Eavesdropping and signal matching in visual courtship displays of spiders

Eavesdropping on communication is widespread among animals, e.g. bystanders observing male-male contests, female mate choice copying and predator detection of prey cues. Some animals also exhibit signal matching, e.g. overlapping of competitors' acoustic signals in aggressive interactions. Fewer studies have examined male eavesdropping on conspecific courtship, although males could increase mating success by attending to others' behaviour and displaying whenever courtship is detected. In this study, we show that field-experienced male Schizocosa ocreata wolf spiders exhibit eavesdropping and signal matching when exposed to video playback of courting male conspecifics. Male spiders had longer bouts of interaction with a courting male stimulus, and more bouts of courtship signalling during and after the presence of a male on the video screen. Rates of courtship (leg tapping) displayed by individual focal males were correlated with the rates of the video exemplar to which they were exposed. These findings suggest male wolf spiders might gain information by eavesdropping on conspecific courtship and adjust performance to match that of rivals. This represents a novel finding, as these behaviours have previously been seen primarily among vertebrates.     

Vibratory Communication in the Jumping Spider Phidippus clarus: Substrate‐borne Courtship Signals are Important for Male Mating Success

Recently, work has shown that multimodal communication is common throughout the animal kingdom but the function of multimodal signals is still poorly understood. Phidippus clarus are jumping spiders in which males produce multimodal (visual and vibrational) signals in both male–male (aggressive) and male–female (courtship) contexts. The P. clarus mating system is complex, with sex ratios and the level of male competition changing over the course of the breeding season. Vibrational signal components have been shown to function in male aggressive contests but their role in courtship has not been investigated. Here, we performed an experiment to test the role of vibrational signaling in courtship by observing mating success for males that were experimentally muted. We show that vibratory courtship signals, and in particular signaling rate, is an important component of mating success and potentially a target of female choice. While the ability to produce vibratory signals significantly increased mating success, some muted males were still able to successfully mate. In these trials, signaling rate also predicted mating success suggesting that redundant signal components may compensate for errors and perturbations in signal transmission or that vibratory signals function to enhance the efficacy of visual signals.     

Male collared lizards,Crotaphytus collaris (Sauria: Crotaphytidae), signal females by broadcasting visual displays

The signalling function of displays broadcast when animals are distant from conspecifics can be difficult to determine. I tested the extent to which visually transmitted broadcast displays given by free‐ranging territorial male collared lizards signalled same‐sex rivals or females. One test involved recording the frequency of broadcast displays, aggressive contests with rivals, and courtship encounters with females during ten reproductive seasons when local sex ratios varied markedly. The frequency of broadcast displays decreased as the ratio of male competitors to females increased. The frequency with which males initiated contests with rivals was not related to the ratio of competitors to females, whereas the frequency of courtship interactions decreased with sex ratio because there were fewer females to court. The behaviour of males that defended territories during two successive seasons showed a similar pattern. Broadcast display frequency was positively correlated with courtship frequency, but not with the frequency of contests with rivals. Lastly, individual males gave more broadcast displays during focal observations when they also engaged in courtship encounters with females than other observations when they engaged in aggressive conflicts with rival males. Although these results do not reject the possibility that broadcast displays may also signal male rivals, they support a major role of these displays in advertisement to females. © 2013 The Linnean Society of London     

Social context influences aggressive and courtship behavior in a cichlid fish

Social interactions require knowledge of the environment and status of others, which can be acquired indirectly by observing the behavior of others. When being observed, animals can also alter their signals based on who is watching. Here we observed how male cichlid fish (Astatotilapia burtoni) behave when being watched in two different contexts. In the first, we show that aggressive and courtship behaviors displayed by subordinate males depends critically on whether dominant males can see them, and in the second, we manipulated who was watching aggressive interactions and showed that dominant males will change their behavior depending on audience composition. In both cases, when a more dominant individual is out of view and the audience consists of more subordinate individuals, those males signal key social information to females by displaying courtship and dominant behaviors. In contrast, when a dominant male is present, males cease both aggression and courtship. These data suggest that males are keenly aware of their social environment and modulate their aggressive and courtship behaviors strategically for reproductive and social advantage.     

The role of chemoreception in sex recognition by male crickets: Acheta domesticus and Teleogryllus oceanicus

ABSTRACT. Contact chemoreception is important in female recognition by Teleogryllus oceanicus (Le Guillou) males. Antennal contact of female conspecifics, body regions, detached antennae and conditioned substrate elicited mostly courtship responses including courtship songs. Aggressive acts were produced only in response to male conspecifics. Male body regions, detached antennae and conditioned substrate elicited very few courtship or aggressive acts and no songs. This suggests that one or several communication modes, in addition to chemical communication, are necessary to elicit aggressive responses. Acheta domesticus (L.) males cannot rely upon chemical cues for recognition of either sex. Responses to conspecifics suggest that A. domesticus males produce aggressive acts immediately after antennal contact with either sex. Aggressive response to males usually persists, but response to females often switches to courtship. Responses to body regions, detached antennae, and conditioned substrate were few, with courtship and aggressive responses elicited by both male- and female-generated stimuli. The importance of contact chemoreception in cricket communication is suggested by (1) failure of hexane-washed antennae to elicit aggressive or courtship acts, and (2) males spending more time in contact with body regions and conditioned substrates than with corresponding controls. Lack of response to male or female odour-laden air suggests that chemical signals are used by males only if directly contacted. Chemical and other signals supplement the obvious use of acoustic signals for intra- and intersexual communication in these crickets. The importance of multimodal communication in sex recognition is discussed.     

Court and spark: electric signals in the courtship and mating of gymnotoid fish

By mimicking tropical rainy season conditions in aquaria, we stimulated two species of gymnotoid electric fish, Eigenmannia virescens and Apteronotus leptorhynchus, to spawn in captivity. Their courtship activity, breeding behaviour and electric social communication were monitored in several groups over 2 years. Groups of both species established dominance hierarchies correlated with electric organ discharge frequency, aggressiveness and size. Spawning was preceded by several nights of courtship during which the male modulated its electric organ discharge to produce ‘chirps’. Continual bouts of chirping lasted for hours on evenings prior to spawning. These electrical signals play a significant role in courtship and spawning, as gravid E. virescens females could be stimulated to spawn by playing back into the tank a tape recording of male courtship chirps. In both species the chirp invovves a slight increase in frequency followed by a cessation of the dominant frequency. This suggests a common mode of signal production in these two different genera of fish. Chirps are short and abrupt during aggressive encounters, but assume a softer and more raspy quality during courtship.     

Bat echolocation calls facilitate social communication

Bat echolocation is primarily used for orientation and foraging but also holds great potential for social communication. The communicative function of echolocation calls is still largely unstudied, especially in the wild. Eavesdropping on vocal signatures encoding social information in echolocation calls has not, to our knowledge, been studied in free-living bats so far. We analysed echolocation calls of the polygynous bat Saccopteryx bilineata and found pronounced vocal signatures encoding sex and individual identity. We showed experimentally that free-living males discriminate approaching male and female conspecifics solely based on their echolocation calls. Males always produced aggressive vocalizations when hearing male echolocation calls and courtship vocalizations when hearing female echolocation calls; hence, they responded with complex social vocalizations in the appropriate social context. Our study demonstrates that social information encoded in bat echolocation calls plays a crucial and hitherto underestimated role for eavesdropping conspecifics and thus facilitates social communication in a highly mobile nocturnal mammal.     

Phenotypic Plasticity and the Ecotypic Differentiation of Aggressive Behavior in Threespine Stickleback

The threespine stickleback fish, Gasterosteus aculeatus, has undergone a remarkable postglacial adaptive radiation in which an ancient oceanic ancestor has given rise to uncountable freshwater populations. The radiation is characterized by repeated, independent evolution of similar derived phenotypes under similar environmental conditions. A common pattern of divergence is caused by differences in habitat that favor morphological and behavioral features that enhance efficiency of feeding on plankton (limnetic ecotypes) vs. those that enhance efficiency of feeding on benthic invertebrates (benthic ecotypes). These two ecotypes exhibit consistently different patterns of courtship and of foraging and cannibalistic behavior (divergent behavioral syndromes). Here, we demonstrate that there also exist differences in aggression toward conspecifics that are likely to be characteristic of the ecotypes. We report differences in patterns of aggression toward rivals between the ecotypes and offer evidence of differences in the patterns of phenotypic plasticity (norms of reaction) for these traits across population types, and of differences in the incorporation of aggressive elements of behavior in courtship. These data support an earlier suggestion that differences in aggressive tendencies could have facilitated assortative mating between the four benthic–limnetic species pairs found in British Columbia lakes, and they demonstrate the need to evaluate divergent behavioral phenotypes in this radiation as phenotypic norms of reaction rather than as fixed traits.     

Energy and predation costs of firefly courtship signals

Animal courtship signals include many highly conspicuous traits and behaviors, and it is generally assumed that such signals must balance the benefits of attracting mates against some fitness costs. However, few studies have assessed the multiple costs potentially incurred by any one courtship signal, so we have limited understanding of the relative importance of different costs. This study provides the first comprehensive assessment of signal costs for Photinus fireflies (Coleoptera: Lampyridae), using controlled experiments to measure both the energy and predation costs associated with their bioluminescent courtship signals. We measured energy required to generate bioluminescent flashes, using differential open-flow respirometry, and found that flash signaling results in only a nominal increase in energy expenditure above resting levels. These results suggest that the energy required to generate bioluminescent flashes represents a minor component of the total cost of firefly courtship. However, controlled field experiments revealed that visually oriented predators imposed major costs on firefly courtship signals, with higher signaling rates significantly increasing the likelihood of predation. Together with previous results demonstrating that female fireflies prefer more conspicuous courtship signals, these results support the importance of multiple-receiver communication networks in driving signal evolution.     

Acoustic signals in insects: A reproductive barrier and a taxonomic character

In singing insects, the song is an important component of the specific mate recognition system (SMRS). In communities of sympatric singing species, there is a partitioning of communication channels, the so-called “acoustic niches.” Within one community, the songs of different species always differ in temporal or frequency characters, i.e. occupy different acoustic niches. However, conspecific songs do not always act as an interspecific reproductive barrier, despite always being a SMRS component. The species that do not communicate acoustically due to allopatry, different timing of vocalization, inhabiting different biotopes, or unmatched food specializations can produce similar songs while forming reproductively isolated communities. Individuals of different sexes need not only to recognize a conspecific mate but also to evaluate its “quality.” The close-range signal (courtship song) provides more opportunities for choosing the “best” male than does the distant signal (calling song). In many species of Orthoptera, courtship includes not only acoustic but also vibrational, visual, chemical, and mechanical signals. An analysis of cricket songs showed the courtship songs to be on average more elaborate and variable than the calling songs. At the same time, due to the difference in mating behavior between the two groups, the acoustic component of courtship is used for mate quality evaluation to a greater extent in grasshoppers than in crickets. The courtship songs of grasshoppers are generally more elaborate in temporal structure than cricket songs; moreover, they may be accompanied by visual displays such as movements of various body parts. Thus, song evolution in grasshoppers is more strongly driven by sexual selection than that in crickets. According to the reinforcement hypothesis, the premating barrier between hybridizing species becomes stronger in response to reduced hybrid fitness. However, our behavioral experiments with two groups of hybridizing grasshopper species did not confirm the reinforcement hypothesis. We explain this, firstly, by a low level of genetic incompatibility between the hybridizing species and secondly, by high hybrid fitness when attracting a mate. A high competitive capability of hybrids may be accounted for by attractiveness of new elements in hybrid courtship songs. When we divide similar forms based on their songs, we in fact distinguish biological species using the criterion of their reproductive isolation. Acoustic differences between species are usually greater than morphological ones. Therefore, song analysis allows one to determine the real status of doubtful species-rank taxa, to distinguish species in a medley of sibling forms, and to reveal cryptic species in the cases when morphological studies fail to provide a univocal result. At the same time, songs are subject to intraspecific variation the range of which is different in different groups. Therefore, it is necessary to study which degree of difference corresponds to the species level before interpreting the status of some forms based on song comparisons. Besides, song similarities cannot indicate conspecificity of acoustically isolated forms; on the other hand, song differences between these forms prove that they are full-rank species.     

Female signals enhance the efficiency of mate assessment in satin bowerbirds (Ptilonorhynchus violaceus)

Recent evidence suggests that males adjust their sexually selecteddisplay traits in response to female behaviors during courtships.Little is known, however, about whether females signal to influencemale displays and whether females benefit from this interaction.Male courtship displays in the satin bowerbird (Ptilonorhynchusviolaceus) are highly intense and aggressive. Females may usethese displays as indicators of mating benefits, but these displaysoften startle females and disrupt courtship. Previous studieshave shown that successful males decrease female startling byadjusting their display intensity according to female crouchingbehaviors, suggesting that crouching behaviors function as signals.Here we address whether female crouching is a signal by usingobservations of natural courtship behaviors. In addition, weexamine why females differ in signaling and whether femalesbenefit from signaling. First, we find that female crouchingis related to the likelihood that females will be startled bymale displays, suggesting that crouching signals the degreeof display intensity that females will tolerate from a malewithout being startled. Second, we find that female tolerancefor intense display increases during successive courtships asfemales assess potential mates, and that female tolerance mayalso be affected by age and condition. Third, we find evidencethat females that reduce startling by signaling their intensitytolerance are more efficient in mate searching. These resultssuggest that females signal to influence how males display theirsexually selected traits, and by doing so, females may increasetheir benefits in mate choice.     

The African cichlid fish Astatotilapia burtoni uses acoustic communication for reproduction: sound production, hearing, and behavioral significance

Sexual reproduction in all animals depends on effective communication between signalers and receivers. Many fish species, especially the African cichlids, are well known for their bright coloration and the importance of visual signaling during courtship and mate choice, but little is known about what role acoustic communication plays during mating and how it contributes to sexual selection in this phenotypically diverse group of vertebrates. Here we examined acoustic communication during reproduction in the social cichlid fish, Astatotilapia burtoni. We characterized the sounds and associated behaviors produced by dominant males during courtship, tested for differences in hearing ability associated with female reproductive state and male social status, and then tested the hypothesis that female mate preference is influenced by male sound production. We show that dominant males produce intentional courtship sounds in close proximity to females, and that sounds are spectrally similar to their hearing abilities. Females were 2-5-fold more sensitive to low frequency sounds in the spectral range of male courtship sounds when they were sexually-receptive compared to during the mouthbrooding parental phase. Hearing thresholds were also negatively correlated with circulating sex-steroid levels in females but positively correlated in males, suggesting a potential role for steroids in reproductive-state auditory plasticity. Behavioral experiments showed that receptive females preferred to affiliate with males that were associated with playback of courtship sounds compared to noise controls, indicating that acoustic information is likely important for female mate choice. These data show for the first time in a Tanganyikan cichlid that acoustic communication is important during reproduction as part of a multimodal signaling repertoire, and that perception of auditory information changes depending on the animal's internal physiological state. Our results highlight the importance of examining non-visual sensory modalities as potential substrates for sexual selection contributing to the incredible phenotypic diversity of African cichlid fishes.     

Underwater vocalizations and associated behavior in captive ringed seals (<Emphasis Type="Italic">Pusa hispida</Emphasis>)

In pinniped species, especially those that mate in the water, acoustic communication is suggested to play an important role in various aspects of behavior. However, little is known about the behavioral context or function of vocalization, principally because direct observation is difficult in the wild. In the present study, we analyzed the seasonality, sexual differences, and behavioral contexts of the vocalizations of captive ringed seals to explore the function of such communication. The behavior of and underwater sounds made by three ringed seals (an adult male, an adult female, and a subadult female) living in Otaru Aquarium, Japan, were recorded for 19 days between August 2011 and April 2012. Six call types (long snort, knock, yelp, bark, click, and woof) were identified in the recordings. The 12 observed social behaviors could be categorized into three categories (male courtship, aggression, and submission). All call types except clicks were vocalized during social behavior. Vocalizations of all types increased during the breeding season. The long snorts were only produced by the adult male toward an adult female during his courtship behavior. All three individuals emitted knocks, yelps, and bark sounds. Of these three call types, knocks were associated with aggressive behavior or the male’s courtship behavior. In contrast, alternate series of yelps and barks were vocalized by the recipients of aggressive behaviors, suggesting their function as submissive signals. This study could be applied to the monitoring of wild ringed seals with passive acoustic recordings to assess not only their distribution but also their behavior.     

Male swordtails court with an audience in mind

Females are usually considered to be the target of male courtship behaviour. In nature, however, social interactions rarely occur without other observers; thus, it is conceivable that some male courtship behaviours are directed not towards females, but rather towards male rivals. The northern swordtail, Xiphophorus birchmanni, is a freshwater fish found in high densities in natural streams. Males court by swimming close to and in parallel with the female, raising their large sail-like dorsal fin, and quivering briefly. Here, we show that females prefer males that display small dorsal fins to those with large ones, and that males are less aggressive to other males with large dorsal fins. Male swordtails also raise their dorsal fins more frequently when courting in the presence of other males. These results suggest that, despite female avoidance of large dorsal fins, males that raise their fin during courtship benefit by intimidating potential competitors; the intended receivers of this signal are thus males, not females. Intrasexual selection can therefore offset the forces of intersexual selection, even in a courtship display.     

Temporal and Spatial Distribution of Male Scent Marks in the Polygynous Greater Sac-Winged Bat

Scent marks are relatively long-lived signals that can be perceived by conspecifics when the producer is absent. Therefore, it is often not obvious to whom the signal is directed. In daytime roosts of the polygynous greater sac-winged bat, males scent mark territories with facial gland secretions. Territories are a valuable resource for males, as they offer exclusive courtship opportunities, which results in increased male reproductive success and, consequently, increased male–male competition over territories. The information encoded in male scent marks could, therefore, be either directed at females as part of an olfactory courtship display or at male competitors as part of territorial behaviour. We expected territorial males to scent mark in the morning, shortly before females return to the territory and close to female roosting sites, if scent marks are directed at females as part of the courtship display. And we expected harem males to scent mark at the territory boundaries, where male–male encounters are most likely to occur, if scent marks are directed at male competitors. We found that males marked more frequently in the afternoon, at a time when all females have already left the territory, and harem males marked at the territory boundaries and not inside their territory in the area where females roost. At boundaries males fan volatiles from specialised wing sacs towards competitors outside the territory. Scent marking of male Saccopteryx bilineata might therefore be congruent with the assessment-hypothesis, which states that scent marks offer intruders the possibility to make an olfactory assessment of the territory owner without direct physical interaction. Thus, scent marks of male S. bilineata are most likely influenced by male–male competition and not by female choice.     

Evolution and function of multimodal courtship displays

Courtship displays are behaviours aimed to facilitate attraction and mating with the opposite sex and are very common across the animal kingdom. Most courtship displays are multimodal, meaning that they are composed of concomitant signals occurring in different sensory modalities. Although courtship often strongly influences reproductive success, the question of why and how males use multimodal courtship to increase their fitness has not yet received much attention. Very little is known about the role of different components of male courtship and their relative importance for females. Indeed, most of the work on courtship displays have focused on effects on female choice, often neglecting other possible roles. Additionally, a number of scientists have recently stressed the importance of considering the complexity of a display and the interactions between its different components in order to grasp all the information contained in those multimodal signals. Unfortunately, these methods have not yet been extensively adapted in courtship studies. The aim of this study was to review what is currently known about the functional significance of courtship displays, particularly about the role of multimodality in the courtship communication context. Emphasis is placed on those cases where a complete picture of the communication system can only be assessed by taking complexity and interaction between different modalities into account.     

Evolutionarily labile responses to a signal of aggressive intent

Males of many swordtail species possess vertical bar pigment patterns that are used both in courtship and agonistic interactions. Expression of the bars may function as a conventional threat signal during conflicts with rival males; bars intensify at the onset of aggression and fade in the subordinate male at contest's end. We used mirror image stimulation and bar manipulations to compare the aggressive responses of the males of four swordtail species to their barred and barless images. We found that having a response to the bars is tightly linked to having genes for bars, while the nature of the response the bars evoked varied across species. Specifically, we report the first known instance where closely related species exhibited differing and contradictory responses to a signal of aggressive motivation. Demonstrating that a signal conveys the same information across species (aggressive intent) while the response to that information has changed among species suggests that the nature of the responses are more evolutionarily labile than the signal.     

Turtles Are Not Just Walking Stones: Conspicuous Coloration and Sexual Selection in Freshwater Turtles

Turtles are among the most intriguing amniotes but their communication and signaling have rarely been studied. Traditionally, they have been seen as basically just silent armored ‘walking stones’ with complex physiology but no altruism, maternal care, or aesthetic perception. Recently, however, we have witnessed a radical change in the perception of turtle behavioral and cognitive skills. In our study, we start by reviewing some recent findings pertaining to various highly developed behavioral and cognitive patterns with special emphasis on turtles. Then we focus on freshwater turtles and use data about their sexual behavior and size sexual dimorphism (SSD) to test whether conspicuous coloration of the head is in these animals related to sexual processes. We found that absence of aggressive mating behavior is statistically associated with the presence of conspicuous coloration on turtles’ heads. It also seems that while species with female-biased SSD are characterised by conspicuously colored head ornaments, in species with male-biased SSD conspicuous coloration is absent. Unlike large females, males thus seem to be under pressure to develop conspicuous coloration and engage in non-aggressive behavior using signaling to succeed in courtship. And finally, we discuss possible roles of head color patterns in turtle communication during mating.     

Acoustic behavior of the damselfish Dascyllus albisella: behavioral and geographic variation

Behavioral and geographic variation in animal communication has been well-studied in insects, frogs, birds, and mammals, but little is known about variation in fishes. We used underwater audio-video recordings of the behavior and associated sounds produced by the domino damselfish, Dascyllus albisella, at Johnston Atoll and Hawaii, which are separated by 1000 km, to study behavioral and geographic variation in communication sounds. Males produced pulsed sounds during the courtship behavior known as the signal jump, visiting by females (during pseudospawning), mating, aggression to heterospecifics and conspecifics, and nest preparation. Females made only aggressive sounds. The following features of the sounds were measured: number of pulses, pulse rate, pulse duration, inter-pulse interval, dominant frequency, and frequency envelope. The only difference between visiting and mating sounds was a small difference in pulse duration. Two types of aggressive sounds were produced, pops and chirps. Pops contained only one or two pulses and were more commonly made towards heterospecifics than conspecifics. Aggressive chirps had between 3–11 pulses and were made most often towards conspecifics. The pulse rate of aggressive chirps was faster than signal jump sounds. The only difference in signal jump sounds made by males from Johnston Atoll and Hawaii, was a small difference in pulse duration, which was likely due to differences in the depths of the recording environment and not in the sounds produced.     

Influence of group rearing on sexual behavior of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis> males

Social interactions are able to strongly influence animal physiology and behavior. As is known, social experience can lead to changes in sexual and aggressive behavior, circadian rhythms and composition of cuticular hydrocarbons in Drosophila. Previously, we have shown that housing Drosophila males in monosexual groups of 20 individuals for 3 days after eclosion leads to a strong and long-term suppression of locomotor activity as revealed at individual testing, in contrast to males kept separately. The present research addressed courtship behavior, and specifically song production, in Drosophila males reared under similar conditions. It was found that rearing males in monosexual groups leads to a suppression of courtship and song production as well as to a simultaneous increase in locomotor activity when tested with a moving female. The latter effect was due to the strong urge of males to avoid interindividual contacts that prevented triggering the courtship ritual. It was suggested that intermale aggression caused by group rearing generates a state similar to conditioned fear.