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Wetland ecosystems host an extensive number of avian species and play therefore a key role in biodiversity conservation. However, avifauna in wetlands is severely impacted by the anthropogenic transformation of natural habitats into other land uses. Understanding how habitat changes influence avian community dynamics is crucial for effective management and conservation of bird habitats. Here, we assessed spatio-temporal variation in taxonomic, functional, and phylogenetic diversity, and assembly structures of the bird community in the Indawgyi Wetland Ecosystem in Myanmar. Bird surveys were seasonally conducted at 120 sample plots, which were equally distributed among four habitat types subject to different level of human influences (natural lake habitat, seasonally flooded grasslands, riparian forest, and agricultural land). Across habitat types, functional and phylogenetic diversity were the highest in the lake habitat, especially during the migration season, and the lowest in the cultivated habitat. Our results indicated over-dispersed functional and phylogenetic structures in the lake habitat, with clustered structures in the remaining habitats. Diversity of species important for conservation was also higher in the lake habitat and in areas far away from human disturbances. Spatial variation of bird community diversity suggests that conservation efforts in this ecosystem should concentrate more on the lake habitat and associated grassland habitats. Moreover, our results suggest the need for habitat restoration in riparian forests and increased sustainable agricultural practices in order to improve the contribution of these habitats to the diversity in the avifauna community. Our study highlighted the importance of integrating functional and phylogenetic dimensions into biodiversity analyses providing broader ecological insights for conservation.

Telomeres, the nucleotide sequences that protect the ends of eukaryotic chromosomes, shorten with each cell division and telomere loss may be increased by factors such as oxidative stress. Telomere length (TL) decreases with age in several species, and TL or the change in TL (∆TL) have emerged as biomarkers reflecting experienced stress. In this study, we tracked changes in TL throughout the natural lifespan (from a few months to almost 9 years) of free-living house sparrows (Passer domesticus) in two different island populations. TL was measured in nestlings and subsequently up to 4 times during their lifetime. TL generally decreased with age (senescence), but we also observed instances of telomere lengthening within individuals. We found some evidence for selective disappearance of individuals with shorter telomeres through life. Early-life TL positively predicted later-life TL, but the within-individual repeatability in TL was low (9.2%). Using genetic pedigrees, we found a moderate heritability of ∆TL (h2 = 0.21), which was higher than the heritabilities of early-life TL (h2 = 0.14) and later-life TL measurements (h2 = 0.15). Cohort effects explained considerable proportions of variation in early-life TL (60%), later-life TL (53%), and ∆TL (37%), which suggests persistent impacts of the early-life environment on lifelong telomere dynamics. Individual changes in TL were independent of early-life TL. Finally, there was weak evidence for population differences in ∆TL that may be linked to ecological differences in habitat types. Combined, our results show that individual telomere biology is highly dynamic and influenced by both genetic and environmental variation in natural conditions.

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., Avian scavengers in African savanna ecosystems are reported to rely exclusively on visual stimuli to locate carcasses. Here, we report on the response of vultures and other avian scavengers to auditory cues indicative of foraging opportunities. Based on our findings, we hypothesize that these birds are able to utilize auditory cues while foraging.

Translocation of conspecific individuals to reduce extinction risk of small, isolated populations and prevent genetic depletion is a powerful tool in conservation biology. An important question is how the translocated individuals influence the long-term genetic composition of the recipient population. Here, we experimentally reinforced a house sparrow (Passer domesticus) population, and examined the impact of this translocation on allele frequencies, levels of heterozygosity and genetic differentiation over six cohorts. We found no permanent increase in the mean number of alleles across loci or levels of observed heterozygosity, but a few alleles private to the translocated individuals remained in the population and we found a short-term increase in heterozygosity. Consequently, genetic differentiation of the recipient population compared to the genetic composition prior to reinforcement was small. The limited genetic impact was due to combined effects of a small probability of establishment and low mating success for the translocated individuals, together with increased genetic drift in the recipient population. Our findings emphasize the importance of selection and genetic drift as forces that may decrease the genetic contribution of reinforcement, especially in small populations. Conservation managers should aim to improve habitat quality in the recipient population to reduce genetic drift following translocation and thereby avoid the need for continued reinforcement. Furthermore, by facilitating establishment success and selecting individuals expected to have high mating success, possibly indicated by sexually selected traits, genetic contribution of released individuals is increased which in turn will decrease reproductive skew and genetic drift.

1. Environmental conditions during early-life development can have lasting effects on individual quality and fitness. Telomere length (TL) may correlate with early-life conditions and may be an important mediator or biomarker of individual quality or pace-of-life, as periods of increased energy demands can increase telomere attrition due to oxidative stress. Thus, knowledge of the mechanisms that generate variation in TL, and the relation between TL and fitness, is important in understanding the role of telomeres in ecology and life-history evolution. 2. Here, we investigate how environmental conditions and morphological traits are associated with early-life TL and if TL predicts natal dispersal probability or components of fitness in two populations of wild house sparrows (Passer domesticus). 3. We measured morphological traits and blood TL in 2746 nestlings from 20 cohorts (1994-2013) and retrieved data on weather conditions. We monitored population fluctuations, and individual survival and reproductive output using field observations and genetic pedigrees. We then used generalized linear mixed-effects models to test which factors affected TL in early-life, and if TL predicted dispersal propensity, or was associated with recruitment probability, mortality risk, or reproductive success. 4. We found a negative effect of population density on TL, but only in one of the populations. There was a curvilinear association between TL and the maximum daily North Atlantic Oscillation (NAO) index during incubation, suggesting that there are optimal weather conditions that result in the longest TL. Dispersers tended to have shorter telomeres than non-dispersers. TL did not predict survival, but we found a tendency for individuals with short telomeres to have higher annual reproductive success. 5. Our study showed how early-life TL is shaped by effects of growth, weather conditions and population density, supporting that environmental stressors negatively affect TL in wild populations. In addition, TL may be a mediator or biomarker of individual pace-of-life, with higher dispersal rates and annual reproduction tending to be associated with shorter early-life TL in this study. However, clear associations between early-life TL and individual fitness seems difficult to establish and may differ between different populations in the wild.

Many animal populations are exposed to disturbance originating from human activities. In response to human disturbance, certain animals display a variety of potentially costly behavioural responses, such as increased antipredator behaviour or relocation to new areas. In contrast, other animals seemingly thrive in the presence of humans and benefit from human‐derived resources. Flight initiation distance (FID: the distance between predator and prey when prey starts to flee) is a measure commonly used to assess animals’ tolerance to humans. In this study, we tested how FID changes in relation to human presence in two hyrax species in Serengeti National Park. Hyraxes living on kopjes (rock outcrops) among human settlements showed a significantly shorter FID than hyraxes living on kopjes without human settlements. In addition, we found that hyraxes feeding before the experiment had shorter FID than hyraxes resting or being vigilant, and hyraxes disturbed during the early morning had shorter FID than hyraxes disturbed during late morning. We did not find any significant effects of group size or species composition on FID. Our results suggest that hyraxes living in the presence of humans are habituated and are not adversely affected by human settlements. © 2019 The Authors. Ethology published by Blackwell Verlag GmbH This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Dispersal has a crucial role determining eco-evolutionary dynamics through both gene flow and population size regulation. However, to study dispersal and its consequences, one must distinguish immigrants from residents. Dispersers can be identified using telemetry, capture-mark-recapture (CMR) methods, or genetic assignment methods. All of these methods have disadvantages, such as, high costs and substantial field efforts needed for telemetry and CMR surveys, and adequate genetic distance required in genetic assignment. In this study, we used genome-wide 200K Single Nucleotide Polymorphism data and two different genetic assignment approaches (GSI_SIM, Bayesian framework; BONE, network-based estimation) to identify the dispersers in a house sparrow (Passer domesticus) metapopulation sampled over 16 years. Our results showed higher assignment accuracy with BONE. Hence, we proceeded to diagnose potential sources of errors in the assignment results from the BONE method due to variation in levels of inter-population genetic differentiation, intra-population genetic variation and sample size. We show that assignment accuracy is high even at low levels of genetic differentiation and that it increases with the proportion of a population that has been sampled. Finally, we highlight that dispersal studies integrating both ecological and genetic data provide robust assessments of the dispersal patterns in natural populations.

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.

Generation time determines the pace of key demographic and evolutionary processes. Quantified as the weighted mean age at reproduction, it can be studied as a life-history trait that varies within and among populations and may evolve in response to ecological conditions. We combined quantitative genetic analyses with age- and density-dependent models to study generation time variation in a bird metapopulation. Generation time was heritable, and males had longer generation times than females. Individuals with longer generation times had greater lifetime reproductive success but not a higher expected population growth rate. Density regulation acted on recruit production, suggesting that longer generation times should be favoured when populations are closer to carrying capacity. Furthermore, generation times were shorter when populations were growing and longer when populations were closer to equilibrium or declining. These results support classic theory predicting that density regulation is an important driver of the pace of life-history strategies.

Investigating space use of wild birds provides important knowledge of bird behavior and ecology, which is crucial in the management and conservation of threatened species. In the present study, we used GPS satellite telemetry to investigate space use and movements of seven adult Eurasian Eagle-Owls (Bubo bubo) in Norway during breeding and post-breeding seasons. Breeding adults had a mean home range size of 42.9 km2 (SD ± 35.1 km2; 95% kernel density estimation Had hoc), and five individuals performed long (>20 km) excursive movements away from their breeding territories during autumn. Such wide-ranging behavior has not previously been reported for adults of the nominate subspecies B. b. bubo. The study demonstrates the need to investigate seasonal movements of species, populations, and age groups even if they are considered residents, and provides valuable information for management and conservation of the Eurasian Eagle-Owl. Paid open access

Most large carnivore populations are declining due to anthropogenic activities including direct persecution, prey depletion, habitat loss and degradation. protected areas (PAs) can help maintain viable large carnivore populations; however, anthropogenic activities occurring near and within PA borders or edges can reduce their effectiveness. We investigated the influence of edge effects on abundance of lions (Panthera leo) and spotted hyenas (Crocuta crocuta) in Maswa Game Reserve (MGR), a part of the Serengeti ecosystem in northern Tanzania. We conducted repeated call-ins to attract and enumerate lions and hyenas at 20 stations in MGR during June–July 2017. We used N-mixture models to estimate hyena and lion abundance in relation to land cover and distance from the south-western MGR borders which are adjacent to villages. We found lowest lion and hyena abundances by the south-western border, with abundance of both species increasing toward the eastern border adjacent to Serengeti National Park. Lions were uniformly distributed among land covers whereas hyenas were more abundant in woodlands. We suggest that reduced lion and hyena abundance near human settlements was in response to depleted prey, due to human actions. We recommend ecologically compatible land uses and effective border patrols to mitigate these adverse effects.

The understanding of how educational and spatial factors influence appreciation of conservation benefits to locals living adjacent to protected areas is a very important aspect in conservation initiatives. In this study we compared how educational and spatial factors affect the appreciation of conservation benefits to local communities adjoining the two Tanzanian national parks; Tarangire National Park (TNP) and Saadani National Park (SANAPA). Questionnaire surveys including closed-ended questions were conducted in eight villages, four near each of the two parks. A total of 400 randomly selected households were interviewed (50 in each village). Results from these questionnaires showed that the local people’s appreciation of benefits was negatively related to increasing distance from the parks boundaries. Moreover, people with secondary education or higher, appreciated to receive more benefits than those with primary and tended to appreciate benefits more often than those with no formal education. In addition, people living around TNP responded that the park itself benefited more in terms of benefits from tourism compared to communities surrounding SANAPA. The underlying mechanisms for establishing strong relationships, e.g., by initiating conservation benefits close to national parks, is key to improve views on conservation practices from local people surrounding national parks.

Human disturbance represents a considerable challenge for the conservation of living organisms and has profound negative impacts globally. There is substantial variation in how species respond to human disturbance. The main aim of the present study is to examine how human activities influence the risk of brood parasitism and nest predation for three passerine species living in a highly fragmented landscape in Central Bangladesh. We use data collected from 518 nests over nine breeding seasons for three host species of the Asian koel (Eudynamys scolopaceus), a non-evicting brood parasite: i) long-tailed shrike (Lanius schach), ii) common myna (Acridotheres tristis) and iii) house crow (Corvus splendens). Two sources of disturbance were considered: i) houses and buildings inside the study area, the Jahangirnagar University campus and ii) the highly urbanized area outside the campus boundary. Additionally, we regarded iii) nest height, iv) number of days after egg laying until discovery (time unmonitored), v) distance to nearest fruit tree and vi) distance to nearest conspecific neighbor, as important habitat and nest characteristics. We found significantly lower predation risk with increasing time unmonitored in all three host species, thus more regular nest checks by fieldworkers lead to more predation. Moreover, we found that the distance to nearest fruit tree was the most important factor for the risk of parasitism. Although we could not find any significant impacts of distance to boundary or buildings on either predation or parasitism risk, we found a trend both in common myna and house crow that nesting success was higher closer to the boundary. Ultimately, increased monitoring time had the most consistent negative impact on nesting success among the host species. This study highlights the effect of disturbance caused by nest monitoring, exceeding the impact of permanent structures in an around the university campus i.e. boundary and buildings. © 2020 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Generation time determines the pace of key demographic and evolutionary processes. Quantified as the weighted mean age at reproduction, it can be studied as a trait that varies within and among populations and may evolve in response to ecological conditions. We combined quantitative genetic analyses with age- and density-dependent models to study generation time variation in a bird metapopulation. Generation time was heritable, and males had longer generation times compared with females. Individuals with longer generation times had a higher lifetime reproductive success but not a higher expected population growth rate. Density regulation acted on recruit production, suggesting that longer generation times should be favored when populations are closer to carrying capacity. Furthermore, generation times were shorter when populations were growing, and longer when populations were closer to equilibrium or declining. These results support classic theory predicting that density regulation is an important driver of the pace of life-history strategies.

Generation time determines the pace of key demographic and evolutionary processes. Quantified as the weighted mean age at reproduction, it can be studied as a trait that may evolve and change in response to ecological conditions. We combined quantitative genetic analyses of individual projection matrices with age- and density-dependent models to study generation time variation in a bird metapopulation. We found that males have longer generation times than females and that it is a heritable trait. Individuals with longer generation times contributed to population growth later in life, lived longer, produced fewer recruits per year, had greater lifetime reproductive success, but not necessarily a higher expected individual growth rate. As predicted by density-dependence theory, generation times were shorter when populations were growing, and longer when populations were closer to equilibrium or declining. These results support classic theory predicting that competitive regimes are key determinants of the pace of life-history strategies.

Inbreeding may increase the extinction risk of small populations. Yet, studies using modern genomic tools to investigate inbreeding depression in nature have been limited to single populations, and little is known about the dynamics of inbreeding depression in subdivided populations over time. Natural populations often experience different environmental conditions and differ in demographic history and genetic composition, characteristics that can affect the severity of inbreeding depression. We utilized extensive long-term data on more than 3,100 individuals from eight islands in an insular house sparrow metapopulation to examine the generality of inbreeding effects. Using genomic estimates of realized inbreeding, we discovered that inbred individuals had lower survival probabilities and produced fewer recruiting offspring than noninbred individuals. Inbreeding depression, measured as the decline in fitness-related traits per unit inbreeding, did not vary appreciably among populations or with time. As a consequence, populations with more resident inbreeding (due to their demographic history) paid a higher total fitness cost, evidenced by a larger variance in fitness explained by inbreeding within these populations. Our results are in contrast to the idea that effects of inbreeding generally depend on ecological factors and genetic differences among populations, and expand the understanding of inbreeding depression in natural subdivided populations.

Body size plays a key role in the ecology and evolution of all organisms. Therefore, quantifying the sources of morphological (co)variation, dependent and independent of body size, is of key importance when trying to understand and predict responses to selection. We combine structural equation modeling with quantitative genetics analyses to study morphological (co)variation in a meta-population of house sparrows (Passer domesticus). As expected, we found evidence of a latent variable "body size," causing genetic and environmental covariation between morphological traits. Estimates of conditional evolvability show that allometric relationships constrain the independent evolution of house sparrow morphology. We also found spatial differences in general body size and its allometric relationships. On islands where birds are more dispersive and mobile, individuals were smaller and had proportionally longer wings for their body size. Although on islands where sparrows are more sedentary and nest in dense colonies, individuals were larger and had proportionally longer tarsi for their body size. We corroborated these results using simulations and show that our analyses produce unbiased allometric slope estimates. This study highlights that in the short term allometric relationships may constrain phenotypic evolution, but that in the long term selection pressures can also shape allometric relationships.

Translocation of individuals from source populations to augment small populations facing risk of extinction is an important conservation tool. Here we examine sex-specific differences between resident and translocated house sparrows Passer domesticus in reproductive success and survival, and the contribution of translocated individuals to the growth of a local population. We found evidence for assortative mating based on origin revealed by fewer parentages between translocated males and resident females than expected, and the total number of fledglings produced by such pairs was lower. The reproductive success of translocated males was positively related to the size of the throat badge (a sexual ornament), such that only translocated males with a large badge size were as successful as resident males. However, offspring with parents of different origin had higher survival than offspring with parents of the same origin, which suggests hybrid vigour. The contribution of resident and translocated individuals to the stochastic component of the long-run growth rate of the population was similar; neither the mean individual contributions in fitness nor the demographic variance differed between the two groups. Thus, this experiment shows that translocated individuals may have a similar demographic influence on the growth of local populations as resident individuals. Still, the intermixing of translocated and resident individuals was low, and fitness differed according to origin in relation to individual differences in a sexually selected trait. In addition, hybrid vigour with respect to offspring recruitment seemed to partially decrease the negative fitness consequences of the assortative mating based on origin. This is the peer reviewed version of the following article: [Demographic influences of translocated individuals on a resident population of house sparrows], which has been published in final form at http://onlinelibrary.wiley.com/doi/10.1111/oik.04065/abstract. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving. Locked until 24.4.2018 due to copyright restrictions.

Human-carnivore encounters are common where humans and wild carnivores share the same landscape. The frequency of such encounters gives insight regarding carnivore density and might correlate with human-carnivore conflict incidences. We interviewed livestock owners in the eastern Serengeti ecosystem and recorded reported carnivore presence and relative abundance. We simultaneously conducted a carnivore survey to assess the potential variability of reported carnivore presence that was recorded during the surveys. The playback surveys attracted 9 lions (Panthera leo), 88 spotted hyenas (Crocuta crocuta) and 47 black-backed jackals (Canis mesomelas schmidti) to 12 call-in stations which were resurveyed three times (36 playbacks in total). Reported encounters with lions, leopards (Panthera pardus), cheetahs (Acinonyx jubatus), spotted hyenas, African wild dogs (Lycaon pictus) and jackals were higher closer to the Serengeti National Park (SNP). Data from carnivore surveys were positively correlated with what people reported in questionnaires. These results indicate that local reports of encounters with wild carnivores may act as an important indicator of carnivore presence. Combining observational data through surveys with data reported by local people in areas where humans and wild carnivores coexist may improve existing data on carnivore abundance and distribution in such areas. This is an open access article distributed under the terms of the Creative Commons CC-BY license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Conservation crimes are posing serious threats to wildlife species and biodiversity of the Rungwa-Kizigo-Muhesi-Game-Reserves. Devising effective strategies to reduce risks to as low as reasonably possible of these crimes, calls for adequate information on factors driving people to commit these crimes. Data for this study were obtained from 315 respondents in 20 villages and 316 people who were arrested for committing different conservation crimes in the Rungwa-Kizigo-Muhesi-Game-Reserves. Our results suggest that arrested respondents were typically young adult males with limited alternative sources of income and owning virtually no livestock or land. There were heterogeneous drivers for each type of conservation crime. To address the challenge of conservation crimes in the Reserve, we recommend, among other strategies, the establishment of effective conservation education programmes, strengthening law enforcements as a deterrence method, and development of entrepreneurship skills to enhance employment. Keywords: Bushmeat poaching, Conservation criminology, Elephant, Illegal timber logging, Wildlife poaching

Understanding the genetic architecture of quantitative traits can provide insights into the mechanisms driving phenotypic evolution. Bill morphology is an ecologically important and phenotypically variable trait, which is highly heritable and closely linked to individual fitness. Thus, bill morphology traits are suitable candidates for gene mapping analyses. Previous studies have revealed several genes that may influence bill morphology, but the similarity of gene and allele effects between species and populations is unknown. Here, we develop a custom 200K SNP array and use it to examine the genetic basis of bill morphology in 1857 house sparrow individuals from a large‐scale, island metapopulation off the coast of Northern Norway. We found high genomic heritabilities for bill depth and length, which were comparable with previous pedigree estimates. Candidate gene and genomewide association analyses yielded six significant loci, four of which have previously been associated with craniofacial development. Three of these loci are involved in bone morphogenic protein (BMP) signalling, suggesting a role for BMP genes in regulating bill morphology. However, these loci individually explain a small amount of variance. In combination with results from genome partitioning analyses, this indicates that bill morphology is a polygenic trait. Any studies of eco‐evolutionary processes in bill morphology are therefore dependent on methods that can accommodate polygenic inheritance of the phenotype and molecular‐scale evolution of genetic architecture. Publisher embargo applies until July 24, 2019

There is at present considerable variation in the level of antiparasite defences among different host species of avian brood parasites, but in many potential hosts some individuals reject poorly matching parasite eggs. Here we present unique absence of egg discrimination behaviour backed up by a lack of egg recognition abilities in a suitable common cuckoo Cuculus canorus host, the skylark Alauda arvensis. Skylarks did not show any clear rejection response to experimentally added highly non-mimetic foreign eggs in any behavioural context, even before they had started laying or when the whole clutch was exchanged with foreign eggs. This absence of antiparasite defence can be explained by the breeding habitat of larks consisting of largely treeless open landscapes where cuckoos have little access to the nests, thereby eroding the possibility of coevolutionary interactions. Our results are strikingly consistent with the spatial habitat structure hypothesis proposed to explain the occurrence and extent of avian host-parasite co-adaptation.

The brood parasitic common cuckoo Cuculus canorus consists of gentes, which typically parasitize only a single host species whose eggs they often mimic. Where multiple cuckoo gentes co-exist in sympatry, we may expect variable but generally poorer mimicry because of host switches or inter-gens gene flow via males if these also contribute to egg phenotypes. Here, we investigated egg trait differentiation and mimicry in three cuckoo gentes parasitizing great reed warblers Acrocephalus arundinaceus, marsh warblers Acrocephalus palustris and corn buntings Miliaria calandra breeding in close sympatry in partially overlapping habitat types. The three cuckoo gentes showed a remarkable degree of mimicry to their three host species in some but not all egg features, including egg size, a hitherto largely ignored feature of egg mimicry. Egg phenotype matching for both background and spot colours as well as for egg size has been maintained in close sympatry despite the possibility for gene flow.

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. © 2012 Fossøy et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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. © 2012 Antonov et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.