Untangling behaviours: independent expressions of female-female aggression and snake-like hissing in the blue tit (Cyanistes caeruleus).

Aggression plays a crucial role in deterring predators and securing resources to promote fitness. Nevertheless, studies focussing on female aggression remain scarce. In songbirds, aggression is prevalent during the breeding season, when same-sex individuals compete for limited resources. Additionally, females of some bird species exhibit snake-like hissing behaviour during incubation presumably to lower predation rates and improve fitness. Such behaviours may co-vary, forming a behavioural syndrome that could constrain trait expression. Here, we investigated a resident population of blue tits (Cyanistes caeruleus), to examine the repeatability and covariation of female-female aggression and hissing behaviour, aiming to determine if these constitute a behavioural syndrome. We quantified female-female aggression during simulated territorial intrusions and measured number of hissing calls in response to a simulated predator intrusion into the nest box. We found that both female-female aggression and hissing behaviour were repeatable traits, and that older females approached the intruder less. However, we found no evidence of covariation between female-female aggression and hissing behaviour. Thus, our findings suggest that female-female aggression and hissing behaviour, although both displayed in a nest defence context, are evolutionarily independent traits in the blue tit.

Females Are More Aggressive Than Males towards Same- and Opposite-Sex Intruders in the Blue Tit (Cyanistes caeruleus).

During the breeding season, aggression is expressed to gain access to resources such as territories and mates and protect offspring. Female aggressiveness has received much less attention than male aggressiveness, and few studies have examined female and male aggressiveness towards intruders of both sexes in the same species. We compared female and male aggressiveness towards same- and opposite-sex intruders during the egg-laying period in blue tits (Cyanistes caeruleus) using simulated territorial intrusions. For each sex, we examined the occurrence of different behavioural responses during agonistic encounters, and compared the intensity and individual consistency of intra- and inter-sexual aggression using same- and opposite-sex taxidermy mounts. Our results show that females are the more aggressive sex. Both sexes showed similar behaviours during simulated intrusions, although females were never observed singing and males never entered the nest box. In females, aggression was predominantly independent of the sex of the intruder, while males sang more from a distance during male-male encounters. The relative levels of aggression (pecking and perching on the mounts) during intra- and intersexual conflicts were consistent for females, but not for males. Females might be under stronger selection for aggressive phenotypes due to nest-hole competition and larger reproductive investments.

Anthropogenic noise and light pollution additively affect sleep behaviour in free-living birds in sex- and season-dependent fashions.

Rapid anthropogenic transformation of environments exposes organisms to diverse disturbance factors, including anthropogenic noise pollution and artificial light at night (ALAN). These sensory pollutants interfere with acquisition of, and response to, environmental cues and can be perceived as stressors. Noise pollution and ALAN are often experienced simultaneously, and are thus likely to jointly affect organisms, either additively or interactively. Yet, combined effects of noise pollution and ALAN remain poorly elucidated. We studied combined effects of noise pollution and ALAN on the sleep behaviour of a free-living songbird, the great tit (Parus major). Sleep is widely conserved across animal taxa and fulfils essential functions, and research has demonstrated independent effects of both noise and ALAN on sleep. We measured noise and light levels at nest boxes and used infrared video-recording to assess sleep behaviour. Results did not support interactive effects of noise and ALAN. However, noise pollution and ALAN were both independently related to variation in sleep behaviour, in sex- and season-dependent fashions. Males, but not females, woke up and left the nest box ∼20 min later in the noisiest as compared to quietest environments (range: 44.2-79.4 dB), perhaps because males are more sensitive to acoustical cues that are masked by noise. Furthermore, as the season progressed from November to early March, birds woke up and left the nest box ∼35 min earlier relative to sunrise on territories with the lowest, but not the highest, light levels (range: 0.01-8.5 lux). Thus, the seasonal difference in sleep duration was dampened on light polluted territories. These effects could arise if ALAN interferes with birds' ability to sense and respond to increasing daylength, and could have fitness ramifications. Our study suggests that noise pollution and ALAN exert additive effects on sleep behaviour, and that these effects can be sex- and season-dependent.

Temperature synchronizes temporal variation in laying dates across European hole-nesting passerines.

Identifying the environmental drivers of variation in fitness-related traits is a central objective in ecology and evolutionary biology. Temporal fluctuations of these environmental drivers are often synchronized at large spatial scales. Yet, whether synchronous environmental conditions can generate spatial synchrony in fitness-related trait values (i.e., correlated temporal trait fluctuations across populations) is poorly understood. Using data from long-term monitored populations of blue tits (Cyanistes caeruleus, n = 31), great tits (Parus major, n = 35), and pied flycatchers (Ficedula hypoleuca, n = 20) across Europe, we assessed the influence of two local climatic variables (mean temperature and mean precipitation in February-May) on spatial synchrony in three fitness-related traits: laying date, clutch size, and fledgling number. We found a high degree of spatial synchrony in laying date but a lower degree in clutch size and fledgling number for each species. Temperature strongly influenced spatial synchrony in laying date for resident blue tits and great tits but not for migratory pied flycatchers. This is a relevant finding in the context of environmental impacts on populations because spatial synchrony in fitness-related trait values among populations may influence fluctuations in vital rates or population abundances. If environmentally induced spatial synchrony in fitness-related traits increases the spatial synchrony in vital rates or population abundances, this will ultimately increase the risk of extinction for populations and species. Assessing how environmental conditions influence spatiotemporal variation in trait values improves our mechanistic understanding of environmental impacts on populations.

An Experimental Approach to Investigating Effects of Artificial Light at Night on Free-Ranging Animals: Implementation, Results, and Directions for Future Research.

Animals have evolved with natural patterns of light and darkness. However, artificial light is being increasingly introduced into the environment from human infrastructure and recreational activity. Artificial light at night (ALAN) has the potential to have widespread effects on animal behavior, physiology, and fitness, which can translate into broader-scale effects on populations and communities. Understanding the effects of ALAN on free-ranging animals is non-trivial due to challenges such as measuring levels of light encountered by mobile organisms and separating the effects of ALAN from those of other anthropogenic disturbance factors. Here we describe an approach that allows us to isolate the effects of artificial light exposure on individual animals by experimentally manipulating light levels inside nest boxes. To this end, a system can be used consisting of light-emitting diode (LED) light(s) adhered to a plate and connected to a battery and timer system. The setup allows exposure of individuals inside nest boxes to varying intensities and durations of ALAN while simultaneously obtaining video recordings, which also include audio. The system has been used in studies on free-ranging great tits (Parus major) and blue tits (Cyanistes caeruleus) to gain insight into how ALAN affects sleep and activity patterns in adults and physiology and telomere dynamics in developing nestlings. The system, or an adaptation thereof, could be used to answer many other intriguing research questions, such as how ALAN interacts with other disturbance factors and affects bioenergetic balance. Furthermore, similar systems could be installed in or near the nest boxes, nests or burrows of a variety of species to manipulate levels of ALAN, evaluate biological responses, and work towards building an interspecific perspective. Especially when combined with other advanced approaches for monitoring the behavior and movement of free-living animals, this approach promises to yield ongoing contributions to our understanding of the biological implications of ALAN.

Serotonin transporter (SERT) polymorphisms, personality and problem-solving in urban great tits.

Understanding underlying genetic variation can elucidate how diversity in behavioral phenotypes evolves and is maintained. Genes in the serotonergic signaling pathway, including the serotonin transporter gene (SERT), are candidates for affecting animal personality, cognition and fitness. In a model species, the great tit (Parus major), we reevaluated previous findings suggesting relationships between SERT polymorphisms, neophobia, exploratory behavior and fitness parameters, and performed a first test of the relationship between single nucleotide polymorphisms (SNPs) in SERT and problem-solving in birds. We found some evidence for associations between SERT SNPs and neophobia, exploratory behavior and laying date. Furthermore, several SNPs were associated with behavioral patterns and success rates during obstacle removal problem-solving tests performed at nest boxes. In females, minor allele homozygotes (AA) for nonsynonymous SNP226 in exon 1 made fewer incorrect attempts and were more likely to problem-solve. In both sexes, there was some evidence that minor allele homozygotes (CC) for SNP84 in exon 9 were more likely to problem-solve. Only one SNP-behavior relationship was statistically significant after correcting for multiple comparisons, but several were associated with substantial effect sizes. Our study provides a foundation for future research on the genetic basis of behavioral and cognitive variation in wild animal populations.

Variable and consistent traffic noise negatively affect the sleep behavior of a free-living songbird.

Anthropogenic noise is a ubiquitous disturbance factor, which, owing to the extensive nature of transportation networks, and ability of sound waves to penetrate distances, has wide-reaching impacts on biological communities. Research effort on biological effects of anthropogenic noise is extensive, but has focused on waking behavior, and to our knowledge, no published experimental study exists on how noise affects sleep in free-living animals. Sleep plays vital functions in processes such as cellular repair and memory consolidation. Thus, understanding the potential for noise to disrupt sleep is a critical research objective. Whether different noise regimes exert distinct effects on behavior also remains poorly understood, as does intraspecific variation in noise sensitivity. To address these knowledge gaps, we used a repeated-measures field experiment involving broad-casting traffic noise recordings at great tit (Parus major) nest boxes over a series of consecutive nights. We evaluated whether increasing the temporal variability and amplitude of traffic noise increased deleterious effects on sleep behavior in free-living great tits, and whether individuals differed in the magnitude of responses. We found that traffic noise reduced sleep duration, proportion, and bout length, and induced birds to exit nest boxes earlier in the morning. There was some support for a stronger effect of more variable noise, and relative to lower amplitude noise, higher amplitude noise resulted in less and more fragmented sleep. Effects of noise on sleep duration were stronger in older adults, and substantial, repeatable variation existed in individual responses. We demonstrate for the first time that anthropogenic noise can have strong effects on sleep in free-living animals, which may have cascading effects on waking behavior, physiology and fitness. Results suggest that reducing the amplitude of traffic noise may be an effective mitigation strategy, and that differences in individual sensitivity are important to consider when evaluating effects of noise exposure. CAPSULE: Experimental exposure to temporally variable and consistent traffic noise negatively affected sleep behavior in a free-living songbird.

The serotonin transporter gene and female personality variation in a free-living passerine.

Quantifying variation in behaviour-related genes provides insight into the evolutionary potential of repeatable among-individual variation in behaviour (i.e. personality). Yet, individuals typically also plastically adjust their behaviour in response to environmental conditions and/or age, thereby complicating the detection of genotype-phenotype associations. Here, using a population of free-living great tits (Parus major), we assessed the association between single nucleotide polymorphisms (SNPs) in the serotonin transporter gene (SERT) and two repeatable behavioural traits, i.e. female-female aggression and female hissing behaviour. For female-female aggression, a trait showing age-related plasticity, we found no evidence for associations with SERT SNPs, even when assessing potential age-dependent effects of SERT genotype on aggression. We also found no strong support for associations between SERT SNPs and hissing behaviour, yet we identified two synonymous polymorphisms (exon 13 SNP66 and exon 12 SNP144) of particular interest, each explaining about 1.3% of the total variation in hissing behaviour. Overall, our results contribute to the general understanding of the biological underpinning of complex behavioural traits and will facilitate further (meta-analytic) research on behaviour-related genes. Moreover, we emphasize that future molecular genetic studies should consider age-dependent genotype-phenotype associations for behavioural trait (co)variation, as this will vastly improve our understanding of the proximate causes and ultimate consequences of personality variation in natural populations.

Little parental response to anthropogenic noise in an urban songbird, but evidence for individual differences in sensitivity.

Anthropogenic noise exposure has well-documented behavioral, physiological and fitness effects on organisms. However, whether different noise regimes evoke distinct responses has rarely been investigated, despite implications for tailoring noise mitigation policies. Urban animals might display low responsiveness to certain anthropogenic noise regimes, especially consistent noise (e.g. freeway noise), but might remain more sensitive to more diverse noise regimes. Additionally, whether individuals differ in noise sensitivity is a rarely explored issue, which is important to fully understand organismal responses to noise. To address these knowledge gaps, we used a field experiment to measure how urban great tits (Parus major) altered parental behaviors in response to two noise regimes: consistent freeway noise, and a diverse anthropogenic noise regime that incorporated variability in noise type and temporal occurrence. We also evaluated whether sex, age, or a well-described personality trait, novel environment exploration behavior, were associated with responses to noise, although our power to assess individual differences in responses was somewhat limited. We found no evidence for mean population-level changes in nestling provisioning behaviors during either noise treatment. However, despite this overall canalization of behavior, there was evidence for individual differences in noise sensitivity, particularly during the diverse noise treatment. Females and birds that explored a novel environment more rapidly (fast explorers) reduced nestling provisioning rate more relative to baseline levels than males and slow explorers during the diverse urban noise, but not during the consistent freeway noise. Furthermore, first year breeders and fast explorers displayed larger increases in latency to return to the nest box relative to baseline conditions during the diverse noise only. Results suggest that urban animal populations might become overall tolerant to anthropogenic noise, but that certain individuals within these populations nonetheless remain sensitive to certain types of noise exposure. CAPSULE: In an urban songbird, we found no population-level changes in nestling provisioning behavior during noise exposure, but did find evidence for individual differences in noise sensitivity.

Sources of individual variation in problem-solving performance in urban great tits (Parus major): Exploring effects of metal pollution, urban disturbance and personality.

Despite growing research effort, we have a limited understanding of how urban disturbance factors affect cognitive traits, such as innovative problem-solving. We performed an initial assessment of how metal pollution and urbanization levels on territories are related to problem-solving performance in urban great tits (Parus major), by presenting an obstacle removal test at nest boxes in three urban nest box populations that are exposed to different levels of metal pollution. We predicted that problem-solving capacity might be reduced within the most polluted population due to pollution-related neurological impairments. On the other hand, we predicted that problem-solving might positively correlate with urban disturbance levels on territories, because some past research suggests that problem-solving promotes persistence in urbanized habitats. We also assessed relationships between exploratory personality type, behavioral patterns during tests, and problem-solving performance, and examined the repeatability and fitness correlates of problem-solving. We found no evidence that behavioral patterns or problem-solving performance were related to metal exposure or exploratory personality, or that problem-solving promotes reproductive success. However, birds on territories exposed to more urban disturbance, as quantified by proximity to paths and roads, were more likely to problem-solve. Moreover, an aggressive problem-solving approach negatively predicted problem-solving success, and behavioral patterns during tests and problem-solving success were repeatable. Thus, rather than indicating negative effects of pollution or urban disturbance on problem-solving performance, our study provides preliminary support for the hypothesis that urbanization favors innovative problem-solving, and suggests that problem-solving could be associated with a personality dimension independent of exploratory personality.

Anthropogenic noise is associated with telomere length and carotenoid-based coloration in free-living nestling songbirds.

Growing evidence suggests that anthropogenic noise has deleterious effects on the behavior and physiology of free-living animals. These effects may be particularly pronounced early in life, when developmental trajectories are sensitive to stressors, yet studies investigating developmental effects of noise exposure in free-living populations remain scarce. To elucidate the effects of noise exposure during development, we examined whether noise exposure is associated with shorter telomeres, duller carotenoid-based coloration and reduced body mass in nestlings of a common urban bird, the great tit (Parus major). We also assessed how the noise environment is related to reproductive success. We obtained long-term measurements of the noise environment, over a ∼24-h period, and characterized both the amplitude (measured by LAeq, LA90, LA10, LAmax) and variance in noise levels, since more stochastic, as well as louder, noise regimes might be more likely to induce stress. In our urban population, noise levels varied substantially, with louder, but less variable, noise characteristic of areas adjacent to a highway. Noise levels were also highly repeatable, suggesting that individuals experience consistent differences in noise exposure. The amplitude of noise near nest boxes was associated with shorter telomeres among smaller, but not larger, brood members. In addition, carotenoid chroma and hue were positively associated with variance in average and maximum noise levels, and average reflectance was negatively associated with variance in background noise. Independent of noise, hue was positively related to telomere length. Nestling mass and reproductive success were unaffected by noise exposure. Results indicate that multiple dimensions of the noise environment, or factors associated with the noise environment, could affect the phenotype of developing organisms, that noise exposure, or correlated variables, might have the strongest effects on sensitive groups of individuals, and that carotenoid hue could serve as a signal of early-life telomere length.

Early-life exposure to artificial light at night elevates physiological stress in free-living songbirds☆.

Artificial light at night (ALAN) can disrupt adaptive patterns of physiology and behavior that promote high fitness, resulting in physiological stress and elevation of steroid glucocorticoids (corticosterone, CORT in birds). Elevated CORT may have particularly profound effects early in life, with the potential for enduring effects that persist into adulthood. Research on the consequences of early-life exposure to ALAN remains limited, especially outside of the laboratory, and whether light exposure affects CORT concentrations in wild nestling birds particularly remains to be elucidated. We used an experimental setup to test the hypothesis that ALAN elevates CORT concentrations in developing free-living birds, by exposing nestling great tits (Parus major) to ALAN inside nest boxes. We measured CORT in feathers grown over the timeframe of the experiment (7 nights), such that CORT concentrations represent an integrative metric of hormone release over the period of nocturnal light exposure, and of development. We also assessed the relationships between feather CORT concentrations, body condition, nestling size rank and fledging success. In addition, we evaluated the relationship between feather CORT concentrations and telomere length. Nestlings exposed to ALAN had higher feather CORT concentrations than control nestlings, and nestlings in poorer body condition and smaller brood members also had higher CORT. On the other hand, telomere length, fledging success, and recruitment rate were not significantly associated with light exposure or feather CORT concentrations. Results indicate that exposure to ALAN elevates CORT concentrations in nestlings, which may reflect physiological stress. In addition, the organizational effects of CORT are known to be substantial. Thus, despite the lack of an effect on telomere length and survivorship, elevated CORT concentrations in nestlings exposed to ALAN may have subsequent impacts on later-life fitness and stress sensitivity.

Carotenoid- but not melanin-based plumage coloration is negatively related to metal exposure and proximity to the road in an urban songbird.

Rapid urbanization is a global phenomenon that is increasingly exposing organisms to novel stressors. These novel stressors can affect diverse aspects of organismal function, including development of condition-dependent ornaments, which play critical roles in social and sexual selection. We investigated the relationship between metal pollution, proximity to roads, and carotenoid- and melanin-based plumage coloration in a common songbird, the great tit (Parus major). We studied populations located across a well-characterized metal pollution gradient and surrounded by roadway networks. Metal exposure and road-associated pollution could reduce carotenoid-based pigmentation by inducing oxidative stress or affecting habitat quality, but metals could also enhance melanin-based pigmentation, through effects on melanogenesis and testosterone concentrations. Using a large sample size (N > 500), we found that birds residing close to a point source for metals had reduced ultraviolet chroma, a component of carotenoid-based pigmentation. Moreover, birds with high feather metal concentrations had lower carotenoid chroma, hue, and ultraviolet chroma, with effects modified by age class. Birds residing closer to roads also had lower carotenoid chroma and hue. Melanin-based pigmentation showed high between-year repeatability, and no association with anthropogenic pollution. Results suggest that carotenoid-, but not melanin-, based pigmentation is negatively affected by multiple anthropogenic stressors. We are the first to demonstrate a negative association between roads and a plumage-based signaling trait, which could have important implications for sexual signaling dynamics in urban landscapes.

An Important Personality Trait Varies with Blood and Plumage Metal Concentrations in a Free-Living Songbird.

Metal pollution is a global problem, which threatens to seriously disrupt behavioral patterns and health in humans and wildlife. Nonetheless, little is known regarding how exposure to metal pollution affects animal personalities, as defined by repeatable among individual differences in behavior. We used a large dataset to investigate the relationship between individual blood and feather metal concentrations and three personality traits (exploration behavior, territorial aggressiveness, and aggressiveness during nest defense) in great tits (Parus major), a model species for animal personality research. We previously demonstrated slower exploration behavior at highly polluted study sites, where exposure to lead, cadmium, and arsenic is high. Here, we demonstrate the across-year repeatability of exploration behavior and aggressiveness during nest defense, providing strong evidence for the existence of personalities in our populations. Furthermore, we demonstrate that individuals with high blood lead concentrations and high concentrations of multiple metals in the feathers exhibit slower exploration behavior but no differences in territorial aggressiveness or nest defense relative to less exposed birds. The mechanism underlying the relationship between metal exposure and exploration behavior remains to be determined but could involve neurotoxic effects. Our study highlights that metal pollution could have underappreciated effects on animal personalities, with implications for individual fitness and societal function.

Nest defence behavioural reaction norms: testing life-history and parental investment theory predictions.

Predation is the primary source of reproductive failure in many avian taxa and nest defence behaviour against predators is hence an important aspect of parental investment. Nest defence is a complex trait that might consistently differ among individuals (personality), while simultaneously vary within individuals (plasticity) according to the reproductive value of the offspring. Both complementary aspects of individual variation can influence fitness, but the causality of links with reproductive success remains poorly understood. We repeatedly tested free-living female great tits (Parus major) for nest defence (hissing) behaviour across the nesting cycle, by presenting them with a model predator. Hissing behaviour was highly repeatable but, despite population-level plasticity, we found no support for individual differences in plasticity. Path analysis revealed that repeatable differences in hissing behaviour had no direct effect on nest success or fledgling number. However, our best supported path-model showed that more fiercely hissing females laid smaller clutches, with clutch size in turn positively influencing fledgling number, suggesting that females are most likely facing a trade-off between investment in nest defence and reproduction. Strong stabilizing selection for optimal plasticity, in combination with life-history trade-offs, might explain the high repeatability of nest defence and its link with reproductive success.

Personality and plasticity in neophobia levels vary with anthropogenic disturbance but not toxic metal exposure in urban great tits: Urban disturbance, metal pollution and neophobia.

Animal personalities, as defined by repeatable among individual differences in behavior, can vary across urbanization gradients. However, how urbanization affects personalities remains incompletely understood, especially because different urban stressors could affect personality traits in opposing ways, whereas most previous studies have considered only one urban disturbance factor. For instance, novel habitat features could favor reduced neophobia, whereas exposure to pollutants could increase risk sensitivity through neurotoxic or hormonal effects. To address this contingency, we studied object neophobia in four urban populations of great tits (Parus major) that vary in exposure to metal pollution and anthropogenic disturbance, as quantified by proximity to roads and pathways. We measured the return latency of incubating females when flushed from the nest and presented with up to two different novel objects, allowing quantification of behavioral repeatability and plasticity. To separate neophobia from sensitivity to disturbance, we also conducted baseline trials, in which females were flushed but no object was presented. We additionally measured exploration behavior and aggression (hissing) during nest defense, to explore whether suites of behaviors covary with urbanization, and examined whether neophobia affects reproductive success. Sensitivity to disturbance and neophobia were repeatable, and thus represent personality traits. Moreover, females occupying territories near roads and pathways had shorter return latencies during novel object but not baseline trials, suggesting a specific reduction in neophobia in disturbed areas. Plasticity in neophobia also increased with disturbance level. In contrast, metal exposure did not affect neophobia or sensitivity to disturbance, despite negatively correlating with exploration behavior. Neophobia correlated with exploration behavior, but not aggression or reproductive success. Results suggest that shifts in personality types in urbanized areas might involve specific reductions in neophobia, rather than general reductions in sensitivity to disturbance, and unexpectedly indicate no effect of toxic metals on risk sensitivity.

Artificial light at night does not affect telomere shortening in a developing free-living songbird: A field experiment: Artificial light at night and telomere dynamics.

Artificial light at night (ALAN) is an increasingly pervasive anthropogenic disturbance factor. ALAN can seriously disrupt physiological systems that follow circadian rhythms, and may be particularly influential early in life, when developmental trajectories are sensitive to stressful conditions. Using great tits (Parus major) as a model species, we experimentally examined how ALAN affects physiological stress in developing nestlings. We used a repeated-measure design to assess effects of ALAN on telomere shortening, body mass, tarsus length and body condition. Telomeres are repetitive nucleotide sequences that protect chromosomes from damage and malfunction. Early-life telomere shortening can be accelerated by environmental stressors, and has been linked to later-life declines in survival and reproduction. We also assayed nitric oxide, as an additional metric of physiological stress, and determined fledging success. Change in body condition between day 8 and 15 differed according to treatment. Nestlings exposed to ALAN displayed a trend towards a decline in condition, whereas control nestlings displayed a trend towards increased condition. This pattern was driven by a greater increase in tarsus length relative to mass in nestlings exposed to ALAN. Nestlings in poorer condition and nestlings that were smaller than their nest mates had shorter telomeres. However, exposure to ALAN was unrelated to telomere shortening, and also had no effect on nitric oxide concentrations or fledging success. Thus, exposure to ALAN may not have led to sufficient stress to induce telomere shortening. Indeed, plasticity in other physiological systems could allow nestlings to maintain telomere length despite moderate stress. Alternatively, the cascade of physiological and behavioral responses associated with light exposure may have no net effect on telomere dynamics.

Personality and artificial light at night in a semi-urban songbird population: No evidence for personality-dependent sampling bias, avoidance or disruptive effects on sleep behaviour.

Light pollution or artificial light at night (ALAN) is an increasing, worldwide challenge that affects many aspects of animal behaviour. Interestingly, the response to ALAN varies widely among individuals within a population and variation in personality (consistent individual differences in behaviour) may be an important factor explaining this variation. Consistent individual differences in exploration behaviour in particular may relate to the response to ALAN, as increasing evidence indicates its relation with how individuals respond to novelty and how they cope with anthropogenic modifications of the environment. Here, we assayed exploration behaviour in a novel environment as a proxy for personality variation in great tits (Parus major). We observed individual sleep behaviour over two consecutive nights, with birds sleeping under natural dark conditions the first night and confronted with ALAN inside the nest box on the second night, representing a modified and novel roosting environment. We examined whether roosting decisions when confronted with a camera (novel object), and subsequently with ALAN, were personality-dependent, as this could potentially create sampling bias. Finally, we assessed whether experimentally challenging individuals with ALAN induced personality-dependent changes in sleep behaviour. Slow and fast explorers were equally likely to roost in a nest box when confronted with either a camera or artificial light inside, indicating the absence of personality-dependent sampling bias or avoidance of exposure to ALAN. Moreover, slow and fast explorers were equally disrupted in their sleep behaviour when challenged with ALAN. Whether other behavioural and physiological effects of ALAN are personality-dependent remains to be determined. Moreover, the sensitivity to disturbance of different behavioural types might depend on the behavioural context and the specific type of challenge in question. In our increasingly urbanized world, determining whether the effects of anthropogenic stressors depend on personality type will be of paramount importance as it may affect population dynamics.

Effects of interspecific coexistence on laying date and clutch size in two closely related species of hole-nesting birds.

Coexistence between great tits Parus major and blue tits Cyanistes caeruleus, but also other hole-nesting taxa, constitutes a classic example of species co-occurrence resulting in potential interference and exploitation competition for food and for breeding and roosting sites. However, the spatial and temporal variations in coexistence and its consequences for competition remain poorly understood. We used an extensive database on reproduction in nest boxes by great and blue tits based on 87 study plots across Europe and Northern Africa during 1957-2012 for a total of 19,075 great tit and 16,729 blue tit clutches to assess correlative evidence for a relationship between laying date and clutch size, respectively, and density consistent with effects of intraspecific and interspecific competition. In an initial set of analyses, we statistically controlled for a suite of site-specific variables. We found evidence for an effect of intraspecific competition on blue tit laying date (later laying at higher density) and clutch size (smaller clutch size at higher density), but no evidence of significant effects of intraspecific competition in great tits, nor effects of interspecific competition for either species. To further control for site-specific variation caused by a range of potentially confounding variables, we compared means and variances in laying date and clutch size of great and blue tits among three categories of difference in density between the two species. We exploited the fact that means and variances are generally positively correlated. If interspecific competition occurs, we predicted a reduction in mean and an increase in variance in clutch size in great tit and blue tit when density of heterospecifics is higher than the density of conspecifics, and for intraspecific competition, this reduction would occur when density of conspecifics is higher than the density of heterospecifics. Such comparisons of temporal patterns of means and variances revealed evidence, for both species, consistent with intraspecific competition and to a smaller extent with interspecific competition. These findings suggest that competition associated with reproductive behaviour between blue and great tits is widespread, but also varies across large spatial and temporal scales.

Cavities shield birds from effects of artificial light at night on sleep.

Light pollution is an ever increasing worldwide problem disrupting animal behavior. Artificial light at night (ALAN) has been shown to affect sleep in wild birds. Even cavity-nesting bird species may be affected when sleeping inside their cavity. Correlational studies suggest that light from outside the cavity/nest box, for example from street lights, may affect sleep. We used an experimental design to study to what extent nest boxes shield animals from effects of ALAN on sleep. We recorded individual sleep behavior of free-living great tits (Parus major) that were roosting in dark nest boxes and exposed their nest box entrance to ALAN the following night (1.6 lux white LED light; a similar light intensity as was found at nest boxes near street lights). Their behavior was compared to that of control birds sleeping in dark nest boxes on both nights. Our experimental treatment did not affect sleep behavior. Sleep behavior of birds in the control group did not differ from that of individuals in the light treated group. Our results suggest that during winter cavities shield birds from some effects of ALAN. Furthermore, given that effects of ALAN and exposure to artificial light are species-, sex-, and season-dependent, it is important that studies using wild animals quantify individual exposure to light pollution, and be cautious in the interpretation and generalization of the effects, or lack thereof, from light pollution. Rigorous studies are necessary to examine individual light exposure and its consequences in cavity- and open-nesting birds.