In many areas where human activities have resulted in the decline of top predators, smaller carnivores--species such as foxes, raccoons, and rats--benefit from no longer being prey items, as well as from the decreased competition over resources in the habitat. Until recently, this "mesopredator release" was thought to occur only when apex carnivores--things like tigers, wolves, and wild dogs--had been locally extirpated, leaving vacancies in the food web that their smaller brethren could step in to fill. However, a new study on pumas suggests that mesopredator release could happen by another mechanism altogether: changes to the feeding behaviors of top predators.
Scientists from the University of California's Santa Cruz campus discovered this by collaring and tracking 30 pumas between 2008 and 2013. Every four hours, the collars collected information on the location of the cats, and the resulting data points were plotted on a map. Where points were clustered, the researchers suspected a kill site and went to investigate for signs of predation. For logistical reasons, it wasn't possible to visit all potential kill sites, so, to overcome this difficulty, the researchers created computer models into which they could plug variables associated with confirmed kills--factors such as how long the pumas stayed there, whether they were there at night or during the day, and how far the cats strayed from the area. These traits were then used to determine the likelihood that the putative kill sites were, in fact, the location of a puma kill.
Close-up of a puma. Image courtesy of Bas Lammers.
What made this information so interesting was that the hunting data could be plotted on maps showing human housing densities. The study area, the Santa Cruz Mountains of California's Central Coast region, encompassed a range of anthropogenic sites, from rural areas with fewer than 1 house per hectare, to suburban locations with nearly 10. Along this disturbance gradient, the hunting behavior of pumas--specifically, female pumas--varied. As housing density increased, these huntresses spent up to 42% less time consuming prey. Their site fidelity was 36% lower, and the farthest distance traveled from the kill site was up to 31% higher.
To make sense of these numbers, it is necessary to understand a bit about the feeding strategies of apex predators like pumas. They can expend quite a lot of energy while stalking, chasing, and taking down prey. These calories can be replaced by eating whatever has just been caught, but big carnivores don't have large enough stomachs to accommodate many of their prey items--things like deer, for example--in a single sitting; instead, the hunters have to drag the body somewhere safe and revisit it until they've had their fill.
Mountain lion eating. Image courtesy of Benjamint444
The current results suggest that this is a much trickier prospect for female pumas living in anthropogenically disturbed areas. These cats weren't able to spend as much time eating, and seemed to roam much farther from where they had stashed their kills; the low site fidelity values suggest that many of the animals left the area altogether. Given these figures, it is, perhaps, unsurprising that some of the hunters in these areas were estimated to kill as many as 20 more deer per year than pumas living in the most rural areas. These data suggest that predators in high-human-density areas are having to target more prey because they are starting over after being interrupted while feeding on their previous kills.
Males seem to get off easy because they already spend less time at kills; they are adapted to eat quickly and head back out to patrol the borders of the large territories they defend. The size of their home ranges (up to 170 square km) also means that if humans become disruptive in one area, the cats can withdraw to more natural spots for a bit of privacy. Given that male territories only have approximately 16 houses per square km, this isn't too hard to do. Female pumas, however, don't have as much flexibility; their territories are smaller (as small as 51 square km), and may comprise only exurban or suburban land; their home ranges sometimes contain as many as 27 houses per square km.
Mountain lion kitten about to be outfitted with a tracking device. Image courtesy of the NPS.
Female pumas must also take care of kittens, a responsibility that requires them to bring down even more prey. To adequately feed their young, mothers may need to make more than a dozen additional kills per year. For females living in high-human-density areas, where disruptions to feeding sessions are already inflating hunting rates, this could be untenable; these mothers could begin to lose weight, suffer poor health, or even be driven to abandon their young. Indeed, the researchers provided anecdotal evidence of the last of these possibilities, suggesting that puma populations in human-disturbed sites may only be viable so long as they are replenished by young pumas migrating in from more rural areas.
While this is bad news for pumas, it is potentially great news for mesopredators. Female pumas are leaving a larger number of kills for longer periods of time, giving scavengers more of an opportunity to swoop in and have a free meal. This increased source of nutrition could allow the ecosystem to sustain larger populations of middle predators and give individual animals the energy boost they need to live longer and/or procreate more successfully. Beneficiaries could include a range of species, from raccoons and foxes all the way up to coyotes. Additional work would need to be conducted to explore whether these species are commoner or more successful in more human-dense areas, and, if so, whether those patterns can be directly attributed to puma behavior rather than other characteristics of anthropogenic environments.
Image courtesy of Tony Hisgett.
As the authors point out, "behavioral responses are often overlooked as ecosystem drivers in modified systems, overshadowed by population declines and extirpations." Their current study, however, shows that behavioral flexibility can allow species to persist in modified environments--but that this persistence may come at a cost, and have widespread implications for the habitat.
Source material: Smith, Justine A., Yiwei Wang, and Christopher C. Wilmers. 2015. Top carnivores increase their kill rates on prey as a response to human-induced fear. Proceedings B. 282: 20142711.