Trends in body condition of native piscivores following invasion of Lakes Erie and Ontario by the round goby

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Implications of shared predation for space use in two sympatric leporids

Spatial variation in habitat riskiness has a major influence on the predator–prey space race. However, the outcome of this race can be modulated if prey shares enemies with fellow prey (i.e., another prey species). Sharing of natural enemies may result in apparent competition, and its implications for prey space use remain poorly studied. Our objective was to test how prey species spend time among habitats that differ in riskiness, and how shared predation modulates the space use by prey species. We studied a one‐predator, two‐prey system in a coastal dune landscape in the Netherlands with the European hare (Lepus europaeus) and European rabbit (Oryctolagus cuniculus) as sympatric prey species and red fox (Vulpes vulpes) as their main predator. The fine‐scale space use by each species was quantified using camera traps. We quantified residence time as an index of space use. Hares and rabbits spent time differently among habitats that differ in riskiness. Space use by predators and habitat riskiness affected space use by hares more strongly than space use by rabbits. Residence time of hare was shorter in habitats in which the predator was efficient in searching or capturing prey species. However, hares spent more time in edge habitat when foxes were present, even though foxes are considered ambush predators. Shared predation affected the predator–prey space race for hares positively, and more strongly than the predator–prey space race for rabbits, which were not affected. Shared predation reversed the predator–prey space race between foxes and hares, whereas shared predation possibly also released a negative association and promoted a positive association between our two sympatric prey species. Habitat riskiness, species presence, and prey species’ escape mode and foraging mode (i.e., central‐place vs. noncentral‐place forager) affected the prey space race under shared predation.     

Body mass relationships affect the age structure of predation across carnivore–ungulate systems: a review and synthesis

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Diet of feral house cats Felis catus, ferrets Mustela furo and stoats M. erminea in grassland surrounding yellow-eyed penguin Megadyptes antipodes breeding areas, South Island, New Zealand

Spring and summer diets of feral house cats ( Felis catus ), ferrets ( Mustela furo ) and stoats ( M. erminea ) were studied in grassland surrounding breeding areas of yellow-eyed penguins ( Megadyptes antipodes ), a regionally threatened native species. All three predator species ate large numbers of young rabbits ( Oryctolagus cuniculus ) and birds. Stoats also relied heavily on mice ( Mus musculus ). Use of rabbits increased in rank order of increasing predator size, and male stoats ate more lagomorphs than female stoats. Diet differences may reflect character displacement as a result of exploitation competition, but interference competition or predation may force the smaller species to exploit micro-habitats with increased ground cover and consequent increased availability of smaller prey. Reduction of predation of native species like yellow-eyed penguins by decreasing or increasing staple mammal prey numbers of the introduced predators may provide lasting conservation benefits, but could also trigger diet changes that increase risk to endangered wildlife.     

Seasonal Effects of Habitat on Sources and Rates of Snowshoe Hare Predation in Alaskan Boreal Forests

Survival and predation of snowshoe hares (Lepus americanus) has been widely studied, yet there has been little quantification of the changes in vulnerability of hares to specific predators that may result from seasonal changes in vegetation and cover. We investigated survival and causes of mortalities of snowshoe hares during the late increase, peak, and decline of a population in interior Alaska. From June 2008 to May 2012, we radio-tagged 288 adult and older juvenile hares in early successional and black spruce (Picea mariana) forests and, using known-fate methods in program MARK, evaluated 85 survival models that included variables for sex, age, and body condition of hares, as well as trapping site, month, season, year, snowfall, snow depth, and air temperature. We compared the models using Akaike’s information criterion with correction for small sample size. Model results indicated that month, capture site, and body condition were the most important variables in explaining survival rates. Survival was highest in July, and more generally during summer, when alternative prey was available to predators of hares. Low survival rates coincided with molting periods, breeding activity in the spring, and the introduction of juveniles to the sample population in the fall. We identified predation as the cause of mortality in 86% of hare deaths. When the source of predation could be determined, hares were killed more often by goshawks (Accipiter gentilis) than other predators in early successional forest (30%), and more often by lynx (Lynx canadensis) than other predators in black spruce forest (31%). Great horned owls (Bubo virginianus) and coyotes (Canis latrans) represented smaller proportions of hare predation, and non-predatory causes were a minor source (3%) of mortality. Because hares rely on vegetative cover for concealment from predators, we measured cover in predation sites and habitats that the hares occupied and concluded that habitat type had a greater influence on the sources of predation than the amount of cover in any given location within a habitat. Our observations illustrate the vulnerability of hares to predators in even the densest coniferous habitat available in the boreal forest, and indicate strong seasonal changes in the rates and sources of predation.     

Predation on exotic zebra mussels by native fishes: effects on predator and prey

SUMMARY 1. Exotic zebra mussels, Dreissena polymorpha, occur in southern U.S. waterways in high densities, but little is known about the interaction between native fish predators and zebra mussels. Previous studies have suggested that exotic zebra mussels are low profitability prey items and native vertebrate predators are unlikely to reduce zebra mussel densities. We tested these hypotheses by observing prey use of fishes, determining energy content of primary prey species of fishes, and conducting predator exclusion experiments in Lake Dardanelle, Arkansas. 2. Zebra mussels were the primary prey eaten by 52.9% of blue catfish, Ictalurus furcatus; 48.2% of freshwater drum, Aplodinotus grunniens; and 100% of adult redear sunfish, Lepomis microlophus. Blue catfish showed distinct seasonal prey shifts, feeding on zebra mussels in summer and shad, Dorosoma spp., during winter. Energy content (joules g⁻¹) of blue catfish prey (threadfin shad, Dorosoma petenense; gizzard shad, D. cepedianum; zebra mussels; and asiatic clams, Corbicula fluminea) showed a significant species by season interaction, but shad were always significantly greater in energy content than bivalves examined as either ash-free dry mass or whole organism dry mass. Fish predators significantly reduced densities of large zebra mussels (>5 mm length) colonising clay tiles in the summers of 1997 and 1998, but predation effects on small zebra mussels (≤5 mm length) were less clear. 3. Freshwater drum and redear sunfish process bivalve prey by crushing shells and obtain low amounts of higher-energy food (only the flesh), whereas blue catfish lack a shell-crushing apparatus and ingest large amounts of low-energy food per unit time (bivalves with their shells). Blue catfish appeared to select the abundant zebra mussel over the more energetically rich shad during summer, then shifted to shad during winter when shad experienced temperature-dependent stress and mortality. Native fish predators can suppress adult zebra mussel colonisation, but are ultimately unlikely to limit population density because of zebra mussel reproductive potential.     

Predation against birds with low immunocompetence

Differences in the phenotypic characteristics between individuals falling prey to predators and conspecifics avoiding predation will reflect the intensity of selection on prey. If prey are generally in poor condition, we predicted that they should have an inferior health status in comparison to individuals dying for other reasons. We investigated this prediction for prey and conspecifics that did not die from predation by comparing the size of the spleen, which is an important immune defence organ reflecting one component of immunocompetence, using 18 species of passerine birds and domestic cat Felis catus predators as a model system. Prey had consistently smaller spleens than non-prey, implying that they had weak immune systems. The data set did not indicate that sex or age, month of death, body mass, body condition, liver mass, wing length or tarsus length differed significantly between prey and non-prey. Thus there was little evidence of confounding factors affecting the results. These observations indirectly suggest that disease and parasitism may play an important role in predator-prey interactions. Received: 4 August 1999 / Accepted: 7 October 1999     

Toad’s tongue for breakfast: exploitation of a novel prey type,the invasive cane toad,by scavenging raptors in tropical Australia

Although interest in the ecological impacts of invasive species has largely focused on negative effects, some native taxa may benefit from invader arrival. In tropical Australia, invasive cane toads (Bufo marinus) have fatally poisoned many native predators (e.g., marsupials, crocodiles, lizards) that attempt to ingest the toxic anurans, but birds appear to be more resistant to toad toxins. We quantified offtake of dead (road-killed) cane toads by raptors (black kites (Milvus migrans) and whistling kites (Haliastur sphenurus)) at a site near Darwin, in the Australian wet-dry tropics. Raptors readily took dead toads, especially small ones, although native frogs were preferred to toads if available. More carcasses were removed in the dry season than the wet season, perhaps reflecting seasonal availability of alternative prey. Raptors appeared to recognize and avoid bufotoxins, and typically removed and consumed only the toads’ tongues (thereby minimizing toxin uptake). The invasion of cane toads thus constitutes a novel prey type for scavenging raptors, rather than (as is the case for many other native predators) a threat to population viability.     

Predator-Mediated Indirect Effects of Snowshoe Hares on Dall's Sheep in Alaska

ABSTRACT Indirect interactions among species can strongly influence population dynamics and community structure but are often overlooked in management of large mammals. We estimated survival of Dall's sheep (Ovis dalli) in the central Alaska Range, USA, during years of differing snowshoe hare (Lepus americanus) abundance to test whether indirect interactions with a cyclic hare population affect Dall's sheep either negatively, by subsidizing predators (apparent competition), or positively, by diverting predation (apparent commensalism). Annual survival of adult female sheep was consistently high (0.85 for all yr and age classes combined). In contrast, annual estimates of lamb survival ranged from 0.15 to 0.63. The main predators of lambs were coyotes (Canis latrans) and golden eagles (Aquila chrysaetos), which rely on hares as their primary food and prey on lambs secondarily. Coyotes and eagles killed 78% of 65 radiocollared lambs for which cause of death was known. Lamb survival was negatively related to hare abundance during the previous year, and lamb survival rates more than doubled when hare abundance declined, supporting the hypothesis of predator-mediated apparent competition between hares and sheep. However, stage-specific predation and delays in predator responses to changes in hare numbers led to a positive relationship between abundance of adult Dall's sheep and hares. Lacking reliable estimates of survival, a manager might erroneously conclude that hares benefit sheep. Thus, support for different indirect effects can be obtained from different types of data, which demonstrates the need to determine the mechanisms that create indirect interactions. Long-term survey data suggest that predation by coyotes is limiting this sheep population below levels typical when coyotes were rare or absent. Understanding the nature of indirect interactions is necessary to effectively manage complex predator–prey communities.     

Predation of migratory Little Stint (<Emphasis Type="Italic">Calidris minuta</Emphasis>) by Barbary Falcon (<Emphasis Type="Italic">Falco pelegrinoides</Emphasis>) is dependent on body mass and duration of stopover time

Shorebird (Charadriiformes) migration phenology is critically synchronised with prey availability at traditional staging sites that allows the birds to stage and complete the migrations. These stopovers, usually in large concentrations, make the migrants vulnerable to local predators. The body mass gained or maintained is considered to be the result of the trade-off between the risks of starvation and predation. Theoretically, heavier birds should be less adept at escaping predators, and experimental evidence showed that flight performance is impaired in heavier birds. During three migration seasons we searched pellets and prey remains taken by a pair of Barbary Falcons (Falco pelegrinoides). We discovered that many pellets contained rings from the ringing program in Eilat, Israel, and the overwhelming majority belonged to Little Stints (Calidris minuta). We checked if the body mass and length of the stopover as expressed by ringing status of the Little Stints in a particular migration season was related to the risk of predation by Barbary Falcons. We found the chance that a heavier bird would be predated was lower than that of a lighter individual, and that Stints retrapped during the same migration season were significantly more endangered by predation than those recorded only once.     

Influence of intra‐ and interspecific variation in predator–prey body size ratios on trophic interaction strengths

1. Predation is a pervasive force that structures food webs and directly influences ecosystem functioning. The relative body sizes of predators and prey may be an important determinant of interaction strengths. However, studies quantifying the combined influence of intra‐ and interspecific variation in predator–prey body size ratios are lacking.
2. We use a comparative functional response approach to examine interaction strengths between three size classes of invasive bluegill and largemouth bass toward three scaled size classes of their tilapia prey. We then quantify the influence of intra‐ and interspecific predator–prey body mass ratios on the scaling of attack rates and handling times.
3. Type II functional responses were displayed by both predators across all predator and prey size classes. Largemouth bass consumed more than bluegill at small and intermediate predator size classes, while large predators of both species were more similar. Small prey were most vulnerable overall; however, differential attack rates among prey were emergent across predator sizes. For both bluegill and largemouth bass, small predators exhibited higher attack rates toward small and intermediate prey sizes, while larger predators exhibited greater attack rates toward large prey. Conversely, handling times increased with prey size, with small bluegill exhibiting particularly low feeding rates toward medium–large prey types. Attack rates for both predators peaked unimodally at intermediate predator–prey body mass ratios, while handling times generally shortened across increasing body mass ratios.
4. We thus demonstrate effects of body size ratios on predator–prey interaction strengths between key fish species, with attack rates and handling times dependent on the relative sizes of predator–prey participants.
5. Considerations for intra‐ and interspecific body size ratio effects are critical for predicting the strengths of interactions within ecosystems and may drive differential ecological impacts among invasive species as size ratios shift.

     

From process to pattern: how fluctuating predation risk impacts the stress axis of snowshoe hares during the 10-year cycle

Predation is a central organizing process affecting populations and communities. Traditionally, ecologists have focused on the direct effects of predation—the killing of prey. However, predators also have significant sublethal effects on prey populations. We investigated how fluctuating predation risk affected the stress physiology of a cyclic population of snowshoe hares (Lepus americanus) in the Yukon, finding that they are extremely sensitive to the fluctuating risk of predation. In years of high predator numbers, hares had greater plasma cortisol levels at capture, greater fecal cortisol metabolite levels, a greater plasma cortisol response to a hormone challenge, a greater ability to mobilize energy and poorer body condition. These indices of stress had the same pattern within years, during the winter and over the breeding season when the hare:lynx ratio was lowest and the food availability the worst. Previously we have shown that predator-induced maternal stress lowers reproduction and compromises offspring’s stress axis. We propose that predator-induced changes in hare stress physiology affect their demography through negative impacts on reproduction and that the low phase of cyclic populations may be the result of predator-induced maternal stress reducing the fitness of progeny. The hare population cycle has far reaching ramifications on predators, alternate prey, and vegetation. Thus, predation is the predominant organizing process for much of the North American boreal forest community, with its indirect signature—stress in hares—producing a pattern of hormonal changes that provides a sensitive reflection of fluctuating predator pressure that may have long-term demographic consequences.     

Ontogenetic variations in the type and size of prey consumed by juvenile coho,Oncorhynchus kisutch,and chinook,O. tshawytscha,salmon

Synopsis Stomach contents of juvenile coho,Oncorhynchus kisutch, and chinook,O. tshawytscha, salmon collected in purse seines off the coast of Washington and Oregon were examined for variations related to predator size. There was a general trend toward increasing consumption of fish with increasing body size, due mainly to the increase in northern anchovy biomass consumed by the larger salmon. Most of the major prey taxa showed significant differences among the size classes examined for both salmon species. There was a direct relationship between predator and prey size for both coho and chinook, but considerable variation was found in prey length consumed within each size class. Prey width did not provide as good a fit as prey length for either species. In general, coho consumed larger fish prey in relation to their body length than chinook but there were substantial differences by month or year of collection.     

Biogeographical patterns in the diet of an opportunistic predator: the red fox Vulpes vulpes in the Iberian Peninsula

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Safety in numbers for secondary prey populations: an experimental test using egg predation by small mammals in New Zealand

New Zealand's native avifauna is threatened by introduced mammalian predators. Native species are often not the primary prey of these predators, which depend on introduced mice and rabbits as their primary food source. Theoretical models predict that predation risk for a subsidiary, or "secondary" prey species is inversely proportional to its population size. This prediction was tested by a quasi-natural experiment in which four different sized prey "colonies" were constructed at four existing sooty shearwater breeding sites. Domestic hens' eggs were placed in shearwater burrows immediately following the shearwater breeding season and egg predation rates monitored at five, ten and fifteen days. Treatments were switched between sites and the experiment run for a second time after a two-week stand-down period. The net effect of increasing colony size was to lower individual risk of predation. The larger number of individuals present served to effectively "buffer," or dilute, per-capita predation risk from predators whose numbers are fixed by extraneous factors: chiefly the abundance of their primary prey. Although eggs were removed more slowly from smaller colonies than from larger ones, each loss had a greater per-capita effect on individual mortality risk. The inverse density dependent relationship found between colony size and predation risk implies that predator population dynamics are largely independent of secondary prey numbers. Abundant introduced predators can therefore easily drive a small secondary prey population to extinction. Control of primary prey populations may be an important management tool in these circumstances.     

Predation and avoidance behaviour in aphid‐ladybird interactions of native and invasive ladybirds in Europe

1. While detrimental effects of invasive predators on native species are well documented, we often lack a mechanistic understanding of the invasion success. Lack of prey avoidance behaviour can lead to higher consumption rates by invasive predators compared to native predators. This competitive advantage is expected to contribute to the invasion success of non‐native predators.
2. We compared aphid consumption and cue avoidance behaviour of aphids between four native ladybird species (Coccinella septempunctata, Adalia bipunctata, Propylea quatuordecimpunctata, and Hippodamia variegata) and the invasive Asian ladybird Harmonia axyridis.
3. The invasive H. axyridis and the native C. septempunctata consumed more aphids than the three smaller native ladybird species. In line with our expectations, aphids avoided leaves bearing cues of most native ladybird species but not of the invasive H. axyridis.
4. Our results indicate that body size rather than ladybird origin determined aphid predation rates. The lack of aphid avoidance behaviour towards cues of H. axyridis indicates that they were not able to recognise the chemical cues of the invasive predator.
5. Relatively large body size and the absence of cue avoidance in aphids might benefit the invasive H. axyridis, particularly in comparison to smaller native ladybird species. The absence of avoidance behaviour in aphids might lead to even higher predation rates of H. axyridis under more natural conditions.

     

Dietary complexity and hidden costs of prey switching in a generalist top predator

1. Variation in predator diet is a critical aspect of food web stability, health, and population dynamics of predator/ prey communities. Quantifying diet, particularly among cryptic species, is extremely challenging, however, and differentiation between demographic subsets of populations is often overlooked.
2. We used prey remains and data taken postmortem from otter Lutra lutra to determine the extent to which dietary variation in a top predator was associated with biotic, spatial, and temporal factors.
3. Biotic data (e.g., sex, weight, and length) and stomach contents were taken from 610 otters found dead across England and Wales between 1994 and 2010. Prey remains were identified to species where possible, using published keys and reference materials. Multi‐model inference followed by model prediction was applied to test for and visualize the nature of associations.
4. Evidence for widespread decline in the consumption of eels (Anguilla anguilla) reflected known eel population declines. An association between eel consumption and otter body condition suggested negative consequences for otter nutrition. Consumption of Cottus gobio and stickleback spp. increased, but was unlikely to compensate (there was no association with body condition). More otters with empty stomachs were found over time. Otter sex, body length, and age‐class were important biotic predictors of the prey species found, and season, region, and distance from the coast were important abiotic predictors.
5. Our study is unique in its multivariate nature, broad spatial scale, and long‐term dataset. Inclusion of biotic data allowed us to reveal important differences in costs and benefits of different prey types, and differences between demographic subsets of the population, overlaid on spatial and temporal variation. Such complexities in otter diet are likely to be paralleled in other predators, and detailed characterization of diet should not be overlooked in efforts to conserve wild populations.

     

Phenotypic variation of an alien species in a new environment: the body size and diet of American mink over time and at local and continental scales

Introduced species must adapt their ecology, behaviour, and morphological traits to new conditions. The successful introduction and invasive potential of a species are related to its levels of phenotypic plasticity and genetic polymorphism. We analysed changes in the body mass and length of American mink (Neovison vison) since its introduction into the Warta Mouth National Park, western Poland, in relation to diet composition and colonization progress from 1996 to 2004. Mink body mass decreased significantly during the period of population establishment within the study area, with an average decrease of 13% from 1.36 to 1.18 kg in males and of 16% from 0.83 to 0.70 kg in females. Diet composition varied seasonally and between consecutive years. The main prey items were mammals and fish in the cold season and birds and fish in the warm season. During the study period the proportion of mammals preyed upon increased in the cold season and decreased in the warm season. The proportion of birds preyed upon decreased over the study period, whereas the proportion of fish increased. Following introduction, the strictly aquatic portion of mink diet (fish and frogs) increased over time, whereas the proportion of large prey (large birds, muskrats, and water voles) decreased. The average yearly proportion of large prey and average‐sized prey in the mink diet was significantly correlated with the mean body masses of males and females. Biogeographical variation in the body mass and length of mink was best explained by the percentage of large prey in the mink diet in both sexes, and by latitude for females. Together these results demonstrate that American mink rapidly changed their body mass in relation to local conditions. This phenotypic variability may be underpinned by phenotypic plasticity and/or by adaptation of quantitative genetic variation. The potential to rapidly change phenotypic variation in this manner is an important factor determining the negative ecological impacts of invasive species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 681–693.     

Diet of Montagu's Harrier Circus pygargus in central Spain: analysis of temporal and geographic variation

The diet of Montagu's Harrier Circus pygargus was studied for 4 years near Madrid, Spain, with the use of pellets and prey remains collected during the breeding season. The diet of the study birds included a wide variety of prey categories, the most important one by weight being lagomorphs (mainly Hares Lepus granatensis ). The diet varied seasonally, with Hares predominating early in the breeding season, while birds increased in importance later. The shift was partly a result of variation in availability (with fledgling birds being more abundant late in the season), but statistical analysis suggested that lagomorphs and then birds were actively chosen in the prelaying and nestling periods, respectively. Montagu's Harrier seemed to be an opportunistic species, hunting a wider range of prey types in more southern latitudes, but specialized in each region in the prey species that was most profitable. This was the only study on Montagu's Harriers to date which found lagomorphs to be the most important part of the diet throughout the season.