O (Environment) Canada!

In “Estimated Number of Birds Killed by House Cats (Felis catus) in Canada,” published late last year in Avian Conservation and Ecology, Environment Canada research scientist Peter Blancher estimates that cats—owned and unowned—“kill between 100 and 350 million birds per year in Canada,” and suggests that this level of predation “is probably the largest human-related source of bird mortality in Canada.” [1]

I submitted the following comments to the journal in response to Blancher’s article, but retracted my submission upon learning that (1) the length is nearly twice as long as what is permitted, and (2) that I would be required to pay an “author fee” of about $340.

I’m not naïve enough to think that posting my comments here is comparable to having them published in ACE, but, given the considerable work involved—and, more important, the obvious policy implications of Blancher’s paper—I think it’s important that they be available. Read more

Red Herrings, White Lies, and Blue Birds

A pair of Eastern Bluebirds in Michigan, USA. Photo courtesy Wikimedia Commons and Sandysphotos2009

As I sift through my growing collection of studies, news stories, press releases, and anything else relevant to the free-roaming cats/TNR debate, it’s not unusual for me to be diverted by a seemingly minor item—a claim, interpretation, or reference that simply doesn’t sit right with me. (The subsequent investigation of which helps explains the almost alarming rate at which the collection continues to grow.) Sometimes these diversions snowball, taking on a momentum all their own and, ultimately, evolve into their own blog post(s). Others remain largely in the background—overshadowed by more pressing issues—but are too compelling to be ignored for long.

This was the case with some comments Lepczyk et al. made about “two species of conservation concern” in their 2003 paper “Landowners and Cat Predation Across Rural-to-Urban Landscapes,” [1] which I discussed some time ago. The study, in which surveys were distributed across three southeastern Michigan landscapes (rural, suburban, and urban) corresponding to established breeding bird survey (BBS) routes, asked respondents to recall the number of birds their cat(s) brought home April through August 2000. Among the authors’ conclusions:

“The fact that both Eastern Bluebirds and Ruby-throated Hummingbirds were listed [among those killed by cats] indicates that some species of concern are being captured.” [1]

Eastern Bluebirds
Of the 137 birds (representing an estimated 23 species) identified by landowners (approximately one-third were not identified), six (4.4%) were Eastern Bluebirds. What piqued my curiosity was not the tally itself, but the authors’ subsequent comment that “the location of the three landscapes represents an area of Michigan where the species is rarest and not always identified on bird atlas survey routes.” [1]

Wait a minute. The people conducting bird counts along these routes rarely, if ever, locate Eastern Bluebirds—but the cats that live nearby managed to find at least six of them over the course of just five months? Where Lepczyk et al. see reason for concern, I see reason for (cautious) optimism: There are, as the cats demonstrated, more Eastern Bluebirds than we thought!

Just to be clear: I don’t mean to dismiss conservation concerns for uncommon or rare species. Nor am I criticizing the efforts of the many dedicated professionals and volunteers responsible for bird counts. But in trying to reconcile these seemingly contradictory findings, one can’t help but wonder: How many birds are there, really?

To find out, I began by consulting The Atlas of Breeding Birds of Michigan, the very source Lepczyk et al. cite, where, sure enough, a map indicates that no more than two Eastern Bluebirds are typically found along survey routes in the southeastern part of the state. Indeed, even in the most abundant parts of Michigan, no more than four of this species are reported. [2] But this atlas was published in 1991, more than 10 years prior to Lepczyk’s dissertation research.

To look at more recent data—and long-term trends—I referred to the website for the North American Breeding Bird Survey, where the survey is explained this way:

“The BBS is a large-scale survey of North American birds. It is a roadside survey, primarily covering the continental United States and southern Canada, although survey routes have recently been initiated in Alaska and northern Mexico. The BBS was started in 1966, and the over 3,500 routes are surveyed in June by experienced birders.”

The BBS site allows visitors to investigate trends (spanning roughly 40 years) by species, region, and survey route. What I found for Eastern Bluebirds along the three routes employed by Lepczyk et al. (49053, 49167, and 49168) was rather surprising. In all three cases, the abundance of Eastern Bluebirds trends upward—in some cases dramatically.

Eastern Bluebird BBS Data
BBS Data: Eastern Bluebirds for Three Michigan Routes (adapted from North American Breeding Bird Survey website)

Route 49053 is of particular interest for a couple reasons. First of all, it is the only one of the three for which data going back to the BBS’s inception is available, allowing the best long-term perspective. Secondly, the increase in Eastern Bluebird abundance corresponds almost perfectly to publication of The Atlas of Breeding Birds of Michigan. Between 1991 and 2000 (the year Lepczyk was conducting his research), the picture changed considerably.

The other two routes—to a lesser degree, certainly—also exhibit notable increases. Why Lepczyk referred to the Atlas rather than to this more recent data is unclear.

Bird Counts
It’s important to recognize that bird counts are not intended to quantify, in any absolute sense, the number of birds in a particular area, a point made clear on the BBS website:

“The survey produces an index of relative abundance rather than a complete count of breeding bird populations.”

In addition, the use of roadside surveys has been criticized for its potential biases. Roadside habitats may not reflect—and/or may change at rates different from—an area’s overall habitat, for example. [3] Also, a number of factors affect an observer’s ability to detect or identify a particular species. Some—sight, hearing, and training, and even clothing color, for example—are associated with the surveyors, while others (e.g., plumage, body size, coloration, and density) have to do with the birds being surveyed. [4] Rosenstock et al. suggest that such impediments raise serious questions about index counts in general:

“Measures of relative abundance derived from index counts… represent an uncertain, confounded combination of detectability and density. Given these weaknesses, index counts should not be expected to provide reliable information or a valid basis for inference.” [4]

All of which makes weighing the six Eastern Bluebirds in Lepczyk’s study against those detected along the survey routes a dodgy proposition. (Dodgier still is the more expansive claim made by Longcore et al. that Lepczyk’s work is “evidence indicat[ing] that cats can play an important role in fluctuations of bird populations.” [5])

House Sparrows
To reiterate, I’m not discounting conservation concerns for rare or protected species (the Eastern Bluebird, it should be noted—and Lepczyk et al. acknowledge—is neither “extremely rare” nor a “species of state or national concern.” [1]) I’m merely pointing out some of the complexities involved in trying to connect predation levels of one species to population levels of another.

Which brings me back to The Atlas of Breeding Birds of Michigan, where I discovered an interesting twist in this story. The Eastern Bluebird, it seems, probably peaked in Michigan during the late 1800s.

“A gradual decline occurred in the early 1900s as the House Sparrow advanced, and favorite nesting places such as wooden fenceposts and old apple orchards were eliminated.” [2]

As it turns out, House Sparrows were at the top of the list in Lepczyk’s study, making up 38% of the identified species of birds taken by cats.

“Although the species group of Sparrows could not be broken down into species, it is very likely that the dominant species observed was the House Sparrow (Passer domesitcus). Sparrows were also the most commonly observed depredated species found in England and Australia [6, 7].” [1]

While I’m not prepared to suggest that the cats’ heavy predation of House Sparrows is responsible for the increasing numbers of Eastern Bluebirds, perhaps it’s not as far-fetched as it sounds. Similar assertions have been made regarding the role of cats in the larger ecosystem (though such claims are rarely in defense of the cats).

Nature’s interconnectedness rarely makes for punchy sound bites or bumper sticker aphorisms. Then, too, such complex relationships are often overlooked, ignored, or dismissed simply because they don’t fit cleanly into one’s argument.

Literature Cited
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. Brewer, R., McPeek, G.A., and Adams, R.J., The atlas of breeding birds of Michigan. 1991, East Lansing, MI: Michigan State University Press.

3. Keller, C.M.E. and Scallan, J.T., “Potential Roadside Biases Due to Habitat Changes along Breeding Bird Survey Routes.” The Condor. 1999. 101(1): p. 50-57.

4. Rosenstock, S.S., et al., “Landbird Counting Techniques: Current Practices and an Alternative.” The Auk. 2002. 119(1): p. 46-53.

5. 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.

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

7. 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.

Repeat After Me

Listening to NPR’s On the Media this weekend, I was struck by a story (first broadcast in 2006) about how certain “sticky” numbers—however dubious—find their way into the media landscape and beyond, as On the Media co-host Brooke Gladstone noted:

“Four years ago, we delved into the mysterious number, said to be 50,000, of child predators online at any given time. It was cited by the NBC Dateline program “To Catch a Predator” and also by then Attorney General Alberto Gonzales.

But spokespersons for the FBI, the National Center for Missing and Exploited Children, and the Crimes against Children Research Center said it was not based on any research they were aware of. The A.G.’s office at the time, well, they said it came from Dateline.”

Wall Street Journal columnist Carl Bialik, who spoke to Gladstone for the story, described the process whereby such slippery figures gain traction:

“An interesting phenomenon of these numbers is that they’ll often be cited to an agency or some government body, and then a study will pick it up, and then the press will repeat it from that study. And then once it appears in the press, public officials will repeat it again, and now it’s become an official number.”

All of which sounds very familiar—Bialik could easily be describing the “official numbers” put out by so many TNR opponents. Among those that have gained the most currency are the predation estimates from the Wisconsin Study, the American Bird Conservancy’s figure for the proportion of birds in the diets of free-roaming cats, and Dauphiné and Cooper’s estimate of free-roaming cats in the U.S.

The Wisconsin Study
Despite its having been discredited long ago (see, for example, “Addressing the Wisconsin Study”), the Wisconsin Study continues to be cited as if its estimate of 8–219 million birds killed by the state’s rural cats [1] was credible. As recently as last year, Longcore et al. cited the work in their essay “Critical Assessment of Claims Regarding Management of Feral Cats by Trap–Neuter–Return.” [2]

This, despite the fact that—15 years earlier—co-author Stanley Temple told the press:

“The media has had a field day with this since we started. Those figures were from our proposal. They aren’t actual data; that was just our projection to show how bad it might be.” [3]

It’s true: the media has had a field day. Among the major newspapers to cite the Wisconsin Study are the Wall Street Journal [4], the New York Times [5], and the Los Angeles Times [6]. However, as I’ve described previously, it’s been the wildlife conservationists and bird advocates who’ve really had a field day with the Wisconsin Study:

  • The American Bird Conservancy (ABC) refers to the study, in its brochure Domestic Cat Predation on Birds and Other Wildlife. And the ABC goes one step further, pointing out that Coleman and Temple’s estimate was for rural cats, and that “suburban and urban cats add to that toll.” [7]
  • A 2009 article in Audubon Magazine suggests “cats were annually knocking off somewhere in the neighborhood of 8 million birds just in rural Wisconsin.” [9] To the magazine’s credit, they used Coleman and Temple’s low estimate—but none of the numbers from the Wisconsin Study are scientifically sound.

Birds Represent 20–30% of the Diet of Free-roaming Cats
According to an ABC report (downloadable from their website), “extensive studies of the feeding habits of domestic, free-roaming cats… show that approximately… 20 to 30 percent [of their diet] are birds.”

This, apparently, is the same report that Ellen Perry Berkeley debunked in her book, TNR Past Present and Future: A history of the trap-neuter-return movement, noting that the ABC’s 20–30% figure was not based on “extensive studies” at all. [10] In fact, just three sources were used: the now-classic “English Village” study by Churcher and Lawton [11], the Wisconsin Study (described above), and Mike Fitzgerald’s contribution to “The Domestic Cat: The Biology of Its Behaviour.” [12]

This gets a little complicated, so bear with me.

When Churcher and Lawton reported, “overall, birds comprised 35% of the total catch,” [11] they were referring to prey tallies recorded by study participants—not to the overall diets of the cats involved. Figures obtained through similar methods for the Wisconsin Study were 20–23%, [1, 13, 14] which the authors suggest—citing Fitzgerald’s comprehensive review of predation and dietary studies—are in line with other work:

“Extensive studies of the feeding habits of free-ranging domestic cats over 50 years and four continents [12] indicate that small mammals make up approximately 70% of these cats’ prey while birds make up about 20%.” [14]

But they’re comparing apples and oranges. Both the English Village and Wisconsin Studies report the percentage of birds returned as a portion of the “total catch,” whereas Fitzgerald reports percentage by frequency (i.e., the occurrence of birds in the stomach contents or scats of free-roaming cats), a point apparently lost on Coleman and Craven. The 21% figure [12] they refer to, then, is simply not comparable to their own (or that of the English Village study, a fact Churcher and Lawton acknowledge in their paper). As Berkeley notes, “this 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!”

To put all of this into more familiar terms, it’s a bit like saying that coffee makes up 20–30% of the American diet versus saying that 20–30% of Americans drink coffee each day.

Nevertheless, 13 years after the ABC first published its report, the myth persists. The report—including the mistaken dietary figures—is still available. And the National Audubon Society has helped perpetuate the error, noting in its Resolution Regarding Control and Management of Feral and Free-Ranging Domestic Cats:

“…it has been estimated that birds represent 20–30% of the prey of feral and free-ranging domestic cats.”

Estimates of Free-roaming Cats
In January, Steve Holmer, the ABC’s Senior Policy Advisor, told the Los Angeles Times, “The latest estimates are that there are about . . . 160 million feral cats [nationwide].” Sounds like an awful lot of cats—nearly one for every two humans in the country. So where does this figure come from?

The source is a paper by Nico Dauphiné and Robert Cooper (which can be downloaded via the ABC website), presented at the Fourth International Partners in Flight conference. In it, Dauphiné and Cooper use some remarkably creative accounting, beginning with an unsubstantiated estimate of unowned cats, to which they add an inflated number of owned cats that spend time outdoors. In the end, they conclude that there are “117–157 million free-ranging cats in the United States.” [15] (For a more thorough explanation, see my previous post on the subject.)

Estimating the number of free-roaming cats wasn’t even the point of their paper. As the title—“Impacts of Free-ranging Domestic Cats (Felis catus) On Birds In the United States: A Review of Recent Research with Conservation and Management Recommendations”—suggests, the primary purpose was to describe the cats’ impact on birds. The authors’ exaggerated figure was merely a convenient route to their estimate of birds killed annually by cats: “a minimum of one billion birds” [15] (which, it should be clear, has the potential to become a very sticky number).

Holmer goes a step further, using only the upper limit of the range published by Dauphiné and Cooper, and making the subtle—but important—shift from free-ranging to feral cats.

When I asked him about this, he explained that those figures were “based on an earlier version of Nico’s latest paper and are now being updated in our materials.” I don’t know that any such changes were made; and in any event, the bogus estimate has already been published in the L.A. Times—as if it were true.

*     *     *

TNR opponents will often point to the vast collection of research studies, government reports, news accounts, and the like, that support their assertions. Drill down a bit into that collection, though, and they all start to look alike: the same familiar sources, the same flawed studies—and the same bogus figures. These figures have become the kind of “official numbers” Bialik refers to: quantitative poseurs owing their popularity to tireless—and irresponsible—repetition more than anything else.

Literature Cited
1. 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

2. 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.

3. Elliott, J., The Accused, in The Sonoma County Independent. 1994. p. 1, 10.

4. Sterba, J.P., Tooth and Claw: Kill Kitty?, in Wall Street Journal. 2002: New York. p. A.1

5. Barcott, B., Kill the Cat That Kills the Bird?, in New York Times. 2007: New York. http://www.nytimes.com/2007/12/02/magazine/02cats-v–birds-t.html

6. Kennedy, J.M., Killer Among Us, in Los Angeles Times. 2003: Los Angeles. http://articles.latimes.com/2003/dec/23/news/os-cat23

7. 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

8. FWS, Migratory Bird Mortality. 2002, U.S. Fish and Wildlife Service: Arlington, VA. www.fws.gov/birds/mortality-fact-sheet.pdf

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

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

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

12. 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.

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

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

15. 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.

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.

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 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.

The Work Speaks—Part 3: Predatory Blending?

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

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

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

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

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

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

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

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

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

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

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

But Baker et al. express caution about their findings:

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

The Work Speaks—Part 2: Sample-Minded Research

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

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

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

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

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

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

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

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

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

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

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

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

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

*     *     *

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

A Critical Assessment of “Critical Assessment”—Part 1

The first in a series of posts that breaks down my critique of the essay “Critical Assessment of Claims Regarding Management of Feral Cats by Trap-Neuter-Return” (Conservation Biology, Volume 23, No. 4, 887–894) by Travis Longcore, Catherine Rich, and Lauren M. Sullivan.

How many birds are killed by cats? It’s a fair question. And if Longcore et al. are to be believed, we actually have a pretty good handle on this issue:

Feral and free-roaming cats are efficient predators, and their abundance results in substantial annual mortality of wildlife. Churcher and Lawton (1987) concluded that cats were responsible for 30% of the mortality of House Sparrows (Passer domesticus) in an English village. May (1988) extrapolated their results to an estimated 100 million birds and small mammals killed per year in England. Although this extrapolation is often criticized for the limited geographic scope and number of cats studied, Woods et al. (2003) confirmed and refined this result with a larger sample size and geographic area that included England, Scotland, and Wales. From a survey of cat owners that documented prey returned by 696 cats, Woods et al. (2003) estimated that the 9 million cats in Britain kill at least 52–63 million mammals, 25–29 million birds, and 4–6 million reptiles each summer. In North America Coleman and Temple (1996) developed estimates of cat densities in Wisconsin and associated mortality of 8–217 million birds per year.

The relationship between cat predation and bird populations is highly complex, and our understanding quite limited—something Longcore et al. only hint at. It doesn’t help matters that results of small, isolated studies are often extrapolated from rural to urban environments, from one region to another, and so forth. In 1995, Churcher himself cautioned against making such leaps: “I’d be very wary about extrapolating our results even for the rest of Britain, let alone America,” he told Catnip, a newsletter published by the Cummings School of Veterinary Medicine at Tufts University.

Actually, Churcher went much further: “I don’t really go along with the idea of cats being a threat to wildlife. If the cats weren’t there, something else would be killing the sparrows or otherwise preventing them from breeding.” [1] Although Longcore et al. seem eager to cite Churcher and Lawton’s now-classic work as “evidence” of the damage cats can do, they make no mention of Churcher’s later comments (just one of many examples of their tendency to “cherry pick” from the literature only the bits and pieces that fit neatly into their argument).

But back to the number of birds killed by cats. Many of the studies on the subject—including those cited by Longcore et al.—are quite flawed. Among the numerous issues that call into question their estimates are assumptions regarding the number of cats that actually hunt, the number of cats allowed outdoors, the number of cats that live in a particular area, and so forth. And then, of course, there are the risks inherent in estimating population numbers and characteristics based on a small sample size.

(In fact, Woods et al. go to some lengths to emphasize the limitations of their study, conceding, for example, that they “may have focused on predatory cats.” [2] This is just one of many reasons the authors cite for requesting that their work be treated “with requisite caution”—a request apparently ignored by Longcore et al.)

By referring uncritically to such studies, Longcore et al. give far greater importance to this work than is warranted. Repeating—and therefore reinforcing—figures known to be erroneous and/or misleading is simply irresponsible.

The fact that the “English village” study and “Wisconsin Study” have been so thoroughly discredited (see, for example, the comments of Nathan Winograd, director of the No Kill Advocacy Center, and a report by Laurie D. Goldstein, Christine L. O’Keefe, and Heidi L. Bickel) raises some unsettling questions about their inclusion in a paper billed as a “critical assessment.” For example: Are the authors interested enough in rigorous scientific inquiry to look beyond “the usual suspects” in their assessment of the key issues?

One might also wonder: Given the important literature that Longcore et al. choose to overlook, ignore, or dismiss (to be addressed in detail in future posts), what is their motivation for writing the essay in the first place? Actually, this question was answered in January, two months after the paper’s publication, when L.A. Superior Court Judge Thomas McKnew decided in favor of an injunction against publicly supported TNR in Los Angeles (LASC BS115483). The Urban Wildlands Group (for which Longcore serves as Science Director, and Rich as Executive Officer) was the lead petitioner in the case.

If “Critical Assessment” is any indication, the case had much more to do with politics, PR, and marketing than with science.

References
1. n.a. (1995). What the Cat Dragged In. Catnip, 4–6.

2. Woods, M., Mcdonald, R. A., & Harris, S. (2003). Predation of wildlife by domestic cats Felis catus in Great Britain. Mammal Review, 33(2), 174-188.

Works cited in “Critical Assessment” excerpt:

• Churcher, P. B., & Lawton, J. H. (1987). Predation by domestic cats in an English village. Journal of Zoology, 212(3), 439-455.
• Coleman, J. S., & Temple, S. A. (1996). On the Prowl. Wisconsin Natural Resources, 20, 4–8.
• May, R. M. (1988). Control of feline delinquency. Nature, 332, 392-393.