Summary1. Why are some common and apparently suitable resources avoided by potential users? This interesting ecological and evolutionary conundrum is vividly illustrated by obligate brood parasites. Parasitic birds lay their eggs into nests of a wide range of host species, including many rare ones, but do not parasitize some commonly co-occurring potential hosts. 2. Attempts to explain the absence of parasitism in common potential hosts are limited and typically focused on single-factor explanations while ignoring other potential factors. We tested why thrushes Turdus spp. are extremely rarely parasitized by common cuckoos Cuculus canorus despite breeding commonly in sympatry and building the most conspicuous nests among forest-breeding passerines. 3. No single examined factor explained cuckoo avoidance of thrushes. Life-history traits of all six European thrush species and the 10 most frequently used cuckoo hosts in Europe were similar except body ⁄ egg size, nest design and nestling diet. 4. Experiments (n = 1211) in several populations across Europe showed that host defences at egg-laying and incubation stages did not account for the lack of cuckoo parasitism in thrushes. However, cross-fostering experiments disclosed that various factors during the nestling period prevent cuckoos from successfully parasitizing thrushes. Specifically, in some thrush species, the nest cup design forced cuckoo chicks to compete with host chicks with fatal consequences for the parasite. Other species were reluctant to care even for lone cuckoo chicks. 5. Importantly, in an apparently phylogenetically homogenous group of hosts, there were interspecific differences in factors responsible for the absence of cuckoo parasitism. 6. This study highlights the importance of considering multiple potential factors and their interactions for understanding absence of parasitism in potential hosts of parasitic birds. In the present study, comparative and experimental procedures are integrated, which represent a novel approach that should prove useful for the understanding of interspecific ecological relationships in general.

All animals flee from potential predators, and the distance at which this happens is optimized so the benefits from staying are balanced against the costs of flight. Because predator diversity and abundance decreases with increasing latitude, and differs between rural and urban areas, we should expect escape distance when a predator approached the individual to decrease with latitude and depend on urbanization. We measured the distance at which individual birds fled (flight initiation distance, FID, which represents a reliable and previously validated surrogate measure of response to predation risk) following a standardized protocol in nine pairs of rural and urban sites along a ca. 3000 km gradient from Southern Spain to Northern Finland during the breeding seasons 2009–2010. Raptor abundance was estimated by means of standard point counts at the same sites where FID information was recorded. Data on body mass and phylogenetic relationships among bird species sampled were extracted from the literature. An analysis of 12,495 flight distances of 714 populations of 159 species showed that mean FID decreased with increasing latitude after accounting for body size and phylogenetic effects. This decrease was paralleled by a similar cline in an index of the abundance of raptors. Urban populations had consistently shorter FIDs, supporting previous findings. The difference between rural and urban habitats decreased with increasing latitude, also paralleling raptor abundance trends. Overall, the latitudinal gradient in bird fear was explained by raptor abundance gradients, with additional small effects of latitude and intermediate effects of habitat. This study provides the first empirical documentation of a latitudinal trend in anti-predator behavior, which correlated positively with a similar trend in the abundance of predators.

Living organisms generally occur at the highest population density in the most suitable habitat. Therefore, invasion of and adaptation to novel habitats imply a gradual increase in population density, from that at or below what was found in the ancestral habitat to a density that may reach higher levels in the novel habitat following adaptation to that habitat. We tested this prediction of invasion biology by analyzing data on population density of breeding birds in their ancestral rural habitats and in matched nearby urban habitats that have been colonized recently across a continental latitudinal gradient. We estimated population density in the two types of habitats using extensive point census bird counts, and we obtained information on the year of urbanization when population density in urban habitats reached levels higher than that of the ancestral rural habitat from published records and estimates by experienced ornithologists. Both the difference in population density between urban and rural habitats and the year of urbanization were significantly repeatable when analyzing multiple populations of the same species across Europe. Population density was on average 30 % higher in urban than in rural habitats, although density reached as much as 100-fold higher in urban habitats in some species. Invasive urban bird species that colonized urban environments over a long period achieved the largest increases in population density compared to their ancestral rural habitats. This was independent of whether species were anciently or recently urbanized, providing a unique cross-validation of timing of urban invasions. These results suggest that successful invasion of urban habitats was associated with gradual adaptation to these habitats as shown by a significant increase in population density in urban habitats over time.

How do birds tell the colours of their own and foreign eggs apart? We demonstrate that perceptual modelling of avian visual discrimination can predict behavioural rejection responses to foreign eggs in the nest of wild birds. We use a photoreceptor noise-limited colour opponent model of visual perception to evaluate its accuracy as a predictor of behavioural rates of experimental egg discrimination in the song thrush Turdus philomelos. The visual modelling of experimental and natural eggshell colours suggests that photon capture from the ultraviolet and short wavelength-sensitive cones elicits egg rejection decisions in song thrushes, while inter-clutch variation of egg coloration provides sufficient contrasts for detecting conspecific parasitism in this species. Biologically realistic sensory models provide an important tool for relating variability of behavioural responses to perceived phenotypic variation.

BackgroundWhy have birds evolved the ability to reject eggs? Typically, foreign egg discrimination is interpreted as evidence that interspecific brood parasitism (IP) has selected for the host’s ability to recognize and eliminate foreign eggs. Fewer studies explore the alternative hypothesis that rejection of interspecific eggs is a by-product of host defenses, evolved against conspecific parasitism (CP). We performed a large scale study with replication across taxa (two congeneric Turdus thrushes), space (populations), time (breeding seasons), and treatments (three types of experimental eggs), using a consistent design of egg rejection experiments (n = 1057 nests; including controls), in areas with potential IP either present (Europe; native populations) or absent (New Zealand; introduced populations). These comparisons benefited from the known length of allopatry (one and a half centuries), with no gene flow between native and introduced populations, which is rarely available in host-parasite systems.ResultsHosts rejected CP at unusually high rates for passerines (up to 60%). CP rejection rates were higher in populations with higher conspecific breeding densities and no risks of IP, supporting the CP hypothesis. IP rejection rates did not covary geographically with IP risk, contradicting the IP hypothesis. High egg rejection rates were maintained in the relatively long-term isolation from IP despite non-trivial rejection costs and errors.ConclusionsThese egg rejection patterns, combined with recent findings that these thrushes are currently unsuitable hosts of the obligate parasitic common cuckoo (Cuculus canorus), are in agreement with the hypothesis that the rejection of IP is a by-product of fine-tuned egg discrimination evolved due to CP. Our study highlights the importance of considering both IP and CP simultaneously as potential drivers in the evolution of egg discrimination, and illustrates how populations introduced to novel ecological contexts can provide critical insights into brood parasite-host coevolution.