Size-assortative pairing in Gammarus pulex (Crustacea : Amphipoda): a test of the timing hypothesis

We present the first empirical test of the timing hypothesis regarding the generation of size-assortative pairing in amphipods. The timing hypothesis proposes that, since large males are better able to afford the costs of mate guarding than small males, the former can take larger females into precopula earlier in the female moult cycle than is feasible for the latter. This leaves small males to form pairs with smaller females closer to moult, thus generating size assortment. We presented male Gammarus pulex, collected both in precopula and as singletons, with females that were (1) previously guarded and therefore near to copulatory moult and (2) previously unguarded and therefore far from copulatory moult. This comparison tested the prediction of the timing hypothesis, that size assortment should break down when the opportunity for time-based male decisions is removed, but that size assortment should occur where timing is not disrupted. Counter to the hypothesis, we found that size assortment did not break down upon removal of the time factor. Large males tended to initiate mate guarding earlier than small males in both female moult groups. However, only in the previously unguarded group did large males guard for longer than small males. This result suggests that, although size assortment occurred in all groups, the causative mechanisms that generated this pattern may differ between these groups. We therefore consider the possible importance of mechanisms such as aggression, simultaneous manipulation of females and female resistance in producing size assortment where males encounter numerous females that are close to moult. We also observed that prior recent guarding experience by males had no effect on latency to guard or size-assortative pairing. (C) 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

Maternal care in the rockpool amphipod Apherusa jurinei: developmental and environmental cues

There is evidence that active, pre-emergence maternal brood care in amphipod crustaceans may be associated with 'harsh' environmental conditions. We examined, in the rockpool amphipod Apherusa jurinei, behavioural activities that may function as a form of active brood care. Only ovigerous females showed 'curl' and 'stretch' activities, with consequent flushing of the brood pouch and cycling of the eggs therein. There was a significant decline in these activities as embryonic development advanced and brood care almost ceased when well-developed embryos showed a heart pulse and self-ventilation. We propose that this pattern of brood care reflects changes in the physiological requirements of embryos as they develop within the egg membrane. In addition, ovigerous females showed significantly higher levels of brood care under lowered oxygen conditions. They achieved this by increasing the average duration of the 'stretch' component, with other brood care components remaining constant. Thus, developmental and environmental cues alter the components of active brood care in distinct ways. Experimental removal showed that the physical presence of eggs in the brood pouch is important in controlling the expression of brood care activities. However, females with all of their eggs removed continued to brood at low levels, suggesting that a maternal state also controls brood care. The sophisticated expression of active maternal brood care in amphipods under 'harsh' environmental conditions such as rockpools has implications both for individual reproductive success and the distribution and abundance of brooding versus nonbrooding species. (C) 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

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.

Responses to interactive playback predict future pairing success in nightingales

Birdsong is a sexually selected trait that serves in territory defence and mate choice. Individual song traits can be affected by the body condition of the male and thus may reflect his quality. Such relations between male quality and general singing performance raise the question whether differences in male quality also affect response strategies used in dyadic interactions. To address this question, we studied the relation between pairing success of male common nightingales, Luscinia megarhynchos, and their responses to rivals posing different levels of threat. Using interactive playback, we exposed males prior to mating to either aggressively or moderately singing rivals (by song overlapping and song alternating, respectively). Males that remained unpaired throughout the season (bachelors) interrupted their singing significantly more often after being overlapped than after alternating playback, whereas subsequently mated males kept the number of singing interruptions more constant across playback treatment. This suggests that subsequently paired males are less discriminative than are bachelors when challenged by rivals varying in aggressiveness. Regardless of playback treatment, males that later became paired responded significantly more strongly than did bachelor males. Thus, an increase in singing after a vocal interaction prior to mating predicted future mating success. (c) 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Early birds are sexy: male age, dawn song and extrapair paternity in blue tits, Cyanistes (formerly Parus) caeruleus

Sexual selection theory predicts that signals reflecting the relative quality of individuals should be used in mate choice. Females could base their choice of copulation partners on male secondary sexual traits that honestly signal male age, as predicted by the age-based indicator mechanism. Studies have shown that female blue tits prefer older males and that aspects of dawn song reflect male quality, but it remains unknown whether dawn song characteristics correlate with male age. We compared dawn song characteristics of second-year (SY) and older (ASY) male blue tits (cross-sectional analysis), and tested for age-related changes within individuals (longitudinal analysis) and differential overwinter survival of SY males. We further investigated the relation between dawn song and paternity gain and loss. We found that ASY male blue tits began to sing earlier relative to sunrise than did SY males. This difference in the onset of dawn singing was due to age-related changes in individual performance rather than differential survival of individuals with varying expression of the trait. Males that began to sing earlier at dawn had more mating partners, and were more likely to gain extrapair paternity. Our findings suggest that the onset of dawn song can provide a simple mechanism for females to assess the relative quality of their mate and of neighbouring males. We propose that females use the onset of singing as a cue for their choice of extrapair partners. (c) 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Seasonal variation in dawn song characteristics in the common nightingale

Many male temperate zone passerines show a marked peak of singing activity before sunrise. The two main functions of this so-called dawn chorus are mate attraction and territory defence. We examined how seasonal patterns of different dawn song characteristics were related to mating status and to the breeding cycle of females in the common nightingale, Luscinia megarhynchos. We investigated two measures of song output: song rate and percentage performance time. We also calculated the proportion of 'whistle songs', a song category that is thought to be important in female choice. We predicted that if the main function of dawn singing in nightingales is to attract a social mate, then mated males should change their dawn singing behaviour after pair formation. In contrast, if dawn singing is mainly used in territory defence, we expected no difference in song traits between mated and unmated males throughout the season. We found that song rate and the proportion of whistle songs were low at the beginning of the season and did not predict future mating status. After arrival of females, all measures of dawn song performance remained largely constant throughout the breeding season, and we did not find significant differences in the seasonal variation between mated and unmated males. These findings are consistent with the hypothesis that song at dawn is important to defend a territory throughout the breeding season. (c) 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

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.

Nociception or pain in a decapod crustacean?

Nociception is the ability to perceive a noxious stimulus and react in a re flexive manner and occurs across a wide range of taxa. However, the ability to experience the associated aversive sensation and feeling, known as pain, is not widely accepted to occur in nonvertebrates. We examined the responses of a decapod crustacean, the prawn, Palaemon elegans, to different noxious stimuli applied to one antenna to assess reflex responses (nociception) and longer-term, specifically directed behavioural responses that might indicate pain. We also examined the effects of benzocaine, a local anaesthetic, on these responses. Noxious stimuli elicited an immediate reflex tail flick response, followed by two prolonged activities, grooming of the antenna and rubbing of the antenna against the side of the tank, with both activities directed specifically at the treated antenna. These responses were inhibited by benzocaine; however, benzocaine did not alter general swimming activity and thus the decline in grooming and rubbing is not due to general anaesthesia. Mechanical stimulation by pinching also resulted in prolonged rubbing, but this was not inhibited by benzocaine. These results indicate an awareness of the location of the noxious stimuli, and the prolonged complex responses indicate a central involvement in their organization. The inhibition by a local anaesthetic is similar to observations on vertebrates and is consistent with the idea that these crustaceans can experience pain.

The less amorous Gammarus: predation risk affects mating decisions in Gammarus duebeni (Amphipoda)

We examined the trade-off between the behaviours associated with predator avoidance and mate acquisition in the mate-guarding amphipod crustacean Gammarus duebeni. We used laboratory experiments to investigate the impact of olfactory predator cues on activity, mate choice and mate-guarding behaviour of males and females. Pair formation declined under perceived risk of predation, reflecting reduced activity of both males and females and hence a reduced likelihood of encountering a mate. We also observed a reduction in the choosiness of both males and females. Under increased perceived predation risk, assessment of the female by the male was more likely to be followed by pair formation, and males showed a nonsignificant trend towards reduced discrimination in favour of large females and were less tenacious in their pair bond when they paired during exposure to predator cues. Females also showed less resistance behaviour, suggesting that both males and females trade off the costs of maximizing current reproductive success against the benefits of predator avoidance for survival and reproduction in the future. We discuss the implications of such context-dependent mating behaviours for ecological interactions between species and suggest that predators, via the effects of perceived predation risk on mate choice and mate guarding in the prey species, induce trait-mediated indirect effects with the potential to influence population dynamics and community structure. (C) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Embryo retrieval and kin recognition in an amphipod (Crustacea)

Active maternal care directed towards embryos within the brood pouch has been identified in amphipod crustaceans from harsh aquatic environments. This involves 'curl' and 'stretch' components and brood flushing that alters in distinct ways in response to developmental and environmental cues. However, a cost of active brood care in crustaceans is the susceptibility to embryo loss, this being further predisposed by the structure of the amphipod brood pouch. We found embryo retrieval by females of the rock-pool amphipod Apherusa jurinei, whereby females inserted experimentally offered embryos into their brood pouches. Females early in brood development retrieved embryos to a greater degree than both nonovigerous and later stage females. In this experiment, all offered embryos were from other females, indicating a motivation to retrieve embryos that often overrides any kin recognition. In a second experiment, we found kin discrimination, with both early stage and late stage females retrieving more of their own embryos than those from other females. Recognition was not simply of embryos of similar developmental stages. There were high levels of embryo cannibalism in both experiments, but females were significantly less likely to consume their own compared to foreign embryos. We thus further show that 'lower' crustaceans such as amphipods engage in elaborate active maternal care including kin recognition and discrimination. Their maternal behaviour appears to balance the costs and benefits of embryo retrieval, minimizing fitness reductions due to embryo loss and adoption of foreign embryos. (C) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. 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.