The diet of feral cats on islands: a review and a call for more studies

Cats are among the most successful and damaging invaders on islands and a significant driver of extinction and endangerment. Better understanding of their ecology can improve effective management actions such as eradication. We reviewed 72 studies of insular feral cat diet from 40 islands worldwide. Cats fed on a wide range of species from large birds and medium sized mammals to small insects with at least 248 species consumed (27 mammals, 113 birds, 34 reptiles, 3 amphibians, 2 fish and 69 invertebrates). Three mammals, 29 birds and 3 reptiles recorded in the diet of cats are listed as threatened by the IUCN. However, a few species of introduced mammals were the most frequent prey, and on almost all islands mammals and birds contributed most of the daily food intake. Latitude was positively correlated with the predation of rabbits and negatively with the predation of reptiles and invertebrates. Distance from landmass was positively correlated with predation on birds and negatively correlated with the predation of reptiles. The broad range of taxa consumed by feral cats on islands suggests that they have the potential to impact almost any native species, even the smallest ones under several grams, that lack behavioral, morphological or life history adaptations to mammalian predators. Insular feral cat??s reliance on introduced mammals, which evolved with cat predation, suggests that on many islands, populations of native species have already been reduced.     

Direct evidence implicates feral cat predation as the primary cause of failure of a mammal reintroduction programme

As evidence mounts that the feral Cat (Felis catus) is a significant threat to endemic Australian biodiversity and impedes reintroduction attempts, uncertainty remains about the impact a residual population of cats following control will have on a mammal reintroduction programme. Also, behavioural interactions between cats and their prey continue to be an area of interest. Within the framework of an ecosystem restoration project, we tested the hypotheses that successful reintroductions of some medium‐sized mammals are possible in locations where feral cats are controlled (but not eradicated) in the absence of European Red Fox (Vulpes vulpes), and that hare‐wallabies that dispersed from their release area are more vulnerable to cat predation compared with those that remain at the release site. We used radiotelemetry to monitor the survivorship and dispersal of 16 Rufous Hare‐wallabies (Lagorchestes hirsutus spp.) and 18 Banded Hare‐wallabies (Lagostrophus fasciatus fasciatus) reintroduced to four sites within Shark Bay, Western Australia. Nearly all foxes were removed and feral cats were subject to ongoing control that kept their indices low relative to prerelease levels. All monitored hare‐wallabies were killed by cats within eight and 10 months following release. Significant predation by feral cats was not immediate: most kills occurred in clusters, with periods of several months where no mortalities occurred. Once a hare‐wallaby was killed, however, predation continued until each population was eliminated. Animals remaining near their release site survived longer than those that dispersed. The aetiology of predation events observed offers new insights into patterns of feral cat behaviour and mammal releases. We propose a hypothesis that these intense per capita predation events may reflect a targeted hunting behaviour in individual feral cats. Even where feral cats are controlled, the outcome from consistent predation events will result in reintroduction failures. Managers considering the reintroduction of medium‐sized mammals in the presence of feral cats should, irrespective of concurrent cat control, consider the low probability of success. We advocate alternative approaches to cat‐baiting alone for the recovery of cat‐vulnerable mammals such as hare‐wallabies.     

Habitat and introduced predators influence the occupancy of small threatened macropods in subtropical Australia

Australia has had the highest rate of mammal extinctions in the past two centuries when compared to other continents. Frequently cited threats include habitat loss and fragmentation, changed fire regimes and the impact of introduced predators, namely the red fox (Vulpes vulpes) and the feral cat (Felis catus). Recent studies suggest that Australia's top predator, the dingo (Canis dingo), may have a suppressive effect on fox populations but not on cat populations. The landscape of fear hypothesis proposes that habitat used by prey species comprises high to low risk patches for foraging as determined by the presence and ubiquity of predators within the ecosystem. This results in a landscape of risky versus safe areas for prey species. We investigated the influence of habitat and its interaction with predatory mammals on the occupancy of medium‐sized mammals with a focus on threatened macropodid marsupials (the long‐nosed potoroo [Potorous tridactylous] and red‐legged pademelon [Thylogale stigmatica]). We assumed that differential use of habitats would reflect trade‐offs between food and safety. We predicted that medium‐sized mammals would prefer habitats for foraging that reduce the risk of predation but that predators would have a positive relationship with medium‐sized mammals. We variously used data from 298 camera trap sites across nine conservation reserves in subtropical Australia. Both dingoes and feral cats were broadly distributed, whilst the red fox was rare. Long‐nosed potoroos had a strong positive association with dense ground cover, consistent with using habitat complexity to escape predation. Red‐legged pademelons showed a preference for open ground cover, consistent with a reliance on rapid bounding to escape predation. Dingoes preferred areas of open ground cover whereas feral cats showed no specific habitat preference. Dingoes were positively associated with long‐nosed potoroos whilst feral cats were positively associated with red‐legged pademelons. Our study highlights the importance of habitat structure to these threatened mammals and also the need for more detailed study of their interactions with their predators.     

Introduced cats Felis catus eating a continental fauna: inventory and traits of Australian mammal species killed

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Predation of insects by feral cats (<Emphasis Type="Italic">Felis silvestris catus</Emphasis> L., 1758) on an oceanic island (La Palma,Canary Island)

Predation of insects by feral cats (Felis silvestris catus) on a heterogeneous oceanic island (La Palma, Canary Islands) was studied. A total of 127 invertebrates were identified in the analysis of 500 scats (100 from each habitat of the Island). Invertebrates appear in 18.00% of the scats, representing an insignificant percentage of the total consumed biomass by feral cats on La Palma Island (0.05%). Insects were the most common invertebrate prey both in percentage of occurrence (90.6%) and invertebrate biomass (93.53%), with a total of 115 prey items. Orthoptera, Lepidoptera and Coleoptera were the main prey groups. Among the five main habitats present in La Palma Island, the temperate forest shows the lowest consumption of invertebrates, although insect consumption did not show statistical differences. However Orthoptera and Lepidoptera were more frequently preyed on in the pine forest and in the xerophytic shrub, respectively. Moreover, applying the Simplified Morisita index, a different insect composition of the diet was observed among habitats. Although, none of insects predated by feral cats are threatened, the identification of invertebrate component of the feral cats’ diet is an important tool for the correct understanding of predation significance and to prevent damage to endangered insect species.     

Trophic habits of feral cats in the high mountain shrublands of the Macaronesian islands (NW Africa,Atlantic Ocean)

Feral cats Felis catus Linnaeus, 1758 have contributed to the extinction of numerous native species on islands, which are clearly sources of global biodiversity. We studied the diet of this introduced predator in the Madeira and Cape Verde archipelagos, which harbour important colonies of endangered seabirds in the high mountain habitats, and compared the results with those obtained in the same habitat in the Canary Islands, Macaronesian archipelago. On Madeira, 461 prey were identified from 143 scat groups. Mammals, overall mice, constituted the basic diet appearing in 95% of cat scats. On Fogo (Cape Verde), 657 prey items were obtained from 145 scats, and mammals were also the most important prey, reaching a frequency of occurrence of 88%. Although introduced mammals were the main prey category on all Macaronesian islands, we observed variation in feral cat diet among these islands. Birds were more frequently consumed on Madeira, lizards on Tenerife (Canaries) and invertebrates on Fogo. No specific differences were observed in relation to La Palma. We suggest that the diet composition on these islands varies according to the respective availability of the different prey types.     

Seasonal and sex-specific prey composition of black-footed catsFelis nigripes

During a 6-year field study on the game farm ‘Benfontein’ in the central Republic of South Africa 1725 prey items were observed consumed by 17 free-ranging habituated black-footed catsFelis nigripes Burchell, 1824. Average prey size was 24.1 g. Eight males fed on significantly larger prey (27.9 g) than 9 females (20.8 g). Fifty-four prey species were classified by their average mass into 8 different size classes, 3 for mammals, 3 for birds, 1 for amphibians/reptiles, and 1 for invertebrates. Small mammals (5–40 g) constituted the most important prey class (39%) of total prey biomass followed by larger mammals (>100 g; 17%) and small birds (<40 g; 16%). Mammals and birds pooled comprised 72% and 26% of total prey biomass, respectively, whereas invertebrates and amphibians/reptiles combined constituted just 2% of total prey mass consumed. Three seasons of 4-months duration were recognized. Heterotherm prey items were unavailable during winter, when larger birds and mammals (> 100 g) were mainly consumed. Small rodents like the large-eared mouseMalacothrix typica, captured 595 times by both sexes, were particularly important during the reproductive season for females with kittens. Male black-footed cats showed less variation between prey size classes consumed among climatic seasons. This sex-specific difference in prey size consumption may help to reduce intra-specific competition.     

Diet of the feral cat,Felis catus,in central Australian grassland habitats: do cat attributes influence what they eat?

The house cat Felis catus was introduced to Australia as a pet and means of rodent control over 200 years ago, but now has established feral populations and has become a serious threat to native wildlife. Using stomach content analysis of 73 feral cats from semi-arid grassland habitats in Queensland, Australia, we aimed to identify dominant prey groups in the cats' diet and to explore associations between the diversity of prey eaten and attributes of the cats including body size, condition, sex, age and coat colour. We also sought to determine any relationships between cat size and the size of the dominant prey in the diet, the long-haired rat Rattus villosissimus. Mammals and reptiles were the dominant prey, with R. villosissimus occurring in 60 % of samples and comprising more than half of all prey by volume. Birds and terrestrial invertebrates were the next most important contributors to the diet, but fish, frogs and freshwater crustaceans also were surprisingly well represented. The dietary diversity of cats was largely unrelated to any of the cat attributes that we measured, although a positive relationship emerged between cat head width and the range of prey types eaten. Our study was conducted during a population irruption of R. villosissimus and confirms that cats are able to exploit an abundant focal prey resource when the opportunity occurs. Further research now is needed to explore associations between diet and cat attributes during periods when rats are scarce.     

Hunters and non-hunters: skewed predation rate by domestic cats in a rural village

Domestic cats Felis catus, as companion animals provided with supplemental food, are not limited by the availability of wild prey and locally occur at extraordinary high densities. There is growing concern about the potential impact of large cat numbers on native prey populations. In the present study, we quantified the minimum number of animals killed in a rural village in Switzerland by asking owners (1) to estimate the predation rate in advance and (2) to record prey animals returned home by their pets. The frequency distribution of the numbers of prey items was markedly skewed: 16% of the cats accounted for 75% of prey, irrespective of sex, age or breed. A large fraction of owners considerably overestimated their cat’s predation, indicating that surveying predation rates by means of a questionnaire alone is not sufficient. The observed average rate of predation within 48 days in spring was 2.29 prey items/cat/month (N = 32 cats); major prey types were rodents (76.1%) and birds (11.1%). The absolute number of prey items taken per area is striking and indicates that cat predation represents an important factor in ecosystems. Its role may be momentous in intensively fragmented urban habitats, where cat densities are especially high. We thus highlight the need to identify the factors determining predation rates of individual cats. Further extended studies, especially in urbanised areas, are needed to quantify the actual impact of cat predation upon the population dynamics of their prey.

     

The dynamics of predation on Gammarus spp. (Crustacea: Amphipoda)

Gammarus spp. (Crustacea: Amphipoda) are widespread throughout a diverse range of marine, freshwater and estuarine/brackish habitats, often dominating benthic macroinvertebrate communities in terms of both numbers and/or biomass. Gammarus spp. are the dominant macroinvertebrate prey items of many fish, whether as a seasonal food source or a year-round staple. Selective predation by fish on Gammarus spp. is often linked to parasitism and the body size of the prey. Gammarus spp. populations are under increasing threat from both pollution and replacement/displacement by introduced species. Loss of populations and species invasions/replacements could have significant impacts on native predator species if the predator(s) cannot successfully adapt their feeding patterns to cope with non-indigenous Gammarus prey species. Despite this, many fish predation studies do not identify Gammarus prey to species level. This lack of precision could be important, as Gammarus spp. exhibit wide variations in physiochemical tolerances, habitat requirements, abilities to invade and susceptibility to replacement. Although rarely acknowledged, the impacts of nonpiscean predators (particularly macroinvertebrates) on Gammarus prey species may frequently be stronger than those exerted by fish. A major aim of this review is to ascertain the current importance of Gammarus as a prey species, such that the implications of changes in Gammarus spp. populations can be more accurately assessed by interested groups such as ecologists and fisheries managers. We also review the dynamics of Gammarus spp. as prey to a diverse array of mammals, birds, amphibians, insects, flatworms, other crustaceans such as crabs and crayfish and, perhaps most importantly, other Gammarus spp.     

Trophic status and feeding habits of the European Polecat Mustela putorius L. 1758

The trophic status of the European Polecat Mustela putorius was studied through a review of the diets of 18 different populations in Europe. Rodents represented the principal prey in 10 sites (55.5%) and were a prey of secondary importance in seven sites (38.8%). Anurans prevailed in three sites (17%) but constituted the second food category in eight others (44%). The ascendant hierarchical classification showed a relatively unvarying diet in Europe. The frequency of birds and invertebrates were important in the diversification of the diet, indicated by the value of the food niche index, whereas rabbits were more significant in southern regions. The constant incidence of predation on woodland rodents and amphibians gave a particular status to the Polecat and showed it to be a generalist feeder well adapted to the mid-European region.     

Abundance and detection of feral cats decreases after severe fire on Kangaroo Island,Australia

Predation by feral cats (Felis catus) has caused the extinction of many native species in Australia and globally. There is growing evidence that the impacts of feral cats can be amplified in post-fire environments, as cats are drawn to hunt in or around recently burnt areas and are also more effective hunters in open habitats. In 2018–2019, we established arrays of camera traps to estimate the abundance of feral cats on Kangaroo Island, South Australia. Much of the island (including five of our seven survey sites) was subsequently burnt in a severe wildfire (December 2019–February 2020). We re-sampled the sites 3–8 months post-fire (seven sites) and 11–12 months post-fire (three sites). At two unburnt sites sampled post-fire, it was possible to produce density estimates of cats using a spatially explicit capture–recapture approach. Where estimating density was not possible (due to low detections or individual cats not being distinguishable), the number of individuals and percentage of trap nights with detections was compared between the sampling periods. Some low-level cat control occurred within 2 km of three of the seven arrays (all within the burn scar) within 3 months of the fire. Across the five burnt sites, there was a decline in cat detections post-fire (including those without post-fire cat control). At 3–8 months post-fire, there was, on average, a 57% reduction in the number of individual cats, and a 65% reduction in the number of nights with cat detections, relative to pre-fire levels. Although cat detections declined following the fire, reduced population sizes of prey species and reduced cover as a result of the fire might still mean that cat predation is a threat to some surviving prey species. Management that reduces feral cat predation pressure on wildlife following wildfire should enhance the likelihood of post-fire wildlife persistence and recovery.     

Comparative analysis of the diet of feral and house cats and wildcat in Europe

Differences in availability of food resources are often manifested in the differentiation of feeding habits of closely related mammal species. Therefore, we assumed that the diet composition and trophic niche of house (i.e., highly dependent on human households), feral (i.e. independent on human households) domestic cats (Felis silvestris catus) and wildcats (Felis s. silvestris) differs. Based on the literature data from Europe (53 study sites), we compared the diet of these three felids analyzed with use of indirect methods (stomach and scat analyses). In the case of the house cat, we additionally compared consumption data obtained directly from prey brought home. Data were expressed as the relative frequency of occurrence to compare dietary patterns. The main prey of the three cat types were small mammals in different ratios. According to the stomach and scat samples, the diet composition of the cat types showed differences in the consumption of rodents, insectivores, wild ungulates, and household food, supporting the “dietary differences originate from varying resources” hypothesis. More opportunistic house cats had a broader trophic niche than feeding specialist wildcats, while feral cats had an intermediate position. The trophic niche breadth of all three cat types increased along a latitudinal gradient from northern to southern areas of Europe. The predation of the house cat which was examined from prey brought home differed from the data obtained by indirect diet analysis; however, it yielded similar results to the diet of the feral cat and the wildcat. Due to their high numbers and similarity of its diet to the wildcat, house cats are a threat to wild animals; therefore, their predation pressure needs to be further investigated.     

Feeding habits of Geoffroy's cat (Leopardus geoffroyi) in southern Brazil

The present study was carried out from 2001 to 2005, with the objective of analyzing the diet of Leopardus geoffroyi in extreme southern Brazil, through analyses of stomach contents and scats. The importance of each prey type found in the scats (n=66) was determined from the frequency of occurrence and percentage of occurrence. In the stomachs (n=9), the importance of each prey was determined using Pianka's index of relative importance (IRI). The food spectrum of L. geoffroyi included mammals, birds, serpents, amphibians and insects. Mammals were the most abundant items, present in 95.5% of the scats and representing 74.4% of the total prey, reaching an IRI of 13296.3 in the stomachs. The principal prey types found in the stomachs and scats were small rodents (Cavia spp., Oligoryzomys spp., Necromys spp. and Holochilus brasiliensis) and large-sized rodents (Myocastor coypus), of terrestrial, arboreal or semi-aquatic habit. Probably, the high percentage of these prey items in the diet is related to their availability and abundance. The index of niche breadth was low in both the stomach and scat analyses (respectively, B_sta=0.18 and 0.17), demonstrating the high degree of specialization of these cats.     

Predicting bait uptake by feral cats,Felis catus,in semi‐arid environments

Feral cat control using aerial broadcasting of toxic baits continues to be used in the rangelands of Western Australia. The effectiveness of these operations has sometimes been compromised by different environmental factors that affect prey and cat numbers. This study demonstrates that the ratio of cats to their preferred prey (small mammals) can be used to predict the most effective time to bait. The regular baiting of three conservation sites offered an opportunity to study the relationship between feral cat abundance, the abundance of their prey and ingestion of toxic baits. Peron Peninsula on the mid‐west coast, Lorna Glen station in the northern Goldfields and the central Gibson Desert of Western Australia are sites where cat control using toxic baits has been routinely applied over the last 15 years. We postulated that bait ingestion by cats was linked to the availability of live prey. Small mammal abundance (capture rates in pit‐fall traps) and relative cat abundance (based on daily track counts) were assessed at these sites and the data used to produce a predator‐prey ratio index (PPRI). We used generalised linear mixed models to test the effect of prey abundance, prebaiting cat abundance and PPRI on baiting efficacy (BE). The best model for predicting efficacy of baiting contained only PPRI. This simple model was able to predict baiting success over the entire range of outcomes, from highly successful ( >75% cat reduction) to unsuccessful (0% cat reduction). The ability to predict feral cat BE in advance of planned toxic baiting operations will provide a valuable tool for wildlife managers involved in cat control.     

Diets of sympatric native and introduced carnivores in the Barrington Tops,eastern Australia

Invasive predators have severe impacts on global biodiversity, and their effects in Australia have been more extreme than on any other continent. The spotted‐tailed quoll (Dasyurus maculatus), an endangered marsupial carnivore, coexists with three eutherian carnivores, the red fox (Vulpes vulpes), feral cat (Felis catus) and wild dog (Canis lupus ssp.) with which it did not coevolve. No previous study has investigated dietary overlap between quolls and the suite of three eutherian carnivores. By analysing scats, we aimed to quantify dietary overlap within this carnivore assemblage in eastern Australia, and to detect any differences that may facilitate coexistence. We also sought evidence of intraguild predation. Dietary overlap between predators was extensive, with the greatest similarity occurring between foxes and cats. However, some differences were apparent. For example, cats mainly consumed smaller prey, and wild dogs larger prey. Quolls showed greater dietary overlap with foxes and cats than with dogs. Intraguild predation was evident, with fox remains occurring in 3% of wild dog scats. Our results suggest wild dogs competitively dominate invasive foxes, which in turn are likely to compete with the endangered quoll.     

A triple threat: high population density,high foraging intensity and flexible habitat preferences explain high impact of feral cats on prey

Alien mammalian carnivores have contributed disproportionately to global loss of biodiversity. In Australia, predation by the feral cat and red fox is one of the most significant causes of the decline of native vertebrates. To discover why cats have greater impacts on prey than native predators, we compared the ecology of the feral cat to a marsupial counterpart, the spotted-tailed quoll. Individual prey are 20–200 times more likely to encounter feral cats, because of the combined effects of cats'' higher population densities, greater intensity of home-range use and broader habitat preferences. These characteristics also mean that the costs to the prey of adopting anti-predator behaviours against feral cats are likely to be much higher than adopting such behaviours in response to spotted-tailed quolls, due to the reliability and ubiquity of feral cat cues. These results help explain the devastating impacts of cats on wildlife in Australia and other parts of the world.     

Detecting frogs as prey in the diets of introduced mammals: a comparison between morphological and DNA‐based diet analyses

Amphibians are currently the most threatened group of vertebrates worldwide, and introduced fauna play a major role in their decline. The control of introduced predators to protect endangered species is often based on predation rates derived from diet studies of predators, but prey detection probabilities using different techniques are variable. We measured the detectability of frogs as prey, using morphological and DNA‐based diet analyses, in the stomachs and faeces of four mammal species that have been introduced to many areas of the world. Frogs (Litoria raniformis) were fed to rats (Rattus norvegicus and R. rattus), mice (Mus musculus) and hedgehogs (Erinaceus europaeus). DNA‐based analysis outperformed morphological analysis, increasing the prey detection rate from 2% to 70% in stomachs and from 0% to 53% in faeces. In most cases, utilizing either stomachs or faeces did not affect the success of prey DNA detection; however, using faeces extended the detectability half‐life from 7 to 21 h. This study is the first to measure prey DNA detection periods in mammalian stomachs, and the first to compare prey DNA detection periods in the stomachs and faeces of vertebrates. The results indicate that DNA‐based diet analysis provides a more reliable approach for detecting amphibians as prey and has the potential to be used to estimate the rate of predation by introduced mammals on endangered amphibians.     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.