Temperature coupling of mate attraction signals and female mate preferences in four populations of Enchenopa treehopper (Hemiptera: Membracidae)

Variation in temperature can affect the expression of a variety of important fitness‐related behaviours, including those involved with mate attraction and selection, with consequences for the coordination of mating across variable environments. We examined how temperature influences the expression of male mating signals and female mate preferences—as well as the relationship between how male signals and female mate preferences change across temperatures (signal–preference temperature coupling)—in Enchenopa binotata treehoppers. These small plant‐feeding insects communicate using plantborne vibrations, and our field surveys indicate they experience significant natural variation in temperature during the mating season. We tested for signal–preference temperature coupling in four populations of E. binotata by manipulating temperature in a controlled laboratory environment. We measured the frequency of male signals—the trait for which females show strongest preference—and female peak preference—the signal frequency most preferred by females—across a range of biologically relevant temperatures (18°C–36°C). We found a strong effect of temperature on both male signals and female preferences, which generated signal–preference temperature coupling within each population. Even in a population in which male signals mismatched female preferences, the temperature coupling reinforces predicted directional selection across all temperatures. Additionally, we found similar thermal sensitivity in signals and preferences across populations even though populations varied in the mean frequency of male signals and female peak preference. Together, these results suggest that temperature variation should not affect the action of sexual selection via female choice, but rather should reinforce stabilizing selection in populations with signal–preference matches, and directional selection in those with signal–preference mismatches. Finally, we do not predict that thermal variation will disrupt the coordination of mating in this species by generating signal–preference mismatches at thermal extremes.     

Divergence in Female Duetting Signals in the Enchenopa binotata Species Complex of Treehoppers (Hemiptera: Membracidae)

Sexual communication often involves signal exchanges between the sexes, or duetting, in which mate choice is expressed through response signals. With both sexes acting as signalers and receivers, variation in the signals of males and females may be important for mate choice, reproductive isolation, and divergence. In the Enchenopa binotata species complex – a case study of sympatric speciation in which vibrational duetting may have an important role – male signals are species‐specific, females choose among males on the basis of signal traits that reflect species and individual differences, and female preferences have exerted divergent selection on male signals. Here, we describe variation in female signals in the E. binotata species complex. We report substantial species differences in the spectral and temporal features of female signals, and in their timing relative to male signals. These differences were similar in range to differences in male signals in the E. binotata complex. We consider processes that might contribute to divergence in female signals, and suggest that signal evolution in the E. binotata complex may be influenced by mate choice in both sexes.     

Males adjust their signalling behaviour according to experience of male signals and male–female signal duets

Sexual signals are conspicuous sources of information about neighbouring competitors, and species in which males and females signal during pair formation provide various sources of public information to which individuals can adjust their behaviour. We performed two experiments with a duetting vibrational insect, Enchenopa binotata treehoppers (Hemiptera: Membracidae), to ask whether males adjust their signalling behaviour according to (1a) their own experience of competitors' signals, (1b) how females adjust their mate preferences on the basis of their experience of male signals (described in prior work), and/or (2) their own experience of female response signals to competitors' signals. We presented males with synthetic male signals of different frequencies and combinations thereof for 2 weeks. We recorded males a day after their last signal exposure, finding that (1a) male signal rate increased in response to experience of attractive competitors, but that (1b) male signal frequency did not shift in a manner consistent with how females adjust their mate preferences in those experience treatments. Second, we presented males with different male–female duets for 2 weeks, finding that (2) male signal length increased from experience of female duets with attractive competitors. Males thus make two types of adjustment according to two sources of public information: one provided by experience of male signals and another by experience of female responses to male signals. Signalling plasticity can generate feedback loops between the adjustments that males and females make, and we discuss the potential consequences of such feedback loops for the evolution of communication systems.     

The function of duetting in magpie-larks: conflict, cooperation, or commitment?

Avian duetting is a poorly understood phenomenon despite many hypotheses as to its function. Contrary to the recent view that duetting functions for mate guarding and is a result of conflict between the sexes, Australian magpie-larks, Grallina cyanoleuca, do not use duetting as a paternity guard. I used a playback experiment to investigate the role of antiphonal duetting in territorial defence and pair bond maintenance, two traditional hypotheses about the function of duetting. The experiment showed that, like many nonduetting species, magpie-larks recognize neighbours on the basis of song. It also provided evidence of functional differences between duetting and solo singing which indicate that temporal coordination of song between partners is used to maintain the territory and pair bond. Duets were more threatening territorial signals than solo songs: males initiated more vocalizations in response to playback of duets than playback of solos. Simulated intrusion also caused males and females to approach the speaker together and coordinate more of their vocalizations to form duets. Females did not engage in sex-specific territorial defence, responding equally strongly to playback of male and female song, and maintaining both territory and pair bond by attempting to exclude intruders of either sex. Males initiated more vocalizations in response to playback of male than female song, and their likelihood of duetting appeared to be related more to threats to the pair bond, in particular desertion by their partner. Copyright 2000 The Association for the Study of Animal Behaviour.     

Female Choice or Male Sex Drive? The Advantages of Male Body Size during Mating in Drosophila Melanogaster

The mating success of larger male Drosophila melanogaster in the laboratory and the wild has been traditionally been explained by female choice, even though the reasons are generally hard to reconcile. Female choice can explain this success by virtue of females taking less time to mate with preferred males, but so can the more aggressive or persistent courtships efforts of large males. Since mating is a negotiation between the two sexes, the behaviors of both are likely to interact and influence mating outcomes. Using a series of assays, we explored these negotiations by testing for the relative influence of male behaviors and its effect on influencing female courtship arousal threshold, which is the time taken for females to accept copulation. Our results show that large males indeed have higher copulation success compared to smaller males. Competition between two males or an increasing number of males had no influence on female sexual arousal threshold;—females therefore may have a relatively fixed ‘arousal threshold’ that must be reached before they are ready to mate, and larger males appear to be able to manipulate this threshold sooner. On the other hand, the females’ physiological and behavioral state drastically influences mating; once females have crossed the courtship arousal threshold they take less time to mate and mate indiscriminately with large and small males. Mating quicker with larger males may be misconstrued to be due to female choice; our results suggest that the mating advantage of larger males may be more a result of heightened male activity and relatively less of female choice. Body size per se may not be a trait under selection by female choice, but size likely amplifies male activity and signal outputs in courtship, allowing them to influence female arousal threshold faster.     

Complex Courtship Behavior in the Striped Ground Cricket, Allonemobius socius (Orthoptera: Gryllidae): Does Social Environment Affect Male and Female Behavior?

Complex courtship in the striped ground cricket, Allonemobius socius, involves a series of behaviors alternating between the sexes. We examined if complex courtship allows either or both genders to evaluate their mate and how mating behavior changes in different social environments. While complex courtship may allow discrimination by both sexes, here only females exhibited a preference. Males did not alter their courtship behavior or change spermatophore size for different size females. In contrast, females initiated copulation more quickly with bigger males possessing bigger spermatophores. In a different social environment (additional male, female, or both), males were less likely to omit courtship songs and female discrimination of mates changed. The distinct differences in male and female behavior suggest that subtle changes in social environment can have important consequences in structuring courtship and mating behavior.     

Function of pair duets in the eastern whipbird: cooperative defense or sexual conflict?

Paired male and female eastern whipbirds, Psophodes olivaceus,sing precisely coordinated, male-led duets. Four broad explanationshave been proposed for the function of duets: 1) cooperativeresource defense, 2) prevention of partner usurpation, 3) defenseof an individual's own position within the partnership, or 4)mate identification and localization. These 4 hypotheses makedifferent predictions about how male and female residents shouldrespond to simulated intrusion by other pairs or individuals.We compared the behavioral and vocal responses of 20 pairs ofeastern whipbirds to simulated territorial intrusions by: 1)a solitary singing male, 2) a solitary singing female, and 3)a duetting pair. Males and females did not coordinate theirapproach to the playback speaker and showed sex-specific responsesto playback. Males did not respond differently to duetting versussolo singing intruders. By contrast, females approached moreclosely during solo female song than during solo male song orduet playback. Females also produced specific vocalizationsonly in response to duet and solo female playback. Both sexesapproached the speaker more closely and quickly during playbackof same-sex solo songs than opposite-sex solo songs. Finally,females answered more of their mate's songs during simulatedintrusion by a lone female than during simulated intrusion bya lone male. Our results suggest that duets in this speciesprimarily function to allow females to defend their exclusiveposition in a partnership. Mate defense by females is unusualin birds but may be promoted in eastern whipbirds by a female-biasedsex ratio and the need for exclusive access to male care. Thus,duets result from independent and conflicting strategies ofmate and territory defense in males and females.     

Emergence, Mating, and Postmating Behaviors of the Oriental Beetle (Coleoptera: Scarabaeidae)

In a previous field-trapping study of the oriental beetle, Exomala orientalis (Waterhouse), by using synthetic sex pheromone on golf course fairways, numerous males were observed and trapped during the hours of peak mating activity. However, very few beetles were observed in the same areas when synthetic pheromone was absent. To investigate the hypothesis that mating in nature occurs cryptically within vegetation at the soil surface, laboratory studies on female emergence and pheromone release, male emergence and mate-locating, and female and male mating behaviors were conducted. Mate acquisition and copulation occurred on the soil surface near the female emergence site, with both sexes engaging in pheromone-mediated behaviors after having emerged from the soil. A highly stereotyped female pheromone release, or calling, behavior was observed, consisting of insertion of the female's head into the soil and elevation of the tip of her abdomen into the air. Bioassays conducted in a wind tunnel that simulated a turf fairway environment showed that walking and flying were both important in the upwind response of males to females. Mating and copulation occurred without an obvious complex courtship, but observations of postmating behaviors suggested that mate guarding occurs.     

Courtship behavior and discrimination between potential mates by maleDelia antiqua (Diptera: Anthomyiidae)

The repertoire of courtship behaviors of male onion maggots,Delia antiqua (Meigen), in a laboratory bioassay chamber, was analyzed by direct observation and by video recordings, in conjunction with a multichannel event recorder. Seven courtship behaviors were categorized: inspection from the substrate, aerial inspection, contact from the substrate, contact from the air, genital alignment, copulation, and male-male interaction. The frequency distribution of copulation bouts was best described by a Poisson distribution; peak mating activity occurred about 1 h into the bioassay. The duration in copulo, however, was extremely variable. On average, males spent 30 sin copulo (n=183); <30% of bouts were >50 s. The ability of males to discriminate between sexes, sexually immature and mature females, and between females ofD. antiqua and the cabbage maggot,Delia radicum (L.), was most pronounced in the elements of genital alignment and attempted copulation. The courtship and mating behavior inD. antiqua is consistent with a sequence that relies initially primarily on indiscriminate visual recognition of a potential mate, followed by species-and sex-specific semiochemical recognition upon contact.     

Mating Behavior of Cephalonomia tarsalis (Ashmead) (Hymenoptera: Bethylidae) and the Effect of Female Mating Frequency on Offspring Production

The courtship behavior of Cephalonomia tarsalis, a solitary semiectoparasitoid of Oryzaephilus surinamensis, was investigated in the laboratory. Courtship behavior includes a series of stereotypic movements. Males play the most active role, executing the majority of courtship action, and females respond with relatively limited observable behaviors. Males typically keep antennae still during encounters with females prior to mounting, which may be correlated with recognition of the female's sexual status. After mounting, males display a series of movements on females, such as antennae touching female's antennae, antennae or mouth touching female's head or thorax, and walking around on female, which may serve to stimulate females towards increased receptivity. Females signal receptivity by assuming a stereotypical posture of remaining stationary, with head down, and antennae still in front of the body. The male then inserts his aedeagus and the pair copulates. After an average of 40.4 s of copulation, females signal the end of copulation by waving the antennae and moving away from the copulation site. Males continue copulating for a short time after females start moving but dismount soon thereafter. After dismounting, the two wasps move away from each other immediately, and they typically begin grooming. Neither males nor females exhibit mating preference based on mate's mating status in both choice and no-choice tests. The male is polygynous and the mated female can mate multiple times within the first 3 days after starting oviposition. However, female mating frequency does not affect the production of female progeny.     

Co-Evolution of the Mating Position and Male Genitalia in Insects: A Case Study of a Hangingfly

Hangingflies are unique for the male providing a nuptial gift to the female during mating and taking a face-to-face hanging copulation with the female. Their male genitalia are peculiar for an extremely elongated penisfilum, a pair of well-developed epandrial lobes (9th tergum), and a pair of degenerated gonostyli. However, the co-evolution of their face-to-face copulation behavior and the male genitalia has rarely been studied hitherto. In this paper the mating behavior of the hangingfly Bittacus planus Cheng, 1949 was observed under laboratory conditions, and the morphology of the male and female external genitalia was investigated using light and scanning electron microscopy. The male provides an insect prey as a nuptial gift to the female in courtship and mating process, and commits a face-to-face copulation. During copulation, the male abdomen twists temporarily about 180° to accommodate their face-to-face mating position. The aedeagal complex has an extremely elongated penisfilum, corresponding to the elongated spermathecal duct of the female. The well-developed epandrial lobes serve as claspers to grasp the female subgenital plate during copulation, replacing the function of gonostyli, which are greatly reduced in Bittacidae. The modified proctiger assists the penisfilum to stretch and to enter into the female spermathecal duct. The possible reasons why this species might mate face-to-face are briefly discussed.     

Firefly Courtship: Behavioral and Morphological Predictors of Male Mating Success in Photinus greeni

Differences in male mating success can generate selection on male morphological traits and courtship behaviors involved in male–male competition or female mate choice. In Photinus fireflies (Coleoptera: Lampyridae), courtship is based on bioluminescent flash signals produced by both sexes. We conducted field observations of Photinus greeni fireflies engaged in competitive courtships, in which females are able to simultaneously assess several males, to identify male morphological traits and courtship behaviors that might predict male mating success. Male morphological traits did not differ between males that successfully mated compared with unsuccessful males (dialoging males that did not mate). However, courtship behavioral interactions differed: successful males tended to have higher flash pattern rates (number of flash patterns per minute), and their courtship flashes were more likely to be answered by females. We also examined how the risk of predation by Photuris fireflies altered courtship behavior of their Photinus prey. When predatory Photuris fireflies were present, P. greeni females were less likely to mate, and showed decreased flash responses to most males. However, P. greeni males that did successfully mate in spite of Photuris presence were males that maintained high flash pattern rates that elicited female responses. These results suggest that both female mate choice and Photuris predation exert strong selective pressures on the evolution of courtship signals in Photinus fireflies.     

The Effect of Limb Loss on the Courtship and Mating Behavior of the Wolf Spider Pardosa milvina (Araneae: Lycosidae)

Limb loss is common in the wolf spider Pardosa milvina, appearing in nearly one third of adult males but occurring less frequently among adult females and juveniles. Since males wave their first pair of legs during courtship displays, the reproductive consequences of limb loss may be significant. We measured the courtship and mating effects of the loss of one, two, or four legs among adult male P. milvina. Missing one or two legs did not significantly reduce a male's ability to mate, but missing four legs was detrimental to mating success, reduced both courtship intensity and copulation duration, and increased cannibalism frequency. Results suggest behavioral flexibility in compensating for limited leg loss and a defensive function of the anteriormost legs to thwart female cannibalism attempts.     

Song repertoire and duetting behaviour of the tropical boubou, Laniarius aethiopicus

Female song and the significance of duetting in birds remain puzzling, in part because of the limited number of species studied. We investigated duetting behaviour in the tropical boubou in West Africa. Birds produced a diverse song repertoire consisting of solos, duets and trios, with duets being the most conspicuous vocalization. We identified 12 distinct duet types in which tonal and broadband notes were combined by males and females in highly synchronized and temporally precise patterns. Molecular sexing and observations of colour-banded birds revealed that duets were initiated by both sexes, with strict sex-specific roles maintained within the duet. An equal number of duet types was initiated by males and females. Seven of 12 duets were terminated by the male. Male-initiated duets were also more common (89% of total) and were repeated more often, i.e. sung more frequently. Solo singing occurred when partners did not respond and was also used by unpaired birds. Trios were produced by subadult birds joining the duet of a resident pair. The wide variety of contexts in which duets were sung suggests that they serve several functions, including territorial defence and mutual mate guarding.     

Loss of complex female song but not duetting in the ancestors of Carolina wrens

Female singing and coordinated male–female duetting are often but not always found in the same species. Both behaviors are more common in tropical than temperate songbirds, but few studies have differentiated between the factors selecting for each. Here we investigate the evolution of female vocal complexity and male–female vocal coordination in Carolina wrens (Thryothorus ludovicianus), one of the few non-tropical members of a songbird family (Troglodytidae) that is well known for producing coordinated male–female duets. Female Carolina wrens are not known to sing; rather, they produce relatively simple, sex-specific chatters, often during territorial encounters. We analyzed field recordings to show that females coordinate these chatters with male songs at rates similar to those observed in some tropical duetting wren species. We then used phylogenetic comparative methods to show that the evolutionary ancestors of Carolina wrens had female songs that were more acoustically complex than the vocalizations of current females, suggesting past selection against female vocal complexity. Levels of vocal coordination with males, in contrast, have changed relatively little from those of tropical ancestors. Our results suggest that these two aspects of female behavior, acoustic complexity and vocal coordination with males, have evolved independently and have different functions in communication.     

Male search behavior of the stonefly,Pteronarcella badia (hagen) (Plecoptera: Pteronarcyidae), in relation to drumming

The mating system of many northern hemisphere stoneflies involves vibrational duetting, yet searching behavior in relation to the percussive communication has remained unreported. The communication-search system of a Colorado population of the ubiquitous western stonefly Pteronarcella badiain an experimental arena entails (1) a ranging search by calling virgin or polygynous males until duet establishment with virgin females, (2) continued intermittent duetting while the male engages in a local search and the female remains stationary, and (3) a tactile-based find of the female, followed by immediate mounting and mating. The average 234-s find time for pairs engaging in strong, continuous duets was significantly shorter than those for nonduetting or anomalously duetting pairs. Males of strongly duetting pairs made significantly more turns toward the female during search than in anomalously duetting pairs, and their local search pattern appears to be a triangulation aided by resource-specific (female) vibrational cues. Males displayed a wide range of fitness, based on searching time, and an increased number of duets significantly reduced finding time. Potential female selection and possible extrapolation of these experimental results to natural field conditions are discussed.     

The evolution of male and female mating preferences in Drosophila speciation*

The relative importance of male and female mating preferences in causing sexual isolation between species remains a major unresolved question in speciation. Despite previous work showing that male courtship bias and/or female copulation bias for conspecifics occur in many taxa, the present study is one of the first large‐scale works to study their relative divergence. To achieve this, we used data from the literature and present experiments across 66 Drosophila species pairs. Our results revealed that male and female mate preferences are both ubiquitous in Drosophila but evolved largely independently, suggesting different underlying evolutionary and genetic mechanisms. Moreover, their relative divergence strongly depends on the geographical relationship of species. Between allopatric species, male courtship and female copulation preferences diverged at very similar rates, evolving approximately linearly with time of divergence. In sharp contrast, between sympatric species pairs, female preferences diverged much more rapidly than male preferences and were the only drivers of enhanced sexual isolation in sympatry and Reproductive Character Displacement (RCD). Not only does this result suggest that females are primarily responsible for such processes as reinforcement, but it also implies that evolved female preferences may reduce selection for further divergence of male courtship preferences in sympatry.     

Breeding in the crayfish, Austropotamobius pallipes: mating patterns, mate choice and intermale competition

1. The breeding behaviour of the white-clawed crayfish, Austropotamobius pallipes, which is in decline throughout Europe, was analysed. 2. Observations and experiments focused on: (i) several aspects of mating behaviour, showing the primary role of the female during courtship, the patterns of mature male receptivity and the possible existence of a mating stimulus produced by either a mating pair or a receptive female; (ii) the potential for mate choice either by males (males were not choosy, mating with the first receptive female they met) or by females (that rejected the smallest males and males with only one cheliped) and (iii) intermale fighting before copulation (larger males copulated more often, with the exception of some ’sneakers‘). 3. Sperm competition occurred when a new male was presented with a female after a previous male‘s spermatophore had been deposited; the new male cleaned the rival male‘s spermatophore by feeding on it before copulation. This behaviour has also been observed in some pseudoscorpions, millipedes and collembolans, but always in cases where the spermatophore is deposited on the substratum. Introduction     

Partners coordinate territorial defense against simulated intruders in a duetting ovenbird

Duets in breeding pairs may reflect a situation of conflict, whereby an individual answers its partner's song as a form of unilateral acoustic mate guarding or, alternatively, it may reflect cooperation, when individuals share in territory defense or safeguard the partnership. The degree of coordination between the sexes when responding to solo versus paired intruders may elucidate the function of songs in duets. We examined this issue in a study with rufous horneros (Furnarius rufus), a duetting, socially monogamous Neotropical species with low levels of extrapair paternity. We exposed social pairs during the nonbreeding season to playbacks of duets, male solos, female solos, and control heterospecific songs. Partners approached all conspecific stimuli together and responded by singing quickly, at higher rates and by coordinating ~80% of their songs into duets. For both sexes, most response variables (seven of nine) did not vary across conspecific treatments. These results suggest that partners duet and coordinate behaviors to cooperatively defend common territories. However, females spent more time in territorial vigilance, and partners were highly coordinated (correlated responses) in response to duets and female solos in comparison with male solos. This indicates that female intrusions (paired or solo) might be more threatening than male intrusions in the nonbreeding season, especially for territorial females, and that females are less cooperative with their partners in territory defense against male intruders.     

Courtship song, male agonistic encounters, and female mate choice in the house cricket,Acheta domesticus (Orthoptera: Gryllidae)

Whether female crickets choose among males based on characteristics of the courtship song is uncertain, but in many species, males not producing courtship song do not mate. In the house cricket,Acheta domesticus, we examined whether a female chose or rejected a male based on his size, latency to chirp, latency to produce courtship song, or rate of the high-frequency pulse of courtship song (“court rate”). We confirmed that females mated only with males that produced courtship song, but we found no evidence that the other factors we measured affected a female’s decision to mate. In addition, we investigated whether the outcome of male agonistic encounters affected the subsequent production of courtship song. In one experiment, we observed courtship and mating behavior when a single female was placed with a pair of males following a 10-min interaction period between the two males. Winners of male agonistic encounters had higher mating success. However, winners and losers of agonistic encounters were not different in their likelihood or latency to produce courtship song or in the number of times they were disrupted by the other male in the pair. In a second experiment, we allowed two males to interact for a 10-min period, but following this interaction period, we placed a female with each male separately and observed courtship and mating behavior. The mating success of winners and losers was not different under these circumstances, and we found no differences between winners and losers in any subsequent courtship or mating behavior examined. We conclude that winning agonistic encounters influences a male’s mating success in ways other than his production of courtship song and this effect is lost when winning and losing males are separated and each is given an opportunity to mate.