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

The Work Speaks—Part 7: Leaky Sink

21 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 6 of this series, I critiqued Christopher Lepczyk’s paper Landowners and cat predation across rural-to-urban landscapes, published in 2003. Here, I’m going to examine two studies conducted by Philip J. Baker and various collaborators.

The Studies
In the first study, Baker et al. distributed questionnaires to 3,494 households across a 4.2 km² area of northwest Bristol (UK), and used responses to estimate cat ownership and predation levels (via prey returned home). [2] This work served as a pilot study for the subsequent study.

The second study, conducted August 2005–July 2006, was also conducted in Bristol. Added to the original 4.2 km² site were nine 1 km² sites. The researchers used very similar sampling methods, but, based on results of their pilot study, had somewhat more specific objectives:

To quantify cat density
To quantify the various species of birds killed by cats.
To estimate the impact of cat predation by species and site.
To determine whether the predation observed was compensatory or additive. [3]

Sources and Sinks
Among the authors’ conclusions from the pilot study was that, at least for three of the ten bird species surveyed:

“…it is possible that cat predation was significantly affecting levels of recruitment and creating a dispersal sink for more productive neighboring areas.” [2]

Dispersal sinks or habitat sinks, are patches of low-quality habitat that are unable to sustain a population of a particular species were it not for immigration from higher quality habitat patches—called sources—nearby. So, what Baker et al. are suggesting is that predation by cats may be extensive enough to deplete populations of certain bird species at their study site, such that at least some of the birds observed there were immigrants from nearby habitat.

But the authors also point out that, “despite occurring at very high densities, the summed effects on prey populations appeared unlikely to affect population size for the majority of prey species.” [2] And even for House sparrows, which were among the three species of concern (and, apparently, in decline throughout the UK’s urban areas), Baker et al. note that their “numbers appear to be stable in Bristol as a whole.”

So, is the area a habitat sink or not?

A cursory look at the theory and empirical measurement of source-sink dynamics reveals great complexity. Variations across time and geography must be taken into account—the ebb and flow of local populations might easily be overlooked or misunderstood by applying a short time horizon (i.e., 12 months) and arbitrary boundaries (i.e., those that define the study site). Annual rainfall, for example, can dramatically influence yearly population levels on a local scale. And it’s been shown that source-sink dynamics can occur over distances of 60–80 km. [4] In fact, the determination of sinks and sources in the field can be problematic enough that sources sometimes appear to be sinks and vice-versa. [5]

Given the complex nature of source-sink dynamics, the suggestion by Baker et al. that cat predation may be creating a habitat sink seems rather premature. Such assertions—despite the requisite disclaimers (the authors note only that “it is possible”)—tend to attract attention and gain traction. Longcore et al., for example, cited the pilot study in their 2009 essay, “Critical Assessment of Claims Regarding Management of Feral Cats by Trap-Neuter-Return.” [6]

Of greater interest to me, though, are the assumptions Baker et al. used to estimate the impact of cat predation.

Counting Cats and Counting Birds
In both studies, the authors quantified the impact of cat predation on bird populations by comparing different levels of predation with different bird densities. Their maximum impacts, for example, assumed that all cats were hunters—despite the fact that 51–74% of the cats included in the two studies brought home no prey at all—and that bird productivity was zero (i.e., no young birds survive to adulthood). As the authors admit:

“This was clearly not realistic, as the estimated maximum numbers of birds killed typically exceeded breeding density and productivity combined, such that the prey populations studied would probably have gone extinct rapidly at a local level or acted as a major sink for birds immigrating from neighbouring areas.” [3]

But how realistic are their other estimates?

A detailed examination of a single species at a one site (taken from the second study, for which such information is available) illustrates some flaws. I looked at House sparrows for the 1 km² site designated as ST5277. Here, 18 participants reported that their 22 cats returned a total of 30 prey items, nine of which were birds (two of them “unidentified”). Of the birds returned home, two were House sparrows.

When it comes to estimating impacts, though, Baker et al. use figures of 332–1,245 House sparrows killed by the cats of ST5277. The maximum, we already know, is “not realistic,” but even the minimum seems awfully high. So, where are these birds coming from?

To start with, two adjustments have to be made to the original predation figure. First, the two unidentified birds are “distributed” across the categories of bird species that were identified. Then, we have to account for participant drop-out; not all of the 22 cats were surveyed for the entire year of the study. Now we’re up to an average of 8.7 House sparrows brought home annually by the cats at this site.

But of course there are more than 22 cats at ST5277. Baker et al. estimate that there are 314 of them (although we know very little about the factors that affect their hunting ability and success—for example, their access to the outdoors, age, etc.). We also know that only seven of the 22 cats included in the study brought home prey. In other words, 32% of the cats surveyed were documented hunters. Based on these numbers, then, we can estimate the yearly predation rate of House sparrows at ST5277 to be roughly 125—well short of the minimum proposed by Baker et al. (and just a quarter of their intermediate rate).

There are some minor differences between their method for estimating predation rates and mine. For the most part, though, the “missing” sparrows can be found in the authors’ use of a correction factor (3.3) proposed by Kays and DeWan to account for prey killed but not returned home. [7] Undoubtedly, cats fail to bring home all the prey they catch (though they also undoubtedly bring home prey they didn’t kill), but there is good reason to doubt Kays and DeWan’s “correction.” Among the flaws in their analysis were small, dissimilar samples of cats, and a failure to account for highly skewed data sets.

So, even setting aside the complexities of source-sink dynamics, these inflated predation rates, combined with the fact that “the estimates of breeding density presented in this manuscript should be regarded as minima,” [3] raise serious doubts about whether the site is in fact a habitat sink (or, if so, to what extent).

Compensatory and Additive Predation
As I’ve discussed previously, even accurately predicted levels of predation can be deceptive. There’s 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.” [8]

When it comes to relating predation to population levels, it’s critical to understand the difference, and know the extent to which each type is occurring.

To get at this critical issue, Baker et al. compared the physical attributes (e.g., muscle mass score, mean fat score, etc.) of 86 birds killed by collisions (e.g., with cars, windows, etc.) to those of 48 birds killed by cats. Although the authors point out, “the relationship between body mass and quality (i.e., likelihood of long-term survival and therefore reproductive potential) in passerines is complex,” they nevertheless conclude that the birds killed by cats “were likely to have had poor long-term survival prospects.” [3] (An earlier study comparing spleen mass arrived at essentially the same conclusion: that birds killed by cats “often have a poor health status.” [9])

Still, Baker et al. express caution 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.”

But their inflated predation rates and low estimates of breeding density combine 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.

* * *

It’s not clear why Longcore et al. cited the pilot study their essay, but left out any mention of the much larger subsequent study. Perhaps it was just a matter of timing—“Cats About Town” was published in August of 2008, while “Critical Assessment” was published in August of 2009. A year is not much time in the world of scientific journals, and it’s possible that the two manuscripts more or less crossed in the mail. On the other hand, the pilot study fits more neatly into the argument put forward by Longcore et al.—an argument that doesn’t even recognize the distinction between compensatory and additive predation.

Of course, Baker et al. did themselves no favors, either. By using inflated predation rates—the result of some peculiar, unjustified assumptions—they virtually buried the most important findings of their study.

References
1. 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.

2. 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.

3. 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.

4. Tittler, R., Fahrig, L., and Villard, M.-A., “Evidence of Large-Scale Source-Sink Dynamics and Long-Distance Dispersal among Wood Thrush Populations.” Ecology. 2006. 87(12): p. 3029-3036.

5. Runge, J.P., Runge, M.C., and Nichols, J.D., “The Role of Local Populations within a Landscape Context: Defining and Classifying Sources and Sinks.” The American Naturalist. 2006. 167(6): p. 925-938.

6. 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.

7. 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.

8. 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.

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

The Work Speaks—Part 4: Mean Spirited

4 Jun 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 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.

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