Bart Kempenaers

Max Planck Institute for Ornithology · Department of Behavioural Ecology and Evolutionary Genetics

Mate guarding is typically considered a male strategy to protect paternity. However, under some circumstances, females might also benefit from guarding their mate. Female mate guarding might be particularly important in socially polyandrous species in which females compete for access to care-giving males. Because males also benefit from being near...

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....

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 s...

In the Anthropocene, the era when the imprint of humans on nature is pervasive across the planet, it is of utmost importance to understand human relationships with other species. The aesthetics of nature, and of species, is one of the values that plays a role in shaping human-nature relationships. Birds are ubiquitous across the world. The beauty o...

Animal coloration is one of the most conspicuous aspects of human-perceived organismal diversity, yet also one of the least understood. In particular, explaining why species have specific colors (e.g., blue vs. red) has proven elusive. Here, we quantify for nearly all bird species, the proportion of the body covered by each of 12 human-visible colo...

The ruff sandpiper Calidris pugnax is a Palearctic lekking shorebird with three genetic morphs determined by an autosomal inversion. Male morphs differ strikingly in body size, ornaments, endocrinology and mating behavior. Aggressive Independents represent the ancestral haplotype, semi-cooperative Satellites and female-mimicking Faeders the inverte...

Across birds, male age is the most consistent predictor of extra-pair siring success, yet little is known about age effects on paternity over the lifetime of individuals. Here, we use data from a 13-year study of a population of blue tits (Cyanistes caeruleus) to investigate how extra-pair siring success changes with age within individuals. Our res...

Artificial light at night significantly alters the predictability of the natural light cycles that most animals use as an essential Zeitgeber for daily activity. Direct light has well-documented local impacts on activity patterns of diurnal and nocturnal organisms. However, artificial light at night also contributes to an indirect illumination of t...

All animals host a microbial community within and on their reproductive organs, known as the reproductive microbiome. In free‐living birds, studies on the sexual transmission of bacteria have typically focused on a few pathogens instead of the bacterial community as a whole, despite a potential link to reproductive function. Theory predicts higher...

The germline-restricted chromosome (GRC) is likely present in all songbird species, but differs widely in size and gene content. This extra chromosome has been described as either a microchromosome with only limited basic gene content or a macrochromosome with enriched gene functions related to female gonad and embryo development. Here, we assemble...

The study of chromosomal inversion polymorphisms has received much recent attention, particularly in cases where inversions have drastic effects on phenotypes and fitness (e.g. lethality of homozygotes). Less attention has been paid to the question of the maintenance of inversion polymorphisms that show only weak effects. Here, we study the mainten...

Urbanization is transforming ecosystems at a global scale and at an increasing rate, and its profound consequences for wildlife have been well documented. Understanding how animals thrive in the urban environment and how this environment affects (co-)evolutionary processes remains an important challenge. Urban environments can provide resources suc...

Differences in the strength of sexual selection between males and females can lead to sexual dimorphism. Extra-pair paternity (EPP) can increase the variance in male reproductive success and hence the opportunity for sexual selection. Previous research on birds suggests that EPP drives the evolution of dimorphism in plumage colour and in body size....

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 ex...

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 la...

The adaptive value of mate retention has been studied in several socially monogamous birds, but evidence of reproductive benefits for short‐lived species is inconclusive. Most studies come from northern latitudes, but more research on tropical birds is needed, as these species typically show higher survival rates and longer pair bonds than those fr...

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 sy...

Most vertebrates, including all mammals and birds, are sexually reproducing organisms. The implication of sexual reproduction is that an adult bird exists because a male transferred sperm to the reproductive tract of a female during copulation, one of the sperm fertilized an egg, and the ensuing embryo developed into an individual male or female th...

Identifying the environmental drivers of variation in fitness‐related traits is a central objective in ecology and evolutionary biology. Temporal fluctuations of these environmental drivers are often synchronized at large spatial scales. Yet, whether synchronous environmental conditions can generate spatial synchrony in fitness‐related trait values...

Brood parasitic cuckoos and their hosts serve as model systems for studying host–parasite coevolution. Egg-rejection behavior constitutes an effective defense against brood parasitism, but some host species show phenotypic plasticity in egg-rejection behavior. Direct exposure to a cuckoo near the nest can increase egg-rejection likelihood, and long...

We describe a new taxon of terrestrial bird of the genus Aphrastura (rayaditos) inhabiting the Diego Ramírez Archipelago, the southernmost point of the American continent. This archipelago is geographically isolated and lacks terrestrial mammalian predators as well as woody plants, providing a contrasted habitat to the forests inhabited by the othe...

Many birds use carotenoids to colour their plumage yellow to red. Because birds cannot synthesise carotenoids, they need to obtain these pigments from food, although some species metabolise dietary carotenoids (which are often yellow) into derived carotenoids (often red). Here, we study the occurrence of yellow and red carotenoid‐based plumage colo...

Understanding the targets of selection associated with changes in behavioral traits represents an important challenge of current evolutionary research. Owls (Strigiformes) are a diverse group of birds, most of which are considered nocturnal raptors. However, a few owl species independently adopted a diurnal lifestyle in their recent evolutionary hi...

In species with biparental care, coordination of parental behaviour between pair members increases reproductive success. Coordination is difficult if opportunities to communicate are scarce, which might have led to the evolution of elaborate nest relief rituals in species facing a low predation risk. However, whether such conspicuous rituals also e...

The phenology of many species shows strong sensitivity to climate change; however, with few large scale intra-specific studies it is unclear how such sensitivity varies over a species’ range. We document large intra-specific variation in phenological sensitivity to temperature using laying date information from 67 populations of two co-familial Eur...

Sex-bias in breeding dispersal is considered the norm in many taxa, and the magnitude and direction of such sex-bias is expected to correlate with the social mating system. We used local return rates in shorebirds as an index of breeding site fidelity, and hence as an estimate of the propensity for breeding dispersal, and tested whether variation i...

Culturally transmitted communication signals – such as human language or bird song – can change over time through cultural drift, and the resulting dialects may consequently enhance the separation of populations. However, the emergence of song dialects has been considered unlikely when songs are highly individual-specific, as in the zebra finch (Ta...

Chromosomal inversions frequently underlie major phenotypic variation maintained by divergent selection within and between sexes. Here we examine whether and how intralocus conflicts contribute to balancing selection stabilizing an autosomal inversion polymorphism in the ruff Calidris pugnax. In this lekking shorebird, three male mating morphs (Ind...

Investigating whether mating patterns are biased in relation to kinship in isolated populations can provide a better understanding of the occurrence of inbreeding avoidance mechanisms in wild populations. Here, we report on the genetic relatedness (r) among breeding pairs in a relict population of Thorn‐tailed Rayadito (Aphrastura spinicauda) in no...

Significance Most if not all songbirds possess a germline-restricted chromosome (GRC) which is believed to be exclusively maternally inherited. However, we show that, in the zebra finch, the GRC can also be paternally inherited and that the potential for paternal inheritance may differ between families. We further show that the genetic diversity of...

Understanding the genomic landscape of adaptation is central to the understanding of microevolution in wild populations. Genomic targets of selection and the underlying genomic mechanisms of adaptation can be elucidated by genome-wide scans for past selective sweeps or by scans for direct fitness associations. We sequenced and assembled 150 haploty...

In socially monogamous species, extrapair paternity typically results from extrapair copulations, but it can also be due to rapid mate switching. Oring, Fleischer, Reed, and Marsden (1992, Nature, 359 (6396), 631–633) proposed a mechanism to explain the occurrence of extrapair paternity in sequentially polyandrous species: sperm stored by females f...

Migratory birds undertake long and challenging journeys that have selected for a suite of adaptations from sensory mechanisms that facilitate orientation to extreme feats of endurance that push physiological limits. Recent work on two distantly related species revealed that migrating individuals increase their flight altitude dramatically during th...

Female mate choice is thought to be responsible for the evolution of many extravagant male ornaments and displays, but the costs of being too selective may hinder the evolution of choosiness. Selection against choosiness may be particularly strong in socially monogamous mating systems, because females may end up without a partner and forego reprodu...

Leaving the nest is a key transition in the life of altricial birds, whereby fledging decisions should depend on multiple factors, including the risk of predation. High postfledging predation risk may favour fledging at a more advanced stage of development, if more developed fledglings are better at escaping predation, or together with others. Whil...

The transition from nestling to fledgling is a key moment in the development of altricial birds. Mortality immediately after fledging is typically high and selection should favour fledging strategies that maximize the chance of survival. While several studies have examined the influence of ecological conditions or nestling development on the timing...

Individual-specific mate preferences are thought to be widespread, but they are still poorly understood in terms of mechanisms and function. Earlier work on a songbird (the zebra finch, Taeniopygia guttata) showed predominantly individual-specific mate preferences and signs of behavioural incompatibility with certain partners. However, the phenotyp...

In many socially monogamous bird species, females produce offspring sired by males other than their social partner. A large body of research has aimed to elucidate the evolutionary causes and consequences of such extra‐pair paternity, but relatively little is known about the underlying behaviour. The number of sperm on the egg's perivitelline layer...

The propulsion of sperm cells via movement of the flagellum is of vital importance for successful fertilization. While the exact mechanism of energy production for this movement varies between species, in avian species energy is thought to come predominantly from the mitochondria located in the sperm midpiece. Larger midpieces may contain more mito...

Biparental care requires coordination between parents. Such coordination might prove difficult if opportunities to communicate are scarce, which might have led to the evolution of elaborate and noisy nest relief rituals in species facing a low risk of predation. However, whether such conspicuous rituals also evolved in species that avoid predation...

Chromosomal inversions frequently underlie major phenotypic variation maintained by divergent selection within and between sexes. Here we examine whether and how intralocus conflicts contribute to balancing selection stabilizing an autosomal inversion polymorphism in the ruff Calidris pugnax. In this lekking shorebird, three male mating morphs (Ind...

Extra‐pair paternity (EPP) is a key aspect of the mating behaviour of birds and its frequency varies widely among populations and species. Several hypotheses predict patterns of geographical variation in the occurrence and frequency of EPP, but a global‐scale study on variation in this trait is still lacking. We collected data on EPP from 663 popul...

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 o...

Individuals differ in the quantity and quality of their associations with conspecifics. The resulting variation in the positions that individuals occupy within their social environment can affect several aspects of life history, including reproduction. While research increasingly shows how social factors can predict dyadic mating patterns (who will...

Female mate choice is thought to be responsible for the evolution of many extravagant male ornaments and displays, but the costs of being too selective may hinder the evolution of choosiness. Selection against choosiness should be strongest in socially monogamous mating systems, because females may end up without a partner and forego reproduction,...

In altricial birds, leaving the nest is a key life history transition associated with a high risk of mortality. Studies of numerous species have shown that young typically fledge early in the day, and it is often asserted that early fledging is important for survival; however, evidence for this hypothesis is limited. We used an automated monitoring...

Classic explanations for polygyny consider habitat, genetic makeup and paternal care, but little attention has been paid to social inertia. We studied facultative social polygyny in a population of Blue Tits Cyanistes caeruleus with a low rate of polygyny (3% of males across 12 years). Occurrence of polygyny was best predicted by social turnover af...

Most birds are socially monogamous with both parents providing offspring care, but sometimes individuals are observed to provision at a nest that is not their own. One possible explanation for this behaviour is that it is a fitness maximising strategy by males who have copulated with the female and hence potentially sired extra‐pair offspring in th...

Many shorebird species are rapidly declining (Piersma et al. 2016; Munro 2017; Studds et al. 2017), but it is not always clear why. Deteriorating and disappearing habitat, e.g. due to intensive agriculture (Donal et al. 2001; Kentie et al. 2013; Kentie et al. 2018), river regulation (Nebel et al. 2008) or mudflat reclamation (Ma et al. 2014; Larson...

Culturally transmitted communication signals, such as human language or bird song, can change over time through a process of cultural drift, and may consequently enhance the separation of populations, potentially leading to reproductive isolation. Local song dialects have been identified in bird species with relatively simple songs where individual...

All songbirds have one special accessory chromosome, the so-called germline-restricted chromosome (GRC), which is only present in germline cells and absent from all somatic tissues. Earlier work on the zebra finch (Taeniopygia guttata castanotis) showed that the GRC is inherited only through the female line - like mitochondrial DNA - and is elimina...

Animals use acoustic signals for communication, implying that the properties of these signals can be under strong selection. The acoustic adaptation hypothesis predicts that species in dense habitats emit lower-frequency sounds than those in open areas because low-frequency sounds propagate further in dense vegetation than high-frequency sounds. Si...

Heterozygosity-fitness correlations (HFCs) have been used to monitor the effects of inbreeding in threatened populations. HFCs can also be useful to investigate the potential effects of inbreeding in isolated relict populations of long-term persistence, and to better understand the role of inbreeding and outbreeding as drivers of changes in genetic...

Significance Many ecological and evolutionary processes strongly depend on the way natural selection varies over time. However, a gap remains when trying to connect theoretical predictions to empirical work on this question: Most theory assumes that adaptation involves tracking a moving optimum phenotype through time, but this is seldom estimated e...

How species will adapt to future climate change is a key question in modern biology. One way to predict such adaptation is to draw from our knowledge of current spatial patterns of phenotypic variation. These are often summarised by different ecogeographical rules that describe how environmental gradients predict geographic variation in form and fu...

Many studies investigated variation in the frequency of extrapair paternity (EPP) among individuals. However, our understanding of within-individual variation in EPP remains limited. Here, we comprehensively investigate variation in EPP at the within-individual level in a population of blue tits (Cyanistes caeruleus). Our study is based on parentag...

1. The integration and synthesis of the data in different areas of science is drastically slowed and hindered by a lack of standards and networking programmes. Long‐term studies of individually marked animals are not an exception. These studies are especially important as instrumental for understanding evolutionary and ecological processes in the w...

Meiotic drivers have been proposed as a potent evolutionary force underlying genetic and phenotypic variation, genome structure, and also speciation. Due to their strong selective advantage, they are expected to rapidly spread through a population despite potentially detrimental effects on organismal fitness. Once fixed, autosomal drivers are crypt...

Studies on extrapair paternity (EPP) are key to understanding the ecological and evolutionary drivers of variation in avian mating strategies, but information is currently lacking for most tropical and subtropical taxa. We describe the occurrence of EPP in two populations of a South American socially monogamous bird, the Thorn-tailed Rayadito, base...

The evolutionary consequences of individual genetic diversity are frequently studied by assessing heterozygosity‐fitness correlations (HFCs). The prevalence of positive and negative HFCs and the predominance of general versus local effects in wild populations are far from understood, partly because comprehensive studies testing for both inbreeding...

1) Moulting strategies in birds have evolved to avoid overlap with, or prepare for, other demanding parts of the annual cycle, such as reproduction or migration. When moulting for the first time after leaving the nest, young birds replace their typically poor-quality plumage during the post-juvenile moult. The extent of this moult varies between sp...

The phenology of many species shows strong sensitivity to climate change; however, with few large scale intra-specific studies it is unclear how such sensitivity varies over a species' range. We document large intra-specific variation in phenological sensitivity to temperature using laying date information from 67 populations of two European songbi...

Psittaciformes (parrots, cockatoos and lorikeets) comprise one of the most colourful clades of birds. Their unique pigments and safe cavity nesting habits are two potential explanations for their colourful character. However, plumage colour varies substantially between parrot species and sometimes also between males and females of the same species....

The integration and synthesis of the data in different areas of science is drastically slowed and hindered by a lack of standards and networking programmes. Long-term studies of individually marked animals are not an exception. These studies are especially important as instrumental for understanding evolutionary and ecological processes in the wild...

Owls (Strigiformes) evolved specific adaptations to their nocturnal predatory lifestyle, such as asymmetrical ears, a facial disc, and a feather structure allowing silent flight. Owls also share some traits with diurnal raptors and other nocturnal birds, such as cryptic plumage patterns, reversed sexual size dimorphism and acute vision and hearing....

Some species show high rates of reproductive failure, which is puzzling because natural selection works against such failure in every generation. Hatching failure is common in both captive and wild zebra finches (Taeniopygia guttata), yet little is known about its proximate causes. Here we analyze data on reproductive performance (the fate of >23,0...

How genetic polymorphisms are maintained in a population is a key question in evolutionary ecology. Previous work on a plumage colour polymorphism in the common buzzard Buteo buteo suggested heterozygote advantage as the mechanism maintaining the co‐existence of three morphs (light, intermediate and dark). We took advantage of 20 years of life hist...

Many animals use acoustic signals for communication, implying that the properties of these signals can be under strong selection. The acoustic adaptation hypothesis predicts that species living in dense habitats emit lower-frequency sounds than those in open areas, because low-frequency sounds generally propagate further in denser vegetation. Signa...

Understanding the population genetic consequences of habitat heterogeneity requires assessing whether patterns of gene flow correspond to landscape configuration. Studies of the genetic structure of populations are still scarce for Neotropical forest birds. We assessed range-wide genetic structure and contemporary gene flow in the thorn-tailed raya...