The Work Speaks—Part 7: Leaky Sink

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:

  1. To quantify cat density
  2. To quantify the various species of birds killed by cats.
  3. To estimate the impact of cat predation by species and site.
  4. 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.

Learning Your ABC

According to an AP story posted on MSNBC, city officials in Barre, VT, are considering a leash law for cats—effectively prohibiting them from roaming. Such stories tend to vary only in their specifics; quotes from embattled citizens, and feline references that are more dismissive than clever (e.g., in Barre, the debate “sparked a hissing match”) are pretty much a given. And, more often than not, there’s a statement from the American Bird Conservancy (ABC) about the threat free-roaming cats pose to wildlife. This story was no exception:

“Scientists estimate that free-roaming cats kill hundreds of millions of birds, small mammals, reptiles and amphibians each year,” the Virginia-based American Bird Conservancy, which runs a “Cats Indoors!” campaign, says on its website. “Cat predation is an added stress to wildlife populations already struggling to survive habitat loss, pollution, pesticides, and other human impacts.”

Although I don’t know anybody who would argue with ABC’s second point, that first one bears closer inspection.

I discussed a similar claim by ABC’s Senior Policy Advisor, Steve Holmer, in one of my first Vox Felina posts. In January, Holmer told the Los Angeles Times:

“The latest estimates are that there are about . . . 160 million feral cats [nationwide] . . .  It’s conservatively estimated that they kill about 500 million birds a year.”

Late last year, an article in Audubon Magazine had published nearly identical figures, citing ABC as its source. [1]

The feral cat estimate comes from a conference paper written by Nico Dauphiné and Robert J. Cooper, available for download via the ABC website. When I pressed Holmer about the authors’ “creative accounting,” he backed off, assuring me that ABC’s materials “should now say”:

There are currently 88 million pet cats in the U.S. according to a pet trade association, and that number is growing. In addition, it is estimated that there may be 60–100 million free-ranging feral cats in the U.S., and that these cats may collectively kill more than one million birds each day. Reducing this mortality even a small amount could potentially save millions of birds each year.

I never received a reply, though, to my inquiries about that “more than one million birds each day” claim. Such incidents are, unfortunately, not uncommon; when it comes to assertions about cat predation and its impact on wildlife, ABC has a rich—and rather shameful—history.

Cats Indoors!
Holmer’s comment to the L.A. Times is just one example of ABC’s concerted effort to use the (largely unquestioning) media in getting their message out. Last year, at a news conference about the “The U.S. State of the Birds” report, ABC’s Darin Schroeder told the press, “education is urgently needed to make the public aware of the toll of pet cats.” Which is precisely what ABC’s Cats Indoors! campaign—launched in 1997—aims to do.

The question is, what kind of education is the public getting from ABC?

  • A 1997 report by ABC claimed, “extensive studies of the feeding habits of domestic, free-roaming cats… show that approximately… 20 to 30 percent [of their diet] are birds.” [cited in 2] In fact, as Ellen Perry Berkeley points out in her book, TNR Past Present and Future: A history of the trap-neuter-return movement, the 20–30% figure was not based on “extensive studies” at all. [2]ABC’s Linda Winter, writing to Berkeley, cited just three sources. Two of them—the now-classic “English village” study by Peter Churcher and John Lawton, and the “Wisconsin Study” by John Coleman and Stanley Temple—have been widely discredited. [3–5] And the third, Mike Fitzgerald’s contribution to “Diet of domestic cats and their impact on prey populations,” [6] was misinterpreted and/or misrepresented by ABC. (As Berkeley notes, Fitzgerald’s data “would put birds, as a portion of the diet of cats, at roughly 7 to 10.5 percent—nowhere near the ‘20 to 30 percent’ figures unleashed on the unscientific public by ABC!” [2])
  • Winter, director of Cats Indoors! (assuming she’s still at ABC; their website does not list her among the staff), and ABC president George Fenwick were among those thanked “for helpful and constructive reviews” in the Acknowledgements section of Christopher Lepczyk’s 2003 paper, “Landowners and cat predation across rural-to-urban landscapes.” As I detailed in my previous post, Lepczyk’s study is flawed both in terms of its method and analysis, and his predation estimates are highly inflated as a result. The fact that Winter and Fenwick were involved in such as study—at any level—raises questions about ABC’s credibility (and its possible influence on research outcomes).
  • In 2004, Winter misrepresented the results of a survey commissioned by ABC. In “Trap-neuter-release programs: The reality and the impacts,” published in the Journal of the American Veterinary Medical Association, she suggested, “66% of cat owners let their cats outdoors some or all of the time.” [7]In fact, the survey indicated that “35% keep their cats indoors all of the time” and “31% keep them indoors mostly with some outside access.” [9] While Winter’s claim isn’t exactly untrue, it certainly paints a very different picture: rather than one-third, two-thirds of cats are free-roaming. Which, apparently, is exactly how Dauphiné and Cooper read it, combining this with an inflated figure for the number of feral cats to come up with their estimate of “117–157 million free-ranging cats in the United States.” [8] (It’s difficult not to see a certain coziness here: Dauphiné and Cooper citing Winter’s “interpretation” of her own survey results, and Holmer’s reliance on Dauphiné and Cooper’s conference paper.
  • To this day, ABC refers to the highly-criticized Wisconsin Study in its brochure Domestic Cat Predation on Birds and Other Wildlife: “Researchers… estimated that rural free-roaming cats kill at least 7.8 million and perhaps as many as 217 million birds a year in Wisconsin. Suburban and urban cats add to that toll.” [9]And, despite Berkeley’s efforts to untangle their erroneous dietary figures, ABC has backed off only slightly: “In an ongoing, but unpublished, study of cat prey items including stomach contents, scat analysis, observations of kills, and prey remains, birds were 19.6% of 1,976 prey captured by 78 outdoor cats (Temple, S.A, Univ. of WI, personal communication, 1/22/04).”

    [Note: Download Laurie D. Goldstein’s Addressing the Wisconsin Study for a comprehensive critique of this work.]

*     *     *

Don’t get me wrong; I’m all for keeping cats indoors. But what about the feral and stray cats out there—what happens to them? Here, ABC doesn’t seem to have a lot of answers. At least not any they’re willing to be up-front about.

In fact, by disseminating information that is at best misleading—and often, just plain wrong—ABC is doing whatever it can to shape policy in such a way that many of these cats will, one way or another, be killed. Intentional or not, Cats Indoors! has become a kind of Trojan horse for those determined to eliminate all free-roaming cats. Attention can very quickly shift from the impact of a proposed leash law, for example, to the “cat problem” in general.

Although it’s packaged as sound advice for cat owners, the Cats Indoors! campaign has probably had a far greater (deadly) impact on unowned cats than on pet cats.

References
1. Williams, T., Felines Fatale, in Audubon Magazine. 2009, National Audubon Society: New York, NY. http://www.audubonmagazine.org/incite/incite0909.html

2. Berkeley, E.P., TNR Past present and future: A history of the trap-neuter-return movement. 2004, Bethesda, MD: Alley Cat Allies.

3. Goldstein, L.D., O’Keefe, C.L., and Bickel, H.L. Addressing “The Wisconsin Study”. 2003.  http://www.straypetadvocacy.org/html/wisconsin_study.html.

4. Clifton, M. Where cats belong—and where they don’t. Animal People 2003. http://www.animalpeoplenews.org/03/6/wherecatsBelong6.03.html.

5. Patronek, G.J., “Free-roaming and feral cats—their impact on wildlife and human beings.” Journal of the American Veterinary Medical Association. 1998. 212(2): p. 218–226.

6. 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. p. 123–147.

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

8. 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. 2009. p. 205–219.

9. ABC, Domestic Cat Predation on Birds and Other Wildlife. n.d., American Bird Conservancy: The Plains, VA. www.abcbirds.org/abcprograms/policy/cats/materials/predation.pdf

The Work Speaks—Part 6: Pain by Numbers

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 5 of this series, I critiqued Cole Hawkins’ 1998 PhD dissertation. Here, I’m going to untangle some of Lepczyk’s own PhD work: Landowners and cat predation across rural-to-urban landscapes, published in 2003.

The Study
In this study, surveys were distributed across three southeastern Michigan landscapes (rural, suburban, and urban) corresponding to established breeding bird survey (BBS) routes. [2] Among the survey questions:

“If you or members of your household own cats that are allowed access to the outside, approximately how many dead or injured birds a week do all the cats bring in during the spring and summer months (April through August) (0, 1, 2–3, 4–5, 6–7, 8–9, 10–15, 16–20, more than 20)?”

Based on 968 surveys returned from 1654 private landowners (a decent response rate of 58.5%), Lepczyk et al. conclude:

“Across the three landscapes there were ~800 to ~3100 cats, which kill between ~16,000 and ~47,000 birds during the breeding season, resulting in a minimum of ~1 bird killed/km/day.”

Increasing Uncertainty
How do Lepczyk and his collaborators arrive at these figures? It’s not entirely clear, actually. Despite numerous attempts, I’ve been unable to follow all of their calculations. However, using their data, I developed my own estimate: 1,119 outdoor cats, 511 of which were reported to be successful hunters.

Using this figure, I then summed across all three landscapes the birds killed or injured, plus those killed or injured by non-respondents’ hunting cats (based on the ratio of hunters to outdoor cats owned by respondents, or about 50%). The resulting estimate is 15,856 birds killed over the 22-week breeding season—close to the low estimate suggested by Lepczyk et al., but just a third of their maximum.

So, why the discrepancy?

One reason is that, at least for some of their estimates, Lepczyk et al. assumed that every landowner who didn’t respond to the survey owned outdoor cats. This, despite their survey results, which indicated that only about one-third of landowners fell into this category.

But the authors go further, generating predation estimates based on pure speculation, specifically that “non-respondents have 150% the number of outdoor cats as respondents.” [2] It should be noted that Lepczyk et al. also ran another scenario in which non-respondents had half the outdoor cats as did respondents—but, again, in both cases they assume that every non-respondent owned outdoor cats.

As a result of this approach, the authors end up in some strange territory: the estimated number of cats owned by non-respondents (based on the assumptions described above) far exceeds the number owned by respondents—by more than a two-to-one margin, in some cases. If the greatest impacts are going to be attributable to non-respondents, then what’s the point of doing the survey in the first place? There are accepted methods by which one can manage uncertainty—statistical analysis, confidence intervals, and the like. What Lepczyk et al. have done serves just one purpose: to inflate apparent predation rates.

Skewed Distributions
In addition to the flaws described above, there are some fundamental errors in the way the authors handle their data. Like so many others, Lepczyk et al. ignore the fact that their data is not normally distributed:

  1. Lepczyk et al. use the average number of birds killed/cat to calculate the total number of bids killed for each of the three landscapes. As I discussed previously), this is a highly positively skewed distribution—using a simple average, therefore, greatly overestimates the cats’ impact (by as much as a factor of two).
  2. A similar error is made when the authors use an average to describe the number of outdoor cats owned by each landowner. Again, because this is a skewed distribution, their use of a simple average exaggerates the extent of predation.
  3. The two inflated figures described in (1) and (2) are multiplied together, further inflating estimated predation rates.

Barratt has suggested that “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.” [3] Accounting for the first point alone, then, my estimate is reduced to 8,000 birds killed over the 22-week breeding season.

Accounting for the second point is somewhat trickier. For one thing, we don’t know what constitutes an outdoor cat here—the survey simply asked respondents if they owned cats “that are allowed access to the outdoors.” [2] However, we do know the results of a 2003 survey, which indicated 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. [4] Although these figures almost certainly reflect owners in urban and suburban landscapes more than those in rural landscapes, it’s clear that a simple yes-or-no question on the subject is insufficient. Indeed, such a question will invariably overestimate the number of “outdoor cats”—which in turn overestimates predation rates.

This, coupled with the error inherent in using a simple average, pushes predation estimates lower. And the third point reduces those estimates further still. Taken together, these corrections could put my estimate closer to 4,000 birds. More important, the upper estimate proposed by Lepczyk et al.—47,000 birds—could easily be 10 times too high.

The Small Print
Despite their inflated figures, Lepczyk et al. suggest—rather absurdly, in light of the substantial flaws described above—that perhaps their estimates are actually too conservative:

“One caveat to our study is that landowners may have underestimated the number of cats they allow access to the outside. Such a result was found in a similar study of landowners in Wisconsin (Coleman and Temple, 1993).” [1] (Note: After reviewing “Rural Residents’ Free-Ranging Domestic Cats: A Survey,” [5] I’ve found no evidence of such a result.)

“… we found that a very common volunteered response among landowners that had no outdoor cats was that either their neighbors owned outdoor cats or that feral cats were present in the vicinity of their land… [suggesting] that at least some landowners under reported or chose not to report the number of outdoor cats they owned.”

But what about their reports of birds brought home killed or injured—how trustworthy were those? After all, the survey (mailed during the first week of October) asked respondents to recall the number of birds their cat(s) brought home April through August. Surely, there was a lot of guesswork involved. In fact, David Barratt found this kind of guesswork to overestimate predation rates. In a study published five years prior to “Landowners and Cat Predation,” Barratt concluded, “predicted rates of predation greater than about ten prey per year generally over-estimated predation observed.” [3]

The two studies cannot be compared directly for a number of reasons, but by way of comparison, the average predation rate used by Lepczyk et al. is approximately 31 birds/cat for the 22-week breeding season. Using Barratt’s work, in which the “heaviest” six continuous months correspond to about 58% of yearly prey totals, [6] I converted this to a yearly rate of 53 birds/cat/year. Barratt has shown that the actual predation rate, at this level, is less than half the rate predicted by cat owners. In other words, predictions of 50 birds/year generally correspond to catches closer to 25 birds/year.

While Lepczyk et al. emphasize the potential for under-estimating predation levels, they never consider the risk of over-estimating these levels—or their most obvious potential source of error: landowners’ recollections of birds killed. The authors question respondents’ reports of outdoor cats, but accept without question their reports of birds injured or killed over the previous six-month period. And, as Barratt indicated, such reports can be inflated by a factor of two or more!

Something else I find troubling comes, of all places, from the Acknowledgements section. Among those thanked “for helpful and constructive reviews” are American Bird Conservancy (ABC) president George Fenwick and Linda Winter, director of ABC’s Cats Indoors! campaign. It’s not clear how Fenwick and Winter contributed to the final paper, but their involvement on any level raises questions about possible bias. Certainly, Winter has credibility issues when it comes to “research” about the impact of free-roaming cats on birds, as I’ve already described (see also pp. 18–24 of TNR Past present and future: A history of the trap-neuter-return movement [7]).

*     *     *

The same year Lepczyk’s paper was published, the American Veterinary Medicine Association held an Animal Welfare Forum “devoted to the management of abandoned and feral cats.” [8] In attendance were more than 200 veterinarians, animal control officials, wildlife conservationists, and animal advocates—each with a different perspective on feral cats in general and TNR in particular.

In welcoming this diverse group, then-President-Elect Bonnie Beaver recognized the range of contentious issues before them:

“Feral cats evoke hot debates about ecological issues, individual cat welfare, human responsibilities, intercat disease transmission, humaneness, zoonosis control, and management and dissolution of unowned cats.” [8]

Amidst the “hot debate,” though, Beaver was optimistic:

“We will not always agree, but we will come away with increased knowledge and a renewed commitment to work for the welfare of all the animals with which we share the earth” [8]

While I tend to share Beaver’s optimism, I think the debate is hurt—if not derailed entirely—by the publication of research aimed not at increasing our collective knowledge, but rather at supporting a particular position. Like Cole Hawkins’ dissertation, “Landowners and Cat Predation” is, at best, an interesting pilot study for subsequent work. And yet, it’s widely—and uncritically—cited in the feral cat/TNR literature. Longcore et al., for example, refer to it as “evidence [indicating] that cats can play an important role in fluctuations of bird populations,” [9] despite the fact that Lepczyk et al. don’t actually address the issue of bird populations at all. More recently, Dauphiné and Cooper use the inflated predation rate suggested by Lepczyk et al. (along with rates proposed by other researchers) to arrive at their “billion birds” figure. [10]

The method employed in “Landowners and Cat Predation”—asking owners of cats to recall the number and species of birds over the previous six-month period—invites overestimation from the very outset. Lepczyk et al. then inflate these numbers through both careless (e.g., using averages to describe skewed data) and deliberate (e.g., assuming all non-respondents owned cats—perhaps 50% more than respondents did) means. Rather than getting us any closer to the truth about cat predation, this study only obscured it further.

Worse, it’s been packaged and sold—and subsequently “bought”—as rigorous science, thereby giving it an undeserved legitimacy. Such efforts are impediments to knowledge and understanding—and therefore, to progress.

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. 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.

3. 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.

4. 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.

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. 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.

7. Berkeley, E.P., TNR Past present and future: A history of the trap-neuter-return movement. 2004, Bethesda, MD: Alley Cat Allies.

8. Kuehn, B.M. and Kahler, S.C. The Cat Debate. JAVMA Online 2004 November 27, 2009 [accessed 2009 December 24].  http://www.avma.org/onlnews/javma/jan04/040115a.asp.

9. 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.

10. 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.

The Work Speaks—Part 5: Jumping to Conclusions

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 4 of this series, I discussed 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.

For the next few posts, I’m going to critique three of the studies most often cited by these researchers, starting with Cole Hawkins’ 1998 PhD dissertation, Impact of a subsidized exotic predator on native biota: Effect of house cats (Felis catus) on California birds and rodents. I mentioned Hawkins’ dissertation previously, but only briefly. Here, I’ll take a closer look, paying particular attention to how he gets from his results to his rather dubious conclusions.

The Study
Hawkins’ two-year study was conducted in Alameda County, CA, spread across two adjacent parks. He started by designating a “cat area” (where, nearby, free-roaming cats were being fed) and a “no-cat area” (where no cats were being fed), and then designated “rodent grids” (nine locations used for trapping and counting rodents) and walking transects (from which bird surveys were conducted) in each area. Hawkins then compared the number of birds and rodents detected in the two areas.

Among Hawkins’ conclusions:

“The differences observed in this study were the results of the cat’s predatory behavior.” [2] (It should be noted that Hawkins tempered this assertion in his 1999 article summarizing the work: “The differences observed in this study may have been due to the cats’ predatory behavior.” [3] (italics mine))

And this:

“The presence of cats in this study area already has caused a shift in the composition of the rodent community; it is possible that a shift in the larger biotic community could follow.”

And, finally:

“It is not prudent to manage for wildlife and allow cat feeding in the same parks.”

Unfair Comparisons
But Hawkins’ findings are insufficient to supports such claims; indeed, his methodology doesn’t allow for them. Hawkins has no idea what the cat area was like prior to his arrival; he merely assumes the populations of birds and rodents would have been identical to those found at the no-cat area, and makes his comparisons accordingly. In fact, there are a number of factors that indicate that the two areas are not as comparable as Hawkins suggests:

  • The cat area was almost a peninsula, with a lake on one side and a residential area (within 0.5 km) on the other. The no-cat area, on the other hand, was located largely in the interior of the parks.
  • Hawkins notes that there were more people in the cat area (of­ten twice as many as were observed at the no-cat area), but dismisses the possibility that their presence may have influenced the numbers of birds and rodents he observed there.
  • The habitat along the 2.2 km transects from which bird counts were conducted varied considerably between the two areas. Compared to the no-cat area, the cat area had 31% less chaparral, 183% more trees, 52% less grass, and 240% more “modified” habitat (it’s not clear what Hawkins means by “modified,” but I assume it refers to habitat that reflects significant human impact).
  • Finally, the presence of pesticides may have played a role. According to a 2002 report (the earliest I was able to find) from the East Bay Regional Park District, “The focus of Lake Chabot’s weed control efforts are vegetation reduction within the two-acre overflow parking lot, picnic sites and firebreaks around park buildings, corp. yard, service yard, and the Lake Chabot classroom.” [4] And it’s clear from Hawkins’ 1999 article that the cat area did include picnic sites: “…over half of the cat scat in this study was collected under and around picnic tables.” [3] Now, Hawkins’ fieldwork was done in 1995 and 1996, but if there was any pesticide use during the study period, it may have affected the results—especially if the pesti­cide was distributed differently across the two areas.

Cats and Birds
“Almost twice as many birds were seen on the no-cat transect as on the cat transect,” writes Hawkins. But it’s not quite as simple as that—the details reveal a rather complex, often uneven count over the course of the study. Nevertheless—and despite the differences between the two areas—Hawkins’ only explanation is the cats. This is especially true for ground-feeders:

The preference of ground feeding birds for the no-cat treatment was striking; for ex­ample, California quail were seen almost daily in the no-cat area, whereas they were never seen in the cat area.

What’s more striking to me is the fact that five of the nine ground-feeding species included in the study showed no preference for either area. But Hawkins scarcely acknowledges the point, and doesn’t even hint at an explanation. “Birds that were known to nest on or near the ground or in shrubs and vines ≤ 1.5 m in height” also showed no preference between the two areas (though no nest counts were conducted).

The picture painted by Hawkins is that bird species absent from the cat area represent species killed off by the cats. But it’s generally accepted that cats are opportunistic hunters, catching whatever prey is readily available and easily caught. [5–8] Fitzgerald and Turner, for ex­ample, argue that “domestic cats (both house and feral ones) are best described as generalist resident preda­tors, exploiting a wide range of prey, and able to switch readily from one prey to another.” [9] So, how is it that some species were present at the cat site while others were not? Again, Hawkins offers no explanation.

In fact, it’s clear from Hawkins’ study that the cats aren’t much of threat at all to the birds—even vulnerable ground-feeding and ground-nesting species—in the cat area. Of the 120 scat samples found by searching the cat area, “65% were found to contain rodent hair and 4% feathers.” [2] This finding comes toward the end of the study, when the cat population was at its greatest—and still, only 4% contained feathers. And this could easily represent one cat and one bird.

One final point about the birds: Hawkins suggests (without explanation) that the olive-sided flycatcher, American robin, and Stellar’s jay—all of which showed no preference for either the no-cat area or the cat area—may have been responding to a “specialized habitat.” Could it be that the birds not seen in the cat area were also responding to a specialized habitat—by “migrating” to a place with less human activity (e.g., the no-cat area), for example? Once again, Hawkins has no comment.

Cats and Rodents
The fact that scats indicated rodents were predated to a greater extent than birds is hardly surprising [5, 6, 9, 10], but it should be noted the 65% figure represents the frequency of occurrence, and not a predation rate (a topic I address in greater detail here).

Hawkins’ analysis didn’t reveal whether the rodent hair was that of deer mice, harvest mice (both of which were found less often in the cat area than in the no-cat area), house mice (found more often in the cat area), or California voles (which showed no preference for either area). In any case, it’s not clear that the cats were responsible for the presence or absence of any of these rodent species. Again, the selective dietary habits suggested by Hawkins simply don’t fit with the domestic cat’s profile as a “generalist resident predator.”

Two additional points that might explain the differences Hawkins observed concern the habitat of the cat area. First, there’s the nearby lake and residences—potential sources of pollution that could affect nearby plant and animal life. Secondly, there’s the issue of possible pesticide use mentioned previously. As I say, it’s largely conjecture on my part; at the same time, though, it’s easy to imagine its potential impact on small mammals (and ground-feeding birds, for that matter).

Finally, Hawkins suggests that certain bird species were responding to specialized habitat—perhaps the rodents were simply doing the same.

Cats
Hawkins used track plates (baited devices that detect the presence of mammals by way of preserved “footprints”) for “estimating a relative cat presence index,” but found only one cat track in 200 track plate nights. And, “in 560 days of exposure, no scat was found in any of the sand boxes.” [2] Now, the cats were seen at feeding stations and on the rodent grids of the cat area—as many as 26 during a one-week period toward the end of the study. But clearly, they were not where Hawkins was expecting them to be. If, after two years at the study site, Hawkins was unable to get a better handle on the presence of the cats, how can he be so sure of their behavior when it comes to predation?

If, as Hawkins argues, the differences observed between the two areas are a result of the cats’ predatory behavior, then one would expect the number of birds and rodents to decrease as the number of cats increases. Yet, Hawkins’ findings don’t bear this out.

And then there are the unanswered questions about the cats—for example:

  • Where did these cats come from—were they illegally dumped, the result of nearby residents’ unsterilized cats breeding? Did they belong to the residents?
  • Were the cats sterilized? (Their increasing numbers would suggest that they weren’t.)
  • Were these cats part of a managed TNR colony? (Local newspaper reports indicate a long-standing battle between TNR advocates and opponents. [11–13])

Considering the central role these cats played in Hawkins’ two-year study, he knew surprisingly little about their behavior—including various factors that surely had an impact on his findings.

*     *     *

In their recent comment, Lepczyk et al. suggest that conservation biologists and wildlife ecologists “look to the evolutionary biology community” [1] for an example of how to influence policy. For feral cat/TNR opponents interested in shaping policy, it seems Hawkins’ study has become quite popular. [14–17] Actually, Nico Dauphiné and Robert J. Cooper take its already-tenuous claims one step further, citing Hawkins’ work (actually a 2004 conference paper that summarizes his dissertation [18]) as evidence that “the continuous predation pressure exerted by exotic predators in exponentially high densities can and has resulted in numerous local extinctions of continental land birds.” [8]

But is Hawkins’ methodology one that evolutionary biologists would advocate—or even recognize? Not likely.

Hawkins draws conclusions—infers important causal relationships—without any evidence of what “pre-treatment” conditions were like. And ignores entirely his own findings when they contradict his conclusions. Rather than beginning his inquiry with questions to answer, it seems Hawkins had his answer from the outset. At best, his work is an interesting pilot study—generating research questions for a more rigorous, less biased investigation.

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. 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.

3. 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.

4. Brownfield, N.T., 2002 Annual Analysis of Pesticide Use East Bay Regional Park District. 2003, East Bay Regional Park District. www.ebparks.org/files/stew_pest_report_02.pdf

5. 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.

6. 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.

7. 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.

8. 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.

9. 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.

10. 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.

11. Chui, G., Stray Cats Live Harsh Lives in Area Parks, in San Jose Mercury News. 1985. p. 1

12. Bogue, G., Those Poor Cats Need a Human Assist, in Contra Costa Times. 1997: Walnut Creek, CA. p. A02

13. n.a., Spring controversy: What to do with feral cats?, in San Mateo Daily Journal, The (CA). 2001.

14. 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.

15. ABC, Domestic Cat Predation on Birds and Other Wildlife. n.d., American Bird Conservancy: The Plains, VA. www.abcbirds.org/abcprograms/policy/cats/materials/predation.pdf

16. Winter, L. and Wallace, G.E., Impacts of Feral and Free-Ranging Cats on Bird Species of Conservation Concern, G.E. Wallace, Editor. 2006, American Bird Conservancy. www.abcbirds.org/newsandreports/NFWF.pdf

17. Ash, S.J. and Adams, C.E., “Public Preferences for Free-Ranging Domestic Cat (Felis catus) Management Options.” Wildlife Society Bulletin. 2003. 31(2): p. 334–339.

18. Hawkins, C.C., Grant, W.E., and Longnecker, M.T. Effect of house cats, being fed in parks, on California birds and rodents. in Proceedings Of The 4th International Symposium On Urban Wildlife Conservation. 2004. Tucson, AZ: University of Arizona. http://cals.arizona.edu/pubs/adjunct/snr0704/snr07042l.pdf

The Work Speaks—Part 4: Mean Spirited

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.

Available: Resources

As some of you may have noticed, I’ve added a new tab to the Vox Felina site. Resources is where you’ll find printer-friendly documents adapted from Vox Felina content.

The first, “Reassessment: A Closer Look at ‘Critical Assessment of Claims Regarding Management of Feral Cats by Trap-Neuter-Return,’” is a brief review 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. It’s based on a four-part series of posts from May. Download PDF