Diel movements of the blacktip shark (<Emphasis Type="Italic">Carcharhinus limbatus</Emphasis>) in a Caribbean nursery

The blacktip shark (Carcharhinus limbatus) is a common coastal species in tropical and subtropical waters around the world. To examine the fine-scale movement ecology of this species in a Caribbean nursery, 17 neonate blacktip sharks were acoustically tagged in May, 2012 and tracked for one year in Coral Bay, St John, USVI. By quantifying linear movement and shifts in position from a fixed Inner harbor location, we identified a diel movement pattern where blacktip sharks spend daylight hours within core habitat of Inner Coral Harbor and move each night to the central and outer portions of the Bay, a linear shift of 174–934 m. When compared to standard home range calculations applied to the overall movement data, these nightly positions were outside of the 95% activity space and, therefore, undetected using traditional space utilization methods, despite their predictable daily occurrence. Cluster analysis and nonmetric multidimensional scaling indicated distinct movement periods and locations: daytime (sun up) within Inner Coral Harbor; nighttime (sun down) in the center of the Bay; and brief periods approximately six hours after sunset at the mouth of the Bay. This diel shift in habitat use is likely associated with nocturnal foraging because it coincides with similar shifts in potential prey species. Habitat and resource management that incorporates the blacktip shark movements described herein is critical to the protection of these vulnerable life stages. The findings of this paper advance the understanding of blacktip shark behavior and acoustic telemetry experimental design.     

Information‐theoretic model selection affects home‐range estimation and habitat preference inference: a case study of male Reeves’s Pheasants Syrmaticus reevesii

Reeves’s Pheasant Syrmaticus reevesii is a vulnerable forest bird inhabiting broadleaved habitats dominated by oaks Quercus spp. in central China. Identifying home‐ranges and habitat associations is important for understanding the biology of this species and developing effective management and conservation plans. We used information‐theoretic criteria to evaluate the relative performance of four parametric (exponential power, one‐mode bivariate normal, two‐mode bivariate normal and two‐mode bivariate circle) and two non‐parametric models (adaptive and fixed kernel) for estimating home‐ranges and habitat associations of Reeves’s Pheasants. For parametric models, Akaike’s information criterion (AIC_c) and the likelihood cross‐validation criterion (CVC) were relatively consistent in ranking the bivariate exponential power model the least acceptable, whereas the two‐mode bivariate models performed better. The CVC suggested that kernel models, particularly the adaptive kernel, performed best among all six models evaluated. The average core area and 95% contour area based on the model with greatest support were 6.1 and 54.9 ha, respectively, and were larger than those estimated from other models. The discrepancy in estimates between models with highest and the lowest support decreased as the contour size increased; however, home‐range shapes differed between models. Minimum convex polygons that removed 5% of extreme data points (MCP95) were roughly half the size of home‐ranges based on kernel models. Estimates of home‐range and model evaluation were not affected by sample size (> 50 observations for each bird). Inference about habitat preference based on composition analysis and home‐range overlap varied between models. That with strongest support suggested that Reeves’s Pheasants selected mature fir and mixed forest, avoided farmland, and had mean among‐individual home‐range overlaps of 20%. We recommend non‐parametric methods, particularly the adaptive kernel method, for estimating home‐ranges and core areas for species with complex multi‐polar habitat preferences in heterogeneous environments with large habitat patches. However, we caution against the traditional convenience of using a single model to estimate home‐ranges and recommend exploration of multiple models for describing and understanding the ecological processes underlying space use and habitat associations.     

Spatial separation without territoriality in shark communities

Spatial separation within predator communities can arise via territoriality but also from competitive interactions among and within species. However, linking competitive interactions to predator distribution patterns is difficult and theoretical models predict different habitat selection patterns dependent on habitat quality and how competition manifests itself. While models generally consider competitors to be either equal in ability, or for one phenotype to have a fixed advantage over the other, few studies consider that an animal may only have a competitive advantage in specific habitats. We used  10 years of telemetry data, habitat surveys and behavioral experiments, to show spatial partitioning between and within two species of reef shark (grey reef Carcharhinus amblyrhinchos and blacktip reef sharks C. melanopterus) at an unfished Pacific atoll. Within a species, sharks remained within small ‘sub‐habitats’ with very few movements of individuals between sub‐habitats, which previous models have suggested could be caused by intra‐specific competition. Blacktip reef sharks were more broadly distributed across habitat types but a greater proportion used lagoon and backreef habitats, while grey reef sharks preferred forereef habitats. Grey reef sharks at a nearby atoll where blacktip reef sharks are absent, were distributed more broadly between habitat types than when both species were present. A series of individual‐based models predict that habitat separation would only arise if there are competitive interactions between species that are habitat‐specific, with grey reefs having a competitive advantage on the forereefs and blacktips in the lagoons and backreef. We provide compelling evidence that competition helps drive distribution patterns and spatial separation of a marine predator community, and highlight that competitive advantages may not be constant but rather dependent on habitats.     

Home range and movements of male feral cats (Felis catus) in a semiarid woodland environment in central Australia

There is a paucity of data on the movement patterns of feral cats in Australia. Such data can be used to refine control strategies and improve track‐based methods of monitoring populations of feral cats. In this study the home ranges and movements of male feral cats were examined over 3.5 years in a semiarid woodland environment in central Australia. Two home range estimators were used in the examination: (i) minimum convex polygon (MCP); and (ii) fixed kernel. The most widely used method of estimating home range in feral cats is MCP, while the fixed kernel method can be used to identify core areas within a home range. On the basis of the MCP method, the long‐term home ranges of feral cats in central Australia were much larger than those recorded elsewhere (mean, 2210.5 ha). Twenty‐four hour home ranges were much smaller (mean, 249.7 ha) and feral cats periodically shifted their 24 h ranges within the bounds of their long‐term home ranges. Core area analysis indicated marked heterogeneity of space use by male feral cats. Several instances where feral cats moved large distances (up to 34 km) were recorded. These long distance movements may have been caused by nutritional stress. Using data from the literature, it is shown that prey availability is a primary determinant of long‐term home range size in feral cats. The relevance of the results to the design of management strategies for feral cats in central Australia is also discussed.     

Running before the storm: blacktip sharks respond to falling barometric pressure associated with Tropical Storm Gabrielle

A small population of juvenile (<1 year old) blacktip sharks Carcharhinus limbatus responded to the approach of a tropical storm by moving to deeper water. Examination of meterological variables suggested that the movement of the blacktip sharks was triggered by a drop in barometric pressure associated with the approach of the storm. This response was consistent for all the fish being studied, and all blacktip sharks returned to the shallow nursery area after the storm's passage, suggesting that this was an innate behaviour.     

The shark nursery of Bulls Bay,South Carolina,with a review of the shark nurseries of the southeastern coast of the United States

Synopsis Shark nurseries, or nursery areas, are geographically discrete parts of a species range where the gravid females of most species of coastal sharks deliver their young or deposit their eggs, and where their young spend their first weeks, months, or years. These areas are usually located in shallow, energy rich coastal areas where the young find abundant food and have little predation by larger sharks. Nurseries are characterized by the presence of both gravid females and free swimming neonates. Neonates are young bearing fresh, unhealed umbilical scars in the case of placental species, or those at or near the birth size in aplacental species. Bulls Bay, South Carolina, is a nursery for the blacknose, spinner, finetooth, blacktip, sandbar, dusky, Atlantic sharpnose, scalloped hammerhead, and smooth dogfish sharks. The lemon shark has its nursery in shallow waters of south Florida and the Bahamas. The bull shark has its nursery in the lagoons of the east coast of central Florida.     

Direct genetic evidence for reproductive philopatry and associated fine‐scale migrations in female blacktip reef sharks (Carcharhinus melanopterus) in French Polynesia

Conservation of top predators has been emphasized as essential in an ecosystem due to their role in trophic chain regulation. Optimizing conservation strategies for these endangered marine top predators requires direct estimates of breeding patterns and connectivity as these are essential to understanding the population dynamics. There have been some attempts to investigate breeding patterns of reef sharks from litter reconstruction using molecular analyses. However, direct fine‐scale migrations of female sharks for parturition as well as connectivity at a medium scale like between islands remain mostly unknown. We used microsatellite DNA markers and a likelihood‐based parentage analysis to determine breeding patterns of female blacktip reef sharks in Moorea (Society Islands, French Polynesia). Most females gave birth at their home island but some migrated to specific nursery areas outside the area they are attached to, sometimes going to another island 50 km away across deep ocean. Our analysis also revealed that females migrated to the same nursery for every birthing event. Many offspring showed a high level of inbreeding indicating an overall reduced population size, restricted movements and dispersal, or specific mating behaviour. Females represent the vectors that transport the genes at nursery grounds, and their fidelity should thus define reproductive units. As females seem to be philopatric, males could be the ones dispersing genes between populations. These results highlight the need to conserve coastal zones where female reef sharks seem to exhibit philopatry during the breeding season.     

Standardising Home Range Studies for Improved Management of the Critically Endangered Black Rhinoceros

Comparisons of recent estimations of home range sizes for the critically endangered black rhinoceros in Hluhluwe-iMfolozi Park (HiP), South Africa, with historical estimates led reports of a substantial (54%) increase, attributed to over-stocking and habitat deterioration that has far-reaching implications for rhino conservation. Other reports, however, suggest the increase is more likely an artefact caused by applying various home range estimators to non-standardised datasets. We collected 1939 locations of 25 black rhino over six years (2004–2009) to estimate annual home ranges and evaluate the hypothesis that they have increased in size. A minimum of 30 and 25 locations were required for accurate 95% MCP estimation of home range of adult rhinos, during the dry and wet seasons respectively. Forty and 55 locations were required for adult female and male annual MCP home ranges, respectively, and 30 locations were necessary for estimating 90% bivariate kernel home ranges accurately. Average annual 95% bivariate kernel home ranges were 20.4 ± 1.2 km², 53 ±1.9% larger than 95% MCP ranges (9.8 km² ± 0.9). When home range techniques used during the late-1960s in HiP were applied to our dataset, estimates were similar, indicating that ranges have not changed substantially in 50 years. Inaccurate, non-standardised, home range estimates and their comparison have the potential to mislead black rhino population management. We recommend that more care be taken to collect adequate numbers of rhino locations within standardized time periods (i.e., season or year) and that the comparison of home ranges estimated using dissimilar procedures be avoided. Home range studies of black rhino have been data deficient and procedurally inconsistent. Standardisation of methods is required.     

Use of space and home range characteristics of Lepilemur mittermeieri,an endangered sportive lemur endemic to the Ampasindava peninsula,north‐west Madagascar

Lepilemur mittermeieri, a little‐studied sportive lemur of north‐west Madagascar, endemic to the Ampasindava Peninsula, faces habitat loss through forest degradation and rapid fragmentation. Understanding its habitat requirement is the first step toward preservation of this threatened forest‐dependent species. In this study, we gathered data on the use of space and home range characteristics of L. mittermeieri. We studied individuals from early March to the end of June 2015 and 2016, in three sites of the Ampasindava peninsula. We radio‐tracked 15 individuals to obtain detailed information on the size and location of home ranges (around 450 hr of tracking). Direct observation and morphometric measurements provided additional data sets. Both kernel density estimation (KDE) and minimum convex polygon (MCP) methods yielded similar home range sizes (an average of 2.01 ha with KDE method and 1.96 ha with MCP method). We did not find differences in home range size between males and females, with respect to forest type or proximity to the forest edge. Home ranges overlapped and individuals showed low levels of territoriality. We highlighted a sexually‐dimorphic trait: males have longer upper canine than females. Our results constitute the first set of ecological information on Lepilemur mittermeieri and could be the basis for a conservation strategy for this endangered species with a very small distribution area.     

Spatial and temporal analysis of juvenile blacktip reef shark (Carcharhinus melanopterus) demographies identifies critical habitats

Reef shark species have undergone sharp declines in recent decades, as they inhabit coastal areas, making them an easy target in fisheries (i.e., sharks are exploited globally for their fins, meat, and liver oil) and exposing them to other threats (e.g., being part of by-catch, pollution, and climate change). Reef sharks play a critical role in coral reef ecosystems, where they control populations of smaller predators and herbivorous fishes either directly via predation or indirectly via behavior, thus protecting biodiversity and preventing potential overgrazing of corals. The urgent need to conserve reef shark populations necessitates a multifaceted approach to policy at local, federal, and global levels. However, monitoring programmes to evaluate the efficiency of such policies are lacking due to the difficulty in repeatedly sampling free-ranging, wild shark populations. Over nine consecutive years, we monitored juveniles of the blacktip reef shark (Carcharhinus melanopterus) population around Moorea, French Polynesia, and within the largest shark sanctuary globally, to date. We investigated the roles of spatial (i.e., sampling sites) and temporal variables (i.e., sampling year, season, and month), water temperature, and interspecific competition on shark density across 10 coastal nursery areas. Juvenile C. melanopterus density was found to be stable over 9 years, which may highlight the effectiveness of local and likely federal policies. Two of the 10 nursery areas exhibited higher juvenile shark densities over time, which may have been related to changes in female reproductive behavior or changes in habitat type and resources. Water temperatures did not affect juvenile shark density over time as extreme temperatures proven lethal (i.e., 33°C) in juvenile C. melanopterus might have been tempered by daily variation. The proven efficiency of time-series datasets for reef sharks to identify critical habitats (having the highest juvenile shark densities over time) should be extended to other populations to significantly contribute to the conservation of reef shark species.     

Home range dynamics,habitat selection,and survival of Greater Roadrunners

ABSTRACT Greater Roadrunners (Geococcyx californianus) are common, poorly studied birds of arid and semi‐arid ecosystems in the southwestern United States. Conservation of this avian predator requires a detailed understanding of their movements and spatial requirements that is currently lacking. From 2006 to 2009, we quantified home‐range and core area sizes and overlap, habitat selection, and survival of roadrunners (N= 14 males and 20 females) in north‐central Texas using radio‐telemetry and fixed kernel estimators. Median home‐range and core‐area sizes were 90.4 ha and 19.2 ha for males and 80.1 ha and 16.7 ha for females, respectively. The size of home range and core areas did not differ significantly by either sex or season. Our home range estimates were twice as large (x?= 108.9 ha) as earlier published estimates based on visual observations (x?= 28–50 ha). Mean percent overlap was 38.4% for home ranges and 13.7% for core areas. Male roadrunners preferred mesquite woodland and mesquite savanna cover types, and avoided the grass‐forb cover type. Female roadrunners preferred mesquite savanna and riparian woodland cover types, and avoided grass‐forb habitat. Kaplan‐Meier annual survival probabilities for females (0.452 ± 0.118[SE]) were twice that estimated for males (0.210 ± 0.108), but this difference was not significant. Mortality rates of male roadrunners were higher than those of females during the spring when males call from elevated perches, court females, and chase competing males. Current land use practices that target woody‐shrub removal to enhance livestock forage production could be detrimental to roadrunner populations by reducing availability of mesquite woodland and mesquite savanna habitat required for nesting and roosting and increasing the amount of grass‐forb habitat that roadrunners avoid.     

Sex-specific indicators of landscape use by servals: Consequences of living in fragmented landscapes

The conversion of wetlands and its associated habitat for farming, residential development and commercial purposes has led to many small disconnected patches of native vegetation surrounded by generally inhospitable human altered land uses. Such human interventions pose a serious threat to the survival of wetland dependent indicator species such as the serval (Leptailurus serval) in the Drakensberg Midlands of KwaZulu-Natal, South Africa. To address the effects of habitat fragmentation on serval space use we used GPS fixes of collared individuals for 100 days between May 2013 and January 2014. We tracked five females and eleven male servals with GPS-UHF collars. Fragmentation indices at the class level (wetland, forest with bushland, grassland and cropland) were measured within the collared serval's minimum convex polygon area (MCP) of home range. Serval use points and fragmentation indices were extracted to each sub-landscape unit of 2 km² falling within the MCP home range. The response of serval population to fragmentation indices was analyzed using generalized linear models at three levels: male, female and the overall population. We found that core area of wetland positively explained landscape use by servals. Effects of forest core area, forest proximity and patch richness were important for landscape use by male servals. Male and female serval use declined with increase in complex shapes of forest and grassland, wetland clumpiness for females and overall, and cropland patches for servals overall. Our results showed that wetland is a key determinant for the survival of male and female servals in fragmented landscapes therefore effective conservation of serval population demands adequate viable native habitat.     

Homing ability of young lemon sharks, <Emphasis Type="Italic">Negaprion brevirostris</Emphasis>

Synopsis We carried out the first experimental study testing an elasmobranchs ability to return home. We displaced juvenile lemon sharks,Negaprion brevirostris, 4–16 km from their observed home ranges at Bimini Islands, Bahamas during daylight and at night. We tracked all sharks except one back to the Bimini Islands and most returned to their home ranges observed before displacement. Even sharks displaced to a site closer to another island with suitable habitat for young lemon sharks returned to their home ranges at Bimini Islands. Sharks displayed a preferred compass direction (PCD) toward the east as their first swimming direction after release, suggesting an innate sense of direction. This bearing was followed shortly afterwards by a home-oriented direction. Swimming speeds prior to reaching shore were approximately twice as fast than the usual cruising speed reported for juvenile lemon sharks. The return of young (0–2 years), inexperienced sharks to their original home range indicate high site fidelity and an ability to home.     

Spatial ecology of juvenile lemon sharks (<Emphasis Type="Italic">Negaprion brevirostris</Emphasis>) in tidal creeks and coastal waters of Eleuthera,The Bahamas

Fisheries exploitation and habitat alteration are threatening lemon shark (Negaprion bevirostris) populations because they use nearshore regions as nursery sites. As such, there is a need for information on the spatial ecology of juvenile lemon sharks to identify critical habitats that require protection, as well as to understand their basic ecology. The purpose of this study was to determine the habitat preferences and movement patterns of juvenile lemon sharks along a sub-section of coastline characterized by coastal flats and tidal creeks of Eleuthera, The Bahamas. Eleven juvenile lemon sharks (766 ± 127 mm total length; mean±SD) were captured from various tidal creeks within the 23 km study area and were surgically implanted with acoustic transmitters. A series of 27 hydrophone receivers acted as a passive monitoring array to detect tagged individuals as they moved among habitats. Findings suggest that juvenile lemon sharks tagged in this study prefer shallow water habitats within tidal creeks, and typically display high site fidelity with occasional forays to alternate habitats or creeks. In fact, more than 90% of tagged lemon sharks had the greatest percentage of detections located at a receiver at or close to the location where they were tagged. There was no evidence of differences in diel or seasonal movement and habitat use. Knowledge gained from this study will be useful for directing future conservation and management strategies including coastal development plans and marine protected areas.     

Residency and movement patterns of bonnethead sharks, Sphyrna tiburo, in a large Florida estuary

The use of a coastal estuary by bonnethead sharks, Sphyrna tiburo, was examined by acoustic monitoring, gillnet sampling and tag- recapture studies. Acoustic monitoring data were used to define the residency and movement patterns of sharks within Pine Island Sound, Charlotte Harbor, Florida. Sharks were monitored for periods of 1–173 days with individuals regularly moving in and out of the detection range of the acoustic system. Patterns of movement could not be correlated with tidal level or time of day. Home range sizes within the Pine Island Sound population were typically small with individuals using core areas on a daily basis. However, core areas shifted within the study site over time resulting in eventual usage of most of the available habitat. Gillnet sampling revealed that S. tiburo were abundant in shallow water near seagrass beds, but that presence of individuals at specific sites was variable. Tag-recapture data showed that most individuals remained within the Pine Island Sound region over time and did not appear to undergo long coastal migrations. The movement and residence patterns of S. tiburo suggest that individuals are resident within the estuary, but do not show site fidelity to specific areas within the estuary.     

A local nearest-neighbor convex-hull construction of home ranges and utilization distributions

We describe a new method for estimating the area of home ranges and constructing utilization distributions (UDs) from spatial data. We compare our method with bivariate kernel and α-hull methods, using both randomly distributed and highly aggregated data to test the accuracy of area estimates and UD isopleth construction. The data variously contain holes, corners, and corridors linking high use areas. Our method is based on taking the union of the minimum convex polygons (MCP) associated with the k−1 nearest neighbors of each point in the data and, as such, has one free parameter k. We propose a "minimum spurious hole covering" (MSHC) rule for selecting k and interpret its application in terms of type I and type II statistical errors. Our MSHC rule provides estimates within 12% of true area values for all 5 data sets, while kernel methods are worse in all cases: in one case overestimating area by a factor of 10 and in another case underestimating area by a factor of 50. Our method also constructs much better estimates for the density isopleths of the UDs than kernel methods. The α-hull method does not lead directly to the construction of isopleths and also does not always include all points in the constructed home range. Finally we demonstrate that kernel methods, unlike our method and the α-hull method, does not converges to the true area represented by the data as the number of data points increase.     

Influences of parturition on home range and microhabitat use of female black-faced impalas

The use of home ranges and microhabitats by female black-faced impalas Aepyceros melampus petersi was investigated using radio-telemetry over a 4-month period that coincided with parturition at Ongava Game Reserve, north-west Namibia. The aim of the study was to test how ranging patterns and microhabitat use were affected by the presence of young. The mean minimum convex polygon (MCP) home range of the 11 tracked female black-faced impalas exceeded the largest of home ranges of impalas elsewhere by at least six times, and females used significantly larger total ranges (MCPs) after birth, although (50% kernel) core areas did not change significantly. Females were more likely to be solitary or alone with a lamb in the first week of their lambs' lives than in the periods after or before that time, and they both shifted their home ranges and changed their microhabitat use after parturition. Decisions on microhabitat and space use by female black-faced impalas appeared to be affected by parturition.     

Utilisation of a tropical bay as a nursery area by sharks of the families Carcharhinidae and Sphyrnidae

Synopsis At least eight species of sharks of the families Carcharhinidae and Sphyrnidae use Cleveland Bay in northern Australia as a communal nursery area.Carcharhinus dussumieri, C. fitzroyensis, C. limbatus andC. tilstoni use the bay as a seasonal primary nursery, with juveniles occurring in it for only a few months each year immediately after birth. Alternatively,Carcharhinus sorrah, Rhizoprionodon acutus andR. taylori use the bay as a year-round primary and secondary nursery, with juveniles remaining in it up to the size at maturity. AdultR. taylori also persist in the bay, a behavioural pattern possibly explained by their small maximum size. While present immediately after birth the type of utilisation pattern displayed bySphyrna lewini could not be clarified in this study. Although diets of these species in the bay are similar, there is probably little direct competition for food due to the highly productive habitats in the bay supporting an abundance of food resources. The highest numbers of juveniles occur when prey species are the most abundant, and when temporal separation of some seasonally-occurring species of sharks in effect.     

Long-term natal site-fidelity by immature lemon sharks (Negaprion brevirostris) at a subtropical island

Although many sharks begin their life confined in nursery habitats, it is unknown how rapidly they disperse away from their natal area once they leave the nursery. We examine this issue in immature lemon sharks ( Negaprion brevirostris ) from the time they leave the nursery (∼ age 3) at a subtropical island (Bimini, Bahamas), through to the onset of sexual maturity (∼ age 12). From 1995 to 2007 we tagged and genotyped a large fraction of the nursery-bound sharks at this location (0–3 years of age, N  = 1776 individuals). From 2003 to 2007 we sampled immature sharks aged from 3 to 11 years ( N  = 150) living around the island and used physical/genetic tag recaptures coupled with kinship analysis to determine whether or not each of these 'large immature sharks' was locally born. We show that many island-born lemon sharks remain close to their natal area for long periods (years) after leaving the nursery; more than half of the sampled sharks up to 135 cm total length (∼6 years old) were locally born. The fraction of locally born sharks gradually declined with increasing shark size, indicating that dispersal is relatively slow and does not primarily occur after sharks reach a threshold size. Local conservation measures (e.g. localized fishery closures, marine protected areas) can therefore help protect island-born lemon sharks even after they leave the nursery habitat.     

Stable isotope analysis of vertebrae reveals ontogenetic changes in habitat in an endothermic pelagic shark

Ontogenetic changes in habitat are driven by shifting life-history requirements and play an important role in population dynamics. However, large portions of the life history of many pelagic species are still poorly understood or unknown. We used a novel combination of stable isotope analysis of vertebral annuli, Bayesian mixing models, isoscapes and electronic tag data to reconstruct ontogenetic patterns of habitat and resource use in a pelagic apex predator, the salmon shark (Lamna ditropis). Results identified the North Pacific Transition Zone as the major nursery area for salmon sharks and revealed an ontogenetic shift around the age of maturity from oceanic to increased use of neritic habitats. The nursery habitat may reflect trade-offs between prey availability, predation pressure and thermal constraints on juvenile endothermic sharks. The ontogenetic shift in habitat coincided with a reduction of isotopic niche, possibly reflecting specialization upon particular prey or habitats. Using tagging data to inform Bayesian isotopic mixing models revealed that adult sharks primarily use neritic habitats of Alaska yet receive a trophic subsidy from oceanic habitats. Integrating the multiple methods used here provides a powerful approach to retrospectively study the ecology and life history of migratory species throughout their ontogeny.