Rapid change in energy status in fighting animals: causes and effects of strategic decisions

Energetic costs of fighting, such as high lactate or low glucose, have been shown in a range of species to correlate with the decisions made by each opponent, particularly the decision by one opponent, the 'loser', to end the fight by 'giving up'. Studies based on complete fights of differing duration, however, do not provide information on the changes in the physiological correlates of fighting that may take place during the course of the encounter, or how these changes may influence the capability and decisions of the contestants. We interrupted fights between hermit crabs, Pagurus bernhardus, at specific points, and related energy status to the preceding activities. Costs rose quickly with a rapid accumulation of lactic acid in attackers and declining muscular glycogen in defenders. Changes in physiological status appeared much earlier than the changes in behaviour that they may have caused. Furthermore, some physiological changes might have been an effect, rather than the cause, of fight decisions. (c) 2005 The Association for the Study of Animal Behaviour Published by Elsevier Ltd. All rights reserved.

Motivational change during shell fights in the hermit crab Pagurus bernhardus

During agonistic interactions the motivation of each contestant is expected to vary because of increased information and changes in fighting ability. In shell fights between hermit crabs over gastropod shells, attackers rap their shell in a series of bouts against that of the defender whereas defenders remain withdrawn into their shells until the encounter is resolved; either the defender is evicted from its shell or the attacker 'gives up' and the defender retains its shell. We assessed the motivational state of attackers for performing rapping by measuring the duration of startle responses elicited by a novel stimulus. We staged fights between pairs of crabs in six different groups defined by the potential gain in shell quality available to attackers (high or low) and by the point at which the novel stimulus was applied (prior to rapping, after one bout or after four bouts). Startle response duration decreased during the first four bouts of fighting and showed a U-shaped relationship with the relative difference in size between the crabs. There was, no difference in startle response duration between high- and low-gain groups. Individuals showing short startle responses were likely to be victorious and we conclude that the relationship between the relative size difference of the opponents and. startle duration reflects that between size difference and the cost of gaining an eviction. (C) 2001 The Association for the Study of Animal Behaviour.

Decision rules, energy metabolism and vigour of hermit-crab fights

Aggressive interactions between animals are often settled by the use of repeated signals that reduce the risk of injury from combat but are expected to be costly. The accumulation of lactic acid and the depletion of energy stores may constrain activity rates during and after fights and thus represent significant costs of signalling. We tested this by analysing the concentrations of lactate and glucose in the haemolymph of hermit crabs following agonistic interactions over the ownership of the gastropod shells that they inhabit. Attackers and defenders play distinct roles of sender and receiver that are fixed for the course of the encounter. Attackers perform bouts of 'shell rapping', which vary in vigour between attackers and during the course of the encounter, and are a key predictor of victory. In contrast to the agonistic behaviour of other species, we can quantify the vigour of fighting. We demonstrate, to our knowledge for the first time, an association between the vigour of aggressive activity and a proximate cost of signalling. We show that the lactate concentration in attackers increases with the amount of shell rapping, and that this appears to constrain the vigour of subsequent rapping. Furthermore, attackers, but not defenders, give up when the concentration of lactate is high. Glucose levels in attackers also increase with the amount of rapping they perform, but do not appear to influence their decision to give up. Defenders are more likely to lose when they have particularly low levels of glucose. We conclude that the two roles use different decision rules during these encounters.

Claw removal and feeding ability in the edible crab, Cancer pagurus: Implications for fishery practice

Feeding ability and motivation were assessed in the edible crab, Cancer pagurus, to investigate how the fishery practice of de-clawing may affect live crabs returned to the sea. Crabs were either induced to autotomise one claw, or were only handled, before they were offered food. Initially, autotomised and handled crabs were offered mussels, Mytilis edulis, a large part of their natural diet. After 3 days, both autotomised and handled crabs were then offered fish, a more readily handled food source. Autotomy induced crabs consumed significantly fewer mussels and less mussel mass, but ate significantly more mass of fish. This indicates that the effect of autotomy was a reduction of ability to feed on mussels rather than a general reduction of feeding motivation. The discontinuation of claw removal needs to be considered, both for the sustainability of the fishery and animal welfare concerns. (C) 2008 Elsevier B.V. All rights reserved.

Signal residuals and hermit crab displays: flaunt it if you have it!

In animal contests selection should favour information gathering regarding the likely costs and benefits of continued conflict, and displays may provide a means for contestants to gain information about the fighting ability or aggressive intent of competitors. However, there is debate over the reliability of such displays and low levels of deception may occur within otherwise honest signalling systems. Hermit crabs use displays involving the chelipeds during agonistic encounters. We examined how variation in chelae size in relation to body size, a determinant of fighting ability, affects their use in displays and the process and outcome of contests over gastropod shells. In accordance with deceptive use of an otherwise honest signal, we found that contestants with large chelipeds for their body size spent more time performing the cheliped presentation display. Moreover, cheliped residuals and displays influenced the escalation level of encounters. There was a positive association between cheliped displays and the occurrence of 'grappling', but a negative association between displays and the occurrence of shell fights, suggesting that displays may signal aggressive intent and a reluctance to back off or accept the more passive defender role in a fight. Furthermore, the smaller of the two contestants in shell fights had larger cheliped residuals compared to those smaller contestants not involved in shell fights, which is consistent with disrupted opponent assessment. This study adds to mounting evidence that when acting as a signaller, individuals for whom the display exaggerates competitive ability attempt to manipulate opponents, using the display more often. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Motivation matters: cheliped extension displays in the hermit crab, Pagurus bernhardus, are honest signals of hunger

Whether animal signals convey honest information is a central evolutionary question, since selection pressures could, in some circumstances, favour dishonesty. A prior study of signalling in hermit crabs proposed that the cheliped extension display of Pagurus bernhardus might represent such an instance of dishonesty. A limitation of this conclusion, however, was that honesty was defined in the context of size assessment, neglecting the potential information that displays might transmit about signallers' variable internal states. Recent analyses of signalling in this same species have shown that its displays provide reliable information about the amount of risk crabs are prepared to tolerate, which therefore might enable signallers to use these displays to honestly convey their motivation to take such risks. Here we test this 'honest advertisement of motivation' hypothesis by varying crabs' need for food and analysing their signalling during simulated feeding conflicts against a model. When crabs were starved for 1-5 days, they dropped significantly in weight. Despite this decrement in resource-holding potential and energy reserves, crabs were more likely to perform cheliped extension displays the longer they were food deprived. Longer-starved crabs, whose subjective resource value was greater, also displayed at a higher rate and were more likely to risk seizing the food from the model. We conclude that cheliped extension is a reliable indicator of crabs' internal state and suggest how this honest signal might operate in conflicts over a variety of other resources in addition to food. We propose that future studies detecting apparent dishonesty should analyse many possible signal-state correlations before concluding a signal is actually dishonest. (c) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Cumulative or sequential assessment during hermit crab shell fights: effects of oxygen on decision rules

Agonistic interactions between animals are often settled by the use of repeated signals which advertise the resource-holding potential of the sender. According to the sequential assessment game this repetition increases the accuracy with which receivers may assess the signal, but under the cumulative assessment model the repeated performances accumulate to give a signal of stamina. These models may be distinguished by the temporal pattern of signalling they predict and by the decision rules used by the contestants. Hermit crabs engage in shell fights over possession of the gastropod shells that they inhabit. During these interactions the two roles of signaller and receiver may be examined separately because they are fixed for the duration of the encounter. Attackers rap their shell against that of the defender in a series of bouts whereas defenders remain tightly withdrawn into their shells for the duration of the contest. At the end of a fight the attacker may evict the defender from its shell or decide to give up without first effecting an eviction; the decision for defenders is either to maintain a grip on its shell or to release the shell and allow itself to be evicted. We manipulated fatigue levels separately for attackers and defenders, by varying the oxygen concentration of the water that they are held in prior to fighting, and examined the effects that this has on the likelihood of each decision and on the temporal pattern of rapping. We show that the vigour of rapping and the likelihood of eviction are reduced when the attacker is subjected to low oxygen but that this treatment has no effect on rates of eviction when applied to defenders. We conclude that defenders compare the vigour of rapping with an absolute threshold rather than with a relative threshold when making their decision. The data are compatible with the cumulative assessment model and with the idea that shell rapping signals the stamina of attackers, but do not fit the predictions of the sequential assessment game.

Simulated predator extinctions: Predator identity affects survival and recruitment of oysters

The rate of species loss is increasing at a global scale, and human-induced extinctions are biased toward predator species. We examined the effects of predator extinctions on a foundation species, the eastern oyster (Crassostrea virginica). We performed a factorial experiment manipulating the presence and abundance of three of the most common predatory crabs, the blue crab (Callinectes sapidus), stone crab (Menippe mercenaria), and mud crab (Panopeus herbstii) in estuaries in the eastern United States. We tested the effects of species richness and identity of predators on juvenile oyster survival, oyster recruitment, and organic matter content of sediment. We also manipulated the density of each of the predators and controlled for the loss of biomass of species by maintaining a constant mass of predators in one set of treatments and simultaneously using an additive design. This design allowed us to test the density dependence of our results and test for functional compensation by other species. The identity of predator species, but not richness, affected oyster populations. The loss of blue crabs, alone or in combination with either of the other species, affected the survival rate of juvenile oysters. Blue crabs and stone crabs both affected oyster recruitment and sediment organic matter negatively. Mud crabs at higher than ambient densities, however, could fulfill some of the functions of blue and stone crabs, suggesting a level of ecological redundancy. Importantly, the strong effects of blue crabs in all processes measured no longer occurred when individuals were present at higher-than-ambient densities. Their role as dominant predator is, therefore, dependent on their density within the system and the density of other species within their guild (e.g., mud crabs). Our findings support the hypothesis that the effects of species loss at higher trophic levels are determined by predator identity and are subject to complex intraguild interactions that are largely density dependent. Understanding the role of biodiversity in ecosystem functioning or addressing practical concerns, such as loss of predators owing to overharvesting, remains complicated because accurate predictions require detailed knowledge of the system and should be drawn from sound experimental evidence, not based on observations or generalized models.

Predatory fish loss affects the structure and functioning of a model marine food web

The rate of species loss is increasing on a global scale and predators are most at risk from human-induced extinction. The effects of losing predators are difficult to predict, even with experimental single species removals, because different combinations of species interact in unpredictable ways. We tested the effects of the loss of groups of common predators on herbivore and algal assemblages in a model benthic marine system. The predator groups were fish, shrimp and crabs. Each group was represented by at least two characteristic species based on data collected at local field sites. We examined the effects of the loss of predators while controlling for the loss of predator biomass. The identity, not the number of predator groups, affected herbivore abundance and assemblage structure. Removing fish led to a large increase in the abundance of dominant herbivores, such as Ampithoids and Caprellids. Predator identity also affected algal assemblage structure. It did not, however, affect total algal mass. Removing fish led to an increase in the final biomass of the least common taxa (red algae) and reduced the mass of the dominant taxa (brown algae). This compensatory shift in the algal assemblage appeared to facilitate the maintenance of a constant total algal biomass. In the absence of fish, shrimp at higher than ambient densities had a similar effect on herbivore abundance, showing that other groups could partially compensate for the loss of dominant predators. Crabs had no effect on herbivore or algal populations, possibly because they were not at carrying capacity in our experimental system. These findings show that contrary to the assumptions of many food web models, predators cannot be classified into a single functional group and their role in food webs depends on their identity and density in 'real' systems and carrying capacities.

The costs of bearing arms and armour in the hermit crab Pagurus bernhardus

Hermit crabs use empty gastropod shells as protective armour and enlarged chelipeds as signals and weapons. However, carrying armour and arms may impose energy costs that result in increased lactate and hence potential fatigue and there may be consequent effects on general activity. We investigated whether variation in shell and cheliped size influences lactate levels in hermit crabs. Lactate was positively related to residual cheliped size for both sexes and was higher in males than females; when we controlled for body size, the former had larger chelipeds. Shell weight unexpectedly had no effect on lactate but crabs in small shells had high lactate, possibly because of reduced ability to maintain a respiratory current. The size of natural shells had no effect on activity but the addition of food odour increased locomotion. However, activity was not related to lactate. We conclude that possession of larger chelipeds than expected for body size imposes significant costs and may limit development of sexual dimorphism. (C) 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

No evidence of morphine analgesia to noxious shock in the shore crab, Carcinus maenas

A number of criteria have been suggested for testing if pain occurs in animals, and these include an analgesic effect of opiates (Bateson, 1991). Morphine reduces responses to noxious stimuli in crustaceans but also reduces responsiveness in a non-pain context. Here we use a paradigm in which shore crabs receive a shock in a preferred dark shelter but not if they remain in an unpreferred light area. Analgesia should thus enhance movement to the preferred dark area because they should not experience 'pain'. However, morphine inhibits rather than enhances this movement even when no shock is given. Morphine produces a general effect of non-responsiveness rather than a specific analgesic effect and this could also explain previous studies claiming analgesia. However, we question the utility of this criterion of pain and suggest instead that behavioural criteria be employed. (C) 2011 Published by Elsevier B.V.

Importance of consumers on exposed and sheltered rocky shores.

It is often suggested that the relative importance of biotic processes, such as recruitment, competition and predation of marine benthic species, varies predictably along a gradient of exposure to wave action. Several established models of community dynamics on rocky shores predict that top-down processes are more important for structuring communities on sheltered than on exposed shores. To test the relative dominance of top-down processes, we first measured the establishment of key benthic species (mussels, barnacles and algae) on 3 sheltered and 3 exposed rocky shores in southwest Ireland over two 6 mo periods. We then manipulated the presence of consumers (e.g. grazing gastropods, crabs, whelks), using caged exclosures, on 2 sheltered and 2 exposed shores to test for an interaction between effects of consumers and shore exposure on the establishment of benthic species. In contrast to predictions, we found that consumers strongly affected establishment of all species regardless of shore exposure. We also found that shore exposure was not a reliable predictor for spatial and temporal variation in rates of establishment of sessile benthic species. Our findings provide experimental evidence which demonstrates the importance of consumers in early post-settlement stages of benthic species-essential for the development of benthic-pelagic models. © 2011 Inter-Research.


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The long-term impacts of fisheries on epifaunal assemblage function and structure, in a special area of conservation

Fisheries can have profound effects on epifaunal community function and structure. We analysed the results from five dive surveys (1975–1976, 1980, 1983, 2003 and 2007), taken in a Special Area of Conservation, Strangford Lough, Northern Ireland before and after a ten year period of increased trawling activity between 1985 and 1995. There were no detectable differences in the species richness or taxonomic distinctiveness before (1975–1983) and after (2003–2007) this period. However, there was a shift in the epifaunal assemblage between the surveys in 1975–1983 and 2003–2007. In general, the slow-moving, or sessile, erect, filterfeeders were replaced by highly mobile, swimming, scavengers and predators. There were declines in the frequency of the fished bivalve Aequipecten opercularis and the non-fished bivalves Modiolus modiolus and Chlamys varia and some erect sessile invertebrates between the surveys in 1975–1983 and 2003–2007. In contrast, there were increases in the frequency of the fished and reseeded bivalves Pecten maximus and Ostrea edulis, the fished crabs Cancer pagurus and Necora puber and the non-fished sea stars Asterias rubens, Crossaster papposus and Henricia oculata between the surveys in 1975–1983 and 2003–2007. We suggest that these shifts could be directly and indirectly attributed to the long-termimpacts of trawl fishing gear, although increases in the supply of discarded bait and influxes of sediment may also have contributed to changes in the frequency of some taxa. These results suggest that despite their limitations, historical surveys and repeat sampling over long periods can help to elucidate the inferred patterns in the epifaunal community. The use of commercial fishing gear was banned from two areas in Strangford Lough in 2011, making it a model ecosystem for assessing the long-term recovery of the epifaunal community from the impacts of mobile and pot fishing gear.

Effects of the organophosphorous pesticide, dimethoate, on cardiac and acetylcholinesterase (AChE) activity in the shore crab Carcinus maenas

The potential use of biochemical and physiological responses as biomarkers of organophosphate exposure and/or effect were assessed in the shore crab (Carcinus maenas). Male crabs were assigned to one of four dimethoate treatments (0, 0.5, 1.0 or 2.0 mg 1(-1)). Cardiac activity was measured non-invasively before and during dimethoate exposure using automated interpulse duration assessment. Heart rates decreased significantly in a concentration-dependent manner (p

Automated interpulse-duration assessment (AIDA): A new technique for detecting disturbances in cardiac activity in selected macroinvertebrates

An Automated Interpulse Duration Assessment system (AIDA) is described which permits detection of irregularities in cardiac rhythms in selected invertebrates. The sensitivity of AIDA was demonstrated by its ability to detect handling stress in mussels (Mytilus edulis) that was not evident when measuring heart rate alone. Changes in cardiac activity patterns of crabs (Carcinus maenas) held in the laboratory for up to 10 wk was also examined using the new technique. The frequency distribution of interpulse duration changed significantly as the nutritional state changed. Potential applications of the AIDA system are discussed.