Turning Shy on a Winter's Day: Effects of Season on Personality and Stress Response in Microtus arvalis

Animal personalities are by definition stable over time, but to what extent they may change during development and in adulthood to adjust to environmental change is unclear. Animals of temperate environments have evolved physiological and behavioural adaptations to cope with the cyclic seasonal changes. This may also result in changes in personality: suites of behavioural and physiological traits that vary consistently among individuals. Winter, typically the adverse season challenging survival, may require individuals to have shy/cautious personality, whereas during summer, energetically favourable to reproduction, individuals may benefit from a bold/risk‐taking personality. To test the effects of seasonal changes in early life and in adulthood on behaviours (activity, exploration and anxiety), body mass and stress response, we manipulated the photoperiod and quality of food in two experiments to simulate the conditions of winter and summer. We used the common voles (Microtus arvalis) as they have been shown to display personality based on behavioural consistency over time and contexts. Summer‐born voles allocated to winter conditions at weaning had lower body mass, a higher corticosterone increase after stress and a less active, more cautious behavioural phenotype in adulthood compared to voles born in and allocated to summer conditions. In contrast, adult females only showed plasticity in stress‐induced corticosterone levels, which were higher in the animals that were transferred to the winter conditions than to those staying in summer conditions. These results suggest a sensitive period for season‐related behavioural plasticity in which juveniles shift over the bold–shy axis.     

The non-consumptive effects of predators and personality on prey growth and mortality

Individual organisms vary in personality, and the ecological consequences of that variation can affect the strength of predator–prey interactions. Prey with bolder tendencies can mitigate the strength of species interactions by altering growth and initiating ontogenetic niche shifts (ONS). While the link between personality and growth has been established, recent research has highlighted the important interplay between ONS and predator cues in community ecology. The objective of this study was to evaluate the effects of prey personality and predator cues on prey growth and ONS. We predicted growth–mortality trade-offs among personalities with higher survival, larger size, and accelerated ONS for bold individuals in comparison with shy individuals. To evaluate this objective, we conducted behavioral assays and a mesocosm experiment to test how southern leopard frog (Rana sphenocephala) tadpole personality and predatory fish (bluegill, Lepomis macrochirus) cues affects tadpole growth and metamorphosis. On average, bold tadpoles had higher mortality across all treatments in comparison with shy tadpoles. The effects of fish cues were dependent on tadpole personality with shy tadpoles metamorphosing significantly later than bold tadpoles. Bold tadpoles were larger than shy tadpoles at metamorphosis; however, that pattern reversed with fish cues as shy individuals metamorphosed larger than bold individuals. Our results suggest personality may be useful for predicting growth and life history for some prey species with predators. Specifically, the threat of predation can interact with personality to incur a benefit (earlier ONS) while also incurring a cost (size at metamorphosis). Hence by incorporating predator cues with personality, ecologists will be able to elucidate growth–mortality trade-offs mediated by personality.     

Shy birds play it safe: personality in captivity predicts risk responsiveness during reproduction in the wild

Despite a growing body of evidence linking personality to life-history variation and fitness, the behavioural mechanisms underlying these relationships remain poorly understood. One mechanism thought to play a key role is how individuals respond to risk. Relatively reactive and proactive (or shy and bold) personality types are expected to differ in how they manage the inherent trade-off between productivity and survival, with bold individuals being more risk-prone with lower survival probability, and shy individuals adopting a more risk-averse strategy. In the great tit (Parus major), the shy–bold personality axis has been well characterized in captivity and linked to fitness. Here, we tested whether ‘exploration behaviour’, a captive assay of the shy–bold axis, can predict risk responsiveness during reproduction in wild great tits. Relatively slow-exploring (shy) females took longer than fast-exploring (bold) birds to resume incubation after a novel object, representing an unknown threat, was attached to their nest-box, with some shy individuals not returning within the 40 min trial period. Risk responsiveness was consistent within individuals over days. These findings provide rare, field-based experimental evidence that shy individuals prioritize survival over reproductive investment, supporting the hypothesis that personality reflects life-history variation through links with risk responsiveness.     

Does it always pay to defend one's nest? A case study in African penguin

Throughout the animal kingdom, individual variation in reproductive success is commonly observed, even under similar environmental conditions. However, the mechanisms behind such differences remain unclear. The notion of behavioural consistency in animals has developed rapidly since the early 21st century partly as an approach to understand among‐individual differences. In this context, a number of studies have highlighted the influence of pair assortment in personality on breeding success. In this study, we related breeding success to individual behaviour, specifically a risk‐taking behaviour, and pair assortment per behaviour in African penguins (Spheniscus demersus) over two breeding seasons of contrasting food availability. On Bird Island, Algoa Bay, South Africa, we used indices of boldness and overall mobility in penguins’ nest defence behaviour as a response to a standard pedestrian approach during chick‐rearing. These behaviours were consistent over the trials and indicated these traits may be related to personality in African penguins. Individuals were categorized as risk‐prone (“bold,” “mobile”) or risk‐averse (“shy,” “non‐mobile”). We then assessed their breeding success through chick growth and survival over 4 weeks in 2015 and 2016. There was weak positive assortment of pairs in relation to nest defence behaviour. However, pair assortment did not significantly influence birds’ breeding success. Shy penguins were generally the most successful (had the highest chick growth rates), which was especially apparent during a food shortage in 2016, possibly reflecting a higher energy investment when foraging. In contrast, chicks from bold parents grew significantly slower, especially in 2016. Bold parents may defend their nest successfully against predation or intra‐specific aggression when food is abundant, but when predation risks are limited and food availability is low, this strategy may not be beneficial. In the context of climate change, where food shortage events may become more frequent, risk‐averse individuals may be favoured and genetic diversity may be reduced in African penguins.     

Personality composition is more important than group size in determining collective foraging behaviour in the wild

Describing the factors that shape collective behaviour is central to our understanding of animal societies. Countless studies have demonstrated an effect of group size in the emergence of collective behaviours, but comparatively few have accounted for the composition/diversity of behavioural phenotypes, which is often conflated with group size. Here, we simultaneously examine the effect of personality composition and group size on nest architecture and collective foraging aggressiveness in the social spider Stegodyphus dumicola. We created colonies of two different sizes (10 or 30 individuals) and four compositions of boldness (all bold, all shy, mixed bold and shy, or average individuals) in the field and then measured their collective behaviour. Larger colonies produced bigger capture webs, while colonies containing a higher proportion of bold individuals responded to and attacked prey more rapidly. The number of attackers during collective foraging was determined jointly by composition and size, although composition had an effect size more than twice that of colony size: our results suggest that colonies of just 10 bold spiders would attack prey with as many attackers as colonies of 110 ‘average’ spiders. Thus, personality composition is a more potent (albeit more cryptic) determinant of collective foraging in these societies.     

Behavioural biology: fortune favours bold and shy personalities

A new study has shown that in the great tit (Parus major) bold males and shy females apparently flourish after rich winter pickings, while shy males and bold females profit from meagre winters. This groundbreaking work exemplifies the approach required for a biological understanding of an apparently common animal trait - personality.     

Correlation between boldness and body mass in natural populations of the poeciliid Brachyrhaphis episcopi

The boldness of individual Brachyrhaphis episcopi , collected from regions of high and low predation, was investigated using two independent assays: (1) the time to emerge from cover and (2) the propensity to leave shoal mates and investigate a novel object. A strong correlation between the two assays was revealed such that fish that emerged from shelter sooner were also more likely to approach a novel object. This is indicative of a boldness personality axis acting across both behavioural contexts. Fish from high-predation areas were bolder than those from low-predation areas and males were bolder than females. A significant correlation between body mass, standard length ( L _S) and boldness score was also found. In general, bold fish had a greater body mass at a given L _S than shy fish. These results suggest that personality traits are strongly influenced by population-specific ecological variables and may have fitness consequences in wild populations.     

Elevating perceived predation risk modifies the relationship between parental effort and song complexity in the song sparrow Melospiza melodia

Adult‐directed predation risk elevates costs of parental care, and may modify relationships between sexually selected ornaments and parental effort by accentuating the tradeoff between survival and parental investment. We assessed multiple hypotheses regarding the relationship between maternal effort, paternal effort, and the sexually selected trait of male song complexity in the song sparrow Melospiza melodia. Further, we explored whether experimentally elevating perceived adult‐directed predation risk near nests affected these relationships. We quantified two dimensions of song complexity: song repertoire size and residual syllable number (the relative number of syllables for a given song repertoire size). Under elevated perceived predation risk, but not in the absence of the predator stimuli, females mated to males with higher residual syllable number displayed higher nestling provisioning rates and performed a greater proportion of nestling provisioning trips. In other words, elevating perceived predation risk induced a pattern of maternal investment consistent with differential allocation. In contrast, under elevated perceived predation risk, only, females performed a lesser proportion of provisioning trips when mated to males with large song repertoire sizes. Further, consistent with the good parent hypothesis, males with large song repertoire sizes displayed lower latencies to return to the nest, independent of the predator stimuli. Results suggest that residual syllable number may reflect some aspect of male genetic quality, such that females are more willing to maintain maternal effort while facing heightened predation risk. On the other hand, females may gain paternal benefits when mated to males with large song repertoires. Our study supports the hypothesis that increased costs of parental care associated with predation risk may induce relationships between sexually selected traits and parental behavior, which may increase the strength of sexual selection. Additionally, results suggest that different aspects of song complexity may fulfill non‐equivalent signaling roles.     

Boldness and Stress Responsiveness as Drivers of Nest‐Site Selection in a Ground‐Nesting Bird

Nest‐site selection involves trade‐offs between the probability of nest discovery by egg predators, danger to the incubating parent from predators and provision of an appropriate microclimate for incubation, and all three components are potentially influenced by nest concealment and distance to habitat edge. Personality traits also affect habitat choice, and the trade‐off hypothesis suggests that variable stress‐coping strategies may evolve because individuals prioritize either productivity (proactive coping) or survival (reactive coping), both of which cannot be simultaneously maximized. Applying this hypothesis to understand nest‐site selection by female eider ducks (Somateria mollissima), we predicted that bold individuals with attenuated stress responsiveness should select concealed nests further away from the shore, thereby promoting clutch survival at the potential expense of reduced escape opportunities for themselves. In testing this prediction, we controlled for individual quality (female body condition and breeding experience) and verified that individual stress responses and nest‐site preferences were consistent by analysing their repeatability. Finally, we analysed how the viable proportion of the clutch was related to nest‐site and female characteristics. Our prediction was supported: bold females [short flight initiation distance (FID)] and those with lower handling‐induced body temperature were found in concealed nests, and bold females and those with lower handling‐induced corticosterone concentrations occupied nests farther from the shore. The viable proportion of the clutch peaked at intermediate proportional nest‐cover and increased linearly with increasing nest distance from the shore. Stress coping styles may thus be related to nest‐site selection, but their fitness consequences may be manifested indirectly through the nest‐site characteristics.     

Personality-dependent inter- and intraspecific foraging competition in the invasive round goby,Neogobius melanostomus

This study examines the impact of boldness on foraging competition of the highly invasive round goby Neogobius melanostomus Pallas 1815. Individual risk tolerance, or boldness, was measured as the time to resume movement after a simulated predation strike. Fish that resumed movement faster were categorized as “bold,” fish that took more time to resume movement were categorized as “shy” and those that fell in between these two categories were determined to have “intermediate” boldness. Competitive impacts of boldness in N. melanostomus were determined in a laboratory foraging experiment in which interspecific (juvenile Atlantic cod Gadus morhua Linnaeus 1758) and intraspecific (intermediate N. melanostomus) individuals were exposed to either bold or shy N. melanostomus competitors. G. morhua consumed fewer prey when competing with bold N. melanostomus than when competing with shy N. melanostomus, whereas intermediately bold N. melanostomus foraging was not affected by competitor boldness. Bold and shy N. melanostomus consumed similar amounts of prey, and the number of interactions between paired fish did not vary depending on the personality of N. melanostomus individuals. Therefore, intraspecific foraging competition was not found to be personality dependent. This study provides evidence that individual differences in boldness can mediate competitive interactions in N. melanostomus; nonetheless, results also show that competition is also governed by other mechanisms that require further study.     

A Selfish Genetic Element Influencing Longevity Correlates with Reactive Behavioural Traits in Female House Mice (Mus domesticus)

According to theory in life-history and animal personality, individuals with high fitness expectations should be risk-averse, while individuals with low fitness expectations should be more bold. In female house mice, a selfish genetic element, the t haplotype, is associated with increased longevity under natural conditions, representing an appropriate case study to investigate this recent theory empirically. Following theory, females heterozygous for the t haplotype (+/t) are hypothesised to express more reactive personality traits and be more shy, less explorative and less active compared to the shorter-lived homozygous wildtype females (+/+). As males of different haplotype do not differ in survival, no similar pattern is expected. We tested these predictions by quantifying boldness, exploration, activity, and energetic intake in both +/t and +/+ mice. +/t females, unlike +/+ ones, expressed some reactive-like personality traits: +/t females were less active, less prone to form an exploratory routine and tended to ingest less food. Taken together these results suggest that differences in animal personality may contribute to the survival advantage observed in +/t females but fail to provide full empirical support for recent theory.     

Birds reveal their personality when singing

Background

Individual differences in social behaviour may have consequences for mate choice and sexual signalling, because partners should develop preferences for personalities that maximize reproductive output. Here we propose that behavioural traits involved in sexual advertisement may serve as good indicators of personality, which is fundamental for sexual selection to operate on temperament. Bird song has a prominent and well-established role in sexual selection, and it displays considerable variation among individuals with a potentially strong personality component. Therefore, we predicted that features of song would correlate with estimates of personality.

Methodology/Principal Findings

In a field study of free-living male collared flycatchers, Ficedula albicollis, we characterised personality based on the exploration of an altered breeding environment, and based on the risk taken when a potential predator was approaching during a simulated territorial interaction. We found that explorative and risk-taker individuals consistently sang at lower song posts than shy individuals in the presence of a human observer. Moreover, males from lower posts established pair-bonds relatively faster than males from higher posts.

Conclusions/Significance

Our results may demonstrate that risk taking during singing correlates with risk taking during aggression and with exploration, thus personality may be manifested in different contexts involving sexual advertisement. These findings are in accordance with the hypothesis that the male''s balance between investment in reproduction and risk taking is reflected in sexual displays, and it may be important information for choosy females that seek partners with personality traits enhancing breeding success.     

Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor

The sexes’ share in parental care and the social mating system in a marked population of the single‐brooded Lesser Spotted Woodpecker Dendrocopos minor were studied in 17 woodpecker territories in southern Sweden during 10 years. The birds showed a very strong mate fidelity between years; the divorce rate was 3.4%. In monogamous pairs, the male provided more parental care than the female. The male did most of the nest building and all incubation and brooding at night. Daytime incubation and brooding were shared equally by the sexes, and biparental care at these early breeding stages is probably necessary for successful breeding. In 42% of the nests, however, though still alive the female deserted the brood the last week of the nestling period, whereas the male invariably fed until fledging and fully compensated for the absent female. Post‐fledging care could not be quantified, but was likely shared by both parents. Females who ceased feeding at the late nestling stage resumed care after fledging. We argue that the high premium on breeding with the same mate for consecutive years and the overall lower survival of females have shaped this male‐biased organisation of parental care. In the six years with best data, most social matings were monogamous, but 8.5% of the females (N=59) exhibited simultaneous multi‐nest (classical) polyandry and 2.9% of the males (N=68) exhibited multi‐nest polygyny. Polyandrous females raised 39% more young than monogamous pairs. These females invested equal amounts of parental care at all their nests, but their investment at each nest was lower than that of monogamous females. The polyandrously mated males fully compensated for this lower female investment. Polygynous males invested mainly in their primary nest and appeared to be less successful than polyandrous females. Polyandry and polygyny occurred only when the population sex ratio was biased, and due to strong intra‐sexual competition this is likely a prerequisite for polygamous mating in Lesser Spotted Woodpeckers.     

Mate Choice and Spawning Success in the Fighting Fish Betta splendens: the Importance of Body Size, Display Behavior and Nest Size

Courtship displays should be exaggerated enough to attract mates and yet tempered so as not to deter them. We tested this hypothesis in the fighting fish Betta splendens by studying courtship displays and body size and their relationships with male parental quality and female fecundity, as well as the effects of display behavior and body size on mate choice decisions and spawning success. Because of their high degree of parental investment, males are expected to be discriminating in their choice of mates. Males who displayed more frequently built larger nests, a measure of parental quality, but larger males did not. When females were paired with males with high display rates, however, the pair had fewer eggs in their nest, even when accounting for female body mass. In a mate choice test using computer‐generated male stimuli that differed only in display behavior, females showed no preferences for displaying males vs. non‐displaying males, or for males with higher display rates vs. lower display rates. In similar tests in which the computer‐generated males differed only in size, females preferred larger males, but also preferred males that differed with respect to body size (negative assortative mating). Males preferred computer‐generated females that performed courtship displays over non‐displaying females, but showed no preferences for female body size. Neither a female's body size nor her display behavior was a significant predictor of her fecundity as estimated by the number of eggs released during spawning. Thus, our results suggest that female B. splendens must balance male parental quality (nest size) with the risk of potentially disruptive or dangerous behavior during spawning, and that females may minimize these risks through negative size‐assortative mating. Female display behavior, while unrelated to fecundity in our study, may attract males because it indicates reproductive readiness or serves a species‐recognition function.     

Relationship between reversed sexual dimorphism,breeding investment and foraging ecology in a pelagic seabird,the masked booby

Reversed sexual dimorphism (RSD) may be related to different roles in breeding investment and/or foraging, but little information is available on foraging ecology. We studied the foraging behaviour and parental investment by male and female masked boobies, a species with RSD, by combining studies of foraging ecology using miniaturised activity and GPS data loggers of nest attendance, with an experimental study where flight costs were increased. Males attended the chick more often than females, but females provided more food to the chick than males. Males and females foraged during similar periods of the day, had similar prey types and sizes, diving depths, durations of foraging trips, foraging zones and ranges. Females spent a smaller proportion of the foraging trip sitting on the water and had higher diving rate than males, suggesting higher foraging effort by females. In females, trip duration correlated with mass at departure, suggesting a flexible investment through control by body mass. The experimental study showed that handicapped females and female partners of handicapped males lost mass compared to control birds, whereas there was no difference for males. These results indicate that the larger female is the main provisioner of the chick in the pair, and regulates breeding effort in relation to its own body mass, whereas males have a fixed investment. The different breeding investment between the sexes is associated with contrasting foraging strategies, but no clear niche differentiation was observed. The larger size of the females may be advantageous for provisioning the chick with large quantities of energy and for flexible breeding effort, while the smaller male invests in territory defence and nest guarding, a crucial task when breeding at high densities. In masked boobies, division of labour appears to be maximal during chick rearing—the most energy-demanding period—and may be related to evolution of RSD.     

Human Perceptions of Coat Color as an Indicator of Domestic Cat Personality

ABSTRACT

Associations between mammalian coat color and behavior have been investigated in a number of species, most notably the study of silver foxes by the Institute of Cytology and Genetics at the Russian Academy of Sciences. However, the few studies conducted regarding a potential relation between coat color and domestic cat personality have shown mixed results, even though many people believe that differently colored cats have distinct personalities. Understanding how humans might perceive personality in relation to coat color may have important ramifications regarding whether cats are relinquished to shelters or adopted from them. In order to assess human perceptions of differently colored cats, we conducted an anonymous, online survey, using a 7-point Likert scale and 10 terms describing personality traits that were chosen based on previous studies of animal personality. This survey examined how people assigned these given terms (active, aloof, bold, calm, friendly, intolerant, shy, stubborn, tolerant, and trainable) to five different colors of cats (orange, tricolored, white, black, and bi-colored). There were significant differences in how participants in this study chose to assign personality terms to differently colored cats. For example, participants (n = 189) were more likely to attribute the trait “friendliness” to orange cats, “intolerance” to tri-colored cats, and “aloofness” to white and tri-colored cats. No significant differences were found for “stubbornness” in any colors of cats. White cats were seen as less bold and active and more shy and calm than other colors of cats. While survey respondents stated that they placed more importance on personality than color when selecting a companion cat, there is some evidence that they believe the two qualities are linked. We anticipate our findings will be relevant to further study in domestic cat personality and to those who work in animal rescue, particularly in how shelters promote differently colored cats and educate potential adopters.     

Reproductive behavior of the Amazonian dwarf cichlid Apistogramma hippolytae Kullander, 1982: offsetting costs and benefits

The reproduction trade-off for an animal is a conflicting choice in which resources (e.g., time and/or energy) allocated to one reproduction trait (e.g., parental care) become unavailable to other traits (e.g., future reproduction events). Here, we tested three hypotheses related to the parental care of the Amazonian dwarf cichlid Apistogramma hippolytae in its natural habitat of Central Amazonia: (1) brood-caring females have a lower feeding frequency than individuals that are not involved in this behavior; (2) females that spend more time on nest defense have lower feeding rates; and (3) females can recognize the species that present the greatest danger to its offspring and move farther from the nest to chase away these piscivorous fishes. We also described for the first time the reproductive behavior (including courtship) and parental care of this species. The results showed that maternal care produces a reduction in the rate of feeding of mothers, a greater amount of time is spent chasing invaders away from the nest, and reproductive females are able to distinguish species-specific predators. These observations support the hypotheses of this study and also suggest a trade-off between current and future reproduction events.     

The Association Between Personality Traits,Morphological Traits and Alternative Mating Behaviour in Male Endler's Guppies,Poecilia wingei

Alternative mating behaviour, personality traits and morphological characters are predicted to be correlated. Bolder, larger and more colourful males are expected to preferentially court females, while shy, small and drab‐coloured individuals are predicted to sneak copulations. We used males of Endler's guppy, Poecilia wingei, to test this association over a long temporal period (hence including ontogenetic changes) and under two social environments (male‐biased and female‐biased). We found that personality traits (exploration, boldness, activity) of P. wingei males were highly repeatable across long time spans, but they were not correlated (formed no behavioural syndrome). Male age and social environment had no effect on any personality trait, despite their effects on alternative mating behaviour. Young males with higher activity levels were more likely to attempt sneaking. In older fish, there was an association between orange coloration, courtship and boldness, but this was not observed in young males. Our results suggest that alternative mating behaviour is more flexible than personality traits and is independent of them. Non‐colour‐based morphological traits (gonopodium length, body length, caudal straps length, dorsal fin length) were not correlated with any particular mating behaviour.     

Plasticity in animal personality traits: does prior experience alter the degree of boldness?

Theoreticians predict that animal 'personality' traits may be maladaptive if fixed throughout different contexts, so the present study aimed to test whether these traits are fixed or plastic. Rainbow trout (Onchorhyncus mykiss) were given emboldening or negative experiences in the forms of watching bold or shy individuals responding to novelty or winning or losing fights to examine whether prior experience affected boldness. Bold individuals that lost fights or watched shy demonstrators became more shy by increasing their latency to approach a novel object, whereas shy observers that watched bold demonstrators remained cautious and did not modify their responses to novelty. Shy winners became bolder and decreased their latency to approach a novel object, but shy losers also displayed this shift. In comparison, control groups showed no change in behaviour. Bold fishes given negative experiences reduced their boldness which may be an adaptive response; however, shy fishes may base their strategic decisions upon self-assessment of their relative competitive ability and increase their boldness in situations where getting to resources more quickly ensures they outcompete better competitors.     

Individual variation in behavioural responsiveness to humans leads to differences in breeding success and long‐term population phenotypic changes

Whether human disturbance can lead to directional selection and phenotypic change in behaviour in species with limited behavioural plasticity is poorly understood in wild animal populations. Using a 19‐year study on Montagu′s harrier, we report a long‐term increase in boldness towards humans during nest visits. The probability of females fleeing or being passive during nest visits decreased, while defence intensity steadily increased over the study period. These behavioural responses towards humans were significantly repeatable. The phenotypic composition of the breeding population changed throughout the study period (4–5 harrier generations), with a gradual disappearance of shy individuals, leading to a greater proportion of bolder ones and a more behaviourally homogeneous population. We further show that nest visit frequency increased nest failure probability and reduced productivity of shy females, but not of bold ones. Long‐term research or conservation programmes needing nest visits can therefore lead to subtle but relevant population compositional changes that require further attention.