Summary1. Urban areas contain high densities of non-native species, which in the UK include the domestic cat Felis catus (Linnaeus 1758) and the grey squirrel Sciurus carolinensis (Gmelin 1788). The direct predation effects of domestic cats on prey populations attract intense debate, and such influences of the nest-predatory grey squirrel are receiving increasing attention. In contrast, theory predicts that sublethal and indirect effects are more important, but empirical evidence is currently lacking. 2. We conducted controlled model presentation experiments at active urban blackbird Turdus merula (Linnaeus 1758) nests to provide the first empirical evidence that quantifies the potential sublethal and indirect effects of predators (domestic cat and grey squirrel) on avian reproductive success. 3. Domestic cat models reduced subsequent parental provisioning rates, a strong indicator of sublethal effects, by one-third relative to a nonpredatory rabbit Oryctolagus cuniculus (Linnaeus 1758) control. There was no compensatory increase in food load size. Previous experiments demonstrate that this magnitude of reduced food delivery will reduce nestling growth rates by c. 40%. The grey squirrel model induced similar but weaker effects. 4. Following the brief presence of the domestic cat model, subsequent daily nest predation rates, chiefly by corvids, increased by an order of magnitude relative to the squirrel and rabbit models. The intensity of parental nest defence elicited in response to model presentations predicts the probability of such third-party predator predation events, and the domestic cat model generated significant increases in nest defence behaviour. 5. Synthesis and applications. The brief presence of a domestic cat at avian nest sites reduces subsequent provisioning rates and induces lethal trait-mediated indirect effects. We provide the first robust evidence for these latter effects in any avian predator-prey system, although they are likely to be common in many avian assemblages with high predator densities. It is imperative that future assessments of the impact of predatory species on avian prey species take lethal trait-mediated indirect effects into account, as so doing will prevent biased estimates of predator effects and facilitate the design of more effective control strategies. Full mitigation of the sublethal and indirect effects of domestic cats would problematically require permanent indoor housing.
The influence of predators on bird populations is controversial and poorly understood, especially in urban areas where predator densities can be particularly high. We assessed if fine‐scale spatial variation in predator activity and proximity have direct and indirect effects on urban songbird distributions and breeding success, by testing the hypotheses that (1) songbirds that are sensitive to nest predation select territories with reduced activity of nest predators; (2) blackbird Turdus merula, a species that experiences high nest predation rates, lays smaller clutches in territories located in areas with higher numbers of nest predators as predicted by Skutch's hypothesis; (3) songbirds that are sensitive to nest predation have higher nest predation rates in areas with greater predator activity. We tested these hypotheses using two sites in urban Sheffield, UK, and focus on nest predatory corvids and grey squirrels Sciurus carolinensis. We found no evidence that songbirds that are most sensitive to nest predation adjust their territory location in response to fine‐scale spatial variation in predator distributions. It thus seems unlikely that urban predators are indirectly regulating urban bird population size by restricting habitat availability. Blackbirds did not vary their clutch size in response to predator distributions. These findings generate the potential for an ecological trap in which prey species fail to avoid areas with the highest concentrations of nest predators, or to exhibit behavioural adjustments to reduce the risk of nest predation at such sites. We find some evidence for such ecological traps as, while fine‐scale variation in grey squirrel occurrence and activity were not associated with nest predation rates, nests located in closer proximity to corvids and in areas with higher indices of corvid activity experienced slightly higher nest predation rates.
Supplementary feeding of birds, particularly in urban areas, is often associated with increased population size and fecundity. In the UK, the non‐native Grey Squirrel Sciurus carolinensis is common in rural and urban habitats. It exploits supplementary feeders and may induce interference competition by excluding birds, but empirical evidence of this is unavailable. Using controlled model presentation experiments, we demonstrate that Grey Squirrels could reduce bird use of supplementary feeders and induce interference competition. Total bird resource use was reduced by 98% and most species exhibited similar sensitivities. The likelihood and magnitude of interference competition will depend on how rapidly displaced birds find alternative food sources; it will be greatest where there are high Grey Squirrel densities and few supplementary feeders. Other studies suggest that supplementary feeding increases Grey Squirrel numbers, and the species is also predicted to expand its non‐native range across most of Europe. Our data indicate that Grey Squirrels may eventually alter the net effect of supplementary feeding on bird populations across the European continent; increased use of squirrel‐proof feeders may help to minimize such effects.
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