Sound production during courtship and spawning of Oreochromis mossambicus: male–female and male–male interactions

The role of sound production of the Mozambique tilapia Oreochromis mossambicus in agonistic and mating interactions observed during hierarchy formation and in established groups was examined. Only territorial males produced sounds, during male–female and male–male courtship interactions and during pit-related activities ( e.g. dig, hover and still in the nest). Sound production rate was positively correlated with courting rate. Although sounds in other cichlids are typically emitted in early stages of courtship, O. mossambicus produced sounds in all phases, but especially during late stages of courtship, including spawning. It is suggested that the acoustic emissions in this species may play a role in advertising the presence and spawning readiness of males and in synchronizing gamete release.     

Sounds Modulate Males’ Aggressiveness in a Cichlid Fish

Acoustic signals are produced in many fish species during agonistic or courtship interactions. A way to test the biological role of these sounds is the use of acoustic playback experiments. However, sounds are usually associated with visual displays and playback experiments performed in fish so far, often failed to match acoustic and visual stimuli. To avoid this mismatch issue, we experimentally separated or coupled visual and acoustic channels to test the role of sounds produced during male–male aggressive interactions in a cichlid fish, Metriaclima zebra. Results show that aggressive behaviour is based on visual stimuli and that acoustic signals alone never trigger aggression. Furthermore, the association between visual and acoustic channels lowers the level of aggressiveness found when fish can only interact visually. This suggests that acoustic signals used during a dispute may complement visual displays to modulate males’ behaviour by reducing their aggressiveness and the risk of escalated fights.     

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.     

Absence of social facilitation of courtship in the wolf spider, Schizocosa ocreata (Hentz) (Araneae: Lycosidae)

Males of many animal species are reproductively limited by the difficulty and time costs of finding mates. Males of such species should be selected to take advantage of any cues that might reveal the location of prospective mates. Cues to female location are not restricted to those produced by females, but might also include the highly apparent courtship displays of males that have already found a female. By “eavesdropping” on these courting rivals, initiating sexual displays when courting rivals are detected (i.e., social facilitation of displays); males might effectively exploit the mate-searching efforts of their rivals. We tested the possibility that male Schizocosa ocreata wolf spiders exhibit social facilitation of courtship behaviors using a combination of live behavioral trials and video playback with single stimulus presentations. When exposed to visual cues from another male, male S. ocreata can discern the presence of another individual whether that individual is courting or not. However, we found no evidence of social facilitation of courtship or chemoexploratory behaviors in response to seismic or visual cues presented in isolation or combined. While complex, multimodal, male courtship signals are important in mate choice by female S. ocreata, males do not appear to use these cues to socially facilitate their own courtship.     

Digital video playback and visual communication in lizards

Experimental analyses of dynamic visual signals have to overcome the technical obstacle of reproducing complex motor patterns such as those found in courtship and threat displays. Video playback offers a potential solution to this problem, but it has recently been criticized because of sensory differences between humans and nonhuman animals, which suggest that video stimuli might be perceived as deficient relative to live conspecifics. Quantitative comparisons are therefore necessary to determine whether video sequences reliably evoke natural responses. Male Jacky dragons, Amphibolurus muricatus, compete for territories using complex displays delivered in a rapid stereotyped sequence. We evaluated video playback as a technique for studying this visual signal. Digital video sequences depicting a life-sized displaying male were indistinguishable from live male conspecifics in the rate and structure of aggressive displays evoked. Other measures of social behaviour suggested that video stimuli were more effective in this context. Lizards produced significantly more appeasement displays and had higher rates of substrate licking and locomotor activity in response to video playback than to confined male opponents, which failed to produce aggressive displays. Lizards tracked temporal changes in the display rate of video stimuli and were also sensitive to individual differences in morphology and behaviour between video exemplars. These results show that video stimuli are appropriate for the experimental analysis of Jacky dragon aggressive displays. We compare the potential shortcomings of video playback with those of other techniques and conclude that no approach offers a panacea, but that several have complementary characteristics. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The Role of Agonistic Sounds in Male Nest Defence in the Painted Goby Pomatoschistus pictus

Animals often vocalize during territorial challenges as acoustic signals may indicate motivation and fighting ability and contribute to reduce aggressive escalation. Here, we tested the function of agonistic sounds in territorial defence in the painted goby. Pomatoschistus pictus, a small vocal marine fish that defends nests during the breeding season. We first measured the number of times a male approached, avoided, explored, entered and exited two unattended nests associated with either conspecific agonistic sounds or a control: silence or white noise. Acoustic stimuli were played back when the male approached a nest. In a second experimental set, we added visual stimuli, consisting of a conspecific male in a small confinement aquarium near each nest. Even though we found no effect of the visual stimuli, the sound playbacks induced similar effects in both experimental conditions. In the sound vs. silence treatment, we found that when males approached a nest, the playback of conspecific sounds usually triggered avoidance. However, this behaviour did not last as in longer periods males visited nests associated with agonistic sounds more often than silent ones. When the control was white noise, we found no significant effect of the playback treatment in male behaviour. Although we cannot exclude the possibility that other sounds may dissuade nest occupation, our results suggest that agonistic sounds act as territorial intrusion deterrents but are insufficient to prevent nest intrusion on their own. Further studies are needed to test the significance of sound production rate, spectral content and temporal patterns to deter territorial intrusion in fish.     

Sound production during agonistic behavior of male Drosophila melanogaster

Male Drosophila fruit flies acquire and defend territories in order to attract females for reproduction. Both, male-directed agonistic behavior and female-directed courtship consist of series of recurrent stereotypical components. Various studies demonstrated the importance of species-specific sound patterns generated by wing vibration as being critical for male courtship success. In this study we analyzed the patterns and importance of sound signals generated during agonistic interactions of male Drosophila melanogaster. In contrast to acoustic courtship signals that consist of sine and pulse patterns and are generated by one extended wing, agonistic signals lack sine-like components and are generally produced by simultaneous movements of both wings. Though intra-pulse oscillation frequencies (carrier frequency) are identical, inter-pulse intervals are twice as long and more variable in aggression signals than in courtship songs, where their precise temporal pattern serves species recognition. Acoustic signals accompany male agonistic interactions over their entire course but occur particularly often after tapping behavior which is a major way to identify the gender of the interaction partner. Since similar wing movements may either be silent or generate sound and wing movements with sound have a greater impact on the subsequent behavior of a receiver, sound producing wing movements seem to be generated intentionally to serve as a specific signal during fruit fly agonistic encounters.     

The significance of sound interception to males of the bicolor damselfish,Pomacentrus partitus,during courtship

Synopsis Playback experiments conducted in the field and the laboratory demonstrated the use of sound interception in bicolor damselfish,Pomacentrus partitus. Two courtship sounds produced by male bicolors: the ‘chirp’ which occurs at the initiation of courting and the ‘grunt’ which occurs near the termination, just prior to spawning, were found to hold different meanings to intercepting male competitors. Males responded to grunt playback with directional swimming towards the sound source and increased courtship behavior. No directional response was observed during chirp playback. The grunt sound appears to indicate that a spawn-ready female is present near the sound source. Males therefore move towards it, likely to interfere with the imminent spawning or to gain a possible spawning partner. Such is not observed in response to chirps, as little advantage would be gained by moving to the source of a sound which is known to serve as an ‘advertisement’ or ‘territorial keep-out’ signal. Thus, upon intercepting a neighbor's chirps, males exhibit courting within their own territory, or directly court a nearby female.     

Behaviours Associated with Acoustic Communication in Nile Tilapia (Oreochromis niloticus)

Background

Sound production is widespread among fishes and accompanies many social interactions. The literature reports twenty-nine cichlid species known to produce sounds during aggressive and courtship displays, but the precise range in behavioural contexts is unclear. This study aims to describe the various Oreochromis niloticus behaviours that are associated with sound production in order to delimit the role of sound during different activities, including agonistic behaviours, pit activities, and reproduction and parental care by males and females of the species.

Methodology/Principal Findings

Sounds mostly occur during the day. The sounds recorded during this study accompany previously known behaviours, and no particular behaviour is systematically associated with sound production. Males and females make sounds during territorial defence but not during courtship and mating. Sounds support visual behaviours but are not used alone. During agonistic interactions, a calling Oreochromis niloticus does not bite after producing sounds, and more sounds are produced in defence of territory than for dominating individuals. Females produce sounds to defend eggs but not larvae.

Conclusion/Significance

Sounds are produced to reinforce visual behaviours. Moreover, comparisons with O. mossambicus indicate two sister species can differ in their use of sound, their acoustic characteristics, and the function of sound production. These findings support the role of sounds in differentiating species and promoting speciation. They also make clear that the association of sounds with specific life-cycle roles cannot be generalized to the entire taxa.     

An evaluation of video playback using Xiphophorus helleri

Video playback is being increasingly used as a technique for behavioural research. The importance of critically evaluating the effectiveness of video playback is clear, as available video technology is not designed for nonhuman visual systems. We discuss several aspects concerning the perception of video images that could lead to inconclusive or erroneous results. Researchers should verify that behaviour observed in response to video playback is comparable to behaviour observed in response to live animals. We conducted such a verification using live and video playback methods to measure female response to swords of varying lengths in the green swordtail, Xiphophorus helleri. Using both methods, female response appeared to be an increasing function of male sword length. Females did not differ in their response to live and video versions of noncourting, noninteractive males, however, females tended to prefer video playbacks of males with longer swords, a result that has also been found in experiments using live males. These results suggest that females express the same qualitative mating preference, but not necessarily the same quantitative preference, for sword length when viewing video stimuli. Several methodological factors that may contribute to an apparent difference in the strength of the preference are discussed. Despite these differences, both methods produced comparable results; female response to sworded males tended to increase as sword length increased. These experiments demonstrate that video playback is an effective method to measure female preferences accurately in X. helleri and provide an example of how video playback can be evaluated in other species. Copyright 2000 The Association for the Study of Animal Behaviour.     

Risky Courtship: Background Contrast,Ornamentation, and Display Behavior of Wolf Spiders Affect Visual Detection by Toad Predators

Males that search widely for females and perform conspicuous courtship displays run a high risk of being detected by their predators. Therefore, gains in reproductive success might be offset by increased mortality due to predation. Male brush‐legged wolf spiders (Schizocosa ocreata) with larger decorative traits (foreleg tufts) are preferred by females as mates, but are more readily detected by predators. However, predation risk may also be influenced by the interaction between components of signals and the environment in which signaling occurs. Courting male spiders were readily accepted as prey by a sympatric predator, the American toad (Anaxyrus americanus). We used video playback to tease apart the interactive effect between visual signals and the signaling environment on the ability of toads to detect courting spiders as a function of distance, background contrast, the presence or absence of male foreleg tufts, and behavioral activity. The response of toads to video sequences of male spiders was similar to their response to live male spiders. Toad response varied over distance toward spiders displayed against high contrast (sunny) vs. low contrast (shaded) backgrounds. Beyond 30 cm, more toads detected courting male spiders against light, ‘sunny’ backgrounds and detected them faster when compared to the same spider stimulus against darker, ‘shady’ backgrounds. In choice tests, toads oriented more often toward courting males with leg tufts than those without. Toad responses also varied with male spider behavior in that only videos of moving males were attacked. Latency to orient and detection by toads was significantly greater for walking males than courting males, and this effect was most evident at distances between 30 cm and 50 cm. Results supported that courting wolf spiders are at significant risk of predation by visually acute predators. Distance, background contrast, and the presence of foreleg decorations influence detection probability. Thus, the same complex visual signals that make males conspicuous and are preferred by females can make males more vulnerable as prey to toads.     

Court Like You Mean It: Male Siamese Fighting Fish are Less Attentive to Courting Males that Have Been Exposed to an Estrogen Mimic

Endocrine disrupting chemicals (EDCs) are ubiquitous in aquatic ecosystems where they have adverse effects on exposed organisms. In addition to causing physiological changes, EDCs often target fitness‐related behaviors such as locomotion and courtship. Ethinylestradiol (EE2) is an estrogen mimic that has been found to reduce courtship and aggression in males. However, the consequences of these reductions are not always explicitly addressed. One way in which EE2 may lead to decreased fitness occurs when males respond differently to exposed vs. unexposed conspecifics. To examine this, video playback was used to determine whether male Siamese fighting fish, Betta splendens, respond differently to exposed and unexposed males. Males were presented with four different combinations of videos of males played simultaneously: exposed swimming and unexposed swimming, exposed courting and unexposed courting, unexposed courting and unexposed courting, and exposed courting and exposed courting. Males directed more behaviors to the unexposed than the exposed courting male when presented simultaneously, likely because these unexposed males were perceived as a greater threat to mating success. Additionally, males spent more time tracking and gill flaring, two behaviors that are indicative of fight intent, when presented with two courting unexposed males. This combination could be considered the most threatening because there are two rival vigorously courting males present. These results suggest that EE2 exposure could result in exposed males receiving decreased attention from other males. However, since EE2 exposure also decreases competitive and courtship abilities, this would be far outweighed by the costs.     

Importance of pause between spider courtship vibrations and general problems using synthetic stimuli in behavioural studies

Effectiveness of natural stimuli (recorded and replayed male vibratory courtship signals) in eliciting a female's vibratory response was compared with that of synthetic stimuli in a plant-dwelling spider (Cupiennius salei). Specifically, the role of interseries (pauses between consecutive male series) was evaluated. Effectiveness was assessed by the number of responding females and the number of responses per female per stimulus. In case of both synthetic and natural stimuli, effectiveness increases with increasing duration of interseries in both measures. Interseries are thus represented in the female's innate releasing mechanism. However, synthetic stimuli are less effective (about 55%) than natural stimuli (set 100%). With decreasing interseries, the number of female responses per unit time sharply increases in case of playback of natural stimuli, but only slightly in case of synthetic stimuli. Since males courting on plants decrease interseries as soon as a female responds, thus facilitating their orientation, the data obtained with natural stimuli much better fit natural male courtship behaviour. This finding brings into question the validity of the synthetic stimulus. In the bioacoustics literature there is neither consensus on the effectiveness of a synthetic stimulus necessary to qualify it for behavioural analysis nor on how to measure effectiveness. We propose three measures : the percentage of responding animals, the action pattern of the response and the number of responses. It may be crucial, particularly when addressing evolutionary questions, to use either natural stimuli or synthetic stimuli (nearly) as effective as natural stimuli to avoid biased sampling of experimental animals and results difficult to interpret in biological terms.     

The Elaborate Postural Display of Courting <Emphasis Type="Italic">Drosophila persimilis</Emphasis> Flies Produces Substrate-Borne Vibratory Signals

Sexual selection has led to the evolution of extraordinary and elaborate male courtship behaviors across taxa, including mammals and birds, as well as some species of flies. Drosophila persimilis flies perform complex courtship behaviors found in most Drosophila species, which consist of visual, air-borne, gustatory and olfactory cues. In addition, Drosophila persimilis courting males also perform an elaborate postural display that is not found in most other Drosophila species. This postural display includes an upwards contortion of their abdomen, specialized movements of the head and forelegs, raising both wings into a “wing-posture” and, most remarkably, the males proffer the female a regurgitated droplet. Here, we use high-resolution imaging, laser vibrometry and air-borne acoustic recordings to analyse this postural display to ask which signals may promote copulation. Surprisingly, we find that no air-borne signals are generated during the display. We show, however, that the abdomen tremulates to generate substrate-borne vibratory signals, which correlate with the female’s immobility before she feeds onto the droplet and accepts copulation.     

Contribution to the study of acoustic communication in two Belgian river bullheads (<Emphasis Type="Italic">Cottus rhenanus and C. perifretum</Emphasis>) with further insight into the sound-producing mechanism

Background

The freshwater sculpins (genus Cottus) are small, bottom-living fishes widely distributed in North America and Europe. The taxonomy of European species has remained unresolved for a long time due to the overlap of morphological characters. Sound production has already been documented in some cottid representatives, with sounds being involved in courtship and agonistic interactions. Although the movements associated with sound production have been observed, the underlying mechanism remains incomplete. Here, we focus on two closely related species from Belgium: C. rhenanus and C. perifretum. This study aims 1) to record and to compare acoustic communication in both species, 2) to give further insight into the sound-producing mechanism and 3) to look for new morphological traits allowing species differentiation.

Results

Both Cottus species produce multiple-pulsed agonistic sounds using a similar acoustic pattern: the first interpulse duration is always longer, making the first pulse unit distinct from the others. Recording sound production and hearing abilities showed a clear relationship between the sound spectra and auditory thresholds in both species: the peak frequencies of calls are around 150 Hz, which corresponds to their best hearing sensitivity. However, it appears that these fishes could not hear acoustic signals produced by conspecifics in their noisy habitat considering their hearing threshold expressed as sound pressure (~ 125 dB re 1 μPa). High-speed video recordings highlighted that each sound is produced during a complete back and forth movement of the pectoral girdle.

Conclusions

Both Cottus species use an acoustic pattern that remained conserved during species diversification. Surprisingly, calls do not seem to have a communicative function. On the other hand, fish could detect substrate vibrations resulting from movements carried out during sound production. Similarities in temporal and spectral characteristics also suggest that both species share a common sound-producing mechanism, likely based on pectoral girdle vibrations. From a morphological point of view, only the shape of the spinelike scales covering the body allows species differentiation.

     

Multiple male traits interact: attractive bower decorations facilitate attractive behavioural displays in satin bowerbirds

Sexually selected male courtship displays often involve multiple behavioural and physical traits, but little is known about the function of different traits in mate choice. Here, we examine female courtship behaviours to learn how male traits interact to influence female mating decisions. In satin bowerbirds (Ptilonorhynchus violaceus), successful males give highly aggressive, intense behavioural displays without startling females. Males do this by modulating their displays in response to female crouching, which signals the display intensity that females will tolerate without being startled. Females typically visit multiple males for multiple courtships before choosing a mate, and females show differing tolerance for intense displays during their first courtship with each male. We test three hypotheses that may explain this: (i) familiarity with the courting male; (ii) the order of the courtship in mate-searching; and (iii) the attractiveness of the courting male. We found that females are more tolerant of intense displays during first courtships with attractive males; this increased female tolerance may allow attractive males to give higher intensity courtship displays that further enhance their attractiveness. We then examined why this is so, finding evidence that females are less likely to be startled by males with better physical displays (bower decorations), and this reduced startling then contributes to male courtship success. This role of physical displays in facilitating behavioural displays suggests a novel mechanism by which multiple physical and behavioural traits may influence female choice.     

Studying Visual Cues in Fish Behavior: A Review of Ethological Techniques

This paper reviews the variety of approaches available to fish ethologists to study the role of visual cues in fish behavior. Examples of studies that have used live fish, mirror images, dummies (i.e. models), or video playback as stimuli to investigate fish behavior are described and discussed. These examples represent a diversity of functional categories of behavior exhibited by fishes, including aggression, courtship, aggregation, or schooling behavior, parent–offspring, predator–prey, and cleaner–host interactions. The specific techniques that fish biologists have used to control or manipulate body shape, size, posture, morphological structures, color, marking patterns, or movement are systematically discussed, and the importance of each of these visual features to fish behavior is documented through examples. Studies that have used these techniques to investigate the interaction between visual and nonvisual cues are also considered. Each section encompassing a general experimental approach ends with a discussion of the advantages and disadvantages of that approach for studying fish behavior.     

Singing above the chorus: cooperative Princess cichlid fish (<Emphasis Type="Italic">Neolamprologus pulcher</Emphasis>) has high pitch

Teleost fishes not only communicate with well-known visual cues, but also olfactory and acoustic signals. Communicating with sound has advantages, as acoustic signals propagate fast, omnidirectionally, around obstacles and over long distances. Heterogeneous environments might favour multimodal communication, especially in socially complex species, as the combination of modalities’ strengths helps overcome their individual limitations. Fishes of the ecologically and morphologically diverse family Cichlidae are known to be vocal. Here we investigated sound production in the socially complex Princess cichlid Neolamprologus pulcher from Lake Tanganyika in East Africa. We show that wild and captive N. pulcher produce only short-duration, broadband high-frequency sounds (mean: 12 kHz), when stimulated by mirror images. The evolutionary reasons for this “low frequency silencing” are still unclear. In laboratory experiments, N. pulcher produced distinct two-pulsed calls mostly, but not exclusively, associated with agonistic displays. Princess cichlids produce these high-frequency sounds both in combination with and independent from visual displays, suggesting that sounds are not a by-product of behavioural displays. Further studies on the hearing abilities of N. pulcher are needed to clarify whether the high-frequency sounds are used in intra- or inter-specific communication.     

A review of acoustic playback techniques for studying avian vocal duets

ABSTRACT Playback experiments involve the broadcast of natural or synthetic sound stimuli and provide a powerful tool for studying acoustic communication in birds. Playback is a valuable technique for exploring vocal duetting behavior because it allows investigators to test predictions of the various hypotheses for duet function. Here, we adopt a methodological perspective by considering various challenges specific to studying duetting behavior, and highlighting the utility of different playback designs for testing duet function. Single‐speaker playback experiments allow investigators to determine how duetting birds react to different stimuli, but do not simulate duets in a spatially realistic manner. Multi‐speaker playback experiments are superior to single‐speaker designs because duet stimuli are broadcast with spatial realism and unique and additional predictions can be generated for testing duet function. In particular, multi‐speaker playback allows investigators to evaluate how birds respond to male versus female duet contributions separately, based on reactions to the different loudspeakers. Interactive playback allows investigators to ask questions about the time‐ and pattern‐specific singing behavior of birds, and to understand how singing strategies correspond to physical behavior during vocal interactions. Although logistically challenging, interactive playback provides a powerful tool for examining specific elements of duets (such as the degree of coordination) and may permit greater insight into their functions from an operational perspective. Interactive playback designs where the investigator simulates half of a duet may be used to describe and investigate the function of pair‐specific and population‐wide duet codes. Regardless of experimental design, all playback experiments should be based on a sound understanding of the natural duetting behavior of the species of interest, and should aim to produce realistic and carefully controlled duet simulations. Future studies that couple playback techniques with other experimental procedures, such as Acoustic Location System recordings for monitoring the position of birds in dense vegetation or multimodal techniques that combine acoustic with visual stimuli, are expected to provide an even better understanding of these highly complex vocal displays.     

Sound production in <Emphasis Type="Italic">Etheostoma oophylax</Emphasis> (Percidae) and call characteristics correlated to body size

Sound production in fishes is common in marine and freshwater species, however there are still many vocal species for which sound production has not been documented. This paper is the first account of sound production in the Guardian Darter (Etheostoma oophylax). Laboratory recordings revealed that males produced several vocalizations, including single pulse knocks, multi-pulsed purrs, and tonal drums. All vocalizations were documented during agonistic and courtship encounters, including spawning. We also investigated possible correlations between call characteristics and male size. Male standard length was found to be correlated to inter-pulse interval of purrs, as well as the slope of the drum vocalizations. Determining a link between male size and acoustic characteristics could be the first step in documenting mate or male-male assessment by acoustic communication in darters.