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.     

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.     

Substitution of top predators: effects of pike invasion in a subarctic lake

1. Invasions of top predators may have strong cascading effects in ecosystems affecting both prey species abundance and lower trophic levels. A recently discussed factor that may enhance species invasion is climate change and in this context, we studied the effects of an invasion of northern pike into a subarctic lake ecosystem formerly inhabited by the native top predator Arctic char and its prey fish, ninespined stickleback. 2. Our study demonstrated a strong change in fish community composition from a system with Arctic char as top predator and high densities of sticklebacks to a system with northern pike as top predator and very low densities of sticklebacks. A combination of both predation and competition from pike is the likely cause of the extinction of char. 3. The change in top predator species also cascaded down to primary consumers as both zooplankton and predator‐sensitive macroinvertebrates increased in abundance. 4. Although the pike invasion coincided with increasing summer temperatures in the study area we have no conclusive evidence that the temperature increase is the causal mechanism behind the pike invasion. But still, our study provides possible effects of future pike invasions in mountain lakes related to climate change. We suggest that future pike invasions will have strong effects in lake ecosystems, both by replacing native top consumers and through cascading effects on lower trophic levels.     

Predator–prey relationships: arctic foxes and lemmings

1. The number of breeding dens and litter sizes of arctic foxes Alopex lagopus were recorded and the diet of the foxes was analysed during a ship-based expedition to 17 sites along the Siberian north coast. At the same time the cyclic dynamics of co-existing lemming species were examined.
2. The diet of arctic foxes was dominated by the Siberian lemming Lemmus sibiricus (on one site the Norwegian lemming L. lemmus ), followed by the collared lemming Dicrostonyx torquatus .
3. The examined Lemmus sibiricus populations were in different phases of the lemming cycle as determined by age profiles and population densities.
4. The numerical response of arctic foxes to varying densities of Lemmus had a time lag of 1 year, producing a pattern of limit cycles in lemming–arctic fox interactions. Arctic fox litter sizes showed no time lag, but a linear relation to Lemmus densities. We found no evidence for a numerical response to population density changes in Dicrostonyx .
5. The functional or dietary response of arctic foxes followed a type II curve for Lemmus , but a type III response curve for Dicrostonyx .
6. Arctic foxes act as resident specialist for Lemmus and may increase the amplitude and period of their population cycles. For Dicrostonyx , on the other hand, arctic foxes act as generalists which suggests a capacity to dampen oscillations.     

Evaluating a predator–prey interaction in the field: the interaction between beetle larvae (predator) and tadpoles (prey)

The larval amphibian community of temporary pond ecosystems has served as a model for studies in community ecology, with a majority of this work being conducted in mesocosms. Recent research has suggested that mesocosms may overestimate ecological effects; therefore, experimental studies conducted under field conditions are required to gauge the results of mesocosm studies. To assess a species interaction under more natural conditions, we conducted a series of field experiments examining the predator–prey interaction between beetle larvae ( Dytiscus sp.; predator) and larval wood frogs Rana sylvatica (prey) in central Pennsylvania, USA. Quantitative sampling of woodland ponds indicated that beetle larvae of the genus Dytiscus were the most common predator of tadpoles. In a field enclosure experiment, dytiscids were effective predators of tadpoles in the pond environment. Moreover, tadpoles avoided areas in a pond containing caged dytiscids, demonstrating that tadpoles recognize the chemical stimuli of predators in complex environments. The results of this study are consistent with data from prior laboratory and mesocosm studies and suggest that these venues can produce reliable interpretations of predator–prey dynamics in this community.     

Multiple predators induce risk reduction in coexisting vole species

Large predators may affect the hunting efficiency of smaller ones directly by decreasing their numbers, or indirectly by altering their behaviour. Either way this may have positive effects on the density of shared prey. Using large outdoor enclosures, we experimentally studied whether the presence of the Tengmalm's owl Aegolius funereus affects the hunting efficiency of the smallest member of the vole-eating predator guild, the least weasel Mustela nivalis, as measured by population responses of coexisting prey species, the field vole Microtus agrestis and the sibling vole M. levis . We compared the density and survival probability of vole populations exposed to no predation, weasel predation or combined predation by a weasel and an owl. The combined predation of both owl and weasel did not result in obvious changes in the density of sibling and field vole populations compared to the control populations without predators, while predation by least weasel alone decreased the densities of sibling voles and induced a similar trend in field vole densities. Survival of field voles was not affected by predator treatment while sibling vole survival was lower in predator treated populations than in control populations. Our results suggest that weasels are intimidated by avian predators, but without changing the effects of predators on competitive situations between the two vole species. Non-lethal effects of intraguild predation therefore will not necessarily change competitive interactions between shared prey species.     

Harbour porpoises respond to climate change

The effects of climate change on marine ecosystems and in particular on marine top predators are difficult to assess due to, among other things, spatial variability, and lack of clear delineation of marine habitats. The banks of West Greenland are located in a climate sensitive area and are likely to elicit pronounced responses to oceanographic changes in the North Atlantic. The recent increase in sea temperatures on the banks of West Greenland has had cascading effects on sea ice coverage, residency of top predators, and abundance of important prey species like Atlantic cod (Gadus morhua). Here, we report on the response of one of the top predators in West Greenland; the harbour porpoise (Phocoena phocoena). The porpoises depend on locating high densities of prey species with high nutritive value and they have apparently responded to the general warming on the banks of West Greenland by longer residence times, increased consumption of Atlantic cod resulting in improved body condition in the form of larger fat deposits in blubber, compared to the situation during a cold period in the 1990s. This is one of the few examples of a measurable effect of climate change on a marine mammal population.     

Effects of predator-specific defence on biodiversity and community complexity in two-trophic-level communities

Summary Antipredator strategies employed by prey may be specific (effective against only one type of predator) or non-specific (effective against all predators). To examine the effects of the specificity of antipredator behaviour on biodiversity and community complexity, we analyse mathematical models including both evolutionary and population dynamics of a system including multiple prey species and multiple predator species. The models assume that all predator species change in their prey choice and all prey species have evolutionary change in their antipredator effort in evolution. The traits of each species change in an adaptive manner, whose rate is proportional to the slope of their fitness function. We calculate community complexity, resource-overlap between predators, an index of biodiversity and other properties of the coevolutionarily stable community for two cases: (1) all prey species have non-specific antipredator behaviour and (2) all prey species have predator-specific defence. Predator-specificity in defence increases community complexity, resource-overlap between predators, the total abundance of predators and the ratio of predator to prey abundance. Specific defence also decreases the number of isolated subwebs within the entire foodweb.     

Strategies of aphidophagous predators: lessons for modelling insect predator–prey dynamics

Both theoretical and empirical evidence indicate that in systems where insect predators have longer developmental times than their prey the predators have little impact on the abundance of their prey. In assessing the 'effectiveness' of a predator for biological control one should take into account that selection maximizes predator fitness, not its effctiveness as a biocontrol agent. Therefore, predators that have a long developmental time relative to their prey are unlikely to be the best biocontrol agents. If these results can be generalized to other predator–prey systems, then it is clear that an understanding of predator–prey dynamics can only be achieved by studying predators.     

Coupled predator–prey oscillations in a chaotic food web

Coupling of several predator–prey oscillations can generate intriguing patterns of synchronization and chaos. Theory predicts that prey species will fluctuate in phase if predator–prey cycles are coupled through generalist predators, whereas they will fluctuate in anti-phase if predator–prey cycles are coupled through competition between prey species. Here, we investigate predator–prey oscillations in a long-term experiment with a marine plankton community. Wavelet analysis of the species fluctuations reveals two predator–prey cycles that fluctuate largely in anti-phase. The phase angles point at strong competition between the phytoplankton species, but relatively little prey overlap among the zooplankton species. This food web architecture is consistent with the size structure of the plankton community, and generates highly dynamic food webs. Continued alternations in species dominance enable coexistence of the prey species through a non-equilibrium 'killing-the-winner' mechanism, as the system shifts back and forth between the two predator–prey cycles in a chaotic fashion.     

Top–down limits on prey populations may be more severe in larger prey species,despite having fewer predators

Variation in the vulnerability of herbivore prey to predation is linked to body size, yet whether this relationship is size‐nested or size‐partitioned remains debated. If size‐partitioned, predators would be focused on prey within their preferred prey size range. If size‐nested, smaller prey species should become increasingly more vulnerable because increasingly more predators are capable of catching them. Yet, whether either of these strategies manifests in top–down prey population limitation would depend both on the number of potential predator species as well as the total mortality imposed. Here we use a rare ecosystem scale ‘natural experiment’ comparing prey population dynamics between a period of intense predator persecution and hence low predator densities and a period of active predator protection and population recovery. We use three decades of data on herbivore abundance and distribution to test the role of predation as a mechanism of population limitation among prey species that vary widely in body size. Notably, we test this within one of the few remaining systems where a near‐full suite of megaherbivores occur in high density and are thus able to include a thirtyfold range in herbivore body size gradient. We test whether top–down limitation on prey species of particular body size leads to compositional shifts in the mammalian herbivore community. Our results support both size‐nested and size‐partitioning predation but suggest that the relative top–down limiting impact on prey populations may be more severe for intermediate sized species, despite having fewer predators than small species. In addition we show that the gradual recovery of predator populations shifted the herbivore community assemblage towards large‐bodied species and has led to a community that is strongly dominated by large herbivore biomass.     

Coleomegilla maculata (Coleoptera: Coccinellidae) predation on pea aphids promoted by proximity to dandelions

The impact of a predator on its prey may depend on the presence of other species in the community. In particular, if predators are attracted to areas containing one prey species, another prey species may suffer greater predation if it occurs in the same areas. If the predator is omnivorous, this may occur even if one prey species is an animal and the other is a plant. We investigated the role of local dandelion densities on the impact of the predator Coleomegilla maculata on pea aphids in alfalfa fields. At small spatial scales, increased dandelion densities were associated with high C. maculata densities, presumably because these omnivorous ladybird beetles aggregated to pollen resources. In turn, the high C. maculata densities were associated with low aphid densities, presumably because of increased predation. We used laboratory cages to simulate C. maculata foraging in two adjacent patches of alfalfa, one with dandelions and one without. As in the field, the laboratory experiment showed that C. maculata aggregated to alfalfa interspersed with dandelions, which resulted in increased predation on aphids on alfalfa. This study demonstrates that a pollen-producing plant can indirectly decrease nearby herbivore densities by attracting an omnivorous predator.     

Food web structure and climate effects on the dynamics of small mammals and owls in semi-arid Chile

Population dynamics of small mammals and predators in semi-arid Chile is positively correlated with rainfall associated with incursions of El Niño (El Niño Southern Oscillation: ENSO). However, the causal relationships between small mammal fluctuations, predator oscillations, and climatic disturbances are poorly understood. Here, we report time series models for three species of small mammal prey and two species of owl predators. The large differences in population fluctuations between the three small mammal species are related to differences in their respective feedback structures. The analyses reveal that per capita growth rate of the leaf-eared mouse is a decreasing function of log density and of log barn owl abundance together with a positive rainfall effect. In turn, per capita population growth rate ( R -function) of the barn owl is a negative function of log barn owl abundance and a positive function of leaf-eared mouse abundance, suggesting a predator–prey interaction. The dramatic population fluctuations exhibited by leaf-eared mouse ( Phyllotis darwini ) are caused by climate effects coupled with a complex food web architecture.     

Prey and predator density‐dependent interactions under different water volumes

Predation is a critical ecological process that directly and indirectly mediates population stabilities, as well as ecosystem structure and function. The strength of interactions between predators and prey may be mediated by multiple density dependences concerning numbers of predators and prey. In temporary wetland ecosystems in particular, fluctuating water volumes may alter predation rates through differing search space and prey encounter rates. Using a functional response approach, we examined the influence of predator and prey densities on interaction strengths of the temporary pond specialist copepod Lovenula raynerae preying on cladoceran prey, Daphnia pulex, under contrasting water volumes. Further, using a population dynamic modeling approach, we quantified multiple predator effects across differences in prey density and water volume. Predators exhibited type II functional responses under both water volumes, with significant antagonistic multiple predator effects (i.e., antagonisms) exhibited overall. The strengths of antagonistic interactions were, however, enhanced under reduced water volumes and at intermediate prey densities. These findings indicate important biotic and abiotic contexts that mediate predator–prey dynamics, whereby multiple predator effects are contingent on both prey density and search area characteristics. In particular, reduced search areas (i.e., water volumes) under intermediate prey densities could enhance antagonisms by heightening predator–predator interference effects.     

Predator personality structures prey communities and trophic cascades

Intraspecific variation is central to our understanding of evolution and population ecology, yet its consequences for community ecology are poorly understood. Animal personality – consistent individual differences in suites of behaviours – may be particularly important for trophic dynamics, where predator personality can determine activity rates and patterns of attack. We used mesocosms with aquatic food webs in which the top predator (dragonfly nymphs) varied in activity and subsequent attack rates on zooplankton, and tested the effects of predator personality. We found support for four hypotheses: (1) active predators disproportionately reduce the abundance of prey, (2) active predators select for predator‐resistant prey species, (3) active predators strengthen trophic cascades (increase phytoplankton abundance) and (4) active predators are more likely to cannibalise one another, weakening all other trends when at high densities. These results suggest that intraspecific variation in predator personality is an important determinant of prey abundance, community composition and trophic cascades.     

Landscapes of fear or competition? Predation did not alter habitat choice by Arctic rodents

In systems where predation plays a key role in the dynamics of prey populations, such as in Arctic rodents, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, habitat selection by collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings depends on intra- and inter-specific densities, and there has been little agreement on the respective influences of food abundance, predators, and competition for habitat on lemming dynamics. Thus, we investigated whether predation affected selection of sedge-meadow versus upland tundra by collared lemmings in the central Canadian Arctic. We first controlled for the effects of competition on lemming habitat selection. We then searched for an additional signal of predation by comparing habitat selection patterns between 12 control plots and one large grid where lemmings were protected from predators by fencing in 1996 and 1997, but not during 5 subsequent years when we monitored habitat use in the grid as well as in the control plots. Dicrostonyx used upland preferentially over meadows and was more numerous in 1996 and 2011 than in other sample years. Lemmus was also more abundant in 1996 than in subsequent years, but its abundance was too low in the exclosure to assess whether exclusion of predators influenced its habitat selection. Contrary to the effects of competition, predation had a negligible impact on the spatial dynamics of Dicrostonyx, at least during summer. These results suggest that any differences in predation risk between the two habitats have little direct influence on the temporal dynamics of Dicrostonyx even if induced through predator–prey cycles.     

Predator density and the functional responses of coral reef fish

Predation is a key process driving coral reef fish population dynamics, with higher per capita prey mortality rates on reefs with more predators. Reef predators often forage together, and at high densities, they may either cooperate or antagonize one another, thereby causing prey mortality rates to be substantially higher or lower than one would expect if predators did not interact. However, we have a limited mechanistic understanding of how prey mortality rates change with predator densities. We re-analyzed a previously published observational dataset to investigate how the foraging response of the coney grouper (Cephalopholis fulva) feeding on the bluehead wrasse (Thalassoma bifasciatum) changed with shifts in predator and prey densities. Using a model-selection approach, we found that per-predator feeding rates were most consistent with a functional response that declines as predator density increases, suggesting either antagonistic interactions among predators or a shared antipredator behavioral response by the prey. Our findings suggest that variation in predator density (natural or anthropogenic) may have substantial consequences for coral reef fish population dynamics.     

Mixed encounters, limited perception and optimal foraging

This article demonstrates how perceptual constraints of predators and the possibility that predators encounter prey both sequentially (one prey type at a time) and simultaneously (two or more prey types at a time) may influence the predator attack decisions, diet composition and functional response of a behavioural predator-prey system. Individuals of a predator species are assumed to forage optimally on two prey types and to have exact knowledge of prey population numbers (or densities) only in a neighbourhood of their actual spatial location. The system characteristics are inspected by means of a discrete-time, discrete-space, individual-based model of the one-predator-two-prey interaction. Model predictions are compared with ones that have been obtained by assuming only sequential encounters of predators with prey and/or omniscient predators aware of prey population densities in the whole environment. It is shown that the zero-one prey choice rule, optimal for sequential encounters and omniscient predators, shifts to abruptly changing partial preferences for both prey types in the case of omniscient predators faced with both types of prey encounters. The latter, in turn, become gradually changing partial preferences when predator omniscience is considered only local.     

Seasonally varying marine influences on the coastal ecosystem detected through molecular gut analysis

Terrestrial predators on marine shores benefit from the inflow of organisms and matter from the marine ecosystem, often causing very high predator densities and indirectly affecting the abundance of other prey species on shores. This indirect effect may be particularly strong if predators shift diets between seasons. We therefore quantified the seasonal variation in diet of two wolf spider species that dominate the shoreline predator community, using molecular gut content analyses with general primers to detect the full prey range. Across the season, spider diets changed, with predominantly terrestrial prey from May until July and predominantly marine prey (mainly chironomids) from August until October. This pattern coincided with a change in the spider age and size structure, and prey abundance data and resource selection analyses suggest that the higher consumption of chironomids during autumn is due to an ontogenetic diet shift rather than to variation in prey abundance. The analyses suggested that small dipterans with a weak flight capacity, such as Chironomidae, Sphaeroceridae, Scatopsidae and Ephydridae, were overrepresented in the gut of small juvenile spiders during autumn, whereas larger, more robust prey, such as Lepidoptera, Anthomyidae and Dolichopodidae, were overrepresented in the diet of adult spiders during spring. The effect of the inflow may be that the survival and growth of juvenile spiders is higher in areas with high chironomid abundances, leading to higher densities of adult spiders and higher predation rates on the terrestrial prey next spring.     

Nonlinearities Lead to Qualitative Differences in Population Dynamics of Predator-Prey Systems

Since typically there are many predators feeding on most herbivores in natural communities, understanding multiple predator effects is critical for both community and applied ecology. Experiments of multiple predator effects on prey populations are extremely demanding, as the number of treatments and the amount of labour associated with these experiments increases exponentially with the number of species in question. Therefore, researchers tend to vary only presence/absence of the species and use only one (supposedly realistic) combination of their numbers in experiments. However, nonlinearities in density dependence, functional responses, interactions between natural enemies etc. are typical for such systems, and nonlinear models of population dynamics generally predict qualitatively different results, if initial absolute densities of the species studied differ, even if their relative densities are maintained. Therefore, testing combinations of natural enemies without varying their densities may not be sufficient. Here we test this prediction experimentally. We show that the population dynamics of a system consisting of 2 natural enemies (aphid predator Adalia bipunctata (L.), and aphid parasitoid, Aphidius colemani Viereck) and their shared prey (peach aphid, Myzus persicae Sulzer) are strongly affected by the absolute initial densities of the species in question. Even if their relative densities are kept constant, the natural enemy species or combination thereof that most effectively suppresses the prey may depend on the absolute initial densities used in the experiment. Future empirical studies of multiple predator – one prey interactions should therefore use a two-dimensional array of initial densities of the studied species. Varying only combinations of natural enemies without varying their densities is not sufficient and can lead to misleading results.