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.     

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.     

Consistency in Context-specific Measures of Shyness and Boldness in Rainbow Trout, Oncorhynchus mykiss

Shyness and boldness has been considered a fundamental axis of human behavioural variation. At the extreme ends of this behavioural continuum subjects vary from being bold and assertive to shy and timid. Analogous patterns of individual variation have been noted in a number of species including fish. There has been debate on the nature of this continuum as to whether it depends on context. That is, whether it is domain‐general (as in humans), or context‐specific. The purpose of our study was to test if shyness and boldness depends on context in rainbow trout, Oncorhynchus mykiss and to this end we estimated boldness in five different situations. Our data provide evidence of a shy–bold behavioural syndrome in rainbow trout. Bold trout tended to be bold in four situations when the context was similar (when the context concerned foraging). However, in a different context, exploring a swim flume, the ranking was entirely different. We suggest that shyness and boldness depends on context in rainbow trout.     

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.     

Shoal composition determines foraging success in the guppy

The composition of an animal group can impact greatly on thesurvival and success of its individual members. Much recentwork has concentrated on behavioral variation within animalpopulations along the bold/shy continuum. Here, we screenedindividual guppies, Poecilia reticulata, for boldness usingan overhead fright stimulus. We created groups consisting of4 bold individuals (bold shoals), 4 shy individuals (shy shoals),or 2 bold and 2 shy individuals (mixed shoals). The performanceof these different shoal types was then tested in a novel foragingscenario. We found that both bold and mixed shoals approacheda novel feeder in less time than shy shoals. Interestingly,we found that more fish from mixed shoals fed than in eitherbold or shy shoals. We suggest that this can be explained bythe fact that nearly all the cases where one fish was followedinto the feeder by another occurred within mixed shoals andthat it was almost always a shy fish following a bold one. Theseresults suggest clear foraging benefits to shy individuals throughassociating with bold ones. Surprisingly, our results also suggestpotential foraging benefits to bold individuals through associatingwith shy individuals. This study highlights a possible mechanismby which interindividual variation in behavioral types is maintainedin a population.     

Effects of group size and personality on social foraging: the distribution of sheep across patches

Group cohesion in social animals foraging in patchy environmentsdepends on 2 opposite forces, intraspecific competition, andattraction. The decision to leave or stay in a group may varyaccording to the individual personality. We investigated therole of personality when feeding competition increases as aresult of increasing group size. Individuals referred as "bold"and "shy" were identified from an indoor exploration test accordingto their propensity to leave the group to explore a novel environment,using 12 novel objects placed at increasing distances from thegroup. Groups of 2, 4, 6, or 8 shy or bold sheep were testedin 45 x 5–m grass arenas, with one 5 x 5–m patchof preferred vegetation at each end. Sheep grazed on or closeto these patches, but the number grazing the patches seldomreached more than 4–5 individuals, suggesting that crowdingmight affect foraging at the highest densities. The smallestgroups grazed together on the same patch, but there was an increasinglikelihood of splitting into subgroups as group size increased,with equal-sized subgroups most commonly grazing the 2 patchessimultaneously. This effect was greatest in bold sheep, whichtended to split into subgroups at smaller group sizes than shysheep. This study provides new insight into the mechanisms bywhich group-living herbivores distribute themselves across patchyresources in a way that minimizes interference competition anddemonstrates the importance of individual variability for spatialorganization at the level of the group.     

Male personality,life‐history strategies and reproductive success in a promiscuous mammal

Recent theoretical work suggests that personality is a component of life history, but links between personality and either age‐dependent reproductive success or life‐history strategy are yet to be established. Using quantitative genetic analyses on a long‐term pedigree we estimated indices of boldness and docility for 105 bighorn sheep rams (Ovis canadensis), born between 1983 and 1999, and compared these indices to their reproductive history from 2 years of age until death. Docility and boldness were highly heritable and negatively genetically correlated. Docile and bold rams survived longer than indocile and shy rams. Docility and boldness had a weak negative effect on reproductive success early in life, but a strong positive effect on older rams. Our findings highlight an important role of personality on reproductive success and suggest that personality could be an important component of life‐history strategy.     

Individual boldness is linked to protective shell shape in aquatic snails

The existence of consistent individual differences in behaviour (‘animal personality’) has been well documented in recent years. However, how such individual variation in behaviour is maintained over evolutionary time is an ongoing conundrum. A well-studied axis of animal personality is individual variation along a bold–shy continuum, where individuals differ consistently in their propensity to take risks. A predation-risk cost to boldness is often assumed, but also that the reproductive benefits associated with boldness lead to equivalent fitness outcomes between bold and shy individuals over a lifetime. However, an alternative or complementary explanation may be that bold individuals phenotypically compensate for their risky lifestyle to reduce predation costs, for instance by investing in more pronounced morphological defences. Here, we investigate the ‘phenotypic compensation’ hypothesis, i.e. that bold individuals exhibit more pronounced anti-predator defences than shy individuals, by relating shell shape in the aquatic snail Radix balthica to an index of individual boldness. Our analyses find a strong relationship between risk-taking propensity and shell shape in this species, with bolder individuals exhibiting a more defended shell shape than shy individuals. We suggest that this supports the ‘phenotypic compensation’ hypothesis and sheds light on a previously poorly studied mechanism to promote the maintenance of personality variation among animals.     

Differences in aggression,activity and boldness between native and introduced populations of an invasive crayfish

Aggressiveness, along with foraging voracity and boldness, are key behavioral mechanisms underlying the competitive displacement and invasion success of exotic species. However, do aggressiveness, voracity and boldness of the invader depend on the presence of an ecologically similar native competitor in the invaded community? We conducted four behavioral assays to compare aggression, foraging voracity, threat response and boldness to forage under predation risk of multiple populations of exotic signal crayfish Pacifastacus leniusculus across its native and invaded range with and without a native congener, the Shasta crayfish P. fortis. We predicted that signal crayfish from the invaded range and sympatric with a native congener (IRS) should be more aggressive to outcompete a close competitor than populations from the native range (NR) or invaded range and allopatric to a native congener (IRA). Furthermore, we predicted that IRS populations of signal crayfish should be more voracious, but less bold to forage under predation risk since native predators and prey likely possess appropriate behavioral responses to the invader. Contrary to our predictions, results indicated that IRA signal crayfish were more aggressive towards conspecifics and more voracious and active foragers, yet also bolder to forage under predation risk in comparison to NR and IRS populations, which did not differ in behavior. Higher aggression/voracity/boldness was positively correlated with prey consumption rates, and hence potential impacts on prey. We suggest that the positive correlations between aggression/voracity/boldness are the result of an overall aggression syndrome. Results of stream surveys indicated that IRA streams have significantly lower prey biomass than in IRS streams, which may drive invading signal crayfish to be more aggressive/voracious/bold to acquire resources to establish a population.     

An Age-Structured Approach to Modelling Behavioural Variation Maintained by Life-History Trade-Offs

There have been numerous empirical studies on the fitness consequences of behavioural syndromes in various animal taxa; however, the ecological and evolutionary implications on a population level are still poorly understood. To better understand these implications, we develop a non-linear age-structured mathematical model to qualitatively examine the evolutionary consequences of a heritable boldness personality trait within an animal population. We assume that this heritable boldness trait is positively correlated with boldness towards predators and intraspecific aggressiveness. This assumption leads to a growth/reproductive success versus mortality trade-off, which is thoroughly investigated and documented in the literature. Another life-history trade-off we include in the model is future versus current reproduction, which was shown by Wolf et al. [1] to be a possible mechanism for the evolution of behavioural syndromes within an animal population. The stability of the system is analysed, whereby the characteristic equation is in the form of a homogeneous Fredholm equation of the second kind which depends on both the perturbation and equilibrium solution. The system is found to be stable due to the competition between individuals of similar boldness acting as a negative feedback mechanism. Using numerical simulations we examine the qualitative features of the solution to the system. In particular, we investigate the interplay between the mutation and competition strength between two individuals with different boldness, whereby we find that an increasing competition range acts to push individuals to both extremes of the shy-bold axis, while an increasing mutation range counteracts this effect. This qualitative trait of aggregation of individuals around the shy and bold extremes is also found when examining different birth, mortality and competition functions.     

Is larger better in sit-and-wait predators? Competitive superiority in Hydra

Contrary to a generalisation arising from many studies, a larger body size is not always the key to competitive superiority amongst animals. An analysis of competition between pairs of Hydra oligactis, Hydra vulgaris and Hydra circumcincta, that simultaneously encountered a single prey item, showed that competitive success in these sessile predators depended on species and clone in inter- and intraspecific competition, respectively. H. oligactis appeared to be competitively superior, even to the larger H. vulgaris individuals. Phenotypic traits important for prey capture, such as the fraction of the nematocyst that penetrates the prey (penetrants), were positively related to success in intraspecific competition. Body size appeared to be a positive key factor in determining foraging success in the competition between pairs of conspecifics from single or different clones. In contrast to the results of the intraspecific competition, body size was not significantly related to the foraging success of competing heterospecifics.     

Predator and prey: the role of the round goby Neogobius melanostomus in the western Baltic

Different studies on the position of the non-indigenous species Neogobius melanostomus within the coastal food web of the Pomeranian Bay (western Baltic) were performed, resulting in a quantitative and qualitative species list of prey organisms found in the stomachs of the invader and an estimation concerning the importance of round goby as prey for different resident predators. It seems that the colonization process is not fully completed yet, but the results reveal that the species is already established in the food web 16 years after the first observation within the study area. The results show that N. melanostomus feed upon a wide range of different resident organisms. While a direct predation effect on native fish species appears rather unlikely, indirect effects such as competition cannot yet be excluded. In addition, our results reveal an ontogenetic diet shift and that the round goby itself already serves as an important prey for piscivorous fish and seabirds. Finally, we formulate different hypotheses based on our results which will require further research.     

Express yourself: bold individuals induce enhanced morphological defences

Organisms display an impressive array of defence strategies in nature. Inducible defences (changes in morphology and/or behaviour within a prey''s lifetime) allow prey to decrease vulnerability to predators and avoid unnecessary costs of expression. Many studies report considerable interindividual variation in the degree to which inducible defences are expressed, yet what underlies this variation is poorly understood. Here, we show that individuals differing in a key personality trait also differ in the magnitude of morphological defence expression. Crucian carp showing risky behaviours (bold individuals) expressed a significantly greater morphological defence response when exposed to a natural enemy when compared with shy individuals. Furthermore, we show that fish of different personality types differ in their behavioural plasticity, with shy fish exhibiting greater absolute plasticity than bold fish. Our data suggest that individuals with bold personalities may be able to compensate for their risk-prone behavioural type by expressing enhanced morphological defences.     

Link between Aggressiveness and Shyness in the Spider <Emphasis Type="Italic">Philodromus albidus</Emphasis> (Araneae,Philodromidae): State Dependency over Stability

Behavioral syndromes, seen as correlations among two or more functionally different behaviors, are well documented in many different animal taxa. They may not be present automatically, however, and their consistency within populations and individuals also varies among studies. Here, we studied a behavioral syndrome comprising foraging aggressiveness and boldness/shyness and its time consistency in the cursorial spider Philodromus albidus. We measured foraging aggressiveness as the number of prey killed per 2-hour period. Boldness/shyness was assessed as a latency of initiating exploration in a novel environment. We found the presence of the behavioral syndrome in P. albidus, as bold individuals were also more aggressive than were shy individuals. The syndrome was consistent through time within the population but not consistent for individuals. We further discuss the possible causes of the pattern.     

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.     

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.     

Dietary niche partitioning in sympatric gadid species in coastal Newfoundland: evidence from stomachs and C-N isotopes

The feeding habits of co-occurring gadid species Atlantic cod (Gadus morhua) and Greenland cod (Gadus ogac) in coastal Newfoundland waters, examined using stable isotope (δ ¹³C and δ ¹⁵N) and stomach content analysis, indicated little dietary niche overlap and interspecific competition for food resources despite similar trophic levels. Both species consumed a variety of invertebrates and fish but showed a preference for different prey items. Polychaetes, fish and small crustaceans dominated G. ogac stomach contents while small crustaceans, in particular hyperiid amphipods and fish, dominated those of G. morhua. In general, G. morhua consumed more pelagic prey and had a significantly more pelagic (more negative) δ ¹³C signature while G. ogac consumed primarily benthic prey and had a more benthic (more positive) δ ¹³C signature. δ ¹⁵N levels were similar in these species suggesting similar trophic positions, with levels increasing with fish length in both species. Dietary overlap was not significant in both stomach and stable isotope analyses. We conclude that interspecific competition for food is low between G. ogac and G. morhua and is unlikely to be a factor in the slow rebuilding of Atlantic cod in this region.     

ASPECTS OF THE FORAGING BEHAVIOUR OF THE SHOEBILL

Guillet, A. 1979. Aspects of the foraging behaviour of the Shoebill. Ostrich 50:252-255.

The foraging behaviour of the Shoebill Balaeniceps rex. in relation to the bird's habitat and morphology, is described. The Shoebill preys on fish in shallow water, and uses platforms of floating vegetation as fishing sites. The Shoebill's behaviour in stalking, detecting and capturing prey is compared with foraging techniques used by herons and storks. The Shoebill uses a peculiar and complicated technique, called “collapsing”, for capturing prey.     

Field study of the relationship between personality and reproductive strategy in the White-collared Blackbird Turdus albocinctus

The discovery of personality traits in animal populations may help to explain individual variation in breeding strategies. In the White-collared Blackbird Turdus albocinctus, females, but not males, exhibited different nest defence behaviours that can be used to classify them into bold and shy personalities. Bold females had higher nest success, lower fecundity and higher parental investment than shy females, and the social mates of bold females had lower parental investment than that of shy females. Our findings suggest that the reproductive strategy of birds may be closely related to their personality traits.