Diverse foraging opportunities drive the functional response of local and landscape-scale bear predation on Pacific salmon

The relationship between prey abundance and predation is often examined in single habitat units or populations, but predators may occupy landscapes with diverse habitats and foraging opportunities. The vulnerability of prey within populations may depend on habitat features that hinder predation, and increased density of conspecifics in both the immediate vicinity and the broader landscape. We evaluated the relative effects of physical habitat, local, and neighborhood prey density on predation by brown bears on sockeye salmon in a suite of 27 streams using hierarchical Bayesian functional response models. Stream depth and width were inversely related to the maximum proportion of salmon killed, but not the asymptotic limit on total number killed. Interannual variation in predation was density dependent; the number of salmon killed increased with fish density in each stream towards an asymptote. Seven streams in two geographical groups with ≥23 years of data in common were then analyzed for neighborhood density effects. In most (12 of 18) cases predation in a stream was reduced by increasing salmon abundance in neighboring streams. The uncertainty in the estimates for these neighborhood effects may have resulted from interactions between salmon abundance and habitat that influenced foraging by bears, and from bear behavior (e.g., competitive exclusion) and abundance. Taken together, the results indicated that predator–prey interactions depend on density at multiple spatial scales, and on habitat features of the surrounding landscape. Explicit consideration of this context dependency should lead to improved understanding of the ecological impacts of predation across ecosystems and taxa.     

Do monk seals exert top‐down pressure in subphotic ecosystems?

Patterns of subphotic fish assemblages on seamounts in the Northwestern Hawaiian Islands were identified and compared for potential structuring influences, including the bottom-up effects of regional oceanic productivity and top-down predation pressure exerted by visiting monk seals ( Monachus schauinslandi ). Patterns in fish size, density, and biomass were evaluated at the deep extreme (350–500 m) of the seals feeding range to avoid confounding effects of diverse shallow habitats ( e.g. , coral reefs). Fish number and size were used to calculate biomass density of the seamount fish assemblages that were then compared to the independent variables of summit depth, substrate type, relief, oceanic productivity, distance to seal colonies, and seal colony population. Only the variables of distance to seal colony and seal colony population were retained in a multiple regression model that explained 31% of the variance. Despite the presence of obvious regional differences in oceanic productivity, the overall patterns in the subphotic fish assemblages are better explained by the top-down hypothesis of predation pressure from monk seals.     

Estimation of In-Water Density and Abundance of Harbor Seals

Ecologists and managers require accurate population estimates of marine mammals to assess potential anthropogenic threats to these animals. We present estimates of in-water density and abundance of a distinct stock of harbor seals (Phoca vitulina richardii) in Hood Canal, Washington, USA. We used aerial line-transect survey data collected from 2013 to 2016 to directly estimate harbor seal density and abundance in the waters of Hood Canal, a deep-water fjord in the Salish Sea. We estimated a correction factor for trackline detection probability from dive and surface time data gathered from regional seal tagging studies, and applied this factor to correct for seals missed on the trackline during surveys. We applied conventional and multiple covariate line-transect approaches in the analysis. The resulting best estimate of in-water density of harbor seals in the Hood Canal study region was 5.80 seals/km², with an estimated abundance of 2,009 seals. We did not derive a correction factor to account for the number of seals on land (i.e., hauled out). Therefore, these estimates do not reflect total stock size but provide a starting point to evaluate potential influences of anthropogenic activities, particularly those involving underwater noise, on this marine mammal stock. © 2021 The Wildlife Society.     

Functional responses of the rough-legged buzzard in a multi-prey system

The functional response is a key element of predator–prey interactions. Basic functional response theory explains foraging behavior of individual predators, but many empirical studies of free-ranging predators have estimated functional responses by using population-averaged data. We used a novel approach to investigate functional responses of an avian predator (the rough legged-buzzard Buteo lagopus Pontoppidan, 1763) to intra-annual spatial variation in rodent density in subarctic Sweden, using breeding pairs as the sampling unit. The rough-legged buzzards responded functionally to Norwegian lemmings (Lemmus lemmus L. 1758), grey-sided voles (Myodes rufocanus Sundevall, 1846) and field voles (Microtus agrestis L. 1761), but different rodent prey were not utilised according to relative abundance. The functional response to Norwegian lemmings was a steep type II curve and a more shallow type III response to grey-sided voles. The different shapes of these two functional responses were likely due to combined effects of differences between lemmings and grey-sided voles in habitat utilisation, anti-predator behaviour and size-dependent vulnerability to predation. Diet composition changed less than changes in relative prey abundance, indicating negative switching, with high disproportional use of especially lemmings at low relative densities. Our results suggest that lemmings and voles should be treated separately in future empirical and theoretical studies in order to better understand the role of predation in this study system.     

Temporal variation in distribution and density of ice-obligated seals in western Hudson Bay, Canada

Recent unidirectional climatic trends and changes in top predator population ecology suggest that long-term modifications may be happening in Hudson Bay, Canada. Effects of such changes on ice-obligated seal populations are expected but long-term studies are required to differentiate climate-induced changes from natural variation. We conducted strip-transect surveys in late spring in 1995–1997, 1999–2000 and 2007–2008 to estimate distribution, density and abundance of ice-obligated ringed (Phoca hispida) and bearded (Erignathus barbatus) seals in western Hudson Bay. When hauled out, ringed seals preferred land-fast and consolidated pack ice, whereas bearded seals preferred unconsolidated pack ice. Bearded and ringed seal density estimates varied from 0.0036 to 0.0229 seals/km² of ice and from 0.46 to 1.60 seals/km² of ice, respectively. Strong inter-annual variations were recorded in the abundance estimates of both species, with the largest abundance estimates in 1995 (104,162 and 1,494 ringed and bearded seals, respectively) and the lowest in 2008 for ringed seals (33,701) and 1997 for bearded seals (278). A sine function best described seal density estimates in western Hudson Bay and suggested a decadal cycle. Previous studies that reported low ringed seal demographic parameters in the 1990s and a recovery in the 2000s supported our interpretation of the survey results. We discuss our results in the context of climate warming and suggest that a long-term decline in ice-obligated seal density estimates may overlay a possible natural decadal cycle.     

Sea lice escape predation on their host

Parasites seldom have predators but often fall victim to those of their hosts. How parasites respond to host predation can have important consequences for both hosts and parasites, though empirical investigations are rare. The exposure of wild juvenile salmon to sea lice (Lepeophtheirus salmonis) from salmon farms allowed us to study a novel ecological interaction: the response of sea lice to predation on their juvenile pink and chum salmon hosts by two salmonid predators-coho smolts and cut-throat trout. In approximately 70% of trials in which a predator consumed a parasitized prey, lice escaped predation by swimming or moving directly onto the predator. This trophic transmission is strongly male biased, probably because behaviour and morphology constrain female movement and transmission. These findings highlight the potential for sea lice to be transmitted up marine food webs in areas of intensive salmon aquaculture, with implications for louse population dynamics and predatory salmonid health.     

Environmental conditions modify density-dependent salmonid recruitment: Insights into the 2016 recruitment crash in Wales

1. Understanding the effects of density-dependent and density-independent factors on recruitment is often inhibited by difficulties quantifying their relative contributions in highly variable recruitment data. Use of data-driven statistical methods with data that include one or more extreme recruitment events could help overcome these difficulties.
2. Juvenile Atlantic salmon and trout abundances in Wales have declined over the last 2 decades, and 2016 was a notably poor recruitment year in rivers around southern Europe, including England and Wales. The 2016 recruitment crash coincided with extreme winter weather conditions, leading to speculation that unusually warm temperatures and high flows adversely affect salmonid recruitment and caused the 2016 crash, although this remains untested.
3. We developed data-driven statistical models to: (1) describe juvenile salmonid recruitment from density-dependent and density-independent factors; and (2) assess whether the density-independent factors probably contributed to the 2016 salmon recruitment crash. We compiled salmon and trout young-of-year juvenile abundances from electrofishing surveys, egg deposition estimates and river flow and air temperature data from 2001–2017 for seven Welsh rivers, broadly representative of rivers around Wales. We used river flow and air temperature data to derive ecologically and behaviourally meaningful density-independent explanatory variables.
4. Salmonid recruitment in Wales was best described using density-dependent and density-independent factors, especially for salmon: after accounting for a concave relationship with egg deposition, salmon juvenile abundance was reduced under (1) warmer spawning temperatures that might inhibit spawning, and (2) higher flood frequencies during pre-emergence and emergence that might washout eggs or alevins. Results were less clear for trout, perhaps because they are behaviourally more plastic.
5. Our findings provide empirical support for general and predictable effects of temperature and flow during spawning and emergence on salmonid—especially salmon—recruitment in Wales. Furthermore, we suggest that the 2016 salmon recruitment crash was caused—in part—by particularly inclement spawning and emergence conditions, which could be more common under future climate change. Our findings suggest that future salmonid stock assessment models could include the effects of density-independent variables on recruitment to improve their predictive power.

     

Assessing Habitat Suitability for Juvenile Atlantic Salmon in Relation to In‐Stream Restoration and Discharge Variability

In‐stream restoration often aims at increasing the availability of the stream habitat suitable for salmonid fishes, thus creating potential for increased fish abundance. We assessed the success of in‐stream restoration of River Kiiminkijoki, northern Finland, by combining River2D habitat hydraulic modeling and fish density monitoring at the same sites, with data from multiple restored and reference reaches for 3 years both before and after restoration. We modeled the effects of restoration on the area suitable (weighted usable area, WUA) for juvenile Atlantic salmon from post‐hatching to age‐1 fish. Wetted width in the restored reaches increased by 8.1% on average compared with only ?0.2% change in the reference reaches. Habitat time series across 10 years showed significant increases in the amount of suitable habitat under summer conditions for both age‐0 and age‐1 salmon. However, improvement of overwintering habitats was marginal or nonexistent. Densities of age‐1 salmon showed no response to restoration. Low river discharge during the winter was correlated with low salmon densities the following summer. It thus appears that variability in wintertime discharge, and associated high interannual variation of WUA values, overrode the almost 20% increase in average post‐ versus pre‐restoration summertime WUA. Our study shows that the combination of hydraulic modeling and biological monitoring is a promising approach to stream restoration assessment.     

Factors affecting early mortality in the South American fur seal (Arctocephalus australis) in Peru: density-related effects and predation

Early mortality in the South American fur seal ( Arctocephatus australis ) in Peru is considerably higher than that seen in any other population of fur seal; 31–49% in the first month as opposed to a maximum of 20% in other populations. In 1987 and 1988, pup mortality was found to be enhanced by density-related effects and predation on pups by the southern sea-lion ( Olaria byronia ). At a high density beach, where 60% of the fur seals bred, mortality correlated with the number of females ashore as did aggression. Female aggression was driven to high levels by a combination of high density and movement within the colony to thermoregulate. However, even at a low density, beach mortality remained high due to a high level of predation by male southern sea-lions (low density beach 5·0–8·3% of all pups; high density beach 0·2%). Predation rates were lower at the high density beach because male fur seals expelled most attacking sea-lions during the breeding season (low density beach 10·0% of attacking sea-lions expelled; high density beach 58·8%).     

First documentation of leopard seal predation of South Georgia pintail duck

Leopard seals are regular winter visitors to Bird Island, South Georgia, where they mostly prey on fur seals and penguins, and to a lesser extent on Antarctic krill and fish. Leopard seals can exploit many different species, but there are no records of predation on flying shorebirds in the wild. On 4 October 2008, an individually identified juvenile leopard seal female was observed killing and eating a South Georgia Pintail duck. It also preyed on Antarctic fur seals and gentoo and macaroni penguins during its 2-month temporary residency around the island. The varied diet of this seal exemplifies the generalist prey utilization typical of its species. Long-term diet studies at Bird Island and the published record suggest that predation on ducks is a rather exceptional finding; individual ducks are more likely to escape leopard seal attacks than penguins and provide a far less substantial ration. This note documents the first observation of this species of duck in the diet of leopard seals.     

Assessing the potential impact of salmon fisheries management on the conservation status of harbour seals (Phoca vitulina) in north-east Scotland

Conservation efforts are often constrained by uncertainty over the factors driving declines in marine mammal populations. In Scotland, there is concern over the potential impact of unrecorded shooting of seals, particularly where this occurs near Special Areas of Conservation. Here, we show that the abundance of harbour seals Phoca vitulina in the Moray Firth, north-east Scotland, declined by 2–5% per annum between 1993 and 2004. Records from local salmon fisheries and aquaculture sites indicated that 66–327 harbour seals were shot each year between 1994 and 2002. Matrix models and estimates of potential biological removal indicate that this level of shooting is sufficient to explain observed declines. Nevertheless, uncertainty over the number and identity of seals shot means that other factors such as changes in food availability may be contributing. Recent conservation measures markedly reduced the recorded levels of shooting in 2003 and 2004. In 2005, a coordinated management plan was developed to protect salmon fisheries interests while minimizing impacts on local seal populations. Comprehensive monitoring of future population trends and improved regulation of culls are now required to provide more robust assessments of the impact of human persecution on harbour seal populations in the Moray Firth and in other parts of the UK.     

Haul-out behavior of harbor seals (Phoca vitulina) in Hood Canal, Washington

The goal of this study was to model haul-out behavior of harbor seals (Phoca vitulina) in the Hood Canal region of Washington State with respect to changes in physiological, environmental, and temporal covariates. Previous research has provided a solid understanding of seal haul-out behavior. Here, we expand on that work using a generalized linear mixed model (GLMM) with temporal autocorrelation and a large dataset. Our dataset included behavioral haul-out records from archival and VHF radio tag deployments on 25 individual seals representing 61,430 seal hours. A novel application for increased computational efficiency allowed us to examine this large dataset with a GLMM that appropriately accounts for temporal autocorellation. We found significant relationships with the covariates hour of day, day of year, minutes from high tide and year. Additionally, there was a significant effect of the interaction term hour of day : day of year. This interaction term demonstrated that seals are more likely to haul out during nighttime hours in August and September, but then switch to predominantly daylight haul-out patterns in October and November. We attribute this change in behavior to an effect of human disturbance levels. This study also examined a unique ecological event to determine the role of increased killer whale (Orcinus orca) predation on haul-out behavior. In 2003 and 2005 these harbor seals were exposed to unprecedented levels of killer whale predation and results show an overall increase in haul-out probability after exposure to killer whales. The outcome of this study will be integral to understanding any changes in population abundance as a result of increased killer whale predation.     

Do shark declines create fear‐released systems?

We illustrate the theoretical plausibility that fishery removal of sharks can indirectly alter predation pressure on different fish species via the behavioural responses of mesoconsumers released from predator intimidation. Our dynamic state variable model of foraging decisions by harbour seals, a mesopredator, predicted indirect effects of the removal of Pacific sleeper sharks on two species consumed by seals, Pacific herring and walleye pollock, as mediated by seal behaviour. Herring, a fatty fish, form near‐surface aggregations that often are ephemeral and widely dispersed. Pollock are found in the deeper strata preferred by sharks and have lower energy density than herring, but also are larger and their more continuous distribution potentially makes them the more predictable resource for seals. During simulations, predation risk from sharks produced an asymmetric trophic cascade mediated by the seal's underutilisation of deeper, riskier strata. Risk management by seals reduced mortality on pollock, which required riskier access in deep strata, while increasing mortality on herring, which could be accessed with less risk in shallow strata. This effect, however, attenuated if herring were scarcer and seal energy state was poor. During shark removal scenarios, seals shifted to deeper strata, increasing pollock consumption and substantially decreasing use of herring; the proportional change was greatest if seals were in good energy state. Prior studies have addressed how shark declines might affect community structure through density responses by species consumed by sharks; earlier models incorporating antipredator behaviour of the mesopredator (i.e. Ecosim/Ecospace) allow for activity reduction but not for the spatial shifts that altered the asymmetric trophic cascade in our simulations. Our results suggest that shark declines might have stronger ecological consequences than previously recognised if we account for spatial and diet shifts by mesoconsumers released from shark intimidation.     

The potential influence of Atlantic salmon Salmo salar and brown trout Salmo trutta on density and breeding of the white‐throated dipper Cinclus cinclus

Interactions between birds and fish are often overlooked in aquatic ecosystems. We studied the influence of Atlantic salmon and brown trout on the breeding population size and reproductive output of the white‐throated dipper in a Norwegian river. Acidic precipitation led to the extinction of salmon, but salmon recolonized after liming was initiated in 1991. We compared the dipper population size and reproductive output before (1978–1992) and after (1993–2014) salmon recolonization. Despite a rapid and substantial increase in juvenile salmon, the breeding dipper population size and reproductive output were not influenced by juvenile salmon, trout, or total salmonid density. This might be due to different feeding strategies in salmonids and dippers, where salmonids are mainly feeding on drift, while the dipper is a benthic feeder. The correlation between the size of the dipper population upstream and downstream of a salmonid migratory barrier was similar before and after recolonization, indicating that the downstream territories were not less attractive after the recolonization of salmon. Upstream dipper breeding success rates declined before the recolonization event and increased after, indicating improved water quality due to liming, and increasing invertebrate prey abundances and biodiversity. Surprisingly, upstream the migratory barrier, juvenile trout had a weak positive effect on the dipper population size, indicating that dippers may prey upon small trout. It is possible that wider downstream reaches might have higher abundances of alternative food, rending juvenile trout unimportant as prey. Abiotic factors such as winter temperatures and acidic precipitation with subsequent liming, potentially mediated by prey abundance, seem to play the most important role in the life history of the dipper.     

ABUNDANCE OF RINGED SEALS (PUSA HISPIDA) IN THE FJORDS OF SPITSBERGEN, SVALBARD, DURING THE PEAK MOLTING PERIOD

Ringed seal (Pusa hispida) abundance in Spitsbergen, Svalbard, was estimated during the peak molting period via aerial, digital photographic surveys. A total of 9,145 images, covering 41.7%–100% of the total fast‐ice cover (1,496 km²) of 18 different fjords and bays, were inspected for the presence of ringed seals. A total of 1,708 seals were counted, and when accounting for ice areas that were not covered by images, a total of 3,254 (95% CI: 3,071–3,449) ringed seals were estimated to be hauled out during the surveys. Extensive behavioral data from radio‐tagged ringed seals (collected in a companion study) from one of the highest density fjords during the molting period were used to create a model that predicts the proportion of seals hauled out on any given date, time of day, and under various meteorological conditions. Applying this model to the count data from each fjord, we estimated that a total of 7,585 (95% CI: 6,332–9,085) ringed seals were present in the surveyed area during the peak molting period. Data on interannual variability in ringed seal abundance suggested higher numbers of seals in Van Keulenfjorden in 2002 compared to 2003, while other fjords with very stable ice cover showed no statistical differences. Poor ice conditions in general in 2002 probably resulted in seals from a wide area coming to Van Keulenfjorden (a large fjord with stable ice in 2002). The total estimated number of ringed seals present in the study area at the time of the survey must be regarded as a population index, or at least a minimum estimate for the area, because it does not account for individuals leaving and arriving, which might account for a considerable number of animals. The same situation is likely the case for many other studies reporting aerial census data for ringed seals. To achieve accurate estimates of population sizes from aerial surveys, more extensive knowledge of ringed seal behavior will be required.     

USE OF PHOTOGRAPHIC IDENTIFICATION IN CAPTURE-RECAPTURE STUDIES OF MEDITERRANEAN MONK SEALS

The use of photo-identification and its reliability in capture-recapture studies of Mediterranean monk seals were assessed using slides collected in the colony at Cap Blanc, western Sahara, from 1993 to 1996. Five tests indicated that researchers involved in photo-identification were proficient in matching slides of identified seals, consistent in classifying the side of the seal shown in slides and in assigning the morphological stage of the seal, and that changes of markings over a period of three years were insufficient to affect matching success. The certainty of identifying a seal was not dependent on the number of slides used but on distinctiveness of the markings and the quality of the slides taken. Capture-recapture abundance estimates were biased upwards when including poor quality slides. The exclusive use of excellent- and good quality slides provided the best estimates. The proportion of distinctive seals varied between morphological stages and was significantly lower in juveniles. When including the identification histories of juveniles, the heterogeneity of capture probabilities was higher. Therefore, abundance estimates were less biased when all juveniles were considered as non-distinctive seals. Reliable abundance estimates required a balance between duration of capture occasions and time interval between these.     

Effects of alternative prey on predation by small mammals on gypsy moth pupae

Previous work shows that predation by small mammals is a dominant cause of mortality of low-density gypsy moths in North America and that declines in small mammal density result in increases in gypsy moth density. Here we examined whether predation by small mammals is density dependent by way of a type III functional response, and how predation is influenced by alternative prey. First we showed that the preference of predators for gypsy moth pupae was low compared to other experimental prey items, such as mealworm pupae and sunflower seeds. Predation on gypsy moth pupae was characterized by a type II functional response with percent predation highest at the lowest prey densities, whereas the functional response to sunflower seeds was characterized by a type III functional response in which predation increased with increasing prey density. These results suggest that predation by small mammals is unlikely to stabilize low-density gypsy moth populations.     

The use (and misuse) of archaeological salmon data to infer historical abundance in North America with a focus on New England

Information about historical animal or plant abundance often either explicitly or implicitly informs current conservation practice. If it can be shown that an organism was not historically abundant in a region, its conservation importance may be downgraded. In contrast to abundant archaeological support for historic importance of salmon in the Pacific Northwest, historic abundance of Atlantic salmon in New England has been called into question based on the rarity of salmon bones in archaeological sites. These data have been used to argue that the importance of salmon to the region has been exaggerated and that expensive restoration efforts in some rivers should be reconsidered. Here, we argue that lack of archaeological bone fragment abundance does not make a convincing case against historical Atlantic salmon abundance in New England for three primary reasons. First, salmon bones were rare or absent at sites that still have large salmon runs. Second, the lack of salmonid bones in general at archaeological sites suggests poor preservation and/or recovery of bone for these species relative to bones of other fishes. Third, given the presence of large numbers of non-salmonid anadromous fish in the site areas where people fished and deposited fish bones, power to detect salmon bones in studies to date may have been generally low. We present reliable historical accounts that help build a convincing case that salmon were historically abundant in New England rivers. We suggest that rarity of salmon bones in the existing archaeological data should not have unwarranted influence on present-day conservation decision-making in New England.     

SUCCESSFUL USE OF A TRANSLOCATION PROGRAM TO INVESTIGATE DIVING BEHAVIOR IN A MALE AUSTRALIAN FUR SEAL, ARCTOCEPHALUS PUSILLUS DORIFERUS

This study reports some of the first foraging behavior data collected for male fur seals. A nonbreeding male Australian fur seal, Arctocephalus pusillus doriferus , captured at a commercial salmon farm in southern Tasmania, Australia, was relocated 450 km from the site of capture. The animal was equipped with a geolocating time-depth recorder that recorded diving behavior and approximate location for the 14.4 d that it took the seal to travel down the east coast of Tasmania and be recaptured at the salmon farm. During its time at sea, the seal spent most of its time over the relatively shallow shelf waters. It spent 30% of its time ashore on a number of different haul-out sites. The deepest dive was 102 m and the maximum duration was 6.8 min. "Foraging" type dives made up 31.2% of the time at sea and had a median duration of 2.5 min and a median depth of 14 m. The seal performed these dives more commonly during the latter part of its time at sea, while it was on the east coast. Unlike other fur seal species studied to date, there was no evidence of a diurnal foraging pattern; it made dives at all times of the day and night.     

Killing of Cape fur seal (Arctocephalus pusillus pusillus) pups by brown hyenas (Parahyaena brunnea) at mainland breeding colonies along the coastal Namib Desert

Brown hyenas (Parahyaena brunnea) scavenge and kill seal pups at mainland Cape fur seal (Arctocephalus pusillus pusillus) colonies. The prey encounter interval and interval between kills depended on seal density, and increased density resulted in an increase of the capture rate and increase in hunting efficiency from 14% in November to 47% in January. The time brown hyenas spent at the seal colony decreased with increasing seal density and increasing air temperatures. Nevertheless, they were regularly active during the day when less adult seals were present at the colony, which indicates that the attendance of adult seals might play a role in the choice of foraging time. Brown hyenas killed seal pups throughout the study period. The predation rate was independent of the availability of non-violent mortalities, but the absolute number of kills was positively density-dependent. Mass kill events were recorded throughout the study period and are therefore not unusual occurrences. The overabundance of easy and vulnerable prey may lead to an over stimulus situation that triggers killing independent of the consumption of the prey or the hunger state.