Video evidence that cuckoos farm their hosts by ejecting nestlings.

When host nests are scarce, avian brood parasites would benefit from behaviours that increase the availability of suitable nests. Several studies reported ejection of host nestlings from nests by brood parasites; however, whether brood parasites do so to induce the host to re-nest and thus increase opportunities for future parasitism (i.e. 'farming' behaviour) remains unclear. Here, we report observational evidence of farming behaviour by a common cuckoo Cuculus canorus female in a Daurian redstart Phoenicurus auroreus population: (1) the cuckoo destroyed a host nest by ejecting all nestlings, (2) the host then produced a new nest and (3) the cuckoo successfully parasitized the replacement nest. We suggest that farming behaviour may be more common, but often goes undetected because it requires intense nest monitoring.

Removal of older males increases extra-pair siring success of yearling males.

In animals, reproductive performance typically improves over time early in life. Several ultimate and proximate mechanisms may contribute to such an age-related improvement and these mechanisms can act in a relative or in an absolute sense. Low performance of young individuals may be the consequence of a comparison or competition with older individuals (relative), or it may be due to specific traits of young individuals and be unrelated to the presence of older competitors (absolute). Here, we perform a test to disentangle whether the effect of age class (yearling or older) on male extra-pair siring success is relative or absolute. Male age is the most consistent predictor of male extra-pair siring success across bird species, yet the mechanisms underlying this pattern are not well understood. Low extra-pair siring success of yearling males may be a consequence of the presence of older ("adult") males (hypothesis 1), because adult males are more successful in intra- and intersexual interactions or because females prefer to copulate with adult males when available (relative preference). Alternatively, low extra-pair siring success of yearlings may be independent of the presence of adult males (hypothesis 2), for example, if yearling males on average invest less in extra-pair behavior or if females avoid them as extra-pair mates, independent of the availability of older males (absolute preference). To distinguish between these 2 hypotheses, we experimentally manipulated the age structure of a nest-box-breeding population of blue tits (Cyanistes caeruleus) by removing almost all adult males, and compared patterns of extra-pair paternity in the experimental year with those from the preceding 15 "control" years. Removal of adult males resulted in a substantial increase in the extra-pair siring success of yearling males compared to the "control" years, but did not affect the population-level frequency of extra-pair paternity or its spatial patterns. Our results provide clear evidence that extra-pair siring success of yearlings can increase and that it depends on the presence of older males in the population, indicating a relative effect of age on reproductive performance. These results suggest that older males outcompete yearling males in direct or indirect interactions, in sperm competition or as a result of differences in attractiveness to females.

Sexual selection for extreme physical performance in a polygynous bird is associated with exceptional sex differences in oxygen carrying capacity.

In many animals, males compete for access to fertile females. The resulting sexual selection leads to sex differences in morphology and behaviour, but may also have consequences for physiology. Pectoral sandpipers are an arctic-breeding polygynous shorebird in which males perform elaborate displays around-the-clock and move over long distances to sample potential breeding sites, implying the need for physiological adaptations to cope with extreme endurance. We examined the oxygen carrying capacity of pectoral sandpipers, measured as the volume percentage of red blood cells in blood (haematocrit, Hct). We found a remarkable sex difference in Hct levels, with males having much higher values (58.9 ± 3.8 s.d.) than females (49.8 ± 5.3 s.d.). While Hct values of male pectoral sandpipers are notable for being among the highest recorded in birds, the sex difference we report is unprecedented and more than double that of any previously described. We also show that Hct values declined after arrival to the breeding grounds in females, but not in males, suggesting that males maintain an aerobic capacity during the mating period equivalent to that sustained during trans-hemispheric migration. We conclude that sexual selection for extreme physical performance in male pectoral sandpipers has led to exceptional sex differences in oxygen carrying capacity.

A dawn and dusk chorus will emerge if males sing in the absence of their mate.

The spring dawn and dusk chorus of birds is a widespread phenomenon, yet its origin remains puzzling. We propose that a dawn and dusk chorus will inevitably arise if two criteria are met: (1) females leave their roost later in the morning and go to roost earlier in the evening than their mate, and (2) males sing more when separated from their mate. Previous studies on blue tits (Cyanistes caeruleus) support the first criterion. We here report that males sing at a higher rate whenever they are separated from their mate and that song rate increases with the duration of female absence. These findings can explain the existence of the dawn and dusk chorus in blue tits, and they can explain why the dawn chorus is more pronounced than the dusk chorus, as is typically observed. An exhaustive literature search provides support for both criteria of the 'absent mate' hypothesis in several passerine birds. We found no evidence contradicting the hypothesis. The new hypothesis is not inconsistent with many of the existing hypotheses about dawn singing, but may be a more general explanation for the occurrence of a dawn and dusk chorus. We describe how the 'absent mate' hypothesis leads to testable predictions about daily and seasonal variation in song output.

Experimentally advancing morning emergence time does not increase extra-pair siring success in blue tit males.

Extra-pair paternity occurs frequently in socially monogamous birds, but there is substantial variation in extra-pair siring success among males. Several studies have shown that siring success relates to the timing of morning activity, with the earliest active males being more successful, suggesting that early activity is important for acquiring extra-pair copulations. However, these studies are correlational, and it, therefore, remains unclear whether the relationship between timing and extra-pair siring success is causal. An alternative explanation is that successful extra-pair sires tend to be active earlier (e.g., because they are of high quality or in good condition), but that early activity in itself does not increase siring success. We experimentally advanced the emergence time of male blue tits by exposing them to light about half an hour before their natural emergence time. Although males that were exposed to the light treatment emerged from their roost substantially earlier than males that were exposed to a control treatment, light-treated males were not more likely to sire extra-pair offspring. Furthermore, whereas control males showed the expected relation between emergence time and siring success (although not statistically significant), there was no relation between emergence time and extra-pair siring success among light-treated males. Our results suggest that the timing of emergence from the roost is not an important factor underlying extra-pair siring success.

Patterns of extra-territorial nest-box visits in a songbird suggest a role in extrapair mating.

Many animals make visits outside of their territory during the breeding period, but these are typically infrequent and difficult to observe. As a consequence, comprehensive data on extra-territorial movements at the population-level are scarce and the function of this behavior remains poorly understood. Using an automated nest-box visit tracking system in a wild blue tit population over six breeding seasons, we recorded all extra-territorial nest-box visits (n = 22 137) related to 1195 individual breeding attempts (761 unique individuals). Sixty-two percent of breeders made at least one extra-territorial visit between the onset of nest building and the day of fledging of their offspring, and individuals visited another nest-box on average on 11% of the days during this period. Visit behavior differed clearly between the sexes, with males making over three times as many extra-territorial forays as females. There was a strong overall seasonal decline in visit behavior, but this was sex dependent, with females showing a strong reduction in the number of extra-territorial visits before the onset of egg laying and males showing a strong and sudden reduction on the day their offspring hatched. The likelihood of visiting a particular nest-box declined sharply with the distance to that box, and blue tits almost exclusively visited direct neighbors. Individuals were more likely to have extrapair offspring with an individual whose box they visited, but they were not more likely to disperse to a box they had visited. Thus, our results are inconsistent with the hypothesis that extra-territorial nest-box visits serve to inform dispersal decisions, but suggest that such visits are linked to extrapair mating opportunities.

Experimental evidence that cuckoos choose host nests following an egg matching strategy.

The arms race between brood parasites and their hosts provides a classic model to study coevolution. Hosts often reject the parasitic egg, and brood parasites should therefore select host nests in which the colour of the eggs best matches that of their own. Although this hypothesis has received some support, direct experimental evidence is still lacking. Here, we report on a study of Daurian redstarts, which show a distinct egg-colour dimorphism, with females laying either blue or pink eggs. Redstarts are often parasitized by common cuckoos, which lay light blue eggs. First, we showed that cuckoo eggs were more similar in spectral reflectance to the blue than to the pink redstart egg morph. Second, we report that the natural parasitism rate was higher in blue than in pink host clutches. Third, we performed a field experiment in which we presented a dummy clutch of each colour morph adjacent to active redstart nests. In this set-up, cuckoos almost always chose to parasitize a blue clutch. Our results demonstrate that cuckoos actively choose redstart nests in which the egg colour matches the colour of their own eggs. Our study thus provides direct experimental evidence in support of the egg matching hypothesis.

Brood parasitism risk drives birds to breed near humans.

Urbanization is transforming ecosystems at a global scale and at an increasing rate,1,2 and its profound consequences for wildlife have been well documented.3,4,5,6 Understanding how animals thrive in the urban environment and how this environment affects (co-)evolutionary processes remains an important challenge.7 Urban environments can provide resources such as food or nest sites (e.g., cavities)10,8,9 and also reduce exposure to predators.11,12 For some species, urban environments may also affect susceptibility to brood parasitism,13,14 but this has never been tested experimentally. Here, we use a combination of field observations and experimental manipulations to show that Daurian redstarts, Phoenicurus auroreus, a common host of the common cuckoo, Cuculus canorus, nest in proximity to humans to avoid brood parasitism. First, redstarts were more likely to be parasitized with increasing distance to the nearest building. Second, redstarts adjusted their nesting location in response to a seasonally predictable change in the risk of brood parasitism. Third, experimentally simulating the presence of cuckoos during a period when they are naturally absent increased the likelihood that redstarts nested indoors or closer to human settlements. These findings suggest that redstarts actively choose to place their nest in the vicinity of a human residence as a defense against cuckoos. Our study exemplifies how animals take advantage of the urban environment by using it as a novel line of defense against detrimental interspecific interactions.

Host personality predicts cuckoo egg rejection in Daurian redstarts Phoenicurus auroreus.

In species that are subject to brood parasitism, individuals often vary in their responses to parasitic eggs, with some rejecting the eggs while others do not. While some factors, such as host age (breeding experience), the degree of egg matching and the level of perceived risk of brood parasitism have been shown to influence host decisions, much of the variation remains unexplained. The host personality hypothesis suggests that personality traits of the host influence its response to parasitic eggs, but few studies have tested this. We investigated the relationship between two personality traits (exploration and neophobia) and a physiological trait (breathing rate) of the host, and egg-rejection behaviour in a population of Daurian redstarts Phoenicurus auroreus in northeast China. We first show that exploratory behaviour and the response to a novel object are repeatable for individual females and strongly covary, indicating distinct personality types. We then show that fast-exploring and less neophobic hosts were more likely to reject parasitic eggs than slow-exploring and more neophobic hosts. Variation in breathing rate-a measure of the stress-response-did not affect rejection behaviour. Our results demonstrate that host personality, along the bold-shy continuum, predicts the responses to parasitic eggs in Daurian redstarts, with bold hosts being more likely to reject parasitic eggs.

Exposure to predator models during the fertile period leads to higher levels of extra-pair paternity in blue tits.

The perceived risk of predation can affect breeding behaviour and reduce reproductive success in prey species. Individuals exposed to predators may also adopt different mating tactics with potential consequences for the distribution of paternity in socially monogamous species that engage in extra-pair copulations. We experimentally increased perceived predation risk during the fertile period in blue tits Cyanistes caeruleus. Every morning between nest completion and the onset of egg laying, we presented a model of either a predator or a non-predator (control) near active nestboxes. Broods from pairs exposed to predators had higher levels of extra-pair paternity than control broods. This mainly resulted from a higher proportion of extra-pair offspring in broods with at least one extra-pair young. Females exposed to predators first emerged from the nestbox later in the morning, stayed away from the nestbox for longer and were less likely to be visited at the nest by their social mate, but we detected no behavioural differences once the model was removed. Our results suggest that the higher rates of extra-pair paternity resulted from the disruption of morning routines, which may have inhibited within-pair copulations or increased opportunities for females to engage in extra-pair copulations. We conclude that the perceived risk of predation can have substantial effects on levels of extra-pair paternity.

Complete brood failure in an altricial bird is almost always associated with the sudden and permanent disappearance of a parent.

A central goal in evolutionary ecology is to identify factors that explain variation in reproductive success, i.e. in the number of offspring produced. In altricial birds, a substantial part of this variation is determined by the number of nestlings that die before fledging, but surprisingly little is known about the proximate causes of offspring mortality during the nestling period. We used a uniquely comprehensive dataset of parental nestbox visits from seven breeding seasons to investigate the association between parental behaviour and nestling mortality in a population of blue tits (Cyanistes caeruleus). In almost all nests that suffered complete brood mortality one of the parents had suddenly disappeared during the nestling stage. In contrast, parental disappearance in nests with partial brood mortality was rare and equally common as in nests with no brood loss. With few exceptions, parents that disappeared during the nestling stage were never observed again and never returned to breed. In contrast, parents that remained after their partner disappeared were equally likely to be observed again or return to breed as parents of nests where both parents stayed. Visit rates at nests where a parent would disappear did not differ from those at nests where both parents stayed. Taken together, our results show that - in contrast to partial brood failure - complete brood failure is almost always associated with the sudden and permanent disappearance of one of the parents, probably due to predation. Partial and complete brood mortality should be treated as distinct processes that have different underlying causes.

Validation of a Fecal Glucocorticoid Assay to Assess Adrenocortical Activity in Meerkats Using Physiological and Biological Stimuli.

In mammals, glucocorticoid (i.e. GC) levels have been associated with specific life-history stages and transitions, reproductive strategies, and a plethora of behaviors. Assessment of adrenocortical activity via measurement of glucocorticoid metabolites in feces (FGCM) has greatly facilitated data collection from wild animals, due to its non-invasive nature, and thus has become an established tool in behavioral ecology and conservation biology. The aim of our study was to validate a fecal glucocorticoid assay for assessing adrenocortical activity in meerkats (Suricata suricatta), by comparing the suitability of three GC enzyme immunoassays (corticosterone, 11ß-hydroxyetiocholanolone and 11oxo-etiocholanolone) in detecting FGCM increases in adult males and females following a pharmacological challenge with adrenocorticotropic hormone (ACTH) and biological stimuli. In addition, we investigated the time course characterizing FGCM excretion, the effect of age, sex and time of day on FGCM levels and assessed the potential effects of soil contamination (sand) on FGCM patterns. Our results show that the group specific 11ß-hydroxyetiocholanolone assay was most sensitive to FGCM alterations, detecting significant and most distinctive elevations in FGCM levels around 25 h after ACTH administration. We found no age and sex differences in basal FGCM or on peak response levels to ACTH, but a marked diurnal pattern, with FGCM levels being substantially higher in the morning than later during the day. Soil contamination did not significantly affect FGCM patterns. Our results emphasize the importance of conducting assay validations to characterize species-specific endocrine excretion patterns, a crucial step to all animal endocrinology studies using a non-invasive approach.

Group augmentation and the evolution of cooperation.

The group augmentation (GA) hypothesis states that if helpers in cooperatively breeding animals raise the reproductive success of the group, the benefits of living in a resulting larger group--improved survival or future reproductive success--favour the evolution of seemingly altruistic helping behaviour. The applicability of the GA hypothesis remains debatable, however, partly owing to the lack of a clear conceptual framework and a shortage of appropriate empirical studies. We conceptualise here the GA hypothesis and illustrate that benefits of GA can accrue via different evolutionary mechanisms that relate closely to well-supported general concepts of group living and cooperation. These benefits reflect several plausible explanations for the evolutionary maintenance of helping behaviour in cooperatively breeding animals.