Sea anemones (Actinia equina) show consistent individual differences in boldness and thoroughness but lack a behavioural syndrome.

Behavioural syndromes are suites of behaviours that corelate between-individuals but the same behaviours may also show within-individual correlations owing to state dependency or trade-offs. Therefore, overall phenotypic behavioural correlations must be separated into their between- and within-individual components. We investigate how startle response duration (an index of boldness) and time taken to reject an inert item (an index of investigation thoroughness) covary in beadlet sea anemones, Actinia equina. Anemones took longer to reject a more complex item compared to a simpler one, validating this measure of investigation thoroughness. We then quantified between- and within-individual correlations using a Bayesian analysis and an alternative frequentist analysis, which returned the same results. Startle responses decreased with anemone size while thoroughness decreased across repeated observations, indicative of simple learning. For each behaviour, repeatability was significant but relatively low and there was no behavioural syndrome. Rather, the two behaviours showed a negative within-individual correlation in most individuals. Thus, boldness and thoroughness are unlikely to be under correlative selection, and they may instead be expressed independently, in line with the general pattern that cross-contextual behavioural syndromes are comparatively rare. It now appears that this pattern may extend broadly across animal diversity.

Boldness is not associated with dynamic performance capacity in hermit crabs.

Boldness, the way an individual reacts to risk, is a commonly studied personality trait in animals. Consistent among-individual differences in startle response durations (latency to recover from a startling stimulus) are frequently assumed to reflect variation in boldness. An alternative explanation is that these latencies are not directly driven by variation in responses to information on risk, but by underlying differences in dynamic performance capacities. Here we investigate this possibility by analysing relationships between locomotory speed, a measure of whole-body dynamic performance capacity in hermit crabs, and startle response duration, a repeatable latency measure used as an index of boldness. Individuals differed in mean startle response duration, in the consistency of their startle responses, in their reaction norms across repeated observations, and mean startle responses increased with crab mass. However, there were no relationships between startle responses and locomotory speed. This indicates that startle responses do not reflect underlying performance capacities and suggests that they provide insight into differences in how individuals respond to risky situations. Since similar latencies are used as measures of boldness in other animals, we suggest that potential relationships between apparent boldness and performance capacity should be tested.

When should we ascribe sentience to animals? A commentary on "Hermit crabs, shells and sentience" (Elwood 2022).

In a new review article, experiments on hermit crab behaviour are discussed in the context of possible animal sentience. Sentience can be defined as the ability to experience feelings such as pleasure or pain but there are also broader definitions that include elements of awareness. Here I suggest that of the different levels of awareness described as components of sentience, only the higher levels (assessment and executive awareness) seem distinct from the basic non-sentient cognitive tasks of gathering, processing and storing information, which are demonstrated by most animals. Studies that attempt to differentiate between basic cognitive functions and higher levels of awareness are rare for most animal taxa, including hermit crabs. Therefore, while results such as those obtained from studies of hermit crab behaviour are compatible with sentience they cannot yet be distinguished from simpler explanations based on basic cognitive functions, which we should prefer for the time-being. Nevertheless, hermit crabs are promising model systems for investigating awareness in animals.

Male song stability shows cross-year repeatability but does not affect reproductive success in a wild passerine bird.

Predictable behaviour (or 'behavioural stability') might be favoured in certain ecological contexts, for example when representing a quality signal. Costs associated with producing stable phenotypes imply selection should favour plasticity in stability when beneficial. Repeatable among-individual differences in degree of stability are simultaneously expected if individuals differ in ability to pay these costs, or in how they resolve cost-benefit trade-offs. Bird song represents a prime example, where stability may be costly yet beneficial when stable singing is a quality signal favoured by sexual selection. Assuming energetic costs, ecological variation (e.g. in food availability) should result in both within- and among-individual variation in stability. If song stability represents a quality signal, we expect directional selection favouring stable singers. For a 3-year period, we monitored 12 nest box plots of great tits Parus major during breeding. We recorded male songs during simulated territory intrusions, twice during their mate's laying stage and twice during incubation. Each preceding winter, we manipulated food availability. Assuming that stability is costly, we expected food-supplemented males to sing more stable songs. We also expected males to sing more stable songs early in the breeding season (when paternity is not decided) and stable singers to have increased reproductive success. We found strong support for plasticity in stability for two key song characteristics: minimum frequency and phrase length. Males were plastic because they became more stable over the season, contrary to expectations. Food supplementation did not affect body condition but increased stability in minimum frequency. This treatment effect occurred only in 1 year, implying that food supplementation affected stability only in interaction with (unknown) year-specific ecological factors. We found no support for directional, correlational or fluctuating selection on the stability in minimum frequency (i.e. the song trait whose stability exhibited cross-year repeatability): stable singers did not have higher reproductive success. Our findings imply that stability in minimum frequency is not a fitness quality indicator unless males enjoy fitness benefits via pathways not studied here. Future studies should thus address the mechanisms shaping and maintaining individual repeatability of song stability in the wild.

Perceived and actual fighting ability: determinants of success by decision, knockout or submission in human combat sports.

Animal contest theory assumes individuals to possess accurate information about their own fighting ability or resource-holding potential (RHP) and, under some models, that of their opponent. However, owing to the difficulty of disentangling perceived and actual RHP in animals, how accurately individuals are able to assess RHP remains relatively unknown. Furthermore, it is not just individuals within a fight that evaluate RHP. Third-party observers evaluate the fight performance of conspecifics in order to make behavioural decisions. In human combat sports, when fights remain unresolved at the end of the allotted time, bystanders take a more active role, with judges assigning victory based on their assessment of each fighter's performance. Here, we use fight data from mixed martial arts in order to investigate whether perceived fighting performance (judges' decisions) and actual fighting success (fights ending in knockout or submission) are based on the same performance traits, specifically striking skill and vigour. Our results indicate that both performance traits are important for victory, but that vigour is more important for fights resolved via decision, even though the effect of vigour is enhanced by skill. These results suggest that while similar traits are important for fighting success across the board, vigour is overvalued in judges' perceptions of RHP.

How does the environment affect fighting? The interaction between extrinsic fighting ability and resource value during contests.

An individual's performance during a fight is influenced by a combination of their capacity and willingness to compete. While willingness to fight is known to be determined by both intrinsic and extrinsic drivers, an individual's capacity to fight is generally thought of as solely intrinsic, being driven by a host of physiological factors. However, evidence indicates that variation in fighting ability can also be generated through exposure to different environmental conditions. Environmental contributions to fighting ability may be particularly important for animals living in spatially and temporally heterogeneous habitats, in which fights can occur between rivals recently exposed to different environmental conditions. The rapidly changing environment experienced within intertidal zones, for example, means that seawater parameters, including dissolved oxygen content and temperature, can vary across small spatial and temporal scales. Here, we investigated the relative importance of these extrinsic contributions to fighting ability and resource value on contest dynamics in the beadlet sea anemone Actinia equina We manipulated the extrinsic fighting ability of both opponents (through dissolved oxygen concentration prior to fights) and resource value (through seawater flow rate during the fight). Our results indicate that the extrinsic fighting ability of both opponents can interact with resource value to drive escalation patterns and that extrinsic drivers can be more important in determining contest dynamics than the intrinsic traits commonly studied. Our study highlights the need to combine data on intrinsic state and extrinsic conditions in order to gain a more holistic view of the factors driving contest behaviour.

Immune function and the decision to deploy weapons during fights in the beadlet anemone, Actinia equina.

The ability to mitigate the costs of engaging in a fight will depend on an individual's physiological state. However, the experience of fighting itself may, in turn, affect an individual's state, especially if the fight results in injury. Previous studies have found a correlation between immune state and fighting success, but the causal direction of this relationship remains unclear. Does immune state determine fighting success? Or does fighting itself influence subsequent immune state? Using the beadlet anemone, Actinia equina, we disentangled the cause and effect of this relationship, measuring immune response once pre-fight and twice post-fight. Contrary to previous findings, pre-fight immune response did not predict fighting success, but rather predicted whether an individual used its weapons during the fight. Furthermore, weapon use and contest outcome significantly affected post-fight immune response. Individuals that used their weapons maintained a stable immune response following the fight, whereas those that fought non-injuriously did not. Furthermore, although winners suffered a reduction in immune response similar to that of losers immediately post-fight, winners began to recover pre-fight levels within 24 h. Our findings indicate that immune state can influence strategic fighting decisions and, moreover, that fight outcome and the agonistic behaviours expressed can significantly affect subsequent immunity.

The role of skill in animal contests: a neglected component of fighting ability.

What attributes make some individuals more likely to win a fight than others? A range of morphological and physiological traits have been studied intensely but far less focus has been placed on the actual agonistic behaviours used. Current studies of agonistic behaviour focus on contest duration and the vigour of fighting. It also seems obvious that individuals that fight more skilfully should have a greater chance of winning a fight. Here, we discuss the meaning of skill in animal fights. As the activities of each opponent can be disrupted by the behaviour of their rival, we differentiate among ability, technique and skill itself. In addition to efficient, accurate and sometimes precise movement, skilful fighting also requires rapid decision-making, so that appropriate tactics and strategies are selected. We consider how these different components of skill could be acquired, through genes, experiences of play-fighting and of real fights. Skilful fighting can enhance resource holding potential (RHP) by allowing for sustained vigour, by inflicting greater costs on opponents and by minimizing the chance of damage. Therefore, we argue that skill is a neglected but important component of RHP that could be readily studied to provide new insights into the evolution of agonistic behaviour.

How does environment influence fighting? The effects of tidal flow on resource value and fighting costs in sea anemones.

An animal's decision to enter into a fight depends on the interaction between perceived resource value (V) and fighting costs (C). Both could be altered by predictable environmental fluctuations. For intertidal marine animals, such as the sea anemone Actinia equina, exposure to high flow during the tidal cycle may increase V by bringing more food. It may also increase C via energy expenditure needed to attach to the substrate. We asked whether simulated tidal cycles would alter decisions in fighting A. equina We exposed some individuals to still water and others to simulated tidal cycles. To gain insights into V, we measured their startle responses before and after exposure to the treatments, before staging dyadic fights. Individuals exposed to flow present shorter startle responses, suggesting that flowing water indicates high V compared with still water. A higher probability of winning against no-flow individuals and longer contests between flow individuals suggests that increased V increases persistence. However, encounters between flow individuals were less likely to escalate, suggesting that C is not directly related to V. Therefore, predictable environmental cycles alter V and C, but in complex ways.

How do anthropogenic contaminants (ACs) affect behaviour? Multi-level analysis of the effects of copper on boldness in hermit crabs.

Natural animal populations are increasingly exposed to human impacts on the environment, which could have consequences for their behaviour. Among these impacts is exposure to anthropogenic contaminants. Any environmental variable that influences internal state could impact behaviour across a number of levels: at the sample mean, at the level of among-individual differences in behaviour ('animal personality') and at the level of within-individual variation in behaviour (intra-individual variation, 'IIV'). Here we examined the effect of exposure to seawater-borne copper on the startle response behaviour of European hermit crabs, Pagurus bernhardus across these levels. Copper exposure rapidly led to longer startle responses on average, but did not lead to any change in repeatability indicating that individual differences were present and equally consistent in the presence and absence of copper. There was no strong evidence that copper exposure led to changes in IIV. Our data show that exposure to copper for 1 week produces sample mean level changes in the behaviour of hermit crabs. However, there is no evidence that this exposure led to changes in repeatability through feedback loops.

Weak rappers rock more: hermit crabs assess their own agonistic behaviour.

Fighting animals use a variety of information sources to make strategic decisions. A neglected potential source of information is an individual's own performance during a fight. Surprisingly, this possibility has yet to be incorporated into the large body of theory concerning the evolution of aggressive behaviour. Here, by experimentally dampening the impact of their shell rapping behaviour, we test for the possibility that attacking hermit crabs monitor their own fight performance. Attackers with dampened raps did not show a reduction in the number of raps used. By contrast, they showed an increased frequency of a less intense agonistic behaviour, shell rocking. This change in behaviour, in attackers that are forced to rap weakly, indicates that they assess their own agonistic behaviour.

Individual quality and personality: bolder males are less fecund in the hermit crab Pagurus bernhardus.

One explanation for animal personality is that different behavioural types derive from different life-history strategies. Highly productive individuals, with high growth rates and high fecundity, are assumed to live life at a fast pace showing high levels of boldness and risk taking, compared with less productive individuals. Here, we investigate among-individual differences in mean boldness (the inverse of the latency to recover from a startling stimulus) and in the consistency of boldness, in male hermit crabs in relation to two aspects of life-history investment. We assessed aerobic scope by measuring the concentration of the respiratory pigment haemocyanin, and we assessed fecundity by measuring spermatophore size. First, we found that individuals investing in large spermatophores also had high concentrations of haemocyanin. Using doubly hierarchical-generalized linear models to analyse longitudinal data on startle responses, we show that hermit crabs vary both in their mean response durations and in the consistency of their behaviour. Individual consistency was unrelated to haemocyanin concentration or spermatophore size, but mean startle response duration increased with spermatophore size. Thus, counter to expectations, it was the most risk-averse individuals, rather than the boldest and most risk prone, that were the most productive. We suggest that similar patterns should be present in other species, if the most productive individuals avoid risky behaviour.

Animal personality as a cause and consequence of contest behaviour.

We review the evidence for a link between consistent among-individual variation in behaviour (animal personality) and the ability to win contests over limited resources. Explorative and bold behaviours often covary with contest behaviour and outcome, although there is evidence that the structure of these 'behavioural syndromes' can change across situations. Aggression itself is typically repeatable, but also subject to high within-individual variation as a consequence of plastic responses to previous fight outcomes and opponent traits. Common proximate mechanisms (gene expression, endocrine control and metabolic rates) may underpin variation in both contest behaviour and general personality traits. Given the theoretical links between the evolution of fighting and of personality, we suggest that longitudinal studies of contest behaviour, combining behavioural and physiological data, would be a useful context for the study of animal personalities.

Anthropogenic noise limits resource distribution without changing social hierarchies.

Increasing evidence demonstrates that anthropogenic noise is a global pollutant that threatens marine ecosystems. Mounting numbers of studies show its diverse effects on individuals and their behaviour. However, little is known about how individual changes in response to anthropogenic noise could cascade through groups and populations affecting resource distribution vital for survival and fitness. Here we test the hypotheses that anthropogenic noise could alter resource distribution, associated hierarchies and consequently individual benefits. We used groups of hermit crabs, a globally distributed model system for assessing impacts of environmental change on wildlife and measured in controlled laboratory conditions the resource distribution of their reusable shelters (gastropod shells) under ship noise and ambient control playbacks. We applied vacancy chain theory to test three predictions about how new resource units create benefits for a population. A new resource unit leads to (i) a cascade of resource abandonments and acquisitions (= chain of vacancy moves) based on an internal (ii) hierarchy (here size-based) which allows (iii) more than one individual to benefit. All three predictions were supported under control sound. Under anthropogenic noise however, fewer individuals benefitted from the arrival of a new, empty shell, while the size-based hierarchy was maintained. The latter was apparent in chain structures, which were concordant between sound treatments. This experiment shows that anthropogenic noise can affect individual behaviours that cascade through groups. This has the potential to disrupt wider resource distribution in populations.

Plastic proteans: reduced predictability in the face of predation risk in hermit crabs.

Variation in behaviour occurs at multiple levels, including between individuals (personality) and between situations (plasticity). Behaviour also varies within individuals, and intra-individual variation (IIV) in behaviour describes within-individual residual variance in behaviour that remains after the effects of obvious external and internal influences on behaviour have been accounted for. IIV thus describes how predictable an individual's behaviour is. Differences in predictability, between individuals and between situations, might be biologically significant. For example, behaving unpredictably under predation threat might reduce the chance of capture. Here, we investigated the duration of startle responses in hermit crabs, in the presence and absence of a predator cue. Individuals differed in startle response duration (personality) and while individuals also varied in their sensitivity to risk, mean response time was greater in the presence of a predator (plasticity). Moreover, IIV was greater in the presence of a predator, providing some of the first evidence that the facultative injection of unpredictability into behaviour might represent a strategy for dealing with risk.

Size specific boldness associated with differences in resource requirements and habitat use: a cross-sectional study in hermit crabs.

Animal personality is often studied within compressed periods of observation that represent narrow windows in comparison to animal lifespans. Although much is known about the relations between repeatable personality traits and cross-situational behavioral plasticity, less is known about how such traits might differ across age classes or life history transitions. We conducted a cross-sectional study of startle response duration in 3 size classes of Pagurus bernhardus, the common European hermit crab. We defined size classes using transitions in the preferred species of gastropod shells that accompany growth, and this change in preference is in turn associated with a transition from intertidal to subtidal habitats. Compared with small- and medium-sized intertidal individuals the larger subtidal hermit crabs behaved cautiously by showing startle responses of greater duration following disturbance. Startle responses were also repeatable within all 3 size classes, confirming the presence of animal personality in intertidal hermit crabs and demonstrating that this pattern is retained within the largest size classes, which have undergone the transition from intertidal to subtidal habitat. Interestingly, there was a trend for the pattern of repeatable startle response durations to increase with size class, with the highest value for repeatability and the greatest range of startle response durations being present within the large subtidal population. The greater range of startle responses indicates that the longer startle response durations in some larger individuals are more likely due to developmental changes with age and habitat use than reflecting selection against the boldest individuals during earlier stages of life.

Evolutionary and spatiotemporal analyses reveal multiple introductions and cryptic transmission of SARS-CoV-2 VOC/VOI in Malta.

IMPORTANCE: Our study provides insights into the evolution of the coronavirus disease 2019 (COVID-19) pandemic in Malta, a highly connected and understudied country. We combined epidemiological and phylodynamic analyses to analyze trends in the number of new cases, deaths, tests, positivity rates, and evolutionary and dispersal patterns from August 2020 to January 2022. Our reconstructions inferred 173 independent severe acute respiratory syndrome coronavirus 2 introductions into Malta from various global regions. Our study demonstrates that characterizing epidemiological trends coupled with phylodynamic modeling can inform the implementation of public health interventions to help control COVID-19 transmission in the community.

Is boldness a resource-holding potential trait? Fighting prowess and changes in startle response in the sea anemone, Actinia equina.

Contest theory predicts the evolution of a stable mixture of different strategies for fighting. Here, we investigate the possibility that stable between-individual differences in startle-response durations influence fighting ability or 'resource-holding potential' (RHP) in the beadlet sea anemone, Actinia equina. Both winners and losers showed significant repeatability of pre-fight startle-response durations but mean pre-fight startle-response durations were greater for eventual losers than for eventual winners, indicating that RHP varies with boldness. In particular, individuals with short startle responses inflicted more attacks on their opponent. Both repeatability and mean-level responses were changed by the experience of fighting, and these changes varied with outcome. In losers, repeatability was disrupted to a greater extent and the mean startle-response durations were subject to a greater increase than in winners. Thus, following a fight, this behavioural correlate of RHP behaves in a way similar to post-fight changes in physiological status, which can also vary between winners and losers. Understanding the links between aggression and boldness therefore has the potential to enhance our understanding of both the evolution of animal personality and the 'winner and loser effects' of post-fight changes in RHP.

Fight tactics in wood ants: individuals in smaller groups fight harder but die faster.

When social animals engage in inter-group contests, the outcome is determined by group sizes and individual masses, which together determine group resource-holding potential ('group RHP'). Individuals that perceive themselves as being in a group with high RHP may receive a motivational increase and increase their aggression levels. Alternatively, individuals in lower RHP groups may increase their aggression levels in an attempt to overcome the RHP deficit. We investigate how 'group RHP' influences agonistic tactics in red wood ants Formica rufa. Larger groups had higher total agonistic indices, but per capita agonistic indices were highest in the smallest groups, indicating that individuals in smaller groups fought harder. Agonistic indices were influenced by relative mean mass, focal group size, opponent group size and opponent group agonistic index. Focal group attrition rates decreased as focal group relative agonistic indices increased and there was a strong negative influence of relative mean mass. The highest focal attrition rates were received when opponent groups were numerically large and composed of large individuals. Thus, fight tactics in F. rufa seem to vary with both aspects of group RHP, group size and the individual attributes of group members, indicating that information on these are available to fighting ants.

Do I stand out or blend in? Conspicuousness awareness and consistent behavioural differences in hermit crabs.

Animals titrate their behaviour against the level of risk and an individual's conspicuousness should influence decisions such as when to flee and for how long to hide. Conspicuousness will vary with variation in substrate colour. Since hermit crabs frequently change the shells they occupy, shell colour will also influence conspicuousness and to be aware of their conspicuousness would require information on both of these factors to be integrated. Reduced boldness in high-contrast shell and substrate combinations compared with situations of low contrast indicates that hermit crabs are aware of current conspicuousness. Differences between individuals remained consistent across conspicuousness levels indicating the presence of animal personalities.