Courtship behaviour

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Latency for facultative expression of male typical courtship behaviour by female bluehead wrasses depends on social rank: the ‘priming/gating’ hypothesis

Latency for facultative expression of male typical courtship behaviour by female bluehead wrasses depends on social rank: the ‘priming/gating’ hypothesis

Behavioural transformation in bluehead wrasses may reflect a rapid, irreversible physiological change triggered by loss of the TP male. Alternatively, the presence of a territorial TP fish may inhibit the expression of male-typical courtship behaviours in IP females that are already physiologically competent to assume the male role. In this ‘ priming/gating hypothesis ’ , we predict that IP fish become more competent to transform as they ascend the social hierarchy. The positions of IP fish in the hierarchy are related to their relative sizes (Warner and Swearer, 1991; Warner and Schultz, 1992), and established through aggressive interactions (e.g. dominant fish chase subordinate fish away from food sources). In this model, the physiological changes that underlie behavioural gender plasticity occur with a longer time course than is generally postulated. Once competent, i.e. ‘ primed ’ , the expression of male-typical courtship behaviour by an IP female can be toggled off or on depending on whether a territorial TP is present or absent, respectively. This model for behavioural ‘ transformation ’ is conceptually similar to classical ethological examples of sensorimotor gating (Pearson, 1993). For example, flight behaviour in insects is gated on or off depending on whether the feet are in contact with the substrate. The gating model could be extended to account for the presence of ‘ non- territorial ’ phenotypes in bluehead wrasses (as proposed by Perry and Grober, 2003) and social regulation of expression of aggressive signalling in cichlids (Fernald and Hirata, 1977; Maruska and Fernald, 2010; Desjardins et al., 2012).

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The role of the antennae during courtship behaviour in the parasitic wasp
Trichopria drosophilae

The role of the antennae during courtship behaviour in the parasitic wasp Trichopria drosophilae

We have studied the courtship behaviour of Trichopria drosophilae Perkins (Hymenoptera: Diapriidae), a pupal parasitoid of the common fruit fly Drosophila melanogaster Meigen (Diptera: Drosophilidae), to understand the role of the antennae. Virgin pairs of the parasitoid perform an intense and stereotyped antennal courtship, which leads to copulation. During antennation, the two male fourth antennomeres come into contact with the two apical female antennomeres, and thus the secretion produced by the sex pheromone gland is spread onto the female receptors. By preventing the transfer of the courtship pheromone from male to female antennae, mating was inhibited. Moreover, selective ablation of single antennae demonstrated that the courtship pheromone acts on contact. When antennae of both sexes were partially removed (ablation at the same side, i.e. right or left) courtship was successful and copulation occurred. In contrast, in the case of antennal ablation at opposite sides, courtship failed despite the short distance between secretion and receptors. These results confirm the hypothesis that T. drosophilae male antennal glands are the release site of a contact courtship pheromone, playing a key role in mating behaviour. The occurrence of male antennal glands in Hymenoptera and other insect orders is discussed.

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A two-fly tracker that solves occlusions by dynamic programming: computational analysis of Drosophila courtship behaviour

A two-fly tracker that solves occlusions by dynamic programming: computational analysis of Drosophila courtship behaviour

In particular, the work of Dankert et al. [8] has similar aspects to this paper as it also introduces a two-fly tracker and tackles similar challenges. Our tracker mainly differs in three aspects: It was initially designed to process unseen videos that are not specif- ically recorded for automated tracking and therefore comes with an arsenal of quality boosting and qual- ity control methods. Second, we spend a huge effort to tackle the occlusion problem, which was probably less critical for the application scenarios of [8]. Finally, our system offers top-down and bottom-up classifiers for courtship behaviour, while the other system offers top- down classifiers only but for both courtship and aggressive behaviours.

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Courtship behaviour of Phlebotomus papatasi the sand fly vector of cutaneous leishmaniasis

Courtship behaviour of Phlebotomus papatasi the sand fly vector of cutaneous leishmaniasis

In P. papatasi, the association between blood-feeding and mating is less clear. A blood meal is not always a prerequisite for egg production in this species [14-16], and the extent to which mating takes place in aggrega- tions on host animals is unknown. Current evidence may also suggest that male P. papatasi do not lek in the same manner as L. longipalpis: females are not attracted to large male aggregations [20], and males have not yet been observed to engage in behaviour which might func- tion in defending territories against other males. Further- more, while there is behavioural evidence to support the existence of male-produced sex pheromones in P. papa- tasi [20], this attractive chemical has not been found stored in large amounts in the body (J.G.C. Hamilton pers. obs.). This may indicate that sex pheromone does not play the same role in mate choice as it does in L. longipalpis, and has therefore not been subject to the same directional selection for greater pheromone production. In addition, while male wing flapping has been found to correlate with mating success in L. longipalpis [22,23], none of the male behaviours described here were identified as predictors of successful courtship, other than attempting to copulate.

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Mate preference in the painted goby: the influence of visual and acoustic courtship signals

Mate preference in the painted goby: the influence of visual and acoustic courtship signals

We tested the hypothesis that females of a small vocal marine fish with exclusive paternal care, the painted goby, prefer high parental-quality mates such as large or high-condition males. We tested the effect of male body size and male visual and acoustic courtship behaviour (playback experiments) on female mating preferences by measuring time spent near one of a two-choice stimuli. Females did not show preference for male size but preferred males that showed higher levels of courtship, a trait known to advertise condition (fat reserves). Also, time spent near the preferred male depended on male courtship effort. Playback experiments showed that when sound was combined with visual stimuli (a male confined in a small aquarium placed near each speaker), females spent more time near the male associated with courtship sound than with the control male (associated with white noise or silence). Although male visual courtship effort also affected female preference in the pre-playback period, this effect decreased during playback and disappeared in the post-playback period. Courtship sound stimuli alone did not elicit female preference in relation to a control. Taken together, the results suggest that visual and mainly acoustic courtship displays are subject to mate preference and may advertise parental quality in this species. Our results indicate that visual and acoustic signals interplay in a complex fashion and highlight the need to examine how different sensory modalities affect mating preferences in fish and other vertebrates.

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Females in Control: Female Sensitivity to Predation Risk Affects Courtship and Reproductive Behaviours

Females in Control: Female Sensitivity to Predation Risk Affects Courtship and Reproductive Behaviours

foraging, but predation risk may not necessarily negatively impact their overall energy intake if prey greatly increase their foraging during times of low risk to prevent starvation according to the „interrupted foraging hypothesis‟ (Lima 1986, Lima & Bednekoff 1999). This idea is part of „risk allocation hypothesis‟ in the foraging literature which describes how prey can change the timing in which they feed according to temporal variations in the level of predation risk (Lima & Bednekoff 1999). Similarly, to mitigate the risks of courtship, individuals can alter when they signal or receive signals (Endler 1980, Sih 1987, Lima & Dill 1990, Hughes et al. 2012). However, temporal variations in risk have yet to be considered in studies examining predation risk effects on courtship behaviour (Hughes et al. 2012). In a separate paper, we presented results demonstrating that the birds in this study gained mass under predation risk, which can be attributed to large increases in foraging during low risk periods as per the „interrupted foraging hypothesis‟ (Walters 2015). The birds in this study may also adjust their courtship behaviour to court more during times of low risk such that overall levels of courtship may not be affected by predation risk. Thus, testing whether predation risk actually affects courtship behaviour to an extent that can affect reproductive success in nature requires presenting prey with predation risk that varies temporally over an extended period of time as I have aimed to do so in the study presented here.

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Pampas deer (Ozotoceros bezoarticus) courtship and mating behavior

Pampas deer (Ozotoceros bezoarticus) courtship and mating behavior

Male courtship behaviors are those that involve an ac- tive sexual display from the male toward a female. The mating period included the behaviors displayed immedi- ately before, during, and after the mating, ending when the female was no longer receptive and the interactions between male and female stopped. We developed a basic sexual ethogram analyzing the real-time data and the videos. Considering the previous descriptions available from other widely studied deer species, and other small ruminants, including red deer [2], mouflon [20] and sheep [21], as well as preliminary information reported in pampas deer [12,16] we performed male courtship and male and female mating ethograms. The frequency in which each behavior was displayed in different con- texts (males during courtship and males and females during mating periods) was calculated. We also analyzed the average number of mounts per mating period, and the length of the receptive period and of each mating.

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Moths are not silent, but whisper ultrasonic courtship songs

Moths are not silent, but whisper ultrasonic courtship songs

Fig.  5. Pressure levels of sounds emitted by various moths for mating, territorial display or bat-defense. (A)  Scatter plot of the sound pressure levels (0  dB  SPL  20   Pa) at a distance of 1  cm from the moths, arranged by family. The levels (mean ± s.d.) of each family are as follows: Noctuidae, 114±24  dB  peSPL, N 7 species; Arctiidae, 103±11  dB  peSPL, N 46 species; Nolidae, 123±0  dB  peSPL, N 2 species; Pyralidae, 92±18  dB  peSPL, N 4 species; Crambidae, 101±27  dB  peSPL, N 6 species; Geometridae, 73  dB  peSPL, N 1 species. The color of symbols denotes the sound-producing mechanism: red, stridulation with a file-scraper; black, clicking with a tymbal; blue, percussion with an alar castanet [stridulation, 111±30  dB peSPL (mean ±s.d.), N 6 species; click, 105±16  dB  peSPL, N 59 species; percussion, 114  dB  peSPL, N 1 species]. Filled circles: sound emissions for advertisement, territorial and bat-defense calls; open circles: unknown function at long distance; stars: sound emission for courtship calls at close distance. See B for descriptive statistics. Numbers near the symbols are the moth species: 1, Rileyana (Thecophora) fovea; 2, Amyna natalis; 3, Hecatesia exultans; 4, Heliothis zea; 5, Amphipyra perflua; 6, Spodoptera litura; 7, Herminia tarsicrinalis; 8, Syntomeida epilais; 9, Cycnia tenera; 10, Arctia caja; 11, Euchaetes egle; 12, Phragmatobia fuliginosa; 13, Pyrrharctia isabella; 14, Euchaetes bolteri; 15, Empyreuma affinis; 16, Spilosoma punctarium; 17, Eilema japonica; 18, 36 arctiid species studied by Barber and Conner (Barber and Conner, 2006) (filled circle and bar indicate median and range, respectively); 19, Bena bicolorana; 20, Pseudoips prasinana; 21, Corcyra cephalonica; 22, Achroia grisella; 23, Galleria mellonella; 24, Plodia interpunctella; 25, Symmoracma minoralis; 26, Chilo suppressalis; 27, Palpita nigropunctalis; 28, Glyphodes pyloalis; 29, Ostrinia furnacalis; 30, Spoladea recurvalis; 31, Ascotis selenaria. Sound pressure levels, normalized at a distance of 1  cm, of species 1–17, 19–31 were obtained from Nakano et al. (Nakano et al., 2008; Nakano et al., 2009). (B)  Difference in sound pressure levels of acoustic signals between effective ranges. ‘Close’ [64±11  dB  peSPL, N 12 species (64±10  dB  peSPL, N 9 species used in the present study), symbolized by stars in A] indicates sound emission within 5  cm during courtship, and ‘far’ (100±11  dB  peSPL, N 54 species, symbolized by circles in A) indicates emission of ultrasounds beyond 5  cm for advertisement to mates and rivals, and for avoidance of bat predation. Box-and-whisker plots show the median, lower and upper quartiles, and adjacent values within 1.5 ⫻ interquartile ranges from the quartiles. A significant difference exists between the ranges (LR test in GLM, Gaussian error, identity link,  2

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Social effects on fruit fly courtship song

Social effects on fruit fly courtship song

The memory mutant strains we used to investigate the role of learn‐ ing in plastic song responses have been shown to display dysfunctional social responses and courtship learning (Griffith, 2014). For instance, amnesiac, but not dunce, do not increase mating duration in response to the presence of conspecific rivals (Rouse et al., 2018). Similarly, both amnesiac and dunce fail to suppress courtship efforts after unsuccess‐ ful mating attempts (Emmons & Lipton, 2003). We found that amnesiac and dunce strains adjusted their singing effort in response to the so‐ cial environment in a similar manner to wild‐type D. melanogaster. We cannot reject the hypothesis that such social responses involve learning and memory, but our data suggest that the learning pathways disrupted by amnesiac and dunce mutations are not involved. We consider it likely that during early adult development, males are sensitive to the pres‐ ence of other males in the environment and adjust allocation to court‐ ship effort, as well as the strategic allocation they make to postmating investment. Allocation to singing effort could therefore represent more of an investment trade‐off modulated by social exposure, rather than an actively learned response. Further studies would be required to as‐ sess costs of such a trade‐off and the contribution of different sensory modalities or signals as cues in this plasticity.

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Through their eyes: selective attention in peahens during courtship

Through their eyes: selective attention in peahens during courtship

Charles Darwin (Darwin, 1871) introduced the concept of sexual selection to explain seemingly ornamental display traits in animals, such as the elaborate trains of peacocks. It is now well established that sexual selection can operate through female choice when females choose their mates based upon these displays (Andersson, 1994; Bradbury and Vehrencamp, 2011). However, the cognitive and sensory processes that females use to assess male displays and choose their mates remain mysterious (Jones and Ratterman, 2009). This is particularly true in species where males have complex displays with multiple components, which could contribute to female choice, competition among males, or both (Hebets and Papaj, 2005). Selection may favor attention mechanisms, which may have originally evolved in response to natural selection (Basolo, 1990; Ryan, 1998), allowing females to selectively and efficiently acquire information relevant to mating decisions (Dukas, 2002). These female attention mechanisms may in turn influence the evolution of male courtship displays, favoring males with traits that effectively capture and hold female attention (Fleishman, 1988; Hebets and Papaj, 2005; Rosenthal, 2007). A promising tool to examine the role of attention mechanisms in mate choice is eye-tracking, which has found strong links between visual orienting and choice behavior in human and non-human primates (Land and Tatler, 2009; Shimojo et al., 2003). Despite a growing literature on visual attention, we still know little about how animals evaluate aspects of their environment that are crucial to their reproduction (Rupp and Wallen, 2007; Yorzinski and Platt, 2010).

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Phylogeny, environment and sexual communication across the Drosophila genus

Phylogeny, environment and sexual communication across the Drosophila genus

Species differences in CHCs are not only correlated to abiotic selection pressures. Even the particular blends of cuticular hydrocarbons appear to be a trait that expresses species identification signals that have higher variation between closely related species, suggesting a link to reinforcement barriers. A classic example is D. melanogaster and D. simulans, two sibling species that often coexist in sympatry (Markow, 2015). Drosophila melanogaster females produce unsaturated CHCs with two double bonds in the carbon chain (dienes) and the males do not, making this species sexually dimorphic in terms of its CHC blends. However, D. simulans does not produce dienes, and dienes exhibit anti- aphrodisiac qualities to D. simulans males (Billeter et al., 2009). Considering the CHC profile of other closely related species reveals that D. sechellia and D. erecta females produce dienes like D. melanogaster, but D. yakuba females do not (Ferveur, 2005; Cobb and Jallon, 1990). Drosophila yakuba and D. erecta are sibling species that are sympatric in western Africa (Lachaise et al., 1988), and this parallels the relationship between D. melanogaster and D. simulans, suggesting character displacement. Although the size and quantity of CHCs a species synthesizes could depend on environmental pressures, such as heat and aridity, the qualitative aspects of CHC profiles (i.e. the range of compounds synthesized) appear sensitive to reproductive isolation mechanisms. Interestingly, an experimental evolution study utilizing D. serrata and D. birchii found that exposing the two species to novel diets (rice and corn) caused a divergence in both CHCs and female mating preference (Rundle et al., 2005). Consequently, the distribution of individual CHC compounds may not hold any phylogenetic correlation and may evolve frequently based on reinforcement, mate selection and shifts in host resources. Accordingly, by compiling a list of species that are known to lack dimorphism in their CHC profile and considering the phylogenetic relationships of these species, we see that this is a trait that appears to have independent origins (Fig. 4). Interestingly, species that show courtship repression in the dark, such as D. grimshawi, D. suzukii, D. simulans and D. mauritiana, are examples of species without dimorphic CHCs (Fig. 4). The lack of dimorphism in their CHCs may partially explain why these species depend on visual cues for courtship and mating. Of special note is that the arid species sampled are exclusively dimorphic in CHCs. This adds credence to our earlier hypothesis of arid repleta species underemphasizing visual signals during courtship. One hypothesis that follows from this observation is that the environmental pressures on arid species are so high that dimorphic CHC profiles became essential for social communication either in dark and cool microhabitats, or at a time of day when sun exposure is reduced.

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Voyeurism, Intrusion and Aggression: the courtship narratives of modern masala

Voyeurism, Intrusion and Aggression: the courtship narratives of modern masala

As the film progresses, however, Zoya‟s apparent independence is revealed to be a ruse. It is an act of intrusion that reveals the truth, as Tiger enters the professor‟s house through a window and finds Zoya downloading secret files. Zoya‟s threat – that of the femme fatale – is at once uncovered and defeated, as Tiger holds a gun to her head. While the revelation of Zoya‟s identity adds a layer of intrigue to her character, it shatters the myth of independence that was initially a feature of her character: Zoya‟s actions are no longer hers but those of her sinister male handlers, her mobility not a sign of her individual freedom but in fact a symptom of their control. After stripping Zoya of her façade, Tiger is able to exert strict control over a courtship that now falls back on the traditional patterns of female restriction and male intrusion. Hearing that Zoya is in Istanbul for a United Nations conference, he decides to attend and finds her dressed in traditional feminine clothes and surrounded by male delegates. After following her from the conference he climbs into her taxi, taking control as he tells the taxi driver where to go. Suddenly traditional dynamics are in the ascendancy: Zoya claims that a relationship is impossible, while Tiger promises to fight off the forces of disunity and make it a reality. Their meeting at the conference leads to a song, “Saiyaara,” (Planet), the lyrics of which celebrate their developing love while acknowledging its difficulties. The accompanying shots show both characters looking emotional as they reflect on the meeting, but a disparity of mobility is evident: the passive Zoya returns to her hotel room and sits idly on her bed, while the active Tiger walks by the vast Bosphorus Strait and looks out from the deck of a ferry.

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A Novel Gene Controlling the Timing of Courtship Initiation in Drosophila melanogaster

A Novel Gene Controlling the Timing of Courtship Initiation in Drosophila melanogaster

during development due to female-specific splicing of fru transcripts. This could result in changes in neurite out- growth and synapse formation that results in aberrant courtship initiation. However, in Tre1 mutant males, the Tre1 neurons are presumably correctly masculinized. The developmental absence of Tre1 would still be expected to result in rapid courtship initiation; however, the reduced copulation success and competitive disadvantage seen in Tre1 mutant males may result from later requirements for Tre1. These hypotheses could be tested by characterizing the critical developmental period for each of the Tre1 court- ship and copulation phenotypes.

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Complex sexual courtship displays by luminescent male marine ostracods

Complex sexual courtship displays by luminescent male marine ostracods

In the western Caribbean Sea, about an hour after the sun sets, a complex and ritualized light show of precise, vertically placed luminescent pulses erupts over shallow grassbeds. These are among the most complex displays known in marine systems. Displays consist of repeated trains of secreted bioluminescent pulses in a specific pattern ejected into the water column as courtship signals by male Vargula annecohenae , which are small (<2 mm) myodocopid ostracod crustaceans. Although these animals display in near darkness, we have used image intensification and infrared videography and three-dimensional analysis in the lab to demonstrate that each luminescent display train, which can be up to 60 cm long, consists of two distinct luminescent and swimming phases. The first, or ‘stationary,’ phase consists of three (usually) bright, longer pulses placed close together, with the male swimming in a looping pattern. We hypothesize that this pattern acts as an attention-grabbing signal for receptive females. The stationary phase is followed by the ‘helical phase,’ which consists of about a dozen evenly placed dimmer, shorter pulses secreted by an individual male rapidly spiraling upward in a helical pattern. We hypothesize that this phase, which has very uniform interpulse intervals and distances, helps an approaching female target and intercept the rapidly moving male. Here we provide details of these two phases, and produce a three-dimensional model of a multiply-displaying male.

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Recognition of variable courtship song in the field cricket Gryllus assimilis

Recognition of variable courtship song in the field cricket Gryllus assimilis

male CV values in excess of 20%. In contrast, only two of nine courtship characters studied in G. texensis and G. rubens and one of eight characters measured in T. oceanicus had CV values exceeding 20% (Balakrishnan and Pollack, 1996; Fitzpatrick and Gray, 2001). Those characters that are variable in G. assimilis are rather consistent in G. texensis and G. rubens. For example, the relative amplitude of chirps and ticks, the most variable character in G. assimilis, shows little variability in G. texensis and G. rubens. The number of pulses per chirp is more variable in G. assimilis than number of pulses per phrase in G. texensis and G. rubens (in which the low-frequency portion of the song is not organized into chirps). At the same time, there are similarities between the songs of these species. Interphrase interval in G. texensis and G. rubens and phrase period in the G. assimilis song (parameters that are likely to be correlated) are relatively variable, whereas pulse period is quite constant in all three species (CV ⬇ 10%), as well as in the courtship song of T. oceanicus (Balakrishnan and Pollack, 1996).

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A study of the courtship song parameter in the 'Drosophila melanogaster' species complex

A study of the courtship song parameter in the 'Drosophila melanogaster' species complex

Hall (1994) examined the effect of mutations in single genes on courtship song. Single genes have been shown to affect the length of the song cycle {period), receptivity of females {spinster), and the cycle number of individual pulses {cacophony). These examples show that changes of single genes could affect the nature of the courtship and comtship song. The process of isolating the genes that control IPI will be complicated if there ai*e no single genes of major effect (Cowling and Burnet, 1981; Ritchie and Kyriacou, 1994; Chapter 3). It is possible that there is a single gene controlling pheromone balance, which could be isolated and characterized, as with the period gene. Characterizing single genes provides a more attractive end result than defining the relative effects of different areas of the genome. However, Liu et al. (1996) have shown, through QTL analysis, that the relative contributions of sections of the genome can be studied in great detail. Their study resulted in the identification of QTL effects of only between 5 and 16% of parental

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quick-to-court, a Drosophila Mutant With Elevated Levels of Sexual Behavior, Is Defective in a Predicted Coiled-Coil Protein

quick-to-court, a Drosophila Mutant With Elevated Levels of Sexual Behavior, Is Defective in a Predicted Coiled-Coil Protein

that courtship behaviors (orientation, following, tapping, wing vibration, licking, or attempted copulation) were observed on average during 42% of a 10-min observation period (Tompkins 1984), whereas CS control males had a CI of only 4 ⫾ 1. CS was chosen as a control because the sexual behavior of CS flies has been thoroughly char- acterized (Tompkins et al. 1980; McRobert et al. 1995); however, since qtc 1 is not in a CS genetic background,

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Evolution of Reproductive Behavior

Evolution of Reproductive Behavior

Premating behaviors allow males and females to recognize and assess partners to determine whether the species, re- productive state, and condition of the partner are appropriate for and conducive to mating. Courtship rituals in Drosophila follow a multimodal pattern, mediated through visual, chem- ical, tactile, and auditory signals, and consist of a sequence of orientation, genital licking, courtship song through wing vi- bration, and attempted copulation [Hall (1994); reviewed in Sokolowski (2001); Figure 1]. Courtship behaviors include locating a potential mate by using visual or olfactory cues, and communicating with and assessing the potential mate by olfactory, auditory, and visual cues during courtship. Behav- iors during mating also are important for reproductive fitness (Markow and O’Grady 2008) and are variable. These include copulation duration as well as interactions that determine whether gametes will be present for fertilization. Although the courtship ritual of D. melanogaster males was once con- sidered a defined stereotypical progression of behavioral el- ements, studies using the DGRP showed extensive heritable variation in courtship patterns (Gaertner et al. 2015). Selec- tion acting on variation in any of the sensory inputs that drive the component behaviors of this multimodal courtship ritual can lead to a reproductive isolation barrier as a scaffold for incipient speciation.

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Male sex pheromone release and female mate choice in a butterfly

Male sex pheromone release and female mate choice in a butterfly

If male P. napi emit citral when courting females, this suggests that it might function as a male sex pheromone. In P. napi, as in other pierids such as Pieris rapae, Pieris brassicae and Anthocharis cardamines, unreceptive females react to male courtship by spreading their wings laterally and lifting their abdomen up in the air, performing the so-called ‘mate-refusal posture’ (Obara, 1964; Wiklund and Forsberg, 1985; Forsberg and Wiklund, 1989). This female behavior appears to have two functions. First, it makes it physically difficult for a courting male to couple with the female. Second, females use this posture to disseminate volatiles that can be perceived easily by the courting male; in many pierids males appear capable of distinguishing between virgin and mated females, and in P. napi, P. rapae and P. brassicae different volatiles are disseminated by virgin and mated females when exhibiting the mate-refusal posture (Andersson et al., 2000; Andersson et al., 2003). Both virgin and mated females initially exhibit the mate-refusal posture when approached by a male, but when a female accepts a courting male she signals her receptivity by closing her wings and acquiescing (Forsberg and Wiklund, 1989). When perceiving this signal the courting male alights next to the female, bends his abdomen sideways to make genital contact, couples with the female, and usually carries the female away in a short post-nuptial flight.

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A reply to Nieberding and Holveck: beyond experimental design and proximate mechanisms - mate choice in the face of sexual conflict

A reply to Nieberding and Holveck: beyond experimental design and proximate mechanisms - mate choice in the face of sexual conflict

We summarise our work on male mating behaviour in the tropical butterfly Bicyclus anynana , responding to the commentary provided by Nieberding and Holveck. We acknowledge that our laboratory studies are not free of shortcomings and potential caveats, though we attempted to address or highlight these within each paper. The concerns raised seem to stem mainly from different notions with respect to the proximate basis of old male mating advantage, and specifically the relative importance of male behaviour versus pheromone blend. In our view, our experiments provided compelling evidence for a prominent role of male behaviour, while we were unable to obtain clear evidence for a major role of male sexual pheromones. In addition to the lack of evidence we argue that a preference of females for older males based on pheromone blend is unlikely, as pheromone titres do not seem to indicate male quality and, more importantly, females actually suffer a fitness cost when mating with older males. The latter suggests that old male mating advantage arises from sexual conflict rather than cooperation. We thus highlight the importance of considering both the proximate and the ultimate level for gaining an integrative understanding of complex behavioural patterns.

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