In April, Conservation Biology published a comment authored by Christopher A. Lepczyk, Nico Dauphiné, David M. Bird, Sheila Conant, Robert J. Cooper, David C. Duffy, Pamela Jo Hatley, Peter P. Marra, Elizabeth Stone, and Stanley A. Temple. In it, the authors “applaud the recent essay by Longcore et al. (2009) in raising the awareness about trap-neuter-return (TNR) to the conservation community,” [1] and puzzle at the lack of TNR opposition among the larger scientific community:

“…it may be that conservation biologists and wildlife ecologists believe the issue of feral cats has already been studied enough and that the work speaks for itself, suggesting that no further research is needed.”

In fact, “the work”—taken as a whole—is neither as rigorous nor as conclusive as Lepczyk et al. suggest. And far too much of it is plagued by exaggeration, misrepresentations, errors, and obvious bias. In a previous post, I presented examples of researchers drawing big conclusions from small sample sizes. Here, I’ll discuss the important distinction between compensatory and additive predation—a point too often left out of the feral cat/TNR discussion.

Sins of Omission
Focusing on the number of prey injured or killed by cats, without also recognizing that there are different types of predation, implies that each and every bird, mammal, reptile, etc. is destined to be part of its species’ breeding population. Of course, that’s not at all how things work out in the natural world—with or without predation by cats.

And yet, numerous studies [2-10], reviews [11], and other published papers [12-14] fail to acknowledge the critical difference between compensatory predation (in which prey would have died even in the absence of a particular predator, due to illness, starvation, other predators, etc.) and additive predation (in which healthy prey are killed). It’s the difference between, as Beckerman et al. put it, the “doomed surplus hypothesis” and the “hapless survivor hypothesis.” [15]

This is a critical point when it comes to connecting predation rates (from cats or any other predator) to population impacts. The more additive the predation, the greater the potential impact on population numbers. Purely compensatory predation, on the other hand, is less likely to affect overall populations. Of course, the connection is seldom so simple and direct, and a number of factors (e.g., habitat area and type, base population numbers, etc.) influence the ultimate outcome—making it quite difficult to tease out specific causal relationships. Nevertheless, if we want to better understand the impact of free-roaming cats on wildlife, we cannot ignore the distinction between—and inherent implications of—these two types of predation.

Honorable Mentions
Although Churcher and Lawton failed to mention the distinction between compensatory and additive predation in their now-classic “English village” study [4], Churcher later suggested that their findings were largely in the compensatory category: “If the cats weren’t there, something else would be killing the sparrows or otherwise preventing them from breeding.” [16]

Woods et al. don’t address the topic directly, but warn against drawing direct connections between predation numbers and potential effects on population dynamics:

“Our estimates of the total numbers of animals brought home by cats throughout Britain should be treated with requisite caution and these figures do not equate to an assessment of the impact of cats on wildlife populations.” [3]

Unfortunately, other researchers have used this study to make exactly that connection. In “Critical Assessment,” for example, Longcore et al. cite Woods et al. (along with Lepczyk et al. 2003, the subject of a future post) when they write, “evidence indicates that cats can play an important role in fluctuations of bird populations.” [11]

Under-Compensating?
In their 2008 study, Baker et al. found that “birds killed by cats in this study had significantly lower fat and pectoral muscle mass scores than those killed by collisions,” [17] suggesting that they may have been among the “doomed surplus” portion of the population. Similar results were reported eight years earlier by Møller and Erritzøe, who found that “small passerine birds falling prey to cats had spleens that were significantly smaller than those of conspecifics that died for other reasons,” concluding ultimately that the birds killed by cats “often have a poor health status.” [18]

But Baker et al. express caution about their findings:

“…the distinction between compensatory and additive mortality does… become increasingly redundant as the number of birds killed in a given area increases: where large numbers of prey are killed, predators would probably be killing a combination of individuals with poor and good long-term survival chances.”

Whatever their concerns, it must be noted that Baker et al. inflated their predation numbers by a factor of 3.3 on the basis of Kays and DeWan’s dubious conclusions [9] (which I discussed in some detail previously). Doing so raises considerable doubts about any level of “redundancy,” as well the authors’ suggestion that cat predation in the area might be “creating a dispersal sink for more productive neighboring areas.” [19] (Such “sinks” can occur when predation outstrips local prey populations, requiring that prey be “recruited” from surrounding areas.)

Implications
Given all the work that’s been done on cat predation, one might expect the subject of compensatory predation to be addressed more fully and more often. By omitting this important issue from the feral cat/TNR discussion, researchers portray a situation both simpler and harsher (in terms of what it implies about the impact of free-roaming cats) than reality suggests. Whether or not such omissions are intentional, I cannot say. I do, however, find it curious—what’s included compared to what’s left out, and by whom.

References
1. Lepczyk, C.A., et al., “What Conservation Biologists Can Do to Counter Trap-Neuter-Return: Response to Longcore et al.” Conservation Biology. 2010. 24(2): p. 627-629.

2. Coleman, J.S. and Temple, S.A., On the Prowl, in Wisconsin Natural Resources. 1996, Wisconsin Department of Natural Resources: Madison, WI. p. 4–8. http://dnr.wi.gov/wnrmag/html/stories/1996/dec96/cats.htm

3. Woods, M., McDonald, R.A., and Harris, S., “Predation of wildlife by domestic cats Felis catus in Great Britain.” Mammal Review. 2003. 33(2): p. 174-188.

4. Churcher, P.B. and Lawton, J.H., “Predation by domestic cats in an English village.” Journal of Zoology. 1987. 212(3): p. 439-455.

5. Coleman, J.S. and Temple, S.A., “Rural Residents’ Free-Ranging Domestic Cats: A Survey.” Wildlife Society Bulletin. 1993. 21(4): p. 381–390.

6. Coleman, J.S. and Temple, S.A., Effects of Free-Ranging Cats on Wildlife: A Progress Report, in Fourth Eastern Wildlife Damaage Control Conference. 1989: University of Nebraska—Lincoln. p. 8–12. http://digitalcommons.unl.edu/ewdcc4/7

7. Hawkins, C.C., Impact of a subsidized exotic predator on native biota: Effect of house cats (Felis catus) on California birds and rodents. 1998. PhD Dissertation, Texas A&M University.

8. Hawkins, C.C., Grant, W.E., and Longnecker, M.T., “Effects of Subsidized House Cats on California Birds and Rodents.” Transactions of the Western Section of the Wildlife Society. 1999. 35: p. 29–33.

9. Kays, R.W. and DeWan, A.A., “Ecological impact of inside/outside house cats around a suburban nature preserve.” Animal Conservation. 2004. 7(3): p. 273-283.

10. Lepczyk, C.A., Mertig, A.G., and Liu, J., “Landowners and cat predation across rural-to-urban landscapes.” Biological Conservation. 2003. 115(2): p. 191-201.

11. Longcore, T., Rich, C., and Sullivan, L.M., “Critical Assessment of Claims Regarding Management of Feral Cats by Trap–Neuter–Return.” Conservation Biology. 2009. 23(4): p. 887–894.

12. Dauphiné, N. and Cooper, R.J., Impacts of Free-ranging Domestic Cats (Felis catus) on birds in the United States: A review of recent research with conservation and management recommendations, in Fourth International Partners in Flight Conference: Tundra to Tropics. 2010. p. 205–219

13. Coleman, J.S., Temple, S.A., and Craven, S.R., Cats and Wildlife: A Conservation Dilemma. 1997, University of Wisconsin, Wildlife Extension. http://forestandwildlifeecology.wisc.edu/wl_extension/catfly3.htm

14. Andersen, M.C., Martin, B.J., and Roemer, G.W., “Use of matrix population models to estimate the efficacy of euthanasia versus trap-neuter-return for management of free-roaming cats.” Journal of the American Veterinary Medical Association. 2004. 225(12): p. 1871-1876.

15. Beckerman, A.P., Boots, M., and Gaston, K.J., “Urban bird declines and the fear of cats.” Animal Conservation. 2007. 10(3): p. 320-325.

16. n.a., What the Cat Dragged In, in Catnip. 1995, Tufts University School of Veterinary Medicine: Boston, MA. p. 4–6

17. Baker, P.J., et al., “Cats about town: is predation by free-ranging pet cats Felis catus likely to affect urban bird populations?“ Ibis. 2008. 150: p. 86-99.

18. Møller, A.P. and Erritzøe, J., “Predation against birds with low immunocompetence.” Oecologia. 2000. 122(4): p. 500-504.