Diel variation in a dynamic sexual display and its association with female mate-searching behaviour

Dynamic sexual signals often show a diel rhythm and may vary substantially with time of day. Diel and short-term fluctuations in such sexual signals pose a puzzle for condition capture models of mate choice, which assume a female preference for male traits that reliably reflect a male's quality. Here we experimentally manipulated the food supply of individual male field crickets Gryllus campestris in their natural habitat in two consecutive seasons to determine (i) the effect of male nutritional condition on the fine-scaled variation of diel investment in acoustic signalling and (ii) the temporal association between the diel variation in male signalling and female mate-searching behaviour. Overall food-supplemented males signalled more often, but the effect was only visible during the daytime. In the evening and the night, signal output was still high but the time spent signalling was unrelated to a male's nutritional condition. Females' mate-searching behaviour also showed a diel rhythm with peak activity during the afternoon, when differences among calling males were highest, and where signal output reliably reflects male quality. These findings suggest that males differing in nutritional condition may optimize their investment in signalling in relation to time of day as to maximize mating success.     

Male responses to conspecific advertisement signals in the field cricket Gryllus rubens (Orthoptera: Gryllidae)

In many species males aggregate and produce long-range advertisement signals to attract conspecific females. The majority of the receivers of these signals are probably other males most of the time, and male responses to competitors' signals can structure the spatial and temporal organization of the breeding aggregation and affect male mating tactics. I quantified male responses to a conspecific advertisement stimulus repeatedly over three age classes in Gryllus rubens (Orthoptera: Gryllidae) in order to estimate the type and frequency of male responses to the broadcast stimulus and to determine the factors affecting them. Factors tested included body size, wing dimorphism, age, and intensity of the broadcast stimulus. Overall, males employed acoustic response more often than positive phonotactic response. As males aged, the frequency of positive phonotactic response decreased but that of the acoustic response increased. That is, males may use positive phonotaxis in the early stages of their adult lives, possibly to find suitable calling sites or parasitize calling males, and then later in life switch to acoustic responses in response to conspecific advertisement signals. Males with smaller body size more frequently exhibited acoustic responses. This study suggests that individual variation, more than any factors measured, is critical for age-dependent male responses to conspecific advertisement signals.     

Mating behaviour and male mating success in the green treefrog

Green treefrogs, Hyla cinerea, at a study site in eastern Georgia behaved as prolonged breeders: there were many more males present each night than females, the arrival of which was unpredictable. Individual males were usually consistent in their behaviour during a single night, and were most often observed calling. There was a weak, negative correlation between male size (snout-vent length, unless otherwise stated) and the frequency of satellite behaviour, defined as a non-calling male's being situated near a calling male. In 2 of 3 years, the mean size of males observed as satellites was less than that of the calling males with which they were associated on a given night; however, many satellites were the same size or larger than calling males. The distributions of matings per male during three seasons were not significantly different than Poisson distributions with the same means. Individual mating success was best predicted by attendance at the breeding site; mating success was negatively related to the frequency of satellite behaviour in some analyses, but very little of the variance was explained. An exhaustive series of analyses failed to show any pattern of size-dependent mating, including weight-assortative mating, even when males that were frequently satellites were excluded. The relative sizes of males and females in amplexus had little effect on fertilization success. On a seasonal basis, males were clumped in their spatial distribution, and matings tended to occur where male density was highest. Individual males were highly variable in their patterns of movement, which were not correlated with size or mating success. Mating success was also unrelated to the frequency (Hz) of the low-frequency spectral peak in the advertisement call of the male; male size and weight were negatively correlated with the frequency of the low-frequency peak.     

Diel Calling Activity and Field Survival of the Bushcricket,Sciarasaga quadrata (Orthoptera: Tettigoniidae): A Role for Sound-locating Parasitic Flies?

In this study I examine the variability in calling activity of Sciarasaga quadrata over both the short (4 d) and long term (life span), and the temporal pattern to host searching by its most significant natural enemy, the ormiine fly Homotrixa alleni. Indeed, few male S. quadrata survive the entire calling season as this acoustically orienting parasitoid fly continually ‘culls’ calling males out of the population. I found that male S. quadrata commence calling over 1 h before sunset, cease around midnight and call, on average, for over 3 h within this period (range 0–11 h 09 min). There was significant concordance and repeatability in calling activity over both the short and long term, enabling me to conclude that relative differences in calling activity among males persist throughout a male's life span. There was a distinct peak to host searching by gravid female flies, with 82 % of all flies collected at acoustic traps between sunset and midnight. A positive association between call duration (time spent calling) and fly attraction was evident from multiple collections of flies at single traps. Counter to expectation, there was no significant decline in average call duration across successive collections of males, indicating that long-call-duration males were not being selectively culled by flies from the calling population. Males at the end of the calling season, when their remaining reproductive potential is low, allocate the majority of their calling time between sunset and midnight despite the high risk of fly attack. The distribution of call durations within the population was not skewed, which if heritable, indicates a lack of history of directional selection acting on call duration. Limited evidence suggests that encounters with females are rare in S. quadrata so that males may be selected, irrespective of natural selection pressure, to call for long periods of time to optimize their chance of attracting mates. The above findings are further discussed in relation to parasitoid foraging and acoustic reproductive strategies.     

Is the later arrival of young male red-breasted flycatchers (Ficedula parva) related to their physical condition?

Intraspecific variation in the arrival time of migratory birds to breeding grounds is common. Although this phenomenon has been explained in various ways, the condition-dependency of arrival is often evoked. I analyzed the arrival time of male red-breasted flycatchers Ficedula parva, a long-distance migratory passerine, in relation to age, body size and body condition. Data were collected over the course of 5 years in the primeval Białowieża Forest of northeast Poland. Each of these years, young males arrived at the breeding grounds significantly later in the year than did older males. Young males also exhibited significantly shorter wings and lower body condition than older males. Settlement speed was significantly related to age, but not to wing length or body condition. Only the arrival time of old males was related to the body mass and condition. Later arrival of young males could be explained by the lack of experience or by avoidance of both aggression and competition from older males, and both explanations are thought to conserve energy for breeding. Young male red-breasted flycatchers exhibit delayed plumage maturation, and this duller plumage supports the strategy of restraint in the arrival time of young males in this species.     

Nightly calling patterns in a Neotropical gladiator frog

In anurans, vocalization is the primary communication form and acoustic parameters are influenced by climatic conditions, but also by social contexts. We investigated calling site use and within-individual variation of acoustic parameters throughout the night in the gladiator-frog Hypsiboas goianus. We expected that large males would call closer to the water and at higher perches to avoid dehydration and maximize sound propagation. Furthermore, we tested the prediction that males would emit more aggressive calls early in the night and more advertisement calls late at night. Male size was not correlated with either distance from the water or perch height. However, as expected, males of H. goianus adopted a calling strategy that consisted of emitting more aggressive calls early in the night and more advertisement calls later in the night. Furthermore, repetition rate and interval between notes of the aggressive calls and repetition rate of the advertisement calls showed within-individual variation throughout the night that agreed with our expectations. The calling strategy of H. goianus is probably related to the establishment of calling sites early in the night and an investment in female attraction in the middle of the night when males’ aggressive interactions have faded away, or due to habituation increasing the males’ aggressive thresholds. This is the first study investigating within-individual patterns of acoustic parameters of calls throughout the night for anurans.     

Artificial selection on male longevity influences age-dependent reproductive effort in the black field cricket Teleogryllus commodus

Although the trade-off between reproductive effort and longevity is central to both sexual selection and evolutionary theories of aging, there has been little synthesis between these fields. Here, we selected directly on adult longevity of male field crickets Teleogryllus commodus and measured the correlated responses of age-dependent male reproductive effort, female lifetime fecundity, and several other life-history traits. Male longevity responded significantly to five generations of divergent selection. Males from downward-selected lines commenced calling sooner and reached their peak calling effort at a younger age. They called more per night and, despite living less than half as long, called more overall than males selected for increased longevity. Females from the downward-selected lines lived significantly shorter lives than females from the upward-selected lines but still produced the same number of offspring. Nymph survival, development time, and body size and weight at eclosion did not show significant correlated response to selection on male longevity, despite evidence for substantial genetic variation in each of these traits. Collectively, our findings directly support the antagonistic pleiotropy model of aging and suggest an important role for sexual selection in the aging process.     

Social Factors Influence the Acoustic Variability in the Long-distance Calls of Male Chimpanzees

Acoustic variability in the species-typical long-distance calls of male chimpanzees was investigated. Analysis of 13 acoustic features revealed that calls varied more within than between individuals. Although only a limited amount of acoustic variability was attributable to different populations, discriminant-function analysis confirmed that males from two populations produce acoustically distinguishable calls. Additional investigation suggested that social factors may contribute to the patterns of acoustic variation both between and within individuals. Matrix-permutation tests revealed a positive association between the amount of time males spent together and a measure of call similarity. Vocal similarities between individuals appeared to result from a dynamic process involving chorusing behaviour: males matched the acoustic characteristics of each other's vocalizations when calling together. Chorusing may also affect the degree of within-individual acoustic variation in calls. Males who chorused often with others produced more variable calls than individuals who chorused less often or called alone.     

Temporally fluctuating selection of sex-limited signaling traits in the Texas field cricket,Gryllus texensis

Abstract.— Gryllus texensis males produce acoustic mating signals and display extensive heritable variation in when and how much time they spend signaling throughout the night. The goal of this research is to elucidate the potential mechanism responsible for maintaining this heritable variation. Mating signals attract female crickets. In low-density spring populations females select males that signal most often; in high-density fall populations mating appears random with respect to signaling time. Mating signals also inadvertently attract acoustically orienting parasitoid flies; parasitoids are prevalent during the first half of the evening in the fall mating season. I hypothesized that mating signals are influenced by sex-limited temporally fluctuating selection. I predicted how mating signals would respond to this pattern of cyclical selection a priori, and then measured the sexual characters over four successive generations. I provide correlative evidence that mating signals appear to respond to sex-limited temporally fluctuating selection. These results indicate that sex-limited temporally fluctuating selection may play a role in the maintenance of variation in these sexual characters.     

Different reproductive tactics in male collared flycatchers signalled by size of secondary sexual character

Most studies of variation in male reproductive tactics have focused on conspicuous categorical differences in mating behaviour (i.e. variation in mating strategies). However, in the presence of trade-offs between investment in competition over matings, parental care and survival, a male''s optimal allocation rule might vary according to his physiological condition and social or ecological environment. Thus, there may also be more subtle variation in male reproductive tactics. Here, I show that the reproductive effort (estimated as residual change in condition) of male collared flycatchers was affected by the size of their forehead patch (a secondary sexual character), age and date of arrival at the breeding grounds. Among early males (i.e. males with a high likelihood of both attracting more than one female and obtaining extra-pair copulations), large-patched males made a relatively large reproductive effort and as a result were in worse condition at the time of feeding offspring as compared to small-patched males. Furthermore, among early breeders, young males and males with experimentally increased forehead patch size made a relatively high effort. By contrast, regardless of age and badge size, there were no such patterns observed among late breeders. These results suggest that collared flycatchers use different reproductive tactics depending on both internal and external factors, and that the size of a secondary sexual trait may not only indicate variation in individual condition but also predict how resources will be allocated between pre- and post-mating reproductive activities.     

Chirp rate is independent of male condition in a synchronising bushcricket

Males of the bushcricket Mecopoda elongata synchronise their chirps with neighbouring males, but because synchrony is imperfect, one male's chirp preceeds the other by some 50-200 ms. Since a male's intrinsic chirp rate is critical for the establishment of the leader role in a duet, and females prefer the leader in a choice situation, we investigated a possible condition dependence of this male trait. In a duet leader males are usually those calling at a higher intrinsic rate; therefore, we investigated whether calling at a higher rate indicates male condition. The calling metabolism was quantified in a respirometer; the factorial slope of males calling at a high rate was three times higher compared to males calling at lower rates. Males produce on average 3.4 singing bouts/per night, and there is a significant increase in chirp periods (CPs) with successive singing bouts. Call properties were investigated throughout a male's life; chirp period increases significantly with age. Two groups of males were reared on either a low- or a high-nutrition diet, and the influence of male condition on different song parameters was investigated. CPs in two feeding regimes did not differ significantly, although males of the low-nutrition diet group were significantly affected by nutrition with respect to mortality, a delayed last moult and reduced weight as adults. We therefore conclude that solo chirp rates do not reflect phenotypic male condition properly.     

Variation in Male Calling Behaviour and Relation to Male Mating Success in the Strawberry Poison Frog (Dendrobates pumilio)

Calling behaviour and the characteristics of the males’ call are important for mate attraction, female mate choice and male mating success in anurans. In this study I investigated the variation in calling activity and the variation in one spectral (dominant frequency) and four temporal properties (call rate, call duration, duty cycle and pulse rate) of the advertisement call of strawberry poison frogs, Dendrobates pumilio, in two Costa Rican populations during two study periods. Regarding all call properties, no differences were found between populations or years, but between‐male variation was significantly higher than within‐recording and within‐male variation. Dominant frequency was less variable within and among males than temporal call properties and I classified the former as static and the latter as intermediate properties of the call. No call property seemed to be strongly affected by body length or weight. Moreover, I used behavioural observations to relate male calling behaviour to mating success. Calling activity was more variable between males and accounted for more variability in mating success in multiple regression models than any call property. Some call properties (call rate, dominant frequency and pulse rate) were also related to male mating success in one or two study periods. These call properties are probably important for females during courtship to assess their prospective mate and/or for competitive male–male interactions. Furthermore, I compared call properties and mating success of some males for which data were available for both study periods. Pulse rate was lower in the second than in the first year, suggesting age‐related changes in this call property. Moreover, male D. pumilio increased their relative mating success from the first to the second study year. Together with other studies, the data of this study suggest that, in anurans, endurance rivalry contributes more to variation in male mating success than does female choice based on male characteristics.     

Androgen and gonadotropin effects on male mate calls in South African clawed frogs, Xenopus laevis

Mate calling is a prominent reproductive behavior of male South African clawed frogs. Calls consist of alternating slow- and fast-amplitude-modulated trills. Each trill is made up of a series of clicks. The effects of administration of exogenous gonadotropin and androgen on mate calling were studied in male Xenopus laevis. Males were paired with unreceptive female frogs to elicit maximal calling. The amount of time each animal spent calling during the testing period, the peak fundamental frequency of the calls, the rate of calling, and the interclick interval (ICI, a measure of the temporal patterning of the calls) were measured in intact, castrated, and hormone-replaced frogs. Injection of human chorionic gonadotropin (HCG) into intact frogs increased the amount of time spent calling and the ICI relative to measures taken after water injection. Castrated males did not call even when given HCG. Testosterone and dihydrotestosterone treatment reinstated calling in castrates and increased circulating levels of androgens. When androgen-replaced castrated males were injected with HCG, the amount of time spent calling increased and approached levels of intact, HCG-injected males. The above results suggest that androgens are necessary for the production of calls. Gonadotropins appear to play an important role in mate calling, a role at least partly independent of effects on testicular androgen synthesis.     

Nutritional effects on male calling behaviour in the variable field cricket

In the variable field cricket, Gryllus lineaticeps, females prefer higher chirp rates and longer chirp durations in male calling song. Higher chirp rates are energetically more expensive to produce, but the energetic cost of calling does not vary with chirp duration. We tested the hypothesis that nutrition affects male chirp rate and chirp duration. Full-sibling brothers of similar age were placed on high- and low-nutrition feeding regimes. There was no effect of feeding regime on male weight; neither group showed a significant change in weight, and the two groups did not differ from each other in weight change. However, males on the high-nutrition feeding regime both called more frequently and called at higher chirp rates when they did call. The two groups did not differ in chirp duration, the duration of pulses within chirps or chirp dominant frequency. These results suggest that females select mates based on one nutrition-dependent call character (chirp rate) and one nutrition-independent call character (chirp duration). In addition, because males in the two groups did not show significant differences in weight change, and because males on the high-nutrition feeding regime engaged in energetically more expensive calling, these results suggest that males invest any excess energy above their basic maintenance requirements in the production of call types that increase their attractiveness to females. The absence of a relationship between body condition and calling song structure for males in the field may be a consequence of this pattern of energy allocation. Copyright 1999 The Association for the Study of Animal Behaviour.     

Variation in the number of sperm transferred during mating among males of the Colorado potato beetle (Coleoptera: Chrysomelidae)

We suggest an explanation for the bimodal distribution in the number of sperm transferred into the female during mating by males of the Colorado potato beetle (Leptinotarsa decemlineata). While in most matings females receive 20,000 sperm, in others they receive less than 1000. A variety of potential proximate causes for this bimodal distribution including experimental procedures and several physiological and behavioral causes were examined: male body size, female body size, male mating history, socio-sexual environments, duration and number of mounts, and duration and number of copulations. Surprisingly, none of these explained the observed distribution. However, we did find that the total time spent in copulation was significantly less variable among males that transferred more than 20,000 sperm than among males that passed less than 1000 sperm. In addition, males transferring large amounts of sperm resulted in proportionally more sperm inside the female’s spermatheca within a short period of time than males passing less than 1000 sperm. The lack of pattern in male sperm ejaculate suggests definite differences in sperm delivery tactics and may be related to the female condition rather than the male.     

Seasonal bat activity related to insect emergence at three temperate lakes

Knowledge of aquatic food resources entering terrestrial systems is important for food web studies and conservation planning. Bats, among other terrestrial consumers, often profit from aquatic insect emergence and their activity might be closely related to such events. However, there is a lack of studies which monitor bat activity simultaneously with aquatic insect emergence, especially from lakes. Thus, our aim was to understand the relationship between insect emergence and bat activity, and investigate whether there is a general spatial or seasonal pattern at lakeshores. We assessed whole‐night bat activity using acoustic monitoring and caught emerging and aerial flying insects at three different lakes through three seasons. We predicted that insect availability and seasonality explain the variation in bat activity, independent of the lake size and characteristics. Spatial (between lakes) differences of bat activity were stronger than temporal (seasonal) differences. Bat activity did not always correlate to insect emergence, probably because other factors, such as habitat characteristics, or bats’ energy requirements, play an important role as well. Aerial flying insects explained bat activity better than the emerged aquatic insects in the lake with lowest insect emergence. Bats were active throughout the night with some activity peaks, and the pattern of their activity also differed among lakes and seasons. Lakes are important habitats for bats, as they support diverse bat communities and activity throughout the night and the year when bats are active. Our study highlights that there are spatial and temporal differences in bat activity and its hourly nocturnal pattern, that should be considered when investigating aquatic–terrestrial interactions or designing conservation and monitoring plans.     

Acoustic defence in an insect: characteristics of defensive stridulation and differences between the sexes in the tettigoniid Poecilimon ornatus (Schmidt 1850)

Many insects exhibit secondary defence mechanisms upon contact with a predator, such as defensive sound production or regurgitation of gut contents. In the tettigoniid Poecilimon ornatus, both males and females are capable of sound production and of regurgitation. However, wing stridulatory structures for intraspecific acoustic communication evolved independently in males and females, and may result in different defence sounds. Here we investigate in P. ornatus whether secondary defence behaviours, in particular defence sounds, show sex-specific differences. The male defence sound differs significantly from the male calling song in that it has a longer syllable duration and a higher number of impulses per syllable. In females, the defence sound syllables are also significantly longer than the syllables of their response song to the male calling song. In addition, the acoustic disturbance stridulation differs notably between females and males as both sexes exhibit different temporal patterns of the defence sound. Furthermore, males use defence sounds more often than females. The higher proportion of male disturbance stridulation is consistent with a male-biased predation risk during calling and phonotactic behaviour. The temporal structures of the female and male defence sounds support a deimatic function of the startling sound in both females and males, rather than an adaptation for a particular temporal pattern. Independently of the clear differences in sound defence, no difference in regurgitation of gut content occurs between the sexes.     

Pheromonally mediated mate attraction by males of the burying beetle Nicrophorus orbicollis: alternative calling tactics conditional on both intrinsic and extrinsic factors

Male burying beetles attract females using a pheromonal signaland can provide parental care and a food resource, vertebratecarrion, for their developing offspring. But males attempt toattract females even when they have no carrion. We examinedthe factors that influence male behavior directed toward findingor attracting mates in both field-caught and laboratory-reared Nicrophorusorbicollis, a North American burying beetle. We investigatedwhether male behavior differed based on both intrinsic (size)and extrinsic (resources held) differences among males. Further,we examined repeatability of individual behaviors and the effectof holding or lacking resources on these repeatabilities. Field-caughtand laboratory-reared individuals differed in overall activitybut not in their behavioral repertoire, making studies of laboratory-rearedmales relevant. The behavior of individual males was very consistentwithin a condition, but plastic between resource conditions.The frequency of calling (adopting a posture that indicatespheromone release to attract females) depended on male sizewhen males did not hold resources, but this relationship disappearedwhen males held resources. Without carrion, smaller males calledmore frequently than did larger males. When holding carrion,smaller males reduced their calling, whereas larger males significantlyincreased the frequency with which they attempted to attractfemales and reduced the amount of time they spent searching.Thus, calling behavior of males was conditional on not only intrinsicand extrinsic factors, but also an interaction between them.We suggest that the changes in calling represent alternativetactics based on the costs and benefits of attracting both potentialmates and competitors, which differ for males of different sizes.     

Effects of a host plant,Brassica juncea, on calling behaviour and egg maturation in the diamondback moth,Plutella xylostella

The influence of the presence of seedlings of a host plant, Oriental mustard,Brassica juncea (L.) Coss., on the calling behaviour and egg maturation in the diamondback moth,Plutella xylostella (L.), was investigated. In the presence of plants, females began calling at a younger age, began calling earlier in the night, and spent more time calling. Females which were kept with plants until after their first night of calling and then were isolated from plants, subsequently called less, at a level not significantly different from females which had never been exposed to plants. Of all females, both with and without plants, which were dissected at the end of the first night of calling (n=150), 93% contained mature eggs. Heavier females were significantly more likely to have mature eggs at adult emergence, to begin calling at a younger age and to spend more time calling on their first night of calling. The number of developed eggs in calling females was not significantly different in the presence or absence of host plants on the first night of calling. However, by the second night, females in the presence of plants contained significantly more eggs, indicating that the presence of plants accelerated egg maturation, both before and after the onset of calling.     

Male monkeys remember which group members have given alarm calls

Primates give alarm calls in response to the presence of predators. In some species, such as the Thomas langur (Presbytis thomasi), males only emit alarm calls if there is an audience. An unanswered question is whether the audience's behaviour influences how long the male will continue his alarm calling. We tested three hypotheses that might explain the alarm calling duration of male Thomas langurs: the fatigue, group size and group member behaviour hypotheses. Fatigue and group size did not influence male alarm calling duration. We found that males only ceased calling shortly after all individuals in his group had given at least one alarm call. This shows that males keep track of and thus remember which group members have called.