Dampening prey cycle overrides the impact of climate change on predator population dynamics: a long‐term demographic study on tawny owls

Predicting the dynamics of animal populations with different life histories requires careful understanding of demographic responses to multifaceted aspects of global changes, such as climate and trophic interactions. Continent‐scale dampening of vole population cycles, keystone herbivores in many ecosystems, has been recently documented across Europe. However, its impact on guilds of vole‐eating predators remains unknown. To quantify this impact, we used a 27‐year study of an avian predator (tawny owl) and its main prey (field vole) collected in Kielder Forest (UK) where vole dynamics shifted from a high‐ to a low‐amplitude fluctuation regime in the mid‐1990s. We measured the functional responses of four demographic rates to changes in prey dynamics and winter climate, characterized by wintertime North Atlantic Oscillation (wNAO). First‐year and adult survival were positively affected by vole density in autumn but relatively insensitive to wNAO. The probability of breeding and number of fledglings were higher in years with high spring vole densities and negative wNAO (i.e. colder and drier winters). These functional responses were incorporated into a stochastic population model. The size of the predator population was projected under scenarios combining prey dynamics and winter climate to test whether climate buffers or alternatively magnifies the impact of changes in prey dynamics. We found the observed dampening vole cycles, characterized by low spring densities, drastically reduced the breeding probability of predators. Our results illustrate that (i) change in trophic interactions can override direct climate change effect; and (ii) the demographic resilience entailed by longevity and the occurrence of a floater stage may be insufficient to buffer hypothesized environmental changes. Ultimately, dampened prey cycles would drive our owl local population towards extinction, with winter climate regimes only altering persistence time. These results suggest that other vole‐eating predators are likely to be threatened by dampening vole cycles throughout Europe.     

Predator–vole interactions in northern Europe: the role of small mustelids revised

The cyclic population dynamics of vole and predator communities is a key phenomenon in northern ecosystems, and it appears to be influenced by climate change. Reports of collapsing rodent cycles have attributed the changes to warmer winters, which weaken the interaction between voles and their specialist subnivean predators. Using population data collected throughout Finland during 1986–2011, we analyse the spatio-temporal variation in the interactions between populations of voles and specialist, generalist and avian predators, and investigate by simulations the roles of the different predators in the vole cycle. We test the hypothesis that vole population cyclicity is dependent on predator–prey interactions during winter. Our results support the importance of the small mustelids for the vole cycle. However, weakening specialist predation during winters, or an increase in generalist predation, was not associated with the loss of cyclicity. Strengthening of delayed density dependence coincided with strengthening small mustelid influence on the summer population growth rates of voles. In conclusion, a strong impact of small mustelids during summers appears highly influential to vole population dynamics, and deteriorating winter conditions are not a viable explanation for collapsing small mammal population cycles.     

Climate change and perishable food hoards of an avian predator: Is the freezer still working?

Changing climate can modify predator–prey interactions and induce declines or local extinctions of species due to reductions in food availability. Species hoarding perishable food for overwinter survival, like predators, are predicted to be particularly susceptible to increasing temperatures. We analysed the influence of autumn and winter weather, and abundance of main prey (voles), on the food‐hoarding behaviour of a generalist predator, the Eurasian pygmy owl (Glaucidium passerinum), across 16 years in Finland. Fewer freeze–thaw events in early autumn delayed the initiation of food hoarding. Pygmy owls consumed more hoarded food with more frequent freeze–thaw events and deeper snow cover in autumn and in winter, and lower precipitation in winter. In autumn, the rotting of food hoards increased with precipitation. Hoards already present in early autumn were much more likely to rot than the ones initiated in late autumn. Rotten food hoards were used more in years of low food abundance than in years of high food abundance. Having rotten food hoards in autumn resulted in a lower future recapture probability of female owls. These results indicate that pygmy owls might be partly able to adapt to climate change by delaying food hoarding, but changes in the snow cover, precipitation and frequency of freeze–thaw events might impair their foraging and ultimately decrease local overwinter survival. Long‐term trends and future predictions, therefore, suggest that impacts of climate change on wintering food‐hoarding species could be substantial, because their ‘freezers’ may no longer work properly. Altered usability and poorer quality of hoarded food may further modify the foraging needs of food‐hoarding predators and thus their overall predation pressure on prey species. This raises concerns about the impacts of climate change on boreal food webs, in which ecological interactions have evolved under cold winter conditions.     

Increased autumn rainfall disrupts predator–prey interactions in fragmented boreal forests

There is a pressing need to understand how changing climate interacts with land‐use change to affect predator–prey interactions in fragmented landscapes. This is particularly true in boreal ecosystems facing fast climate change and intensification in forestry practices. Here, we investigated the relative influence of autumn climate and habitat quality on the food‐storing behaviour of a generalist predator, the pygmy owl, using a unique data set of 15 850 prey items recorded in western Finland over 12 years. Our results highlighted strong effects of autumn climate (number of days with rainfall and with temperature <0 °C) on food‐store composition. Increasing frequency of days with precipitation in autumn triggered a decrease in (i) total prey biomass stored, (ii) the number of bank voles (main prey) stored, and (iii) the scaled mass index of pygmy owls. Increasing proportions of old spruce forests strengthened the functional response of owls to variations in vole abundance and were more prone to switch from main prey to alternative prey (passerine birds) depending on local climate conditions. High‐quality habitat may allow pygmy owls to buffer negative effects of inclement weather and cyclic variation in vole abundance. Additionally, our results evidenced sex‐specific trends in body condition, as the scaled mass index of smaller males increased while the scaled mass index of larger females decreased over the study period, probably due to sex‐specific foraging strategies and energy requirements. Long‐term temporal stability in local vole abundance refutes the hypothesis of climate‐driven change in vole abundance and suggests that rainier autumns could reduce the vulnerability of small mammals to predation by pygmy owls. As small rodents are key prey species for many predators in northern ecosystems, our findings raise concern about the impact of global change on boreal food webs through changes in main prey vulnerability.     

Mortality and lamb body mass growth in free‐ranging domestic sheep – environmental impacts including lethal and non‐lethal impacts of predators

The management and recovery of large predator populations in areas where human persecution has driven them to ecological extinction requires a solid understanding of the effects of both predation and food limitation on prey populations. We used 11 yr of data on reported losses among 17.3 million free‐ranging sheep Ovis aries in the Norwegian farming industry to elucidate the relative roles of climate, vegetation characteristics, sheep densities, lamb body mass and densities of predators and alternative prey on the number of lambs and ewes lost on summer pastures. We first examined whether predator densities predicted autumn lamb body mass through possible impacts of predators on body growth (non‐lethal effects) but found no evidence for such effects in our study system. This might be due to weak anti‐predator behavioral responses in domesticated sheep. However, autumn lamb body mass was predicted by both sheep density and winter and spring weather conditions, probably through food availability. Losses of both lambs and ewes were positively and strongly related to the density of Eurasian lynx Lynx lynx, wolverine Gulo gulo and brown bear Ursus arctos. In addition, food availability and spring weather conditions were associated to losses of lambs, while precipitation in May predicted losses of ewes. There was little evidence for interaction effects of predator species on losses, suggesting that most of the effects of the predators were additive to each other. Given the strong effect of predator densities on sheep losses, we conclude that changing livestock husbandry practices towards a system that actively protects sheep and/or active management of predator densities may be necessary to reduce sheep losses where predators are recolonizing.     

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.     

Density dependence,prey accessibility and prey depletion by fisheries drive Peruvian seabird population dynamics

In marine ecosystems top predator populations are shaped by environmental factors affecting their prey abundance. Coupling top predators’ population studies with independent records of prey abundance suggests that prey fluctuations affect fecundity parameters and abundance of their predators. However, prey may be abundant but inaccessible to their predators and a major challenge is to determine the relative importance of prey accessibility in shaping seabird populations. In addition, disentangling the effects of prey abundance and accessibility from the effects of prey removal by fisheries, while accounting for density dependence, remains challenging for marine top predators. Here, we investigate how climate, population density, and the accessibility and removal of prey (the Peruvian anchovy Engraulis ringens) by fisheries influence the population dynamics of the largest sedentary seabird community (≈ 4 million individuals belonging to guanay cormorant Phalacrocorax bougainvillii, Peruvian booby Sula variegata and Peruvian pelican Pelecanus thagus) of the northern Humboldt Current System over the past half‐century. Using Gompertz state–space models we found strong evidence for density dependence in abundance for the three seabird species. After accounting for density dependence, sea surface temperature, prey accessibility (defined by the depth of the upper limit of the subsurface oxygen minimum zone) and prey removal by fisheries were retained as the best predictors of annual population size across species. These factors affected seabird abundance the current year and with year lags, suggesting effects on several demographic parameters including breeding propensity and adult survival. These findings highlight the effects of prey accessibility and fishery removals on seabird populations in marine ecosystems. This will help refine management objectives of marine ecosystems in order to ensure sufficient biomass of forage fish to avoid constraining seabird population dynamics, while taking into account of the effects of environmental variability.     

Mechanisms of coexistence in diverse herbivore–carnivore assemblages: demographic,temporal and spatial heterogeneities affecting prey vulnerability

Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.     

Predator–prey relationships in a Mediterranean vertebrate system: Bonelli’s eagles,rabbits and partridges

How predators impact on prey population dynamics is still an unsolved issue for most wild predator–prey communities. When considering vertebrates, important concerns constrain a comprehensive understanding of the functioning of predator–prey relationships worldwide; e.g. studies simultaneously quantifying ‘functional’ and ‘numerical responses’ (i.e., the ‘total response’) are rare. The functional, the numerical, and the resulting total response (i.e., how the predator per capita intake, the population of predators and the total of prey eaten by the total predators vary with prey densities) are fundamental as they reveal the predator’s ability to regulate prey population dynamics. Here, we used a multi-spatio-temporal scale approach to simultaneously explore the functional and numerical responses of a territorial predator (Bonelli’s eagle Hieraaetus fasciatus) to its two main prey species (the rabbit Oryctolagus cuniculus and the red-legged partridge Alectoris rufa) during the breeding period in a Mediterranean system of south Spain. Bonelli’s eagle responded functionally, but not numerically, to rabbit/partridge density changes. Type II, non-regulatory, functional responses (typical of specialist predators) offered the best fitting models for both prey. In the absence of a numerical response, Bonelli’s eagle role as a regulating factor of rabbit and partridge populations seems to be weak in our study area. Simple (prey density-dependent) functional response models may well describe the short-term variation in a territorial predator’s consumption rate in complex ecosystems.     

Combined climate‐ and prey‐mediated range expansion of Humboldt squid (Dosidicus gigas), a large marine predator in the California Current System

Climate‐driven range shifts are ongoing in pelagic marine environments, and ecosystems must respond to combined effects of altered species distributions and environmental drivers. Hypoxic oxygen minimum zones (OMZs) in midwater environments are shoaling globally; this can affect distributions of species both geographically and vertically along with predator–prey dynamics. Humboldt (jumbo) squid (Dosidicus gigas) are highly migratory predators adapted to hypoxic conditions that may be deleterious to their competitors and predators. Consequently, OMZ shoaling may preferentially facilitate foraging opportunities for Humboldt squid. With two separate modeling approaches using unique, long‐term data based on in situ observations of predator, prey, and environmental variables, our analyses suggest that Humboldt squid are indirectly affected by OMZ shoaling through effects on a primary food source, myctophid fishes. Our results suggest that this indirect linkage between hypoxia and foraging is an important driver of the ongoing range expansion of Humboldt squid in the northeastern Pacific Ocean.     

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.     

Circumpolar dynamics of a marine top‐predator track ocean warming rates

Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long‐lived, wide‐ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black‐legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea‐surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large‐scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom‐up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large‐scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.     

The cost of migratory prey: seasonal changes in semi‐domestic reindeer distribution influences breeding success of Eurasian lynx in northern Norway

Migratory prey is a widespread phenomenon that has implications for predator–prey interactions. By creating large temporal variation in resource availability between seasons it becomes challenging for carnivores to secure a regular year‐round supply of food. Some predators may respond by following their migratory prey, however, most predators are sedentary and experience strong seasonal variation in resource availability. Increased predation on alternative prey may dampen such seasonal resource fluctuations, but reduced reproduction rates in predators is a predicted consequence of migratory primary prey behavior that has received little empirical attention. We used data from 23 GPS collared Eurasian lynx Lynx lynx monitored during 2007–2013 in northern Norway, to examine how spatio‐temporal variation in the migratory behavior of semi‐domestic reindeer Rangifer tarandus influences lynx spatial organization and reproductive success using estimates of seasonal home range overlap and breeding success. We found that lynx of both sexes maintained seasonally stable home ranges and exhibited site fidelity across years, independent of whether they had access to reindeer throughout the year or experienced a scarcity of reindeer in winter due to migration. However, lynx without access to reindeer in winter showed a decreased probability of reproducing and a tendency for lowered kitten survival into their first winter, when compared to female lynx with reindeer available year around. This supports the hypothesis that sedentary predators experience demographic costs in systems with migratory primary prey. Changes in the migratory behavior of ungulates, including disrupted migrations, is therefore likely to have bottom–up effects on the population dynamics of sedentary predators as well as the previously documented consequences for ungulate population dynamics.     

Warming up the system: higher predator feeding rates but lower energetic efficiencies

Predictions on the consequences of the rapidly increasing atmospheric CO₂ levels and associated climate warming for population dynamics, ecological community structure and ecosystem functioning depend on mechanistic energetic models of temperature effects on populations and their interactions. However, such mechanistic approaches combining warming effects on metabolic (energy loss of organisms) and feeding rates (energy gain by organisms) remain a key, yet elusive, goal. Aiming to fill this void, we studied the metabolic rates and functional responses of three differently sized, predatory ground beetles on one mobile and one more resident prey species across a temperature gradient (5, 10, 15, 20, 25 and 30 °C). Synthesizing metabolic and functional‐response theory, we develop novel mechanistic predictions how predator–prey interaction strengths (i.e., functional responses) should respond to warming. Corroborating prior theory, warming caused strong increases in metabolism and decreases in handling time. Consistent with our novel model, we found increases in predator attack rates on a mobile prey, whereas attack rates on a mostly resident prey remained constant across the temperature gradient. Together, these results provide critically important information that environmental warming generally increases the direct short‐term per capita interaction strengths between predators and their prey as described by functional‐response models. Nevertheless, the several fold stronger increase in metabolism with warming caused decreases in energetic efficiencies (ratio of per capita feeding rate to metabolic rate) for all predator–prey interactions. This implies that warming of natural ecosystems may dampen predator–prey oscillations thus stabilizing their dynamics. The severe long‐term implications; however, include predator starvation due to energetic inefficiency despite abundant resources.     

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.     

Mammalian predator–prey interaction in a fragmented landscape: weasels and voles

The relationship between predators and prey is thought to change due to habitat loss and fragmentation, but patterns regarding the direction of the effect are lacking. The common prediction is that specialized predators, often more dependent on a certain habitat type, should be more vulnerable to habitat loss compared to generalist predators, but actual fragmentation effects are unknown. If a predator is small and vulnerable to predation by other larger predators through intra-guild predation, habitat fragmentation will similarly affect both the prey and the small predator. In this case, the predator is predicted to behave similarly to the prey and avoid open and risky areas. We studied a specialist predator’s, the least weasel, Mustela nivalis nivalis, spacing behavior and hunting efficiency on bank voles, Myodes glareolus, in an experimentally fragmented habitat. The habitat consisted of either one large habitat patch (non-fragmented) or four small habitat patches (fragmented) with the same total area. The study was replicated in summer and autumn during a year with high avian predation risk for both voles and weasels. As predicted, weasels under radio-surveillance killed more voles in the non-fragmented habitat which also provided cover from avian predators during their prey search. However, this was only during autumn, when the killing rate was also generally high due to cold weather. The movement areas were the same for both sexes and both fragmentation treatments, but weasels of both sexes were more prone to take risks in crossing the open matrix in the fragmented treatment. Our results support the hypothesis that habitat fragmentation may increase the persistence of specialist predator and prey populations if predators are limited in the same habitat as their prey and they share the same risk from avian predation.     

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.     

Sacrificial males: the potential role of copulation and predation in contributing to copepod sex‐skewed ratios

Predation is thought to play a selective role in the emergence of behavioural traits in prey. Differences in behaviour between prey demographics may, therefore, be driven by predation with select components of the population being less vulnerable to predators. While under controlled conditions prey demography has been shown to have consequences for predation success, investigations linking these implications to natural prey population demographics are scarce. Here we assess predator–prey dynamics between notonectid predators (backswimmers) and Lovenula raynerae (Copepoda), key faunal groups in temperate ephemeral pond ecosystems. Using a combination of field and experimental approaches we test for the development and mechanism of predation‐induced sex‐skewed ratios. A natural population of L. raynerae was tracked over time in relation to their predator (notonectid) and prey (Cladocera) numbers. In the laboratory, L. raynerae sex ratios were also assessed over time but in the absence of predation pressure. Predation success and prey performance experiments evaluating differences between L. raynerae male, female, gravid female and copulating pairs exposed to notonectid predation were then examined. Under natural conditions, a female dominated copepod population developed over time and was correlated to predation pressure, while under predator‐free conditions non sex‐skewed prey population demographics persisted. Predator–prey laboratory trials showed no difference in vulnerability and escape performance for male, female and gravid female copepods, but pairs in copula were significantly more vulnerable to predation. This vulnerability was not shared by both sexes, with only female copepods ultimately escaping from successful predation on a mating pair. These results suggest that contact periods during copula may contribute to the development of sex‐skewed copepod ratios over time in ecosystems dominated by hexapod predators. This is discussed within the context of vertebrate and invertebrate predation and how these dissimilar types of predation are likely to have acted as selective pressures for copepod mating systems.     

Variation in foraging success among predators and its implications for population dynamics

The effects of the expected predation rate on population dynamics have been studied intensively, but little is known about the effects of predation rate variability (i.e., predator individuals having variable foraging success) on population dynamics. In this study, variation in foraging success among predators was quantified by observing the predation of the wolf spider Pardosa pseudoannulata on the cricket Gryllus bimaculatus in the laboratory. A population model was then developed, and the effect of foraging variability on predator–prey dynamics was examined by incorporating levels of variation comparable to those quantified in the experiment. The variability in the foraging success among spiders was greater than would be expected by chance (i.e., the random allocation of prey to predators). The foraging variation was density‐dependent; it became higher as the predator density increased. A population model that incorporates foraging variation shows that the variation influences population dynamics by affecting the numerical response of predators. In particular, the variation induces negative density‐dependent effects among predators and stabilizes predator–prey dynamics.     

Predator population dynamics under a cyclic prey regime: numerical responses,demographic parameters and growth rates

The consequences of cyclic fluctuations in abundance of prey species on predator continue to improve our understanding of the mechanisms behind population regulation. Among predators, vole‐eating raptors usually respond to changes in prey abundance with no apparent time‐lag and therefore contradict predictions from the predator–prey theory. In such systems, the interplay between demographic traits and population growth rate in relation to prey abundance remains poorly studied, yet it is crucial to characterize the link between ecological processes and population changes. Using a mechanistic approach, we assessed the demographic rates associated to the direct and indirect numerical responses of a specialist raptor (Montagu's harrier) to its cyclic prey (common vole), using long term data from two adjacent study sites in France. First‐year survival rates were weakly affected by vole abundance, probably due to the fact that Montagu's harriers are trans‐Saharan migrants and thus escape the vole collapse occurring in autumn–winter. Recruitment of yearling as well as breeding propensity of experienced adult females were strongly affected by vole abundance and at least partially shaped the trajectory of the breeding population. We argued that the strong density dependent signal detected in predator time series was mostly the phenomenological consequence of the positive direct numerical response of harriers to vole abundance. Accounting for this, we proposed a method to assess density dependence in predator relying on a cyclic prey. Finally, the variation in Montagu's harrier population growth rates was best explained by overwinter growth rates of the prey population and to a lesser extent by previous residual predator density.