Search

Space-based tracking technology using low-cost miniature tags is now delivering data on fine-scale animal movement at near-global scale. Linked with remotely sensed environmental data, this offers a biological lens on habitat integrity and connectivity for conservation and human health; a global network of animal sentinels of environmental change. Publisher PDF

In species with separate sexes, females and males often differ in their morphology, physiology and behaviour. Such sex-specific traits are functionally linked to variation in reproductive competition, mate choice and parental care, which have all been linked to sex roles. At the 150th anniversary of Darwin's theory on sexual selection, the question of why patterns of sex roles vary within and across species remains a key topic in behavioural and evolutionary ecology. New theoretical, experimental and comparative evidence suggests that variation in the adult sex ratio (ASR) is a key driver of variation in sex roles. Here, we first define and discuss the historical emergence of the sex role concept, including recent criticisms and rebuttals. Second, we review the various sex ratios with a focus on ASR, and explore its theoretical links to sex roles. Third, we explore the causes, and especially the consequences, of biased ASRs, focusing on the results of correlational and experimental studies of the effect of ASR variation on mate choice, sexual conflict, parental care and mating systems, social behaviour, hormone physiology and fitness. We present evidence that animals in diverse societies are sensitive to variation in local ASR, even on short timescales, and propose explanations for conflicting results. We conclude with an overview of open questions in this field integrating demography, life history and behaviour. © 2022 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.

Traffic noise leads to vocal learning deficits in young songbirds, the primary model for speech development in humans., Noise pollution has been linked to learning and language deficits in children, but the causal mechanisms connecting noise to cognitive deficiencies remain unclear because experimental models are lacking. Here, we investigated the effects of noise on birdsong learning, the primary animal model for vocal learning and speech development in humans. We found that traffic noise exposure retarded vocal development and led to learning inaccuracies. In addition, noise suppressed immune function during the sensitive learning period, indicating that it is a potent stressor for birds, which is likely to compromise their cognitive functions. Our results provide important insights into the consequences of noise pollution and pave the way for future studies using birdsong as an experimental model for the investigation of noise-induced learning impairments.

Background The connection between testosterone and territoriality in free-living songbirds has been well studied in a reproductive context, but less so outside the breeding season. To assess the effects of seasonal androgenic action on territorial behavior, we analyzed vocal and non-vocal territorial behavior in response to simulated territorial intrusions (STIs) during three life-cycle stages in free-living male black redstarts: breeding, molt and nonbreeding. Concurrently, we measured changes in circulating testosterone levels, as well as the mRNA expression of androgen and estrogen receptors and aromatase in the preoptic, hypothalamic and song control brain areas that are associated with social and vocal behaviors. Results Territorial behavior and estrogen receptor expression in hypothalamic areas did not differ between stages. But plasma testosterone was higher during breeding than during the other stages, similar to androgen receptor and aromatase expression in the preoptic area. The expression of androgen receptors in the song control nucleus HVC was lower during molt when birds do not sing or sing rarely, but similar between the breeding and the nonbreeding stage. Nevertheless, some song spectral features and the song repertoire differed between breeding and nonbreeding. Territorial behavior and song rate correlated with the expression of steroid receptors in hypothalamic areas, and in the song control nucleus lMAN. Conclusions Our results demonstrate seasonal modulation of song, circulating testosterone levels, and brain sensitivity to androgens, but a year-round persistency of territorial behavior and estrogen receptor expression in all life-cycle stages. This suggests that seasonal variations in circulating testosterone concentrations and brain sensitivity to androgens is widely uncoupled from territorial behavior and song activity but might still affect song pattern. Our study contributes to the understanding of the complex comparative neuroendocrinology of song birds in the wild.

Most studies on sexual size dimorphism address proximate and functional questions related to adults, but sexual size dimorphism usually develops during ontogeny and developmental trajectories of sexual size dimorphism are poorly understood. We studied three bird species with variation in adult sexual size dimorphism: black coucals (females 69% heavier than males), white‐browed coucals (females 13% heavier than males) and ruffs (males 70% heavier than females). Using a flexible Bayesian generalized additive model framework (GAMM), we examined when and how sexual size dimorphism developed in body mass, tarsus length and bill length from hatching until fledging. In ruffs, we additionally examined the development of intrasexual size variation among three morphs (Independents, Satellites and Faeders), which creates another level of variation in adult size of males and females. We found that 27–100% of the adult inter‐ and intrasexual size variation developed until fledging although none of the species completed growth during the observational period. In general, the larger sex/morph grew more quickly and reached its maximal absolute growth rate later than the smaller sex/morph. However, when the daily increase in body mass was modelled as a proportion, growth patterns were synchronized between and within sexes. Growth broadly followed sigmoidal asymptotic models, however only with the flexible GAMM approach, residual distributions were homogeneous over the entire observation periods. These results provide a platform for future studies to relate variation in growth to selective pressures and proximate mechanisms in these three species, and they highlight the advantage of using a flexible model approach for examining growth variation during ontogeny.

Artificial light at night could have widespread and detrimental impacts on sleep. To reduce disruptive effects of artificial light on sleep in humans, most smartphones and computers now have software that reduces blue light emissions at night. Little is known about whether reducing blue light emissions from city lights could also benefit urban wildlife. We investigated the effects of blue-rich (white) and blue-reduced (amber) LED streetlights on accelerometry-defined rest, electrophysiologically-identified sleep, and plasma melatonin in a diurnal bird, the black swan (Cygnus atratus). Urban swans were exposed to 20 full nights of each lighting type in an outdoor, naturalistic environment. Contrary to our predictions, we found that night-time rest was similar during exposure to amber and white lights but decreased under amber lights compared with dark conditions. By recording brain activity in a subset of swans, we also demonstrated that resting birds were almost always asleep, so amber light also reduced sleep at night. We found no effect of light treatment on total (24 h) daily rest or plasma melatonin. Our study provides the first electrophysiologically-verified evidence for effects of streetlights on sleep in an urban animal, and furthermore suggests that reducing blue wavelengths of light might not mitigate these effects., Frontiers in Ecology and Evolution, 8, ISSN:2296-701X

Lay Summary Human-induced environmental change can disturb animals, reducing individual and population health. We found no general relationship between urbanization and hormones associated with stressors across birds and reptiles, suggesting hormonal responses to disturbance vary across species and even individuals. Information beyond hormone levels is needed to inform conservation efforts., Rates of human-induced environmental change continue increasing with human population size, potentially altering animal physiology and negatively affecting wildlife. Researchers often use glucocorticoid concentrations (hormones that can be associated with stressors) to gauge the impact of anthropogenic factors (e.g. urbanization, noise and light pollution). Yet, no general relationships between human-induced environmental change and glucocorticoids have emerged. Given the number of recent studies reporting baseline and stress-induced corticosterone (the primary glucocorticoid in birds and reptiles) concentrations worldwide, it is now possible to conduct large-scale comparative analyses to test for general associations between disturbance and baseline and stress-induced corticosterone across species. Additionally, we can control for factors that may influence context, such as life history stage, environmental conditions and urban adaptability of a species. Here, we take a phylogenetically informed approach and use data from HormoneBase to test if baseline and stress-induced corticosterone are valid indicators of exposure to human footprint index, human population density, anthropogenic noise and artificial light at night in birds and reptiles. Our results show a negative relationship between anthropogenic noise and baseline corticosterone for birds characterized as urban avoiders. While our results potentially indicate that urban avoiders are more sensitive to noise than other species, overall our study suggests that the relationship between human-induced environmental change and corticosterone varies across species and contexts; we found no general relationship between human impacts and baseline and stress-induced corticosterone in birds, nor baseline corticosterone in reptiles. Therefore, it should not be assumed that high or low levels of exposure to human-induced environmental change are associated with high or low corticosterone levels, respectively, or that closely related species, or even individuals, will respond similarly. Moving forward, measuring alternative physiological traits alongside reproductive success, health and survival may provide context to better understand the potential negative effects of human-induced environmental change.

The 24 h geophysical light-dark cycle is the main organizer of daily rhythms, scheduling physiology and behavior. This cycle attenuates greatly during the continuous light of summer at polar latitudes, resulting in species-specific and even individual-specific patterns of behavioral rhythmicity, but the physiological mechanisms underlying this variation are poorly understood. To address this knowledge gap and to better understand the roles of the hormones melatonin and corticosterone in rhythmic behavior during polar day, we exploited the behavior of thick-billed murres (Uria lomvia), a charadriiform seabird with sexually opposite ('antiphase') activity-rhythms on a 24 h cycle during the continuous light of polar summer. Melatonin concentration in the plasma of inactive males was unexpectedly high around midday and subsequently fell during a sudden decrease in light intensity as the colony became shaded. Corticosterone concentration in plasma did not vary with time of day or activity in either sex. While the reasons for these unusual patterns remain unclear, we propose that a flexible melatonin response and little diel variation of corticosterone may be adaptive in thick-billed murres, and perhaps other polar birds and mammals, by stabilizing glucocorticoids' role of modulating energy storage and mobilization across the diel cycle and facilitating the appropriate reaction to unexpected stimuli experienced across the diel cycle while attending the colony.

The impact of human activity on the acoustic environment is overwhelming, with anthropogenic noise reaching even remote areas of the planet. The World Health Organization has identified noise pollution as one of the leading environmental health risks in humans, and it has been linked to a myriad of short- and long-term health effects in exposed individuals. However, less is known about the health effects of anthropogenic noise exposure on animals. We investigated long- and short-term effects of traffic noise on zebra finches breeding in small communal aviaries, using a repeated measures design. Birds bred in both noise and no-noise conditions, and we measured baseline plasma glucocorticoid levels before, during and after breeding. In addition, we assayed immune function, measured reproductive success and offspring growth and compared rates of extra-pair paternity of breeding adults. Breeding birds had significantly lower baseline plasma corticosterone levels when exposed to traffic noise than when they were not exposed to noise playback. In addition, the nestlings reared during noise exposure were lighter than nestlings of the same parents when breeding in control conditions. Our results suggest that traffic noise poses a more severe hurdle to birds at more vulnerable stages of their life history, such as during reproductive events and ontogeny. While chronic exposure to traffic noise in our birds did not, by itself, prove to be a sufficient stressor to cause acute effects on health or reproductive success in exposed individuals, it did result in disruptions to normal glucocorticoid profiles and delayed offspring growth. However, animals living in urban habitats are exposed to a multitude of anthropogenic disturbances, and it is likely that even species that appear to be thriving in noisy environments may suffer cumulative effects of these multiple disturbances that may together impact their fitness in urban environments.

Providing parental care often reduces additional mating opportunities. Paternal care becomes easier to understand if trade-offs between mating and caring remain mild. The black coucal Centropus grillii combines male-only parental care with 50% of all broods containing young sired by another male. To understand how much caring for offspring reduces a male's chance to sire additional young in other males' nests, we matched the production of extra-pair young in each nest with the periods during which potential extra-pair sires were either caring for offspring themselves or when they had no own offspring to care for. We found that males which cared for a clutch were not fully excluded from the pool of competitors for siring young in other males' nests. Instead, the relative siring success showed a temporary dip. Males were approximately 17% less likely to sire young in other males' nests while they were incubating, about 48% less likely to do so while feeding nestlings, followed by 26% when feeding fledglings, compared to the success of males that currently did not care for offspring. These results suggest that real-life care situations by males may involve trade-off structures that differ from, and are less strict than those frequently employed in theoretical considerations of operational sex ratios, sex roles and parenting decisions.

Hormones are central regulators of organismal function and flexibility that mediate a diversity of phenotypic traits from early development through senescence. Yet despite these important roles, basic questions about how and why hormone systems vary within and across species remain unanswered. Here we describe HormoneBase, a database of circulating steroid hormone levels and their variation across vertebrates. This database aims to provide all available data on the mean, variation, and range of plasma glucocorticoids (both baseline and stress-induced) and androgens in free-living and un-manipulated adult vertebrates. HormoneBase ( www.HormoneBase.org ) currently includes >6,580 entries from 476 species, reported in 648 publications from 1967 to 2015, and unpublished datasets. Entries are associated with data on the species and population, sex, year and month of study, geographic coordinates, life history stage, method and latency of hormone sampling, and analysis technique. This novel resource could be used for analyses of the function and evolution of hormone systems, and the relationships between hormonal variation and a variety of processes including phenotypic variation, fitness, and species distributions. Machine-accessible metadata file describing the reported data (ISA-Tab format)

Environmental cues, such as photoperiod, regulate the timing of major life-history events like breeding through direct neuroendocrine control. Less known is how supplementary environmental cues (e.g., nest sites, food availability) interact to influence key hormones and behaviors involved in reproduction, specifically in migratory species with gonadal recrudescence largely occurring at breeding sites. We investigated the behavioral and physiological responses of male European starlings to the sequential addition of nest boxes and nesting material, green herbs, and female conspecifics and how these responses depend on the availability of certain antioxidants (anthocyanins) in the diet. As expected, cloacal protuberance volume and plasma testosterone of males generally increased with photoperiod. More notably, testosterone levels peaked in males fed the high antioxidant diet when both nest box and herbal cues were present, while males fed the low antioxidant diet showed no or only a muted testosterone response to the sequential addition of these environmental cues; thus our results are in agreement with the oxidation handicap hypothesis. Males fed the high antioxidant diet maintained a constant frequency of breeding behaviors over time, whereas those fed the low antioxidant diet decreased breeding behaviors as environmental cues were sequentially added. Overall, sequential addition of the environmental cues modulated physiological and behavioral measures of reproductive condition, and dietary antioxidants were shown to be a key factor in affecting the degree of response to each of these cues. Our results highlight the importance of supplementary environmental cues and key resources such as dietary antioxidants in enhancing breeding condition of males, which conceivably aid in attraction of high quality females and reproductive success.

During migration, birds need to optimize their time and/or energy management, especially during stop-overs. Previous studies with caged birds under controlled condition support the notion that departure decisions are condition-dependent, but they did not take into account the availability or the actual intake of food. In the study reported here we investigated whether food intake interacts with condition in influencing migratory disposition in temporarily caged wild migrants. We conducted the study on Garden Warblers (Sylvia borin) and European Robins (Erithacus rubecula) at a spring stop-over site that is reached after a long non-stop flight over the Mediterranean Sea. The birds were held in cages during the day and the first night following capture, and their locomotor activity, food intake, and body mass change were recorded. In both species, food intake was positively correlated with body mass change. Body condition was positively correlated with migratory restlessness (Zugunruhe) in both species and was also negatively related to diurnal activity in Garden Warblers. Food intake was negatively correlated with diurnal activity in Garden Warblers. When only birds with low fat scores were considered, we found a positive effect of food intake on Zugunruhe in Garden Warblers. In general, European Robins consumed less food than Garden Warblers and no significant effects on Zugunruhe were found. Our results show that food intake interacts with condition in affecting migratory behavior during stop-overs. In particular, food intake may induce Garden Warblers with low fat stores at arrival to leave a stop-over site earlier. The large individual variability in food consumption supports the notion that birds vary in their capacity to refuel soon after arrival at a stop-over site. Overall, our work suggests that the decision to stay at the stop-over site or to resume migration depends on the interaction of condition, refueling capacity and rate, and migratory strategy.

During migration, birds spend more than 80 % of the time at stopover sites to rest and refuel before and after crossing ecological barriers such as deserts or seas. Since stopover has intrinsic costs in terms of energy and time, birds should try to minimize its duration, which is dependent on the combined effects of environmental factors, endogenous programmes, and physiological conditions. Previous studies on long-distance migrants caught after crossing an ecological barrier have indicated that body condition strongly influences the decision whether to prolong stopover or resume migration, with lean birds staying longer than fat birds. In short-distance migrants, evidence is still scarce regarding a determinant role for physiological condition in stopover behaviour after crossing an ecological barrier. Here, we studied whether migratory restlessness (Zugunruhe) at a stopover site is dependent on physiological condition in three European short-distance migratory passerines: black redstarts, European robins, and European stonechats. In all species, an integrated measure of condition based on body mass, amount of subcutaneous fat, and thickness of pectoral muscles predicted the intensity of Zugunruhe. Overall, our results confirmed the importance of energy reserves in determining stopover duration, illustrating similar stopover strategies in short- and long-distance migrants.

Vocal signals such as calls play a crucial role for survival and successful reproduction, especially in group-living animals. However, call interactions and call dynamics within groups remain largely unexplored because their relation to relevant contexts or life-history stages could not be studied with individual-level resolution. Using on-bird microphone transmitters, we recorded the vocalisations of individual zebra finches (Taeniopygia guttata) behaving freely in social groups, while females and males previously unknown to each other passed through different stages of the breeding cycle. As birds formed pairs and shifted their reproductive status, their call repertoire composition changed. The recordings revealed that calls occurred non-randomly in fine-tuned vocal interactions and decreased within groups while pair-specific patterns emerged. Call-type combinations of vocal interactions changed within pairs and were associated with successful egg-laying, highlighting a potential fitness relevance of calling dynamics in communication systems. DOI: http://dx.doi.org/10.7554/eLife.07770.001, eLife digest As the name implies, songbirds produce song, but they may also emit large numbers of shorter calls. Because calls are often given in social situations, they are difficult to record and to assign to the correct individual. Therefore, it is still unclear what information is communicated by these calls and how important they are. Zebra finches are highly vocal songbirds, with males singing and both females and males producing calls. In their natural habitat, Australia, these chatty birds pair for life and live in groups. To ensure successful breeding, zebra finches need to begin breeding activities as soon as the unpredictable environment allows. Therefore, even in captivity, they will readily breed when given nesting material. To find out about the role of zebra finch calls in relation to different environmental or social factors, Gill et al. brought together in groups female and male zebra finches that had not met before, and followed their individual calls during different breeding stages. This was done using a technique called microphone telemetry that involves placing tiny wireless microphones on the birds. The finches quickly formed breeding pairs, and when provided with nesting material, began building nests and laying eggs. While doing so, and especially when pairs began building nests, the birds changed how often they used certain calls and started using different call types; for example, they made more breeding-related ‘cackles’. Calls often featured precisely timed back-and-forth calling interactions, and, over time, were directed more and more towards their partner than other members of the group. Pairs that performed more of these call exchanges during nesting were more likely than others to lay a clutch of eggs. Overall, Gill et al. show that both the timing and types of calls used in pair communication are important for successful breeding. Future research could investigate the role of calls in group communication in more detail—possibly even in the wild—and how calling behaviour is reflected in the brain. DOI: http://dx.doi.org/10.7554/eLife.07770.002

During migration, many songbirds encounter large ecological barriers, like deserts and seas that require substantial fuel to cross and can lead to dehydration during passage. If muscle is not catabolized to generate metabolic water, birds must seek free water on a subsequent stopover to replenish the water lost. Yet, no work has examined if birds crossing large migration barriers use access to free water in concert with energy or protein stores to make stopover decisions. We captured 61 free-living Garden Warblers (Sylvia borin) in spring at a frequently used stopover site in the Mediterranean Sea, housed them with or without drinking water, and measured nocturnal rest- lessness (Zugunruhe) in relation to energy stores at capture. Both groups lost equivalent flight muscle mass overnight, suggesting that water-deprived birds did not preferentially catabolize this tissue to produce metabolic water. Fat score and body mass, but not flight muscle mass, were positively correlated with nocturnal activity in both treatment groups. However, the slope of the relationship between nocturnal activity and fat score differed between groups, with water- deprived birds of high fat score showing the highest Zu- gunruhe activity. Our results suggest that birds with large energy stores use access to drinking water to inform their decisions about when to depart from a stopover site. Indi- viduals with higher fat scores might be expected, regardless of flight muscle size, to depart a dry stopover site more readily than a site with freely available water. We suggest follow-up studies to further elucidate the mechanisms of this phenomenon and establish its prevalence in free-living birds.

Behavior of wild vertebrate individuals can vary in response to environmental or social factors. Such within-individual behavioral variation is often mediated by hormonal mechanisms. Hormones also serve as a basis for among-individual variations in behavior including animal personalities and the degree of responsiveness to environmental and social stimuli. How do relationships between hormones and behavioral traits evolve to produce such behavioral diversity within and among individuals? Answering questions about evolutionary processes generating among-individual variation requires characterizing how specific hormones are related to variation in specific behavioral traits, whether observed hormonal variation is related to individual fitness and, whether hormonal traits are consistent (repeatable) aspects of an individual's phenotype. With respect to within-individual variation, we need to improve our insight into the nature of the quantitative relationships between hormones and the traits they regulate, which in turn will determine how they may mediate behavioral plasticity of individuals. To address these questions, we review the actions of two steroid hormones, corticosterone and testosterone, in mediating changes in vertebrate behavior, focusing primarily on birds. In the first part, we concentrate on among-individual variation and present examples for how variation in corticosterone concentrations can relate to behaviors such as exploration of novel environments and parental care. We then review studies on correlations between corticosterone variation and fitness, and on the repeatability over time of corticosterone concentrations. At the end of this section, we suggest that further progress in our understanding of evolutionary patterns in the hormonal regulation of behavior may require, as one major tool, reaction norm approaches to characterize hormonal phenotypes as well as their responses to environments.In the second part, we discuss types of quantitative relationships between hormones and behavioral traits within individuals, using testosterone as an example. We review conceptual models for testosterone-behavior relationships and discuss the relevance of these models for within-individual plasticity in behavior. Next, we discuss approaches for testing the nature of quantitative relationships between testosterone and behavior, concluding that again reaction norm approaches might be a fruitful way forward.We propose that an integration of new tools, especially of reaction norm approaches into the field of behavioral endocrinology will allow us to make significant progress in our understanding of the mechanisms, the functional implications and the evolution of hormone–behavior relationships that mediate variation both within and among individuals. This knowledge will be crucial in light of already ongoing habitat alterations due to global change, as it will allow us to evaluate the mechanisms as well as the capacity of wild populations to adjust hormonally-mediated behaviors to altered environmental conditions. published

Unfavourable conditions throughout the period of parental care can severely affect growth, reproductive performance, and survival. Yet, individuals may be affected differently, depending on the developmental period during which constraints are experienced. Here we tested whether the nestling phase compared to the fledgling phase is more susceptible to nutritional stress by considering biometry, physiology, sexually selected male ornaments and survival using zebra finches (Taeniopygia guttata) as a model species. As nestlings (day 0-17) or fledglings (day 17-35), subjects were raised either on low or high quality food. A low quality diet resulted in significantly elevated baseline corticosterone titres in both nestlings and fledglings. Subjects showed substantial compensatory growth after they had experienced low quality food as nestlings but catch-up growth did neither lead to elevated baseline corticosterone titres nor did we detect long term effects on biometry, male cheek patch, or survival. The compensation for temporally unfavourable environmental conditions reflects substantial phenotypic plasticity and the results show that costs of catch-up growth were not mediated via corticosterone as a physiological correlate of allostatic load. These findings provide new insights into the mechanisms and plasticity with which animals respond to periods of constraints during development as they may occur in a mistiming of breeding.

During migration, a number of bird species rely on stopover sites for resting and feeding before and after crossing ecological barriers such as deserts or seas. The duration of a stopover depends on the combined effects of environmental factors, endogenous programmes and physiological conditions. Previous studies indicated that lean birds prolong their refuelling stopover compared with fat birds; however, the quantitative relationship between physiological conditions and stopover behaviour has not been studied yet. Here, we tested in a large sample of free-living birds of three European passerines (whinchats, Saxicola rubetra , garden warblers, Sylvia borin and whitethroats, Sylvia communis ) whether the amount of migratory restlessness (Zugunruhe) shown at a stopover site depends on physiological conditions. An integrated measure of condition based on body mass, amount of subcutaneous fat and thickness of pectoral muscles strongly predicted the intensity of Zugunruhe shown in recording cages in the night following capture. These results provide novel and robust quantitative evidence in support of the hypothesis that the amount of energy reserves plays a major role in determining the stopover duration in migratory birds.

Birdsong serves to attract mates and to deter territorial rivals. Even though song is not restricted to males, this dual function has almost exclusively been demonstrated for male song. To test the generality of hypotheses on birdsong, we investigated female song in the sex-role reversed, classically polyandrous African black coucal (Centropus grillii) in the context of female–female competition. We compared spontaneously vocalizing females with females vocally responding to a playback simulating a conspecific intruder. Females changed vocal parameters in response to playbacks: They lowered the pitch of their vocalizations and enhanced the duration of song elements when being challenged. Also, the composition of the vocalizations was altered. There was no significant correlation between pitch and body size parameters in spontaneous song, but there was for response songs, with larger females having a lower pitch. These changes in vocal properties suggest that the vocalizations are important for mutual assessment of competitive abilities in females. Our findings confirm the general role of intrasexual competition in vocal communication of birds. Electronic supplementary material The online version of this article (doi:10.1007/s00265-009-0836-0) contains supplementary material, which is available to authorized users.

Testosterone is assumed to be the key hormone related to resource-defence aggression. While this role has been confirmed mostly in the context of reproduction in male vertebrates, the effect of testosterone on the expression of resource-defence aggression in female vertebrates is not so well established. Furthermore, laboratory work suggests that progesterone inhibits aggressive behaviour in females. In this study, we investigated the hormonal changes underlying territorial aggression in free-living female African black coucals, Centropus grillii (Aves; Cuculidae). Females of this sex-role reversed polyandrous bird species should be particularly prone to be affected by testosterone because they aggressively defend territories similar to males of other species. We show, however, that territorial aggression in female black coucals is modulated by progesterone. After aggressive territorial challenges female black coucals expressed lower levels of progesterone than unchallenged territorial females and females without territories, suggesting that progesterone may suppress territorial aggression and is downregulated during aggressive encounters. Indeed, females treated with physiological concentrations of progesterone were less aggressive than females with placebo implants. This is one of the first demonstrations of a corresponding hormone–behaviour interaction under challenged and experimental conditions in free-living females. We anticipate that our observation in a sex-role reversed species may provide a more general mechanism, by which progesterone—in interaction with testosterone—may regulate resource-defence aggression in female vertebrates.

Ecological immunologists are interested in how immune function changes during different seasons and under different environmental conditions. However, an obstacle to answering such questions is discerning the effects of biological factors of interest and investigation artifacts such as handling stress. Here we examined handling stress and its effects on constitutive (noninduced) immune function via two protocols on captive red knots (Calidris canutus). We investigated how constitutive immunity responds to handling stress, how quickly these changes take place, and the practical implications for researchers interested in sampling baseline immune levels. We found that Staphylococcus aureus and Candida albicans killing increased with handling stress while total leukocyte and lymphocyte concentrations decreased. However, although corticosterone increased significantly and rapidly in response to handling stress, none of the 10 measures of constitutive immunity that we tested differed significantly from baseline within 20 or 30 min of capture. Thus, researchers interested in baseline immune function should sample animals as soon as possible after capture, but studies in species not easily sampled in less than 3 min (such as red knots) could still yield useful results.

Previous investigations suggest that male tropical birds have lower plasma testosterone concentrations than northern latitude species. To test whether this generalization is valid, we analyzed all currently available plasma testosterone data of tropical birds. We focused on peak breeding testosterone levels using phylogenetic and conventional statistics. Explanatory variables considered were social mating system, type of territoriality, breeding season length, and altitude. On average, tropical birds had lower mean peak testosterone levels than northern temperate birds. However, in several tropical species, testosterone levels were well within the range of northern latitude birds. Without controlling for phylogeny, breeding season length, type of territoriality, and altitude explained a significant proportion of the variance in testosterone levels. The shorter the breeding season, the higher the testosterone levels. Tropical birds that defend a breeding season territory had higher testosterone levels than birds that were year-round territorial or colonial, and testosterone levels were positively correlated with altitude. When controlling for phylogeny, only breeding season length predicted testosterone levels. In conclusion, we propose to refine previous notions of low plasma testosterone levels in tropical birds: short breeding seasons and perhaps environmental conditions at high altitudes precipitate conditions under which high testosterone levels are beneficial in the tropics.

The challenge hypothesis (Wingfield et al. in Am Nat 136:829–846, 1990) predicts that circulating testosterone increases when socially monogamous male birds are challenged during breeding. Although the challenge hypothesis has been confirmed in large-scale interspecific comparisons of seasonal hormone profiles, experimental tests of the challenge hypothesis are still uncommon and the results equivocal. We tested one of the predictions of the challenge hypothesis by investigating the behavioural and hormonal response of free-living European robins during simulated territorial intrusions (STIs) in the breeding season. We conducted STIs by placing a stuffed decoy in a territory and playing robin song. After the behaviour of the focal male had been recorded for at least 10 min, it was captured and a blood sample was taken immediately. Controls were caught within 10 min of the first response of the territory owner. Hormone concentrations were measured by radio-immunoassay. Although previous studies have shown that testosterone has an impact on aggression, European robins do not respond to STIs by increasing circulating levels of testosterone.

In many birds, the male reproductive tract is asymmetric with the left testis being larger than the right one. Coucals (Centropodidae) represent an exception as the asymmetry is typically reversed. Here, we describe the functional morphology of the reproductive tract of the African Black Coucal (Centropus grillii), a bird species with reversed sex-roles and a classical polyandrous mating system. In this species, the left testis has been reduced to a tiny, disk-like vestige that in almost all cases examined was not visible macroscopically. The vestigial left testis apparently does not participate in sperm production but has retained a vestigial left excurrent duct system. By contrast, the right deferent duct was engorged with sperm, suggesting a sperm storage function. Both deferent ducts opened on the tip of spectacularly long, erectile deferent duct papillae into the urodaeum of the male cloaca. Behavioural observations suggest that they may function as “pseudophalli”. Testis mass represented 0.49% of body mass, less than half the size of other classical polyandrous bird species. Assuming moderate to high levels of sperm competition this represents a paradox. Heavy demands on the energy budget while caring for altricial young may have prevented males from maintaining two testes and large bilateral sperm storage devices. Reduced testis mass may be compensated for by frequent transfer of small amounts of sperm. Female Black Coucals have not evolved any corresponding cloacal specializations, but like all other birds have sperm storage tubules.

International audience; Bacteria are fundamental associates of animals, and recent studies have highlighted their major role in host behaviour, immunity or reproductive investment. Thus, any environmental factor modifying bacterial community may affect host fitness. In birds, trace metals emitted by anthropogenic activities accumulate onto the plumage where they may alter bacterial community and ultimately affect bird fitness. Although trace metals are current major environmental issues in urban habitats, their effects on feather bacterial community have never been investigated. Here, we supplemented feral pigeons Columba livia, an emblematic urban species, with zinc and/or lead in drinking and bath water. As expected, lead and zinc supplementations modified plumage bacterial community composition. Zinc decreased bacterial load, while lead decreased bacterial richness and the frequency of preening behaviour in birds, known to regulate feather bacteria. Our results demonstrate for the first time the effects of common urban trace metals on plumage bacterial community and shed light on one of the mechanisms by which trace metals can affect bird fitness. Further studies are now needed to investigate how this effect modulates avian life history traits known to depend on plumage bacterial community.