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,”  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.
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.”  (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.”  (italics mine))
“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.”
“It is not prudent to manage for wildlife and allow cat feeding in the same parks.”
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.”  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.”  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.”  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.”  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.
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.”  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”  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 ) 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.” 
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.
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