Response of an arctic predator guild to collapsing lemming cycles

Alpine and arctic lemming populations appear to be highly sensitive to climate change, and when faced with warmer and shorter winters, their well-known high-amplitude population cycles may collapse. Being keystone species in tundra ecosystems, changed lemming dynamics may convey significant knock-on effects on trophically linked species. Here, we analyse long-term (1988–2010), community-wide monitoring data from two sites in high-arctic Greenland and document how a collapse in collared lemming cyclicity affects the population dynamics of the predator guild. Dramatic changes were observed in two highly specialized lemming predators: snowy owl and stoat. Following the lemming cycle collapse, snowy owl fledgling production declined by 98 per cent, and there was indication of a severe population decline of stoats at one site. The less specialized long-tailed skua and the generalist arctic fox were more loosely coupled to the lemming dynamics. Still, the lemming collapse had noticeable effects on their reproductive performance. Predator responses differed somewhat between sites in all species and could arise from site-specific differences in lemming dynamics, intra-guild interactions or subsidies from other resources. Nevertheless, population extinctions and community restructuring of this arctic endemic predator guild are likely if the lemming dynamics are maintained at the current non-cyclic, low-density state.     

Bottom-up regulation of a pole-ward migratory predator population

As the effects of regional climate change are most pronounced at polar latitudes, we might expect polar-ward migratory populations to respond as habitat suitability changes. The southern elephant seal (Mirounga leonina L.) is a pole-ward migratory species whose populations have mostly stabilized or increased in the past decade, the one exception being the Macquarie Island population which has decreased continuously over the past 50 years. To explore probable causes of this anomalous trend, we counted breeding female seals annually between 1988 and 2011 in order to relate annual rates of population change (r) to foraging habitat changes that have known connections with atmospheric variability. We found r (i) varied annually from −0.016 to 0.021 over the study period, (ii) was most effected by anomalous atmospheric variability after a 3 year time lag was introduced (R = 0.51) and (iii) was associated with sea-ice duration (SID) within the seals’ foraging range at the same temporal lag. Negative r years may be extrapolated to explain, at least partially, the overall trend in seal abundance at Macquarie Island; specifically, increasing SID within the seals foraging range has a negative influence on their abundance at the island. Evidence is accruing that suggests southern elephant seal populations may respond positively to a reduced sea-ice field.     

Antarctic Climate Change: Extreme Events Disrupt Plastic Phenotypic Response in Adélie Penguins

In the context of predicted alteration of sea ice cover and increased frequency of extreme events, it is especially timely to investigate plasticity within Antarctic species responding to a key environmental aspect of their ecology: sea ice variability. Using 13 years of longitudinal data, we investigated the effect of sea ice concentration (SIC) on the foraging efficiency of Adélie penguins (Pygoscelis adeliae) breeding in the Ross Sea. A ‘natural experiment’ brought by the exceptional presence of giant icebergs during 5 consecutive years provided unprecedented habitat variation for testing the effects of extreme events on the relationship between SIC and foraging efficiency in this sea-ice dependent species. Significant levels of phenotypic plasticity were evident in response to changes in SIC in normal environmental conditions. Maximum foraging efficiency occurred at relatively low SIC, peaking at 6.1% and decreasing with higher SIC. The ‘natural experiment’ uncoupled efficiency levels from SIC variations. Our study suggests that lower summer SIC than currently observed would benefit the foraging performance of Adélie penguins in their southernmost breeding area. Importantly, it also provides evidence that extreme climatic events can disrupt response plasticity in a wild seabird population. This questions the predictive power of relationships built on past observations, when not only the average climatic conditions are changing but the frequency of extreme climatic anomalies is also on the rise.     

Effect of climatic warming on the Pacific walrus, and potential modification of its helminth fauna

The decreasing extent of sea-ice in the arctic basin as a consequence of climatic warming is modifying the behavior and diets of pagophilic pinnipeds, including the Pacific walrus, Odobenus rosmarus divergens Illiger, the species emphasized here. Mammals such as the walrus and bearded seal, Erignathus barbatus (Erxleben), cannot remain associated with the sea-ice, and continue to feed on their usual diet of benthic invertebrates inhabiting coastal waters to a depth of approximately 100 m, when the northwestward retreating ice reaches deep waters beyond the margins of the continental shelf. With reduction of their customary substrate (ice), the walrus has become more pelagic and preys more often on ringed seals, Phoca hispida Schreber. Dietary changes, with modifications of helminth faunas, may be induced by various factors. Increased consumption of mammals or their remains by walruses may lead to a higher prevalence of trichinellosis in them and to more frequent occurrence in indigenous peoples inhabiting the arctic coasts. To assess predicted effects on the composition of helminth fauna of the walrus, we recommend systematic surveys of their helminths as part of research on effects of climatic warming.     

‘Manipulation’ without the parasite: altered feeding behaviour of mosquitoes is not dependent on infection with malaria parasites

Previous studies have suggested that Plasmodium parasites can manipulate mosquito feeding behaviours such as probing, persistence and engorgement rate in order to enhance transmission success. Here, we broaden analysis of this ‘manipulation phenotype’ to consider proximate foraging behaviours, including responsiveness to host odours and host location. Using Anopheles stephensi and Plasmodium yoelii as a model system, we demonstrate that mosquitoes with early stage infections (i.e. non-infectious oocysts) exhibit reduced attraction to a human host, whereas those with late-stage infections (i.e. infectious sporozoites) exhibit increased attraction. These stage-specific changes in behaviour were paralleled by changes in the responsiveness of mosquito odourant receptors, providing a possible neurophysiological mechanism for the responses. However, we also found that both the behavioural and neurophysiological changes could be generated by immune challenge with heat-killed Escherichia coli and were thus not tied explicitly to the presence of malaria parasites. Our results support the hypothesis that the feeding behaviour of female mosquitoes is altered by Plasmodium, but question the extent to which this is owing to active manipulation by malaria parasites of host behaviour.     

Activity profiles and hook-tool use of New Caledonian crows recorded by bird-borne video cameras

New Caledonian crows are renowned for their unusually sophisticated tool behaviour. Despite decades of fieldwork, however, very little is known about how they make and use their foraging tools in the wild, which is largely owing to the difficulties in observing these shy forest birds. To obtain first estimates of activity budgets, as well as close-up observations of tool-assisted foraging, we equipped 19 wild crows with self-developed miniature video cameras, yielding more than 10 h of analysable video footage for 10 subjects. While only four crows used tools during recording sessions, they did so extensively: across all 10 birds, we conservatively estimate that tool-related behaviour occurred in 3% of total observation time, and accounted for 19% of all foraging behaviour. Our video-loggers provided first footage of crows manufacturing, and using, one of their most complex tool types—hooked stick tools—under completely natural foraging conditions. We recorded manufacture from live branches of paperbark (Melaleuca sp.) and another tree species (thought to be Acacia spirorbis), and deployment of tools in a range of contexts, including on the forest floor. Taken together, our video recordings reveal an ‘expanded’ foraging niche for hooked stick tools, and highlight more generally how crows routinely switch between tool- and bill-assisted foraging.     

Drosophila male sex peptide inhibits siesta sleep and promotes locomotor activity in the post-mated female

Quiescence, or a sleep-like state, is a common and important feature of the daily lives of animals from both invertebrate and vertebrate taxa, suggesting that sleep appeared early in animal evolution. Recently, Drosophila melanogaster has been shown to be a relevant and powerful model for the genetic analysis of sleep behaviour. The sleep architecture of D. melanogaster is sexually dimorphic, with females sleeping much less than males during day-time, presumably because reproductive success requires greater foraging activity by the female as well as the search for egg-laying sites. However, this loss of sleep and increase in locomotor activity will heighten the risk for the female from environmental and predator hazards. In this study, we show that virgin females can minimize this risk by behaving like males, with an extended afternoon ‘siesta’. Copulation results in the female losing 70 per cent of day-time sleep and becoming more active. This behaviour lasts for at least 8 days after copulation and is abolished if the mating males lack sex peptide (SP), normally present in the seminal fluid. Our results suggest that SP is the molecular switch that promotes wakefulness in the post-mated female, a change of behaviour compatible with increased foraging and egg-laying activity. The stress resulting from SP-dependent sleep deprivation might be an important contribution to the toxic side-effects of male accessory gland products that are known to reduce lifespan in post-mated females.     

Global warming and arctic terns: Estimating climate change impacts on the world's longest migration

Climate change is one of the top three global threats to seabirds, particularly species that visit polar regions. Arctic terns migrate between both polar regions annually and rely on productive marine areas to forage, on sea ice for rest and foraging, and prevailing winds during flight. Here, we report 21st-century trends in environmental variables affecting arctic terns at key locations along their Atlantic/Indian Ocean migratory flyway during the non-breeding seasons, identified through tracking data. End-of-century climate change projections were derived from Earth System Models and multi-model means calculated in two Shared Socioeconomic Pathways: ‘middle-of-the-road’ and ‘fossil-fuelled development’ scenarios. Declines in North Atlantic primary production emerge as a major impact to arctic terns likely to affect their foraging during the 21st century under a ‘fossil-fuelled development’ scenario. Minimal changes are, however, projected at three other key regions visited by arctic terns (Benguela Upwelling, Subantarctic Indian Ocean and the Southern Ocean). Southern Ocean sea ice extent is likely to decline, but the magnitude of change and potential impacts on tern survival are uncertain. Small changes (<1 m s⁻¹) in winds are projected in both scenarios, but with minimal likely impacts on migration routes and duration. However, Southern Ocean westerlies are likely to strengthen and contract closer to the continent, which may require arctic terns to shift routes or flight strategies. Overall, we find minor effects of climate change on the migration of arctic terns, with the exception of poorer foraging in the North Atlantic. However, given that arctic terns travel over huge spatial scales and live for decades, they integrate minor changes in conditions along their migration routes such that the sum effect may be greater than the parts. Meeting carbon emission targets is vital to slow these end-of-century climatic changes and minimise extinction risk for a suite of polar species.     

Longevity record of arctic skua (Stercorarius parasiticus)

The arctic skua (Stercorarius parasiticus) is one of the most long‐lived bird species. In 2010, we captured in Finland an adult, female arctic skua which had been ringed as a nestling in 1987. We tagged it also with a color ring. The bird has last been seen in July 2020 at the age of 33 years, making it most likely the oldest known arctic skua of the world. In 2010–2011 the bird carried a light‐level measuring geolocator, the data of which revealed that the bird had spent the nonbreeding season in the Canary Current area on the west coast of Africa. Breeding populations of arctic skuas have declined recently especially in British Isles, thus it is useful to get longevity data of this species with a high breeding site fidelity.     

Global change effects on the long‐term feeding ecology and contaminant exposures of East Greenland polar bears

Rapid climate changes are occurring in the Arctic, with substantial repercussions for arctic ecosystems. It is challenging to assess ecosystem changes in remote polar environments, but one successful approach has entailed monitoring the diets of upper trophic level consumers. Quantitative fatty acid signature analysis (QFASA) and fatty acid carbon isotope (δ¹³C‐FA) patterns were used to assess diets of East Greenland (EG) polar bears (Ursus maritimus) (n = 310) over the past three decades. QFASA‐generated diet estimates indicated that, on average, EG bears mainly consumed arctic ringed seals (47.5 ± 2.1%), migratory subarctic harp (30.6 ± 1.5%) and hooded (16.7 ± 1.3%) seals and rarely, if ever, consumed bearded seals, narwhals or walruses. Ringed seal consumption declined by 14%/decade over 28 years (90.1 ± 2.5% in 1984 to 33.9 ± 11.1% in 2011). Hooded seal consumption increased by 9.5%/decade (0.0 ± 0.0% in 1984 to 25.9 ± 9.1% in 2011). This increase may include harp seal, since hooded and harp seal FA signatures were not as well differentiated relative to other prey species. Declining δ¹³C‐FA ratios supported shifts from more nearshore/benthic/ice‐associated prey to more offshore/pelagic/open‐water‐associated prey, consistent with diet estimates. Increased hooded seal and decreased ringed seal consumption occurred during years when the North Atlantic Oscillation (NAO) was lower. Thus, periods with warmer temperatures and less sea ice were associated with more subarctic and less arctic seal species consumption. These changes in the relative abundance, accessibility, or distribution of arctic and subarctic marine mammals may have health consequences for EG polar bears. For example, the diet change resulted in consistently slower temporal declines in adipose levels of legacy persistent organic pollutants, as the subarctic seals have higher contaminant burdens than arctic seals. Overall, considerable changes are occurring in the EG marine ecosystem, with consequences for contaminant dynamics.     

Arctic warming: nonlinear impacts of sea‐ice and glacier melt on seabird foraging

Arctic climate change has profound impacts on the cryosphere, notably via shrinking sea‐ice cover and retreating glaciers, and it is essential to evaluate and forecast the ecological consequences of such changes. We studied zooplankton‐feeding little auks (Alle alle), a key sentinel species of the Arctic, at their northernmost breeding site in Franz‐Josef Land (80°N), Russian Arctic. We tested the hypothesis that little auks still benefit from pristine arctic environmental conditions in this remote area. To this end, we analysed remote sensing data on sea‐ice and coastal glacier dynamics collected in our study area across 1979–2013. Further, we recorded little auk foraging behaviour using miniature electronic tags attached to the birds in the summer of 2013, and compared it with similar data collected at three localities across the Atlantic Arctic. We also compared current and historical data on Franz‐Josef Land little auk diet, morphometrics and chick growth curves. Our analyses reveal that summer sea‐ice retreated markedly during the last decade, leaving the Franz‐Josef Land archipelago virtually sea‐ice free each summer since 2005. This had a profound impact on little auk foraging, which lost their sea‐ice‐associated prey. Concomitantly, large coastal glaciers retreated rapidly, releasing large volumes of melt water. Zooplankton is stunned by cold and osmotic shock at the boundary between glacier melt and coastal waters, creating new foraging hotspots for little auks. Birds therefore switched from foraging at distant ice‐edge localities, to highly profitable feeding at glacier melt‐water fronts within <5 km of their breeding site. Through this behavioural plasticity, little auks maintained their chick growth rates, but showed a 4% decrease in adult body mass. Our study demonstrates that arctic cryosphere changes may have antagonistic ecological consequences on coastal trophic flow. Such nonlinear responses complicate modelling exercises of current and future polar ecosystem dynamics.     

Sea-ice use by arctic foxes in northern Alaska

The extensive use of sea-ice by three arctic foxes (Alopex lagopus) in northern Alaska was documented using satellite telemetry during the winter of 2005–2006. Here we present the first detailed data on movements of individual foxes while on the sea-ice. Two juvenile males and one juvenile female traveled long distances (904, 1,096, and 2,757 km) and remained on the sea-ice for extended periods of time (76, 120, and 156 days). Average distances traveled per day ranged from 7.5 to 17.6 km and foxes achieved maximum rates of travel of up to 61 km/day. These findings verify the use of sea-ice by arctic foxes and raise concerns that the diminishing arctic ice cover may negatively impact populations by limiting access to marine food sources.     

Polar bear predatory behaviour reveals seascape distribution of ringed seal lairs

Ringed seal (Pusa hispida) breeding distribution has been extensively studied across near-shore habitats, but has received limited attention at a seascape scale due to the difficulty in accessing offshore sea ice environments. Employing highly visible predation attempts by polar bears (Ursus maritimus) on ringed seals in subnivean lairs observed by helicopter, the spatial relationship between predatory behaviour and ringed seal breeding habitat was examined. Resource selection functions were used to determine the relative probability of predation attempts on ringed seals in lairs as a function of habitat during a period of low ringed seal natality (2004–2006). Ringed seal pup kill locations were compared between years of low (2003–2006) and high (2007–2011) natality to assess the effect of reproductive output on habitat use. During years of low natality, polar bear hunting attempts were more likely in near-shore fast ice, and pup kills were observed predominately in fast ice (fast = 65 %, pack = 29 %, P = 0.002) at a median distance of 36 km from shore. In years of high natality, pup kills were observed farther from shore (median = 46 km, P = 0.03), and there was no difference in the proportion of observations in fast ice and pack ice (fast = 43 %, pack = 52 %, P = 0.29). These results suggest that the facultative use of adjacent offshore pack ice by breeding ringed seals may be influenced by natality. This study illustrates how documenting the behaviour of a predator can facilitate insight into the distribution of a cryptic prey.     

Avian scavengers and the threat from veterinary pharmaceuticals

Veterinary use of the non-steroidal anti-inflammatory drug diclofenac on domesticated ungulates caused populations of resident Gyps vultures in the Indian sub-continent to collapse. The birds died when they fed on carrion from treated animals. Veterinary diclofenac was banned in 2006 and meloxicam was advocated as a ‘vulture-safe’ alternative. We examine the effectiveness of the 2006 ban, whether meloxicam has replaced diclofenac, and the impact of these changes on vultures. Drug residue data from liver samples collected from ungulate carcasses in India since 2004 demonstrate that the prevalence of diclofenac in carcasses in 2009 was half of that before the ban and meloxicam prevalence increased by 44%. The expected vulture death rate from diclofenac per meal in 2009 was one-third of that before the ban. Surveys at veterinary clinics show that diclofenac use in India began in 1994, coinciding with the onset of rapid Gyps declines ascertained from measured rates of declines. Our study shows that one pharmaceutical product has had a devastating impact on Asia''s vultures. Large-scale research and survey were needed to detect, diagnose and quantify the problem and measure the response to remedial actions. Given these difficulties, other effects of pharmaceuticals in the environment may remain undetected.     

Alternative mating tactics in the yellow dung fly: resolving mechanisms of small-male advantage off pasture

Recent work suggests that the yellow dung fly mating system may include alternative patroller–competitor mating tactics in which large males compete for gravid females on dung, whereas small, non-competitive males search for females at foraging sites. Small males obtain most matings off pasture, yet the behavioural mechanism(s) giving rise to this pattern are unknown. We investigated the male and female behaviours that determine mating success in this environment by conducting field mating experiments and found small males to benefit from several attributes specific to the off-pasture mating environment. First, small males from foraging sites exhibited higher mating propensity, indicating that large males away from dung may be depleted of energy and/or sperm. Second, small males were more discriminating, being significantly less likely to attempt with non-gravid females, which are absent on dung but common off pasture. Third, non-gravid females were generally more likely to actively struggle and reject mating attempts; however, such behaviours occurred disproportionately more often with large males. Female Scathophaga stercoraria thus appear to preferentially mate with small males when off pasture. These findings challenge assumptions about male–female interactions in systems with alternative mating tactics and reveal hidden processes that may influence selection patterns in the field.     

Does coping style predict optimization? An experimental test in a wild passerine bird

A number of studies have suggested that avian brood size is individually optimized. Yet, optimal reproductive decisions likely vary owing to among-individual differences in environmental sensitivity. Specifically, ‘proactive’ individuals who do not track environmental changes may be less able to produce optimal brood sizes than ‘reactive’ individuals who have more precise local environmental knowledge. To test this, we quantified exploratory behaviour (a proxy for proactivity) in a great tit (Parus major) population, manipulated brood sizes (reduced, control, enlarged) and evaluated whether individuals of dissimilar coping style differed in their level of optimization. If reactive females behaved optimally, any deviation from their original brood size should lower fitness, whereas this should not be the case for proactive females. Reactive females indeed performed best at their natural brood size, whereas proactive females performed best when raising an enlarged brood. These findings imply that proactive females produced sub-optimal brood sizes. We speculate that proactive females might (i) take decisions based on biased perception of their environment, (ii) face energetic constraints in offspring production and/or (iii) be more willing to invest into current reproduction when given the option. Our findings provide experimental evidence for coping style-related differences in optimal reproductive decisions and life-history strategies.     

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter‐specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic‐feeding ice‐associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983–1999 and 2000–2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic‐level productivity in these marine ecosystems.     

Evaluation of Dry Ice as a Potential Cryonematicide for Meloidogyne incognita in Soil

Solid CO₂ (dry ice) was added to pots containing soil that was infested either with eggs of the root-knot nematode, Meloidogyne incognita, or with tomato (Lycopersicon esculentum ''Rutgers'') root fragments that were infected with various stages of the nematode. Two hours after dry ice was added, thermocouples in the soil recorded temperatures ranging from -15 °C to -59 °C. One day after treatment with the dry ice, the temperature of the soil was allowed to equilibrate with that of the greenhouse, and susceptible tomato seedlings were planted in pots containing infested soil treated or untreated (controls) with dry ice. After 5 weeks, roots were removed from the pots and nematode eggs were extracted and counted. Plants grown in soil infested with eggs and receiving dry ice treatment had less than 1% of the eggs found in the controls; plants from soil infested with root fragments and receiving dry ice treatment had less than 4% of the eggs found in controls. Dry ice used to lower soil temperature may have potential as a cryonematicide.     

Bearded seal (Erignathus barbatus) birth mass and pup growth in periods with contrasting ice conditions in Svalbard,Norway

Global climate warming has caused major reductions in sea ice in the Arctic, posing a serious threat to ice-associated marine mammals. Herein, novel data on birth mass and pup growth rates over a 15-year period (1993–2007; 10 years with growth data) are reported for bearded seals, as well as initial behavioral responses by this species to major, local declines in sea-ice in Svalbard, Norway. In total, 205 pups were captured; 64 of which were recaptured, some repeatedly, producing 85 growth intervals for nursing pups. Average birth mass of pups was 37.1 ± 3.8 (SD) kg (range 33–47 kg, n = 25); birth mass before vs. after the sea ice collapse (2006 onward) were not found to differ. Pups grew at a rate of 3.0 ± 0.7 kg/day (1.8–4.8 kg/day, n = 64) during the nursing period. LME models suggest that ice concentration did not affect the growth rate of pups. Most females shifted from traditional first-year ice floes to glacier-ice pieces for birthing and nursing their young, following the regional sea ice collapse. However, retraction of tidal glaciers will likely eliminate this replacement birthing and nursing habitat for bearded seals in Svalbard in the coming decades.     

An island-wide predator manipulation reveals immediate and long-lasting matching of risk by prey

Anti-predator behaviour affects prey population dynamics, mediates cascading effects in food webs and influences the likelihood of rapid extinctions. Predator manipulations in natural settings provide a rare opportunity to understand how prey anti-predator behaviour is affected by large-scale changes in predators. Here, we couple a long-term, island-wide manipulation of an important rodent predator, the island fox (Urocyon littoralis), with nearly 6 years of measurements on foraging by deer mice (Peromyscus maniculatus) to provide unequivocal evidence that prey closely match their foraging behaviour to the number of fox predators present on the island. Peromyscus maniculatus foraging among exposed and sheltered microhabitats (a measure of aversion to predation risk) closely tracked fox density, but the nature of this effect depended upon nightly environmental conditions known to affect rodent susceptibility to predators. These effects could not be explained by changes in density of deer mice over time. Our work reveals that prey in natural settings are cognizant of the dynamic nature of their predators over timescales that span many years, and that predator removals spanning many generations of prey do not result in a loss of anti-predator behaviour.