Luis A. Hückstädt

Foraging Behavior and Success of a Mesopelagic Predator in the Northeast Pacific Ocean: Insights from a Data-Rich Species, the Northern Elephant Seal

The mesopelagic zone of the northeast Pacific Ocean is an important foraging habitat for many predators, yet few studies have addressed the factors driving basin-scale predator distributions or inter-annual variability in foraging and breeding success. Understanding these processes is critical to reveal how conditions at sea cascade to population-level effects. To begin addressing these challenging questions, we collected diving, tracking, foraging success, and natality data for 297 adult female northern elephant seal migrations from 2004 to 2010. During the longer post-molting migration, individual energy gain rates were significant predictors of pregnancy. At sea, seals focused their foraging effort along a narrow band corresponding to the boundary between the sub-arctic and sub-tropical gyres. In contrast to shallow-diving predators, elephant seals target the gyre-gyre boundary throughout the year rather than follow the southward winter migration of surface features, such as the Transition Zone Chlorophyll Front. We also assessed the impact of added transit costs by studying seals at a colony near the southern extent of the species’ range, 1,150 km to the south. A much larger proportion of seals foraged locally, implying plasticity in foraging strategies and possibly prey type. While these findings are derived from a single species, the results may provide insight to the foraging patterns of many other meso-pelagic predators in the northeast Pacific Ocean.

Estimates of the Southern Ocean general circulation improved by animal‐borne instruments

Over the last decade, several hundred seals have been equipped with conductivity-temperature-depth sensors in the Southern Ocean for both biological and physical oceanographic studies. A calibrated collection of seal-derived hydrographic data is now available, consisting of more than 165,000 profiles. The value of these hydrographic data within the existing Southern Ocean observing system is demonstrated herein by conducting two state estimation experiments, differing only in the use or not of seal data to constrain the system. Including seal-derived data substantially modifies the estimated surface mixed-layer properties and circulation patterns within and south of the Antarctic Circumpolar Current. Agreement with independent satellite observations of sea ice concentration is improved, especially along the East Antarctic shelf. Instrumented animals efficiently reduce a critical observational gap, and their contribution to monitoring polar climate variability will continue to grow as data accuracy and spatial coverage increase.

Oceanic controls on the mass balance of Wilkins Ice Shelf, Antarctica

� 0.8 m a � 1 , driven by a mean basal melt rate of 〈wb〉 = 1.3 � 0.4 m a � 1 . Interannual variability was large, associated with changes in both surface mass accumulation and 〈wb〉. Basal melt rate declined significantly around 2000 from 1.8 � 0.4 m a � 1 for 1992–2000 to � 0.75 � 0.55 m a � 1 for 2001–2008; the latter value corresponding to approximately steady-state ice-shelf mass. Observations of ocean temperature T obtained during 2007–2009 by instrumented seals reveal a cold, deep halo of Winter Water (WW; T ≈ � 1.6°C) surrounding WIS. The base of the WW in the halo is � 170 m, approximately the mean ice draft for WIS. We hypothesize that the transition in 〈wb〉 in 2000 was caused by a small perturbation (� 10–20 m) in the relative depths of the ice base and the bottom of the WW layer in the halo. We conclude that basal melting of thin ice shelves like WIS is very sensitive to upper-ocean and coastal processes that act on shorter time and space scales than those affecting basal melting of thicker West Antarctic ice shelves such as George VI and Pine Island Glacier.

Seals map bathymetry of the Antarctic continental shelf

We demonstrate the first use of marine mammal dive-depth data to improve maps of bathymetry in poorly sampled regions of the continental shelf. A group of 57 instrumented elephant seals made on the order of 2 x 10(5) dives over and near the continental shelf on the western side of the Antarctic Peninsula during five seasons, 2005-2009. Maximum dive depth exceeded 2000 m. For dives made near existing ship tracks with measured water depths H<700 m, similar to 30% of dive depths were to the seabed, consistent with expected benthic foraging behavior. By identifying the deepest of multiple dives within small areas as a dive to the seabed, we have developed a map of seal-derived bathymetry. Our map fills in several regions for which trackline data are sparse, significantly improving delineation of troughs crossing the continental shelf of the southern Bellingshausen Sea.

Latitudinal range influences the seasonal variation in the foraging behavior of marine top predators.

Non-migratory resident species should be capable of modifying their foraging behavior to accommodate changes in prey abundance and availability associated with a changing environment. Populations that are better adapted to change will have higher foraging success and greater potential for survival in the face of climate change. We studied two species of resident central place foragers from temperate and equatorial regions with differing population trends and prey availability associated to season, the California sea lion (Zalophus californianus) (CSL) whose population is increasing and the endangered Galapagos sea lion (Zalophus wollebaeki) (GSL) whose population is declining. To determine their response to environmental change, we studied and compared their diving behavior using time-depth recorders and satellite location tags and their diet by measuring C and N isotope ratios during a warm and a cold season. Based on latitudinal differences in oceanographic productivity, we hypothesized that the seasonal variation in foraging behavior would differ for these two species. CSL exhibited greater seasonal variability in their foraging behavior as seen in changes to their diving behavior, foraging areas and diet between seasons. Conversely, GSL did not change their diving behavior between seasons, presenting three foraging strategies (shallow, deep and bottom divers) during both. GSL exhibited greater dive and foraging effort than CSL. We suggest that during the warm and less productive season a greater range of foraging behaviors in CSL was associated with greater competition for prey, which relaxed during the cold season when resource availability was greater. GSL foraging specialization suggests that resources are limited throughout the year due to lower primary production and lower seasonal variation in productivity compared to CSL. These latitudinal differences influence their foraging success, pup survival and population growth reflected in contrasting population trends in which CSL are more successful and potentially more resilient to climate change.

Investigating foraging utilization distribution of female New Zealand sea lions, Auckland Islands

Detrimental interactions between marine mammals and fisheries are increasing worldwide. The ability to manage these interactions requires the knowledge of where and how interactions occur and the effects they have on species. Many pinnipeds are central place foraging colonial breeders who are restricted in foraging range during breeding. Here, we use a utilization distribution approach to examine the foraging habitats of lactating New Zealand (NZ) sea lions (Phocarctos hookeri) from Dundas and Enderby Islands, Auckland Islands. Annually, the NZ sea lions which breed on these two islands produce 83% of this Nationally Critical species’ pups. Satellite transmitters were attached to 55 females during 2001–2007. Data showed that NZ sea lions utilize the entire Auckland Island shelf with partial habitat partitioning between females from the two breeding islands. This habitat partitioning results in differing degrees of overlap with fisheries and therefore possible differing fishery-related impacts on breeding areas.

Seals and sea lions are what they eat, plus what? Determination of trophic discrimination factors for seven pinniped species

RATIONALE Mixing models are a common method for quantifying the contribution of prey sources to the diet of an individual using stable isotope analysis; however, these models rely upon a known trophic discrimination factor (hereafter, TDF) that results from fractionation between prey and animal tissues. Quantifying TDFs in captive animals is ideal, because diet is controlled and the proportional contributions and isotopic values of all prey items are known. METHODS To calculate TDFs for the Hawaiian monk seal, northern elephant seal, bearded seal, ringed seal, spotted seal, harbor seal, and California sea lion, we obtained whiskers, serum, plasma, red blood cells, and prey items from nine captive individuals. We obtained δ(13) C and δ(15) N values using continuous-flow isotope-ratio mass spectrometry. The average δ(13) C and δ(15) N values from bulk and lipid-corrected prey from the diet were subtracted from the δ(13) C and δ(15) N values of each blood and whisker sample to calculate tissue-specific TDFs for each individual (∆(13) C or ∆(15) N). RESULTS The ∆(13) C values ranged from +1.7 to +3.2‰ (bulk prey) and from +0.8 to +1.9‰ (lipid-corrected prey) for the various blood components, and from +3.9 to +4.6‰ (bulk prey) or +2.6 to +3.9‰ (lipid-corrected prey) for whiskers. The ∆(15) N values ranged from +2.2 to +4.3‰ for blood components and from +2.6 to +4.0‰ for whiskers. The TDFs tended to group by tissue, with whiskers having greater ∆(13) C values than blood components. In contrast, the ∆(15) N values were greater in serum and plasma than in red blood cells and whiskers. CONCLUSIONS By providing the first TDF values for five seal species (family Phocidae) and one otariid species (family Otariidae), our study facilitates more accurate mixing models for these species. These values are particularly important for critically endangered Hawaiian monk seals and the three Arctic seal species (bearded, ringed, and spotted) that are faced with a rapidly changing environment.

Regional variability in diving physiology and behavior in a widely distributed air-breathing marine predator, the South American sea lion (Otaria byronia).

ABSTRACT Our understanding of how air-breathing marine predators cope with environmental variability is limited by our inadequate knowledge of their ecological and physiological parameters. Because of their wide distribution along both coasts of the sub-continent, South American sea lions (Otaria byronia) provide a valuable opportunity to study the behavioral and physiological plasticity of a marine predator in different environments. We measured the oxygen stores and diving behavior of South American sea lions throughout most of its range, allowing us to demonstrate that diving ability and behavior vary across its range. We found no significant differences in mass-specific blood volumes of sea lions among field sites and a negative relationship between mass-specific oxygen storage and size, which suggests that exposure to different habitats and geographical locations better explains oxygen storage capacities and diving capability in South American sea lions than body size alone. The largest animals in our study (individuals from Uruguay) were the shallowest and shortest duration divers, and had the lowest mass-specific total body oxygen stores, while the deepest and longest duration divers (individuals from southern Chile) had significantly larger mass-specific oxygen stores, despite being much smaller animals. Our study suggests that the physiology of air-breathing diving predators is not fixed, but that it can be adjusted, to a certain extent, depending on the ecological setting and or habitat. These adjustments can be thought of as a ‘training effect’: as the animal continues to push its physiological capacity through greater hypoxic exposure, its breath-holding capacity increases. Summary: Regional variation in diving ability and behavior of the South American sea lion suggests that exposure to different habitats and geographical locations better explains oxygen storage capacities and diving capability than body size alone.

Using Satellite Tracking and Isotopic Information to Characterize the Impact of South American Sea Lions on Salmonid Aquaculture in Southern Chile.

Apex marine predators alter their foraging behavior in response to spatial and/or seasonal changes in natural prey distribution and abundance. However, few studies have identified the impacts of aquaculture that represents a spatially and temporally predictable and abundant resource on their foraging behavior. Using satellite telemetry and stable isotope analysis we examined the degree of spatial overlap between the South American sea lion (SASL) and salmon farms, and quantify the amount of native prey versus farmed salmonids in SASL diets. We instrumented eight SASL individuals with SRDL-GPS tags. Vibrissae, hair and skin samples were collected for δ13C and δ15N analyses from five of the tagged individuals and from four males captured in a haul-out located adjacent to salmon farms. Tracking results showed that almost all the foraging areas of SASL are within close proximity to salmon farms. The most important prey for the individuals analyzed was farmed salmonids, with an estimated median (±SD) contribution of 19.7 ± 13.5‰ and 15.3 ± 9.6‰ for hair and skin, respectively. Using vibrissae as a temporal record of diet for each individual, we observed a remarkable switch in diet composition in two SASL, from farmed salmonids to pelagic fishes, which coincided with the decrease of salmon production due to the infectious salmon anemia virus that affected salmon farms in Chile at the end of 2008. Our study demonstrates the usefulness of integrating stable isotope derived dietary data with movement patterns to characterize the impacts of a non-native prey on the foraging ecology of an apex marine predator, providing important applied implications in situations where interactions between aquaculture and wildlife are common.

A Dynamic State Model of Migratory Behavior and Physiology to Assess the Consequences of Environmental Variation and Anthropogenic Disturbance on Marine Vertebrates

Integrating behavior and physiology is critical to formulating new hypotheses on the evolution of animal life-history strategies. Migratory capital breeders acquire most of the energy they need to sustain migration, gestation, and lactation before parturition. Therefore, when predicting the impact of environmental variation on such species, a mechanistic understanding of the physiology of their migratory behavior is required. Using baleen whales as a model system, we developed a dynamic state variable model that captures the interplay among behavioral decisions, energy, reproductive needs, and the environment. We applied the framework to blue whales (Balaenoptera musculus) in the eastern North Pacific Ocean and explored the effects of environmental and anthropogenic perturbations on female reproductive success. We demonstrate the emergence of migration to track prey resources, enabling us to quantify the trade-offs among capital breeding, body condition, and metabolic expenses. We predict that periodic climatic oscillations affect reproductive success less than unprecedented environmental changes do. The effect of localized, acute anthropogenic impacts depended on whales’ behavioral response to the disturbance; chronic, but weaker, disturbances had little effect on reproductive success. Because we link behavior and vital rates by modeling individuals’ energetic budgets, we provide a general framework to investigate the ecology of migration and assess the population consequences of disturbance, while identifying critical knowledge gaps.