The effect of environmental variation on the relationship between survival and risk-taking behaviour in a migratory songbird.

Temporal changes in environmental conditions may play a major role in the year-to-year variation in fitness consequences of behaviours. Identifying environmental drivers of such variation is crucial to understand the evolutionary trajectories of behaviours in natural contexts. However, our understanding of how environmental variation influences behaviours in the wild remains limited. Using data collected over 14 breeding seasons from a collared flycatcher (Ficedula albicollis) population, we examined the effect of environmental variation on the relationship between survival and risk-taking behaviour, a highly variable behavioural trait with great evolutionary and ecological significance. Specifically, using annual recapture probability as a proxy of survival, we evaluated the specific effect of predation pressure, food availability, and mean temperature on the relationship between annual recapture probability and risk-taking behaviour (measured as flight initiation distance [FID]). We found a negative trend, as the relationship between annual recapture probability and FID decreased over the study years and changed from positive to negative. Specifically, in the early years of the study, risk-avoiding individuals exhibited a higher annual recapture probability, whereas in the later years, risk-avoiders had a lower annual recapture probability. However, we did not find evidence that any of the considered environmental factors mediated the variation in the relationship between survival and risk-taking behaviour.

Are brood sex ratios adaptive?-The effect of experimentally altered brood sex ratio on nestling growth, mortality and recruitment.

Brood sex ratios (BSRs) have often been found to be nonrandom in respect of parental and environmental quality, and many hypotheses suggest that nonrandom sex ratios can be adaptive. To specifically test the adaptive value of biased BSRs, it is crucial to disentangle the consequences of BSR and maternal effects. In multiparous species, this requires cross-fostering experiments where foster parents rear offspring originating from multiple broods, and where the interactive effect of original and manipulated BSR on fitness components is tested. To our knowledge, our study on collared flycatchers (Ficedula albicollis) is the first that meets these requirements. In this species, where BSRs had previously been shown to be related to parental characteristics, we altered the original BSR of the parents shortly after hatching by cross-fostering nestlings among trios of broods and examined the effects on growth, mortality and recruitment of the nestlings. We found that original and experimental BSR, as well as the interaction of the two, were unrelated to the fitness components considered. Nestling growth was related only to background variables, namely brood size and hatching rank. Nestling mortality was related only to hatching asynchrony. Our results therefore do not support that the observed BSRs are adaptive in our study population. However, we cannot exclude the possibility of direct effects of experimentally altered BSRs on parental fitness, which should be evaluated in the future. In addition, studies similar to ours are required on various species to get a clearer picture of the adaptive value of nonrandom BSRs.

Plato's cube and the natural geometry of fragmentation.

Plato envisioned Earth's building blocks as cubes, a shape rarely found in nature. The solar system is littered, however, with distorted polyhedra-shards of rock and ice produced by ubiquitous fragmentation. We apply the theory of convex mosaics to show that the average geometry of natural two-dimensional (2D) fragments, from mud cracks to Earth's tectonic plates, has two attractors: "Platonic" quadrangles and "Voronoi" hexagons. In three dimensions (3D), the Platonic attractor is dominant: Remarkably, the average shape of natural rock fragments is cuboid. When viewed through the lens of convex mosaics, natural fragments are indeed geometric shadows of Plato's forms. Simulations show that generic binary breakup drives all mosaics toward the Platonic attractor, explaining the ubiquity of cuboid averages. Deviations from binary fracture produce more exotic patterns that are genetically linked to the formative stress field. We compute the universal pattern generator establishing this link, for 2D and 3D fragmentation.

Functional Integration of Multiple Sexual Ornaments: Signal Coherence and Sexual Selection.

AbstractThe sexual ornamentation of animals typically consists of multiple distinct traits. The classical research approach focuses on differences among these traits, but this approach may often be misleading because of correlations among distinct sexual traits of similar origins. There are many published studies on the correlation structures of sexual traits, but the way receivers take into account the components of an integrated, multicomponent trait system remains mostly unknown. Here, we propose a general analytical framework to assess the possible sexual selection consequences of within-individual coherence in the expression of multiple correlated sexual traits. We then apply this framework to a long-term mutual plumage coloration data set from a wild bird population. The results suggest that the coherence of component plumage color traits is not sexually selected. However, component trait coherence affects sexual selection on integrated plumage color. When assessing across-spectrum plumage reflectance, receivers choosing mates apparently disregard a component trait if it is inconsistent with the overall expression of other components. This indicates that separately examining and manipulating distinct sexual traits may often be misleading. Theoretical and empirical studies should further explore the effects of coherence on the ornament-preference coevolution.

Gravity Governs Shear Localization in Confined Dense Granular Flows.

The prediction of flow profiles of slowly sheared granular materials is a major geophysical and industrial challenge. Understanding the role of gravity is particularly important for future planetary exploration in varying gravitational environments. Using the principle of minimization of energy dissipation, and combining experiments and variational analysis, we disentangle the contributions of the gravitational acceleration, confining pressure, and layer thickness on shear strain localization induced by moving fault boundaries at the bottom of a granular layer. The flow profile is independent of the gravity for geometries with a free top surface. However, under a confining pressure or if the sheared layer withstands the weight of the upper layers, increasing gravity promotes the transition from closed shear zones buried in the bulk to open ones that intersect the top surface. We show that the center position and width of the shear zone and the axial angular velocity at the top surface follow universal scaling laws when properly scaled by the gravity, applied pressure, and layer thickness. Our finding that the flow profiles lie on a universal master curve opens the possibility to predict the quasistatic shear flow of granular materials in varying gravitational environments.

Evolution of shear zones in granular packings under pressure.

Stress transmission in realistic granular media often occurs under external load and in the presence of boundary slip. We investigate shear localization in a split-bottom Couette cell with smooth walls subject to a confining pressure experimentally and by means of numerical simulations. We demonstrate how the characteristics of the shear zone, such as its center position and width, evolve as the confining pressure and wall slip modify the local effective friction coefficient of the material. For increasing applied pressure, the shear zone evolves toward the center of the cylinder and grows wider and the angular velocity reduces compared to the driving rate of the bottom disk. Moreover, the presence of slip promotes the transition from open shear zones at the top surface to closed shear zones inside the bulk. We also systematically vary the ratio of the effective friction near the bottom plate and in the bulk in simulations and observe the resulting impact on the surface flow profile. Besides the boundary conditions and external load, material properties such as grain size are also known to influence the effective friction coefficient. However, our numerical results reveal that the center position and width of the shear zone are insignificantly affected by the choice of the grain size as far as it remains small compared to the radius of the rotating bottom disk.

Host dispersal shapes the population structure of a tick-borne bacterial pathogen.

Birds are hosts for several zoonotic pathogens. Because of their high mobility, especially of longdistance migrants, birds can disperse these pathogens, affecting their distribution and phylogeography. We focused on Borrelia burgdorferi sensu lato, which includes the causative agents of Lyme borreliosis, as an example for tick-borne pathogens, to address the role of birds as propagation hosts of zoonotic agents at a large geographical scale. We collected ticks from passerine birds in 11 European countries. B. burgdorferi s.l. prevalence in Ixodes spp. was 37% and increased with latitude. The fieldfare Turdus pilaris and the blackbird T. merula carried ticks with the highest Borrelia prevalence (92 and 58%, respectively), whereas robin Erithacus rubecula ticks were the least infected (3.8%). Borrelia garinii was the most prevalent genospecies (61%), followed by B. valaisiana (24%), B. afzelii (9%), B. turdi (5%) and B. lusitaniae (0.5%). A novel Borrelia genospecies "Candidatus Borrelia aligera" was also detected. Multilocus sequence typing (MLST) analysis of B. garinii isolates together with the global collection of B. garinii genotypes obtained from the Borrelia MLST public database revealed that: (a) there was little overlap among genotypes from different continents, (b) there was no geographical structuring within Europe, and (c) there was no evident association pattern detectable among B. garinii genotypes from ticks feeding on birds, questing ticks or human isolates. These findings strengthen the hypothesis that the population structure and evolutionary biology of tick-borne pathogens are shaped by their host associations and the movement patterns of these hosts.

Digit ratio predicts the number of lifetime recruits in female collared flycatchers.

The early environment in which an organism grows can have long-lasting impacts on both its phenotype and fitness. However, assessing this environment comprehensively is a formidable task. The relative length of the second to the fourth digit (2D : 4D) is a broadly studied skeletal trait that is fixed for life during ontogeny. 2D : 4D has been shown to indicate various early effects including the perinatal steroid milieu in both humans and non-human animals. However, the fitness relevance of the early effects indicated by 2D : 4D remains unknown. Here, we investigated hindlimb 2D : 4D and measures of lifetime performance in wild collared flycatcher ( Ficedula albicollis) females. We found that females with higher 2D : 4D had a greater number of recruiting offspring to the breeding population. This was the case despite the fact that such females did not lay more eggs or breed more frequently during their reproductive life. Our results support the suggestion that 2D : 4D, known to be a retrospective marker of perinatal development, positively associates with female quality in the collared flycatcher.

Relationship between oxidative stress and sexual coloration of lizards depends on thermal habitat.

Sexual signals serve as an honest indicator of individual quality, reflecting either developmental and/or maintenance costs. A possible underlying physiological mechanism is oxidative stress, which could mediate energy trade-offs between sexual signals and other quality traits. In ectotherms, thermal performance acts as a key indicator of individual quality and influence signal intensity. We investigated how oxidative state is reflected in visual signals of lizards from different thermal habitats. According to our hypothesis, efficient thermoregulation requires different strategies in different thermal environments. In a habitat with predictable temperature changes, animals are less exposed to suboptimal temperature ranges and selection will, therefore, be stronger on the maximum oxidative damage at optimal body temperature. Contrarily, in a habitat with rather stochastic thermal shifts, individuals are often constricted by suboptimal thermal conditions, and oxidative damage can be limiting on a wide temperature range. We used Iberolacerta cyreni and Psammodromus algirus inhabiting stochastic and predictable thermal environments respectively. We examined two aspects of oxidative stress: the level of reactive oxygen metabolites at the preferred temperature (maximal ROM) and the temperature range in which animals produce at least 80% of the maximum level of reactive oxygen metabolites (effective ROM range). In I. cyreni, we found that duller coloration was related to a wider effective ROM range, while expression of coloration in P. algirus was negatively correlated with the maximal ROM. Our results suggest that different thermal constraints affect different aspects of oxidative damage which can indicate individual quality and are, therefore, represented in sexual ornaments.

Male territorial aggression and fitness in collared flycatchers: a long-term study.

Aggressive behaviour plays a fundamental role in the distribution of limiting resources. Thereby, it is expected to have consequences for fitness. Here, we explored the relationship between aggression and fitness in a long-term database collected in a wild population of the collared flycatcher (Ficedula albicollis). We quantified the aggression of males during nest-site defence by conducting simulated territorial intrusions in the courtship period. We estimated the fitness of males based on their pairing success, breeding output and survival to next year. Earlier arriving and older males had a higher probability to establish pair-bond, and males that started to breed earlier fledged more young. Aggression did not predict pairing and breeding performances. However, the probability of a male to return in the next year was significantly related to aggression in an age-dependent manner. Among subadult males, more aggressive individuals had higher chances to return, while among adult males, less aggressive ones did so. This finding is in harmony with our general observation that subadult collared flycatcher males behave more aggressively than adult males when confronted with a conspecific intruder. Subadult males may be socially inexperienced, so they should be more aggressive to be successful. In contrast, if adult males suffer from higher physiological costs, a lower level of aggression may be more advantageous for them. Our study shows that aggressive behaviour can be a fitness-related trait, and to understand its role in determining fitness, age should be taken into account.

Linked selection and recombination rate variation drive the evolution of the genomic landscape of differentiation across the speciation continuum of Ficedula flycatchers.

Speciation is a continuous process during which genetic changes gradually accumulate in the genomes of diverging species. Recent studies have documented highly heterogeneous differentiation landscapes, with distinct regions of elevated differentiation ("differentiation islands") widespread across genomes. However, it remains unclear which processes drive the evolution of differentiation islands; how the differentiation landscape evolves as speciation advances; and ultimately, how differentiation islands are related to speciation. Here, we addressed these questions based on population genetic analyses of 200 resequenced genomes from 10 populations of four Ficedula flycatcher sister species. We show that a heterogeneous differentiation landscape starts emerging among populations within species, and differentiation islands evolve recurrently in the very same genomic regions among independent lineages. Contrary to expectations from models that interpret differentiation islands as genomic regions involved in reproductive isolation that are shielded from gene flow, patterns of sequence divergence (d(xy) and relative node depth) do not support a major role of gene flow in the evolution of the differentiation landscape in these species. Instead, as predicted by models of linked selection, genome-wide variation in diversity and differentiation can be explained by variation in recombination rate and the density of targets for selection. We thus conclude that the heterogeneous landscape of differentiation in Ficedula flycatchers evolves mainly as the result of background selection and selective sweeps in genomic regions of low recombination. Our results emphasize the necessity of incorporating linked selection as a null model to identify genome regions involved in adaptation and speciation.

MHC-mediated sexual selection on birdsong: Generic polymorphism, particular alleles and acoustic signals.

Several hypotheses predict that the major histocompatibility complex (MHC) drives mating preference in females. Olfactory, colour or morphological traits are often found as reliable signals of the MHC profile, but the role of avian song mediating MHC-based female choice remains largely unexplored. We investigated the relationship between several MHC and acoustic features in the collared flycatcher (Ficedula albicollis), a European passerine with complex songs. We screened a fragment of the class IIB second exon of the MHC molecule, of which individuals harbour 4-15 alleles, while considerable sequence diversity is maintained at the population level. To make statistical inferences from a large number of comparisons, we adopted both null-hypothesis testing and effect size framework in combination with randomization procedures. After controlling for potential confounding factors, neither MHC allelic diversity nor the presence of particular alleles was associated remarkably with the investigated qualitative and quantitative song traits. Furthermore, genetic similarity among males based on MHC sequences was not reflected by the similarity in their song based on syllable content. Overall, these results suggest that the relationship between features of song and the allelic composition and diversity of MHC is not strong in the studied species. However, a biologically motivated analysis revealed that individuals that harbour an MHC allele that impairs survival perform songs with broader frequency range. This finding suggests that certain aspects of the song may bear reliable information concerning the MHC profile of the individuals, which can be used by females to optimize mate choice.

Phenological sensitivity to climate change is higher in resident than in migrant bird populations among European cavity breeders.

Many organisms adjust their reproductive phenology in response to climate change, but phenological sensitivity to temperature may vary between species. For example, resident and migratory birds have vastly different annual cycles, which can cause differential temperature sensitivity at the breeding grounds, and may affect competitive dynamics. Currently, however, adjustment to climate change in resident and migratory birds have been studied separately or at relatively small geographical scales with varying time series durations and methodologies. Here, we studied differential effects of temperature on resident and migratory birds using the mean egg laying initiation dates from 10 European nest box schemes between 1991 and 2015 that had data on at least one resident tit species and at least one migratory flycatcher species. We found that both tits and flycatchers advanced laying in response to spring warming, but resident tit populations advanced more strongly in relation to temperature increases than migratory flycatchers. These different temperature responses have already led to a divergence in laying dates between tits and flycatchers of on average 0.94 days per decade over the current study period. Interestingly, this divergence was stronger at lower latitudes where the interval between tit and flycatcher phenology is smaller and winter conditions can be considered more favorable for resident birds. This could indicate that phenological adjustment to climate change by flycatchers is increasingly hampered by competition with resident species. Indeed, we found that tit laying date had an additional effect on flycatcher laying date after controlling for temperature, and this effect was strongest in areas with the shortest interval between both species groups. Combined, our results suggest that the differential effect of climate change on species groups with overlapping breeding ecology affects the phenological interval between them, potentially affecting interspecific interactions.

Reflectance variation in the blue tit crown in relation to feather structure.

Structural plumage colour is one of the most enigmatic sexually selected traits. The information content of structural colour variation is debated, and the heterogeneity of the findings is hard to explain because the proximate background of within-species colour differences is very scarcely studied. We combined measurements of feather macrostructure and nanostructure to explain within-population variability in blue tit crown reflectance. We found that sexual dichromatism in aspects of crown reflectance was explained only by feather macrostructure, whereas nanostructural predictors accounted for some of the age-related differences in reflectance. Moreover, we found that both mean reflectance and spectral shape traits reflected a combination of quantity and regularity aspects in macrostructure and nanostructure. This rich proximate background provides ample scope for reflectance to convey various types of information on individual quality.

Frustrated packing in a granular system under geometrical confinement.

Optimal packings of uniform spheres are solved problems in two and three dimensions. The main difference between them is that the two-dimensional ground state can be easily achieved by simple dynamical processes while in three dimensions, this is impossible due to the difference in the local and global optimal packings. In this paper we show experimentally and numerically that in 2 + ε dimensions, realized by a container which is in one dimension slightly wider than the spheres, the particles organize themselves in a triangular lattice, while touching either the front or rear side of the container. If these positions are denoted by up and down the packing problem can be mapped to a 1/2 spin system. At first it looks frustrated with spin-glass like configurations, but the system has a well defined ground state built up from isosceles triangles. When the system is agitated, it evolves very slowly towards the potential energy minimum through metastable states. We show that the dynamics is local and is driven by the optimization of the volumes of 7-particle configurations and by the vertical interaction between touching spheres.

Experience during development triggers between-individual variation in behavioural plasticity.

Behavioural consistency within and across behaviours (animal personality and behavioural syndrome, respectively) has been vigorously studied in the last decade, leading to the emergence of "animal personality" research. It has been proposed recently that not only mean behaviour (behavioural type), but the environmentally induced behavioural change (behavioural plasticity) might also differ between individuals within populations. While case studies presenting between-individual variation in behavioural plasticity have started to accumulate, the mechanisms behind its emergence are virtually unknown. We have recently demonstrated that ecologically relevant environmental stimuli during ontogeny are necessary for the development of animal personality and behavioural syndromes. However, it is unknown whether between-individual variation in behavioural plasticity is hard-wired or induced. Here, we tested whether experience with predation during development affected predator-induced behavioural plasticity in Rana dalmatina tadpoles. We ran a common garden experiment with two ontogenetic predation treatments: tadpoles developed from hatching in either the presence or absence of olfactory predator stimuli. Then, we assayed all tadpoles repeatedly for activity and risk-taking both in the absence and presence of olfactory predator stimuli. We found that (a) between-individual variation in predator-induced behavioural plasticity was present only in the group that developed in the presence of olfactory stimuli from predators and (b) previous experience with predatory stimuli resulted in lower plastic response at the group level. The latter pattern resulted from increased between-individual variation and not from universally lower individual responses. We also found that experience with predation during development increased the predictability (i.e. decreased the within-individual variation unrelated to environmental change) of activity, but not risk-taking. In line with this, tadpoles developing under perceived predatory risk expressed their activity with higher repeatability. We suggest that ecologically relevant environmental stimuli are not only fundamental for the development of animal personality and behavioural syndromes, but also for individual variation in behavioural plasticity. Thus, experience is of central importance for the emergence of individual behavioural variation at many levels.

Side-specific effect of yolk testosterone elevation on second-to-fourth digit ratio in a wild passerine.

Second-to-fourth digit ratio is a widely investigated sexually dimorphic morphological trait in human studies and could reliably indicate the prenatal steroid environment. Conducting manipulative experiments to test this hypothesis comes up against ethical limits in humans. However, oviparous tetrapods may be excellent models to experimentally investigate the effects of prenatal steroids on offspring second-to-fourth digit ratio. In this field study, we injected collared flycatcher (Ficedula albicollis) eggs with physiological doses of testosterone. Fledglings from eggs with elevated yolk testosterone, regardless of their sex, had longer second digits on their left feet than controls, while the fourth digit did not differ between groups. Therefore, second-to-fourth digit ratio was higher in the testosterone-injected group, but only on the left foot. This is the first study which shows experimentally that early testosterone exposure can affect second-to-fourth digit ratio in a wild population of a passerine bird.

Haemoproteus infection status of collared flycatcher males changes within a breeding season.

In ecological studies of haemosporidian parasites, prevalence is typically considered as a stable attribute. However, little is known about the possible within-host dynamics of these parasites that may originate from environmental fluctuations, parasite life cycles and the ability of hosts to suppress or clear infection. We sampled the blood of male collared flycatchers Ficedula albicollis twice within a breeding season and investigated the determinants of initial infection status and change in infection status. We found that older males tended to be initially more infected at courtship. Change in infection status was unrelated to male traits, but a widespread disappearance of Haemoproteus pallidus infection from the blood was detected between courtship and nestling rearing. The probability of change in infection status increased with the time elapsed between sampling occasions. This suggests that the disappearance of infection from the blood was due to either an active parasite suppression mechanism or the beginning of the latent phase in the parasite life cycle. Initial infection status or disappearance of infection from the blood showed no correlation with breeding success. These results show that H. pallidus infection status and thus prevalence are dynamically changing attributes and this has widespread practical and ecological implications.

Behavioural consistency and life history of Rana dalmatina tadpoles.

The focus of evolutionary behavioural ecologists has recently turned towards understanding the causes and consequences of behavioural consistency, manifesting either as animal personality (consistency in a single behaviour) or behavioural syndrome (consistency across more behaviours). Behavioural type (mean individual behaviour) has been linked to life-history strategies, leading to the emergence of the integrated pace-of-life syndrome (POLS) theory. Using Rana dalmatina tadpoles as models, we tested if behavioural consistency and POLS could be detected during the early ontogenesis of this amphibian. We targeted two ontogenetic stages and measured activity, exploration and risk-taking in a common garden experiment, assessing both individual behavioural type and intra-individual behavioural variation. We observed that activity was consistent in all tadpoles, exploration only became consistent with advancing age and risk-taking only became consistent in tadpoles that had been tested, and thus disturbed, earlier. Only previously tested tadpoles showed trends indicative of behavioural syndromes. We found an activity-age at metamorphosis POLS in the previously untested tadpoles irrespective of age. Relative growth rate correlated positively with the intra-individual variation of activity of the previously untested older tadpoles. In previously tested older tadpoles, intra-individual variation of exploration correlated negatively and intra-individual variation of risk-taking correlated positively with relative growth rate. We provide evidence for behavioural consistency and POLS in predator- and conspecific-naive tadpoles. Intra-individual behavioural variation was also correlated to life history, suggesting its relevance for the POLS theory. The strong effect of moderate disturbance related to standard behavioural testing on later behaviour draws attention to the pitfalls embedded in repeated testing.

Laying date and polygyny as determinants of annual reproductive success in male collared flycatchers (Ficedula albicollis): a long-term study.

Annual reproductive success (ARS) is one of the main components of lifetime reproductive success, a reliable measure of individual fitness. Previous studies often dealt with ARS and variables potentially affecting it. Among them, long-term studies that consider multiple factors at the same time are particularly important in understanding the adaptive value of different phenotypes. Here, we used an 18-year dataset to quantify the ARS of male collared flycatchers (Ficedula albicollis) on the basis of recruited offspring. We simultaneously assessed the effect of start of breeding, age, polygyny, body size and the expression of forehead patch (a sexually selected trait). The success of early breeding individuals was appreciably higher than late birds; however, breeding too early was also disadvantaged, and males that bred around the yearly median breeding date had the highest ARS. Polygynous males were more successful in years with good food supply, while in years with low food availability, they did not produce more recruits than monogamous males. The age of males, their forehead patch size and body size did not affect the number of recruits. Our findings support the importance of breeding date and suggest stabilizing selection on it in the long term. We also show that polygyny is not always advantageous for males, and its fitness pay-off may depend on environmental quality.