The Influence of Personality on a Group‐Level Process: Shy Hermit Crabs Make Longer Vacancy Chains

Animal personality has been investigated in a variety of contexts but few studies have investigated the effects of individual‐level traits on processes that occur at the group or population level. Vacancy chains describe sequences of linked vacancy moves in systems where individuals use discrete and reuseable resource units. When a new resource unit becomes available, this can cause a cascade of moves between resource units, beyond the first individual that encounters the new unit. Thus, vacancy chains can drive the distribution of resources through a population. Here, I investigate the effect of variation in boldness on movements between vacant gastropod shells in the hermit crab Pagurus bernhardus. Boldness was quantified using the duration of a startle response, which varied consistently between individuals. In groups of five crabs, introduction of a single new shell led to aggregate benefits, in terms of improving shell quality, across all group members. Boldness did not influence the extent of these benefits or the extent to which they were distributed between crabs of different sizes. Initially, vacancy chains were longer, indicating that more individuals moved between shells, for bold groups compared with shy groups. After 24 h, however, the chain length was longer for shy groups. Thus, although participation in a vacancy chain may lead to similar benefits, these could be more widespread for groups comprising shy individuals that show long startle responses, low inquisitiveness and low aggression. Similar effects on resource acquisition could be present in other examples of animals that utilise reusable resource units and show consistent between‐individual differences in behaviour.     

Vacancy Chains Provide Aggregate Benefits To Coenobita clypeatus Hermit Crabs

Vacancy chain theory describes a unique mechanism for the sequential distribution of animal resources across multiple individuals. This theory applies to any resources, such as shelters or nest sites, that are discrete, reusable, and limited in use to single individuals or groups at one time. Hermit crabs rely on gastropod shells for shelter, and a single vacant shell can initiate a chain of sequential shell switches that distributes new resources across many individuals. Using the terrestrial hermit crab Coenobita clypeatus , we examined the previously untested theoretical prediction that this process will yield trickle-down resource benefits to vacancy chain participants (aggregate benefits). In laboratory experiments, we measured improvements in shell quality when a single vacant shell was provided to groups of eight crabs. We found that crabs participating in vacancy chains (averaging 3.2 individuals) gained significant reductions in their shell crowding. In addition, vacancy chains terminated early when experimental groups included a single crab occupying a damaged shell, because damaged vacancies always remained unoccupied. Hermit crabs in damaged shells were more likely to win resource contests for high quality shells against size-matched hermit crabs in crowded shells. Finally, field additions of many new shells to an island population of C. clypeatus hermit crabs reduced average shell crowding for crabs of all sizes, possibly from propagation of benefits through vacancy chains. These results provide empirical support for the theoretical prediction that vacancy chains should provide benefits distributed across many vacancy chain participants. Since shelter-based vacancy chains likely occur in other animals, additional studies of vacancy chain processes should provide new insights into resource acquisition behaviors in diverse animal groups.     

The vacancy chain process: a new mechanism of resource distribution in animals with application to hermit crabs

A number of resources important to humans such as jobs in bureaucracies, houses and apartments are allocated through a mechanism known as a vacancy chain. In a vacancy chain process an initial, vacant resource unit entering a population of users is taken by a first individual who leaves his/her previous resource unit behind, which is taken by a second individual, and so on. In this process an initial resource unit works both directly and indirectly to provide opportunities for several individuals to gain new and better resources. Vacancy chains are hypothesized to be important in resource distribution for a variety of non-human animals, and it is documented, in particular, that the hermit crab Pagurus longicarpus gets the gastropod shells in which it lives through this mechanism. The direct and indirect effects of vacancy chains on hermit crabs and the systematic ways in which chains flow through groups of crabs and their resources are indicated. In systems where they occur, it is further hypothesized that vacancy chains have unique implications for the ecology of resource users. Several of these hypotheses are explored using the example of hermit crabs.     

Chains of opportunity: A Markov model for acquisition of reusable resources

Summary A vacancy chain is a unique type of resource acquisition process composed of an interconnected series of events in which the gaining of a particular resource unit by one individual depends directly on prior acquisition events by other individuals. Taken from the sociological literature, vacancy chains may also describe the distribution of many types of animal resources such as burrows, dwellings and shelters. Using data on hermit crabs, we present a Markov model simulating a vacancy chain process, and test the model against field data. Our results show that a simple Markov model adequately describes shell acquisition in hermit crabs, and that models combining shell size and crude estimates of quality fit the data extremely well. We illustrate in detail how to generate vacancy chain models from ecological data, how to determine the number and size of organisms gaining new resource units from resource introductions of specific sizes, and how to statistically evaluate the accuracy of Markov models. Not recognizing the presence of a vacancy chain system may lead to serious errors in estimating resource dynamics and therefore in demographic and competition models based on these dynamics. Finally, we suggest some ways in which vacancy chain models can aid studies of competition, population dynamics, life histories, and conservation in species using this type of resource acquisition process.     

Relaxed selection when you least expect it: why declining bird populations might fail to respond to phenological mismatches

Ongoing climate change threatens to cause mismatches between the phenology of many organisms and their resources. Populations of migratory birds may need to undergo ‘evolutionary rescue’ if resource availability moves to earlier dates in the year, as shifted arrival dates at the breeding grounds may be required for persistence under new environmental schedules. Here we show a counterintuitive process that can reduce the strength of selection for early arrival when the resource peaks earlier. This happens when two processes combine to determine selection for early arrival: breeding success is higher if a bird does not miss the resource peak, but this occurs together with a ‘zero‐sum game’ where birds acquire good territories ahead of their competitors if they arrive early. The latter process can relax if the population has experienced a recent decline. Therefore, climate change can have two opposing effects: its direct effect on breeding success strengthens selection for early arrival, but this combines with an indirect effect of relaxed selection due to population declines, if territoriality is a significant determinant of population dynamics and fitness. We show that the latter process can predominate, and this can cause a failure for a population to adapt to a new schedule under changing environmental conditions.     

Effects of Spider Chemotactile Cues on Arthropod Behavior

Predation risk can strongly affect the behavior of prey species. However, empirical evidence for changes in behavior driven by spider cues is restricted to relatively few taxa. Here, we conducted a series of behavioral experiments to test for changes in activity among a wide range of terrestrial arthropods. We confronted 13 insect and eight spider species with chemotactile cues of three spider species. We applied two different experimental setups: In the ‘no-choice experiment’ prey individuals were either put on control filter papers or on filter papers previously occupied by a spider. In the ‘choice experiment’, the prey individuals were able to choose between filter paper halves with and without spider cues. In both setups, the response to spider cues depended significantly on prey species, with some species increasing and others decreasing their activity. Surprisingly few prey species responded to the spider cues at all. Our results indicate that predator recognition upon contact with cue bearing filter papers is strongly prey-specific and that behavioral effects driven by spider chemotactile cues are an exception rather than the rule among terrestrial arthropods.     

An Experimental Test of Predator Detection Rates Using Groups of Free-living Emus

Abstract Improved predator detection is often stated to be one of the principal benefits of social foraging. However, actual field evidence supporting this assumption is scarce. This may be the result of the fact that most observations are conducted on social animals acting in the absence of an acute predation threat, yet the benefits of grouping come to the fore in that brief moment when an individual's life is at risk. As predation attempts are typically rare in nature, experimental manipulations are necessary to further explore the costs and benefits of social foraging. This study utilizes simple predator simulations (by humans) to experimentally test the predator‐detecting abilities and escape strategies of groups of free‐living emus Dromaius novaehollandiae. Emus in larger groups spent less time in vigilance and more time foraging. Nonetheless, the combined vigilance of group members ensured that emus detected the ‘simulated predator’ sooner as group size increased. After detecting the ‘predator’, larger groups waited longer until opting to flee, and then spent less time and energy doing so. Thus, the results of this study provide experimental evidence that emus benefit from grouping in terms of both the ‘many‐eyes effect’ and the ‘dilution effect’.     

Climate change enhances the negative effects of predation risk on an intermediate consumer

Predators are a major source of stress in natural systems because their prey must balance the benefits of feeding with the risk of being eaten. Although this ‘fear’ of being eaten often drives the organization and dynamics of many natural systems, we know little about how such risk effects will be altered by climate change. Here, we examined the interactive consequences of predator avoidance and projected climate warming in a three‐level rocky intertidal food chain. We found that both predation risk and increased air and sea temperatures suppressed the foraging of prey in the middle trophic level, suggesting that warming may further enhance the top‐down control of predators on communities. Prey growth efficiency, which measures the efficiency of energy transfer between trophic levels, became negative when prey were subjected to predation risk and warming. Thus, the combined effects of these stressors may represent an important tipping point for individual fitness and the efficiency of energy transfer in natural food chains. In contrast, we detected no adverse effects of warming on the top predator and the basal resources. Hence, the consequences of projected warming may be particularly challenging for intermediate consumers residing in food chains where risk dominates predator‐prey interactions.     

Signalling of information that is neither cryptic nor private

It is commonly assumed that in order for animal signals to be advantageous, the information being signalled could not have been obtained otherwise, and is therefore ‘cryptic’ or ‘private’. Here, we suggest a scenario in which individuals can gain an advantage by signalling ‘public’ information that is neither cryptic nor private. In that scenario, signalling increases the efficiency with which that ‘public’ information is transmitted. We formalize our idea with a game in which offspring can signal their condition to their parents. Specifically, we consider a resource‐strapped parent who can only invest in one of its two offspring, and we allow offspring the chance to influence parental investment through a signal. A parent in the game seeks to invest in the higher‐quality offspring, which it could identify either through a publicly available cue, such as body size, or by relying on a signal provided by the offspring. We find that if the signal can convey information about offspring quality more efficiently than cues, then signalling of condition between offspring and parents can be favoured by selection, even though parents could potentially have acquired the same information from the cue. Our results suggest that the biological function of signals may be broader than currently considered, and provide a scenario where low cost signalling can be favoured. More generally, efficiency benefits could explain signalling across a range of biological and economic scenarios.     

Territory size in the moorhen (Gallinula chloropus): An outcome of RHP asymmetry between neighbours

Territory size can be considered as the outcome of contests between an owner and its neighbours for the resource units in the defended area. Variation in territory size could be dependent upon three possible asymmetries: (1) difference in resource holding potential (RHP); (2) difference in resource value to the competitors; and (3) difference in ‘ownership’ status. These possibilities were tested in a 3-year study of moorhens which defend linear territories along ditches in a grazing marsh habitat. Of the three, relative difference in RHP between an owner and its two neighbours provided the best correlation with territory size. There was no relationship between the amount of resources per unit area and territory size nor between the time of establishing a territory and its later size. Heavier moorhens are more likely to win contests in winter flocks and are thought to have a greater RHP. The ratio of the weight of a male territory owner to the weight of both its male neighbours was highly correlated with territory size. Birds with larger territories may gain more potential nesting sites with good cover from predators.     

Calling as an ownership convention on pied wagtail territories

Winter feeding territories were more profitable to owners than intruders. Even when intruders landed on a territory undetected they failed to achieve a profitable feeding rate because they fed in areas recently depleted by the owner. For intruders therefore, ownership was a sign of unprofitability. Intruder scalled ‘chis-ick’, apparently to find out if a territory was occupied. Owners replied ‘chee-wee’ and this was usually sufficient to cause the intruder to retreat. Only owners called ‘chee-wee’. Owners reacted more aggressively to tape-recorded broadcasts of the ‘chee-wee’ call played on their territory than to broadcasts of ‘chis-ick’.     

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.     

Sensory cues of a top‐predator indirectly control a reef fish mesopredator

Behavioural trophic cascades highlight the importance of indirect/risk effects in the maintenance of healthy trophic‐level links in complex ecosystems. However, there is limited understanding on how the loss of indirect top–down control can cascade through the food‐web to modify lower level predator–prey interactions. Using a reef fish food‐web, our study examines behavioural interactions among predators to assess how fear elicited by top‐predator cues (visual and chemical stimuli) can alter mesopredator behaviour and modify their interaction with resource prey. Under experimental conditions, the presence of any cue (visual, chemical, or both) from the top‐predator (coral trout Plectropomus leopardus) strongly restricted the distance swum, area explored and foraging activity of the mesopredator (dottyback Pseudochromis fuscus), while indirectly triggering a behavioural release of the resource prey (recruits of the damselfish Pomacentrus chrysurus). Interestingly, the presence of a large non‐predator species (thicklip wrasse Hemigymnus melapterus) also mediated the impact of the mesopredator on prey, as it provoked mesopredators to engage in an ‘inspection’ behaviour, while significantly reducing their feeding activity. Our study describes for the first time a three‐level behavioural cascade of coral reef fish and stresses the importance of indirect interactions in marine food‐webs.     

Signal Interception and the Use of Soft Song in Aggressive Interactions

Song sparrows, like many species of songbirds, produce songs of especially low amplitude during aggressive contests. Such ‘soft songs’ have been shown to be reliable signals of intention to attack in song sparrows. Low intensity is a paradoxical feature in a highly aggressive signal, in that it seems likely to make the signal less intimidating to an opponent. A hypothesis that resolves this paradox is that use of soft songs benefits a signaler by limiting the interception of his signals by third party receivers. Here, we test this hypothesis with respect to song sparrows and two classes of third party receivers, predators and conspecific males. We tested whether selection to avoid predation is responsible for use of soft song by examining whether male song sparrows increase production of soft song in an aggressive context when they also receive signals (alarm calls) that indicate a predator is present. We found that the proportion of soft song produced by male song sparrows was actually significantly lower in the predator context than in a control context, directly contradicting the prediction. We tested whether avoiding eavesdropping by conspecific males is a benefit of soft song by removing territory owners and simulating interactions on their territories using playback from two loudspeakers. We found that intrusions by third party males were more likely in trials in which the simulated owner countered an intruder's songs using soft songs than in trials in which the simulated owner countered with loud song, again directly contradicting the hypothesis. We conclude that limiting interception by predators and conspecific males does not provide an explanation for use of soft song by song sparrows.     

Migration of two calidrid sandpiper species on the predator landscape: how stopover time and hence migration speed vary with geographical proximity to danger

The effects of relative fuel load on migration speed and on vulnerability have been investigated, but the effects of seasonal variation in predation danger on the amount of fuel and duration of stopover have not been considered. We analyzed seasonal patterns of stopover residence times for western and semipalmated sandpipers Calidris mauri and C. pusilla on southward migration in relation to the passage of migratory peregrine falcons Falco peregrinus. We predicted that individuals on stopover far in advance of the seasonal arrival of falcons would adjust stopover length and hence relative fuel load to migrate slowly and cautiously. We predicted that individuals on stopover later in the season would increase migratory speed as the arrival of migratory falcons came closer, while individuals on stopover under or behind the passage of falcons would migrate slowly. Adult and juvenile semipalmated and adult western sandpipers migrated prior to seasonal increases in peregrine abundance, and as predicted, the seasonal patterns of their stopover durations are consistent with an increase in the speed of migration as the date of peregrine arrival approached. Juvenile western sandpipers, in contrast, migrating concurrently with falcons, slowed their speed of migration as predator abundance increased. Stopover patterns differ between species due to different relative fuel loads. The results fit predictions made based on a ‘mortality‐minimizing’ migration strategy.     

Differential responses to predator cues between two mosquito species breeding in different habitats

1. Natural selection favours females who can correctly assess the predation risk and hence avoid high‐risk oviposition sites and reduce the mortality rate of their offspring. In spite of the potential significance of such behaviour, relatively few studies have assessed the relationship between oviposition behaviour and predation risk. 2. The present study aimed to determine the sublethal effects of predators on oviposition site selection by gravid females, the foraging activity of larvae, and the life history traits of two mosquito species that breed in different habitats, Aedes albopictus Skuse (container breeder) and Culex tritaeniorhynchus Giles (wetland breeder). 3. Female C. tritaeniorhynchus avoided laying eggs at oviposition sites in the presence of a predator cue. In contrast, female A. albopictus laid eggs in both the absence and presence of the predator cue. 4. To examine the effects of predator cues on larval behaviour, experiments were conducted in the absence and presence of a predator cue. Although larval activity was lower in the presence of the predator cue than that in its absence in both species, C. tritaeniorhynchus responded to the predator cue more strongly than A. albopictus. Female A. albopictus that had been reared with caged predators exhibited an extended larval development period, whereas the adult C. tritaeniorhynchus reared in the presence of predators were smaller than those reared in their absence. 5. This finding might explain why C. tritaeniorhynchus avoid laying eggs in predator‐conditioned water, for example to increase the fitness of their offspring, but A. albopictus either cannot detect predator cues or are not sensitive to them.     

Effect of Willow Tit Poecile montanus alarm calls on attack rates by Pygmy Owls Glaucidium passerinum

One suggested anti‐predator function of alarm calls is to deliver a message to a predator that it has been detected. Moreover, giving the alarm call could provide a signal to the predator that capturing the individual giving the alarm is more difficult than capturing its silent group members, as the caller is probably the most aware of the predator's location. In an aviary experiment using stuffed dummy Willow Tits Poecile montanus, we assessed whether an authentic alarm call given by Willow Tit affected Pygmy Owl Glaucidium passerinum prey preference. In the experiment, the Owls attacked only the ‘silent’ dummy individuals, suggesting that alarm calling could offer direct fitness benefits to the caller by decreasing the attack risk of the caller relative to its group members.     

Fitness costs and benefits of antipredator behavior mediated by chemotactile cues in the wolf spider Pardosa milvina (Araneae: Lycosidae)

Animals may exhibit a variety of defensive behaviors in thepresence of indirect predator cues. Such behavior offers immediatefitness benefits but may also incur substantial foraging andreproductive costs. We measured shifts in space use (verticalclimbing) by the wolf spider Pardosa milvina induced by chemotactilecues (silk and excreta) from a co-occurring predatory wolfspider Hogna helluo. We then measured foraging and reproductive costs, as well as survival benefits, of this behavior. For 2weeks, we maintained mated adult female Pardosa in plasticcontainers with one of three treated peat moss substrates:a container previously occupied by a conspecific for 3 days,a container previously occupied by an adult Hogna for 3 days,and a container devoid of either cue (control). We measuredprey capture efficiency, body condition, egg sac production,egg sac weight, and egg number for individuals in each treatment.We also counted the number of Pardosa that survived and exhibitedclimbing behavior in the presence of a live Hogna with andwithout silk and excreta cues. Pardosa climbed container wallssignificantly more often in the presence of Hogna silk andexcreta relative to other treatments. Pardosa exposed to Hognacues coupled with live Hogna survived significantly longerthan spiders that had no predator cues available. Pardosa placedin containers with Hogna cues, but no Hogna, lost weight morequickly, ate fewer prey, were in poorer body condition, producedlighter egg sacs, and produced fewer eggs than spiders in controlor conspecific treatments.     

Depth distribution of Daphnia in response to a deep-water algal maximum: the effect of body size and temperature gradient

1. In the absence of fish predation, Daphnia exploiting a deep‐water algal maximum are faced with a trade‐off. They can either dwell in the epilimnion where development in the warm water is fast, but food shortage causes low egg production, or in the hypolimnion, where food availability is high but development is slow because of low temperatures. 2. We tested the hypotheses that (i) depth distributions of various ontogenetic stages (size classes and egg‐bearing females) differ because daphnids react to light with size‐specific diel vertical migration (DVM) even in the absence of fish (residual predator avoidance hypothesis) and (ii) differently sized daphnids select different depths because the relative importance of temperature and food varies for ontogenetic stages (physiological hypothesis). We used large indoor mesocosms (Plankton Towers) to test these hypotheses experimentally. 3. Temperature was the strongest factor governing the distribution, with larger proportions of the population dwelling in the food‐rich hypolimnion if the temperature gradient was shallow. There were small but significant differences between ontogenetic stages during the day, but not at night. This suggested the existence of a ‘residual’ effect of light on depth distribution in the absence of a fish cue. 4. Although large individuals exhibited greater amplitude of DVM, the physiological hypothesis had to be rejected. A stage‐specific physiological effect is unlikely to be directly triggered by light, hence vertical movement of the individuals should not be synchronised. Rather, being forced into deeper layers by the residual light response during the day, large and egg‐bearing females experience a lower average temperature during day than juveniles. They probably compensate for this by spending longer time periods in warm waters at night.     

The logic of asymmetric contests

A theoretical analysis is made of the evolution of behavioural strategies in contest situations. It is assumed that behaviour will evolve so as to maximize individual fitness. If so, a population will evolve an ‘evolutionarily stable strategy’, or ESS, which can be defined as a strategy such that, if all members of a population adopt it, no ‘mutant’ strategy can do better. A number of simple models of contest situations are analysed from this point of view. It is concluded that in ‘symmetric’ contests the ESS is likely to be a ‘mixed’ strategy; that is, either the population will be genetically polymorphic or individuals will be behaviourally variable. Most real contests are probably asymmetric, either in pay-off to the contestants, or in size or weapons, or in some ‘uncorrelated’ fashion; i.e. in a fashion which does not substantially bias either the pay-offs or the likely outcome of an escalated contest. An example of an uncorrelated asymmetry is that between the ‘discoverer’ of a resource and a ‘late-comer’. It is shown that the ESS in asymmetric contests will usually be to permit the asymmetric cue to settle the contest without escalation. Escalated contests will, however, occur if information to the contestants about the asymmetry is imperfect.