Realized trophic niche driven by apparent competition: an example with marsupials

According to apparent competition theory, the co‐occurrence of two species that share the same predators appears to affect each other's population growth and abundance. However, due to habitat loss and over‐hunting, top predators are being made rare worldwide. Considering that apparent competitors share similar resources, we would expect the absence of top predators to reflect in changes on prey realized trophic niches. To test our hypothesis, we developed a model to predict the abundance ratio of apparent competitor species based on changes in their realized trophic niches. We tested our model against field data on the Neotropical marsupials Didelphis aurita and Metachirus nudicaudatus. Our results revealed that D. aurita and M. nudicaudatus are two species under apparent competition and their realized trophic niche and diet overlap change according to the presence of top predators. The model was able to predict the actual relative abundances of D. aurita and M. nudicaudatus in the three empirical studies analyzed. Our study presents quantitative support to the apparent competition theory; however, the model's applications to other groups still need to be verified. Additionally, our study shows that the lack of top predators has consequences on the realized trophic niche of their prey, and therefore, we reinforce that conservation plans need to focus on the effects of top predator loss on ecosystems.     

Does a top predator suppress the abundance of an invasive mesopredator at a continental scale?

Aim We examined evidence for the mesopredator release hypothesis at a subcontinental scale by investigating the relationship between indices of abundance of the dingo Canis lupus dingo (top‐order predator) and the invasive red fox Vulpes vulpes (mesopredator) in three large regions across mainland Australia. The red fox is known to be one of the major threats to the persistence of small and medium‐sized native vertebrates across the continent. Location Australia. Methods Indices of abundance were calculated from three independently collected datasets derived from bounty returns and field surveys. Data were analysed using univariate parametric, semi‐parametric and nonparametric techniques. Results Predator abundance indices did not conform to a normal distribution and the relationships between dingo and fox abundance indices were not well described by linear functions. Semi‐parametric and nonparametric techniques revealed consistently negative associations between indices of dingo and fox abundance. Main conclusions The results provide evidence that mesopredator suppression by a top predator can be exerted at very large geographical scales and suggest that relationships between the abundances of top predators and mesopredators are not linear. Our results have broad implications for the management of canid predators. First, they suggest that dingoes function ecologically to reduce the activity or abundance of red foxes and thus are likely to dampen the predatory impacts of foxes. More generally, they provide support for the notion that the mesopredator‐suppressive effects of top predators could be incorporated into broad‐scale biodiversity conservation programmes in many parts of the world by actively maintaining populations of top predators or restoring them in areas where they are now rare. Determining the population densities at which the interactions of top predators become ecologically effective will be a critical goal for conservation managers who aim to maintain or restore ecosystems using the ecological interactions of top predators.     

Does a top predator reduce the predatory impact of an invasive mesopredator on an endangered rodent?

The mesopredator release hypothesis (MRH) predicts that reduced abundance of top‐order predators results in an increase in the abundance of smaller predators (mesopredators) due to a reduction in intra‐guild predation and competition. The irruption of mesopredators that follows the removal of top‐order predators can have detrimental impacts on the prey of the mesopredators. Here we investigated the mechanisms via which the presence of a top‐order predator can benefit prey species. We tested predictions made according to the MRH and foraging theory by contrasting the abundances of an invasive mesopredator (red fox Vulpes vulpes) and an endangered prey species (dusky hopping mouse Notomys fuscus), predator diets, and N. fuscus foraging behaviour in the presence and absence of a top‐predator (dingo Canis lupus dingo). As predicted by the MRH, foxes were more abundant where dingoes were absent. Dietary overlap between sympatric dingoes and foxes was extensive, and fox was recorded in 1 dingo scat possibly indicating intra‐guild predation. Notomys fuscus were more likely to occur in fox scats than dingo scats and as predicted by the MRH N. fuscus were less abundant in the absence of dingoes. The population increase of N. fuscus following rainfall was dampened in the absence of dingoes suggesting that mesopredator release can attenuate bottom‐up effects, although it remains conceivable that differences in grazing regimes associated with dingo exclusion could have also influenced N. fuscus abundance. Notomys fuscus exhibited lower giving‐up densities in the presence of dingoes, consistent with the prediction that their perceived risk of predation would be lower and foraging efficiency greater in the presence of a top‐predator. Our results suggest that mesopredator suppression by a top predator can create a safer environment for prey species where the frequency of fatal encounters between predators and prey is reduced and the non‐consumptive effects of predators are lower.     

Occasional intraguild predation structuring small mammal assemblages: the marsupial Didelphis aurita in the Atlantic Forest of Brazil

The didelphid marsupial, Didelphis aurita, is suggested as an intraguild predator and as key‐species in small mammal assemblages of the Atlantic Forest of Brazil. The field experiments required to test this hypothesis are complex to implement, but the recent revival of regression methods offers a viable alternative. Here we use the dynamic and static regression methods to determine the importance of D. aurita as a competitor and intraguild predator. Capture–recapture data from two localities in the Rio de Janeiro State were used, Garrafão (municipality of Guapimirim), a coastal forest of the Serra do Mar, and Barra de Maricá, a costal sand dune vegetation. Population and microhabitat variables were monitored from April 1997 to April 2003 in Garrafão, and from January 1986 to July 1990 in Barra de Maricá. Microhabitat variables were related to Canopy, Plant, Litter and Rock covers, Obstruction from 0 to 1.5 m, and Number of logs. Exploitation competition was tested by the dynamic method, which models the effects of D. aurita on the per capita growth rate of a species. Interference by predation or competition was tested by the static method, where the abundance of D. aurita at trap stations was regressed against the abundance of other small mammals, after removal of any variation associated with microhabitat factors. Exploitation competition was not detected, but the interference of D. aurita was pervasive, affecting all small mammals studied in the two localities. The clear avoidance of D. aurita by all small mammals tested in two localities of different physiognomies indicates that it functions as an intraguild predator, even if actual predation by D. aurita is an occasional event.     

Ocelot Population Status in Protected Brazilian Atlantic Forest

Forest fragmentation and habitat loss are detrimental to top carnivores, such as jaguars (Panthera onca) and pumas (Puma concolor), but effects on mesocarnivores, such as ocelots (Leopardus pardalis), are less clear. Ocelots need native forests, but also might benefit from the local extirpation of larger cats such as pumas and jaguars through mesopredator release. We used a standardized camera trap protocol to assess ocelot populations in six protected areas of the Atlantic forest in southeastern Brazil where over 80% of forest remnants are < 50 ha. We tested whether variation in ocelot abundance could be explained by reserve size, forest cover, number of free-ranging domestic dogs and presence of top predators. Ocelot abundance was positively correlated with reserve size and the presence of top predators (jaguar and pumas) and negatively correlated with the number of dogs. We also found higher detection probabilities in less forested areas as compared to larger, intact forests. We suspect that smaller home ranges and higher movement rates in smaller, more degraded areas increased detection. Our data do not support the hypothesis of mesopredator release. Rather, our findings indicate that ocelots respond negatively to habitat loss, and thrive in large protected areas inhabited by top predators.     

Does lethal control of top‐predators release mesopredators? A re‐evaluation of three Australian case studies

Top‐predators can sometimes be important for structuring fauna assemblages in terrestrial ecosystems. Through a complex trophic cascade, the lethal control of top‐predators has been predicted to elicit positive population responses from mesopredators that may in turn increase predation pressure on prey species of concern. In support of this hypothesis, many relevant research papers, opinion pieces and literature reviews identify three particular case studies as supporting evidence for top‐predator control‐induced release of mesopredators in Australia. However, many fundamental details essential for supporting this hypothesis are missing from these case studies, which were each designed to investigate alternative aims. Here, we re‐evaluate the strength of evidence for top‐predator control‐induced mesopredator release from these three studies after comprehensive analyses of associated unpublished correlative and experimental data. Circumstantial evidence alluded to mesopredator releases of either the European Red Fox (Vulpes vulpes) or feral Cat (Felis catus) coinciding with Dingo (Canis lupus dingo) control in each case. Importantly, however, substantial limitations in predator population sampling techniques and/or experimental designs preclude strong assertions about the effect of lethal control on mesopredator populations from these studies. In all cases, multiple confounding factors and plausible alternative explanations for observed changes in predator populations exist. In accord with several critical reviews and a growing body of demonstrated experimental evidence on the subject, we conclude that there is an absence of reliable evidence for top‐predator control‐induced mesopredator release from these three case studies. Well‐designed and executed studies are critical for investigating potential top‐predator control‐induced mesopredator release.     

Trophic niche of the Pacific Sierra (Scomberomorus sierra) in the southeastern Gulf of California: Assessing its importance as a predator and prey (Mesopredator) in the food web

The identification of interspecific links (trophic niche) is important to characterize resource use of a predator, and to know its trophic role (for example, mesopredator or top predator) in the food web. In this study, we examined: a) the trophic ecology of Scomberomorus sierra as a predator (niche breadth, trophic overlap, and trophic position) and b) its presence as a prey in the diets of the region´s top predators, to evaluate the critical link of S. sierra as a probable mesopredator in the food web of the southeastern Gulf of California (GC). Based on %PSIRI, the dominant diet of S. sierra were engraulids and cephalopods. The diet was similar between sexes and among size-classes. However, the isotopic niche breadth values and δ¹⁵N variance (>1) reflect a broad niche for young adults likely related to a) changes in morphology (e.g., size of the mouth), b) development of the visual system, and c) changes in the energy requirements of the species reproductive stages. Seasonal changes in prey species’ availability and abundance resulted in isotopic variations, indicating that S. sierra is an opportunistic predator. A wide range in trophic position value (from 3.8 to 4.2) indicated that it also is an intermediary carnivore, with a high degree of trophic plasticity. Although S. sierra has not a dominant role in top predators’ diets, they share some prey species such as anchovies and other fish, depending on predator size. Therefore, S. sierra is a species with many prey-predator relationships in the southeastern GC food webs that may be considered a critical trophic link. This information is crucial for an ecosystem-based fisheries management in the Gulf of California.     

Optimal predator management for mountain sheep conservation depends on the strength of mesopredator release

Large predators often suppress ungulate population growth, but they may also suppress the abundance of smaller predators that prey on neonatal ungulates. Antagonistic interactions among predators may therefore need to be integrated into predator–prey models to effectively manage ungulate–predator systems. We present a modeling framework that examines the net impact of interacting predators on the population growth rate of shared prey, using interactions among wolves Canis lupus, coyotes Canis latrans and Dall sheep Ovis dalli dalli as a case study. Wolf control is currently employed on approximately 16 million ha in Alaska to increase the abundance of ungulates for human harvest. We hypothesized that the positive effects of wolf control on Dall sheep population growth could be counteracted by increased levels of predation by coyotes. Coyotes and Dall sheep adult females (ewes) and lambs were radiocollared in the Alaska Range from 1999–2005 to estimate fecundity, age‐specific survival rates, and causes of mortality in an area without wolf control. We used stage‐structured population models to simulate the net effect of wolf control on Dall sheep population growth (λ). Our models accounted for stage‐specific predation rates by wolves and coyotes, compensatory mortality, and the potential release of coyote populations due to wolf control. Wolves were the main predators of ewes, coyotes were the main predators of lambs, and wolves were the main source of mortality for coyotes. Population models predicted that wolf control could increase sheep λ by 4% per year in the absence of mesopredator release. However, if wolf control released coyote populations, our models predicted that sheep λ could decrease by up to 3% per year. These results highlight the importance of integrating antagonistic interactions among predators into predator–prey models, because the net effect of predator management on shared prey can depend critically on the strength of mesopredator release.     

Are Cecropia trees ecosystem engineers? The effect of decomposing Cecropia leaves on arthropod communities

Ecosystem engineers structure species richness and the composition of biological communities. Although several studies have uncovered the importance of engineering environments, few studies have evaluated the effect of pioneering plants as ecosystem engineers, especially in tropical environments. When dead, Cecropia leaves become architecturally complex, acquiring a tridimensional shape due to desiccation, and may facilitate other organisms. Here, we evaluate the role of these dead leaves in structuring species richness, abundance, biomass, and composition of macroinvertebrate communities on leaf litter in six protected areas of Brazilian Atlantic Rainforest. Predators were larger, more abundant, and presented higher standing stock in the presence of dead Cecropia leaves compared to soil debris (i.e., common leaf litter); however, detritivores had the opposite patterns. This resulted in shifts in body size structure of the assemblage, thus causing inversion of biomass pyramids to top‐heavy in advanced stages of Cecropia leaves desiccation. Dead Cecropia leaves did not influence species richness and abundance of species, but they influenced the biomass of detritivores and predators in the communities. Our results demonstrated that pioneer trees can act as ecosystem engineers, by facilitating communities of invertebrate predators. In addition, our results suggest that the presence of Cecropia leaves can mediate trophic interactions and shape food web structure on the forest floor. Abstract in Portuguese is available with online material.     

Life history of a neotropical marsupial: Evaluating potential contributions of survival and reproduction to population growth rate

The relative effect of survival and reproductive rates to population growth rate is expected to be similar across species with similar life-histories. We employed a matrix population model and sensitivity and elasticity analysis to assess the absolute and relative importance of age-specific survival and fertility to population growth rate of Didelphis aurita (Didelphimorphia, Didelphidae) in a rural area of Rio de Janeiro, southeastern Brazil. The results were compared to expectations for mammals that mature early and have short generation time, such as D. aurita. Prospective analysis showed that changes in pouch young and juveniles survival would have large effects on population growth rate, relative to other vital rates, being the most critical time periods in the life cycle of D. aurita, whereas the effect of fertilities were always low. These findings do not fit to the observed pattern in mammals that mature early, where reproductive parameters have the largest relative influence on population growth rate. Although reproductive rates were characterized by a relatively small influence on population growth rate, they are still relevant because of their high variability and response to potential environmental disturbances. The first application of matrix population models to a neotropical rainforest marsupial provides information on marsupial demography and life-history strategy, increasing comprehension of this unknown group.     

Keystone nonconsumptive effects within a diverse predator community

The number of prey killed by diverse predator communities is determined by complementarity and interference among predators, and by traits of particular predator species. However, it is less clear how predators' nonconsumptive effects (NCEs) scale with increasing predator biodiversity. We examined NCEs exerted on Culex mosquitoes by a diverse community of aquatic predators. In the field, mosquito larvae co‐occurred with differing densities and species compositions of mesopredator insects; top predator dragonfly naiads were present in roughly half of surveyed water bodies. We reproduced these predator community features in artificial ponds, exposing mosquito larvae to predator cues and measuring resulting effects on mosquito traits throughout development. Nonconsumptive effects of various combinations of mesopredator species reduced the survival of mosquito larvae to pupation, and reduced the size and longevity of adult mosquitoes that later emerged from the water. Intriguingly, adding single dragonfly naiads to ponds restored survivorship of larval mosquitoes to levels seen in the absence of predators, and further decreased adult mosquito longevity compared with mosquitoes emerging from mesopredator treatments. Behavioral observations revealed that mosquito larvae regularly deployed “diving” escape behavior in the presence of the mesopredators, but not when a dragonfly naiad was also present. This suggests that dragonflies may have relaxed NCEs of the mesopredators by causing mosquitoes to abandon energetically costly diving. Our study demonstrates that adding one individual of a functionally unique species can substantially alter community‐wide NCEs of predators on prey. For pathogen vectors like mosquitoes, this could in turn influence disease dynamics.     

Density‐dependent increase in superpredation linked to food limitation in a recovering population of northern goshawks Accipiter gentilis

A better understanding of the mechanisms driving superpredation, the killing of smaller mesopredators by larger apex predators, is important because of the crucial role superpredation can play in structuring communities and because it often involves species of conservation concern. Here we document how the extent of superpredation has changed over time, and assessed the impact of such temporal variation on local mesopredator populations using 40 yr of dietary data collected from a recovering population of northern goshawks Accipiter gentilis, an archetypical avian superpredator. We then assessed which mechanisms were driving variation in superpredation, e.g. was it opportunistic, a response to food becoming limited (due to declines in preferred prey) or to reduce competition. Raptors comprised 8% of goshawk diet on average in years when goshawk abundance was high, which is higher than reported elsewhere. Additionally, there was a per capita increase in superpredation as goshawks recovered, with the proportion of goshawk diet comprising raptors increasing from 2 to 8% as the number of goshawk home‐ranges increased from ≤ 14 to ≥ 25. This increase in superpredation coincided with a population decline in the most commonly killed mesopredator, the Eurasian kestrel Falco tinnunculus, which may represent the reversal of the ‘mesopredator release’ process (i.e. mesopredator suppression) which occurred after goshawks and other large raptors declined or were extirpated. Food limitation was the most likely driver of superpredation in this system given: 1) the substantial decline of two main prey groups in goshawk diet, the increase in diet diversity and decrease in goshawk reproductive success are all consistent with the goshawk population becoming food‐limited; 2) it's unlikely to be purely opportunistic as the increase in superpredation did not reflect changes in the availability of mesopredator species; and 3) the majority of mesopredators killed by goshawks do not compete with goshawks for food or nest sites.     

Apex predator suppression is linked to restructuring of ecosystems via multiple ecological pathways

Removal of apex predators can drive ecological regime shifts owing to compensatory positive and negative population level responses by organisms at lower trophic levels. Despite evidence that apex predators can influence ecosystems though multiple ecological pathways, most studies investigating apex predators’ effects on ecosystems have considered just one pathway in isolation. Here, we provide evidence that lethal control of an apex predator, the dingo Canis dingo, drives shifts in the structure of Australia's tropical‐savannah ecosystems. We compared mammal assemblages and understorey structure at seven paired‐sites. Each site comprised an area where people poisoned dingoes and an area without dingo control. The effects of dingo control on mammals scaled with body size. Where dingoes were poisoned, we found greater activity of herbivorous macropods and feral cats, a mesopredator, but sparser understorey vegetation and lower abundances of native rodents. Our study suggests that ecological cascades arising from apex predators’ suppressive effects on herbivores and mesopredators occur simultaneously. Concordant effects of dingo removal across tropical‐savannah, forest and desert biomes suggest that dingoes once exerted ubiquitous top–down effects across Australia and provides support for calls that top–down forcing should be considered a fundamental process governing ecosystem structure.     

Identifying mesopredator release in multi-predator systems: a review of evidence from North America

1. The mesopredator release hypothesis, defined as the change in distribution, abundance or behaviour of a middle-ranking predator in response to a decrease in density or distribution of an apex predator, is an increasingly popular topic in ecology. Terrestrial mesopredators have been reported as being released in multiple systems globally, particularly in North America, over the past century.
2. We reviewed 2687 scientific articles, of which we determined that 38 met our criteria for investigating mesopredator release (MR) in terrestrial North American mammalian predators.
3. We observed no support or mixed support for MR in 46% of all relevant studies, including conflicting evidence between measures (mesopredator distribution, abundance or behaviour) within a given study and among studies of the same community in different settings.
4. To advance the study of MR, we provide a conceptual framework that 1) highlights the multiple spatial, temporal and ecological scales at which mesopredator responses can occur; 2) suggests the relative weight of evidence for MR that is provided by measures of mesopredator responses at each scale; and 3) clearly defines the threshold for determining when MR is occurring.
5. In increasingly reshuffled predator communities with declining apex predators, there is a need for future studies to assess in more detail the contexts in which mesopredator behavioural responses scale up to the population-level processes and species-level distribution changes needed to identify these responses as MR.

     

Guardian or threat: does golden eagle predation risk have cascading effects on forest grouse?

Previous studies on intraguild predation have mainly focused on within-class assemblages, even though avian top predators may also influence mammalian mesopredator prey. By using nation-wide long-term data from Finland, northern Europe, we examined the impacts of golden eagles (Aquila chrysaetos) together with red foxes (Vulpes vulpes) and pine martens (Martes martes) on forest-dwelling herbivores, black grouse (Tetrao tetrix) and hazel grouse (Tetrastes bonasia). We hypothesized that eagles may alleviate the overall predation pressure on grouse by imposing intraguild predation risk on mesopredators. The predation impact of eagle was modelled using eagle density estimates and distance to eagle nest. Wildlife triangle counts were used as predation impact proxies of mammalian mesopredators and as measures of response in grouse. Our results show that eagle density correlated negatively with black grouse abundance indices while being positively associated with the proportion of juveniles in both grouse species, irrespective of the abundance of mesopredators. Yet, foxes and martens alone had a negative effect on the abundance indices and the proportion of young in the two grouse species. This suggests that the possible cascading effects of eagles are not mediated by decreased mesopredator numbers, but instead by fear effects. Alternatively, they may be mediated by other species than fox or marten studied here. In conclusion, we found support for the hypothesis that eagles provide protection for juvenile black and hazel grouse, whereas they are a threat for adult grouse. This important information helps us to better understand the role of avian top predators in terrestrial ecosystems.     

Sensory cues of a top‐predator indirectly control a reef fish mesopredator

Behavioural trophic cascades highlight the importance of indirect/risk effects in the maintenance of healthy trophic‐level links in complex ecosystems. However, there is limited understanding on how the loss of indirect top–down control can cascade through the food‐web to modify lower level predator–prey interactions. Using a reef fish food‐web, our study examines behavioural interactions among predators to assess how fear elicited by top‐predator cues (visual and chemical stimuli) can alter mesopredator behaviour and modify their interaction with resource prey. Under experimental conditions, the presence of any cue (visual, chemical, or both) from the top‐predator (coral trout Plectropomus leopardus) strongly restricted the distance swum, area explored and foraging activity of the mesopredator (dottyback Pseudochromis fuscus), while indirectly triggering a behavioural release of the resource prey (recruits of the damselfish Pomacentrus chrysurus). Interestingly, the presence of a large non‐predator species (thicklip wrasse Hemigymnus melapterus) also mediated the impact of the mesopredator on prey, as it provoked mesopredators to engage in an ‘inspection’ behaviour, while significantly reducing their feeding activity. Our study describes for the first time a three‐level behavioural cascade of coral reef fish and stresses the importance of indirect interactions in marine food‐webs.     

Ecological parameters of the leaf‐litter frog community of an Atlantic Rainforest area at Ilha Grande,Rio de Janeiro state,Brazil

Abstract The Atlantic Rainforest originally covered much of the Brazilian coast and is now reduced to approximately only 7% of its original area. Data on abundance distribution and microhabitat characteristics of anuran amphibians living on the forest floor leaf litter in the Atlantic Rainforest are scarce. In this study, we analysed the effect of litter depth and structure on the abundance and species richness of leaf‐litter frogs in an area of Atlantic Rainforest at Ilha Grande, Rio de Janeiro State, south‐eastern Brazil. We performed monthly samples (nocturnal and diurnal) from August 1996 to October 1997 using small (2 m × 1 m) plots. We sampled 234 plots, totalling 468 m² of forest leaf litter. We estimated leaf‐litter depth and the proportion of leaves in the plot and tested their effect on the total abundance of frogs and species richness using multiple regression analysis. We found 185 frogs from eight species: Brachycephalus (=Psyllophryne) didactylus (Izecksohn, 1971) (Brachycephalidae), Dendrophryniscus brevipollicatus Jiménez de la Espada 1871 (Bufonidae), Adenomera marmorata Steindachner 1867, Eleutherodactylus parvus (Girard 1853), Eleutherodactylus guentheri (Steindachner 1864), Eleutherodactylus binotatus (Spix 1824) and Zachaenus parvulus (Girard 1853) (Leptodactylidae), and Chiasmocleis sp. (Microhylidae). Brachycephalus didactylus was the most abundant species, with 91 individuals, whereas Dendrophryniscus brevipollicatus was the rarest, with two individuals. Mean litter depth and the proportion of leaves in the leaf litter were significantly related to frog abundance (R² = 0.17; F_2,107 = 10.779; P = 0.0001) and species richness (R² = 0.11; F_2,107 = 6.375; P = 0.002) indicating that microhabitat characteristics may affect local distribution and abundance of frogs in the forest floor.     

Song sparrows, top carnivores and nest predation: a test of the mesopredator release hypothesis

Ground-nesting North American landbirds have declined in the longterm, including species with a variety of migratory strategies. The mesopredator release hypothesis explains declines by suggesting that the virtual elimination of top carnivores (large-bodied canids and felids) from much of North America has “released” populations of nest-destroying mesopredators (i.e., medium-sized terrestrial omnivores such as the raccoon Procyon lotor). The hypothesis predicts (1) higher nest success in the presence than in the absence of top carnivores, and (2) a positive relationship between mesopredator abundance and nest predation. Results from a 4-year "natural experiment" in the agricultural landscape of southern Michigan tended to support prediction 1. When coyotes (Canis latrans) were absent in 1993 and present in 1994, 1995 and 1996, Mayfield nest survival in song sparrows (Melospiza melodia), a ground-nesting landbird, increased significantly during the same 4-year interval. Relative frequency of nest predation, the most common cause of nest failure, declined significantly over the four years. Coyotes may have decreased nest predation in 1994–1996 by reducing the abundance of raccoons, apparently the main nest predator in the study area. In an experiment testing prediction 2, mesopredator abundance and predation rate on artificial nests were positively related, as predicted by the hypothesis. Although the present study is not wholly conclusive by itself, we cautiously suggest it contributes to a growing body of evidence derived from a number of studies supporting the mesopredator release hypothesis. Received: 23 May 1997 / Accepted: 12 January 1998     

Higher nest predation risk in association with a top predator: mesopredator attraction?

Breeding close to top predators is a widespread reproductive strategy. Breeding animals may gain indirect benefits if proximity to top predators results in a reduction of predation due to suppression of mesopredators. We tested if passerine birds gain protection from mesopredators by nesting within territories of a top predator, the Ural owl (Strix uralensis). We placed nest boxes for pied flycatchers (Ficedula hypoleuca) in Ural owl nest sites and in control sites (currently unoccupied by owls). The nest boxes were designed so that nest predation risk could be altered (experimentally increased) after flycatcher settlement; we considered predation rate as a proxy of mesopredator abundance. Overall, we found higher nest predation rates in treatment than in control sites. Flycatcher laying date did not differ between sites, but smaller clutches were laid in treatment sites compared to controls, suggesting a response to perceived predation risk. Relative nest predation rate varied between years, being higher in owl nest sites in 2 years but similar in another; this variation might be indirectly influenced by vole abundance. Proximity to Ural owl nests might represent a risky habitat for passerines. High predation rates within owl territories could be because small mesopredators that do not directly threaten owl nests are attracted to owl nest sites. This could be explained if some mesopredators use owl territories to gain protection from their own predators, or if top predators and mesopredators independently seek similar habitats.     

Facilitative interactions among the pelagic community of temperate migratory terns,tunas and dolphins

We studied habitat and behavioral interactions among the marine community of top pelagic predators over the Atlantic continental shelf, observed from shipboard surveys off the northeastern United States. We hypothesized that foraging seabirds, specifically common terns Sterna hirundo and roseate terns S. dougallii, associate positively with the distribution and abundance of large, easily‐detected subsurface marine predators, as a result of facilitative interactions. Few rigorous tests examine the effect of interspecific interactions on seabird distributions, though many papers note the importance of environmental influences. Fewer, still, assess facilitation, defined as positive interactions among multiple taxa (birds, fish, mammals), where individuals use foraging neighbors for improved prey detection or enhanced prey availability. Our use of spatiotemporal‐structured Bayesian hierarchical models allowed us to test for fine‐scale associations of common and roseate terns with aggregations of tunas and cetaceans, and with standard oceanographic parameters. High tern abundance was linked to relatively high tuna densities and low dolphin densities, as well as high sea surface temperatures, shallow water, and proximity to shore. The fact that terns foraged when in the presence of tunas or relatively dense dolphin pods supports our hypothesis that the mechanism behind this spatial association involves positive interspecific interactions (attraction), a phenomenon that has been described but rarely quantified. This study reveals that community structure depends on static and dynamic covariates alike, ranging from bathymetry to the movement of other guild members. To improve the efficacy of predictive modeling, ecosystem approaches to applied conservation management should integrate not only habitat, but also behavior and community factors into analyses, especially in the case of marine spatial planning.