and management of diving vertebrates.

Taking the sting out of darting : Risks, restraint drugs and procedures for the chemical restraint of Southern Hemisphere otariids

The need to manage otariid populations has necessitated the development of a wide range of capture methods. Chemical restraint by remote drug delivery (i.e., darting) is a highly selective method that can be used to facilitate otariid capture in a range of scenarios, when other methods may be impracticable. However, the risks associated with darting otariids are not widely known and guidelines necessary to promote and refine best practice do not exist. We review the risks associated with darting and in light of our findings, develop darting guidelines to help practitioners assess and minimize risks during capture, anesthesia and recovery. Published studies reveal that mortalities associated with darting predominantly result from complications during anesthetic maintenance (e.g., prolonged respiratory depression, apnea, or hyperthermia), rather than from complications during capture or recovery. In addition to monitoring vital signs and proper intervention, the risk of irreversible complications during anesthesia can be reduced by administering drug doses that are sufficient to enable the capture and masking of animals, after which anesthetic depth can be regulated using gas anesthesia.

Disentangling the cause of a catastrophic population decline in a large marine mammal

Considerable uncertainties often surround the causes of long-term changes in population abundance. One striking example is the precipitous decline of southern sea lions (SSL; Otariaflavescens) at the Falkland Islands, from 80 555 pups in the mid 1930s to just 5506 pups in 1965. Despite an increase in SSL abundance over the past two decades, the population has not recovered, with the number of pups born in 2014 (minimum 4443 pups) less than 6% of the 1930s estimate. The order-of-magnitude decline is primarily attributed to commercial sealing in Argentina. Here, we test this established paradigm and alternative hypotheses by assessing (1) commercial sealing at the Falkland Islands, (2) winter migration of SSL from the Falkland Islands to Argentina, (3) whether the number of SSL in Argentina could have sustained the reported level of exploitation, and (4) environmental change. The most parsimonious hypothesis explaining the SSL population decline was environmental change. Specifically, analysis of 160 years of winter sea surface temperatures revealed marked changes, including a period of warming between 1930 and 1950 that was consistent with the period of SSL decline. Sea surface temperature changes likely influenced the distribution or availability of SSL prey and impacted its population dynamics. We suggest that historical harvesting may not always be the "smoking gun" as is often purported. Rather, our conclusions support the growing evidence for bottom-up forcing on the abundance of species at lower trophic levels (e.g., plankton and fish) and resulting impacts on higher trophic levels across a broad range of ecosystems.