Interplays between pre- and post-natal environments affect early-life mortality, body mass and telomere dynamics in the wild.

Early-life conditions are crucial determinants of phenotype and fitness. The effects of pre- and post-natal conditions on fitness prospects have been widely studied but their interactive effects have received less attention. In birds, asynchronous hatching creates challenging developmental conditions for the last-hatched chicks, but differential allocation in last-laid eggs might help to compensate this initial handicap. The relative importance and potential interaction between pre- and post-hatching developmental conditions for different fitness components remains mostly unknown. We manipulated hatching order in wild pied flycatchers (Ficedula hypoleuca), creating three groups: natural asynchrony (last-laid eggs hatching last), reversed asynchrony (last-laid eggs hatching first) and hatching synchrony (all eggs hatching at once). We examined the effects of these manipulations on early-life survival, growth and telomere length, a potential cellular biomarker of fitness prospects. Mortality was mostly affected by hatching order, with last-hatched chicks being more likely to die. Early-life telomere dynamics and growth were influenced by the interplays between laying and hatching order. Last-laid but first-hatched chicks were heavier but had shorter telomeres 5 days after hatching than their siblings, indicating rapid early growth with potential adverse consequences on telomere length. Synchronous chicks did not suffer any apparent cost of hatching synchronously. Impaired phenotypes only occurred when reversing the natural hatching order (i.e. developmental mismatch), suggesting that maternal investment in last-laid eggs might indeed counterbalance the initial handicap of last-hatched chicks. Our experimental study thus highlights that potential interplays between pre- and post-natal environments are likely to shape fitness prospects in the wild.

Wild zebra finches that nest synchronously have long-term stable social ties.

Many animals live and breed in colonies, and yet, with just a few exceptions, the value of the social bonds between colony members has rarely been examined. Social ties are important for group coordination at collective tasks, and social coordination can facilitate synchronized reproduction among colony members. Synchronized reproduction in turn can amplify the benefits of coloniality, such as social foraging and predator avoidance. We conducted a field study to investigate whether synchronized reproduction among individuals in replicated colonies is linked to the strength of their social bond, and whether these strong bonds are maintained beyond the reproductive period. We PIT-tagged wild zebra finches (Taeniopygia guttata), monitoring their reproduction and social foraging over two consecutive years. We then used social network analysis to characterize the strength of social bonds among birds in the population. We show that birds that are more synchronized in their reproductive timing (and breed in the same colony) had significantly stronger social ties both during and after reproduction than expected by chance. Our long-term sampling also revealed that the strong social ties between synchronized breeders were carried over across years. Our study reveals a strong correspondence between synchronized breeding and the social structure of the breeding colony. This suggests that the synchrony between pairs is not a simple process based on opportunity, but a mechanism underpinned by more complex sociality, which could be carried over to other behavioural contexts. The maintenance of cross-contextual social ties across years suggests that social structure could have extensive consequences on the overall life history of individuals in addition to playing a key role for the reproductive dynamics of colonial breeders.

Early-life social environment predicts social network position in wild zebra finches.

Early-life experience can fundamentally shape individual life-history trajectories. Previous research has suggested that exposure to stress during development causes differences in social behaviour later in life. In captivity, juvenile zebra finches exposed to elevated corticosterone levels were less socially choosy and more central in their social networks when compared to untreated siblings. These differences extended to other aspects of social life, with 'stress-exposed' juveniles switching social learning strategies and juvenile males less faithfully learning their father's song. However, while this body of research suggests that the impacts of early-life stress could be profound, it remains unknown whether such effects are strong enough to be expressed under natural conditions. Here, we collected data on social associations of zebra finches in the Australian desert after experimentally manipulating brood sizes. Juveniles from enlarged broods experienced heightened sibling competition, and we predicted that they would express similar patterns of social associations to stress-treated birds in the captive study by having more, but less differentiated, relationships. We show striking support for the suggested consequences of developmental stress on social network positions, with our data from the wild replicating the same results in 9 out of 10 predictions previously tested in captivity. Chicks raised in enlarged broods foraged with greater numbers of conspecifics but were less 'choosy' and more central in the social network. Our results confirm that the natural range of variation in early-life experience can be sufficient to predict individuals' social trajectories and support theory highlighting the potential importance of developmental conditions on behaviour.

Nosy neighbours: large broods attract more visitors. A field experiment in the pied flycatcher, Ficedula hypoleuca.

Life is uncertain. To reduce uncertainty and make adaptive decisions, individuals need to collect information. Individuals often visit the breeding sites of their conspecifics (i.e., "prospect"), likely to assess conspecifics' reproductive success and to use such information to identify high-quality spots for future breeding. We investigated whether visitation rate by prospectors and success of visited sites are causally linked. We manipulated the reproductive success (enlarged, reduced, and control broods) in a nest-box population of migratory pied flycatchers, Ficedula hypoleuca, in Finland. We measured the visitation rates of prospectors at 87 nest-boxes continuously from manipulation (day 3 after hatching) to fledging. 302 adult pied flycatchers prospected 9194 times on these manipulated nests (at least 78% of detected prospectors were successful breeders). While the number of visitors and visits was not influenced by the relative change in brood size we induced, the resulting absolute brood size predicted the prospecting behaviour: the larger the brood size after manipulation, the more visitors and visits a nest had. The parental provisioning rate at a nest and brood size pre-manipulation did not predict the number of visitors or visits post-manipulation. More visitors, however, inspected early than late nests and broods in good condition. Our study suggests that individuals collect social information when visiting conspecific nests during breeding and provides evidence that large broods attract more visitors than small broods. We discuss the results in light of individual decision-making by animals in their natural environments.

Predator encounters have spatially extensive impacts on parental behaviour in a breeding bird community.

Predation risk has negative indirect effects on prey fitness, partly mediated through changes in behaviour. Evidence that individuals gather social information from other members of the population suggests that events in a community may impact the behaviour of distant individuals. However, spatially wide-ranging impacts on individual behaviour caused by a predator encounter elsewhere in a community have not been documented before. We investigated the effect of a predator encounter (hawk model presented at a focal nest) on the parental behaviour of pied flycatchers (Ficedula hypoleuca), both at the focal nest and at nearby nests different distances from the predator encounter. We show that nest visitation of both focal pairs and nearby pairs were affected, up to 3 h and 1 h, respectively. Parents also appeared to compensate initial disrupted feeding by later increasing nest visitation rates. This is the first evidence showing that the behaviour of nearby pairs was affected away from an immediate source of risk. Our results indicate that the impacts of short-term predator encounters may immediately extend spatially to the broader community, affecting the behaviour of distant individuals. Information about predators is probably quickly spread by cues such as intra- and heterospecific alarm calls, in communities of different taxa.

Life-history trade-offs mediate 'personality' variation in two colour morphs of the pea aphid, Acyrthosiphon pisum.

Life-history trade-offs are considered a major driving force in the emergence of consistent behavioural differences (personality variation); but empirical tests are scarce. We investigated links between a personality trait (escape response), life-history and state variables (growth rate, size and age at first reproduction, age-dependent reproductive rates, lifetime reproductive success, life span) in red and green colour morphs of clonal pea aphids, Acyrthosiphon pisum. Escape response (dropping/non-dropping off a plant upon a predatory attack) was measured repeatedly to classify individuals as consistent droppers, consistent nondroppers or inconsistents. Red morphs experienced stronger trade-offs between early reproduction and life span than green morphs; and red consistent (non)droppers had highest lifetime reproductive success. Red droppers followed a risk-averse life-history strategy (high late reproduction), red nondroppers a risk-prone strategy (high early reproduction), while reproductive rates were equivalent for all green behavioural types and red inconsistents. This suggests that red morphs suffer the highest costs of dropping (they are most conspicuous to predators), which 'equivalates' fitness payoffs to both risk-takers (red non-droppers) and risk-averse red droppers. The strong trade-off also means that committing to a particular lifestyle (being consistent) maximises fitness. Our study suggests that life-history trade-offs likely mediate personality variation but effects might depend on interactions with other organismal characteristics (here: colour morph).

Heatwave-like events affect drone production and brood-care behaviour in bumblebees.

Climate change is currently considered one of the major threats to biodiversity and is associated with an increase in the frequency and intensity of extreme weather events, such as heatwaves. Heatwaves create acutely stressful conditions that may lead to disruption in the performance and survival of ecologically and economically important organisms, such as insect pollinators. In this study, we investigated the impact of simulated heatwaves on the performance of queenless microcolonies of Bombus terrestris audax under laboratory conditions. Our results indicate that heatwaves can have significant impacts on bumblebee performance. However, contrary to our expectations, exposure to heatwaves did not affect survival. Exposure to a mild 5-day heatwave (30-32 °C) resulted in increased offspring production compared to those exposed to an extreme heatwave (34-36 °C) and to the control group (24 °C). We also found that brood-care behaviours were impacted by the magnitude of the heatwave. Wing fanning occurred occasionally at temperatures of 30-32 °C, whereas at 34-36 °C the proportion of workers engaged in this thermoregulatory behaviour increased significantly. Our results provide insights into the effects of heatwaves on bumblebee colony performance and underscore the use of microcolonies as a valuable tool for studying the effects of extreme weather events. Future research, especially field-based studies replicating natural foraging conditions, is crucial to complement laboratory-based studies to comprehend how heatwaves compromise the performance of pollinators. Such studies may potentially help to identify those species more resilient to climate change, as well as those that are most vulnerable.

Environmental transmission of a personality trait: foster parent exploration behaviour predicts offspring exploration behaviour in zebra finches.

Consistent behavioural differences among individuals are common in many species and can have important effects on offspring fitness. To understand such 'personality' variation, it is important to determine the mode of inheritance, but this has been quantified for only a few species. Here, we report results from a breeding experiment in captive zebra finches, Taeniopygia guttata, in which we cross-fostered offspring to disentangle the importance of genetic and non-genetic transmission of behaviour. Genetic and foster-parents' exploratory type was measured in a novel environment pre-breeding and offspring exploratory type was assessed at adulthood. Offspring exploratory type was predicted by the exploratory behaviour of the foster but not the genetic parents, whereas offspring size was predicted by genetic but not foster-parents' size. Other aspects of the social environment, such as rearing regime (uni- versus biparental), hatching position, brood size or an individual's sex did not influence offspring exploration. Our results therefore indicate that non-genetic transmission of behaviour can play an important role in shaping animal personality variation.

Personality variation in a clonal insect: the pea aphid, Acyrthosiphon pisum.

Individuals are often consistent in their behavior but vary from each other in the level of behavior shown. Despite burgeoning interest in such animal personality variation, studies on invertebrates are scarce, and studies on clonal invertebrates nonexistent. This is surprising given the obvious advantages of using invertebrates/clones to tackle the crucial question why such consistent behavioral differences exist. Here we show that individuals of clonal pea aphids exhibit consistent behavioral differences in their escape responses to a predator attack (dropping vs. nondropping off a plant). However, behavior was not repeatable at the clonal level. Genetically identical clones expressed various phenotypes but different clones produced different proportions of each phenotype (dropper, nondropper, and inconsistent). Manipulations of early environmental conditions had little qualitative impact on such patterns. We discuss the importance of our findings for future studies of the evolutionary and ecological consequences of personality variation.

Sex differences in morphology across an expanding range edge in the flightless ground beetle, Carabus hortensis.

Species' ranges are dynamic, changing through range shifts, contractions, and expansions. Individuals at the edge of a species' shifting range often possess morphological traits that increase movement capacity, that are not observed in individuals farther back within the species' range. Although morphological traits that increase in proportion toward the range edge may differ between the sexes, such sex differences are rarely studied.Here, we test the hypotheses that body size and condition increase with proximity to an expanding range edge in the flightless ground beetle, Carabus hortensis, and that these trait changes differ between the sexes.Male, but not female, body size increased with proximity to the range edge. Body size was positively correlated with male front and mid tibia length and to female hind tibia length, indicating that body size is indicative of movement capacity in both sexes. Body condition (relative to body size) decreased with increasing population density in males but not females. Population density was lowest at the range edge.Our results indicate that sex is an important factor influencing patterns in trait distribution across species' ranges, and future studies should investigate changes in morphological traits across expanding range margins separately for males and females. We discuss the implications for sex differences in resource allocation and reproductive rates for trait differentiation across species' shifting ranges.

Sex-specific covariance between metabolic rate, behaviour and morphology in the ground beetle Carabus hortensis.

BACKGROUND: Individuals within the same species often differ in their metabolic rates, which may covary with behavioural traits (such as exploration), that are consistent across time and/or contexts, and morphological traits. Yet, despite the frequent occurrence of sexual dimorphisms in morphology and behaviour, few studies have assessed whether and how sexes differ in metabolic trait covariances. METHODS: We investigated sex-specific relationships among resting or active metabolic rate (RMR and AMR, respectively) with exploratory behaviour, measured independently of metabolic rate in a novel environment, body size and body mass, in Carabus hortensis ground beetles. RESULTS: RMR, AMR and exploratory behaviour were repeatable among individuals across time, except for male RMR which was unrepeatable. Female RMR neither correlated with exploratory behaviour nor body size/body mass. In contrast, AMR was correlated with both body size and exploratory behaviour. Males with larger body sizes had higher AMR, whereas females with larger body sizes had lower AMR. Both male and female AMR were significantly related to exploratory behaviour, though the relationships between AMR and exploration were body mass-dependent in males and temperature-dependent in females. DISCUSSION: Differences between sexes exist in the covariances between metabolic rate, body size and exploratory behaviour. This suggests that selection acts differently on males and females to produce these trait covariances with potentially important consequences for individual fitness.

Exploratory behaviour is not related to associative learning ability in the carabid beetle Nebria brevicollis.

Recently, it has been hypothesised that as learning performance and animal personality vary along a common axis of fast and slow types, natural selection may act on both in parallel leading to a correlation between learning and personality traits. We examined the relationship between risk-taking, exploratory behaviour and associative learning ability in carabid beetle Nebria brevicollis females by quantifying the number of trials individuals required to reach criterion during an associative learning task ('learning performance'). The associative learning task required the females to associate odour and direction with refugia from light and heat in a T-maze. Further, we assessed learning performance in a reversal task by quantifying the number of correct trials when the reinforcement was switched to previously unrewarding stimuli. We found that N. brevicollis females can associate conditional stimuli with a reward. No female was able to reverse the learned association within the number of trials given, however individuals differed in the number of correct trials in the reversal phase. Contrary to previous predictions neither exploratory behaviour, which was repeatable, nor risk-taking were correlated with learning performance. Our results suggest that the relationship between learning and personality may not take a common form across species.

Estimating food resource availability in arid environments with Sentinel 2 satellite imagery.

BACKGROUND: In arid environments, plant primary productivity is generally low and highly variable both spatially and temporally. Resources are not evenly distributed in space and time (e.g., soil nutrients, water), and depend on global (El Niño/ Southern Oscillation) and local climate parameters. The launch of the Sentinel2-satellite, part of the European Copernicus program, has led to the provision of freely available data with a high spatial resolution (10 m per pixel). Here, we aimed to test whether Sentinel2-imagery can be used to quantify the spatial variability of a minor tussock grass (Enneapogon spp.) in an Australian arid area and whether we can identify different vegetation cover (e.g., grass from shrubs) along different temporal scenarios. Although short-lasting, the Enneapogon grassland has been identified as a key primary food source to animals in the arid environment. If we are able to identify and monitor the productivity of this species remotely, it will provide an important new tool for examining food resource dynamics and subsequent animal responses to them in arid habitat. METHODS: We combined field vegetation surveys and Sentinel2-imagery to test if satellite spectral data can predict the spatial variability of Enneapogon over time, through GLMMs. Additionally, a cluster analysis ('gower' distance, 'complete' method), based on Enneapogon seed-productivity, and total vegetation cover in October 2016, identified three clusters: bare ground, grass dominated and shrub dominated. We compared the vegetation indices between these different clusters from October 2016 to January 2017. RESULTS: We found that MSAVI2 and NDVI correlated with the proportion of Enneapogon with seeds across the landscape and this relationship changed over time. Both vegetation indices (MSAVI2 and NDVI) were higher in patches with high seed-productivity of Enneapogon than in bare soil, but only in October, a climatically-favorable period during which this dominant grass reached peak seed-productivity. DISCUSSION: MSAVI2 and NDVI provided reliable estimates of the heterogeneity of vegetation type across the landscape only when measured in the Austral spring. This means that grass cover is related to seed-productivity and it is possible to remotely and reliably predict food resource availability in arid habitat, but only in certain conditions. The lack of significant differences between clusters in the summer was likely driven by the short-lasting nature of the vegetation in the study and the sparseness of the grass-dominated vegetation, in contrast to the shrub vegetation cluster that was particularly well measured by the NDVI. CONCLUSIONS: Overall, our study highlights the potential for Sentinel2-imagery to estimate and monitor the change in grass seed availability remotely in arid environments. However, heterogeneity in grassland cover is not as reliably measured as other types of vegetation and may only be well detected during periods of peak productivity (e.g., October 2016).

No male mate choice for female boldness in a bi-parental West African cichlid, the rainbow krib (Pelvicachromis pulcher).

BACKGROUND: In many species, males have a lower reproductive investment than females and are therefore assumed to increase their fitness with a high number of matings rather than by being choosy. However, in bi-parental species, also males heavily invest into reproduction. Here, reproductive success largely depends on costly parental care; with style and amount of parental effort in several cases being associated with personality differences (i.e., consistent between-individual differences in behaviour). Nonetheless, very little is known about the effect of personality differences on (male) mate choice in bi-parental species. METHODS: In the present study, we tested male mate choice for the level and consistency of female boldness in the rainbow krib, Pelviachromis pulcher, a bi-parental and territorial West African cichlid. Individual boldness was assumed to indicate parental quality because it affects parental defence behaviour. For all males and females, boldness was assessed twice as the activity under simulated predation risk. Mate choice trials were conducted in two steps. First, we let a male observe two females expressing their boldness. Then, the male could choose between these two females in a standard mate choice test. RESULTS: We tested for a male preference for behavioural (dis-)similarity vs. a directional preference for boldness but our data support the absence of effects of male and/or female boldness (level and consistency) on male mating preference. DISCUSSION: Our results suggest female personality differences in boldness may not be selected for via male mate choice.

Predictability is attractive: Female preference for behaviourally consistent males but no preference for the level of male aggression in a bi-parental cichlid.

Although personality traits can largely affect individual fitness we know little about the evolutionary forces generating and maintaining personality variation. Here, we investigated the hypothesis that personality variation in aggression is sexually selected in the monogamous, bi-parental cichlid Pelvicachromis pulcher. In this species, breeding pairs form territories and they aggressively defend their territory and offspring against con- and heterospecific intruders. In our mate choice study, we followed up two alternative hypotheses. We either expected females to show a directional preference for a high level and high consistency of aggression (potentially indicating mate choice for male parental quality). Alternatively, we expected females to choose males for (dis-)similarity in the level/consistency of aggression (potentially indicating mate choice for compatibility). Individual level and consistency of aggression were assessed for males and females using mirror tests. After eavesdropping on aggressive behaviour of two males (differing in level and consistency of aggression) females were then allowed to choose between the two males. Males, but not females, showed personality variation in aggression. Further, females generally preferred consistent over inconsistent males independent of their level of aggression. We did not detect a general preference for the level of male aggression. However, we found an above average preference for consistent high-aggression males; whereas female preference for inconsistent high-aggression did not deviate from random choice. Our results suggest behavioural consistency of aggression in male rainbow kribs is selected for via female mate choice. Further, our study underlines the importance of considering both the level and the consistency of a behavioural trait in studies of animal behaviour.

Evolution of sex-specific pace-of-life syndromes: genetic architecture and physiological mechanisms.

Sex differences in life history, physiology, and behavior are nearly ubiquitous across taxa, owing to sex-specific selection that arises from different reproductive strategies of the sexes. The pace-of-life syndrome (POLS) hypothesis predicts that most variation in such traits among individuals, populations, and species falls along a slow-fast pace-of-life continuum. As a result of their different reproductive roles and environment, the sexes also commonly differ in pace-of-life, with important consequences for the evolution of POLS. Here, we outline mechanisms for how males and females can evolve differences in POLS traits and in how such traits can covary differently despite constraints resulting from a shared genome. We review the current knowledge of the genetic basis of POLS traits and suggest candidate genes and pathways for future studies. Pleiotropic effects may govern many of the genetic correlations, but little is still known about the mechanisms involved in trade-offs between current and future reproduction and their integration with behavioral variation. We highlight the importance of metabolic and hormonal pathways in mediating sex differences in POLS traits; however, there is still a shortage of studies that test for sex specificity in molecular effects and their evolutionary causes. Considering whether and how sexual dimorphism evolves in POLS traits provides a more holistic framework to understand how behavioral variation is integrated with life histories and physiology, and we call for studies that focus on examining the sex-specific genetic architecture of this integration.

Ground beetles in city forests: does urbanization predict a personality trait?

BACKGROUND: Urbanization leads to substantial changes in natural habitats with profound effects on wildlife. Understanding behavioural responses to such environmental change is essential for identifying which organisms may adapt, as behaviour is often the first response to altered conditions. Individuals in more urbanized habitats may be expected to be more exploratory and bolder than their conspecifics in less urbanized habitats as they may be better able to cope with novel challenges. METHODS: In a two-year field study we tested ground beetles from differently urbanized forests for their exploratory behaviour (in a novel environment) and their risk-taking (death-feigning). In total, we tested ca. 3,000 individuals of four forest-dwelling ground beetle species from eight within-city forest patches. In the second year, we also transferred ca. 800 tested individuals of two species to the laboratory to test for consistent behavioural differences (i.e. personality differences) under standardised conditions. RESULTS: Individuals were generally more exploratory in more urbanized than in less urbanized areas but only in one year of the study. Exploratory behaviour was not predicted by population density but increased with temperature or showed a temperature optimum. Exploration was consistent over time and individuals that were more exploratory also took higher risks. DISCUSSION: We demonstrated that species which are generally less directly exposed to human activities (e.g., most invertebrates) show behavioural responses to urbanization. Effects of urbanization were year-dependent, suggesting that other environmental conditions interacted with effects of urbanization on beetle behaviour. Furthermore, our results indicate that different personality compositions might cause behavioural differences among populations living in differently urbanized habitats.

Sexual selection and animal personality.

Consistent individual behavioural tendencies, termed "personalities", have been identified in a wide range of animals. Functional explanations for personality have been proposed, but as yet, very little consideration has been given to a possible role for sexual selection in maintaining differences in personality and its stability within individuals. We provide an overview of the available literature on the role of personality traits in intrasexual competition and mate choice in both human and non-human animals and integrate this into a framework for considering how sexual selection can generate and maintain personality. For this, we consider the evolution and maintenance of both main aspects of animal personality: inter-individual variation and intra-individual consistency.