The importance of marine vs. human-induced subsidies in the maintenance of an expanding mesocarnivore in the arctic tundra

1. Most studies addressing the causes of the recent increases and expansions of mesopredators in many ecosystems have focused on the top-down, releasing effect of extinctions of large apex predators. However, in the case of the northward expansion of the red fox into the arctic tundra, a bottom-up effect of increased resource availability has been proposed, an effect that can counteract prey shortage in the low phase of the multi-annual rodent cycle. Resource subsidies both with marine and with terrestrial origins could potentially be involved. 2. During different phases of a multi-annual rodent cycle, we investigated the seasonal dynamics and spatial pattern of resource use by red foxes across a coast to inland low arctic tundra gradient, Varanger Peninsula, Norway. We employed two complementary methods of diet analyses: stomach contents and stable isotope analysis. 3. We found that inland red foxes primarily subsisted on reindeer carrions during the low phase of a small rodent population cycle. Lemmings became the most important food item towards the peak phase of the rodent cycle, despite being less abundant than sympatric voles. Isotopic signatures of tissue from both predator and prey also revealed that red foxes near the coast used marine-derived subsidies in the winter, but these allochthonous resources did not spillover to adult foxes living beyond 20-25 km from the coast. 4. Although more needs to be learned about the link between increasing primary productivity due to climatic warming and trophic dynamics in tundra ecosystems, we suggest that changes in reindeer management through a bottom-up effect, at least regionally, may have paved the way towards the establishment of a new mesopredator in the tundra biome.     

An avian terrestrial predator of the Arctic relies on the marine ecosystem during winter

Top predators of the arctic tundra are facing a long period of very low prey availability during winter and subsidies from other ecosystems such as the marine environment may help to support their populations. Satellite tracking of snowy owls, a top predator of the tundra, revealed that most adult females breeding in the Canadian Arctic overwinter at high latitudes in the eastern Arctic and spend several weeks (up to 101 d) on the sea‐ice between December and April. Analysis of high‐resolution satellite images of sea‐ice indicated that owls were primarily gathering around open water patches in the ice, which are commonly used by wintering seabirds, a potential prey. Such extensive use of sea‐ice by a tundra predator considered a small mammal specialist was unexpected, and suggests that marine resources subsidize snowy owl populations in winter. As sea‐ice regimes in winter are expected to change over the next decades due to climate warming, this may affect the wintering strategy of this top predator and ultimately the functioning of the tundra ecosystem.     

Strength of asymmetric competition between predators in food webs ruled by fluctuating prey: the case of foxes in tundra

In food webs heavily influenced by multi‐annual population fluctuations of key herbivores, predator species may differ in their functional and numerical responses as well as their competitive ability. Focusing on red and arctic fox in tundra with cyclic populations of rodents as key prey, we develop a model to predict how population dynamics of a dominant and versatile predator (red fox) impacted long‐term growth rate of a subdominant and less versatile predator (arctic fox). We compare three realistic scenarios of red fox performance: (1) a numerical response scenario where red fox acted as a resident rodent specialist exhibiting population cycles lagging one year after the rodent cycle, (2) an aggregative response scenario where red fox shifted between tundra and a nearby ecosystem (i.e. boreal forest) so as to track rodent peaks in tundra without delay, and (3) a constant subsidy scenario in which the red fox population was stabilized at the same mean density as in the other two scenarios. For all three scenarios it is assumed that the arctic fox responded numerically as a rodent specialist and that the mechanisms of competition is of a interference type for space, in which the arctic fox is excluded from the most resource rich patches in tundra. Arctic fox is impacted most by the constant subsidy scenario and least by the numerical response scenario. The differential effects of the scenarios stemmed from cyclic phase‐dependent sensitivity to competition mediated by changes in temporal mean and variance of available prey to the subdominant predator. A general implication from our result is that external resource subsidies (prey or habitats), monopolized by the dominant competitor, can significantly reduce the likelihood for co‐existence within the predator guild. In terms of conservation of vulnerable arctic fox populations this means that the likelihood of extinction increases with increasing amount of subsidies (e.g. carcasses of large herbivores or marine resources) in tundra and nearby forest areas, since it will act to both increase and stabilize populations of red fox.     

The fluctuating world of a tundra predator guild: bottom‐up constraints overrule top‐down species interactions in winter

Global warming is predicted to change ecosystem functioning and structure in Arctic ecosystems by strengthening top‐down species interactions, i.e. predation pressure on small herbivores and interference between predators. Yet, previous research is biased towards the summer season. Due to greater abiotic constraints, Arctic ecosystem characteristics might be more pronounced in winter. Here we test the hypothesis that top‐down species interactions prevail over bottom‐up effects in Scandinavian mountain tundra (Northern Sweden) where effects of climate warming have been observed and top‐down interactions are expected to strengthen. But we test this ‘a priori’ hypothesis in winter and throughout the 3–4 yr rodent cycle, which imposes additional pulsed resource constraints. We used snowtracking data recorded in 12 winters (2004–2015) to analyse the spatial patterns of a tundra predator guild (arctic fox Vulpes lagopus, red fox Vulpes vulpes, wolverine Gulo gulo) and small prey (ptarmigan, Lagopus spp). The a priori top‐down hypothesis was then tested through structural equation modelling, for each phase of the rodent cycle. There was weak support for this hypothesis, with top‐down effects only discerned on arctic fox (weakly, by wolverine) and ptarmigan (by arctic fox) at intermediate and high rodent availability respectively. Overall, bottom‐up constraints appeared more influential on the winter community structure. Cold specialist predators (arctic fox and wolverine) showed variable landscape associations, while the boreal predator (red fox) appeared strongly dependent on productive habitats and ptarmigan abundance. Thus, we suggest that the unpredictability of food resources determines the winter ecology of the cold specialist predators, while the boreal predator relies on resource‐rich habitats. The constraints imposed by winters and temporary resource lows should therefore counteract productivity‐driven ecosystem change and have a stabilising effect on community structure. Hence, the interplay between summer and winter conditions should determine the rate of Arctic ecosystem change in the context of global warming.     

Changes in the Spatial Configuration and Strength of Trophic Control Across a Productivity Gradient During a Massive Rodent Outbreak

Understanding the determinants of spatial and temporal differences in the relative strength of consumer–resource interactions is an important endeavour in ecology. Here, we explore the necessary conditions for temporal shifts in the relative strength of rodent–plant interactions in an area characterised by profound spatial differences in trophic control, with predator–prey interactions prevailing in productive habitats and rodent–plant interactions dominating unproductive habitats of the forest–tundra ecotone. We report data obtained during the exceptionally massive rodent outbreak of 2010–2012 in northernmost Fennoscandia, including an experimental manipulation of herbivore access to vegetation plots across a large-scale productivity gradient, multiple observational measures of plant–rodent interactions linked to rodent abundance data and a large-scale survey of breeding avian predators and mammalian predator activity. Unexpectedly, rodent grazing impacts documented during the rodent outbreak were uniformly strong across the landscape, regardless of habitat productivity. The runaway response in rodent populations was facilitated by a high population growth rate in the early phase of the outbreak due to the extended absence of predators in productive habitats, concomitant with an exceptionally long-lasting lemming outbreak in unproductive habitats. Our results showed that spatio-temporal variation in trophic control also occurs in ecosystems structured according to the exploitation ecosystems hypothesis and emphasises the importance of long-term studies to capture nonlinear and stochastic features that shape ecosystem functioning. In this context, the temporary release from top–down regulation in productive habitats caused strong grazing impacts that may be crucial for the resilience of tundra ecosystems under the threat of climate change-driven shrub encroachment.     

The influence of coastal access on isotope variation in Icelandic arctic foxes

To quantify the ecological effects of predator populations, it is important to evaluate how population-level specializations are dictated by intra- versus inter-individual dietary variation. Coastal habitats contain prey from the terrestrial biome, the marine biome and prey confined to the coastal region. Such habitats have therefore been suggested to better support predator populations compared to habitats without coastal access. We used stable isotope data on a small generalist predator, the arctic fox, to infer dietary strategies between adult and juvenile individuals with and without coastal access on Iceland. Our results suggest that foxes in coastal habitats exhibited a broader isotope niche breadth compared to foxes in inland habitats. This broader niche was related to a greater diversity of individual strategies rather than to a uniform increase in individual niche breadth or by individuals retaining their specialization but increasing their niche differentiation. Juveniles in coastal habitats exhibited a narrower isotope niche breadth compared to both adults and juveniles in inland habitats, and juveniles in inland habitats inhabited a lower proportion of their total isotope niche compared to adults and juveniles from coastal habitats. Juveniles in both habitats exhibited lower intra-individual variation compared to adults. Based on these results, we suggest that foxes in both habitats were highly selective with respect to the resources they used to feed offspring, but that foxes in coastal habitats preferentially utilized marine resources for this purpose. We stress that coastal habitats should be regarded as high priority areas for conservation of generalist predators as they appear to offer a wide variety of dietary options that allow for greater flexibility in dietary strategies.     

How ecological neighbourhoods influence the structure of the scavenger guild in low arctic tundra

Aim How the ecological neighbourhoods of coast and forest affect arctic tundra ecosystems is a pressing question as the circumpolar tundra belt is shrinking under global warming. Mobile facultative scavengers are likely to negatively impact tundra biodiversity as dominant competitors or predators, if they spill over into tundra. Here, we provide the first quantitative assessments of the structure of a scavenger guild in low arctic tundra with emphasis on how it changes along spatial gradients from neighbouring ecosystems (i.e. forest and coast) and with altitude (i.e. productivity gradients). We also assess the likelihood of interactions between guild members that may negatively impact vulnerable tundra species. Location North‐eastern part of Norway. Methods Extensive records of scavenger prevalence were obtained by deploying automatic digital cameras at experimental carcasses in tundra regions covering several thousand square kilometres and three winters in northern Norway. Main conclusions We found short‐range neighbourhood effects of forest and coast within the tundra scavenger guild. Species richness declined steeply with decreasing distance from the neighbouring ecosystems, in particular subarctic forest, and with increasing altitude. Bird species with strongholds in forest (golden eagle Aquila chrysaetos and hooded crow Corvus cornix) or along the coast (white‐tailed eagle Haliaeetus albicilla) were mostly responsible for short‐range neighbourhood effects on guild structure. However, the two most abundant guild members, the common raven Corvus corax and the red fox Vulpes vulpes, exhibited no spatial patterns within the range of neighbourhoods and altitudes examined. There was a clear diurnal segregation in the use of carcasses between birds and mammals reducing the likelihood of direct interactions between these two taxa. Presence of red fox appeared to exclude the arctic fox Vulpes lagopus, the only endemic tundra species within the guild, from carcasses.     

Functional and numerical responses of four lemming predators in high arctic Greenland

The high‐arctic tundra ecosystem has the world's simplest vertebrate predator–prey community, with only four predators preying upon one rodent species, the collared lemming (Dicrostonyx groenlandicus). We document the functional and numerical responses of all the four predators in NE Greenland. Using these data, we assess the impact of predation on the dynamics of the collared lemming with a 4 yr cycle and >100‐fold difference between maximum and minimum densities. All predator species feed mostly (>90%) on lemmings when lemming density is >1 ha^?1, but the shapes of the predators’ responses vary greatly. The snowy owl (Nyctea scandiaca) is present and breeds only when lemming densities at snowmelt are >2 ha^?1, giving rise to a step‐like numerical response. The long‐tailed skua (Stercorarius longicaudus) has a type III functional response and shifts from alternate food (mainly berries and insects) to lemmings with increasing lemming density. The skua surpasses all the other predators in summer by its total response. The type III functional response of the Arctic fox (Alopex lagopus) starts to increase at much lower lemming densities than the responses of the avian predators, but it has only a weak numerical response. Finally, the stoat (Mustela erminea) is the most specialized predator and the only one with a clearly delayed numerical response. According to their specific functional and numerical responses, each predator plays a key role at some point of the lemming cycle, but only the stoat has the potential to drive the lemming cycle. Stoat predation is greatly reduced in the winter preceding the lemming peak, and it reaches a maximum in the winter preceding the lowest lemming summer density. Stoat predation appears to maintain low lemming densities for at least two successive years. Our study provides empirical support for the specialist predator hypothesis about small mammal population cycles.     

Marine subsidies alter the diet and abundance of insular and coastal lizard populations

We evaluated the impact of marine materials on the diet and abundance of arthropodivorous lizards inhabiting islands and the coast of the Gulf of California. Here, marine materials are brought onto land by seabirds and by tidal action, and both subsidy pathways cause arthropod abundance to increase. We evaluated Uta stansburiana (side-blotched lizard) diets in three habitats defined by having: (1) no marine subsidies available, (2) only seabird-derived subsidies, and (3) only tidal-derived subsidies. Stable isotope data indicated that lizard diets are subsidized indirectly by seabird and tidal activity. For example, in coastal areas we determined that 40% of a lizard's diet contains arthropods that have consumed algae. Such subsidies may explain why we found that lizards in coastal areas occupy a significantly higher trophic position than lizards in unsubsidized areas. We analyzed eight years of survey data on all arthropodivorous lizards to determine if diet subsidies result in increased lizard abundance. We found that lizards were more abundant in coastal areas than they were in inland habitats, and that they were more abundant on islands with seabirds than on islands without seabird populations. This study provides insight into the importance and effect of marine-derived nutrients from multiple sources on vertebrate consumers inhabiting islands and coastal areas.     

Indirect food web interactions affect predation of Tengmalm's Owls Aegolius funereus nests by Pine Martens Martes martes according to the alternative prey hypothesis

Although population cycles of rodents are geographically widespread and occur in a number of rodent species, higher‐order food web interactions mediated by predator–rodent dynamics have primarily been described from boreal and arctic biomes. During periods of low rodent abundance, predators may switch to alternative prey, which may affect other predators directly or indirectly. Using a long‐term dataset, we assessed the frequency of Pine Marten Martes martes predation on the nests of Tengmalm's Owl Aegolius funereus during periods of fluctuating rodent abundance in Central Europe. The number of nests predated by Pine Martens was positively correlated with the annual number of nests available. The probability of predation by Pine Martens on Tengmalm's Owl nests decreased with increasing spring abundance index of Apodemus mice, but was not related to the abundance index of Myodes and Microtus voles, pooled rodent abundance or age of the nestbox. Additionally, we found no relationship between the breeding frequency (i.e. the number of nesting attempts per nestboxes available) and an abundance index of Microtus and Myodes voles, Apodemus mice or overall rodent abundance. Our results demonstrate, for the first time in a temperate area, that during periods of low Apodemus mouse abundance, the switching response of an opportunistic mammalian predator can lead to indirect food web interactions through an increase in nest predation on a sympatric avian predator.     

Movement tactics of a mobile predator in a meta‐ecosystem with fluctuating resources: the arctic fox in the High Arctic

Animal movement is a fundamental process shaping ecosystems at multiple levels, from the fate of individuals to global patterns of biodiversity. The spatio‐temporal dynamic of food resources is a major driver of animal movement and generates patterns ranging from range residency to migration and nomadism. Arctic tundra predators face a strongly fluctuating environment marked by cyclic microtine populations, high seasonality, and the potential availability of sea ice, which gives access to marine resources in winter. This type of relatively poor and highly variable environment can promote long‐distance movements and resource tracking in mobile species. Here, we investigated the winter movements of the arctic fox, a major tundra predator often described as a seasonal migrant or nomad. We used six years of Argos satellite telemetry data collected on 66 adults from Bylot Island (Nunavut, Canada) tracked during the sea ice period. We hypothesized that long‐distance movements would be influenced by spatio‐temporal changes in resource availability and individual characteristics. Despite strong annual and seasonal changes in resource abundance and distribution, we found that a majority of individuals remained resident, especially those located in an area characterized by highly predictable pulse resources (goose nesting colony) and abundant cached food items (eggs). Foxes compensated terrestrial food shortage by commuting to the sea ice rather than using long‐distance tracking or moving completely onto the sea ice for winter. Individual characteristics also influenced movement patterns: age positively influenced the propensity to engage in nomadism, suggesting older foxes may be driven out of their territories. Our results show how these mammalian predators can adjust their movement patterns to favor range residency despite strong spatio‐temporal fluctuations in food resources. Understanding the movement responses of predators to prey dynamics helps identifying the scales at which they work, which is a critical aspect of the functioning and connectivity among meta‐ecosystems.     

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.     

Climate‐associated tundra thaw pond formation and range expansion of boreal zooplankton predators

Most of the freshwater component of the Earth's surface is composed of shallow tundra ponds. These high latitude ecosystems have been exposed to a variety of abiotic disturbances associated with recent environmental change. However, the biological significance of these changes remains poorly understood. Here, we characterize the abiotic disturbance to the shallow tundra ponds of northwest Alaska. We used historical aerial imagery to determine that up to 53% of the sampled ponds have formed during the recent warmer decades (since the 1970s). We discovered that two top predator species (phantom midges of the genus Chaoborus) of the freshwater zooplankton have recently undergone range expansion, forming widespread (a scale of hundreds of km) stable tundra populations. We assessed the population persistence of these boreal predators by longitudinal sampling over 14 yr. Recent thaw ponds had significantly dissimilar zooplankton communities to communities of ponds that formed before 1950. Both predator and herbivore species differed by age of pond. Younger pond ages and warmer surface temperatures were the significant predictors of the presence of temperate Chaoborus americanus in tundra ponds. Ponds containing tundra populations of C. americanus and C. cf. flavicans were associated with recent formation (83–90%). Recent ponds in river valleys appeared more important than recent ponds near roads as colonization corridors for C. americanus. Only 24% of the tundra keystone predator, Heterocope septentrionalis, populations were from recent ponds. Our results suggest that climate‐associated disturbance can lead to a widespread stable range expansion of boreal species despite the propinquity of older ponds with top‐down control exerted by an endemic keystone predator.     

Influence of pulsed resources and marine subsidies on insular rodent populations

Inputs of marine resources contribute to high consumer densities on small, barren islands in the arid Gulf of California, Mexico. Episodes of rainfall associated with El Niño Southern Oscillation (ENSO) events, however, stimulate plant growth and seed production and replenish soil detritus and seed banks. We studied the effects of marine inputs and pulsed ENSO resources on rodent populations on islands before, during and after the strong 1997–1998 ENSO event. Stable isotope analysis was used to determine rodent diets. During dry years, P. maniculatus consumed marine prey near shore and on small islands, and were much more abundant in these areas than farther inland and on larger islands. Captures were particularly concentrated near shore on three islands inhabited by Chaetodipus rudinoris, a granivore that was numerous in inland areas. Following the ENSO, Peromyscus maniculatus increased in abundance by 408%, compared to only 160% for C. rudinoris, and new individuals invaded inland areas. P. maniculatus populations declined sharply the following year, however, whereas C. rudinoris populations remained relatively stable. In response to ENSO conditions, P. maniculatus ate more seeds on islands with C. rudinoris than on islands without C. rudinoris. P. maniculatus also increased less markedly and declined more quickly in abundance where C. rudinoris was present. We speculate that P. maniculatus is capable of rapid numerical response to resource pulses but depends on marine foods during dry years. C. rudinoris is restricted to islands with sufficient seed resources, but maintains stable populations by exploiting low‐density seeds and caching seeds from wet years. On islands with both species, C. rudinoris restricts P. maniculatus to near‐shore areas via interference, but spillover consumption of seeds by subsidized P. maniculatus may limit the response of C. rudinoris to ENSO resources. Combined with differences in life history and dispersal abilities, these ecological factors help explain the distinct biogeographic distributions of these rodents on Gulf islands.     

Rough-Legged Buzzards,Arctic Foxes and Red Foxes in a Tundra Ecosystem without Rodents

Small rodents with multi-annual population cycles strongly influence the dynamics of food webs, and in particular predator-prey interactions, across most of the tundra biome. Rodents are however absent from some arctic islands, and studies on performance of arctic predators under such circumstances may be very instructive since rodent cycles have been predicted to collapse in a warming Arctic. Here we document for the first time how three normally rodent-dependent predator species—rough-legged buzzard, arctic fox and red fox – perform in a low-arctic ecosystem with no rodents. During six years (in 2006-2008 and 2011-2013) we studied diet and breeding performance of these predators in the rodent-free Kolguev Island in Arctic Russia. The rough-legged buzzards, previously known to be a small rodent specialist, have only during the last two decades become established on Kolguev Island. The buzzards successfully breed on the island at stable low density, but with high productivity based on goslings and willow ptarmigan as their main prey – altogether representing a novel ecological situation for this species. Breeding density of arctic fox varied from year to year, but with stable productivity based on mainly geese as prey. The density dynamic of the arctic fox appeared to be correlated with the date of spring arrival of the geese. Red foxes breed regularly on the island but in very low numbers that appear to have been unchanged over a long period – a situation that resemble what has been recently documented from Arctic America. Our study suggests that the three predators found breeding on Kolguev Island possess capacities for shifting to changing circumstances in low-arctic ecosystem as long as other small - medium sized terrestrial herbivores are present in good numbers.     

Feedback between size balance and consumption strongly affects the consequences of hatching phenology in size‐dependent predator–prey interactions

In many size‐dependent predator–prey systems, hatching phenology strongly affects predator–prey interaction outcomes. Early‐hatched predators can easily consume prey when they first interact because they encounter smaller prey. However, this process by itself may be insufficient to explain all predator–prey interaction outcomes over the whole interaction period because the predator–prey size balance changes dynamically throughout their ontogeny. We hypothesized that hatching phenology influences predator–prey interactions via a feedback mechanism between the predator–prey size balance and prey consumption by predators. We experimentally tested this hypothesis in an amphibian predator–prey model system. Frog tadpoles Rana pirica were exposed to a predatory salamander larva Hynobius retardatus that had hatched 5, 12, 19 or 26 days after the frog tadpoles hatched. We investigated how the salamander hatch timing affected the dynamics of prey mortality, size changes of both predator and prey, and their subsequent life history (larval period and size at metamorphosis). The predator–prey size balance favoured earlier hatched salamanders, which just after hatching could successfully consume more frog tadpoles than later hatched salamanders. The early‐hatched salamanders grew rapidly and their accelerated growth enabled them to maintain the predator‐superior size balance; thus, they continued to exert strong predation pressure on the frog tadpoles in the subsequent period. Furthermore, frog tadpoles exposed to the early‐hatched salamanders were larger at metamorphosis and had a longer larval period than other frog tadpoles. These results suggest that feedback between the predator‐superior size balance and prey consumption is a critical mechanism that strongly affects the impacts of early hatching of predators in the short‐term population dynamics and life history of the prey. Because consumption of large nutrient‐rich prey items supports the growth of predators, a similar feedback mechanism may be common and have strong impacts on phenological shifts in size‐dependent trophic relationships.     

Representation of tundra vegetation by pollen in lake sediments of northern Alaska

Aim To understand better the representation of arctic tundra vegetation by pollen data, we analysed pollen assemblages and pollen accumulation rates (PARs) in the surface sediments of lakes. Location Modern sediment samples were collected from seventy‐eight lakes located in the Arctic Foothills and Arctic Coastal Plain regions of northern Alaska. Methods For seventy of the lakes, we analysed pollen and spores in the upper 2 cm of the sediment and calculated the relative abundance of each taxon (pollen percentages). For eleven of the lakes, we used ²¹⁰Pb analysis to determine sediment accumulation rates, and analysed pollen in the upper 10–15 cm of the sediment to estimate modern PARs. Using a detailed land‐cover map of northern Alaska, we assigned each study site to one of five tundra types: moist dwarf‐shrub tussock‐graminoid tundra (DST), moist graminoid prostrate‐shrub tundra (PST) (coastal and inland types), low‐shrub tundra (LST) and wet graminoid tundra (WGT). Results Mapped pollen percentages and multivariate comparison of the pollen data using discriminant analysis show that pollen assemblages vary along the main north–south vegetational and climatic gradients. On the Arctic Coastal Plain where climate is cold and dry, graminoid‐dominated PST and WGT sites were characterized by high percentages of Cyperaceae and Poaceae pollen. In the Arctic Foothills where climate is warmer and wetter, shrub‐dominated DST, PST and LST were characterized by high percentages of Alnus and Betula pollen. Small‐scale variations in tundra vegetation related to edaphic variability are also represented by the pollen data. Discriminant analysis demonstrated that DST sites could be distinguished from foothills PST sites based on their higher percentages of Ericales and Rubus chamaemorus pollen, and coastal PST sites could be distinguished from WGT sites based on their higher percentages of Artemisia. PARs appear to reflect variations in overall vegetation cover, although the small number of samples limits our understanding of these patterns. For coastal sites, PARs were higher for PST than WGT, whereas in the Arctic Foothills, PARs were highest in LST, intermediate in DST, and lowest in PST. Main conclusion Modern pollen data from northern Alaska reflect patterns of tundra vegetation related to both regional‐scale climatic gradients and landscape‐scale edaphic heterogeneity.     

Top–down control by great blue herons Ardea herodias regulates seagrass‐associated epifauna

Top predators can influence the structure and function of plant and animal communities. In coastal marine systems, fish, shark and mammal population declines are major drivers of recent ecosystem‐level change. Cascading effects of predatory wading birds, however, are less understood, even though wading bird populations have declined in many regions. We quantified the effects of predation by the piscivorous great blue heron Ardea herodias fannini on fish, invertebrates and epiphytes living in eelgrass Zostera marina. We found that herons forage on benthic fish in seagrass meadows, and foraging intensity increased from late spring until midsummer. When we experimentally excluded herons, benthic fish abundance increased, and the invertebrate assemblage shifted to more shrimp‐dominated assemblages while grazing gammarid amphipod abundance declined. These shifts were associated with reduced epiphyte abundance when herons were excluded, reflecting a four‐level trophic cascade and mediated by shifts in the grazer assemblage. In summary, we found that a piscivorous wading bird species exerts top down control in a subtidal seagrass ecosystem. Losses and recovery of wading birds could have ecosystem‐level ecological consequences that may need to be considered in the context of concern for overfishing and predator recovery in marine coastal management.     

Local genetic adaptation generates latitude‐specific effects of warming on predator–prey interactions

Temperature effects on predator–prey interactions are fundamental to better understand the effects of global warming. Previous studies never considered local adaptation of both predators and prey at different latitudes, and ignored the novel population combinations of the same predator–prey species system that may arise because of northward dispersal. We set up a common garden warming experiment to study predator–prey interactions between Ischnura elegans damselfly predators and Daphnia magna zooplankton prey from three source latitudes spanning >1500 km. Damselfly foraging rates showed thermal plasticity and strong latitudinal differences consistent with adaptation to local time constraints. Relative survival was higher at 24 °C than at 20 °C in southern Daphnia and higher at 20 °C than at 24 °C, in northern Daphnia indicating local thermal adaptation of the Daphnia prey. Yet, this thermal advantage disappeared when they were confronted with the damselfly predators of the same latitude, reflecting also a signal of local thermal adaptation in the damselfly predators. Our results further suggest the invasion success of northward moving predators as well as prey to be latitude‐specific. We advocate the novel common garden experimental approach using predators and prey obtained from natural temperature gradients spanning the predicted temperature increase in the northern populations as a powerful approach to gain mechanistic insights into how community modules will be affected by global warming. It can be used as a space‐for‐time substitution to inform how predator–prey interaction may gradually evolve to long‐term warming.     

Interactions of exotic and native carnivores in an ecotone, the coast of the Beagle Channel, Argentina

In coasts bordering highly productive seas, there can be a flux of resources to the terrestrial ecosystem, and terrestrial carnivores can use marine prey extensively. Two native, endangered species (otter Lontra provocax and culpeo fox Pseudalopex culpaeus lycoides) and two exotic species (mink Neovison vison and grey fox Pseudalopex griseus) inhabit the Beagle coast. The objectives of this paper are: (1) to describe the diet, habitat use and distribution of otters, mink and foxes on the coast of the Beagle channel and (2) to discuss the role of marine resources in the ecological interactions among these species. Diet was determined from the analysis of 245 faeces, and distribution was established from sign surveys. Marine prey occurred in the scats of 98.3, 70.4, 35.5 and 18.2% of otters, mink, culpeo and grey foxes, respectively. Other terrestrial species also use marine resources in Southern Patagonia. All this evidence suggests that the Fueguian coastal channels provide an illuminating example of allochthonous food subsidies from the sea. In the community of four sympatric predators, two native (and endangered) and two exotic, coexistence appears to be facilitated by a renewing marine food subsidy.