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.

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.

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.

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.

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.

Beyond spider personality: The relationships between behavioral, physiological, and environmental factors.

Spiders are useful models for testing different hypotheses and methodologies relating to animal personality and behavioral syndromes because they show a range of behavioral types and unique physiological traits (e.g., silk and venom) that are not observed in many other animals. These characteristics allow for a unique understanding of how physiology, behavioral plasticity, and personality interact across different contexts to affect spider's individual fitness and survival. However, the relative effect of extrinsic factors on physiological traits (silk, venom, and neurohormones) that play an important role in spider survival, and which may impact personality, has received less attention. The goal of this review is to explore how the environment, experience, ontogeny, and physiology interact to affect spider personality types across different contexts. We highlight physiological traits, such as neurohormones, and unique spider biochemical weapons, namely silks and venoms, to explore how the use of these traits might, or might not, be constrained or limited by particular behavioral types. We argue that, to develop a comprehensive understanding of the flexibility and persistence of specific behavioral types in spiders, it is necessary to incorporate these underlying mechanisms into a synthesized whole, alongside other extrinsic and intrinsic factors.

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.

The role of spatial accuracy and precision in hermit crab contests.

Understanding the determinants of fighting ability (or resource-holding potential, RHP) is key to elucidating the evolution of aggressive behaviour, as current tests of contest theory rely on realistic proxies for overall RHP. Traditionally, RHP is considered equivalent to body size but it is increasingly clear that a wider range of morphological and physiological traits contribute to fighting ability. In situations analogous to contests, such as courtship displays in animals and competitive sport in humans, the role of skill has long been appreciated but this component has been neglected in analyses of animal fights. Here, we investigated two spatial components of skill, accuracy and precision, during shell fights in hermit crabs, where an attacker repeatedly strikes (raps) its shell against that of a defender. By analysing the points of impact of these strikes, we found that attackers that rapped with coarse-scale accuracy were more likely to win the fight, indicating that the ability to target a 'sweet spot' on the defender's shell is an important determinant of contest success. Furthermore, we found that this element of skill correlated with temporal performance (vigour). Taken together these results show that spatial skill is an RHP component. Moreover, in contrast to the traditional assumption that fighting ability is equivalent to body size, RHP is actually underpinned by a suite of interlinked traits including performance capacities, morphology and skill.

Analysis of direct and indirect genetic effects in fighting sea anemones.

Theoretical models of animal contests such as the Hawk-Dove game predict that variation in fighting behavior will persist due to mixed evolutionarily stable strategies (ESS) under certain conditions. However, the genetic basis for this variation is poorly understood and a mixed ESS for fighting can be interpreted in more than one way. Specifically, we do not know whether variation in aggression within a population arises from among-individual differences in fixed strategy (determined by an individual's genotype-direct genetic effects [DGEs]), or from within-individual variation in strategy across contests. Furthermore, as suggested by developments of the original Hawk-Dove model, within-individual variation in strategy may be dependent on the phenotype and thus genotype of the opponent (indirect genetic effects-IGEs). Here we test for the effect of DGEs and IGEs during fights in the beadlet sea anemone Actinia equina. By exploiting the unusual reproductive system of sea anemones, combined with new molecular data, we investigate the role of both additive (DGE + IGE) and non-additive (DGE × IGE) genetic effects on fighting parameters, the latter of which have been hypothesized but never tested for explicitly. We find evidence for heritable variation in fighting ability and that fight duration increases with relatedness. Fighting success is influenced additively by DGEs and IGEs but we found no evidence for non-additive IGEs. These results indicate that variation in fighting behavior is driven by additive indirect genetic effects (DGE + IGE), and support a core assumption of contest theory that strategies are fixed by DGEs.

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.

Responses to threat in a freshwater invader: longitudinal data reveal personality, habituation, and robustness to changing water temperatures in the "killer shrimp" Dikerogammarus villosus (Crustacea: Amphipoda).

Freshwater biodiversity and ecosystem integrity are under threat from biological invasions. The "killer shrimp" Dikerogammarus villosus is a highly predatory amphipod that has spread readily across Central Europe and recently the UK and its arrival has been associated with the significant loss of resident species. Despite this, studies of its behavioral ecology are sparse, even though its behavior may contribute to its invasion success. For the first time, we investigated antipredator "fleeing" behavior in D. villosus and how this changed with water temperature. Three key patterns emerged from our analysis. First, within a particular temperature condition there are moderate but consistent among-individual differences in behavior. These are driven by a combination of mean level among-individual differences and within-individual relative consistency in behavior, and provide the key marker for animal personalities. Second, the fleeing responses were not influenced by temperature and third, regardless of temperature, all individuals appeared to habituate to a repeated nondangerous stimulus, indicating a capacity for individual learning. We suggest that the antipredator behavior of D. villosus contributes to its rapid spread and that consistent among-individual differences in behavior may promote biological invasions across heterogeneous conditions. Robustness to changing water temperatures may also be potentially advantageous, particularly in an era of global climate change, where average temperatures could be elevated and less predictable.

Exaggerated displays do not improve mounting success in male seaweed flies Fucellia tergina (Diptera: Anthomyiidae).

Signals of individual quality are assumed to be difficult to exaggerate, either because they are directly linked to underlying traits (indices) or because they are costly to perform (handicaps). In practise advertisement displays may consist of conventional and costly components, for instance where a morphological structure related to body size is used in visual displays. In this case, there is the potential for dishonest displays, due to the population level variance around the relationship between body size and display structures. We examine the use of wing flicking displays that we observed in situ in a strandline dwelling seaweed fly Fucellia tergina, using overall body size and the size of their eyes as underlying indicators of condition. Males displayed far more frequently than females, and were also observed to frequently mount other flies, a behaviour that was rare in females. The rate of display was greater for males that had positive residual values from relationships between wing length and body length. In other words those males with larger than expected wings for their underlying quality displayed more frequently, indicating that these displays are open to exaggeration. Males with larger than expected wings (for the size of their body or eyes), however, mounted less frequently. We suggest that small bodied males are less successful in terms of mounting, but that those small males with relatively large wings may attempt to compensate for this through increased display effort.

Parasites and personality in periwinkles (Littorina littorea): Infection status is associated with mean-level boldness but not repeatability.

We demonstrate the presence of animal personality in an inter-tidal gastropod, Littorina littorea, both in a sample of individuals infected by the trematode Cryptocotyle lingua and in an uninfected sample. On average infected individuals behaved more cautiously than individuals free of infection, but the parasite did not affect repeatability. Although the parasite is not associated with greater diversity of behaviour amongst infected individuals, infection might be associated with state-dependent personality differences between infected and non-infected individuals.

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.