Local adaptation and sexual selection: a reciprocal transfer experiment with the grasshopper Chorthippus biguttulus

Immigration into locally adapted populations has been suggested, among other potential causes, to maintain genetic variance in fitness necessary for good-genes models. Using a reciprocal transplant experiment we examined whether females prefer native to transferred males in the grasshopper Chorthippus biguttulus. On average, native and transferred males did not differ in their attractiveness, measured as female response rate to playbacks of male acoustic courtship signals. In line with this result, we found no significant effect of transfer on body size, condition, fluctuating asymmetry or song traits. However, the reciprocal transplant experiment showed that environmental conditions did influence body condition and maximum loudness of the calling song, but that the genetic origin of male grasshoppers had no significant effect on any of the analysed traits.Communicated by L. Simmons     

A dynamic trait affects continuous pair assessment in the blue-footed booby,<Emphasis Type="Italic"> Sula nebouxii</Emphasis>

Bright colours of male birds have often been shown to be the target of sexual selection through female choice, yet few studies have looked at the role of colour expressed after pairing on female motivation and behaviour. Here we analyse the role of an integumentary colour in the spectral range of 400–700 nm, the foot colour in male blue-footed boobies, Sula nebouxii, which is prominently displayed during pair courtship. Measurements early in the breeding season showed that foot colour of courting males is pale (high values of brightness) and has an aqua-blue chroma, and females in better body condition were mated to males with brighter feet. We carried out an experimental manipulation which modified the foot colour of males in courtship, making it closer to the foot colour of males in low nutritional state. We found that females paired to experimental males courted less and were less likely to copulate than females in the control group. Male behaviour was apparently unaffected by the manipulation; thus the change in female behaviour can be attributed exclusively to foot colour manipulation. These results strongly support the hypothesis of female preference for an integumentary colour and suggest that this dynamic trait is used as a male ornament after pairing.Communicated by J. Graves     

Dynamic female preference for multiple signals in <Emphasis Type="Italic">Rhinogobius brunneus</Emphasis>

Many species base their choice of mates on multiple signals which provide them with different kinds of information. Choosers may assess the signals together to evaluate the overall quality of potential mates, but individuals often pay attention to different signals in different contexts. In Rhinogobius brunneus, a fish displaying exclusive male parental care, females generally prefer males showing larger first dorsal fins (FDF) and more active courtship displays as mates. Females choosing a mate usually initially assess the FDF and later utilize courtship for the final decision. In our experiments, females with different hunger states used different signals when selecting mates. Females in both hunger states preferred males with larger FDF in the first stage. In the second stage, well-fed females showed highly repeatable choice, whereas poorly fed females responded only to variation in the courtship activity of males. The males preferred by poorly fed females exhibited significantly higher offspring survival than nonpreferred males. Under conditions of food shortage, males allocate more energy to future reproduction at the expense of the present brood, and females may prioritize signals predictive of offspring survivorship over signals reflecting other aspects in male quality to minimize the losses in direct benefits. We conclude that R. brunneus females may employ information from both signals but dynamically adjust their prioritization of each signal to current conditions to ensure the choice that is currently most adaptive.     

Courtship signalling with a labile bilateral signal: males show their best side

Asymmetries in courtship signals can result from both developmental instability during ontogeny and from temporary or permanent damage following mating, fighting, or interactions with predators. These two types of asymmetries, which can be divided into fluctuating asymmetry (FA) and damage asymmetry (DA), have both been suggested to play an important role in mate choice as potential honest indicators of phenotypic and/or genetic quality, while at the same time, DA may affect ornament asymmetry in a random manner. Interestingly, despite the massive research effort that has been devoted to the study of asymmetry during the past decades, very little is known about how an individual’s behaviour relates to asymmetry. Here, we measure and characterise asymmetry in morphological courtship signals in Corynopoma riisei, a fish where males carry elaborate paddle-like appendices on each side of the body that they display in front of females during courtship. Moreover, we investigate whether male courtship display, employing this bilateral morphological trait, reflects trait asymmetry. Finally, we assess whether males respond to phenotypic manipulations of DA with corresponding changes in courtship behaviour. We show that male display behaviour is asymmetric in a manner that reflects asymmetry of their morphological courtship trait and that male display behaviour responds to manipulations of asymmetry of these paddles. Our results thus suggest that males preferentially use their best side and, hence, that males respond adaptively to temporary changes in signal trait asymmetry.     

Alternative mating strategies in the water strider Gerris elongatus (Heteroptera,Gerridae)

Summary Males of the water strider Gerris elongatus established territories which included copulation and oviposition sites (small pieces of fallen bamboo). Males were aggressive and competition for territory and females was observed frequently. Male midlegs were more developed than female midlegs and were used as weapons. Reproductive behaviour changed as the breeding season advanced. Early in the season immature females were attracted by male surface wave courtship signals, then copulated white floating on the water surface without ovipositing (type 1). In midseason, males established territories, produced calling signals and attracted females which copulated and oviposited there with male postcopulatory guarding (type 2). In late season, many females oviposited without postcopulatory guarding on pondweed mats near fallen bamboo. Non-territorial males waiter for the arrival of these females and copulated without courtship, but mating success was low (type 3). These alternative mating strategies appeared to depend on differences in male size. Larger males were superior to smaller males in many ways (establishing territory, fighting, mating etc.). The largest males defended territories and had higher mating success than small non-territorial males. Medium sized males used all three strategies according to the number of empty territories and seasonal femald distribution.     

Vocalizations reveal body condition and are associated with visual display traits in great frigatebirds (Fregata minor)

Acoustic displays are known to advertise aspects of male quality and to affect female choice of mates in a variety of birds, including not only songbirds but some seabirds as well. Male great frigatebirds (Fregata minor) produce a rapid warble vocalization that forms a prominent part of their courtship display. We investigated the relationships between aspects of this vocalization and male quality and pairing success in 103 great frigatebirds from a population in the northwestern Hawaiian Islands. We found that across males, frequency bandwidth of the display was negatively related to repetition rate, enabling us to use deviation from the upper bound regression of bandwidth on rate (??vocal deviation??) as one measure of vocal performance. We used vocal consistency, defined as the coefficient of variation of the time interval between successive peaks in the warble, as a second measure of performance. Vocal deviation was significantly positively associated with body condition of male great frigatebirds, but vocal consistency was not. We found that peak frequency of the warble was not significantly associated with body size but was negatively associated with size of the gular pouch, the most prominent visual ornament. Male great frigatebirds showed significant individual differences in all three vocal measures??peak frequency, vocal deviation, and vocal consistency??but none of these measures was a significant predictor of pairing success in our study population. These results suggest that vocalizations provide honest information about male body condition and gular pouch size in great frigatebirds but do not influence male success in pairing with females.     

Social feedback and attractiveness in zebra finches

The attractiveness of an individual (i.e. its ability to arouse interest in a potential mate) has important implications for its reproductive success. However, attractiveness is not a fixed trait. Previous work has shown that male birds can adjust the intensity of their courtship display in response to variation in female behaviour, but little is known about how males adjust their behaviour during mate choice in response to social feedback about their own attractiveness independent of their intrinsic quality. Such information may help to maximize the potential mating success of males. Here, we provide experimental evidence that the amount of attention given by male zebra finches to females is dependent upon the manipulated attractiveness of males. This demonstrates that, in this socially monogamous species of bird, attractiveness of males could be considered to be a social construct, at least partially determined via social feedback from females.     

Advertisement-call modification,male competition,and female preference in the bird-voiced treefrog <Emphasis Type="Italic">Hyla avivoca</Emphasis>

Senders and receivers influence dynamic characteristics of the signals used for mate attraction over different time scales. On a moment-to-moment basis, interactions among senders competing for a mate influence dynamic characteristics, whereas the preferences of receivers of the opposite gender exert an influence over evolutionary time. We observed and recorded the calling patterns of the bird-voiced treefrog Hyla avivoca to assess how the dynamic characters of calls vary during interactions among groups of males in a chorus. This question was also addressed using playback experiments with males. Playback experiments with females showed how changes in dynamic call properties are likely to affect male mating success. Frogs calling in pairs, groups, or in response to playbacks produced longer calls than did isolated males. During call overlap, males often increased the duration of the silent interval (gaps) between the pulses of their calls so that the pulses of the calls of two neighbors interdigitated. This change resulted in increased variability of pulse rate, a traditionally static acoustic property; however, males also produced high proportions of non-overlapped calls in which variability in pulse rate was low and had species-typical values. Females preferred long calls to short- and average-duration calls, and non-overlapped calls to overlapped calls. Given a choice between pairs of overlapped calls, females preferred pairs in which the proportion of overlap was low and pairs in which the pulses of such calls interdigitated completely. The observed patterns of vocal competition thus reflect the preferences of conspecific females, which have influenced the evolution of the calling behavior of H. avivoca. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Correlates of male reproductive success in the browncheek blenny,Acanthemblemaria crockeri (Blennioidea: Chaenopsidae)

Summary Acanthemblemaria crockeri exhibits a resource defense polygyny mating system with male parental care. Females preferred to mate with the larger of two males in laboratory experiments, and male size was positively correlated with the number of eggs defended by males at two sites in the Gulf of California, Baja California, Mexico. Females appeared to avoid mating with males defending heavily-fouled shelters. The role of other factors including the intensity of male courtship coloration and displays in determining male reproductive success was studied in Bahia San Carlos, Sonora, Mexico by providing glass vials as shelters. This provided a non-destructive technique for counting of eggs defended by males and allowed repeated assessment of the reproductive success of individual males. Males varied greatly in their duration of residency of shelters and in their color score and intensity of courtship displays. The number of eggs received by males was positively correlated with their duration of residency in shelters, and females appeared to avoid mating with new residents. Mated males deserted shelters less frequently and were more likely to receive future matings than were unmated males. The mean color score of males was unrelated to their reproductive success, while the intensity of male courtship displays was negatively correlated with the number of eggs received. This may have resulted because female mate choice is based on multiple criteria, including some that more accurately reflect the quality of parental care afforded by males.     

Visual cues elicit courtship signals in a nocturnal anuran

In lekking species, the allocation of effort into mate attraction signals is not uniform over time, and signalers may expend the greatest effort when potential mates are nearby. Close-range courtship interactions are critical determinants of male fitness and the study of these interactions can therefore answer important questions in sexual selection. In anurans, attention has largely focused on long-range mate attraction mediated by acoustic signaling. However, many species also engage in courtship behaviors at close range, and the cues that elicit these behaviors are unknown but likely to be non-acoustic. I performed an experiment in which I assessed the role of female visual cues in eliciting courtship calls by males of the nocturnal treefrog Hyla versicolor. Males that could see an approaching female were more likely to give courtship calls than those that could not. These results provide some of the first evidence for an effect of vision on calling behavior in a nocturnal anuran and demonstrate that multiple sensory modalities are involved in the final stages of mate attraction.     

Female mate choice in the gray treefrog (Hyla versicolor) in three experimental environments

We studied female mate choice by Hyla versicolor in three venues to examine how acoustic and spatial complexity, background noise, and the calling behavior of males might influence preferences manifest in previous laboratory two-stimulus choice tests. Our laboratory-based two-stimulus choice tests with and without broadcasts of chorus noise demonstrated that females prefer long calls relative to short calls when calling efforts of alternatives are equivalent. Background noise impaired the ability of females to discriminate in favor of longer over shorter calls, but the magnitude of the effect was small. Captures of females at eight speakers broadcasting 6- to 27-pulse calls at the edge of a pond revealed strong discrimination against only the shortest call variant. In natural choruses, females may only rarely encounter males using such unattractive vocalizations. Female phonotaxis at an artificial pond with caged and electronically monitored calling males also indicated that consequences of female preferences for temporal aspects of calling observed in two-stimulus choice tests are much attenuated in choruses and explain only small portions (<10%) of the variation in male mating success. Nevertheless, relatively high call duration and calling effort increased male attractiveness. Acoustic interference emerged as another significant factor influencing male mating success and possibly the differences in female choice observed in laboratory and chorus settings. We suggest that the bias of females against both overlapped and very short calls may help explain why males lengthen their calls but lower their rate of delivery in response to increases in chorus size.     

Differences in acoustic directionality among vocalizations of the male red-winged blackbird (<Emphasis Type="Italic">Agelaius pheoniceus</Emphasis>) are related to function in communication

Studies of animal acoustic communication have found that the frequency and temporal structure of acoustic signals can be shaped by selection for efficient communication. The directionality of acoustic radiation may also be adapted for communication, but we know relatively little about how directionality varies with signal function, sender morphology, and the environment in which the sound is transmitted. We tested the hypothesis that the directionality of a vocalization is adapted to its function in communication. This hypothesis predicts that vocalizations that are directed to multiple conspecifics (e.g., advertisements and alarms) will be relatively omnidirectional because this will maximize the number of neighbors and mates that receive the signal, and that vocalizations directed to particular individuals will be relatively directional because this will maximize detection of the signal by the targeted receiver and minimize eavesdropping. To test these predictions, we measured the directionality and amplitude of red-winged blackbird (Agelaius pheoniceus) vocalizations in the field by recording vocalizations simultaneously on eight calibrated microphones encircling the bird. We found significant variation in directionality among vocalizations. Supporting our predictions, we found that the most omnidirectional vocalizations were those used to alert conspecifics to danger, and the most directional vocalizations are those used during courtship and solicitation of copulation, when the costs of eavesdropping are likely to be high. These results suggest that the directionality of red-winged blackbird vocalizations is shaped by selection for effective communication. This study is the first to provide statistical support for the hypothesis that directionality is related to the function of a signal in communication.     

Do microsporidian parasites affect courtship in two-spotted gobies?

Courtship and body condition of male two-spotted gobies (Gobiusculus flavescens) harbouring naturally acquired microsporidian infections were compared with those of microsporidian-free males under standardized conditions in the lab. Although parasite infection had no apparent effect on individual condition, it significantly affected male courtship intensity. This effect, however, was affected by the intensity of the female courtship. Our study is one of the first to demonstrate a sub-lethal behavioural effect of a microsporidian parasite that could negatively affect the reproductive success of infected individuals. Our results also suggest that secondary sexual traits like courtship may be more sensitive to moderately detrimental effects of parasite infection than classical condition indices.     

No evidence for female mate choice based on genetic similarity in the túngara frog Physalaemus pustulosus

In most sexually reproducing animals, the behavior of one or both sexes during courtship critically influences the success at mating of the opposite sex. This behavior is often interpreted as “mate choice,” and there is great interest in why such choices are exercised. The explanation for the evolution of mate choice that has received the most attention and generated the most controversy is based on assumed genetic effects. In this study, we investigated whether female túngara frogs, which choose mates based on acoustic cues, have a preference for genetically less related males. Specifically, we determine if there is disassortive mating based on microsatellite markers, if there is information in the advertisement call that could be used to assess genetic similarity, and if females exhibit acoustic-based mating preferences that would promote choice for genetic diversity. Using seven microsatellite markers, we found no correlation of male call similarity and male genetic relatedness. Female choice experiments showed no female preference for calls of less related males, and there was no evidence for inbreeding avoidance in the field. Our results do not support the hypothesis of mate choice based on information about genetic relatedness conveyed by acoustic signals in túngara frogs.     

Female mate selection in the natterjack toad: active choice or passive atraction?

Summary Variation in seasonal mating success among male natterjack toads (Bufo calamita) was influenced by the number of nights that males spent at the pond and by male body size. Large males produced louder and lower frequency calls than small males, and maintained larger acoustic territories. After arriving at the pond, one half of all observed females mated with the first male they encountered. The remainder visited several males before initiating amplexus, but no criteria could be identified that females might have used when deciding whether to accept or reject a male. Movements between several males seem to be best explained by low female responsiveness to male advertisement calls on cold nights which were nonoptimal for oviposition. Females attempted to reject non-calling males both before and after amplexus, but this may be a mechanism to avoid mismating with males of the common toad (Bufo bufo), an explosive breeder that utilised the same pond. In two-choice playback experiments using synthetic advertisement calls, females showed no preference for calls based on their frequency. Females preferred calls of intermediate pulse repetition rate equivalent to those produced by a male at the same body temperature. Pulse rate is thus potentially subject to stabilishing selection and may be an important character for species-recognition. Females preferred fast rather than slow call rates, but only when the alternative rates were extreme. They also preferred calls which they perceived at the highest sound pressure level, but did not discriminate between absolute sound pressure levels of alternative stimuli at different distances. Since females that delay mating and oviposition may suffer predation, it is suggested that female preference for loud, rapidly repeated calls may be adaptive in the sense of minimizing the costs of locating conspecific males, rather than maximizing the probability of obtaining a high quality mate. Competition between males to maintain large acoustic territories and produce calls that can be easily detected by females would seem to be a sufficient mechanism to explain the evolution of the striking calls produced by male natterjacks.     

Reproductive success of male bank voles (<Emphasis Type="Italic">Clethrionomys glareolus</Emphasis>): the effect of operational sex ratio and body size

The operational sex ratio (OSR) may influence the intensity of competition for mates and mate choice and is therefore thought to be a major factor predicting the intensity and direction of sexual selection. We studied the opportunity for sexual selection, i.e., the variance in male reproductive success and the direction and intensity of sexual selection on male body mass in bank vole (Clethrionomys glareolus) enclosure populations with experimentally manipulated sex ratios. The opportunity for sexual selection was high among male-biased OSRs and decreased towards female-biased OSRs. Paradoxically, selection for large male body mass was strongest in female-biased OSRs and also considerable at intermediate OSRs, whereas at male-biased OSRs, only a weak relationship between male size and reproductive success was found. Litters in male-biased OSRs were more likely to be sired by multiple males than litters in female-biased OSRs. Our results suggest that the intensity and direction of sexual selection in males differs among different OSRs. Although the direction of sexual selection on male body mass was opposite than predicted, large body mass can be favored by sexual selection. Naturally varying OSRs may therefore contribute to maintain variation in male sexually selected traits.     

Territorial counter-singing in male sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>): low-frequency songs trigger a stronger response

In many animal species, individuals compete for resources but avoid escalated conflicts by threat displays, i.e. a mutual signalling behaviour that enables the opponents to predict the outcome of the conflict without the necessity of actual fighting. For example, territory holders may use acoustic signals to communicate not only their own identity and the borders of their territory but also their competitive quality, fighting ability and motivation. Here, we show that male sac-winged bats, Saccopteryx bilineata, adjust their vocal territorial displays according to the fundamental frequency of territorial songs of their opponents. In playback experiments with territorial males, low-frequency stimuli elicited a higher territorial song rate and length than high-frequency stimuli. Male S. bilineata that sing more often and with lower fundamental frequencies have been shown to sire more offspring than their competitors. Fundamental frequency of territorial songs, hence, may reveal male quality and, consequently, the resulting threat posed to competing males. We argue that this is reflected in the increased response of competitors to low-frequency territorial songs shown here. Such competitive signalling behaviour has been shown in a few mammal species like red deer and baboons but, thus far, not in bats. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The interval between sexual encounters affects male courtship tactics in a desert-dwelling fish

Courtship displays are often important in determining male mating success but can also be costly. Thus, instead of courting females indiscriminately, males might be expected to adjust their signalling effort strategically. Theory, however, predicts that such adjustments should depend on the rate with which males encounter females, a prediction that has been subject to very little empirical testing. Here, we investigate the effects of female encounter rate on male courtship intensity by manipulating the time interval between sequential presentations of large (high quality) and small (low quality) females in a fish, the Australian desert goby Chlamydogobius eremius. Males that were presented with a small female immediately after a large female reduced their courtship intensity significantly. However, males courted large and small females with equal intensity if the interval between the sequential presentations was longer. Our results suggest that mate encounter rate is an important factor shaping male reproductive decisions and, consequently, the evolutionary potential of sexual selection.     

Postcopulatory mate guarding by vocalization in the Formosan squirrel

The Formosan squirrel, Callosciurus erythraeus thaiwanensis, emitted different vocalizations in response to terrestrial and aerial predators and snakes. Each vocalization caused nearby individuals to adopt a different type of anti-predator behaviour. In mating bouts, males produced two types of loud calls: precopulatory calls, emitted before copulations, and postcopulatory calls, emitted after copulations. The latter continued for 17 min on average. The estrous female and other males attending the mating bouts stopped moving during the postcopulatory call, so that the calling male was able to tend the female without interruption. The sound characteristics of anti-terrestrial-predator and postcopulatory calls recorded in the captivity were compared, and none of the ten characters of duration and frequency measured differed between the two calls. Playback experiments also showed that responses to the sounds in two different contexts, escape behaviour and defensive immobility, did not differ. The similarity between anti-predator and postcopulatory calls is discussed with reference to the possibility of manipulation and other explanatory hypotheses.     

Multimodal signaling in the Small Torrent Frog (Micrixalus saxicola) in a complex acoustic environment

Many animals use multimodal (both visual and acoustic) components in courtship signals. The acoustic communication of anuran amphibians can be masked by the presence of environmental background noise, and multimodal displays may enhance receiver detection in complex acoustic environments. In the present study, we measured sound pressure levels of concurrently calling males of the Small Torrent Frog (Micrixalus saxicola) and used acoustic playbacks and an inflatable balloon mimicking a vocal sac to investigate male responses to controlled unimodal (acoustic) and multimodal (acoustic and visual) dynamic stimuli in the frogs’ natural habitat. Our results suggest that abiotic noise of the stream does not constrain signal detection, but males are faced with acoustic interference and masking from conspecific chorus noise. Multimodal stimuli elicited greater response from males and triggered significantly more visual signal responses than unimodal stimuli. We suggest that the vocal sac acts as a visual cue and improves detection and discrimination of acoustic signals by making them more salient to receivers amidst complex biotic background noise.