Precocious Courtship and Play in Emydid Turtles

In many mammal species, precocious sexual behaviour is a component of play. A recent model for the evolution of play fighting behaviour in muroid rodents by Pellis (1993) proposes that social play in those species has its origin in precocious sexual behaviour. Captive juvenile emydid turtles perform precocious sexual behaviour. After reviewing its occurrence, we focus on detailed observations of Pseudemys nelsoni. Precocious sexual behaviour in these turtles is characterized by the striking ‘titillation’ display, usually observed only when adult males court females. In this display, the digits of the front feet are rapidly vibrated above the eyes of the female. In our observations, both sexes performed the display, which was energetic, spontaneous, and inconclusive (in that sequences of precocious courtship typically ended when either the displayer or recipient stopped and rested, left, or started some other unrelated non-social behaviour). Displaying individuals clearly preferred conspecific recipients and often appeared to seek out specific individuals. We discuss several plausible explanations for precocious courtship in these turtles but find none entirely satisfying. The characteristics of the precocious courtship behaviour we observed most closely match those commonly listed for social play. Thus, we conclude that the most likely explanation for this behaviour is that it is a component of social play.     

Courtship herding in the fiddler crab <Emphasis Type="Italic">Uca elegans</Emphasis>

Male and female animals are not always complicit during reproduction, giving rise to coercion. One example of a system that is assumed to involve sexual coercion is the mate herding behaviour of fiddler crabs: males push females towards the home burrow with the goal of forcing copulation at the burrow entrance. We recorded and analysed in detail the courtship behaviour of a North Australian species of fiddler crab Uca elegans. Courtship was composed of four main phases: broadcast waving, outward run, herding and at burrow display. During interactions males produced claw-waving displays which were directed posteriorly towards the female and which varied in timing and structure depending on the courtship phase. We suggest that courtship herding in U. elegans is driven primarily by mate choice for the following reasons, (1) females can evade herding, (2) no other reproductive strategies were observed, (3) males broadcast their presence and accompany courtship with conspicuous claw waves, and (4) the behaviour ends with the female leading the male into the home burrow. As an alternative function for herding in U. elegans we suggest that the behaviour represents a form of courtship guiding, in which males direct complicit females to the correct home burrow.     

The evolution of waving displays in fiddler crabs (Uca spp., Crustacea: Ocypodidae)

Male fiddler crabs are commonly recognized by the presence of a single massive claw used in a variety of contexts, including territorial defence, agonistic interactions, and courtship behaviour. The most common behavioural context involving these enlarged chelipeds is their use in waving displays, which are remarkably diverse among species. Although the waving display is one of the most obvious behavioural features of male fiddler crabs, little is known about their main evolutionary trends during the diversification of the genus. The present study employed phylogenetic comparative methods to investigate the evolution of waving behaviour in a sample of 19 species of Uca from Central and North America. Digital recordings were used to quantify the temporal dynamics of waving behaviour in each species. Multivariate ordination methods were used to assess whether different elements of the display showed distinct evolutionary dynamics, particularly with respect to body size and the environment where species are most commonly found. Most of the interspecific variation in displays involves differences in the overall waving velocity, with no correspondence to their local environments, nor their body size. Interestingly, despite the strong concentration of variance in the first two ordination axes, there was no statistically significant evidence for phylogenetic signals in their respective scores. These results suggest that the overall structure of waving displays is evolutionarily labile, at the same time as being concentrated in a few particular axes of variation, possibly indicating evolution along lines of least resistance. The approach employed in the present study highlights the utility of phylogenetic comparative methods for elucidating the evolution of complex behavioural characteristics, such as the waving display in male fiddler crabs. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 307–315.     

Visual Displays in Brachyuran Crabs: Field and Laboratory Studies

Study of the visual displays of more than 30 species of crabsallows some generalizations on the Brachyrhyncha. The displaysfall into two groups, the Lateral Merus and the Chela Forward.Each group contains three major subtypes of display, which originatedas discrete signals of intensity. Several special ritualizationsof the Lateral Merus have evolved in certain genera. The LateralMerus seems to be the primitive type of display in the Brachyuraand still exists in most crabs. It is speculated that the ChelaForward arose from a specialized courtship display but has beenadopted for general agonistic use in some genera.     

Behavioural evidence for a female sex pheromone in Cotesia flavipes (Hymenoptera: Braconidae)

Abstract.  The mating behaviour of Cotesia flavipes Cameron, 1891 (Hymenoptera: Braconidae) was observed under laboratory conditions. The results demonstrate that chemical stimuli play an important role in bringing males and females together for mating. Females from this species release chemical compounds to attract males, which exhibit a distinct courtship behaviour divided into three main steps: (i) antennation, (ii) lateral bouncing and (iii) wing fluttering. Ethological tests using hexane extracts from two different parts of the female's body (head plus thorax and abdomen) demonstrate that males are more attracted and also display a courtship behaviour when a filter paper containing hexane extracts from the abdomen is presented to them, indicating that the abdomen is the source of the sex pheromone of this species. Naive males of this species respond to hexane extracts of this gland by displaying courtship behaviour. Furthermore, additional tests show that C. flavipes females mate only once, whereas virgin males do not show any preference for virgin or mated females, suggesting that mated females of this species continue to produce sex attractants after mating.     

The use of open burrows to estimate abundances of intertidal estuarine crabs

Estimation of absolute (or true) abundances of intertidal burrowing crabs is a difficult problem in some estuarine habitats because the nature of the substratum and behaviour of crabs can restrict researchers to the use of sampling methods which at best estimate only apparent, or relative, abundances. One method that has been used is to count open burrows to estimate population densities. This paper discusses the results of a test examining the validity of using burrows to estimate apparent abundances of a temperate ocypodid crab, Heloecius cordiformis. inhabiting mangrove forests of south-eastern Australia. Under appropriate circumstances, this method may provide a quick and reliable estimate of abundance of crabs.     

It Takes Two to Tango: Relative Influence of Male and Female Identity and Morphology on Complex Courtship Display in a Newt Species

Consistency in behaviour is currently receiving a renewed interest. Although courtship display is generally consistent in terms of behavioural sequence and structure, there is also commonly important variation in the intensity of courtship display between and within males of a given species. Indeed, not all males have the same ability to perform courtship display (variation between males), and each male can potentially adjust his courtship effort in response to the environment (variation within a male). Although the study of male courtship display has received considerable attention in recent years, it is still unclear which part of the variation can be explained by male ability or motivation. We investigated this issue on two phases of the complex courtship display of the palmate newt Lissotriton helveticus. Overall, we found that both male and female identities affected courtship behaviour, but the relative influence of each sex depended on the courtship phase. Male identity explained variation in fan and creep‐quiver display, whereas female identity explained variation in creep‐quiver only. Interestingly, we did not find any link between the expression of courtship display and male or female morphological traits. Our study showed consistency of male courtship display in newts and successfully dissects the different sources of variation that can affect behavioural repeatability/consistency of courtship display.     

Costs increase as ritualized fighting progresses within and between phases in the sierra dome spider, Neriene litigiosa

Magnitudes and patterns of energy expenditure in animal contests are seldom measured, but can be critical for predicting contest dynamics and understanding the evolution of ritualized fighting behaviour. In the sierra dome spider, males compete for sexual access to females and their webs. They show three distinct phases of fighting behaviour, escalating from ritualized noncontact display (phase 1) to cooperative wrestling (phase 2), and finally to unritualized, potentially fatal fighting (phase 3). Using CO₂ respirometry, we estimated energetic costs of male-male combat in terms of mean and maximum metabolic rates and the rate of increase in energy expenditure. We also investigated the energetic consequences of age and body mass, and compared fighting metabolism to metabolism during courtship. All three phases involved mean energy expenditures well above resting metabolic rate (3.5×, 7.4× and 11.5×). Both mean and maximum energy expenditure became substantially greater as fights escalated through successive phases. The rates of increase in energy use during phases 2 and 3 were much higher than in phase 1. In addition, age and body mass affected contest energetics. These results are consistent with a basic prediction of evolutionarily stable strategy contest models, that sequences of agonistic behaviours should be organized into phases of escalating energetic costs. Finally, higher energetic costs of escalated fighting compared to courtship provide a rationale for first-male sperm precedence in this spider species.     

On display behaviour and courtship in the praying mantis Ephestiasula amoena (Bolivar)

Display behaviour in a small species of Indian praying mantis, Ephestiasula amoena , is described and the possible role of this display in courtship is discussed. The display involves exposure of the bright pattern on the inside face of the expanded plate-like profemora. The profemoral patterns found in closely related species are discussed and it is suggested that the display may also play a role in species recognition. The courtship behaviour of E. amoena is contrasted and compared with the courtship of other species of praying mantis.     

Coordination and neuromuscular control of rhythmic behaviors in the blue crab,Callinectes sapidus

The stereotypical courtship display (CD) behavior of the male blue crab, Callinectes sapidus, includes an unusual component: the rhythmic waving of the swimming appendages above the carapace. This behavior occurs in a unique context but it resembles two other rhythmic behaviors performed using the swimming legs: sideways swimming and backward swimming. As a first step to understanding the mechanisms that allow the expression of apparently different rhythmic motor patterns, we have examined these behaviors using slow motion video analysis and electromyography of the basal muscles of the swimming legs in freely behaving crabs. The results show that these behaviors are distinguished by four parameters: the frequency of leg waving, the phase relationship between the legs, the presence of a stationary pause in basal muscle activity combined with rotation of the distal leg during CD, and an extended range of motion of these legs during CD and backward swimming, relative to sideways swimming. EMG analysis revealed that during sideways swimming, the sequence of muscular activity between the two legs was different. In contrast, during CD and backward swimming the sequence of activity for these legs is identical.Abbreviations CD courtship display - EMGs electromyograms - CD AMP courtship display in crabs with amputated fifth legs - CD1 crabs that voluntarily used one leg to perform courtship display waving - CD 1–3 courtship waving in cycles 1–3 - CD MID courtship waving after cycles 1–3 - M-C meral-carpal joint     

The Problem of Adult Play Fighting: A Comparative Analysis of Play and Courtship in Primates

Although play fighting, like play generally, is predominantly a feature of the juvenile phase, such behavior persists in the adults of many species. There are two major contexts in which adults engage in play fighting – with juveniles and with other adults. The least attention has been given to adult–adult play. However, one pattern that has been noted by several authors is that the most commonly occurring context of adult–adult play fighting is during courtship, and that this is more likely to occur in solitary species. Supposedly, such play could function to overcome the aggressiveness of potential pairmates unfamiliar with one another, or as a means of evaluating mate quality by one or both partners. By contrasting the presence and degree of play fighting during courtship with the degree of male–female familiarity, the hypothesis that the former is influenced by the latter is tested. Data on 35 species of primates, from 15 families, were compiled from the literature and compared using a method of independent contrasts that incorporates information on phylogenetic relationships. A significant regression was found, with the degree of male–female familiarity accounting for 40% of the variance in courtship play. Therefore, our data support the hypothesis that play fighting in courtship is influenced by male–female patterns of association. However, the data also indicate that other factors must influence the occurrence of play fighting amongst adults, not only during courtship, but also in nonsexual contexts. The broader context of adult–adult play in mammals is discussed.     

Keeping up appearances: male fiddler crabs wave faster in a crowd

Courtship displays are often energetically and temporally costly as well as highly conspicuous to predators. Selection should therefore favour signalling tactics that minimize courtship costs while maintaining or increasing signal attractiveness. In fiddler crabs, males court females by waving their one greatly enlarged claw in a highly conspicuous and costly display. Here, we investigate whether courting males adjust their wave rate, and therefore the cost of courtship, to the current level of competition. We show that display rate increases as competition increases and that when competition is removed, males reduce their display rate by 30 per cent. These results suggest that male fiddler crabs actively reduce the cost of courtship by adjusting their wave rate in response to the immediate level of competition.     

The feeding behaviour of the sand crab Scopimera inflata (Decapoda, Ocypodidae)

A sequence of activities all related to the feeding of Scopimera inflata H. Milne-Edwards, 1873 can be distinguished during the daytime periods of low tide. Emergence begins one to two hours after a feeding area is uncovered, reaches a peak after a further three hours but some crabs are still emerging after the tide has turned. Before feeding commences the burrow is cleared of waste sand down to the water-table. While feeding S. inflata forms a feeding trench which acts as an escape route back to the hole and is defended against other crabs of the same species. Agonistic behaviour becomes evident as feeding progresses. Threat displays, fighting, displacement feeding, and agitation feeding are recognizable behaviour patterns which have been observed.     

Eavesdropping in crabs: an agency for lady detection

Although conspicuous courtship displays are an effective way of attracting the attention of receptive females, they could provide valuable information to rival males on the location of these females. In fiddler crabs, males that see a receptive female wave their single, greatly enlarged claw in a highly conspicuous courtship display. We test whether other males use this courtship display to alert them to the presence of receptive females that they cannot directly see. We show that male fiddler crabs (Uca mjoebergi) eavesdrop on the courtship displays of nearby males to detect mate-searching females. This allows males to begin waving before a female becomes visible. Furthermore, males appear to adjust their waving according to the information available: eavesdropping males wave 12 times faster than non-courting males but only 1.7 times slower than males in full visual contact with the female.     

The unusual inheritance pattern of the courtship songs in closely related grasshopper species of the Chorthippus albomarginatus-group (Orthoptera: Gomphocerinae)

Sibling species of the Chorthippus albomarginatus-group are exceptional among all European grasshopper species because they have an extremely elaborate courtship behaviour. Here, we present a genetic analysis of the courtship song differences between two closely related grasshopper species Ch. albomarginatus and Ch. oschei. Measurements of seven courtship song characters and one parameter of courtship visual display were compared among parentals, F(1) and F(2) hybrids, and backcrosses. Variation in one morphological character, the number of stridulatory pegs, was studied as well. The distributions of song phenotypes were more consistent with a type-III genetic architecture, which involves complementary or duplicate pairs of loci. We suggest that this type of architecture may provide the novel elements of the hybrid songs, even more complex than the parental ones, which may offer a new starting point for sexual selection. One parameter of the visual display appeared to be controlled by a single locus, but differences in other characters of visual display could be explained by a more complex inheritance pattern. Inheritance of the peg number differences was consistent with a simple polygenic additive model. Sex linkage and/or maternal effects were not detected for any trait. We found a disproportionate influence of the Ch. albomarginatus parent on most hybrid song characteristics and the visual display, and this may support a previously suggested idea that Ch. albomarginatus is an ancestral form and Ch. oschei is a derived species.     

Comparative Studies of Reproductive Behavior in Mantis Shrimps and Fiddler Crabs

Comparative ethological studies of closely related species canplace into clear perspective the functions of behavioral traitsas species diverge and adapt to different environments. In thispaper we contrast the reproductive behavior of species withintwo crustacean groups: the mantis shrimps (stomatopods) andthe fiddler crabs (genus Uca). For the stomatopods, we identifyprobable selective relationships between features of species'ecology and their mating systems. Population density, the kindand availability of shelters in which these shrimps live, theintensity of predation and spawning cycles all play importantroles in molding reproductive behavior in this group. Associationsbetween the ecology and mating systems of fiddler crabs havebeen discussed recently elsewhere. Here we focus on a comparativeanalysis of sexual communication in these crabs. A study ofcompetitive courtship signaling and mate choice in the fiddlercrab Uca beebei has shown that certain male courtship signalsare highly attractive probably because they exploit female sensory-responsesystems that have been molded by selection for escape from predators.Interspecific comparison of male courtship displays and theresponses of females to these displays suggest that sensoryexploitation may play an important role in the evolution ofsexual signals in the genus. Comparative studies have advancedour understanding of how natural and sexual selection affectthe reproductive behavior of both stomatopods and fiddler crabs.     

Ecology and signal structure drive the evolution of synchronous displays*

Animal synchrony is found in phylogenetically distant animal groups, indicating behavioral adaptations to different selective pressures and in different signaling modalities. A notable example of synchronous display is found in fiddler crabs in that males wave their single enlarged claw during courtship. They present species-specific signals, which are composed of distinctive movement signatures. Given that synchronous waving has been reported for several fiddler crab species, the display pattern could influence the ability of a given species to sufficiently adjust wave timing to allow for synchrony. In this study, we quantified the wave displays of fiddler crabs to predict their synchronous behavior. We combined this information with the group's phylogenetic relationships to trace the evolution of display synchrony in an animal taxon. We found no phylogenetic signal in interspecific variation in predicted wave synchrony, which mirrors the general nonphylogenetic pattern of synchrony across animal taxa. Interestingly, our analyses show that the phenomenon of synchronization stems from the peculiarities of display pattern, mating systems, and the complexity of microhabitats. This is the first study to combine mathematical simulations and phylogenetic comparative methods to reveal how ecological factors and the mechanics of animal signals affect the evolution of the synchronous phenomena.     

Evolution of Courtship Behaviour Patterns and Reproductive Isolation in the Desmognathus ochrophaeus Complex

The extent to which differences in courtship behaviour patterns act as mechanisms of reproductive isolation is critical to understanding both speciation and the evolution of these behaviour patterns. While numerous studies have investigated intraspecific and interspecific differences in courtship, fewer interpret results in a phylogenetic framework. We describe and analyse geographic variation in the courtship behaviour patterns of the Allegheny Dusky salamander ( Desmognathus ochrophaeus ). We then examine courtship among closely related species in the D. ochrophaeus complex in a phylogenetic context. We found that populations of D. ochrophaeus separated by extensive geographic distances show little variation in courtship behaviour patterns and are sexually compatible. This contrasts with significant levels of sexual isolation between D. ochrophaeus and other species in the complex. Mapping behaviour patterns onto a phylogeny that we generated from cytochrome b sequences indicates that two behaviour patterns present in the courtship sequence of other members in the complex have either been lost in D. ochrophaeus or gained independently in other species in the complex. Loss of these behaviour patterns may result in reproductive isolation between D. ochrophaeus and its sister taxon, D. orestes .