No evidence of host-specific egg mimicry in Asian koels.

Avian brood parasitism is costly for the host, in many cases leading to the evolution of defenses like discrimination of parasitic eggs. The parasite, in turn, may evolve mimetic eggs as a counter-adaptation to host egg rejection. Some generalist parasites have evolved host-specific races (gentes) that may mimic the eggs of their main hosts, while others have evolved 'jack-of-all-trades' egg phenotypes that mimic key features of the eggs of several different host species. The Asian koel (Eudynamys scolopaceus) is a widely distributed generalist brood parasite that exploits a wide range of host species. Based on human vision, previous studies have described Asian koel eggs as resembling those of its main host, the house crow (Corvus splendens). Using measurements of egg length and breadth, digital image analysis, reflectance spectrophotometry and avian visual modelling, we examined Asian koel egg variation and potential mimicry in egg size and shape, and eggshell pattern and color in three sympatrically occurring host species in Bangladesh: the common myna (Acridotheres tristis), house crow, and long-tailed shrike (Lanius schach). We found some differences among Asian koel eggs laid in different host nests: a) Asian koel eggs in long-tailed shrike nests were larger than those laid in common myna and house crow nests, and b) Asian koel eggs in house crow nests were less elongated than those in common myna nests. However, these changes in Asian koel egg volume and shape were in the opposite direction with respect to their corresponding host egg characteristics. Thus, our study found no evidence for Asian koel host-specific egg mimicry in three sympatrically occurring host species.

Paint it black: Efficacy of increased wind turbine rotor blade visibility to reduce avian fatalities.

As wind energy deployment increases and larger wind-power plants are considered, bird fatalities through collision with moving turbine rotor blades are expected to increase. However, few (cost-) effective deterrent or mitigation measures have so far been developed to reduce the risk of collision. Provision of "passive" visual cues may enhance the visibility of the rotor blades enabling birds to take evasive action in due time. Laboratory experiments have indicated that painting one of three rotor blades black minimizes motion smear (Hodos 2003, Minimization of motion smear: Reducing avian collisions with wind turbines). We tested the hypothesis that painting would increase the visibility of the blades, and that this would reduce fatality rates in situ, at the Smøla wind-power plant in Norway, using a Before-After-Control-Impact approach employing fatality searches. The annual fatality rate was significantly reduced at the turbines with a painted blade by over 70%, relative to the neighboring control (i.e., unpainted) turbines. The treatment had the largest effect on reduction of raptor fatalities; no white-tailed eagle carcasses were recorded after painting. Applying contrast painting to the rotor blades significantly reduced the collision risk for a range of birds. Painting the rotor blades at operational turbines was, however, resource demanding given that they had to be painted while in-place. However, if implemented before construction, this cost will be minimized. It is recommended to repeat this experiment at other sites to ensure that the outcomes are generic at various settings.

A dead giveaway: Foraging vultures and other avian scavengers respond to auditory cues.

Carrion represents an unpredictable and widely distributed primary food source for vultures and other avian scavengers. Avian scavengers in African savanna ecosystems are reported to rely exclusively on visual stimuli to locate carcasses. However, carnivores' predation of large mammalian herbivores and subsequent competition for access to the carcass can result in considerable noise, often audible over long distances and for prolonged periods. Vultures and other avian scavengers may therefore detect and respond to these auditory cues, as do the mammalian carnivores alongside which vultures have coevolved, but this has not been investigated to date. Working in the Serengeti ecosystem, Tanzania, we used diurnal auditory broadcasts to simulate predation and competitive carnivore feeding interactions. Based on the current understanding of avian scavenger ecology, we hypothesized that avian responses to call-in stations would be evoked exclusively by visual, rather than auditory, cues. We therefore predicted that (a) the arrival of avian scavengers at call-in stations should be preceded and facilitated by mammalian carnivores and that (b) the arrival of avian scavengers would be positively correlated with the number of mammalian scavengers present, which would increase detectability. We recorded 482 birds during 122 separate playback events. In 22% of these instances, avian scavengers arrived first, ruling out responses based exclusively on visual observations of mammalian carnivores, thereby contradicting our first prediction. Furthermore, the first avian arrivals at survey sessions were inversely related to the number of hyenas and jackals present, contradicting our second prediction. Since no bait or carcasses were used during the experiments, these responses are indicative of the birds' ability to detect and respond to audio stimuli. Our findings challenge the current consensus of sensory perception and foraging in these species and provide evidence that avian scavengers have the ability to use sound to locate food resources.

Effect of tower base painting on willow ptarmigan collision rates with wind turbines.

Birds colliding with turbine rotor blades is a well-known negative consequence of wind-power plants. However, there has been far less attention to the risk of birds colliding with the turbine towers, and how to mitigate this risk.Based on data from the Smøla wind-power plant in Central Norway, it seems highly likely that willow ptarmigan (the only gallinaceous species found on the island) is prone to collide with turbine towers. By employing a BACI-approach, we tested if painting the lower parts of turbine towers black would reduce the collision risk.Overall, there was a 48% reduction in the number of recorded ptarmigan carcasses per search at painted turbines relative to neighboring control (unpainted) ones, with significant variation both within and between years.Using contrast painting to the turbine towers resulted in significantly reduced number of ptarmigan carcasses found, emphasizing the effectiveness of such a relatively simple mitigation measure.

Adaptive responses of animals to climate change are most likely insufficient.

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

From Svalbard to Siberia: Passerines breeding in the High Arctic also endure the extreme cold of the Western Steppe.

Few species are adapted to high latitudes, and many over-winter in milder climates with migrations involving extensive barrier crossings. By escaping extreme conditions for the majority of the year, physiological and behavioural adaptations presumably need to be less pronounced. The snow bunting Plectrophenax nivalis is the most northerly breeding passerine. We tracked the Svalbard population using geolocators to reveal that these individuals not only breed in environmental extremes, but also spend the winters in the severe cold and highly stochastic weather conditions of the Siberian steppe. Migratory strategies appeared to be flexible between individuals and years. However, common wintering grounds in the Asian Western Steppe were identified, where birds could utilise vast crop- and grasslands while enduring low ambient temperatures. The timing of significant long distance movements was consistent among individuals, and the autumn routing of the >1000 km open water flight to Novaya Zemlya incurred favourable wind assistance and lower risk of precipitation, compared to the shorter route between Svalbard and Norway used in spring. Presumably, Svalbard snow buntings are physiologically well-adapted to extreme conditions and their migration, rather being a retreat from physiologically demanding conditions, allows utilisation of an abundance of resources in the Asian Steppe.

Characteristics determining host suitability for a generalist parasite.

Host quality is critical for parasites. The common cuckoo Cuculus canorus is a generalist avian brood parasite, but individual females show strong preference for a specific host species. Here, we use three extensive datasets to investigate different host characteristics determining cuckoo host selection at the species level: (i) 1871 population-specific parasitism rates collected across Europe; (ii) 14 K cases of parasitism in the United Kingdom; and (iii) 16 K cases of parasitism in Germany, with data collected during the period 1735-2013. We find highly consistent effects of the different host species traits across our three datasets: the cuckoo prefers passerine host species of intermediate size that breed in grass- or shrubland and that feed their nestlings with insects, and avoids species that nest in cavities. Based on these results, we construct a novel host suitability index for all passerine species breeding in Europe, and show that host species known to have a corresponding cuckoo host race (gens) rank among the most suitable hosts in Europe. The distribution of our suitability index shows that host species cannot be classified as suitable or not but rather range within a continuum of suitability.

Ancient origin and maternal inheritance of blue cuckoo eggs.

Maternal inheritance via the female-specific W chromosome was long ago proposed as a potential solution to the evolutionary enigma of co-existing host-specific races (or 'gentes') in avian brood parasites. Here we report the first unambiguous evidence for maternal inheritance of egg colouration in the brood-parasitic common cuckoo Cuculus canorus. Females laying blue eggs belong to an ancient (∼2.6 Myr) maternal lineage, as evidenced by both mitochondrial and W-linked DNA, but are indistinguishable at nuclear DNA from other common cuckoos. Hence, cuckoo host races with blue eggs are distinguished only by maternally inherited components of the genome, which maintain host-specific adaptation despite interbreeding among males and females reared by different hosts. A mitochondrial phylogeny suggests that blue eggs originated in Asia and then expanded westwards as female cuckoos laying blue eggs interbred with the existing European population, introducing an adaptive trait that expanded the range of potential hosts.

Egg arrangement in avian clutches covaries with the rejection of foreign eggs.

In birds, the colour, maculation, shape, and size of their eggs play critical roles in discrimination of foreign eggs in the clutch. So far, however, no study has examined the role of egg arrangement within a clutch on host rejection responses. We predicted that individual females which maintain consistent egg arrangements within their clutch would be better able to detect and reject foreign eggs than females without a consistent egg arrangement (i.e. whose eggs change positions more often across incubation). We tested this "egg arrangement hypothesis" in blackbirds (Turdus merula) and song thrush (T. philomelos). Both species are suitable candidates for research on egg rejection, because they show high inter-individual variation and individual repeatability in egg rejection responses. As predicted, using our custom-defined metrics of egg arrangement, rejecter females' clutches showed significantly more consistent patterns in egg arrangement than acceptor females' clutches. Only parameters related to blunt pole showed consistent differences between rejecters and acceptors. This finding makes biological sense because it is already known that song thrush use blunt pole cues to reject foreign eggs. We propose that a disturbance of the original egg arrangement pattern by the laying parasite may alert host females that maintain a consistent egg arrangement to the risk of having been parasitized. Once alerted, these hosts may shift their discrimination thresholds to be more restrictive so as to reject a foreign egg with higher probability. Future studies will benefit from experimentally testing whether these two and other parasitized rejecter host species may rely on the use of consistent egg arrangements as a component of their anti-parasitic defence mechanisms.

Sex allocation in relation to host races in the brood-parasitic common cuckoo (Cuculus canorus).

Sex allocation theory and empirical evidence both suggest that natural selection should favour maternal control of offspring sex ratio in relation to their ability to invest in the offspring. Generalist parasites constitute a particularly interesting group to test this theory as different females commonly utilize different host species showing large variation in provisioning ability. The common cuckoo (Cuculus canorus) is a generalist brood parasite that lays its eggs in the nest of many different passerine birds, but each female tends to specialize on one particular host species giving rise to highly specialized host races. The different host species show large variation in their ability to invest in the parasitic offspring, presenting an opportunity for female cuckoos to bias offspring sex ratio in relation to host species quality. Here, we investigate host-race specific sex allocation controlling for maternal identity in the common cuckoo. We found no evidence of any significant relationship between host race and sex ratio in one sympatric population harbouring three different host races, or in a total of five geographically separated populations. There was also no significant association between host quality, as determined by species-specific female host body mass, and cuckoo sex ratio. Finally, we found no significant relationship between individual cuckoo maternal quality, as determined by her egg volume, and sex ratio within each host race. We conclude that the generalist brood-parasitic common cuckoo show no significant sex-ratio bias in relation to host race and discuss this finding in light of gene flow and host adaptations.

Are cuckoos maximizing egg mimicry by selecting host individuals with better matching egg phenotypes?

BACKGROUND: Avian brood parasites and their hosts are involved in complex offence-defense coevolutionary arms races. The most common pair of reciprocal adaptations in these systems is egg discrimination by hosts and egg mimicry by parasites. As mimicry improves, more advanced host adaptations evolve such as decreased intra- and increased interclutch variation in egg appearance to facilitate detection of parasitic eggs. As interclutch variation increases, parasites able to choose hosts matching best their own egg phenotype should be selected, but this requires that parasites know their own egg phenotype and select host nests correspondingly. METHODOLOGY/PRINCIPAL FINDINGS: We compared egg mimicry of common cuckoo Cuculus canorus eggs in naturally parasitized marsh warbler Acrocephalus palustris nests and their nearest unparasitized conspecific neighbors having similar laying dates and nest-site characteristics. Modeling of avian vision and image analyses revealed no evidence that cuckoos parasitize nests where their eggs better match the host eggs. Cuckoo eggs were as good mimics, in terms of background and spot color, background luminance, spotting pattern and egg size, of host eggs in the nests actually exploited as those in the neighboring unparasitized nests. CONCLUSIONS/SIGNIFICANCE: We reviewed the evidence for brood parasites selecting better-matching host egg phenotypes from several relevant studies and argue that such selection probably cannot exist in host-parasite systems where host interclutch variation is continuous and overall low or moderate. To date there is also no evidence that parasites prefer certain egg phenotypes in systems where it should be most advantageous, i.e., when both hosts and parasites lay polymorphic eggs. Hence, the existence of an ability to select host nests to maximize mimicry by brood parasites appears unlikely, but this possibility should be further explored in cuckoo-host systems where the host has evolved discrete egg phenotypes.

Egg phenotype matching by cuckoos in relation to discrimination by hosts and climatic conditions.

Although parasites and their hosts often coexist in a set of environmentally differentiated populations connected by gene flow, few empirical studies have considered a role of environmental variation in shaping correlations between traits of hosts and parasites. Here, we studied for the first time the association between the frequency of adaptive parasitic common cuckoo Cuculus canorus phenotypes in terms of egg matching and level of defences exhibited by its reed warbler Acrocephalus scirpaceus hosts across seven geographically distant populations in Europe. We also explored the influence of spring climatic conditions experienced by cuckoos and hosts on cuckoo-host egg matching. We found that between-population differences in host defences against cuckoos (i.e. rejection rate) covaried with between-population differences in degree of matching. Between-population differences in host egg phenotype were associated with between-population differences in parasitism rate and spring climatic conditions, but not with host level of defences. Between-population differences in cuckoo egg phenotype covaried with between-population differences in host defences and spring climatic conditions. However, differences in host defences still explained differences in mimicry once differences in climatic conditions were controlled, suggesting that selection exerted by host defences must be strong relative to selection imposed by climatic factors on egg phenotypes.

Female crowing and differential responses to simulated conspecific intrusion in male and female Hainan partridge (Arborophila ardens).

Vocalizations of Hainan partridges (Arborophila ardens) during the breeding season were studied, by making use of playback experiments, in Yinggeling Nature Reserve, Hainan Island, China. Both males and females were found to crow and to produce duet calls during the breeding season, and there was a sex difference in crowing as shown by spectrogram analyses. Both sexes responded strongly to simulated intrusion from pairs of conspecifics. However, it was found that males responded to intrasexual intrusion to a greater extent than to intersexual intrusion. Moreover, females responded strongly to both intra- and intersexual intrusion. Female-female response is proposed as a behavioral mechanism that favors monogamy in the Hainan partridge, and a generally high response to any kind of vocal stimuli indicates that females may play an important role in territory defense in this species. To our knowledge, this is the first report for female crowing in the genus Arborophila, and the second case for female crowing in the family Phasianidae.

Constraints on host choice: why do parasitic birds rarely exploit some common potential hosts?

1. 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.

Genetic differentiation among sympatric cuckoo host races: males matter.

Generalist parasites regularly evolve host-specific races that each specialize on one particular host species. Many host-specific races originate from geographically structured populations where local adaptations to different host species drive the differentiation of distinct races. However, in sympatric populations where several host races coexist, gene flow could potentially disrupt such host-specific adaptations. Here, we analyse genetic differentiation among three sympatrically breeding host races of the brood-parasitic common cuckoo, Cuculus canorus. In this species, host-specific adaptations are assumed to be controlled by females only, possibly via the female-specific W-chromosome, thereby avoiding that gene flow via males disrupts local adaptations. Although males were more likely to have offspring in two different host species (43% versus 7%), they did not have significantly more descendants being raised outside their putative foster species than females (9% versus 2%). We found significant genetic differentiation for both biparentally inherited microsatellite DNA markers and maternally inherited mitochondrial DNA markers. To our knowledge, this is the first study that finds significant genetic differentiation in biparentally inherited markers among cuckoo host-specific races. Our results imply that males also may contribute to the evolution and maintenance of the different races, and hence that the genes responsible for egg phenotype may be found on autosomal chromosomes rather than the female-specific W-chromosome as previously assumed.

Coevolution in action: disruptive selection on egg colour in an avian brood parasite and its host.

BACKGROUND: Trait polymorphism can evolve as a consequence of frequency-dependent selection. Coevolutionary interactions between hosts and parasites may lead to selection on both to evolve extreme phenotypes deviating from the norm, through disruptive selection. METHODOLOGY/PRINCIPAL FINDING: Here, we show through detailed field studies and experimental procedures that the ashy-throated parrotbill (Paradoxornis alphonsianus) and its avian brood parasite, the common cuckoo (Cuculus canorus), have both evolved egg polymorphism manifested in discrete immaculate white, pale blue, and blue egg phenotypes within a single population. In this host-parasite system the most common egg colours were white and blue, with no significant difference in parasitism rates between hosts laying eggs of either colour. Furthermore, selection on parasites for countering the evolution of host egg types appears to be strong, since ashy-throated parrotbills have evolved rejection abilities for even partially mimetic eggs. CONCLUSIONS/SIGNIFICANCE: The parrotbill-cuckoo system constitutes a clear outcome of disruptive selection on both host and parasite egg phenotypes driven by coevolution, due to the cost of parasitism in the host and by host defences in the parasite. The present study is to our knowledge the first to report the influence of disruptive selection on evolution of discrete phenotypes in both parasite and host traits in an avian brood parasitism system.

Evidence for egg discrimination preceding failed rejection attempts in a small cuckoo host.

Given the high costs of avian obligate brood parasitism, host individuals are selected to reject parasitic eggs they recognize as foreign. We show that rejection may not necessarily follow egg discrimination when selective removal of the parasitic egg is difficult. We studied egg rejection behaviour in a small host of the common cuckoo Cuculus canorus, the eastern olivaceous warbler Hippolais pallida, by experimental parasitism with model and real non-mimetic cuckoo eggs and video recordings of host behaviour. Hosts pecked 87 per cent (20 out of 23) of the model eggs but eventually accepted 43.5 per cent (10 out of 23) of them. A similar pattern was found for real cuckoo eggs, which were all pecked, but as many as 47 per cent (7 out of 15) of them were accepted. To our knowledge, this is the first demonstration of a cuckoo host discriminating against real parasitic eggs but often accepting them. Our results also show that in host species experiencing difficulties in performing puncture ejection, non-mimetic cuckoo eggs may avoid rejection by means of their unusually high structural strength.

The importance of clutch characteristics and learning for antiparasite adaptations in hosts of avian brood parasites.

There is considerable variation in rejection rates of parasitic eggs among hosts of avian brood parasites. In this article, we develop a model that can be used to predict host egg rejection behavior in brood parasite-host systems in general, by considering both intra- and interclutch variation in host egg appearance; clutch characteristics that may be important in calculating the fitness of individuals adopting rejecter or acceptor strategies. In addition, we consider the importance of learning the appearance of own eggs during the first breeding attempt and host probability of survival between breeding seasons on evolution of rejection behavior. Based on this model we can predict at which level of parasitism fitness of rejecter individuals is higher than that of acceptor individuals and vice versa. The model analyses show that variation in egg appearance can be a key factor for the evolution of host defense against parasitism. In more detail, analyses show that we should expect to find a prolonged learning period only in hosts that have a high intraclutch variation in egg appearance, because such hosts may potentially experience high costs in terms of recognition errors. Furthermore, learning is in general more adaptive in parasite-host systems in which hosts do have some reproductive success even when parasitized, and when parasitism rates are moderate. By including variables that have not been considered in previous models, our model represents a useful tool in investigations of host rejection behavior in various host-parasite systems.

First evidence of regular common cuckoo, Cuculus canorus, parasitism on eastern olivaceous warblers, Hippolais pallida elaeica.

Coevolution is defined as specialized relationships between species that lead to a reciprocal evolutionary change. A particularly suitable model system for studying coevolution is the interactions between obligate avian brood parasites and their hosts. The common cuckoo (Cuculus canorus, hereafter cuckoo) is a well-known brood parasite, which utilizes a range of smaller passerines as hosts. However, warblers of the genus Hippolais have rarely been reported as being victims of cuckoos, and furthermore, few data exist on the occurrence of antiparasite defenses in these hosts. In this study, we examined possible host-parasite coevolution between cuckoos and eastern olivaceous warblers (Hippolais pallida elaeica, hereafter olivaceous warblers) in three closely situated areas in northwestern Bulgaria. The olivaceous warbler has never been reported to be a regular cuckoo host. However, the present study, carried out in 2001-2003 shows that the olivaceous warbler is regularly and heavily parasitized by the cuckoo in this area. Parasitism rate was high (26.6%, 34/128) and consistent among years, with some variation between areas. The cuckoo egg mimicry was moderately good, and olivaceous warbler rejection rate of such eggs was 50%. Cuckoo eggs laid in olivaceous warbler nests had a whitish to whitish-green ground color, and the majority appeared to be distinctly different from cuckoo eggs found in other host species in the area. The olivaceous warbler proved to be a rather good host for cuckoos as 20.6% (7/34) of cuckoo eggs laid produced fledglings, a breeding success comparable to other suitable hosts in Europe. This is the first in-depth study of brood parasitism in a warbler of the genus Hippolais, and cuckoos parasitizing olivaceous warblers probably represent a previously unknown gens.

Obligate brood parasites as selective agents for evolution of egg appearance in passerine birds.

Many passerine host species have counteracted the parasite egg mimicry in their coevolutionary arms race with the common cuckoo (Cuculus canorus) by evolving increased interclutch and reduced intraclutch variation in egg appearance. Such variations make it easier for hosts to recognize a foreign egg, reduce the possibility of making recognition errors, and reduce the ability of the cuckoo to mimic the eggs of a particular host. Here, we investigate if such clutch characteristics have evolved among North American passerines. We predict that due to the absence of brood parasites with egg mimicry on this continent, these passerines should (1) not show any relationship between rejection rates and intra- or interclutch variation, and (2) intraclutch variation should be lower and interclutch variation higher in European hosts exposed to cuckoo parasitism as compared to North American hosts parasitized by cowbirds. Here we present data that show support for most of these and other predictions, as well as when controlling statistically for effects of common descent. However, the effect of continent on intraclutch variation was less than predicted and we discuss a possible reason for this. All things considered, the results demonstrate that parasitism by a specialist brood parasite with egg mimicry is a powerful selective force regarding the evolution of egg characteristics in passerine birds.