Spatial variation in mink and muskrat interactions in Canada

We investigated the spatial attributes of mink ( Mustela vison ) and muskrat ( Ondatra zibethicus ) interactions in Canada using 160 geographically paired historic time series of mink ( n =80) and muskrat ( n =80) harvest data obtained from Hudson's Bay Co. Archives. All series were 25 years in length (1925–1949) and were distributed primarily throughout five ecozones. We used autoregressive models and cross-correlation analysis to characterize the interactions between mink and muskrat. Model selection results did not differ among ecozones, and indicated that a predator-prey autoregressive model incorporating a delayed density-dependent term best described both the mink and muskrat harvest time series. Subsequent analysis of autoregressive coefficients and estimated lags indicated that mink and muskrat interactions vary throughout Canada. In western Canada, the trophic interactions appear to be strong, and mink population cycles lag behind muskrats 2–3 years. In central Canada, mink harvests lagged behind muskrats 1 year, and mink and muskrat interactions in central Canada, with the exception of the Hudson Plains ecozone, were intermediate. In eastern Canada, the trophic interactions appeared weakest, and there were no distinct time lags between mink and muskrat. Stronger interactions in western Canada may be a result of decreased prey diversity, forcing mink to specialize more on muskrats, whereas comparatively stronger perturbations stemming from other trophic interactions may alter the estimated interaction between mink and muskrat in eastern Canada.     

Size and scaling of predator-prey dynamics

We propose a scaled version of the Rosenzweig–MacArthur model using both Type I and Type II functional responses that incorporates the size dependence of interaction rates. Our aim is to link the energetic needs of organisms with the dynamics of interacting populations, for which survival is a result of a game-theoretic struggle for existence. We solve the scaled model of predator–prey dynamics and predict population level characteristics such as the scaling of coexistence size ranges and the optimal predator–prey size ratio. For a broad class of such models, the optimal predator–prey size ratio given available prey of a fixed size is constant. We also demonstrate how scaling predictions of prey density differ under resource limitation vs. predator drawdown. Finally, we show how evolution of predator size can destabilize population dynamics, compare scaling of predator–prey cycles to previous work, as well as discuss possible extensions of the model to multispecies communities.     

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.     

Predator–prey interactions under climate change: the importance of habitat vs body temperature

Habitat temperature is often assumed to serve as an effective proxy for organism body temperature when making predictions of species distributions under future climate change. However, the determinants of body temperature are complex, and organisms in identical microhabitats can occupy radically different thermal niches. This can have major implications of our understanding of how thermal stress modulates predator–prey relationships under field conditions. Using body temperature data from four different sites on Santa Cruz Island, California, we show that at two sites the body temperatures of a keystone predator (the seastar Pisaster ochraceus ) and its prey (the mussel Mytilus californianus ) followed very different trajectories, even though both animals occupied identical microhabitats. At the other two sites, body temperatures of predator and prey were closely coupled across a range of scales. The dynamical differences between predator and prey body temperatures depended on the location of pairs of sites, at the extremes of a persistent landscape-scale weather pattern observed across the island. Thus, the well understood predator–prey interaction between Pisaster and Mytilus cannot be predicted based on habitat-level information alone, as is now commonly attempted with landscape-level ('climate envelope') models.     

A synergistic trio of invasive mammals? Facilitative interactions among beavers,muskrats, and mink at the southern end of the Americas

With ecosystems increasingly having co-occurring invasive species, it is becoming more important to understand invasive species interactions. At the southern end of the Americas, American beavers (Castor canadensis), muskrats (Ondatra zibethicus), and American mink (Neovison vison), were independently introduced. We used generalized linear models to investigate how muskrat presence related to beaver-modified habitats on Navarino Island, Chile. We also investigated the trophic interactions of the mink with muskrats and beavers by studying mink diet. Additionally, we proposed a conceptual species interaction framework involving these invasive species on the new terrestrial community. Our results indicated a positive association between muskrat presence and beaver-modified habitats. Model average coefficients indicated that muskrats preferred beaver-modified freshwater ecosystems, compared to not dammed naturally flowing streams. In addition, mammals and fish represented the main prey items for mink. Although fish were mink’s dominant prey in marine coastal habitats, muskrats represented >50 % of the biomass of mink diet in inland environments. We propose that beavers affect river flow and native vegetation, changing forests into wetlands with abundant grasses and rush vegetation. Thus, beavers facilitate the existence of muskrats, which in turn sustain inland mink populations. The latter have major impacts on the native biota, especially on native birds and small rodents. The facilitative interactions among beavers, muskrats, and mink that we explored in this study, together with other non-native species, suggest that an invasive meltdown process may exist; however further research is needed to confirm this hypothesis. Finally, we propose a community-level management to conserve the biological integrity of native ecosystems.     

Individual‐based models of cod movement and population dynamics

Understanding the strength and diversity of predator‐prey interactions among species is essential to understand ecosystem consequences of population‐level variation. Directly quantifying the predatory behaviour of wild fishes at large spatial scales (>100 m) in the open sea is fraught with difficulties. To date the only empirical approach has been to search for correlations in the abundance of predators and their putative prey. As an example we use this approach to search for predators of the keystone crown‐of‐thorns starfish. We show that this approach is unlikely to detect predator–prey linkages because the theoretical relationship is non‐linear, resulting in multiple possible prey responses for single given predator abundance. Instead we suggest some indication of the strength and ecosystem importance of a predator–prey relationship can be gained by using the abundance of both predators and their putative prey to parameterize functional response models.     

Modelling the dynamic spatio-temporal response of predators to transient prey patches in the field

The spatio-temporal dynamics of two aphid species ( Metopolophium dirhodum and Sitobion avenae ) and a generalist predator ( Pterostichus melanarius ) were observed in a field-scale study using a grid of 256 sampling locations with a 12-m spacing. Using Spatial Analysis by Distance Indices we demonstrate that populations show ephemeral spatial pattern at the field scale. We observed a positive, lagged beetle response to this aphid pattern; conversely, the aphids displayed a negative, lagged response to beetle spatial pattern. Examination of the local structure of the spatio-temporal dynamics revealed a strong response by the beetle population to aphid patches. The temporal structure of spatial associations between the species shows a strong correspondence with those from a conceptual model of predator–prey spatial interaction. The spatially coupled dynamics were sufficiently strong for the predator to have a negative effect on the intrinsic rate of increase of their prey.     

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.     

Predator selectivity alters the effect of dispersal on coexistence among apparent competitors

While the majority of studies on dispersal effects on patterns of coexistence among species in a metacommunity have focused on resource competitors, dispersal in systems with predator–prey interactions may provide very different results. Here, we use an analytical model to study the effect of dispersal rates on coexistence of two prey species sharing a predator (apparent competition), when the traits of that predator vary. Specifically, we explore the range in immigration rates where apparent competitors are able to coexist, and how that range changes with predator selectivity and efficiency. We find that if the inferior apparent competitor has a higher probability of being consumed, it will require less immigration to invade and to exclude the superior prey as the predator becomes more opportunistic. However, if the inferior apparent competitor has a lower probability of being consumed (and lower growth rates), higher immigration is required for the inferior prey to invade and exclude the superior prey as the predator becomes more opportunistic. We further find that the largest range of immigration rates where prey coexist occurs when predator selectivity is intermediate (i.e. they do not show much bias towards consuming one species or the other). Increasing predator efficiency generally reduces the immigration rates necessary for the inferior apparent competitor to invade and exclude the superior apparent competitor, but also reduces the range of immigration rates where the two apparent competitors can coexist. However, when the superior apparent competitor has a higher probability of being consumed, increased predator efficiency can increase the range of parameters where the species can coexist. Our results are consistent with some of the variation observed in the effect of dispersal on prey species richness in empirical systems with top predators.     

From simple rules to cycling in community assembly

Simulation studies of community assembly have frequently observed two related phenomena: (1) the humpty dumpty effect in which communities can not be reconstructed by "sequential" invasions (i.e. single species invasions separated by long intervals of time) and (2) cycling between sub-communities. To better understand the mechanisms underlying these phenomena, we analyze a system consisting of two predators and two prey competing for a shared resource. We show how simple dominance rules (i.e. R^* and P^* rules) lead to cycling between sub-communities consisting of predator–prey pairs; predator and prey invasions alternatively lead to prey displacement via apparent competition and predator displacement via exploitative competition. We also show that these cycles are often dynamically unstable in the population phase space. More specifically, while for too slow invasion rates (i.e. "sequential" invasions) the system cycles indefinitely, faster invasion rates lead to coexistence of all species. In the later case, the assembly dynamics exhibit transient cycling between predator-prey subcommunities and the length of these transients decreases with the invasion rate and increases with habitat productivity.     

ECOLOGICAL CONDITIONS AFFECT EVOLUTIONARY TRAJECTORY IN A PREDATOR–PREY SYSTEM

The arms race of adaptation and counter adaptation in predator–prey interactions is a fascinating evolutionary dynamic with many consequences, including local adaptation and the promotion or maintenance of diversity. Although such antagonistic coevolution is suspected to be widespread in nature, experimental documentation of the process remains scant, and we have little understanding of the impact of ecological conditions. Here, we present evidence of predator–prey coevolution in a long-term experiment involving the predatory bacterium Bdellovibrio bacteriovorus and the prey Pseudomonas fluorescens , which has three morphs (SM, FS, and WS). Depending on experimentally applied disturbance regimes, the predator–prey system followed two distinct evolutionary trajectories, where the prey evolved to be either super-resistant to predation (SM morph) without counter-adaptation by the predator, or moderately resistant (FS morph), specialized to and coevolving with the predator. Although predation-resistant FS morphs suffer a cost of resistance, the evolution of extreme resistance to predation by the SM morph was apparently unconstrained by other traits (carrying capacity, growth rate). Thus we demonstrate empirically that ecological conditions can shape the evolutionary trajectory of a predator–prey system.     

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.     

Landscape heterogeneity shapes predation in a newly restored predator-prey system

Because some native ungulates have lived without top predators for generations, it has been uncertain whether runaway predation would occur when predators are newly restored to these systems. We show that landscape features and vegetation, which influence predator detection and capture of prey, shape large-scale patterns of predation in a newly restored predator–prey system. We analysed the spatial distribution of wolf ( Canis lupus ) predation on elk ( Cervus elaphus ) on the Northern Range of Yellowstone National Park over 10 consecutive winters. The influence of wolf distribution on kill sites diminished over the course of this study, a result that was likely caused by territorial constraints on wolf distribution. In contrast, landscape factors strongly influenced kill sites, creating distinct hunting grounds and prey refugia. Elk in this newly restored predator–prey system should be able to mediate their risk of predation by movement and habitat selection across a heterogeneous risk landscape.     

Spatial co-occurrence networks predict the feeding histories of polyphagous arthropod predators at field scales

In agricultural systems, polyphagous beetles and spiders are abundant components of the beneficial arthropod community. Although data on the dietary ranges of these groups is increasing, remarkably little is understood regarding how individuals interact with their prey at small spatial scales. We demonstrate the utility of a spatially-explicit network model that integrates predator behaviour using predator-prey co-occurrences. Three co-occurrence matrices, one each for June, July and August, were generated using Vortis suction sample data collected from an 80 point grid imposed on a field of winter wheat. Heuristic predator-prey linkages, based on positive spatial co-occurrence, were imposed on these three matrices to create networks. It was found that primary consumers were highly aggregated and showed a strong tendency to co-occur. This contrasted with patterns of predator–predator or predator–prey co-occurrences that either aggregated to their prey or were weak and more scattered. These patterns could not be explained by either competition for resources or body size differences. Procrustean methods indicated that the networks were temporally dynamic, consistently achieving rates of turnover >60%. A negative relationship was found between decreasing predator–prey co-occurrence in the network and the number of prey positives in the guts of those predators. For large polyphagous beetles, the closer they were to their prey at the field scale, the more likely they were to have eaten them. This simple underlying relationship suggests that spatial co-occurrence networks can be used to predict feeding behaviour and could make a valuable contribution to food web structuring.     

Temperature effects on chemical signalling in a predator–prey system

SUMMARY 1. We investigated the effect of temperature on chemical signalling in a predator–prey model system (planktivorous fish and Daphnia galeata ). Life-history changes in Daphnia in response to chemical cues (kairomones) derived from fish have become a paradigm for chemically induced anti-predator defences.
2. As temperature can affect both predator and prey, we carried out two experiments to disentangle these effects. In order to test for temperature effects on the predator, we kept prey at a single temperature and exposed them to kairomones from fish exposed to two different temperatures. Daphnia exhibited a higher intrinsic rate of population increase ( r ) when exposed to fish kairomones produced at high rather than low temperature. Assuming a positive correlation between r (because of an earlier maturation and/or increased clutch sizes) and kairomone concentration, our results suggest that kairomone production increases with rising temperature.
3. In the second experiment, to study the influence of temperature on the prey, Daphnia were kept at two different temperatures and exposed to fish kairomones produced at one constant temperature. We found no interaction between the effects of fish kairomone and temperature on Daphnia life history, suggesting that temperature does not directly alter life-history responses to fish kairomones.
4. Our results suggest that temperature influences Daphnia life history through its effects on fish kairomone concentration, but that temperature does not affect the strength of the response of Daphnia to the presence of fish.     

Species on the menu of a generalist predator, the eastern red bat (Lasiurus borealis): using a molecular approach to detect arthropod prey

One of the most difficult interactions to observe in nature is the relationship between a predator and its prey. When direct observations are impossible, we rely on morphological classification of prey remains, although this is particularly challenging among generalist predators whose faeces contain mixed and degraded prey fragments. In this investigation, we used a polymerase chain reaction and sequence-based technique to identify prey fragments in the guano of the generalist insectivore, the eastern red bat ( Lasiurus borealis ), and evaluate several hypotheses about prey selection and prey defences. The interaction between bats and insects is of significant evolutionary interest because of the adaptive nature of insect hearing against echolocation. However, measuring the successes of predator tactics or particular prey defences is limited because we cannot normally identify these digested prey fragments beyond order or family. Using a molecular approach, we recovered sequences from 89% of the fragments tested, and through comparison to a reference database of sequences, we were able to identify 127 different species of prey. Our results indicate that despite the robust jaws of L. borealis , most prey taxa were softer-bodied Lepidoptera. Surprisingly, more than 60% of the prey species were tympanate, with ears thought to afford protection against these echolocating bats. Moths of the family Arctiidae, which employ multiple defensive strategies, were not detected as a significant dietary component. Our results provide an unprecedented level of detail for the study of predator–prey relationships in bats and demonstrate the advantages which molecular tools can provide in investigations of complex ecological systems and food-web relationships.     

A simultaneous test of synchrony causal factors in muskrat and mink fur returns at different scales across Canada

BACKGROUND: Synchrony among populations has been attributed to three major hypotheses: dispersal, the Moran effect, and trophic-level interactions. Unfortunately, simultaneous testing of these hypotheses demands complete and detailed data, which are scarce for ecological systems. METHODOLOGY/PRINCIPAL FINDINGS: Hudson's Bay Company data on mink and muskrat fur returns in Canada represent an excellent opportunity to test these hypotheses because of the detailed spatial and temporal data from this predator-prey system. Using structural equation modelling, support for each hypothesis was evaluated at two spatial scales: across Canada and dividing the country into three regions longitudinally. Our results showed that at both scales mink synchrony is a major factor determining muskrat synchrony, supporting the hypothesis of trophic-level interactions, but the influence of winter precipitation synchrony is also important in eastern Canada. Moreover, mink synchrony is influenced principally by winter precipitation synchrony at the level of all Canada (Moran effect), but by distance at regional level, which might suggest some influence of dispersal at this level. DISCUSSION/SIGNIFICANCE: Our result is one of the few reports of synchrony mediated by trophic-level interactions, highlighting the importance of evaluation of scale effects in population synchrony studies.     

Predator‐Naïve Brown Trout (Salmo trutta) Show Antipredator Behaviours to Scent from an Introduced Piscivorous Mammalian Predator Fed Conspecifics

Introduced mammalian predators may pose a high risk for native and naïve prey populations, but little is known about how native fish species may recognize and respond to scents from introduced mammalian predators. We investigated the role of diet‐released chemical cues in facilitating predator recognition, hypothesizing that native brown trout (Salmo trutta) would exhibit antipredator behaviours to faeces scents from the introduced American mink (Neovision vison) fed conspecifics, but not to non‐trout diets. In treatments‐control and replicate stream tank experiments, brown trout showed significant antipredator responses to faeces scent from mink fed conspecifics, but not to faeces scent from mink fed a non‐trout diet (chicken), or the non‐predator food control, Eurasian beaver (Castor fiber). We conclude that native and naïve brown trout show relevant antipredator behaviours to an introduced mammalian predator, presumably based on diet‐released conspecific alarm cues and thereby estimate the predation risk.     

Risk induced by a native top predator reduces alien mink movements

1. Nonlethal predation effects may have stronger impacts on prey populations than direct predation impacts, and this should also apply to intraguild predation. The consequences of such interactions become especially important if invasive, and potentially destructive alien predators act as intraguild prey. 2. We studied the predation-risk impacts of a re-colonizing native top predator, Haliaeetus albicilla (white-tailed sea eagle), on the movements of Mustela vison (American mink), an alien predator in Europe. We radiocollared 20 mink in two study areas in the outer archipelago of the Baltic Sea, South-west Finland, during 2004 and 2005. In the archipelago, mink home ranges incorporate many islands, and mink are most predisposed to eagle predation while swimming between islands. Observed swimming distances of mink were compared to distances expected at random, and deviations from random swimming were explained by mink distance from nearest eagle nest, number of eagle observations near mink location, and mink home-range size. 3. Mink reduced their swimming distances with increasing sea eagle predation risk: for females, the reduction was 10% for an increase of 10 eagle observations, and 5% for each kilometre towards an eagle nest. Conclusions for males were restricted by their small sample size. 4. Our results suggest that female mink modify their behaviour according to eagle predation risk, which may reduce their population growth and have long-term cascading effects on lower trophic levels including bird, mammal and amphibian populations in the archipelago. Ecosystem restoration by bringing back the top predators may be one way of mitigating alien predator effects on native biota.     

Multi-behavioral strategies in a predator–prey game: an evolutionary algorithm analysis

Behavioral games between predators and prey often involve two sub-games: 'pre-encounter' games affecting the rate of encounter between predators and prey (e.g. predator–prey space games, Sih 2005 ), and 'post-encounter' games that influence the outcome of encounters (e.g. waiting games at prey refugia, Hugie 2003 , and games of vigilance, Brown et al. 1999 ). Most models, however, focus on only one or the other of these two sub-games.
I investigated a multi-behavioral game between predators and prey that integrated both pre-encounter and post-encounter behaviors. These behaviors included landscape-scale movements by predators and prey, a type of prey vigilance that increases immediately after an encounter and then decays over time ('ratcheting vigilance'), and predator management of prey vigilance. I analyzed the game using a computer-based evolutionary algorithm. This algorithm embedded an individual-based model of ecological interactions within a dynamic adaptive process of mutation and selection. I investigated how evolutionarily stable strategies (ESS) varied with the predators' learning ability, killing efficiency, density and rate of movement. I found that when predators learn prey location, random prey movement can be an ESS. Increased predator killing efficiency reduced prey movement, but only if the rate of predator movement was low. Predators countered ratcheting vigilance by delaying their follow-up attacks; however, this delay was reduced in the presence of additional predators. The interdependence of pre-and post-encounter behaviors revealed by the evolutionary algorithm suggests an intricate co-evolution of multi-behavioral predator–prey behavioral strategies.