The effects of predator intimidation on habitat use and behavior of prey species are rarely quantified for large marine vertebrates over ecologically relevant scales. Using state space movement ...models followed by a series of step selection functions, we analyzed movement data of concurrently tracked prey, bowhead whales (Balaena mysticetus; n = 7), and predator, killer whales (Orcinus orca; n = 3), in a large (63,000 km²), partially ice-covered gulf in the Canadian Arctic. Our analysis revealed pronounced predator-mediated shifts in prey habitat use and behavior over much larger spatiotemporal scales than previously documented in any marine or terrestrial ecosystem. The striking shift from use of open water (predator-free) to dense sea ice and shorelines (predators present) was exhibited gulf-wide by all tracked bowheads during the entire 3-wk period killer whales were present, constituting a nonconsumptive effect (NCE) with unknown energetic or fitness costs. Sea ice is considered quintessential habitat for bowhead whales, and ice-covered areas have frequently been interpreted as preferred bowhead foraging habitat in analyses that have not assessed predator effects. Given the NCEs of apex predators demonstrated here, however, unbiased assessment of habitat use and distribution of bowhead whales and many marine species may not be possible without explicitly incorporating spatiotemporal distribution of predation risk. The apparent use of sea ice as a predator refuge also has implications for how bowhead whales, and likely other ice-associated Arctic marine mammals, will cope with changes in Arctic sea ice dynamics as historically ice-covered areas become increasingly ice-free during summer.
Range expansions and increases in the frequency of killer whale (Orcinus orca) sightings have been documented in the eastern Canadian Arctic, presumably the result of climate change‐related sea‐ice ...declines. However, the effects of increased predator occurrence on this marine ecosystem remain largely unknown. We explore the consequences of climate change‐related range expansions by a top predator by estimating killer whale abundance and their possible consumptive effects on narwhal (Monodon monoceros) in the Canadian Arctic. Individual killer whales can be identified using characteristics such as acquired scars and variation in the shape and size of their dorsal fins. Capture–mark–recapture analysis of 63 individually identifiable killer whales photographed between 2009 and 2018 suggests a population size of 163 ± 27. This number of killer whales could consume >1,000 narwhal during their seasonal residency in Arctic waters. The effects of such mortality at the ecosystem level are uncertain, but trophic cascades caused by top predators, including killer whales, have been documented elsewhere. These findings illustrate the magnitude of ecosystem‐level modifications that can occur with climate change‐related shifts in predator distributions.
We explore the consequences of climate change‐related predator range expansions by estimating killer whale (Orcinus orca) abundance and their possible consumptive effects on narwhal (Monodon monoceros) in the Canadian Arctic. Our findings suggest a population size of 163 ± 27, a number of killer whales that could consume >1,000 narwhal during their seasonal residency in Arctic waters. These findings illustrate the magnitude of ecosystem‐level modifications that can occur with climate change‐related shifts in predator distributions.
Arctic marine mammals (AMMs) are icons of climate change, largely because of their close association with sea ice. However, neither a circumpolar assessment of AMM status nor a standardized metric of ...sea ice habitat change is available. We summarized available data on abundance and trend for each AMM species and recognized subpopulation. We also examined species diversity, the extent of human use, and temporal trends in sea ice habitat for 12 regions of the Arctic by calculating the dates of spring sea ice retreat and fall sea ice advance from satellite data (1979–2013). Estimates of AMM abundance varied greatly in quality, and few studies were long enough for trend analysis. Of the AMM subpopulations, 78% (61 of 78) are legally harvested for subsistence purposes. Changes in sea ice phenology have been profound. In all regions except the Bering Sea, the duration of the summer (i.e., reduced ice) period increased by 5–10 weeks and by >20 weeks in the Barents Sea between 1979 and 2013. In light of generally poor data, the importance of human use, and forecasted environmental changes in the 21st century, we recommend the following for effective AMM conservation: maintain and improve comanagement by local, federal, and international partners; recognize spatial and temporal variability in AMM subpopulation response to climate change; implement monitoring programs with clear goals; mitigate cumulative impacts of increased human activity; and recognize the limits of current protected species legislation.
Segregation of adult males from adult females and immature animals is known to occur in some beluga whale populations, but it is unclear if such segregation occurs in Hudson Bay, where the largest ...summering population in the world is found. Using imagery from a photographic aerial survey conducted in August 2015, we examined spatial distribution by age class with respect to several environmental variables near two of three main estuaries, Churchill and Seal River, used by Western Hudson Bay belugas in the summer season. Belugas photographed during aerial surveys were classified by age manually using an identification decision tree, and GPS coordinates of their locations were plotted in ArcGIS. Distribution by age class was examined in relation to five habitat characteristics: distance to coastal habitat, bathymetry, sea surface temperature, and extent of river plume. Habitat characteristics and the proportion of animals by age classes were similar in both estuaries, indicating no segregation, and suggesting the environmental data assessed were not associated with patterns of distribution and density of age classes at the spatial and temporal scale being investigated. Overall density of calves was almost three times higher at the Seal River; however, suggesting this location may be preferred for calf rearing in the summer season. Results provide a greater understanding of spatial patterns of beluga whale habitat use in western Hudson Bay, and information useful in conservation and management advice.
Ecotypes are groups within a species with different ecological adaptations than their conspecifics. Eastern North Pacific (ENP) killer whale (Orcinus orca) ecotypes differ in their diet, behavior, ...and morphology, but the same is not known for this species in the eastern Canadian Arctic (ECA) and Northwest Atlantic (NWA). Using compound-specific stable isotope analysis (CSIA) of amino acids (AAs), we compared δ15N patterns of the primary trophic and source AA pair, glutamic acid/glutamine (Glx) and phenylalanine (Phe), in dentine collagen of (1) sympatric ENP killer whale ecotypes with well-characterized diet differences and (2) ECA/NWA killer whales with unknown diets. δ15NGlx-Phe was significantly higher in the ENP fish-eating (FE) than mammal-eating (ME) ecotype (19.2 ± 0.4‰ vs. 13.5 ± 0.7‰, respectively). Similar bimodal variation in δ15NGlx-Phe indicated analogous dietary divisions among ECA/NWA killer whales, with two killer whales having higher δ15NGlx-Phe (16.5 ± 0.0‰) than the others (13.5 ± 0.6‰). Inferences of dietary divisions between these killer whales were supported by parallel differences in threonine δ15N (-33.5 ± 1.6‰ and -40.4 ± 1.1‰, respectively), given the negative correlation between δ15NThr and TP across a range of marine consumers. CSIA-AA results for ECA/NWA whales, coupled with differences in tooth wear (a correlate for diet), are consistent with ecotype characteristics reported in ENP and other killer whale populations, thus adding to documented ecological divergence in this species worldwide.
Abstract Ecological applications of compound-specific stable isotope analysis (CSIA) of amino acids (AAs) include 1) tracking carbon pathways in food webs using essential AA (AA ESS ) δ 13 C values, ...and 2) estimating consumer trophic position (TP) by comparing relative differences of ‘trophic’ and ‘source’ AA δ 15 N values. Despite the significance of these applications, few studies have examined AA-specific SI patterns among tissues with different AA compositions and metabolism/turnover rates, which could cause differential drawdown of body AA pools and impart tissue-specific isotopic fractionation. To address this knowledge gap, especially in the absence of controlled diet studies examining this issue in captive marine mammals, we used a paired-sample design to compare δ 13 C and δ 15 N values of 11 AAs in commonly sampled tissues (skin, muscle, and dentine) from wild beluga whales ( Delphinapterus leucas ). δ 13 C of two AAs, glutamic acid/glutamine (Glx, a non-essential AA) and, notably, threonine (an essential AA), differed between skin and muscle. Furthermore, δ 15 N of three AAs (alanine, glycine, and proline) differed significantly among the three tissues, with glycine δ 15 N differences of approximately 10 ‰ among tissues supporting recent findings it is unsuitable as a source AA. Significant δ 15 N differences in AAs such as proline, a trophic AA used as an alternative to Glx in TP estimation, highlight tissue selection as a potential source of error in ecological applications of CSIA-AA. Amino acids that differed among tissues play key roles in metabolic pathways (e.g., ketogenic and gluconeogenic AAs), pointing to potential physiological applications of CSIA-AA in studies of free-ranging animals. These findings underscore the complexity of isotopic dynamics within tissues and emphasize the need for a nuanced approach when applying CSIA-AA in ecological research.
Although predators influence behavior of prey, analyses of electronic tracking data in marine environments rarely consider how predators affect the behavior of tracked animals. We collected an ...unprecedented dataset by synchronously tracking predator (killer whales, N = 1; representing a family group) and prey (narwhal, N = 7) via satellite telemetry in Admiralty Inlet, a large fjord in the Eastern Canadian Arctic. Analyzing the movement data with a switching-state space model and a series of mixed effects models, we show that the presence of killer whales strongly alters the behavior and distribution of narwhal. When killer whales were present (within about 100 km), narwhal moved closer to shore, where they were presumably less vulnerable. Under predation threat, narwhal movement patterns were more likely to be transiting, whereas in the absence of threat, more likely resident. Effects extended beyond discrete predatory events and persisted steadily for 10 d, the duration that killer whales remained in Admiralty Inlet. Our findings have two key consequences. First, given current reductions in sea ice and increases in Arctic killer whale sightings, killer whales have the potential to reshape Arctic marine mammal distributions and behavior. Second and of more general importance, predators have the potential to strongly affect movement behavior of tracked marine animals. Understanding predator effects may be as or more important than relating movement behavior to resource distribution or bottom-up drivers traditionally included in analyses of marine animal tracking data.
To understand beluga whale (Delphinapterus leucas) estuarine use in the Nelson River estuary, southwest Hudson Bay, we recorded and examined beluga movements and habitat associations for the July ...through August period in 2002-2005. We compared locations of belugas fitted with satellite transmitters ("tags") (2002-2005) and aerial-surveyed (2003 and 2005) belugas for years of differing freshwater flow from the Nelson River which is influenced by hydroelectric activity. Using the beluga telemetry location data, we estimated an early August behavioral shift in beluga distribution patterns from local estuarine use to a progressively more migratory behavior away from the estuary. The timing of this shift in behavior was also apparent in results of beluga aerial surveys from the 1940s-1960s, despite environmental changes including later freeze-up and warming ocean temperatures. Overall, during the higher than average discharge ("wet") year of 2005, the three tagged belugas ranged farther from the Nelson River but not farther from the nearest shore along southwestern Hudson Bay, compared to the 10 tagged belugas tracked during the "dry" years of 2002-2004 with below average discharges. Aerial survey data for 2003 and 2005 display a similar dry vs. wet year shift in spatial patterns, with no significant change in overall density of belugas within the study area. In the Nelson estuary, proximity to the fresh-salt water mixing area may be more important than the shallow waters of the upper estuary. Killer whales (Orcinus orca) were observed in the Churchill area (200 km northwest) during each year of study, 2002-05, and belugas may benefit from the proximity to shallow estuary waters that provide protection from the larger-bodied predator. Study results contribute to an understanding of the influence of environmental variation on how and why belugas use estuaries although considerable uncertainties exist and additional research is required.
Management boundaries that define populations or stocks of fish form the basis of fisheries planning. In the Arctic, decreasing sea ice extent is driving increasing fisheries development, ...highlighting the need for ecological data to inform management. In Cumberland Sound, southwest Baffin Island, an indigenous community fishery was established in 1987 targeting Greenland halibut (Reinhardtius hippoglossoides) through the ice. Following its development, the Cumberland Sound Management Boundary (CSMB) was designated and a total allowable catch (TAC) assigned to the fishery. The CSMB was based on a sink population of Greenland halibut resident in the northern section of the Sound. Recent fishing activities south of the CSMB, however, raised concerns over fish residency, the effectiveness of the CSMB and the sustainability of the community-based winter fishery. Through acoustic telemetry monitoring at depths between 400 and 1200 m, and environmental and fisheries data, this study examined the movement patterns of Greenland halibut relative to the CSMB, the biotic and abiotic factors driving fish movement and the dynamics of the winter fishery. Greenland halibut undertook clear seasonal movements between the southern and northern regions of the Sound driven by temperature, dissolved oxygen, and sea ice cover with most fish crossing the CSMB on an annual basis. Over the lifespan of the fishery, landfast ice cover initially declined and then became variable, limiting accessibility to favored fisher locations. Concomitantly, catch per unit effort declined, reflecting the effect of changing ice conditions on the location and effort of the fishery. Ultimately, these telemetry data revealed that fishers now target less productive sites outside of their favored areas and, with continued decreases in ice, the winter fishery might cease to exist. In addition, these novel telemetry data revealed that the CSMB is ineffective and led to its relocation to the entrance of the Sound in 2014. The community fishery can now develop an open-water fishery in addition to the winter fishery to exploit the TAC, which will ensure the longevity of the fishery under projected climate-change scenarios. Telemetry shows great promise as a tool for understanding deep-water species and for directly informing fisheries management of these ecosystems that are inherently complex to study.
Bowhead whales (Balaena mysticetus) have a nearly circumpolar distribution, and occasionally occupy warmer shallow coastal areas during summertime that may facilitate molting. However, relatively ...little is known about the occurrence of molting and associated behaviors in bowhead whales. We opportunistically observed whales in Cumberland Sound, Nunavut, Canada with skin irregularities consistent with molting during August 2014, and collected a skin sample from a biopsied whale that revealed loose epidermis and sloughing. During August 2016, we flew a small unmanned aerial system (sUAS) over whales to take video and still images to: 1) determine unique individuals; 2) estimate the proportion of the body of unique individuals that exhibited sloughing skin; 3) determine the presence or absence of superficial lines representative of rock-rubbing behavior; and 4) measure body lengths to infer age-class. The still images revealed that all individuals (n = 81 whales) were sloughing skin, and that nearly 40% of them had mottled skin over more than two-thirds of their bodies. The video images captured bowhead whales rubbing on large rocks in shallow, coastal areas-likely to facilitate molting. Molting and rock rubbing appears to be pervasive during late summer for whales in the eastern Canadian Arctic.
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