Relaxed predation theory: size,sex and brains matter

Australia's wildlife is being considerably impacted by introduced mammalian predators such as cats (Felis catus), dogs (Canis lupus familiaris), and foxes (Vulpes vulpes). This is often attributed to native wildlife being naïve to these introduced predators. A systematic review of the literature reveals that native metatherians (body mass range 0.02–25 kg) do not recognise, and show relaxed antipredator behaviours towards, native and some introduced mammalian predators. Native eutherians (all with body mass < 2 kg), however, do appear to recognise and exhibit antipredator behaviours towards both native and introduced predators. Based on our findings, we propose a novel theory, the ‘Relaxed Predation Theory’. Our new theory is based on the absence of large mammalian predators leading to reduced predation pressure in Australia during the past 40000–50000 years, and on three key differences between Australian metatherians and eutherians: size, sex, and brains. In light of this Relaxed Predation Theory, we make a number of recommendations for the conservation of Australian wildlife: (i) predator avoidance training of suitable species; (ii) exclusion fencing to exclude some, but not all, predators to facilitate the development of antipredator behaviours; (iii) captive breeding programs to prevent the extinction of some species; and (iv) reintroduction of Australia's larger predators, potentially to compete with and displace introduced predators. A more detailed understanding of the responses of Australian mammals to predators will hopefully contribute to the improved conservation of susceptible species.     

A native apex predator limits an invasive mesopredator and protects native prey: Tasmanian devils protecting bandicoots from cats

Apex predators can limit the abundance and behaviour of mesopredators, thereby reducing predation on smaller species. We know less about whether native apex predators are effective in suppressing invasive mesopredators, a major global driver of vertebrate extinctions. We use the severe disease‐induced decline of an apex predator, the Tasmanian devil, as a natural experiment to test whether devils limit abundance of invasive feral cats and in turn protect smaller native prey. Cat abundance was c. 58% higher where devils had declined, which in turn negatively affected a smaller native prey species. Devils had a stronger limiting effect on cats than on a native mesopredator, suggesting apex predators may have stronger suppressive effects on evolutionarily naive species than coevolved species. Our results highlight how disease in one species can affect the broader ecosystem. We show that apex predators not only regulate native species but can also confer resistance to the impacts of invasive populations. Apex predators could therefore be a powerful but underutilised tool to prevent biodiversity loss.     

Morphological and life‐history responses of anurans to predation by an invasive crayfish: an integrative approach

Predator‐induced phenotypic plasticity has been widely documented in response to native predators, but studies examining the extent to which prey can respond to exotic invasive predators are scarce. As native prey often do not share a long evolutionary history with invasive predators, they may lack defenses against them. This can lead to population declines and even extinctions, making exotic predators a serious threat to biodiversity. Here, in a community‐wide study, we examined the morphological and life‐history responses of anuran larvae reared with the invasive red swamp crayfish, Procambarus clarkii, feeding on conspecific tadpoles. We reared tadpoles of nine species until metamorphosis and examined responses in terms of larval morphology, growth, and development, as well as their degree of phenotypic integration. These responses were compared with the ones developed in the presence of a native predator, the larval dragonfly Aeshna sp., also feeding on tadpoles. Eight of the nine species altered their morphology or life history when reared with the fed dragonfly, but only four when reared with the fed crayfish, suggesting among‐species variation in the ability to respond to a novel predator. While morphological defenses were generally similar across species (deeper tails) and almost exclusively elicited in the presence of the fed dragonfly, life‐history responses were very variable and commonly elicited in the presence of the invasive crayfish. Phenotypes induced in the presence of dragonfly were more integrated than in crayfish presence. The lack of response to the presence of the fed crayfish in five of the study species suggests higher risk of local extinction and ultimately reduced diversity of the invaded amphibian communities. Understanding how native prey species vary in their responses to invasive predators is important in predicting the impacts caused by newly established predator–prey interactions following biological invasions.     

A review on the effects of alien rodents in the Balearic (Western Mediterranean Sea) and Canary Islands (Eastern Atlantic Ocean)

Invasions of alien rodents have shown to have devastating effects on insular ecosystems. Here we review the ecological impacts of these species on the biodiversity of the Balearic and the Canary Islands. A total of seven species of introduced rodents (two rats, three mice, one dormouse, and one squirrel) have been recorded (six in the Balearics and four in the Canaries). Some of them can occasionally be important predators of nesting seabirds, contributing to the decline of endangered populations in both archipelagos. Rats are also known to prey upon terrestrial birds, such as the two endemic Canarian pigeons. Furthermore, rats actively consume both vegetative and reproductive tissues of a high number of plants, with potential relevant indirect effects on vegetation by increasing erosion and favoring the establishment of alien plants. In the Balearics, rats and mice are important seed predators of endemic species and of some plants with a restricted distribution. In the Canaries, rats intensively prey upon about half of the fleshy-fruited tree species of the laurel forest, including some endemics. In both archipelagos, alien rodents disrupt native plant–seed dispersal mutualisms, potentially reducing the chances of plant recruitment at the same time that they modify the structure of plant communities. We further suggest that alien rodents played (and play) a key role in the past and present transformation of Balearic and Canarian native ecosystems.     

Your worst enemy could be your best friend: predator contributions to invasion resistance and persistence of natives

Native predators are postulated to have an important role in biotic resistance of communities to invasion and community resilience. Effects of predators can be complex, and mechanisms by which predators affect invasion success and impact are understood for only a few well-studied communities. We tested experimentally whether a native predator limits an invasive species’ success and impact on a native competitor for a community of aquatic insect larvae in water-filled containers. The native mosquito Aedes triseriatus alone had no significant effect on abundance of the invasive mosquito Aedes albopictus. The native predatory midge Corethrella appendiculata, at low or high density, significantly reduced A. albopictus abundance. This effect was not caused by trait-mediated oviposition avoidance of containers with predators, but instead was a density-mediated effect caused by predator-induced mortality. The presence of this predator significantly reduced survivorship of the native species, but high predator density also significantly increased development rate of the native species when the invader was present, consistent with predator-mediated release from interspecific competition with the invader. Thus, a native predator can indirectly benefit its native prey when a superior competitor invades. This shows the importance of native predators as a component of biodiversity for both biotic resistance to invasion and resilience of a community perturbed by successful invasion.     

Exploratory behaviour and novel predator recognition: behavioural correlations across contexts

It was hypothesized that the exploratory behaviour of an individual measured in a novel environment could predict its behaviour in response to a novel predator. This study examined novel predator recognition in the western mosquitofish Gambusia affinis, a species with individual differences in risk‐taking, activity and exploration in novel environments. Prey responded with characteristic shoaling and avoidance in response to native predators, but did not show characteristic antipredator behaviour towards novel predators. Furthermore, G. affinis exhibited individual‐level behavioural correlations across contexts but only when prey were tested with native predators. This could be the result of native predatory selection on behavioural correlations in the prey species.     

Biodiversity in tropical agroforests and the ecological role of ants and ant diversity in predatory function

Abstract.  1. Intensive agricultural practices drive biodiversity loss with potentially drastic consequences for ecosystem services. To advance conservation and production goals, agricultural practices should be compatible with biodiversity. Traditional or less intensive systems (i.e. with fewer agrochemicals, less mechanisation, more crop species) such as shaded coffee and cacao agroforests are highlighted for their ability to provide a refuge for biodiversity and may also enhance certain ecosystem functions (i.e. predation).
2. Ants are an important predator group in tropical agroforestry systems. Generally, ant biodiversity declines with coffee and cacao intensification yet the literature lacks a summary of the known mechanisms for ant declines and how this diversity loss may affect the role of ants as predators.
3. Here, how shaded coffee and cacao agroforestry systems protect biodiversity and may preserve related ecosystem functions is discussed in the context of ants as predators. Specifically, the relationships between biodiversity and predation, links between agriculture and conservation, patterns and mechanisms for ant diversity loss with agricultural intensification, importance of ants as control agents of pests and fungal diseases, and whether ant diversity may influence the functional role of ants as predators are addressed. Furthermore, because of the importance of homopteran-tending by ants in the ecological and agricultural literature, as well as to the success of ants as predators, the costs and benefits of promoting ants in agroforests are discussed.
4. Especially where the diversity of ants and other predators is high, as in traditional agroforestry systems, both agroecosystem function and conservation goals will be advanced by biodiversity protection.     

Antipredator Behavioral Responses of Native and Exotic Tadpoles to Novel Predator

Factors related to the invasion process, such as high abundance of invaders, residence time, and functional distinctiveness, are well documented, but less attention has been given to the effects of antipredator strategy of invasive species during colonization. In this study, we explored the antipredator strategy of an introduced species by comparing the predator avoidance behaviors of two native anuran species and one introduced("exotic") species in the presence of different predators. The two native anuran species used in the study were Black-spotted Pond Frog Rana nigromaculata and Terrestrial Frog Rana limnocharis. The introduced(invasive) species used was American bullfrog Lithobates catesbeianus. Chinese pond turtle Chinemys reevesii, Red-backed rat snake Elaphe rufodorsata, and Big-headed turtle Platysternon megacephalum were used as predator species. Chinese pond turtles and Red-backed rat snakes are native predators of Black-spotted Pond Frogs and Terrestrial Frogs, while Big-headed turtles are novel("unfamiliar") to the two frogs. All three predator species are novel("unfamiliar") to the American bullfrog. The results show that tadpoles of the two native species displayed behaviors of recognizing the two native predators, but did not display the capability of identifying the novel predator. Results from our study also suggest that American bullfrog tadpoles exhibited strong antipredator behavioral responses by displaying the capability of identifying "unfamiliar" predators without cohabitation history and prior exposure to them. Such antipredator behavioral responses could have resulted in more favorable outcomes for an invading species during the invasive introductory process.     

Ecological effects of invasive alien species on native communities,with particular emphasis on the interactions between aphids and ladybirds

The ecological effects of introduced species on native organisms can sometimes, but not always be significant. The risks associated with invasive alien pests are difficult to quantify. This paper concentrates on the ecological effects of invasive insect predators that feed on pest insects, because the former may potentially affect the biological control of the latter. The literature indicates that invasive predatory insects generally are resistant to changes in environmental conditions, long-lived and voracious with a high reproductive rate, high dispersal ability, able to spread very rapidly across landscapes and exhibit phenotypic plasticity. Their colonization of patches of prey may induce native predators to leave, but the evidence that invaders negatively affect the abundance of the native species is scarce and not persuasive. Insect predators do not substantially affect the abundance of their prey, if the ratio of generation time of the predator to that of the prey is large (the generation time ratio hypothesis), therefore the effect of an invasion by long-lived alien predators on systems consisting of long-lived native predators and short-lived prey on the abundance of the prey is hard to detect.     

Trophic cascades in an invaded ecosystem: native keystone predators facilitate a dominant invader in an estuarine community

Determining how various factors contribute to the invasibility of systems is essential for both understanding community formation and informing management of natural areas. Research demonstrating that predators can provide biotic resistance to invasions by consuming invasive species has led to the presence of healthy predator populations being associated with reduced invasion potential of ecosystems. However, predators structure communities in many ways and their presence could also potentially facilitate invasions if they decrease populations of native species that compete with or consume an invader. We considered these two impacts of predators on invasion by analyzing the effects of two keystone predators (Pisaster spp. and Enhydra lutris nereis) on two foundation species (a native mussel Mytilus californianus and the invasive exotic bryozoan Watersipora subtorquata, a putative competitor for space with Mytilus californianus). Both native predators were found to facilitate the invasion of the exotic bryozoan, and the rate of invasion was highest when both predators were present. Facilitation of W. subtorquata occurred via indirect mechanisms that partly involved the removal of a competitor (mussels) via predation. These results illustrate that although predators can provide biotic resistance to invasion, healthy predator populations do not always confer this advantage and in fact may facilitate invasions. Therefore, implementation of management actions to enhance populations of top predators could also potentially increase the invasibility of some ecosystems.     

Comparison of susceptibility to a toxic alien toad (Bufo japonicus formosus) between predators in its native and invaded ranges

1. To manage biological invasions effectively, the impacts of alien species on the demography and traits of native species must be known, but determining those impacts can be challenging. We used a comparative approach to gain insight into the impacts that an alien toad (Bufo japonicus formosus) might have on native Japanese predatory amphibians. We compared the susceptibility of native predator species to alien toad toxins in the alien-invaded range and the susceptibility of closely related native predator species to the toxins in the alien toad's native range to investigate the impacts of an alien on a native species.
2. Bufo japonicus formosus is native to Honshu, but was recently introduced to Hokkaido and Sado. In laboratory experiments, we compared individual mortality of predators exposed to a toad hatchling between novel predators on the toad-invaded islands and ecologically similar congeneric or conspecific species on Honshu, where the toad is native. We also compared (1) the percentage of individuals that consumed a toad hatchling and (2) toxin resistance (i.e. survival and growth of individuals after toad consumption) between these two groups of predators, as mechanistic components behind the susceptibility of the predators to the toxic prey.
3. The mortality of Rana pirica from all populations after consumption of a toad hatchling was almost 100%, and that of Hynobius retardatus ranged from 14 to 90%, depending on the population. In contrast, the mortality of Rana ornativentris and Hynobius nigrescens was near 0% regardless of population. These differences between congeneric predators were mostly due to differences in their toxin resistance.
4. These results suggest that the alien toad is a potential threat to the novel amphibian predators on Hokkaido, although they also imply that the novel predators on Hokkaido have the potential to develop toxin resistance through adaptive evolution. However, this counteradaptation may have a higher chance of evolving in H. retardatus than in R. pirica because of differences in their genetic backgrounds.

     

Recognition and Response to Native and Novel Predators in the Largespring mosquitofish,Gambusia geiseri

The introduction of predator species into new habitats is an increasingly common consequence of human activities, and the persistence of native prey species depends upon their response to these novel predators. In this study, we examined whether the Largespring mosquitofish, Gambusia geiseri exhibited antipredator behavior and/or an elevation of circulating stress hormones (cortisol) to visual and chemical cues from a native predator, a novel predator, or a non‐predatory control fish. Prey showed the most pronounced antipredator response to the native predator treatment, by moving away from the stimulus, while the prey showed no significant changes in their vertical or horizontal position in response to the novel or non‐predator treatments. We also found no significant difference in water‐borne cortisol release rates following any of the treatments. Our results suggest the prey did not recognize and exhibit antipredator behavior to the novel predator, and we infer that this predator species could be detrimental if it expands into the range of this prey species. Further, our study demonstrates prey may not respond to an invasive predator that is phylogenetically, behaviorally, and morphologically dissimilar from the prey species' native predators.     

Hidden costs to an invasive intraguild predator from chemically defended native prey

Studies of invasive species tend to concentrate on the reasons for their success, rather than on the potential costs that they face. Invasive predators are potentially vulnerable to chemical defences of native prey that they have not previously encountered, and evolved to tolerate, within their natural range. While novel chemical defences have been cited as a factor facilitating invasion by exotic prey, the potential costs of chemical novelty to invasive predators have been ignored or remained hidden. We here consider one such example. Although a dominant intraguild predator in interactions with native ladybird beetles, the invasive ladybird Harmonia axyridis finds the defensive alkaloids of certain native ladybirds either distasteful or toxic, although laboratory and field evidence suggests it readily attacks these prey; these natives appear to possess novel alkaloids not previously encountered by H. axyridis . This effect has hitherto remained hidden, due to the methodological approaches used in earlier studies.     

Winning the biodiversity arms race among freshwater gastropods: competition and coexistence through shell variability and predator avoidance

Explanations for the coexistence of many closely related species in inland waters continue to be generated more than 50 years after Hutchinson’s question: why are there so many kinds of animals? This review focuses on the hypothesis that high species diversity of freshwater gastropods results, in part, from predators maintaining biodiversity across a range of deep- and shallow-water habitats. Invertebrate predators, such as aquatic insects, and leeches consume soft tissue of pulmonate snails by penetrating shells of various shapes and sizes. Crayfish and large prawns chip around the shell aperture to enter thick shells and crush small shells with their mandibles. Crabs use their strong chelae to crush thin and thick shells. Fishes with pharyngeal teeth are major shell-breaking predators that combine with other vertebrate predators such as turtles and wading birds to increase the diversity of gastropod communities by regulating the abundance of dominant species. Although the generalized diets of most freshwater predators preclude tight co-evolutionary patterns of responses, there are combinations of predators that modify gastropod behavior and shell morphology in aquatic assemblages of different ages and depths. This combination of invertebrate and vertebrate predatory impacts led to competitive advantages among individual gastropods with different adaptations: (1) less vulnerable shell morphologies and sizes; (2) predator-avoidance behaviors; or (3) rapid and widespread dispersal with variable life histories. Some individuals develop thicker and/or narrow-opening shells or shells with spines and ridges. Other thin-shelled species crawl out of the water or burrow to lower their risk to shell-breaking or shell-entering predators. Some alter their age at first reproduction and grow rapidly into a size refuge. Fluctuations in water levels and introductions of non-native species can change competitive dominance relationships among gastropods and result in major losses of native species. Many different gastropod predators control species that are human disease vectors. Most snails and their predators provide other ecosystem services such as nutrient cycling and transfer of energy to higher trophic levels. Their persistence and diversity of native species require adaptive management and coordinated study.     

A global review of the impacts of invasive cats on island endangered vertebrates

Cats are generalist predators that have been widely introduced to the world's ~179 000 islands. Once introduced to islands, cats prey on a variety of native species many of which lack evolved defenses against mammalian predators and can suffer severe population declines and even extinction. As islands house a disproportionate share of terrestrial biodiversity, the impacts of invasive cats on islands may have significant biodiversity impacts. Much of this threatened biodiversity can be protected by eradicating cats from islands. Information on the relative impacts of cats on different native species in different types of island ecosystems can increase the efficiency of this conservation tool. We reviewed feral cat impacts on native island vertebrates. Impacts of feral cats on vertebrates have been reported from at least 120 different islands on at least 175 vertebrates (25 reptiles, 123 birds, and 27 mammals), many of which are listed by the International Union for the Conservation of Nature. A meta‐analysis suggests that cat impacts were greatest on endemic species, particularly mammals and greater when non‐native prey species were also introduced. Feral cats on islands are responsible for at least 14% global bird, mammal, and reptile extinctions and are the principal threat to almost 8% of critically endangered birds, mammals, and reptiles.     

Spider community responses to grassland restoration: balancing trade‐offs between abundance and diversity

Spiders (Araneae) play key roles in ecosystems, not only as common and abundant generalist predators, but also as major contributors to biodiversity in many areas. In addition, due to their short generation times and high mobility, spiders respond rapidly to small changes in their environment, potentially making them useful indicators for restoration monitoring. However, few studies have focused on spider responses to grassland restoration in the United States. We compared degraded, native, and restored grassland sites to examine how spider communities and habitat respond to arid grassland restoration. We also examined how responses varied with the age of the restoration project. Spider communities in native sites differed from those in restored and degraded sites in several ways: native sites had fewer spiders and a different community composition than degraded and restored sites. However, native and restored sites had more species than degraded sites. Chronosequence data showed trends for lower abundance, higher species richness, and changing community composition as restoration projects mature. Several habitat variables were closely linked to variation in spider communities including cover of invasive annual grasses, litter, and biological soil crusts. Our data suggest that spider and vegetation responses to grassland restoration efforts can be successful in the long term—with resulting communities becoming more similar to native ones—and that spiders are useful indictors of grassland restoration. Our results also suggest that restoration may involve balancing trade‐offs between ecosystem services, with potential losses in predatory control offset by increases in biodiversity with restoration effort.     

Predator–prey naïveté, antipredator behavior,and the ecology of predator invasions

We present a framework for explaining variation in predator invasion success and predator impacts on native prey that integrates information about predator–prey naïveté, predator and prey behavioral responses to each other, consumptive and non‐consumptive effects of predators on prey, and interacting effects of multiple species interactions. We begin with the ‘naïve prey’ hypothesis that posits that naïve, native prey that lack evolutionary history with non‐native predators suffer heavy predation because they exhibit ineffective antipredator responses to novel predators. Not all naïve prey, however, show ineffective antipredator responses to novel predators. To explain variation in prey response to novel predators, we focus on the interaction between prey use of general versus specific cues and responses, and the functional similarity of non‐native and native predators. Effective antipredator responses reduce predation rates (reduce consumptive effects of predators, CEs), but often also carry costs that result in non‐consumptive effects (NCEs) of predators. We contrast expected CEs versus NCEs for non‐native versus native predators, and discuss how differences in the relative magnitudes of CEs and NCEs might influence invasion dynamics. Going beyond the effects of naïve prey, we discuss how the ‘naïve prey’, ‘enemy release’ and ‘evolution of increased competitive ability’ (EICA) hypotheses are inter‐related, and how the importance of all three might be mediated by prey and predator naïveté. These ideas hinge on the notion that non‐native predators enjoy a ‘novelty advantage’ associated with the naïveté of native prey and top predators. However, non‐native predators could instead suffer from a novelty disadvantage because they are also naïve to their new prey and potential predators. We hypothesize that patterns of community similarity and evolution might explain the variation in novelty advantage that can underlie variation in invasion outcomes. Finally, we discuss management implications of our framework, including suggestions for managing invasive predators, predator reintroductions and biological control.     

Keystone effects of an alien top-predator stem extinctions of native mammals

Alien predators can have catastrophic effects on ecosystems and are thought to be much more harmful to biodiversity than their native counterparts. However, trophic cascade theory and the mesopredator release hypothesis predict that the removal of top predators will result in the reorganization of trophic webs and loss of biodiversity. Using field data collected throughout arid Australia, we provide evidence that removal of an alien top-predator, the dingo, has cascading effects through lower trophic levels. Dingo removal was linked to increased activity of herbivores and an invasive mesopredator, the red fox (Vulpes vulpes), and to the loss of grass cover and native species of small mammals. Using species distribution data, we predict that reintroducing or maintaining dingo populations would produce a net benefit for the conservation of threatened native mammals across greater than 2.42 × 10⁶ km² of Australia. Our study provides evidence that an alien top predator can assume a keystone role and be beneficial for biodiversity conservation, and also that mammalian carnivores more generally can generate strong trophic cascades in terrestrial ecosystems.     

Rarity of a top predator triggers continent-wide collapse of mammal prey: dingoes and marsupials in Australia

Top predators in terrestrial ecosystems may limit populations of smaller predators that could otherwise become over abundant and cause declines and extinctions of some prey. It is therefore possible that top predators indirectly protect many species of prey from excessive predation. This effect has been demonstrated in some small-scale studies, but it is not known how general or important it is in maintaining prey biodiversity. During the last 150 years, Australia has suffered the world's highest rate of mammal decline and extinction, and most evidence points to introduced mid-sized predators (the red fox and the feral cat) as the cause. Here, we test the idea that the decline of Australia's largest native predator, the dingo, played a role in these extinctions. Dingoes were persecuted from the beginning of European settlement in Australia and have been eliminated or made rare over large parts of the continent. We show a strong positive relationship between the survival of marsupials and the geographical overlap with high-density dingo populations. Our results suggest that the rarity of dingoes was a critical factor which allowed smaller predators to overwhelm marsupial prey, triggering extinction over much of the continent. This is evidence of a crucial role of top predators in maintaining prey biodiversity at large scales in terrestrial ecosystems and suggests that many remaining Australian mammals would benefit from the positive management of dingoes.     

Native predator eats invasive toxic prey: evidence for increased incidence of consumption rather than aversion-learning

Contemporary adaptation of native prey species to invasive predators has been relatively well documented, but that of native predators to invasive prey has received less attention. Because the level of impact an invasive species will have on its predators versus its prey will determine changes in community trophic structure, it is important to understand how native predators respond to novel prey. Here we examine the response of native fence lizards to the invasion of red imported fire ants, a novel toxic prey. Examining invaded and uninvaded lizard populations, we tested whether or not aversion-learning occurs in juvenile fence lizards over successive feedings (within lifetime), how previous fire ant exposure may affect avoidance behavior (over generations), and whether population differences are consistent when prey choice exists. We also examine rates of phenotypic divergence in traits associated with the native species as both predator and prey. Aversion-learning did not occur in either population. Instead, the incidence of fire ant consumption increased over both successive feedings and generations. Lizards from the fire ant invaded population had a higher propensity to eat fire ants than fire ant-naïve lizards, even when given a choice between prey items. We found greater phenotypic divergence in traits associated with the native species as predator on, versus as prey to, fire ants. Although the strategy of eating these novel toxic prey can impose survival costs in the short term, over the longer-term, eating fire ants may cost little or even benefit survivors.