Exceptional Predator

10 Feb 2011

Using Google to translate the page’s contents, it seems this bird—despite “mock[ing] the cat and with loud cries of diving at him from the branches of acacia”—was yet another one that got away.

In the third edition of his massive book Ornithology—“the classic text for the undergraduate ornithology course,” according to the description on Amazon.com—Frank Gill writes:

“Natural predators are a major source of annual mortality among birds, especially nestlings, incubating females, and young birds in their first year. Relentless predation is a driving force of natural selection for escape behaviors, camouflage plumage, and social behavior. With some conspicuous exceptions, however, predators don’t limit or regulate the bird populations on which they prey [1]. Instead, they take weak, sick, and young birds, many of which are part of the surplus that exceeds locally limiting food supplies.” [2, p 545]

For Gill, it seems, it’s all very straightforward; this, after all, is how Nature works. (It should be noted that, just one paragraph later, the author makes a clear distinction between islands and other habitats: “The endangerment and extinction of island birds by introduced predators is a conspicuous exception to the statement that predators don’t limit bird populations.”)

Unequal Treatment Under the (Natural) Law
Nobody opposed to TNR would deny that cats are predators—so why won’t they admit that the birds and other wildlife killed by cats are generally among, as Gill puts it, the “weak, sick, and young”?

The Carolina Raptor Center, for example, describes the role of predatory birds targeting bird feeders this way:

“Songbirds are part of the food chain just like other animals and their predators are going to look for the easiest targets. The birds that hawks are usually able to catch at feeders are the slow and sick ones. The strong and healthy ones escape, allowing their survival to produce more healthy babies.”

Cats, however, are a different matter altogether. According to the Carolina Raptor Center, they “kill a lot more birds then hawks do because hawks only kill for food, where cats kill for the sport of it.” I’ve never seen any scientific evidence to support such a claim, which may explain why so many have instead argued—again, without any support—that cats compete with raptors for food.

Who’s Crazed Now?
It wasn’t Gill’s book that got me thinking about this, though, but a comment posted last month on the Bountiful Films blog, following the release of their documentary Cat Crazed.

After listening to a CBC interview with director Maureen Palmer, whose “science” was clearly coming straight from the American Bird Conservancy, I posted a comment, stating in part:

“What you won’t find [from organizations opposing TNR] is any mention of the studies that show rather convincingly that birds killed by cats tend to be unhealthy compared to those killed by building collisions, say. Even high predation rates do not equate to population declines—as many scientists have noted.”

I also included a link to my “Predatory Blending” post. Which promptly drew fire from somebody calling him/herself “Catbird”:

“Where cats cause documented extinctions and extirpations, cat predation is additive (e.g., Hawkins 1998, Crooks and Soule 1999, Nogales et al. 2004). Researchers are interested in knowing if some cat predation is compensatory (that is, killing animals that would die anyway) (Beckerman et al. 2007, Baker et al. 2008, van Heezik et al. 2010). The purported evidence of compensatory predation is a study showing that cat-killed birds have smaller spleens (indicating that they are less healthy) than birds killed by other sources (e.g., windows) (Moller and Erritzoe 2000). Other researchers found that birds killed by cats had less fat reserves and lower muscle mass than those killed in collisions (Baker et al. 2008), but warned against assuming that this corresponded with lower fitness of these individuals. In neither instance is it possible to conclude that individuals killed by cats would have died otherwise.”

Actually, Møller & Erritzoe don’t suggest that the birds captured by cats “would have died otherwise.” But, they are quite clear about the implications of their research:

“The present study has suggested that predators like the domestic cat may select against individuals with a weak immune system, leaving a disproportionate fraction of immunocompetent individuals as survivors.” [3]

What Møller & Erritzoe observed is very much in line with what Gill describes as typical predatory behavior.

Still, though, I’m not necessarily surprised with Catbird’s “interpretation” of the science, given his/her comments and tone elsewhere in the discussion. What’s far more troubling is that so few studies on the predatory habits of cats address the topic in any meaningful way.

Sins of Omission
Take that 2008 study by Baker et al., for example. The authors are, just as Catbird suggests, quite cautious about their findings:

“The distinction between compensatory and additive mortality does, however, 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. The predation rates estimated in this study would suggest that this was likely to have been the case for some species on some sites.” [4]

But, as I’ve pointed out previously, the authors’ predation rates are inflated—in part due to their unquestioning application of the dubious multiplier proposed by Kays and DeWan. [5] Baker et al. also use low estimates of breeding density—all of which combines to diminish the apparent level of compensatory predation. Were these estimates adjusted to better reflect the conditions at the site, the “redundancy” the authors refer to would be reduced considerably.

(Frankly, Baker and his colleagues seemed quite eager to demonstrate that Bristol’s cats were negatively affecting bird populations; in an earlier study, they suggested—based, I would argue, on insufficient information—that the area might be a “dispersal sink for more productive neighboring areas.” [6])

On the other hand, at least Baker et al. acknowledge Møller and Erritzoe’s work. Many other studies don’t even go that far.

Coleman and Temple, [7] for example, failed to consider the role of compensatory predation—despite the fact that they cite sources/studies that do. [8–10] And Temple himself addresses this very topic in his 1987 paper Do Predators Always Capture Substandard Individuals Disproportionately From Prey Populations?

Using a trained Red-tailed hawk to prey on eastern chipmunks, cottontail rabbits, and gray squirrels, Temple developed the “proposition that substandard individuals are captured disproportionately when the type of prey is relatively difficult to capture but not when it is relatively easy to capture.” [11]

Which seems a very fitting description for the general case of a cat attempting to capture an adult bird. (Ground-nesting and ground-feeding birds would likely be easier prey, though Hawkins’ PhD dissertation work [12] suggests that even this assumption deserves careful scrutiny.)

Longcore et al. never mention Møller and Erritzoe (one of many shortcomings I address in “Reassessment”); neither do Dauphine and Cooper. [13]

And ABC doesn’t go near the topic of compensatory predation. (Ironic since, unlike cats, most of the “threats to birds” listed by ABC (e.g., pesticides, pollution, oil spills, collisions with towers, buildings, wind turbines, and power lines, etc.) are clearly nondiscriminatory in terms of bird mortality.)

• • •

Is it any wonder that a reasonable discussion about the impacts of free-roaming cats on wildlife is so elusive? The same stakeholders that condemn these cats for their predatory nature too often refuse to acknowledge the nature of predation itself.

Literature Cited
1. Newton, I., Population limitation in birds. 1998, San Diego: Academic.

2. Gill, F.B., Ornithology. 3rd ed. 2007, New York: W.H. Freeman.

3. Møller, A.P. and Erritzøe, J., “Predation against birds with low immunocompetence.” Oecologia. 2000. 122(4): p. 500–504. http://www.springerlink.com/content/ghnny9mcv016ljd8/

4. 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. http://www.ingentaconnect.com/content/bsc/ibi/2008/00000150/A00101s1/art00008

5. 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. http://dx.doi.org/10.1017/S1367943004001489

www.nysm.nysed.gov/staffpubs/docs/15128.pdf

6. Baker, P.J., et al., “Impact of predation by domestic cats Felis catus in an urban area.” Mammal Review. 2005. 35(3/4): p. 302-312. http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2907.2005.00071.x/abstract

7. Coleman, J.S. and Temple, S.A., How Many Birds Do Cats Kill?, in Wildlife Control Technology. 1995. p. 44. http://www.wctech.com/WCT/index99.htm

8. Fitzgerald, B.M., Diet of domestic cats and their impact on prey populations, in The Domestic cat: The biology of its behaviour, D.C. Turner and P.P.G. Bateson, Editors. 1988, Cambridge University Press: Cambridge; New York. p. 123–147.

9. Churcher, P.B. and Lawton, J.H., “Predation by domestic cats in an English village.” Journal of Zoology. 1987. 212(3): p. 439-455. http://dx.doi.org/10.1111/j.1469-7998.1987.tb02915.x

10. Errington, P.L., “Notes on Food Habits of Southwestern Wisconsin House Cats.” Journal of Mammalogy. 1936. 17(1): p. 64–65. http://www.jstor.org/stable/1374554

11. Temple, S.A., “Do Predators Always Capture Substandard Individuals Disproportionately From Prey Populations?“ Ecology. 1987. 68(3): p. 669–674. http://www.esajournals.org/doi/abs/10.2307/1938472

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

13. Dauphine, 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. 2009. p. 205–219. http://www.pwrc.usgs.gov/pif/pubs/McAllenProc/articles/PIF09_Anthropogenic%20Impacts/Dauphine_1_PIF09.pdf

Revisiting “Reassessment”

5 Jan 2011

“Reassessment: A Closer Look at ‘Critical Assessment of Claims Regarding Management of Feral Cats by Trap-Neuter-Return’” has been revised and expanded!

This paper, a brief review and critique of the essay “Critical Assessment of Claims Regarding Management of Feral Cats by Trap-Neuter-Return” by Travis Longcore, Catherine Rich, and Lauren M. Sullivan, now includes sections on Toxoplasma gondii, the mesopredator release phenomenon, and more. In addition, links and downloadable PDFs have been added to the list of references.

Over the past year, “Critical Assessment” has gotten a great deal of traction among TNR opponents, despite its glaring omissions, blatant misrepresentations, and obvious bias. “Reassessment”—intended to be a resource for a broad audience, including, wildlife and animal control professionals, policymakers, and the general public—shines a bright spotlight on these shortcomings, thereby bringing the key issues back into focus.

Act Locally
Politics is, as they say, local. This is certainly true of the debate surrounding TNR. Policies endorsing TNR, the feeding of feral cats, etc. typically begin with “Town Hall” meetings, or even meetings of neighborhood associations. “Reassessment” provides interested parties with a rigorous, science-based counter-argument to those using “Critical Assessment” as a weapon against feral cats/TNR.

So, once you’ve had a look for yourself, please share generously! Together, we can—in keeping with the mission of Vox Felina—improve the lives of feral cats through a more informed, conscientious discussion of feral cat issues in general, and TNR in particular.

Download PDF

The Work Speaks—Part 4: Mean Spirited

4 Jun 2010

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 Part 3 of this series, I discussed the distinction between compensatory and additive predation. Here, I’ll focus on how feral cat/TNR researchers often misuse averages to characterize skewed distributions, and how that error overestimates the impact of free-roaming cats on wildlife.

Something’s Askew
When a data set is skewed, it is inappropriate to use the mean, or average, as a measure of central tendency. The mean should be used only when the data set can be considered normal—the familiar bell curve. As Woods et al put it:

“the simple average number of animals brought home is not a useful measure of central tendency because of the skewed frequency distribution of the numbers of prey items brought home…” [2]

Studies of cat predation routinely reveal a positively skewed distribution; a few cats are responsible for many kills, while many of the cats kill few, if any, prey. So when researchers use the mean to calculate the total number of prey killed by cats in a particular area, they overestimate the cats’ impact.

How common is this? Very [see, for example, 3-9]. Of the many cat predation studies I’ve read, only a few [2, 10, 11] properly account for the skewed nature of this distribution. And others [12-17] often take these inflated figures at face value—as evidence of the impact cats have on wildlife. Published repeatedly, the erroneous estimates take on an undeserved legitimacy.

The proper method for handling skewed distributions involves data transformations, the details of which I won’t go into here. The important point is this: in the case of a positively skewed distribution, the back-transformed mean will always be less than the simple mean of the same data set.

Big Deal
Depending on the particular distribution, the difference between the simple mean and the back-transformed mean can be considerable. Let’s use the 2003 study by Woods et al. [2] to illustrate. In the case of mammals killed and returned home by pet cats, the back-transformed mean was 28.3% less than the simple mean. Or, put another way, the simple mean would have overestimated the number of mammals killed by 39.5%. Similarly, when all prey items were totaled (as depicted in the illustration above), the simple mean would have overestimated the total number off all prey (mammals, birds, herpetofauna, and “others”) by 46.9%.

On the other hand, the figures for birds appear to break the rule mentioned above. In this case, the back-transformed mean (4.1) is actually a bit higher than the simple mean (4.0). How can this be? In order to log-transform the data set, Woods et al. had to first eliminate all the instances where cats returned home no prey—you can’t take the logarithm of 0. So, they were actually working with two data sets. Now, the second data set—which includes only those cats that returned at least one prey item—is also highly positively skewed. As a point of reference, its simple mean was approximately 5.6 birds/cat, which, compared to the back-transformed mean, is an overestimation of 37.5%.

By now, it should be apparent that log-transformed means have another important advantage over simple means: because you have to eliminate those zeros from the data set, you are forced to focus only on the cats that returned prey home—which, of course, is the whole point of such studies! And in the case of this study, Woods et al. found that 20–30% of cats brought home either no birds or no mammals. And 8.6% of the cats brought home no prey at all over the course of the study.

Transforming a data set (and then back-transforming its mean) is simpler than it sounds, but Barratt offers a useful alternative, rule-of-thumb method (one echoed by Fitzgerald and Turner [18]):

“…median numbers of prey estimated or observed to be caught per year are approximately half the mean values, and are a better representation of the average predation by house cats based on these data.” [10]

So, whereas Dauphiné and Cooper (and others) suggest increasing such estimates by factors of two and three (“predation rates measured through prey returns may represent one half to less than one third of what pet cats actually kill…” [14]), they should, in fact, be reducing them by half.

Cat Ownership
There are other instances in which simple averages are used to describe similarly skewed distributions—with similar results. That is, they overestimate a particular characteristic—and not in the cats’ favor.

Cat ownership, for example, is not a normal distribution. Many people own one or two cats; a few people own many cats. This is precisely what Lepczyk et al. found in their 2003 study:

“The total number of free-ranging cats across all landscapes was 656, ranging from 1 to 30 per landowner…” [6]

In fact, about 113 (I’m estimating from the histogram printed in the report) of those landowners owned just one cat apiece. About 70 of them owned two cats. Only one—maybe two—owned 30 cats. And yet, Lepczyk et al. calculate an average of 2.59 cats/landowner (i.e., 656 cats/253 landowners who allow their cats outdoors), thereby substantially overestimating cat ownership—and, in turn, predation rates (which calculations are based upon the average number of cats/landowner).

Lepczyk et al. are not the only ones to make this mistake; several other researchers have done the same. [4, 5, 7-9]

Outdoor Access
The amount of time cats spend outdoors is also highly positively skewed, as is apparent from the 2003 survey conducted by Clancy, Moore, and Bertone. [19] Their work showed that nearly half of the cats with outdoor access were outside for two or fewer hours a day. And 29% were outdoors for less than an hour each day.

Among those researchers to overlook the skewed nature of this distribution are Kays and DeWan, who calculate an average of 8.35 hours/day. This greatly overestimates potential predation, and leads them to conclude—erroneously—that the actual number of prey killed by cats was “3.3 times greater than the rate estimated from prey brought home,” [9] as was discussed previously.

Compound Errors
Clearly, these errors are substantial—in some cases, doubling the apparent impact of cats on wildlife. Of course the errors are even more significant when one inflated figure is multiplied by another—as when Lepczyk et al. [6] multiply the average number of prey items returned by the average number of outdoor cats per owner. The resulting predation figures may well be four times greater than they should be! (Actually, there are additional problems with the authors’ predation estimates, which I’ll address in a future post).

* * *

The fact that such a fundamental mistake—one a student couldn’t get away with in a basic statistics course—is made so often is shocking. The fact that such errors slip past journal reviewers is inexcusable.

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. 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.

3. 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

4. Baker, P.J., et al., “Impact of predation by domestic cats Felis catus in an urban area.” Mammal Review. 2005. 35(3/4): p. 302-312.

5. 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.

6. 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.

7. Crooks, K.R., et al., “Exploratory Use of Track and Camera Surveys of Mammalian Carnivores in the Peloncillo and Chiricahua Mountains of Southeastern Arizona.” The Southwestern Naturalist. 2009. 53(4): p. 510-517.

8. van Heezik, Y., et al., “Do domestic cats impose an unsustainable harvest on urban bird populations?“ Biological Conservation. 143(1): p. 121-130.

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. Barratt, D.G., “Predation by house cats, Felis catus (L.), in Canberra, Australia. II. Factors affecting the amount of prey caught and estimates of the impact on wildlife.” Wildlife Research. 1998. 25(5): p. 475–487.

11. Barratt, D.G., “Predation by House Cats, Felis catus (L.), in Canberra, Australia. I. Prey Composition and Preference.” Wildlife Research. 1997. 24(3): p. 263–277.

12. May, R.M., “Control of feline delinquency.” Nature. 1988. 332(March): p. 392-393.

13. Jessup, D.A., “The welfare of feral cats and wildlife.” Journal of the American Veterinary Medical Association. 2004. 225(9): p. 1377-1383.

14. 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

15. 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.

16. Winter, L., “Trap-neuter-release programs: the reality and the impacts.” Journal of the American Veterinary Medical Association. 2004. 225(9): p. 1369-1376.

17. Clarke, A.L. and Pacin, T., “Domestic cat “colonies” in natural areas: a growing species threat.” Natural Areas Journal. 2002. 22: p. 154–159.

18. Fitzgerald, B.M. and Turner, D.C., Hunting Behaviour of domestic cats and their impact on prey populations, in The Domestic Cat: The biology of its behaviour, D.C. Turner and P.P.G. Bateson, Editors. 2000, Cambridge University Press: Cambridge, U.K.; New York. p. 151–175.

19. Clancy, E.A., Moore, A.S., and Bertone, E.R., “Evaluation of cat and owner characteristics and their relationships to outdoor access of owned cats.” Journal of the American Veterinary Medical Association. 2003. 222(11): p. 1541-1545.

The Work Speaks—Part 3: Predatory Blending?

31 May 2010

“…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.

The Work Speaks—Part 2: Sample-Minded Research

27 May 2010

“…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 my previous post, I presented examples of researchers “reinterpreting” the work of others to better fit their own arguments. For the next few posts, I’ll focus on some of the major flaws in the feral cat/TNR research itself—beginning with the reliance, by some, on small sample sizes.

Size Does Matter
There are all kinds of reasons for small sample sizes, perhaps the most common being limited resources (e.g., time, funding, etc.). And they are often a fact of life in real-world research, where investigators have less control over conditions than they might in a laboratory environment. Studies employing small sample sizes are not without value; indeed, they often serve as useful pilot studies for future, more comprehensive, work. They do become problematic, though, when broad conclusions are drawn from their results. Below are three (among many!) examples of such studies.

Impressive Estimates
In “Free-Ranging Domestic Cat Predation on Native Vertebrates in Rural and Urban Virginia,” [2] published in 1992, the authors estimated that the state’s 1,048,704 cats were killing between 3,146,112 and 26,217,600 songbirds each year. “This number,” they note, “is certainly inaccurate to some degree, although the estimates are impressive.” [2] Impressive? I suppose. Maybe incredible is more fitting—since the study from which they were derived included exactly five cats, four “urban” and one “rural.”

Mitchell and Beck acknowledged “the limitations of extrapolation to large areas from relatively small data sets such as ours,” suggesting that their work was intended to provoke future “careful and detailed studies that can reveal truer estimates of the impact of this introduced species.” Hawkins [3] and Dauphiné and Cooper [4], however, seem to take them at their word, regardless of any disclaimers.

Many Cats, Multiple Seasons
In a recent study on Catalina Island, the researchers “examined the home-range behavior and movements of sterilized and intact radiocollared feral cats living in the interior” [5] of the island. Although Guttilla and Stapp concede that “sample sizes, especially for males, were relatively low” despite having “tracked many cats across multiple seasons,” they nevertheless come to some rather dramatic conclusions. Among them: “sterilization likely would not reduce the impact of feral cats on native prey.” [5]

So what do the authors mean by many and multiple? Actually, there were just 27 cats in the study (of an estimated 614–732 on the island). “Four cats were tracked during all four seasons, 9 cats were tracked for three consecutive seasons, 4 cats were tracked for 2 consecutive seasons, and the remaining cats were tracked for 1 season.” [5] And these numbers were effectively cut in half, because the researchers were comparing sterilized and non-sterilized cats. At best, this is a pilot study—though it’s already morphed into something more substantial in the mainstream media.

Myth vs. Math
In their 2004 study, “Ecological Impact of Inside/Outside House Cats Around a Suburban Nature Preserve,” Kays and DeWan observed hunting cats, concluding that their kill rate (13%) is “3.3 times greater than the rate estimated from prey brought home.” [6] Not surprisingly, this figure has been used as an instant multiplier (much in the same way William George’s work has been misused) for researchers interested in “correcting” (inflating?) prey numbers. [4, 7-11]

But this ratio, 3.3, hinges on the hunting behaviors of just 24 cats—12 that returned prey home, and another 12 (11 pets and 1 feral) that were observed hunting for a total of 181 hours (anywhere from 4.8–46.5 hours per cat). It’s interesting to note that the cat observed the most (46.5 hours) was only a year old—the youngest of the 12 observed, and likely the most active hunter. This factor alone could have had a significant influence on the outcome of the study.

Also, as several studies have shown [7,8,12,13], the distribution of prey catches tends to be highly skewed (many cats catch few/no prey, while a few catch a lot). In other words, the distribution is not the familiar bell curve at all—making it inappropriate to use a simple average for calculating estimations (a topic I’ll address in detail later). What’s more, with only 12 cats being monitored, how can we be sure their behaviors accurately represent any real distribution at all?

But the key to their calculation is the average time spent outdoors. This, too, tends to be a highly skewed distribution [14, 15], although—curiously—Kays and DeWan’s data suggest otherwise. By way of example, a 2003 survey conducted by Clancy, Moore, and Bertone [15] revealed that nearly half of the cats with outdoor access were outside for two or fewer hours a day. And 29% were outdoors for less than an hour each day. A survey conducted by the American Bird Conservancy revealed similar behavior, reporting that “35% keep their cats indoors all of the time” and “31% keep them indoors mostly with some outside access.” [14]

Kays and DeWan’s average of 8.35 hours/day, then, seems rather out of line with other studies. This, in addition to a number of unknowns (e.g., influence of time of day/night on hunting success, actual time spent hunting by each cat, etc.) raises serious questions about their conclusions.

By way of comparison, using an average of 2.5 hours/day (which is not out of line with the surveys described above) would yield a ratio of 1:1. In other words, no difference between predation rates predicted by actual hunting observation and those predicted by way of prey returned home. Which is not to say that I agree with Kays and DeWan’s underlying methods—we don’t know the possible effects of seasonal variation, for example, or differences in habitat. I’m only pointing out how sensitive this one factor—with its enormous consequences—is to the amount of time cats actually spend outdoors (and, just to introduce one more complication: I’d be very surprised if the amount of outdoor time cats spend hunting is normally distributed; it, too, is probably skewed).

Ironically, while the authors express disappointment that “biologists have rarely sampled both cat and prey populations in such a way that direct effects on prey populations can be shown,” [6] they seem to have had no misgivings about how their work—suffering from its own sampling issues—might be used to misrepresent those same effects.

* * *

Next, I’ll discuss the difference between compensatory and additive predation, and how that affects predictions of feral cat impacts on wildlife.

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. Mitchell, J.C. and Beck, R.A., “Free-Ranging Domestic Cat Predation on Native Vertebrates in Rural and Urban Virginia.” Virginia Journal of Science. 1992. 43(1B): p. 197–207.

3. 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.

4. 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

5. Guttilla, D.A. and Stapp, P., “Effects of sterilization on movements of feral cats at a wildland-urban interface.” Journal of Mammalogy. 2010. 91(2): p. 482-489.

6. 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.

7. Baker, P.J., et al., “Impact of predation by domestic cats Felis catus in an urban area.” Mammal Review. 2005. 35(3/4): p. 302-312.

8. 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.

9. van Heezik, Y., et al., “Do domestic cats impose an unsustainable harvest on urban bird populations?“ Biological Conservation. 143(1): p. 121-130.

10. Nelson, S.H., Evans, A.D., and Bradbury, R.B., “The efficacy of collar-mounted devices in reducing the rate of predation of wildlife by domestic cats.” Applied Animal Behaviour Science. 2005. 94(3-4): p. 273-285.

11. MacLean, M.M., et al., “The usefulness of sensitivity analysis for predicting the effects of cat predation on the population dynamics of their avian prey.” Ibis. 2008. 150(Suppl. 1): p. 100-113.

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

13. 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.

14. ABC, Human Attitudes and Behavior Regarding Cats. 1997, American Bird Conservancy: Washington, DC. http://www.abcbirds.org/abcprograms/policy/cats/materials/attitude.pdf

15. Clancy, E.A., Moore, A.S., and Bertone, E.R., “Evaluation of cat and owner characteristics and their relationships to outdoor access of owned cats.” Journal of the American Veterinary Medical Association. 2003. 222(11): p. 1541-1545.