Numerical and Thermal Response of the Bacterivorous Ciliate Colpidium kleini, a Species Potentially at Risk of Extinction by Rising Water Temperatures.

We investigated the food-dependent growth and thermal response of the freshwater ciliate Colpidium kleini using numerical response (NR) experiments. This bacterivorous ciliate occurs in lotic water and the pelagial of lakes and ponds. The C. kleini strain used in this work was isolated from a small alpine lake and identified by combining detailed morphological inspections with molecular phylogeny. Specific growth rates (rmax) were measured from 5 to 21 °C. The ciliate did not survive at 22 °C. The threshold bacterial food levels (0.3 - 2.2 × 106 bacterial cells mL-1) matched the bacterial abundance in the alpine lake from which C. kleini was isolated. The food threshold was notably lower than previously reported for C. kleini and two other Colpidium species. The threshold was similar to levels reported for oligotrich and choreotrich ciliates if expressed in terms of bacterial biomass (0.05 - 0.43 mg C L-1). From the NR results, we calculated physiological mortality rates at zero food concentration. The mean mortality (0.55 ± 0.17 d-1) of C. kleini was close to the mean estimate obtained for other planktonic ciliates that do not encyst. We used the data obtained by the NR experiments to fit a thermal performance curve (TPC). The TPC yielded a temperature optimum at 17.3 °C for C. kleini, a maximum upper thermal tolerance limit of 21.9 °C, and a thermal safety margin of 4.6 °C. We demonstrated that combining NR with TPC analysis is a powerful tool to predict better a species' fitness in response to temperature and food.

Filial cannibalism of Nabis pseudoferus is not evolutionarily optimal foraging strategy.

Using a recursion model with real parameters of Nabis pseudoferus, we show that its filial cannibalism is an optimal foraging strategy for life reproductive success, but it is not an evolutionarily optimal foraging strategy, since it cannot maximize the descendant's number at the end of the reproductive season. Cannibalism is evolutionarily rational, when the number of newborn offspring produced from the cannibalized offspring can compensate the following two effects: (a) The cannibalistic lineage wastes time, since the individuals hatched from eggs produced by cannibalism start to reproduce later. (b) Cannibalism eliminates not only one offspring, but also all potential descendants from the cannibalized offspring during the rest of reproductive season. In our laboratory trials, from conspecific prey Nabis pseudoferus did not produce newborn nymphs enough to compensate the above two effects.

Pollen provisioning attenuates pesticide side-effects on a phytoseiid predator.

BACKGROUND: Biological control with predatory mites is applied against pests in greenhouse crops. Chemical control with the use of selective, reduced-risk pesticides, is an important component of Integrated Pest Management (IPM) programs, that often needs to be combined with biological control. Here, we evaluated the effect of plant pollen when used as supplementary food on the survival, reproduction and predation of the predatory mite Amblydromalus limonicus (Acari: Phytoseiidae) after exposing young larvae and adults to flonicamid, an insecticide of moderate toxicity to phytoseiids. Pollen is an important alternative food for generalist phytoseiids ensuring survival and supporting populations build-up during periods of prey scarcity. Two regimes of cattail (Typha angustifolia L.) pollen differing in application frequency were used. In the first, the total amount of pollen was supplied once, within 30 min after insecticide application, whereas in the second regime, the same amount of pollen was supplied gradually, i.e., every 48 h. RESULTS: Regardless of the frequency of application, pollen provisioning results in a reduction in prey (thrips) consumption relative to the control (no pollen provisioning). Nevertheless, when adult mites were directly exposed to flonicamid residues, pollen provisioning attenuated the reduction in prey consumption as compared to the control. In addition, the gradual (every 48 h) provisioning of pollen to adult predators exposed to flonicamid residues impacted positively the intrinsic rate of population increase (rm) of A. limonicus as compared to when feeding on prey. CONCLUSION: Our results reveal an unexpected role of pollen provisioning in alleviating pesticides side-effects on phytoseiids. © 2024 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Predator home range size mediates indirect interactions between prey species in an arctic vertebrate community.

Indirect interactions are widespread among prey species that share a common predator, but the underlying mechanisms driving these interactions are often unclear, and our ability to predict their outcome is limited. Changes in behavioural traits that impact predator space use could be a key proximal mechanism mediating indirect interactions, but there is little empirical evidence of the causes and consequences of such behavioural-numerical response in multispecies systems. Here, we investigate the complex ecological relationships between seven prey species sharing a common predator. We used a path analysis approach on a comprehensive 9-year data set simultaneously tracking predator space use, prey densities and prey mortality rate on key species of a simplified Arctic food web. We show that high availability of a clumped and spatially predictable prey (goose eggs) leads to a twofold reduction in predator (arctic fox) home range size, which increases local predator density and strongly decreases nest survival of an incidental prey (American golden plover). On the contrary, a scattered cyclic prey with potentially lower spatial predictability (lemming) had a weaker effect on fox space use and an overall positive impact on the survival of incidental prey. These contrasting effects underline the importance of studying behavioural responses of predators in multiprey systems and to explicitly integrate behavioural-numerical responses in multispecies predator-prey models.

Predator-mediated interactions through changes in predator home range size can lead to local prey exclusion.

The strength of indirect biotic interactions is difficult to quantify in the wild and can alter community composition. To investigate whether the presence of a prey species affects the population growth rate of another prey species, we quantified predator-mediated interaction strength using a multi-prey mechanistic model of predation and a population matrix model. Models were parametrized using behavioural, demographic and experimental data from a vertebrate community that includes the arctic fox (Vulpes lagopus), a predator feeding on lemmings and eggs of various species such as sandpipers and geese. We show that the positive effects of the goose colony on sandpiper nesting success (due to reduction of search time for sandpiper nests) were outweighed by the negative effect of an increase in fox density. The fox numerical response was driven by changes in home range size. As a result, the net interaction from the presence of geese was negative and could lead to local exclusion of sandpipers. Our study provides a rare empirically based model that integrates mechanistic multi-species functional responses and behavioural processes underlying the predator numerical response. This is an important step forward in our ability to quantify the consequences of predation for community structure and dynamics.

Functional ecology of planktonic ciliates: Measuring mortality rates in response to starvation.

Population dynamics of aquatic ciliates are controlled "bottom-up" via food supply and "top-down" by grazing and parasitism. While intrinsic growth rates of ciliates under saturating food conditions have been studied in some detail, mortality rates induced by starvation have received little attention thus far. To this end, we examined the response of three algivorous freshwater ciliate species to starvation using three different optical methods. Two of these methods, i.e. ciliate mortality rates (δ) estimated from (i) numerical response experiments and (ii) the rate of decline (ROD) in cell numbers, investigated the response of the ciliate population using conventional light microscopy. The third method, imaging cytometry using a FlowCAM instrument, monitored single cells during the starvation experiment. Like light microscopy, the FlowCAM approach estimated δ based on ROD in the experimental containers. However, imaging cytometry also measured the relative cellular chlorophyll a content in the ciliates' food vacuoles as a proxy for the nutritional status of the cells. The linear decline of the cellular chl. a yielded an independent estimate of δ that was similar to δ calculated from ROD. Additionally, the FlowCAM measurements revealed a high degree of phenotypic plasticity of the ciliates when exposed to starvation.

[Numerical Response Analysis of PM2.5-O3 Compound Pollution in Beijing].

The multi-scale variation trend of PM2.5-O3 compound pollution events was analyzed based on air quality data, meteorological data, and COVID-19 data in Beijing from 2015 to 2020. For the threshold of compound pollution, a compound pollution index was proposed, and the numerical response trend was evaluated based on the generalized additive model. A distributed lag nonlinear model was introduced to analyze the risk response relationship between compound pollution and influencing factors. The results showed that the events of PM2.5-O3 compound pollution in Beijing decreased annually. At the same time, due to the influence of pollutant emissions and meteorological conditions, there were obvious seasonal effects, week effects, holiday effects, and epidemic effects. The composite pollution index had no correlation with rainfall but had a linear positive correlation with O3 and air temperature and a nonlinear correlation with other explanatory variables. Air pollutants and meteorological conditions had obvious lag effects on the composite pollution index, and the lag effects were mainly concentrated in 1-3 d. PM2.5, PM10, O3, SO2, and air temperature in high-value areas significantly increased the risk of compound pollution. The CO (1-6 mg·m-3), NO2 (38-118 µg·m-3), and relative humidity (54%-87%) in the median section would also increase the risk of compound pollution, as would low wind speed. The compound pollution events showed a trend of multi-day continuous pollution in the numerical response. Compared with PM2.5 and PM10, compound pollution events were more dependent on O3, and the compound pollution rate in high-value areas was 30.7%-47.5%. CO and relative humidity had little effect on compound pollution events. The air temperature had the greatest impact, and 84.7% of the composite pollution incidents occurred at 20-30℃.

Foraging behaviour of Scymnus coccivora Ayyar against cotton mealybug Phenacoccus solenopsis Tinsley.

Predation is one of the significant biotic mortality factors reducing the insect pest population as functional response and the numerical response of the predator are the key factor regulating the population dynamics of predator prey species. This study is aimed to evaluate the functional response of all the developmental stages of Scymnus coccivora Ayyar (Coleoptera: Coccinellidae) against the different densities of cotton mealybug, Phenacoccus solenopsis (Tinsley) (Hemiptera: Pseudococcidae) and the numerical response of female predator. Experiments were carried out in controlled environment laboratory conditions at 25 ± 1 °C temperature, 60 ± 5% relative humidity and photoperiod of 16 h. Number of eggs consumed, number of eggs laid and the Efficiency of Conversion of Ingested food (ECI) were recorded daily. Results from the study revealed that all the developmental stages of S. coccivora exhibited a Type II response. Different parameters such as attack rate (a'), handling time (Th) and the maximum rate of predation were estimated using Roger's random attack equation and Holling Disc equation in which Rogers random attack equation was found best fit. Female has shown the highest attack rate (a') followed by IVth instar grub, male, IIIrd, IInd and Ist instar grub. With low handling time, IVth instar grub has shown maximum predation rate of 76.40 per day followed by female (75.86), male (58.79), IIIrd (22.84), IInd (19.65) and Ist instar grub (15.39). The numerical response increase was curvilinearly related to different prey densities with the highest number of eggs (11.8 ± 3.44) produced at highest prey density (160). The Efficiency of Conversion of Ingested food (ECI) was highest (64.49 ± 8.03) at prey density of 10. Understanding the factors that lead to variation in functional response of predator in natural population will advance our understanding of the effects of predation on individual and the effectiveness of coccinellid predators as biocontrol agent against cotton mealybug.

A comprehensive picture of foraging strategies of Neoseiulus cucumeris and Amblyseius swirskii on western flower thrips.

BACKGROUND: This study evaluated the potential of Neoseiulus cucumeris Oudemans and Amblyseius swirskii Athias-Henriot (Acari: Phytoseiidae) for controlling western flower thrips (WFT), Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), by determining the functional response, numerical response, prey switching and mutual interference behaviors of the predators. RESULTS: A type II functional response was detected for two predator species fed on WFT first instars. The attack rate was higher for A. swirskii than N. cucumeris, but the handling time of N. cucumeris was shorter. For two predator species, numerical response was affected by both prey density and oviposition time. The per capita searching efficiencies of the two predators decreased significantly with increasing predator densities. The interference coefficient for A. swirskii was less than that of N. cucumeris. Although N. cucumeris switched from feeding on WFT first instars to feeding on second instars, this behavior was not observed in A. swirskii. CONCLUSIONS: Comparison of the two predator species foraging behaviors indicated optimal performance by A. swirskii at lower F. occidentalis densities. N. cucumeris was found to perform optimally at relatively higher prey densities. Moreover, N. cucumeris could be more effective in stabilizing the predator-prey system by switching behavior compared with A. swirskii. At higher predator densities, A. swirskii seemed to be an effective agent owing to its reduced intraspecific competition compared with N. cucumeris. Findings from this study will be useful in selecting appropriate agents and developing effective biocontrol programs to control WFT. © 2021 Society of Chemical Industry.

Variable effects of mycorrhizal fungi on predator-prey dynamics under field conditions.

Interactions between herbivores and their predators are shaped, in part, by plant phenotype. Consequently, ubiquitous symbionts of plants below-ground, such as arbuscular mycorrhizal fungi (AMF), may influence interactions above-ground between predators and their prey by altering plant phenotype. However, the ecological relevance of below-ground organisms on predator-prey interactions under field conditions remains unclear. We assessed how AMF influence herbivore-predator interactions through a field experiment. We planted two milkweed species (Asclepias curassavica and Asclepias incarnata) provided with different amounts of AMF inoculum (zero, medium, and high) in a randomized block design. We added aphids to plants and reduced predator pressure weekly for 5 weeks to evaluate effects of AMF on predator recruitment. We then allowed herbivore-predator interactions to re-establish naturally for the remainder of the season to examine whether AMF-mediated variation in predator recruitment influenced the suppression of aphid populations. Arbuscular mycorrhizal fungi availability in soils mediated interactions between predaceous aphid midge flies Aphidoletes aphidimyza and their aphid prey Aphis nerii, but the effects were plant species-specific. On A. curassavica, by mid-season, midges were recruited most strongly on plants under medium AMF availability and least on plants under high AMF availability. In contrast, each midge killed fewer aphids with increasing aphid density on medium AMF plants, but killed more aphids with increasing aphid density on high AMF plants. In combination, aphid mortality rates imposed by midges were greatest on medium AMF plants, followed by high and zero AMF plants. By comparison, on A. incarnata, the recruitment of midges was strongest on high AMF plants and weakest on medium AMF plants. AMF had no effect on the number of aphids killed per midge, relative to aphid density, so mortality rates of aphids imposed by midges mirrored recruitment. Rates of decline in aphid populations following predator recolonization were associated with midge densities, as well as lacewing and syrphid densities, which were unaffected by AMF availability. Therefore, the effects of AMF on aphid population decline were not a simple function of AMF-midge interactions. Our findings demonstrate that the availability of AMF in soils has pervasive, but complex, effects on predator-herbivore dynamics in the field.

Functional Ecology of Two Contrasting Freshwater Ciliated Protists in Relation to Temperature.

We conducted microcosm experiments with two contrasting freshwater ciliates on functional traits (FTs) related to their growth rate (numerical response, NR) and ingestion rate (functional response, FR) over a range of ecologically relevant temperatures. Histiobalantium bodamicum and Vorticella natans are common planktonic ciliates but their abundance, swimming behavior, and temperature tolerance are different. In contrast to most sessile peritrich species, the motile V. natans is not strictly bacterivorous but also voraciously feeds upon small algae. We observed three main alterations in the shape of NR of both species with temperature, that is, change in the maximum growth rate, in the initial slope and in the threshold food level needed to sustain the population. Similarly, maximum ingestion rate, gross growth efficiency (GGE), and cell size varied with temperature and species. These findings caution against generalizing ciliate performance in relation to the ongoing global warming. Our results suggest that V. natans is the superior competitor to H. bodamicum in terms of temperature tolerance and bottom-up control. However, the abundance of V. natans is usually low compared to H. bodamicum and other common freshwater ciliates, suggesting that V. natans is more strongly top-down controlled via predation than H. bodamicum. The taxonomic position of V. natans has been debated. Therefore, to confirm species and genus affiliation of our study objects, we sequenced their small subunit ribosomal RNA (SSU rDNA) gene.

Consumer Responses to Experimental Pulsed Subsidies in Isolated versus Connected Habitats.

AbstractIncreases in consumer abundance following a resource pulse can be driven by diet shifts, aggregation, and reproductive responses, with combined responses expected to result in faster response times and larger numerical increases. Previous work in plots on large Bahamian islands has shown that lizards (Anolis sagrei) increased in abundance following pulses of seaweed deposition, which provide additional prey (i.e., seaweed detritivores). Numerical responses were associated with rapid diet shifts and aggregation, followed by increased reproduction. These dynamics are likely different on isolated small islands, where lizards cannot readily immigrate or emigrate. To test this, we manipulated the frequency and magnitude of seaweed resource pulses on whole small islands and in plots within large islands, and we monitored lizard diet and numerical responses over 4 years. We found that seaweed addition caused persistent increases in lizard abundance on small islands regardless of pulse frequency or magnitude. Increased abundance may have occurred because the initial pulse facilitated population establishment, possibly via enhanced overwinter survival. In contrast with a previous experiment, we did not detect numerical responses in plots on large islands, despite lizards consuming more marine resources in subsidized plots. This lack of a numerical response may be due to rapid aggregation followed by disaggregation or to stronger suppression of A. sagrei by their predators on the large islands in this study. Our results highlight the importance of habitat connectivity in governing ecological responses to resource pulses and suggest that disaggregation and changes in survivorship may be underappreciated drivers of pulse-associated dynamics.

Interactions between different predator-prey states: a method for the derivation of the functional and numerical response.

In this paper we introduce a formal method for the derivation of a predator's functional response from a system of fast state transitions of the prey or predator on a time scale during which the total prey and predator densities remain constant. Such derivation permits an explicit interpretation of the structure and parameters of the functional response in terms of individual behaviour. The same method is also used here to derive the corresponding numerical response of the predator as well as of the prey.

Density dependence forces divergent population growth rates and alters occupancy patterns of a central place foraging Antarctic seabird.

Density-dependent regulation is an important process in spatio-temporal population dynamics because it can alter the effects of synchronizing processes operating over large spatial scales. Most frequently, populations are regulated by density dependence when higher density leads to reduced individual fitness and population growth, but inverse density dependence can also occur when small populations are subject to higher extinction risks. We investigate whether density-dependent regulation influences population growth for the Antarctic breeding Adélie penguin Pygoscelis adeliae. Understanding the prevalence and nature of density dependence for this species is important because it is considered a sentinel species reflecting the impacts of fisheries and environmental change over large spatial scales in the Southern Ocean, but the presence of density dependence could introduce uncertainty in this role. Using data on population growth and indices of resource availability for seven regional Adélie penguin populations located along the East Antarctic coastline, we find compelling evidence that population growth is constrained at some locations by the amount of breeding habitat available to individuals. Locations with low breeding habitat availability had reduced population growth rates, higher overall occupancy rates, and higher occupancy of steeper slopes that are sparsely occupied or avoided at other locations. Our results are consistent with evolutionary models of avian breeding habitat selection where individuals search for high-quality nest sites to maximize fitness returns and subsequently occupy poorer habitat as population density increases. Alternate explanations invoking competition for food were not supported by the available evidence, but strong conclusions on food-related density dependence were constrained by the paucity of food availability data over the large spatial scales of this region. Our study highlights the importance of incorporating nonconstant conditions of species-environment relationships into predictive models of species distributions and population dynamics, and provides guidance for improved monitoring of fisheries and climate change impacts in the Southern Ocean.

Pulsed seaweed subsidies drive sequential shifts in the effects of lizard predators on island food webs.

Most prominent theories of food web dynamics imply the simultaneous action of bottom-up and top-down forces. However, transient bottom-up effects resulting from resource pulses can lead to sequential shifts in the strength of top-down predator effects. We used a large-scale field experiment (32 small islands sampled over 5 years) to probe how the frequency and magnitude of pulsed seaweed inputs drives temporal variation in the top-down effects of lizard predators. Short-term weakening of lizard effects on spiders and plants (the latter via a trophic cascade) were associated with lizard diet shifts, and were more pronounced with larger seaweed inputs. Long-term strengthening of lizard effects was associated with lizard numerical responses and plant fertilisation. Increased pulse frequency reinforced the strengthening of lizard effects on spiders and plants. These results underscore the temporally variable nature of top-down effects and highlight the role of resource pulses in driving this variation.

Are tintinnids picky grazers: Feeding experiments on a mixture of mixotrophic dinoflagellates and implications for red tide dynamics.

To understand and predict the outbreak of red tides, which are often dominated by mixotrophic dinoflagellates (MTDs), the effects of "top-down" control by co-occurring predators on red-tide MTDs should be taken into consideration. We studied the numerical and functional responses of the tintinnid ciliate Favella ehrenbergii feeding on two red-tide MTDs, Scrippsiella trochoidea and Heterocapsa triquetra, under single and mixed prey conditions. Our results suggest that a mixed diet could support a better growth of predators compared to a monodiet. In addition, the predators preferred to graze S. trochoidea in the mixed diets, suggesting that predators may switch their feeding preference. The grazing by tintinnid predators could potentially inhibit the outbreaks of red tides dominated by MTDs. The findings in this study provide basic data and new insights for understanding the complex predator-prey relationships in marine microbial food webs, and the dynamics of red tides dominated by MTDs.

Flexibility in a changing arctic food web: Can rough-legged buzzards cope with changing small rodent communities?

Indirect effects of climate change are often mediated by trophic interactions and consequences for individual species depend on how they are tied into the local food web. Here we show how the response of demographic rates of an arctic bird of prey to fluctuations in small rodent abundance changed when small rodent community composition and dynamics changed, possibly under the effect of climate warming. We observed the breeding biology of rough-legged buzzards (Buteo lagopus) at the Erkuta Tundra Monitoring Site in southern Yamal, low arctic Russia, for 19 years (1999-2017). At the same time, data on small rodent abundance were collected and information on buzzard diet was obtained from pellet dissection. The small rodent community experienced a shift from high-amplitude cycles to dampened fluctuations paralleled with a change in species composition toward less lemmings and more voles. Buzzards clearly preferred lemmings as prey. Breeding density of buzzards was positively related to small rodent abundance, but the shift in small rodent community lead to lower numbers relative to small rodent abundance. At the same time, after the change in small rodent community, the average number of fledglings was higher relative to small rodent abundance than earlier. These results suggest that the buzzard population adapted to a certain degree to the changes in the major resource, although at the same time density declined. The documented flexibility in the short-term response of demographic rates to changes in structure and dynamics of key food web components make it difficult to predict how complex food webs will be transformed in a warmer Arctic. The degree of plasticity of functional responses is indeed likely to vary between species and between regions, depending also on the local food web context.

Cross-scale effects of spruce budworm outbreaks on boreal warblers in eastern Canada.

Insect outbreaks are major natural disturbance events that affect communities of forest birds, either directly by affecting the food supply or indirectly by changing the vegetation composition of forest canopies. An examination of correlations between measures of bird and insect abundance across different spatial scales and over varying time lag effects may provide insight into underlying mechanisms. We developed a hierarchical Bayesian model to assess correlations between counts of eight warbler species from the Breeding Bird Survey in eastern Canada, 1966 to 2009, with the presence of spruce budworm (Choristoneura fumiferana Clem.) at immediate local scales and time-lagged regional scales, as measured by extent of defoliation on host tree species. Budworm-associated species Cape May warbler (Setophaga tigrina), bay-breasted warbler (Setophaga castanea), and Tennessee warbler (Oreothlypis peregrina) responded strongly and positively to both local and regional effects. In contrast, non-budworm-associated species, Blackburnian warbler (Setophaga fusca), magnolia warbler (Setophaga magnolia), Canada warbler (Cardellina canadensis), black-throated blue warbler (Setophaga caerulescens), and black-throated green warbler (Setophaga virens), only responded to regional effects in a manner that varied across eastern Canada. The complex responses by forest birds to insect outbreaks involve both increased numerical responses to food supply and to longer term responses to changes in forest structure and composition. These effects can vary across spatial scales and be captured in hierarchical population models, which can serve to disentangle common trends from data when examining drivers of population dynamics like forest management or climate change.

Numerical response of mammalian carnivores to rodents affects bird reproduction in temperate forests: A case of apparent competition?

Resource pulses such as mast seeding in temperate forests may affect interspecific interactions over multiple trophic levels and link different seed and nonseed consumers directly via predation or indirectly via shared predators. However, the nature and strength of interactions among species remain unknown for most resource pulse-driven ecosystems. We considered five hypotheses concerning the influence of resource pulses on the interactions between rodents, predators, and bird reproduction with data from northern Switzerland collected between 2010 and 2015. In high-rodent-abundance-years (HRAYs), wood warbler (Phylloscopus sibilatrix) nest survival was lower than in low-rodent-abundance-years, but rodents were not important nest predators, in contrast to rodent-hunting predators. The higher proportion of nests predated by rodent-hunting predators and their increased occurrence in HRAYs suggests a rodent-mediated aggregative numerical response of rodent-hunting predators, which incidentally prey on the wood warbler's ground nests. There was no evidence that rodent-hunting predators responded behaviorally by switching prey. Lastly, nest losses caused by nonrodent-hunting predators were not related to rodent abundance. We show that wood warblers and rodents are linked via shared predators in a manner consistent with apparent competition, where an increase of one species coincides with the decrease of another species mediated by shared predators. Mast seeding frequency and annual seed production appear to have increased over the past century, which may result in more frequent HRAYs and generally higher peaking rodent populations. The associated increase in the magnitude of apparent competition may thus at least to some extent explain the wood warbler's decline in much of Western Europe.

Vole abundance and reindeer carcasses determine breeding activity of Arctic foxes in low Arctic Yamal, Russia.

BACKGROUND: High latitude ecosystems are at present changing rapidly under the influence of climate warming, and specialized Arctic species at the southern margin of the Arctic may be particularly affected. The Arctic fox (Vulpes lagopus), a small mammalian predator endemic to northern tundra areas, is able to exploit different resources in the context of varying tundra ecosystems. Although generally widespread, it is critically endangered in subarctic Fennoscandia, where a fading out of the characteristic lemming cycles and competition with abundant red foxes have been identified as main threats. We studied an Arctic fox population at the Erkuta Tundra Monitoring site in low Arctic Yamal (Russia) during 10 years in order to determine which resources support the breeding activity in this population. In the study area, lemmings have been rare during the last 15 years and red foxes are nearly absent, creating an interesting contrast to the situation in Fennoscandia. RESULTS: Arctic fox was breeding in nine of the 10 years of the study. The number of active dens was on average 2.6 (range 0-6) per 100 km2 and increased with small rodent abundance. It was also higher after winters with many reindeer carcasses, which occurred when mortality was unusually high due to icy pastures following rain-on-snow events. Average litter size was 5.2 (SD = 2.1). Scat dissection suggested that small rodents (mostly Microtus spp.) were the most important prey category. Prey remains observed at dens show that birds, notably waterfowl, were also an important resource in summer. CONCLUSIONS: The Arctic fox in southern Yamal, which is part of a species-rich low Arctic food web, seems at present able to cope with a state shift of the small rodent community from high amplitude cyclicity with lemming dominated peaks, to a vole community with low amplitude fluctuations. The estimated breeding parameters characterized the population as intermediate between the lemming fox and the coastal fox ecotype. Only continued ecosystem-based monitoring will reveal their fate in a changing tundra ecosystem.