Habitat and behaviour of yellowfin tuna Thunnus albacares in the Gulf of Mexico determined using pop-up satellite archival tags

This study presents the first data on movement, habitat use and behaviour for yellowfin tuna Thunnus albacares in the Atlantic Basin. Six individuals were tracked in the Gulf of Mexico using pop-up satellite archival tags. Records up to 80 days in length were obtained, providing information on depth and temperature preferences as well as horizontal movements. Thunnus albacares in the Gulf of Mexico showed a strong preference for the mixed layer and thermocline, consistent with findings for this species in other ocean basins. Fish showed a diel pattern in depth distribution, remaining in surface and mixed layer waters at night and diving to deeper waters during the day. The vertical extent of T. albacares habitat appeared to be temperature limited, with fish generally avoiding waters that were >6° C cooler than surface waters. The vertical and thermal habitat usage of T. albacares differs from that of bigeye Thunnus obesus and bluefin Thunnus thynnus , Thunnus orientalis and Thunnus maccoyii tunas. These results are consistent with the results of earlier studies conducted on T. albacares in other oceans.     

Insights into Young of the Year White Shark, Carcharodon carcharias, Behavior in the Southern California Bight

A young of the year female white shark, Carcharodon carcharias, was tagged with a pop-up satellite archival tag off Southern California in early June of 2000. The tag was recovered after 28 days, and records of temperature, depth and light intensity were extracted. Depth and temperature records indicate a number of interesting behaviors, including a strong diurnal pattern. At night the shark remained in the top 50 m, often making shallow repetitive vertical excursions. Most dives below the mixed layer were observed during the day, 91% of which occurred from 05:00 to 21:00 h, with depths extending to 240 m. Many of the dives exhibited secondary vertical movements that were consistent with the shark swimming at the bottom (at depths from 9 to 165 m) where it was most likely foraging. The white shark experienced dramatic and rapid changes in temperature, and demonstrated a considerable tolerance for cold waters. Temperatures ranged from 9°C to 22°C, and although 89% of the total time was spent in waters 16–22°C, on some days the small shark spent as much as 32% of the time in 12°C waters. The deep dives into cold waters separate the white sharks from mako sharks, which share the California Bight nursery ground but appear to remain primarily in the mixed layer and thermocline. Movement information (derived from light-based geolocation, bottom depths and sea surface temperatures) indicated that the white shark spent the 28 days in the Southern California Bight, possibly moving as far south as San Diego, California. While the abundance and diversity of prey, warm water and separation from adults make this region an ideal nursery ground, the potential for interaction with the local fisheries should be examined.     

Satellite Telemetry in Avian Research and Management: Sample Size Considerations

Abstract: Satellite tracking is currently used to make inferences to avian populations. Cost of transmitters and logistical challenges of working with some species can limit sample size and strength of inferences. Therefore, careful study design including consideration of sample size is important. We used simulations to examine how sample size, population size, and population variance affected probability of making reliable inferences from a sample and the precision of estimates of population parameters. For populations of >100 individuals, a sample >20 birds was needed to make reliable inferences about questions with simple outcomes (i.e., 2 possible outcomes). Sample size demands increased rapidly for more complex problems. For example, in a problem with 3 outcomes, a sample of >75 individuals will be needed for proper inference to the population. Combining data from satellite telemetry studies with data from surveys or other types of sampling may improve inference strength.     

Telemetry and Satellite Tracking of Whale Sharks, Rhincodon Typus, in the Sea of Cortez, Mexico, and the North Pacific Ocean

We used satellite-linked radio telemetry to document the geographic and vertical movements and thermal habitats of whale sharks in the Sea of Cortez and as they migrated into the north Pacific Ocean. Of 17 sharks tagged between 1994 and 1996, six dispersed widely in the Sea of Cortez during 12–39 days of tracking. Four others left the Sea of Cortez and ranged extensively in the north Pacific Ocean. Indeed, one whale shark migrated to the western north Pacific Ocean, covering over 13000km in 37 months of tracking. The sharks generally occupied areas where sea surface water temperatures were between 28 and 32°C, though several ranged to depths of 240m or deeper where water temperature reached 10°C or colder. Whale sharks may segregate by size and sex, and their movement patterns appear to be related to oceanographic features, such as sea mounts and boundary currents, where primary productivity may be enhanced. These results have important implications for the global conservation of the world's largest yet least known fish. We think that satellite telemetry is a exceptionally promising tool for learning more about the ecology of whale sharks, especially when combined with conventional methods of telemetry and molecular biology.authorship arranged alphabetically     

Intraspecific killing among Leopards (Panthera pardus) in Iran (Mammalia: Felidae)

Intraspecific aggression is one of the most common causes of death in leopards. Here, we report four cases of intraspecific killing amongst Persian Leopards (Panthera pardus saxicolor) in Iran. A young male leopard was found on 7 June 2008 which, according to camera trap images, had been killed by an adult male over a Red Deer (Cervus elaphus) kill, with trauma to his neck in Dorfak No-Hunting Area. A young female that had been fitted with a satellite GPS collar on 6 December 2015 in Tandoureh National Park died on 29 January 2016 at a site where an Urial Sheep (Ovis orientalis) ram had been freshly killed. Necropsy results, footprints at the scene of death and camera trap footage all supported the deduction that the animal was killed by a larger female leopard at the kill site. On 13 January 2017, a young, partially eaten female leopard was found with double puncture on the side of her throat. Finally, a rehabilitated adult female fitted with a satellite GPS collar found on 19 December 2017 with a double puncture on her head with several trauma and haemorrhages on her back. These instances seem to be the first documented reports of intraspecific killing among free-ranging leopards in Asia.     

Migratory connectivity of American woodcock derived using satellite telemetry

American woodcock (Scolopax minor; woodcock) migratory connectivity (i.e., association between breeding and wintering areas) is largely unknown, even though current woodcock management is predicated on such associations. Woodcock are currently managed in the Eastern and Central management regions in the United States with the boundary between management regions analogous to the boundary between the Atlantic and Mississippi flyways, based largely on analysis of band returns from hunters. Factors during migration influence survival and fitness, and existing data derived from banding and very high frequency telemetry provide only coarse-scale information to assess factors influencing woodcock migratory movement patterns and behavior. To assess whether current management-region boundaries correspond with woodcock migratory connectivity in the Central Management Region and to describe migration patterns with higher resolution than has been previously possible, we deployed satellite transmitters on 73 woodcock (25 adult and 28 juvenile females, and 8 adult and 12 juvenile males) and recorded 87 autumn or spring migration paths from 2014 to 2016. Marked woodcock used 2 primary migrations routes: a Western Route and a Central Route. The Western Route ran north-south, connecting the breeding and wintering grounds within the Central Management Region. The hourglass-shaped Central Route connected an area on the wintering grounds reaching from Texas to Florida, to sites throughout northeastern North America in both the Eastern Management Region and Central Management Region and woodcock following this route migrated through the area between the Appalachian Mountains and the Mississippi Alluvial Valley in western Tennessee during autumn and spring. Two of 17 woodcock captured associated with breeding areas in Michigan, Wisconsin, or Minnesota migrated to wintering sites in the Eastern Management Region and 12 marked woodcock captured on wintering areas in Texas and Louisiana migrated to breeding sites in the Eastern Management Region. Woodcock that used the Western Route exhibited high concentrations of stopovers during spring in the Arkansas Ozark Mountains and northern Missouri, and along the Mississippi River on the border between Wisconsin and Minnesota, and autumn concentrations of stopovers in southwestern Iowa, central Missouri, the Arkansas portion of the Ozark Mountains, and around the junction of Texas, Louisiana, Oklahoma, and Arkansas. Woodcock that used the Central Route exhibited high concentrations of stopovers during spring in northern Mississippi through western Tennessee, western Kentucky, and the Missouri Bootheel, and autumn concentrations of stopovers in northern Illinois, southwestern Ohio, and the portions of Kentucky and Tennessee west of the Appalachian Mountains. We suggest that current management of woodcock based on 2 management regions may not be consistent with the apparent lack of strong migratory connectivity we observed. Our results also suggest where management of migration habitat might be most beneficial to woodcock. © 2019 The Wildlife Society.     

Intra‐African movements of the African cuckoo Cuculus gularis as revealed by satellite telemetry

Despite many bird species migrating regularly within the African continent, in response to rainfall and breeding opportunities, documented evidence of the spatiotemporal patterns of such movements is scarce. We use satellite telemetry to document the year round movement of an intra‐African migrant breeding in the savannah zone of sub‐Saharan Africa, the African cuckoo. After breeding in central Nigeria, the birds migrated to more forested sites in the Adamawa region of Cameroon (n = 2) and western Central African Republic (n = 1). Departure from the breeding ground coincided with deteriorating environmental conditions whereas arrival at the non‐breeding sites matched period of increasing vegetation greenness. Migratory movements generally occurred during dark hours. In total, an average distance of 748 km in 66 d was covered during the post‐breeding migration and 744 km in 27 d during return journey with considerable individual variation and with more stopover sites used during post‐breeding migration. The diversity of migration routes followed suggests a relatively variable or flexible initial migration strategy, high individual route consistency as well as high fidelity for non‐breeding grounds.     

GPS tracking of the foraging movements of Manx Shearwaters Puffinus puffinus breeding on Skomer Island, Wales

We report the first successful use of miniature Global Positioning System loggers to track the ocean‐going behaviour of a c. 400 g seabird, the Manx Shearwater Puffinus puffinus. Breeding birds were tracked over three field seasons during the incubation and chick‐rearing periods on their foraging excursions from the large colony on Skomer Island, Pembrokeshire, UK. Foraging effort was concentrated in the Irish Sea. Likely foraging areas were identified to the north, and more diffusely to the west of the colony. No foraging excursions were recorded significantly to the south of the colony, conflicting with the conclusions of earlier studies based on ringing recoveries and observations. We discuss several explanations including the hypothesis that foraging may have shifted substantially northwards in recent decades. We found no obvious relationship between birds’ positions and water depth, although there was a suggestion that observations at night were in shallower water than those during the day. We also found that, despite the fact that Shearwaters can be observed rafting off‐shore from their colonies in the hours prior to making landfall at night, breeding birds are usually located much further from the colony in the last 8 h before arrival, a finding that has significance for the likely effectiveness of marine protection areas if they are only local to the colony. Short sequences of precise second‐by‐second fixes showed that movement speeds were bimodal, corresponding to sitting on the water (most common at night and around midday) and flying (most common in the morning and evening), with flight behaviour separable into erratic (indicative of searching for food) and directional (indicative of travelling). We also provide a first direct measurement of mean flight speed during directional flight (c. 40 km/h), slower than a Shearwater's predicted maximum range velocity, suggesting that birds are exploiting wave or dynamic soaring during long‐distance travel.     

Navigational challenges in the oceanic migrations of leatherback sea turtles

The open-sea movements of marine animals are affected by the drifting action of currents that, if not compensated for, can produce non-negligible deviations from the correct route towards a given target. Marine turtles are paradigmatic skilful oceanic navigators that are able to reach remote goals at the end of long-distance migrations, apparently overcoming current drift effects. Particularly relevant is the case of leatherback turtles (Dermochelys coriacea), which spend entire years in the ocean, wandering in search of planktonic prey. Recent analyses have revealed how the movements of satellite-tracked leatherbacks in the Indian, Atlantic and Pacific Oceans are strongly dependent on the oceanic currents, up to the point that turtles are often passively transported over long distances. However, leatherbacks are known to return to specific areas to breed every 2–3 years, thus finding their way back home after long periods in the oceanic environment. Here we examine the navigational consequences of the leatherbacks'' close association with currents and discuss how the combined reliance on mechanisms of map-based navigation and local orientation cues close to the target may allow leatherbacks to accomplish the difficult task of returning to specific sites after years spent wandering in a moving medium.     

Accuracy of Lightweight Satellite Telemetry: A Case Study in the Iberian Peninsula

Abstract: Here we provide the first assessment of the accuracy of lightweight satellite transmitters (<80 g) under actual operating conditions and the performance of the Argos system in southern Europe. To estimate transmitter accuracy we used transmitters equipped with a Global Positioning System (GPS) and compared the location estimates provided by Argos with the estimates provided by the GPS. Using the 68th percentile to define the accuracy of locations estimates, observed accuracy was 4 km for Location Class (LC) 1, 15 km for LC 0, 20 km for LC A, and 59 km for LC B, which is in line with estimates reported by other authors. Yet, the error of the remaining 32% of the data ranged between 4 km and 11 km, 15 km and 217 km, 20 km and 145 km, and 59 km and 493 km, respectively, suggesting that using the 68th percentile to estimate accuracies might give misleading confidence on the accuracy of location estimates. Using the 90th percentile is probably more appropriate. Less than 10% of the locations we obtained corresponded to the more accurate LCs (3, 2, and 1), with Argos failing to provide a position estimate in 45% of the attempts. The low number of high-quality location estimates is likely a consequence of the electromagnetic interference reported for our study area, rather than a defect of the Platform Transmitter Terminals (PTTs), which under good conditions of signal reception seem to be as reliable as heavier ones. The recent advent of lightweight GPS transmitters overrides most of these problems. Yet, whereas the smallest Argos-GPS PTTs weigh 30 g, which restricts their use to animals weighting > 1,000 g, conventional PTTs can be as small as 9.5 g, allowing their use with animals weighting 250-300 g.     

Movement patterns of the White-tailed Sea Eagle (Haliaeetus albicilla): post-fledging behaviour,natal dispersal onset and the role of the natal environment

Exploratory movements and natal dispersal form essential processes during early life history stages of raptors, but identifying the factors shaping individual movement decisions is challenging. Global positioning system (GPS) telemetry thereby provides a promising technique to study movement patterns on adequate spatio-temporal scales. We analysed data of juvenile White-tailed Sea Eagles Haliaeetus albicilla (WTSE) in north-east Germany (n = 24) derived from GPS tracking to extensively analyse movements between fledging and emigration from the natal territory. Our goal was to determine the time point of fledging, characterize pre-emigration movements and the onset of natal dispersal while investigating the influence of the natal environment. WTSE fledged at an average age of 72 days and showed strong excursive behaviour during the post-fledging period regarding the number, distance and duration of excursions, yet with high individual variability. Excursive behaviour did not differ between sexes. On average, WTSE left the parental territory 93 days after fledging. The quantity of excursive behaviour delayed the timing of emigration and WTSE tended to postpone their emigration when foraging water was accessible within the boundaries of their parental territory. The overall results suggest that young WTSE assess the quality of the natal environment via pre-emigration movements and stay in their territory of origin for as long as internal and external conditions allow for it. Our study is one of the first to characterize post-fledging and natal dispersal movements of young WTSE to such an extent and applies modern techniques to understand related movements in relation to the natal environment. The results emphasize the urgent necessity for the extension of currently existing nest protection periods and guaranteeing sustainable management of potential breeding and foraging grounds for WTSE. Ultimately, the results are relevant for all large raptor species sensitive to human-related disturbance, as they support the increasing importance of regulations with spatio-temporal specifications for breeding populations of large raptors in densely human-inhabited areas with increasing alteration of land.     

What drives long‐distance movements in the nomadic Grey Teal Anas gracilis in Australia?

In contrast to northern temperate environments, where day length and temperature changes are obvious proximate cues for movement to resource-rich breeding habitats, the cues for movement used by birds in an often resource-poor, stochastic environment are less obvious. We recorded long-distance movements of 23 Grey Teal Anas gracilis using satellite telemetry for up to 879 days and examined the relationship between those movements and environmental factors, such as heavy rainfall and flooding, at the destination site. We identified 32 long-distance (> 150 km) movements that met our criterion for minimally interrupted flight between origin and destination. Thirteen of these flights coincided with rainfall and/or flooding events up to 1050 km from the origin. However, some ducks moved without any clear beneficial conditions at the destination onto small wetlands in regions with little surface water. The data suggest that there are two types of long-distance movement – ranging and directed. These flights occurred over distances up to 1200 km across the arid inland. The rates and distances of movement suggest that long-distance movements of Grey Teal entail high energy costs as in waterfowl elsewhere. We conclude that the proximate controls of directed movements need not be very different from those of their temperate counterparts.     

Refuges and risks: Evaluating the benefits of an expanded MPA network for mobile apex predators

Aim

Concurrently, assessing the effectiveness of marine protected areas and evaluating the degree of risk from humans to key species provide valuable information that can be integrated into conservation management planning. Tiger sharks (Galeocerdo cuvier) are a wide‐ranging ecologically important species subject to various threats. The aim of this study was to identify “hotspots” of tiger shark habitat use in relation to protected areas and potential risks from fishing.

Location

Southwest Indian Ocean, east coast of South Africa and Mozambique.

Methods

Satellite tags were fitted to 26 tiger sharks. A subset of 19 sharks with an average period at liberty of 197 (SD = 110) days were analysed using hotspot analysis to identify areas of core habitat use. The spatial and temporal overlap of significant hotspots with current and planned marine protected areas as well as risks from fishing and culling was then calculated.

Results

There was a 5.97% spatial overlap between tiger shark hotspots and marine protected areas, which would increase significantly (p < .05) to 24.36% with the expansion of planned protected areas in South Africa and could be as high as 41.43% if Mozambique similarly expanded neighbouring protected area boundaries. Tiger sharks remained largely coastal, but only showed a spatial overlap of 5.12% with shark culling nets in South Africa. Only three sharks undertook open ocean migrations during which they were more likely to interact with longline fisheries in the region.

Main conclusions

This study demonstrates how spatial information can be used to assess the overlap between marine protected areas and the core habitats of top marine predators and highlights how congruent transnational conservation management can improve the effectiveness of protected areas. Core habitat use of marine apex predators may also be indicative of productive habitats, and therefore, predators such as tiger sharks could act as surrogate species for identifying key habitats to prioritize for conservation planning.

     

Effects of implanted satellite transmitters on behavior and survival of female common eiders

Surgically implanted satellite transmitters have been widely used in studies of avian ecology, yet little is known about their potential impacts on birds. We implanted satellite transmitters with percutaneous antennae (approx. 50 g) in 17 female common eiders (Somateria mollissima) at a breeding colony in Arctic Canada. Among females implanted during incubation, 11 of 12 nests were abandoned within 1 week of being radioed. We observed no differences in the proportion of time that implanted female eiders allocated to basic behaviors. Radioed birds were more likely to pick or preen their abdominal (site of surgical incision) and posterior–dorsal (site of antenna exit) regions than unmarked females, although these behaviors were rare (approx. 0.3% of total time budget). Three of 10 females re-observed had a pronounced limp following surgery, but we observed no walking difficulties among these females in subsequent seasons, and we observed some implanted eiders nesting in subsequent years. Mark–resighting models suggest eiders with transmitters had lower apparent survival the year after implantation (67.0%; 85% CI: 47.8–81.9%) than did color-banded eiders (87.5%, 85% CI: 82.5–91.2%), but there was no model support for a survival difference in subsequent years. We conclude that transmitter implantation in common eiders leads to short-term changes in behavior and a decline in first year survival. We encourage researchers to collect similar data on their study subjects where possible and use it to determine the degree to which data are representative of the greater population. © 2011 The Wildlife Society.     

Behavioral response of a mobile marine predator to environmental variables differs across ecoregions

Animal movement and habitat selection are in part a response to landscape heterogeneity. Many studies of movement and habitat selection necessarily use environmental covariates that are readily available over large‐scales, which are assumed representative of functional habitat features such as resource availability. For widely distributed species, response to such covariates may not be consistent across ecosystems, as response to any specific covariate is driven by its biological relevance within the context of each ecosystem. Thus, the study of any widely distributed species within a limited geographic region may provide inferences that are not widely generalizable. Our goal was to evaluate the response of a marine predator to a suite of environmental covariates across a wide ecological gradient. We identified two behavioral states (resident and transient) in the movements of shortfin mako sharks Isurus oxyrinchus tracked via satellite telemetry in two regions of the western North Atlantic Ocean: the tropical Caribbean/Gulf of Mexico marginal sea (CGM), and the temperate waters of the open western Atlantic Ocean (OWA). We compared patterns of resident behavior between regions, and modeled relationships between oceanographic variables and resident behavior. We tracked 39 sharks during 2013–2015. Resident behavior was associated with shallow, continental shelf and slope waters in both regions. In the OWA resident behavior was associated with low sea surface temperature and high primary productivity, however, sharks exhibited no response to either variable in the CGM. There was a negative relationship between sea‐surface height gradient (a proxy for oceanic fronts) and resident behavior in the OWA, and a positive relationship in the CGM. Our observations likely reflect shark responses to regional variability in factors responsible for the distribution and availability of prey. Our study illustrates the importance of studying widely distributed species in a consistent manner over large spatial scales.     

DIVE CHARACTERISTICS OF SATELLITE-MONITORED BLUE WHALES (BALAENOPTERA MUSCULUS) OFF THE CENTRAL CALIFORNIA COAST

Dive habits of four Northeast Pacific blue whales ( Balaenoptera musculus ) were studied using satellite-monitored radio tags. Tags summarized dive-duration data into eight 3-h periods daily. One tag additionally summarized dive depth and time-at-depth information for these same periods. Tracking periods ranged from 0.6 to 12.7 d and provided data for 17 three-hour summary periods, representing 2,007 dives (788 of which provided depth information). Total number of dives during a 3-h summary period ranged from 83 to 128. Seventy-two percent of dives were ≤ 1 min long. All whales spent >94% of their time submerged. Average duration of true dives (dives >1 min) ranged from 4.2 to 7.2 min. Seventy-five percent of depth-monitored dives were to ≤16 m, accounting for 78% of that animal's time. Average depth of dives to >16 m was 105 ± 13 m.     

Migration cues and timing in leatherback sea turtles

Atlantic leatherback sea turtles migrate annually from foraginggrounds off eastern Canada and the northeastern United Statesto southern foraging and breeding areas. Using Cox's proportionalhazards model, we investigated the individual timing of thesouthward migrations of 27 turtles equipped with satellite-linkedtransmitters off Nova Scotia compared with turtle characteristicsand satellite-measured ocean variables. Latitude, longitude,1-week lagged average sea surface temperature, and 1-week laggedaverage chlorophyll-a concentration appear to influence theprobability of departure. Higher temperature and, in the northernrange of the study, higher chlorophyll concentration increaseddeparture rates, perhaps due to the acceleration of the lifecycle of the leatherback's gelatinous prey and/or increasedfeeding efficiency in these areas. This study highlights theopportunity to use satellite telemetry and environmental datato examine the cues for and timing of animal migrations andexpands the study of migration timing to include a new speciesand environment.     

MOVEMENTS OF NORTH PACIFIC BLUE WHALES DURING THE FEEDING SEASON OFF SOUTHERN CALIFORNIA AND THEIR SOUTHERN FALL MIGRATION1

The satellite-acquired locations of 10 blue whales (Balaenoptera musculus) tagged off southern California with Argos radio tags were used to identify (1) their movements during the late summer feeding season; (2) the routes and rate of travel for individuals on their southern fall migration; and (3) a possible winter calving/breeding area. Whales were tracked from 5.1 to 78.1 d and from 393 to 8,668 km. While in the Southern California Bight, most of the locations for individual whales were either clumped or zigzagged in pattern, suggesting feeding or foraging (searching for prey). Average speeds ranged from 2.4 to 7.2 km/h. One whale moved north to Cape Mendocino, and four migrated south along the Baja California, Mexico coast, two passing south of Cabo San Lucas on the same day. One of the latter whales traveled an additional 2,959 km south in 30.5 d to within 450 km of the Costa Rican Dome (CRD), an upwelling feature. The timing of this migration suggests the CRD may be a calving/breeding area for North Pacific blue whales. Although blue whales have previously been sighted in the Eastern Tropical Pacific (ETP), this is the first evidence that whales from the feeding aggregation off California range that far south. The productivity of the CRD may allow blue whales to feed during their winter calving/breeding season, unlike gray whales (Eschrichtius robustus) and humpbacks (Megaptera novaeangliae) which fast during that period.     

Foraging behavior of adult female Steller sea lions during the breeding season in Southeast Alaska

During the 1990s, the Steller sea lion ( Eumetopias jubatus Schreber) Western Alaska stock (WS) suffered steep population decline while the Eastern Alaska stock (ES) steadily increased. One bottom-up forcing hypothesis explaining this decline predicted lactating adult female foraging behavior would be different between stocks. To investigate this effect, we monitored 11 ES females at two breeding rookeries using satellite dive recorders (SDR) during the early breeding seasons of 1992–1993, examined their behavior with respect to prey, physiological limitations, and habitat, and made limited comparisons to observations of WS female behavior reported in the literature. ES females were not operating at the extremes of ability, with most diving within the limits of aerobic metabolism, less than one-quarter of possible foraging time during trips spent submerged and most foraging trips requiring less than one-half the lipid store fasting ability of dependent pups. Thus, females may have some capacity to alter behavior to accommodate future changes in foraging conditions, but the extent of this plasticity is unknown. Because recent work suggests WS recovery is impeded by low natality, future studies should test differences between reproductive and non-reproductive mature females in order to properly assess the contribution of foraging ecology to SSL population dynamics.