PDF The Impact of Free-Ranging Domestic Cats on Wildlife of the United States

ARTICLE Received 6 Sep 2012 | Accepted 12 Dec 2012 | Published 29 Jan 2013 | Updated 12 Dec 2013 DOI: 10.1038/ncomms2380 The impact of free-ranging domestic cats on wildlife of the United States Scott R. Loss1, Tom Will2 & Peter P. Marra1 Anthropogenic threats, such as collisions with man-made structures, vehicles, poisoning and predation by domestic pets, combine to kill billions of wildlife annually. Free-ranging domestic cats have been introduced globally and have contributed to multiple wildlife extinctions on islands. The magnitude of mortality they cause in mainland areas remains speculative, with large-scale estimates based on non-systematic analyses and little consideration of scientiﬁc data. Here we conduct a systematic review and quantitatively estimate mortality caused by cats in the United States. We estimate that free-ranging domestic cats kill 1.3–4.0 billion birds and 6.3–22.3 billion mammals annually. Un-owned cats, as opposed to owned pets, cause the majority of this mortality. Our ﬁndings suggest that free-ranging cats cause substantially greater wildlife mortality than previously thought and are likely the single greatest source of anthropogenic mortality for US birds and mammals. Scientiﬁcally sound conservation and policy intervention is needed to reduce this impact. 1 Migratory Bird Center, Smithsonian Conservation Biology Institute, National Zoological Park, P.O. Box 37012 MRC 5503, Washington, District of Columbia 20013, USA. 2 U.S. Fish and Wildlife Service, Division of Migratory Birds, Midwest Regional Ofﬁce, 3815 American Boulevard East, Bloomington, Minnesota 20013, USA. Correspondence and requests for materials should be addressed to S.R.L. (email: [email protected]). NATURE COMMUNICATIONS | 4:1396 | DOI: 10.1038/ncomms2380 | www.nature.com/naturecommunications 1 & 2013 Macmillan Publishers Limited. All rights reserved. ARTICLE NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2380 D omestic cats (Felis catus) are predators that humans have owned and un-owned cats—and running 10,000 calculation introduced globally1,2 and that have been listed among iterations. We augmented US predation data by incorporating the 100 worst non-native invasive species in the world3. predation rate estimates from other temperate regions Free-ranging cats on islands have caused or contributed to 33 (Supplementary Table S1). For birds, we generated three US (14%) of the modern bird, mammal and reptile extinctions mortality estimates based on predation data from studies in: recorded by the International Union for Conservation of Nature (1) the United States, (2) the United States and Europe and (IUCN) Red List4. Mounting evidence from three continents (3) the United States, Europe, and other temperate regions (pri- indicates that cats can also locally reduce mainland bird and marily Australia and New Zealand). Owing to a lack of US studies mammal populations5–7 and cause a substantial proportion of un-owned cat predation on mammals, we estimated mammal of total wildlife mortality8–10. Despite these harmful effects, mortality using data groupings 2 and 3. We based all other policies for management of free-ranging cat populations and probability distributions on US studies (distribution details in regulation of pet ownership behaviours are dictated by animal Table 1; data in Supplementary Table S2). welfare issues rather than ecological impacts11. Projects to The three estimates of bird mortality varied moderately, with a manage free-ranging cats, such as Trap-Neuter-Return (TNR) 19% difference among median estimates (Table 2). We focus colonies, are potentially harmful to wildlife populations, but interpretation on the estimate generated using US and European are implemented across the United States without widespread predation data because it is the lowest value. Furthermore, this public knowledge, consideration of scientiﬁc evidence or the estimate is more likely to be representative of the US than the environmental review processes typically required for actions estimate based on incorporation of data from Australia and New with harmful environmental consequences11,12. Zealand, where the wildlife fauna and climate are less similar A major reason for the current non-scientiﬁc approach to to the United States. We estimate that cats in the contiguous management of free-ranging cats is that total mortality from cat United States annually kill between 1.3 and 4.0 billion birds predation is often argued to be negligible compared with other (median ¼ 2.4 billion) (Fig. 1a), with B69% of this mortality anthropogenic threats, such as collisions with man-made caused by un-owned cats. The predation estimate for un-owned structures and habitat destruction. However, assessing the cats was higher primarily due to predation rates by this group conservation importance of a mortality source requires identiﬁca- averaging three times greater than rates for owned cats. tion of which species are being killed (for example, native versus non-native invasive species and rare versus common species) in The magnitude of mammal mortality caused by cat predation. addition to estimation of total numbers of fatalities. Estimates of Our estimate of mammal mortality was robust to the choice of annual US bird mortality from predation by all cats, including predation data as evidenced by a 1.6% difference between the two both owned and un-owned cats, are in the hundreds of median estimates (Table 2). We focus interpretation on the lower millions13,14 (we deﬁne un-owned cats to include farm/barn estimate, which was based on United States and European pre- cats, strays that are fed by humans but not granted access to dation data and US values of other parameters. We estimate habitations, cats in subsidized colonies and cats that are annual mammal mortality in the contiguous United States at completely feral). This magnitude would place cats among the between 6.3 and 22.3 billion (median ¼ 12.3 billion) (Fig. 1b) with top sources of anthropogenic bird mortality; however, window 89% of this mortality caused by un-owned cats. The estimate that and building collisions have been suggested to cause even greater incorporated European data (but not data from Australia and mortality15–17. Existing estimates of mortality from cat predation New Zealand) may be slightly lower because wildlife across much are speculative and not based on scientiﬁc data13–16 or, at best, of Europe were historically exposed to predation by a similarly- are based on extrapolation of results from a single study18. In sized wild cat (Felis sylvestris) and, therefore, may be less naive to addition, no large-scale mortality estimates exist for mammals, predation by domestic cats. However, it is unlikely that European which form a substantial component of cat diets. wildlife have fully adapted to the unusually high densities of We conducted a data-driven systematic review of studies that domestic cats in much of this continent9. estimate predation rates of owned and un-owned cats, and estimated the magnitude of bird and mammal mortality caused by all cats across the contiguous United States (all states excluding Factors explaining estimate uncertainty. For both birds and Alaska and Hawaii). We estimate that free-ranging domestic cats mammals, sensitivity analyses indicated that un-owned cat kill 1.3–4.0 billion birds and 6.3–22.3 billion mammals annually, parameters explained the greatest variation in total mortality and that un-owned cats cause the majority of this mortality. This estimates (Fig. 2). Un-owned cat population size explained the magnitude of mortality is far greater than previous estimates of greatest variation in mortality estimates (42% for birds and 51% cat predation on wildlife and may exceed all other sources of for mammals), and the un-owned cat predation rate explained the anthropogenic mortality of US birds and mammals. second greatest variation (24% for birds and 40% for mammals). The only other parameters that explained 45% of variation in mortality estimates were the owned cat predation rate on birds Results (16%) and the correction factor for imperfect detection of owned The magnitude of bird mortality caused by cat predation. After cats’ prey items (8%). excluding studies that did not meet a priori inclusion criteria designed to increase the accuracy of our analysis, we developed Discussion probability distributions of predation rates on birds and mam- Our estimate of bird mortality far exceeds any previously mals. We combined predation rate distributions with literature- estimated US ﬁgure for cats13,14,16, as well as estimates for any derived probability distributions for US cat population sizes, and other direct source of anthropogenic mortality, including we also accounted for the proportion of owned cats allowed collisions with windows, buildings, communication towers, outdoors, the proportion of owned and un-owned cats that hunt, vehicles and pesticide poisoning13,15–21. Systematic reviews like and imperfect detection of owned cats’ prey items. ours, which includes protocol formulation, a data search strategy, We generated an estimated range of bird and mammal mor- data inclusion criteria, data extraction and formal quantitative tality caused by cat predation by incorporating the above analyses22, are scarce for other anthropogenic mortality sources.21 distributions—including separate predation rate distributions for Increased rigour of mortality estimates should be a high priority 2 NATURE COMMUNICATIONS | 4:1396 | DOI: 10.1038/ncomms2380 | www.nature.com/naturecommunications & 2013 Macmillan Publishers Limited. All rights reserved. NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2380 ARTICLE Table 1 | Probability distributions used for parameters in cat predation model. Parameter Number of studies used* Distribution type Distribution parameters Owned cats Number of owned cats in contiguous United States 2 Normal Mean ¼ 84 M, s.d. ¼ 2.5 M Proportion of owned cats with outdoor access 8 Uniform Min ¼ 0.4, max ¼ 0.7 Proportion of outdoor owned cats that hunt 3 Uniform Min ¼ 0.5, max ¼ 0.8 Correction for owned cats not returning prey 3 Uniform Min ¼ 1.2, max ¼ 3.3 BIRD return rate per cat per year US studies 4 Uniform Min ¼ 1.0, max ¼ 34.1 US and Europe studies 11 Uniform Min ¼ 4.2, max ¼ 18.3 All temperate studies 17 Uniform Min ¼ 3.4, max ¼ 13.2 MAMMAL return rate per cat per year US studies 1 NAw NA US and Europe studies 7 Uniform Min ¼ 11.1, max ¼ 29.5 All temperate studies 13 Uniform Min ¼ 8.7, max ¼ 21.8 REPTILE return rate per cat per year US studies 0 NA NA US and Europe studies 1 NA NA All temperate studies 8 Uniform Min ¼ 0.4, max ¼ 2.21 AMPHIBIAN return rate per cat per year US studies 0 NA NA US and Europe studies 1 NA NA All temperate studies 5 Uniform Min ¼ 0.05, max ¼ 0.5 Un-owned cats Number of un-owned cats in contiguous United States 5 Uniform Min ¼ 30 M, max ¼ 80 M Proportion of un-owned cats that hunt 2 Uniform Min ¼ 0.8, max ¼ 1.0 BIRD predation rate per cat per year US studies 8 Uniform Min ¼ 24.4, max ¼ 51.4 US and Europe studies 11 Uniform Min ¼ 23.2, max ¼ 46.2 All temperate studies 19 Uniform Min ¼ 30.0, max ¼ 47.6 MAMMAL predation rate per cat per year US studies 6 Uniform Min ¼ 162.3, max ¼ 354.9 US and Europe studies 7 Uniform Min ¼ 139.4, max ¼ 328.6 All temperate studies 13 Uniform Min ¼ 177.3, max ¼ 299.5 REPTILE predation rate per cat per year US studies 1 NA NA US and Europe studies 2 NA NA All temperate studies 10 Uniform Min ¼ 4.2, max ¼ 12.4 AMPHIBIAN predation rate per cat per year US studies 0 NA NA US and Europe studies 0 NA NA All temperate studies 3 Uniform Min ¼ 1.9, max ¼ 4.7 *Number of studies found that include an estimate of the model parameter. wNo calculation was conducted for this data grouping because of limited data. Table 2 | Median estimates of annual wildlife mortality caused by cat predation in the contiguous United States. Predation data used Mortality estimate (millions) Owned cats Un-owned cats Total Bird US 1,053 (104–3,039)* 1,792 (861–3,276) 2,967 (1,358–5,324) US and Europe 684 (221–1,682)w 1,652 (803–2,955) 2,407 (1,306–3,992) All temperate 508 (172–1,226) 1,876 (983–3,124) 2,437 (1,378–3,814) Mammal US and Europe 1,249 (512–2,862) 10,903 (4,991–20,874) 12,269 (6,259–22,257) All temperate 958 (397–2,117) 11,426 (5,874–19,451) 12,473 (6,874–20,421) Both Taxa 1,933 (733–4,544) 12,555 (5,794–23,829) 14,676 (7,565–26,249) *Values in parentheses indicate central 95% of estimates. wBold face indicates estimates from which inference is drawn in the text. NATURE COMMUNICATIONS | 4:1396 | DOI: 10.1038/ncomms2380 | www.nature.com/naturecommunications 3 & 2013 Macmillan Publishers Limited. All rights reserved. ARTICLE NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2380 and will allow increased comparability of mortality sources23. 228 to 871 million reptiles (median ¼ 478 million) and between Nonetheless, no estimates of any other anthropogenic mortality 86 and 320 million amphibians (median ¼ 173 million) could be source approach the value we calculated for cat predation, and killed by cats in the contiguous United States each year. Reptile our estimate is the ﬁrst for cats to be based on rigorous data- and amphibian populations, and, therefore, cat predation rates, driven methods. Notably, we excluded high local predation rates may differ between the regions where we gathered predation data and used assumptions that led to minimum predation rate for these taxa and the United States. Furthermore, reptiles and estimates for un-owned cats; therefore, actual numbers of birds amphibians are unavailable as prey during winter across much of killed may be even greater than our estimates. the United States. Additional research is needed to clarify impacts Free-roaming cats in the United States may also have a of cats on US herpetofauna, especially given numerous anthro- substantial impact on reptiles and amphibians. However, US pogenic stressors that threaten their populations (for example, studies of cat predation on these taxa are scarce. To generate a climate change, habitat loss and infectious diseases) and ﬁrst approximation of US predation rates on reptiles and documented extinctions of reptiles and amphibians due to cat amphibians, we used the same model of cat predation along predation in other regions4,24. with estimates of cat predation rates on these taxa from studies in The exceptionally high estimate of mammal mortality from cat Europe, Australia and New Zealand. We estimate that between predation is supported by individual US studies that illustrate high annual predation rates by individual un-owned cats in excess of 200 mammals per year6,25–28 and the consistent ﬁnding that a cats preferentially depredate mammals over other taxa Median=2.4 billion (Supplementary Table S1). Even with a lower yearly predation 600 rate of 100 mammals per cat, annual mortality would range from Estimate frequency 500 3–8 billion mammals just for un-owned cats, based on a 400 population estimate of between 30 and 80 million un-owned 300 cats. This estimated level of mortality could exceed any other 200 direct source of anthropogenic mortality for small mammals; 100 however, we are unaware of studies that have systematically 0 quantiﬁed direct anthropogenic mortality of small terrestrial 0 1,000 2,000 3,000 4,000 5,000 mammals across large scales. Native species make up the majority of the birds preyed upon b by cats. On average, only 33% of bird prey items identiﬁed to 600 Median=12.3 billion species were non-native species in 10 studies with 438 specimens Estimate frequency 500 of 58 species (Supplementary Table S3). For mammals, patterns 400 of predation on native and non-native species are less clear and 300 appear to vary by landscape type. In densely populated urban 200 areas where native small mammals are less common, non-native 100 species of rats and mice can make up a substantial component of 0 mammalian prey29. However, studies of mammals in suburban and rural areas found that 75–100% of mammalian prey were 0 5,000 10,000 15,000 20,000 25,000 native mice, shrews, voles, squirrels and rabbits26,30,31. Further Mortality (millions) research of mammals is needed to clarify patterns of predation by Figure 1 | Estimates of cat predation on US birds and mammals. both owned and un-owned cats on native and non-native (a) Probability distribution of estimated bird mortality caused by all free- mammals, and across different landscape types. ranging cats in mainland areas of the contiguous United States. Sensitivity analyses indicate that additional research of (b) Probability distribution of estimated mammal mortality caused by all un-owned cats will continue to improve precision of mortality free-ranging cats in mainland areas of the contiguous United States. estimates. Our ﬁnding that un-owned cat population size and Number of owned pet cats Mammal Proportion of pet cats outdoors Bird Proportion of outdoor pet cats hunting Outdoor pet cat predation rate Detectability correction for pet cat prey items Number of un-owned cats Proportion of un-owned cats hunting Un-owned cat predation rate 0% 10 % 20 % 30 % 40 % 50 % 60 % Figure 2 | Factors explaining uncertainty in estimates of wildlife mortality from cat predation. Amount of variation in estimates of wildlife mortality in the contiguous United States contributed by each parameter in the cat predation model (percentages represent adjusted R2 values from multiple regression models). 4 NATURE COMMUNICATIONS | 4:1396 | DOI: 10.1038/ncomms2380 | www.nature.com/naturecommunications & 2013 Macmillan Publishers Limited. All rights reserved. NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2380 ARTICLE predation rate explained the greatest variation in mortality substantial in all parts of the world where free-ranging cats estimates reﬂects the current lack of knowledge about un-owned occur. This mortality is of particular concern within the context cats. No precise estimate of the un-owned cat population exists of steadily increasing populations of owned cats, the potential for for the United States because obtaining such an estimate is cost increasing populations of un-owned cats12, and an increasing prohibitive, and feral un-owned cats are wary of humans and tend abundance of direct and indirect mortality sources that threaten to be solitary outside of urban areas. In addition, human wildlife in the United States and globally. subsidized colonies of un-owned cats are maintained without widespread public knowledge. For example, in Washington DC Methods alone there are 4300 managed colonies of un-owned cats and an Literature search. We searched JSTOR, Google Scholar, and the Web of Science unknown number of unmanaged colonies. Population size database (formerly ISI Web of Science) within the Web of Knowledge search estimates can be improved by incorporating observations of engine published by Thomson Reuters to identify studies that document cat pre- dation on birds and mammals. We initially focused this search on US studies, but free-ranging cats into a wildlife mortality reporting database23. due to a limited sample of these studies, we expanded the search to include pre- Context for the population impact of a mortality source depends dation research from other temperate regions. We also searched for studies pro- on comparing mortality estimates to estimates of population viding estimates of cat population sizes at the scale of the contiguous United States abundance of individual species. However, continental-scale and for US studies that estimate the proportion of owned cats with outdoor access and the proportion of cats that hunt wildlife. The search terms we used included: estimates of wildlife population abundance are uncertain due to ‘domestic cat’ in combination with ‘predation,’ ‘prey,’ ‘diet,’ ‘food item’ and spatio-temporal variation in numbers. For mammals, clariﬁcation ‘mortality’; all previous terms with ‘domestic cat’ replaced by ‘Felis catus,’ ‘feral,’ of the population impacts of cat predation is hindered by the ‘stray,’ ‘farm,’ ‘free-ranging,’ and ‘pet’; ‘trap-neuter-return colony’; ‘TNR colony’; absence of nationwide population estimates. For all North and ‘cat predation’ in combination with ‘wildlife,’ ‘bird,’ ‘mammal,’ and ‘rodent’. American land birds, the group of species most susceptible to We checked reference lists of articles to identify additional relevant studies. Lead authors of three studies were also contacted to enquire whether they knew of mainland cat predation (Supplementary Table S3), existing ongoing or completed unpublished studies of cat predation in the United States. estimates range from 10–20 billion individuals in North America32. A lack of detail about relative proportions of Classiﬁcation of cat ranging behaviour. We grouped studies based on the ran- different bird species killed by cats and spatio-temporal variation ging behaviour of cats investigated. We deﬁned owned cats to include owned cats in of these proportions makes it difﬁcult to identify the species and both rural and urban areas that spend at least some time indoors and are also populations that are most vulnerable. The magnitude of our granted outdoor access. We deﬁned un-owned cats to include all un-owned cats mortality estimates suggest that cats are likely causing population that spend all of their time outdoors. The un-owned cat group includes semi-feral cats that are sometimes considered pets (for example, farm/barn cats and strays declines for some species and in some regions. Threatened and that are fed by humans but not granted access to habitations), cats in subsidized endangered wildlife species on islands are most susceptible to the (including TNR) colonies, and cats that are completely feral (that is, completely effects of cat predation, and this may also be true for vulnerable independent and rarely interacting with humans). We did not classify cats by species in localized mainland areas5 because small numbers of landscape type or whether they receive food from humans because the amount of time cats spend outdoors is a major determinant of predation rates33,34 and fatalities could cause signiﬁcant population declines. Threatened because predation is independent of whether cats are fed by humans6,34,35. species in close proximity to cat colonies—including managed TNR colonies11,12—face an especially high level of risk; therefore, Study inclusion criteria. Studies were only included if: (1) they clearly reported cat cat colonies in such locations comprise a wildlife management ranging behaviour (that is, a description of whether cats were owned or un-owned priority. Claims that TNR colonies are effective in reducing cat and whether they were outdoor cats or indoor-outdoor cats), and (2) the group of populations, and, therefore, wildlife mortality, are not supported cats investigated ﬁt exclusively into one of the two groups we deﬁned above (that is, by peer-reviewed scientiﬁc studies11. we excluded studies that lumped owned and un-owned cats in a single predation Our estimates should alert policy makers and the general rate estimate). For some studies, we extracted a portion of data that met these criteria but excluded other data from cats with unknown ranging behaviour. We public about the large magnitude of wildlife mortality caused by only included mainland and large island (New Zealand and United Kingdom) free-ranging cats. Structured decisions about actions to reduce predation studies, because cat predation on small islands is often exceptionally wildlife mortality require a quantitative evidence base. We high36,37 and focused on colony nesting seabirds38. We excluded studies from provide evidence of large-scale cat predation impacts based on outside temperate regions and those with predation rate estimates based on fewer than 10 cats, o1 month of sampling, or on cats that were experimentally systematic analysis of multiple data sources. Future speciﬁc manipulated (for example, by ﬁtting them with bells or behaviour altering bibs). management decisions, both in the United States and globally, We included studies that used cat owners’ records of prey returns, but we excluded must be further informed by ﬁne scale research that allows those that asked owners to estimate past prey returns because such questionnaires analysis of population responses to cats and assessment of the may lead to bias in estimation of predation rates39. (For a list of all included and excluded studies, see Supplementary Table S1). success of particular management actions. We are not suggesting that other anthropogenic threats that kill fewer individuals are biologically unimportant. Virtually nothing is known about the Data extraction and standardization of predation rates. Most studies report an estimate of cat predation rate (that is, daily, monthly or annual prey killed per cat) cumulative population impacts of multiple mortality sources. or present data that allowed us to calculate this rate. When studies only reported Furthermore, comparison of total mortality numbers has limited predation rate estimates for all wildlife combined, we calculated separate predation use for prioritization of risks and development of conservation rates by extracting taxa-speciﬁc prey counts from tables or ﬁgures and multiplying objectives. Combining per species estimates of mortality with the total predation rate by the proportion of prey items in each taxon. If taxa- population size estimates will provide the greatest information speciﬁc counts were not provided, we directly contacted authors to obtain this information. For studies that presented low, medium and high estimates or low and about the risk of population-level impacts of cat predation. high estimates, we used the medium and average values, respectively. For studies Although our results suggest that owned cats have relatively less that presented more than one predation estimate for cats with similar ranging impact than un-owned cats, owned cats still cause substantial behaviour (for example, owned cats in rural and urban areas), we calculated the wildlife mortality (Table 2); simple solutions to reduce mortality average predation rate. Nearly all studies of un-owned cats report numbers or frequencies of occurrence caused by pets, such as limiting or preventing outdoor access, of different taxa in stomachs and/or scats. For studies reporting numbers of prey should be pursued. Efforts to better quantify and minimize items, we estimated annual predation rates by assuming one stomach or scat mortality from all anthropogenic threats are needed to increase sample represented a cat’s average daily prey intake (for example, an average of one sustainability of wildlife populations. prey item per stomach or scat ¼ 365 prey per cat per year). This assumption likely resulted in conservative estimates because cats generally digest prey within 12 h The magnitude of wildlife mortality caused by cats that we (ref. 28) and can produce two or more scats each day29. For studies reporting report here far exceeds all prior estimates. Available evidence occurrence frequencies of prey items, we assumed this proportion represented a suggests that mortality from cat predation is likely to be cat’s average daily prey intake (for example, a 10% bird occurrence rate ¼ 0.1 bird NATURE COMMUNICATIONS | 4:1396 | DOI: 10.1038/ncomms2380 | www.nature.com/naturecommunications 5 & 2013 Macmillan Publishers Limited. All rights reserved. ARTICLE NATURE COMMUNICATIONS | DOI: 10.1038/ncomms2380 per stomach or scat ¼ 36.5 birds per cat per year). This assumption results in coarse 6. Hawkins, C. C., Grant, W. E. & Longnecker, M. T. in Proceedings of the 4th predation rate estimates, but estimates from this approach are even more International Urban Wildlife Symposium. (eds Shaw, W. W., Harris, L. K. & conservative than those from the ﬁrst assumption because many stomachs and Vandruff, L.) 164–170 (University of Arizona, Tucson, AZ, 2004). scats undoubtedly included more than one bird or mammal. 7. van Heezik, Y., Smyth, A., Adams, A. & Gordon, J. Do domestic cats impose an Predation rate estimates from many studies were based on continuous year- unsustainable harvest on urban bird populations? Biol. Conserv. 143, 121–130 round sampling or multiple sampling occasions covering all seasons. However, (2010). seasonal coverage of some studies was incomplete. To generate full-year predation 8. Churcher, P. B. & Lawton, J. H. Predation by domestic cats in an English rate estimates in these cases, we adjusted partial-year predation estimates according to the average proportion of prey taken in each month as determined from year- village. J. Zool. London 212, 439–455 (1987). round studies reporting monthly data (birds and mammals8,33, birds only7,40). For 9. Baker, P. J., Molony, S., Stone, E., Cuthill, I. C. & Harris, S. Cats about town: is partial-year estimates from the northern hemisphere, we offset monthly estimates predation by free-ranging pet cats (Felis catus) likely to affect urban bird from southern hemisphere studies by 6 months. The ﬁnal annual predation rate populations? IBIS 150(Suppl. 1): 86–99 (2008). estimates for all studies are presented in Supplementary Table S1. The year-round 10. Balogh, A. L., Ryder, T. B. & Marra, P. P. Population demography of Gray studies we used represent different geographical regions (for birds—England, Catbirds in the suburban matrix: sources, sinks, and domestic cats. Kansas (US), Australia and New Zealand; for mammals—England and Australia) J. Ornitholol. 152, 717–726 (2011). with varying climates and slightly varying seasonal patterns of predation. For both 11. Longcore, T., Rich, C. & Sullivan, L. M. Critical assessment of claims regarding birds and mammals, averaging across full-year studies resulted in higher management of feral cats by trap-neuter-return. Conserv. Biol. 23, 887–894 proportions of predation in the spring and summer compared with fall and winter, (2009). an expected pattern for much of the United States. The reference studies we used, 12. Lepczyk, C. A. et al. What conservation biologists can do to counter therefore, provide a reasonable baseline for correcting to full-year mortality trap-neuter-return: response to Longcore et al.. Conserv. Biol. 24, 627–629 estimates. This approach greatly improves upon the assumption that mortality is (2010). negligible during the period of the year not covered by sampling. 13. Gill, F. B. Ornithology, 2nd edn. (W.H. Freeman Publishers, 1994). 14. Dauphiné, N. & Cooper, R. J. Impacts of free-ranging domestic cats Quantiﬁcation of annual mortality from cat predation. We estimated wildlife (Felis catus) on birds in the United States: a review of recent research with mortality in the contiguous United States by multiplying data-derived probability conservation and management recommendations. Proceedings of the Fourth distributions of predation rates by distributions of estimated cat abundance, International Partners in Flight Conference: Tundra to Tropics 205–219 following41. Quantiﬁcation was conducted separately for owned and un-owned cats (Partners in Flight, US, 2009). and for birds and mammals. As there was a relatively small sample of US studies 15. Banks, R. C. Human related mortality of birds in the United States. Special that estimated predation rates (n ¼ 14 and 10 for birds and mammals, respectively), Scientiﬁc Report—Wildlife N. 215 (US Dept. of the Interior—Fish and Wildlife we repeated calculations using predation rate distributions that were augmented Service, 1979). with predation rates from Europe and all temperate zones. However, we only used 16. Erickson, W. P., Johnson, G. D. & Young, Jr D. P. A summary and comparison studies from the contiguous United States to construct all other probability of bird mortality from anthropogenic causes with an emphasis on collisions. distributions (listed below). Tech. Rep PSW-GTR-191 1029–1042 (US Dept. of Agriculture—Forest Service, We estimated mortality using the following model of cat predation: 2005). Annual mortality from owned catsðmpÞ ¼ npcpodpphpprcor ð1Þ 17. Klem, Jr D. Avian mortality at windows: the second largest human source of bird mortality on earth. Proceedings of the Fourth International Partners in Flight Conference: Tundra to Tropics 244–251 (Partners in Flight, 2009). Annual mortality from unowned catsðmf Þ ¼ nfcpfhfpr ð2Þ 18. Coleman, J. S. & Temple, S. A. On the Prowl. Wisconsin Nat. Res. Mag. (1996). 19. Pimentel, D., Greiner, A. & Bashore, T. Economic and environmental costs of Total annual mortality from all cats ¼ mp þ mf ð3Þ pesticide use. Arch. Environ. Con. Tox. 21, 84–90 (1991). where npc is the number of owned cats in the contiguous United States, pod is the 20. Manville, II A. Towers, turbines, power lines, and buildings—steps being proportion of owned cats granted outdoor access, pph is the proportion of outdoor taken by the U.S. Fish and Wildlife Service to avoid or minimize take of owned cats that hunt wildlife, ppr is the annual predation rate by owned cats, cor is migratory birds at these structures. Proceedings of the Fourth International a correction factor to account for owned cats not returning all prey to owners, nfc Partners in Flight Conference: Tundra to Tropics 262–272 (Partners in Flight, is the number of un-owned cats in the contiguous United States, pfh is the US, 2009). proportion of un-owned cats that hunt wildlife, and fpr is the annual predation 21. Longcore, T. et al. An estimate of mortality at communication towers in the rate by un-owned cats. From the probability distribution of each parameter United States and Canada. PLoS one 7, e34025 (2012). (see Table 1 and Supplementary Methods for details about the speciﬁc probability 22. Pullin, A. S. & Stewart, G. B. Guidelines for systematic review in conservation distributions used), we randomly drew one value and used the above formulas to and environmental management. Conserv. Biol. 20, 1647–1656 (2006). calculate mortality. Random draws were made using distribution functions in 23. Loss, S. R., Will, T. & Marra, P. P. Direct human-caused mortality of birds: Programme R (rnorm and runif commands for normal and uniform distributions, improving quantiﬁcation of magnitude and assessment of population impact. respectively). We conducted 10,000 random draws to estimate a potential Front. Ecol. Environ. 20, 357–364 (2012). range of annual predation on each wildlife taxa. For all analyses, we report median 24. Henderson, R. W. Consequences of predator introductions and habitat mortality estimates and lower and upper estimates bracketing the central 95% of destruction on amphibians and reptiles in the Post-Columbus West Indies. values. Caribb. J. Sci. 28, 1–10 (1992). 25. Nilsson, N. N. The role of the domestic cat in relation to game birds in the Sensitivity analyses. We used multiple linear regression analysis to assess how Willamette Valley, Oregon. Thesis (Oregon State College, 1940). much variance in mortality estimates was explained by the probability distribution 26. Llewellyn, L. L. & Uhler, F. M. The foods of fur animals of the Patuxent for each parameter. We treated total mortality estimates as the dependent variable Research Refuge, Maryland. Am. Midl. Nat. 48, 193–203 (1952). (n ¼ 10,000) and we deﬁned a predictor variable for each parameter that consisted 27. Parmalee, P. W. 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Res. 4, 249–262 (1977). not necessarily represent the views of the Smithsonian or US Fish and Wildlife Service. 37. Bramley, G. N. A small predator removal experiment to protect North Island All data used for this analysis is available in the Supplementary Materials. Weka (Gallirallus australis greyi) and the case for single-subject approaches in determining agents of decline. NZ J. Ecol. 20, 37–43 (1996). 38. Bonnaud, E. et al. The diet of feral cats on islands: a review and a call for more Author contributions S.R.L. designed the study, collected and analysed data, and wrote the paper. T.W. and studies. Biol. Conserv. 13, 581–603 (2011). P.P.M. designed the study and contributed to paper revisions. All authors discussed the 39. Tschanz, B., Hegglin, D., Gloor, S. & Bontadina, F. Hunters and non-hunters: results and commented on the manuscript. skewed predation rate by domestic cats in a rural village. Eur. J. Wildl. Res. 57, 597–602 (2011). 40. Fiore, C. A. Domestic cat (Felis catus) predation of birds in an urban environment. Thesis (Wichita State University, 2000). Additional information Supplementary Information accompanies this paper on http://www.nature.com/ 41. Blancher, P. J. Estimated number of birds killed by house cats (Felis catus) in naturecommunications Canada. (Avian Conservation and Ecology, in press). Competing ﬁnancial interests: The authors claim no competing ﬁnancial interests Acknowledgements associated with this paper. S.R.L. was supported by a postdoctoral fellowship funded by the US Fish and Wildlife Reprints and permission information is available online at http://npg.nature.com/ Service through the Smithsonian Conservation Biology Institute’s Postdoctoral Fellow- reprintsandpermissions/ ship programme. P. Blancher provided insight for development of the model of cat predation magnitude, and R. Kays, C. Lepczyk and Y. van Heezik provided raw data from How to cite this article: Loss S.R. et al. The impact of free-ranging domestic cats on their publications. C. Machtans facilitated data sharing, and participants in the 2011 wildlife of the United States. Nat. Commun. 4:1396 doi: 10.1038/ncomms2380 (2012). NATURE COMMUNICATIONS | 4:1396 | DOI: 10.1038/ncomms2380 | www.nature.com/naturecommunications 7 & 2013 Macmillan Publishers Limited. All rights reserved. DOI: 10.1038/ncomms3961 Corrigendum: The impact of free-ranging domestic cats on wildlife of the United States Scott R. Loss, Tom Will & Peter P. Marra Nature Communications 4:1396 doi:10.1038/ncomms2380 (2013); Published 29 Jan 2013; Updated 12 Dec 2013 The original version of this Article contained incorrect estimates of the number of animals killed by free-ranging domestic cats, which were based on 90% conﬁdence levels rather than the stated 95% conﬁdence levels. Reanalysis of the original data after publication using 95% conﬁdence levels resulted in larger ranges for the estimated number of animals killed. For example, in the ﬁfth sentence of the Abstract, the number of birds killed by free-ranging domestic cats required revision from 1.4–3.7 billion to 1.3–4.0 billion and the number of mammals killed by free-ranging domestic cats required revision from 6.9–20.7 billion to 6.3–22.3 billion. The estimates have now been corrected throughout the PDF and HTML versions of the Article. Unrelated to the changes above, four estimates of cat predation rates on wildlife from temperate zone studies in Supplementary Table S1 were based on partial year values that had not been adjusted to year-round estimates. The values have now been revised in Supplementary Table S1. The original conclusions are not affected by any of the above changes. NATURE COMMUNICATIONS | 4:2961 | DOI: 10.1038/ncomms3961 | www.nature.com/naturecommunications 1 & 2013 Macmillan Publishers Limited. All rights reserved.

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