Tracing early stages of species differentiation: Ecological, morphological and genetic divergence of Galapagos sea lion populations

Background: Oceans are high gene flow environments that are traditionally believed to hamper the build-up of genetic divergence. Despite this, divergence appears to occur occasionally at surprisingly small scales. The Galápagos archipelago provides an ideal opportunity to examine the evolutionary processes of local divergence in an isolated marine environment. Galápagos sea lions (Zalophus wollebaeki) are top predators in this unique setting and have an essentially unlimited dispersal capacity across the entire species range. In theory, this should oppose any genetic differentiation.
Results: We find significant ecological, morphological and genetic divergence between the western colonies and colonies from the central region of the archipelago that are exposed to different ecological conditions. Stable isotope analyses indicate that western animals use different food sources than those from the central area. This is likely due to niche partitioning with the second Galápagos eared seal species, the Galápagos fur seal (Arctocephalus galapagoensis) that exclusively dwells in the west. Stable isotope patterns correlate with significant differences in foraging-related skull morphology. Analyses of mitochondrial sequences as well as microsatellites reveal signs of initial genetic differentiation.
Conclusion: Our results suggest a key role of intra- as well as inter-specific niche segregation in the evolution of genetic structure among populations of a highly mobile species under conditions of free movement. Given the monophyletic arrival of the sea lions on the archipelago, our study challenges the view that geographical barriers are strictly needed for the build-up of genetic divergence. The study further raises the interesting prospect that in social, colonially breeding mammals additional forces, such as social structure or feeding traditions, might bear on the genetic partitioning of populations.

Pinniped Populations at Islas ae Guadalupe, San Benito, Cedros, and Natividad, Baja California, in 1968

In April and June 1968, the Pacific Ocean Biological Survey Program (POBSP) of the Smithsonian Institution conducted surveys on breeding marine birds and pinnipeds on various Mexican islands. Between 18 to 26 April and 21 to 29 June, pinniped populations were surveyed at Islas de Guadalupe, San Benito, Cedros, and Natividad off Baja California. Species observed were the California sea lion, Zalophus californianus, Guadalupe fur seal, Arctocephalus townsendi, harbor seal, Phoca vitulina, and northern elephant seal, Mirounga angustirostris

Recovery plan for the Steller sea lion (Eumetopias jubatus) /

Mode of access: Internet.

Deciduous dentition of a Californian sea lion pup (Zalophus californianus)

Veterinarians are increasingly faced with examining and/or treating marine mammals during strandings or following requests from theme parks and zoos. A common request involving seals and sea lions regards dentition, either to age a wild animal or as part of the routine health of captive animals. The major problem with examining dentition is the lack of information available in the literature on normal dentition, particularly in juveniles, where eruption patterns may be used to age the animal. The information presented in this report details typical deciduous dentition in one Californian sea lion, a species commonly encountered in captivity.

Characterization of TRPC2, an Essential Genetic Component of VNS Chemoreception, Provides Insights into the Evolution of Pheromonal Olfaction in Secondary-Adapted Marine Mammals

Pheromones are chemical cues released and sensed by individuals of the same species, which are of major importance in regulating reproductive and social behaviors of mammals. Generally, they are detected by the vomeronasal system (VNS). Here, we first investigated and compared an essential genetic component of vomeronasal chemoreception, that is, TRPC2 gene, of four marine mammals varying the degree of aquatic specialization and related terrestrial species in order to provide insights into the evolution of pheromonal olfaction in the mammalian transition from land to water. Our results based on sequence characterizations and evolutionary analyses, for the first time, show the evidence for the ancestral impairment of vomeronasal pheromone signal transduction pathway in fully aquatic cetaceans, supporting a reduced or absent dependence on olfaction as a result of the complete adaptation to the marine habitat, whereas the amphibious California sea lion was found to have a putatively functional TRPC2 gene, which is still under strong selective pressures, reflecting the reliance of terrestrial environment on chemical recognition among the semiadapted marine mammals. Interestingly, our study found that, unlike that of the California sea lion, TRPC2 genes of the harbor seal and the river otter, both of which are also semiaquatic, are pseudogenes. Our data suggest that other unknown selective pressures or sensory modalities might have promoted the independent absence of a functional VNS in these two species. In this respect, the evolution of pheromonal olfaction in marine mammals appears to be more complex and confusing than has been previously thought. Our study makes a useful contribution to the current understanding of the evolution of pheromone perception of mammals in response to selective pressures from an aquatic environment.

The brain of the fur seal, Callorhinus ursinus : with a comparative description of those of Zalophus californianus, Phoca vitulina, Ursus americanus and Monachus tropicalis /

Caption title.

Food habits of Steller sea lions (Eumetopias jubatus) off Oregon and northern California, 1986–2007

We described the diet of the eastern stock of Steller sea lions (Eumetopias jubatus) from 1416 scat samples collected from five sites in Oregon and northern California from 1986 through 2007. A total of 47 prey types from 30 families were identified. The most common prey was Pacific hake (Merluccius productus), followed by salmonids (Oncorhynchus spp.), skates (Rajidae), Pacific lamprey (Lampetra tridentata), herrings (Clupeidae), rockfish (Sebastes spp.), and northern anchovy (Engraulis mordax). Steller sea lion diet composition varied seasonally, annually, and spatially. Hake and salmonids were the most commonly identified prey in scats collected during the summer (breeding season), whereas hake and skate were most common in the nonbreeding season. Continued research on Steller sea lion diet and foraging behavior in the southern extent of their range is necessary to address issues such as climate change, interaction with competing California sea lions, and predation impacts on valuable or sensitive fish stocks.

Chemical contaminants, pathogen exposure and general health status of live and beach-cast Washington sea otters (Enhydra lutris kenyoni)

Analyses of blood and liver samples from live captured sea otters and liver samples from beachcast sea otter carcasses off the remote Washington coast indicate relatively low exposure to contaminants, but suggest that even at the low levels measured, exposure may be indicated by biomarker response. Evidence of pathogen exposure is noteworthy - infectious disease presents a potential risk to Washington sea otters, particularly due to their small population size and limited distribution. During 2001 and 2002, 32 sea otters were captured, of which 28 were implanted with transmitters to track their movements and liver and blood samples were collected to evaluate contaminant and pathogen exposure. In addition, liver samples from fifteen beachcast animals that washed ashore between 1991 and 2002 were analyzed to provide historical information and a basis of reference for values obtained from live otters. The results indicate low levels of metals, butyltins, and organochlorine compounds in the blood samples, with many of the organochlorines not detected except polychlorinated biphenyls (PCBs), and a few aromatic hydrocarbons detected in the liver of the live captured animals. Aliphatic hydrocarbons were measurable in the liver from the live captured animals; however, some of these are likely from biogenic sources. A significant reduction of vitamin A storage in the liver was observed in relation to PCB, dibutyltin and octacosane concentration. A significant and strong positive correlation in vitamin A storage in the liver was observed for cadmium and several of the aliphatic hydrocarbons. Peripheral blood mononuclear cell (PBMC) cytochrome P450 induction was elevated in two of 16 animals and may be potentially related to aliphatic and aromatic hydrocarbon exposure. Mean concentration of total butyltin in the liver of the Washington beach-cast otters was more than 15 times lower than the mean concentration reported by Kannan et al. (1998) for Southern sea otters in California. Organochlorine compounds were evident in the liver of beach-cast animals, despite the lack of large human population centers and development along the Washington coast. Concentrations of PCBs and chlordanes (e.g., transchlordane, cis-chlordane, trans-nonachlor, cis-nonachlor and oxychlordane) in liver of Washington beach-cast sea otters were similar to those measured in Aleutian and California sea otters, excluding those from Monterey Bay, which were higher. Mean concentrations of 1,1,1,- trichloro-2,2-bis(p-chlorophyenyl)ethanes (DDTs) were lower, and mean concentrations of cyclohexanes (HCH, e.g., alpha BHC, beta BHC, delta BHC and gamma BHC) were slightly higher in Washington beach-cast otters versus those from California and the Aleutians. Epidemiologically, blood tests revealed that 80 percent of the otters tested positive for morbillivirus and 60 percent for Toxoplasma, the latter of which has been a significant cause of mortality in Southern sea otters in California. This is the first finding of positive morbillivirus titers in sea otters from the Northeast Pacific. Individual deaths may occur from these diseases, perhaps more so when animals are otherwise immuno-compromised or infected with multiple diseases, but a population-threatening die-off from these diseases singly is unlikely while population immunity remains high. The high frequency of detection of morbillivirus and Toxoplasma in the live otters corresponds well with the cause of death of stranded Washington sea otters reported herein, which has generally been attributable to infectious disease. Washington’s sea otter population continues to grow, with over 1100 animals currently inhabiting Washington waters; however, the rate of growth has slowed over recent years. The population has a limited distribution and has not yet reached its carrying capacity and as such, is still considered at high risk to catastrophic events. (PDF contains 189 pages)

California Cooperative Fisheries Investigations. Hydrographic data report, Monterey Bay, January to December 1977

The data contained in this report were obtained as a continuance of the nearly bi-weekly hydrographic observations initiated by personnel at Hopkins Marine Station over two decades ago. These observations have been supported through the years by the State of California Marine Research Committee, California Cooperative Oceanic Fisheries Investigations. Since July 1974, the hydrographic sampling program has been carried out by the investigators at Moss Landing Marine Laboratories. From July 1974 to June 1976, this work was done in conjunction with an interdisciplinary study of the squid, Loligo opalescens, supported by the National Office of Sea Grant 'via the University of California Sea Grant College Project Number R/F-15. Five of the original CalCOFI stations (2201, 2202, 2203, 2204 and 2205) have been-retained in our sampling routine and additional inner-bay stations have been added (1154 and 1121) Sampling was conducted on a monthly basis for the entire year. All observations were made ab9ard R/V OCONOSTOTA. (PDF contains 93 pages)

Effects of commercial fishing on local abundance of Pacific cod (Gadus macrocephalus) in the Bering Sea

Groundfish fisheries in the southeast Bering Sea in Alaska have been constrained in recent years by management measures to protect the endangered Steller sea lion (Eumetopias jubatus). There is concern that the present commercial harvest may produce a localized depletion of groundfish that would affect the foraging success of Steller sea lions or other predators. A three-year field experiment was conducted to determine whether an intensive trawl fishery in the southeast Bering Sea created a localized depletion in the abundance of Pacific cod (Gadus macrocephalus). This experiment produced strongly negative results; no difference was found in the rate of seasonal change in Pacific cod abundance between stations within a regulatory no-trawl zone and stations in an immediately adjacent trawled area. Corollary studies showed that Pacific cod in the study area were highly mobile and indicated that the geographic scale of Pacific cod movement was larger than the spatial scale used as the basis for current no-trawl zones. The idea of localized depletion is strongly dependent on assumed spatial and temporal scales and contains an implicit assumption that there is a closed local population. The scale of movement of target organisms is critical in determining regional effects of fishery removals.

Diets of Steller sea lions (Eumetopias jubatus) in Southeast Alaska, 1993−1999

The diet of Steller sea lions (Eumetopias jubatus) was determined from 1494 scats (feces) collected at breeding (rookeries) and nonbreeding (haulout) sites in Southeast Alaska from 1993 to 1999. The most common prey of 61 species identified were walleye pollock (Theragra chalcogramma), Pacific herring (Clupea pallasii), Pacific sand lance (Ammodytes hexapterus), Pacific salmon (Salmonidae), arrowtooth flounder (Atheresthes stomias), rockfish (Sebastes spp.), skates (Rajidae), and cephalopods (squid and octopus). Steller sea lion diets at the three Southeast Alaska rookeries differed significantly from one another. The sea lions consumed the most diverse range of prey categories during summer, and the least diverse during fall. Diet was more diverse in Southeast Alaska during the 1990s than in any other region of Alaska (Gulf of Alaska and Aleutian Islands). Dietary differences between increasing and declining populations of Steller sea lions in Alaska correlate with rates of population change, and add credence to the view that diet may have played a role in the decline of sea lions in the Gulf of Alaska and Aleutian Islands.

Survey- and fishery-derived estimates of Pacific cod (Gadus macrocephalus) biomass: implications for strategies to reduce interactions between groundfish fisheries and Steller sea lions (Eumetopias jubatus)

Survey- and fishery-derived biomass estimates have indicated that the harvest indices for Pacific cod (Gadus macrocephalus) within a portion of Steller sea lion (Eumetopias jubatus) critical habitat in February and March 2001 were five to 16 times greater than the annual rate for the entire Bering Sea-Aleutian Islands stock. A bottom trawl survey yielded a cod biomass estimate of 49,032 metric tons (t) for the entire area surveyed, of which less than half (23,329 t) was located within the area used primarily by the commercial fishery, which caught 11,631 t of Pacific cod. Leslie depletion analyses of fishery data yielded biomass estimates of approximately 14,500 t (95% confidence intervals of approximately 9,000–25,000 t), which are within the 95% confidence interval on the fished area survey estimate (12,846–33,812 t). These data indicate that Leslie analyses may be useful in estimating local fish biomass and harvest indices for certain marine fisheries that are well constrained spatially and relatively short in duration (weeks). In addition, fishery effects on prey availability within the time and space scales relevant to foraging sea lions may be much greater than the effects indicated by annual harvest rates estimated from stock assessments averaged across the range of the target spec

Distribution and Abundance of Steller Sea Lions, Eumetopias jubatus, on the Asian Coast, 1720's-2005

We analyzed published and archived records for the past 250 years to assess changes in distribution and abundance of Steller sea lions, Eumetopias jubatus, along the Asian coast from the Bering Strait to the Korean Peninsula. We found that the northern extent of Steller sea lion distribution has not changed but that the southern limit has moved north by some 500–900 km (~300–500 n.mi.) over the past 50 years. Additionally, the number of animals and their distribution has changed on the Commander Islands, Kuril Islands, and Kamchatka Peninsula. We found no changes in the number of rookeries in the northern Sea of Okhotsk, but a new rookery was established at Tuleny Island on the eastern coast of Sakhalin Island. We estimate that the total abundance of Steller sea lions along the Asian coast in the late 19th century was about 115,000 animals; during the 1960’s, the total estimate was about 27,000 (including pups), most of which were in the Kuril Islands. The fewest number of Steller sea lions occurred in the northwestern Pacific in the late 1980’s–early 1990’s when only about 13,000 individuals (including pups) were estimated in the entire region. During the 1990’s, and especially in early 2000, an increasing trend in abundance occurred in most areas. Present estimated abundance of Steller sea lions in Asia is about 16,000 individuals (including about 5,000 pups), about half of which occur in the Kuril Islands. Changes in abundance occurred during all time periods but varied by site and period. Specifically, over the past 150 years Steller sea lion abundance at most sites has changed. There were no rookeries on the Commander Islands between 1850 and 1960 and abundance was low, but by 1977, abundance increased to 4,800 individuals and a rookery was established in the mid 1980’s; abundance there has declined since the early 1980’s and in 2004 only 895 individuals (including 221 pups) were counted during the breeding season. Between 1940 and 2004, abundance along the eastern coast of Kamchatka declined from ~7,000 to ~600 individuals, an overall reduction of 90%. Steller sea lion abundance on the Kuril Islands declined by >90% from the 1800’s to 2005; the most severe decline there occurred during 1969–1981. Steller sea lion numbers in the northern part of the Sea of Okhotsk declined during 1930–2002 from 7,200 to 3,100 individuals. Numbers at Tuleny Island have increased since establishment of a rookery there during 1983–2005 and by immigration from other sites.

The Threatened Status of Steller Sea Lions, Eumetopias jubatus, under the Endangered Species Act: Effects on Alaska Groundfish Fisheries Management

In April 1990, the Steller sea lion, Eumetopias jubatus, was listed as threatened under the U.S. Endangered Species Act by emergency action. Competitive interactions with the billion-dollar Alaska commercial groundfish fisheries have been suggested as one of the possible contributing factors to the Steller sea lion population decline. Since the listing, fisheries managers have attempted to address the potential impacts of the groundfish fisheries on Steller sea lion recovery. In this paper, we review pertinent Federal legislation, biological information on the Steller sea lion decline, changes in the Alaska trawl fishery for walleye pollock, Theragra chalcogramma, since the late 1970's, andpossible interactions between fisheries and sea lions. Using three cases, we illustrate how the listing of Steller sea lions has affected Alaska groundfish fisheries through: I) actions taken at the time of listing designed to limit the potential for directhuman-related sea lion mortality, 2) actions addressing spatial and temporal separation of fisheries from sea lions, and 3) introduction of risk-adverse stock assessment methodologies and Steller sea lion conservation considerations directly in the annual quota-setting process. This discussion shows some of the ways that North Pacific groundfish resource managers have begun to explicitly consider the conservation ofmarine mammal and other nontarget species.