Electric signalling and reproductive behaviour in a mormyrid fish, the bulldog Marcusenius macrolepidotus (South African form)

Bulldog fish (Marcusenius macrolepidotus) generate short (<1 ms) electric-organ discharges (EODs), separated by much longer and highly variable interdischarge intervals (IDIs). We observed overt behaviour and electrical activity during reproductive behaviour in a male and in a female bulldog, and identified IDI patterns with putative signal functions. In contrast to Pollimyrus adspersus and Pollimyrus isidori, in which an elaborate and extended courtship precedes spawning proper, our fish started spawning almost immediately when we allowed the female to enter the males territory. The male did not construct a nest, and neither parent provided parental care. The male showed very little aggression towards the intruding female. Fish spawned in bouts near the males hiding place, and eggs were scattered by the females vigorous tail flips as she left the spawning site, only to return shortly thereafter. During spawning bouts, both fish generated highly stereotyped IDI patterns: the male generated a series of IDIs gradually decreasing from about 200 ms to about 55 ms that was abruptly terminated by a long IDI. The female generated a series of relatively regular IDIs (about 54 ms) that was followed by a marked increase in IDI duration (the probable time of spawning). Finally, a sharp decrease in IDIs to about 20 ms accompanied the females sudden escape from the spawning site. In between spawning bouts, both fish generated series of very short IDIs (high discharge rate, HD) that alternated abruptly with very low-rate inter-HD activity (especially in the male). IDIs as short as 9 ms (male) or 11 ms (female) occurred during HD displays. No visible aggression, in fact very little overt behaviour, occurred during these HD displays in both fish. Agonistic interactions between male and female, outside a reproductive context, were similar to those previously described in male pairs, including overt behavioural patterns such as parallel swimming, antiparallel display and attack, as well as HD displays. When not interacting, fish did not generate HD displays. We suggest the HD display is a communication signal in both reproductive and agonistic contexts.     

The visual control of courtship behaviour in the fly Poecilobothrus nobilitatus

1. Poecilobothrns nobilitatus has a spectacular courtship in which the males perform a complex aerial display directed towards the females. This begins with short bouts of wing-waving, after which the male hovers in front of the female. This is followed by two types of display flight. In one the male circles in an arc around one side of the female, and in the second he flies over her, rotating through 180° in 40ms. The display flights are not modified once they have begun, and are thus pre-coordinated rather than feedback driven, as in tracking behaviour.
2. Courtship behaviour is confined to the males, and the extra tasks the male nervous system is required to perform are discussed in terms of the information needed for the execution of the behaviour.

     

Courtship in Drosophila quadrilineata with a unique male behavioral element, abdomen bending

This paper describes courtship of Drosophila quadrilineata of the immigrans group and reports a species-specific element, abdomen bending, that has not been described in Drosophila. The male bends his abdomen right and left as a display to the female. During courtship the male circles around the female, bends his abdomen, and taps her. The male licks the female ovipositor repeatedly, often after the females extrusion. Although the females ovipositor extrusion is thought to indicate rejection of a courting male in many species, the male of D. quadrilineata continues courtship. It is possible that the extrusion is a females mating signal to the courting male. The male of D. quadrilineata mounts the female in a rearward position and the genitalia of both sexes couple after mounting. This mounting position is shared by the species belonging to the immigrans group. We never observed behavior in which it appeared as if the male displayed the black stripes to the female. Males court one another and a courtship chain is frequently formed even in the absence of females. The males abdomen bending and the females extrusion followed by the males licking are discussed.     

Courtship and intersexual signaling in the parasitic waspCotesia rubecula (Hymenoptera: Braconidae)

The courtship behavior of the parasitic wasp Cotesia rubeculawas studied in a flight tunnel using standard quantitative ethological techniques. Emission of a female sex pheromone induces searching and signaling behavior in males. Males combine wing-fanning, which produces low-frequency airborne sound, with pulsing behavior, which transmits a vibrational signal through the substrate to the female and induces her receptivity. Female receptivity is indicated by a stereotyped antennal position, which may provide a visual or tactile signal to courting males. Comparison of successful and unsuccessful courtships indicated that courtship success was dependent primarily on the effective production or reception of the male pulse signal. Overall, the sequence of courtship behavior was similar to that reported for other parasitic wasps.     

Cues for male phonotaxis in the duetting bushcricketLeptophyes punctatissima

Summary A computer controlled setup is introduced which allows the song analysis of both male and femaleLeptophyes punctatissima during duetting in a laboratory situation. The essential acoustical parameters for the initiation of the male's phonotactic approach towards the stationary female are described. The female responds reflex-like to the male song after a remarkably short delay time of about 28 ms. The male only performs phonotaxis if he perceives the female reply above an intensity value of about 50 dB SPL and if the female response falls within a critical time window from 25 to a maximum of 55 ms after the onset of his song (Figs. 3 and 5). The sound intensity and overall time delay of the female response can be varied independently, so that the relationship between both parameters and their limitations for maximum phonotaxis distance can be described.     

Courtship inDrosophila sechellia: Its structure,functional aspects,and relationship to those of other members of theDrosophila melanogaster species subgroup

The courtship behavior of Drosophila sechelliais described. Male wing displays are mainly vibration and scissoring, with low levels of rowing. As courtship proceeds the proportion of courtship spent in male wing vibration and licking increases, whereas female movement decreases. The male courtship song of sechelliacontains pulse song but no sine song. This species also shows a distinctive copulation song associated with mounting and copulation. The main cuticular hydrocarbon in females is 7,11-heptacosadiene. The number of copulations increased when flies were placed in the presence of food. Visual and acoustic stimuli appear to be important for mating. A multidimensional comparison was used to compare members of the melanogaster species subgroup, based upon courtship behavior, song characteristics, and cuticular hydrocarbons. A multidimensional comparison of courtship sequences in sechellia, melanogaster, simulans,and mauritianashowed differences in variability between the two island species as compared to the two cosmopolitan species. The courtship song of D. orenais described: it shows both sine and pulse song; there is also a copulation song in this species.     

Zur Balz des Mikado-Fasans(Syrmaticus mikado) in Gefangenschaft

Zusammenfassung Drei Jahre lang wurde das Balzverhalten von 20 in Volieren gehaltenen Mikado-Fasanen (Syrmaticus mikado) beobachtet. Die Hauptbalzzeit war bei den ganzjährig im Freien gehaltenen Fasanen in Mainz (50° N.) März-April. Die Balz der bestand nicht wie bisher angegeben im wesentlichen aus einer Lateralpräsentation. Zwar wurde eine solche auch im Rahmen des Balzverhaltens gezeigt, doch stellte sie keine eigentliche Balzleistung dar. Sie trat vielmehr während des ganzen Jahres hindurch mehr oder weniger auf, und zwar bei beiden Geschlechtern. Ein wesentliches Element der Balz war dagegen die Frontalpräsentation, die hier erstmals beschrieben wird. Sie trat nur während der Balz auf und wurde nur von gezeigt. Die beantworteten sie nie mit Flucht. Die ins Balzverhalten eingeschaltete Futtervermittlung durch das wurde bisher nur in echter, nie in symbolischer Form beobachtet. Dabei hielten die das Futter nie im Schnabel, um es den gezielt anzubieten.

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The courtship behaviour of the Mikado Pheasant (Syrmaticus mikado) in captivity

Summary The courtship behaviour of 20 Mikado pheasants (Syrmaticus mikado) was observed over a period of three years. The birds were kept in aviaries in the open the whole year round. In Mainz (50° N), the main mating period occurred in March and April. The courtship of the cocks did not consist essentially in lateral display, as has been previously reported for species of the genusSyrmaticus. Although lateral display did take place during courtship, it was not really part of the cock's courtship behaviour. It was rather the case that lateral display occurs more or less during the whole year and in both sexes. Frontal display was, however, an essential element in the Mikado pheasant's courtship and is described here for the first time. It occurred only during the mating period and was performed exclusively by the cocks. The hens never responded to frontal display by fleeing. The provision of food, which the cocks interpolate into courtship, has been observed so far only in the genuine and never in the symbolic form. It could never be observed that the cocks held the food in their beaks in order to offer it to the hens directly.

     

The structure and function of songs emitted by southern green stink bugs from Brazil, Florida, Italy and Slovenia

Nezara viridula (L.) (Pentatomidae: Heteroptera) from Brazil, Florida, Italy and Slovenia, communicate by vibratory songs associated with long‐range calling and close‐range courting, rivalry and repelling. Each song is composed of spectrally and temporally different units. Spectrally different pulses of duration less than 300 ms are present in the male calling song. The female calling song is characterized by pulse trains composed of pulses shorter than 150 ms and pulse trains composed of a longer (> 700 ms) and shorter (< 250 ms) pulse. Shorter and longer pulses have different spectral characteristics. The male and female courtship songs are characterized by fusion of shorter (< 150 ms) pulses into a pulse train usually followed by a shorter (< 200 ms) postpulse in the case of the male courtship song. The female repelling song is a several seconds long vibration of irregular temporal structure. The short (< 400 ms) male rival song pulses are frequency modulated. The dominant frequency peaks of the songs investigated lie between 70 and 130 Hz. The dominant frequency and the microstructure of song spectra show no population specificity. The average duration varies more in calling than in courtship songs. The repetition time varies extensively in songs of different populations. Normal communication followed by copulation was observed between mates from Slovenia and Brazil and between mates from Florida and Italy. The potential role of different temporal and spectral parameters for species recognition and mate location is discussed in view of the expected distortion of the characteristic signal structure during transmission through plants.     

The extent of visual control in the courtship tracking of D. melanogaster

The courtship tracking behaviour of male D. melanogaster was filmed in a range of visual mutants and in wild type flies under white and infra-red light. The absence of visual input severely restricts the velocity at which the female may be tracked but does not remove the male's ability to discriminate the female's abdomen from her head. Degradation of visual processing, such as occurs in certain visual mutants, leads mainly to disruption of the control of orientation in tracking. The gain control of translation may only be meaningfully assessed when vision is enabled. Courtship tracking does not depend upon the presence of a binocular field of vision; one eyed flies court with a lateral displacement of the body, and loose visual contact with the female readily when her image moves from the sighted towards the blind eye. The selection of the left or right wing for extension follows a distinct pattern with the male's position and orientation around the female; this may be controlled collateral to locomotor output and after interlateral comparison of sensory input.     

A Glandular Gift in the German Cockroach, Blattella germanica (L.) (Dictyoptera: Blattellidae): The Courtship Feeding of a Female on Secretions from Male Tergal Glands

In the courtship behavior of the German cockroach, the male presents tergal glands to the female and feeds her with glandular secretions to place her in the appropriate precopulatory position. The phagostimulant activity of the secretions was quantitatively examined using the polyethylene glycol film method. The methanol extract of the glands on the eighth tergite induced a potent feeding response in 6-day-old virgin females (EC ₅₀ = 0.0037 male equivalent/40 g PEG spot). However, there was no temporal relation between the feeding response and the sexual receptivity of the females. Moreover, besides virgin females, the extract induced a feeding response in gravid or mated females, males, and the last-instar nymphs. These results strongly suggest that the secretions function as a dietary feeding stimulant in principle but as a courtship pheromone in the context of courtship behavior where the stimulants are offered as a nuptial gift.     

Female responses to isolated signals from multimodal male courtship displays in the wolf spider genus Schizocosa (Araneae: Lycosidae)

Male wolf spiders within the genus Schizocosa display considerable variation in foreleg ornamentation as well as in courtship communication. Multiple modes of male signalling have evolved in a number of species. Divergence in courtship signals among species within this genus may be directly associated with variation in the sensory sensitivities of conspecific females. We isolated the visual and vibratory courtship cues of four species of Schizocosa and recorded conspecific female receptivity to each isolated cue. We also examined female receptivity to complete multimodal courtship signals. We found that the sensory sensitivities of conspecific females were associated with the predominant modes of male courtship communication. Species in which females use mostly stridulatory cues in assessing conspecific males tended to have stridulation-based male courtship displays (S. duplex and S. uetzi) while the opposite was true for species in which females used more visual cues in male assessment (S. stridulans and S. crassipes). This study suggests coevolution between male signal design and female sensory design. We discuss possible scenarios that could be driving this coevolution, including hypotheses of sensory bias and environmental constraints. Copyright 1999 The Association for the Study of Animal Behaviour.     

Pheromonal inhibition of male courtship behaviour in the brown tree snake, Boiga irregularis: a mechanism for the rejection of potential mates

Pheromones play a central role in coordinating the events leading up to copulation in snakes. We report here a novel pheromone system in the brown tree snake in which females release a pheromone that inhibits male courtship behaviour. In a previous study, we made observations of female brown tree snakes releasing cloacal secretions (CS) during courtship that appeared to cause courting males to cease courtship. All snakes have glands that release CS through ducts located along the cloacal orifice. Although CS have been studied for many years, their function in the mediation of snake behaviour has not been experimentally well determined. We examined the role of CS in the reproductive behaviour of male and female brown tree snakes. We conducted four experiments to test the effect of both male and female CS on brown tree snake behaviour under two behavioural contexts, courtship and male-male ritualized combat. Within each experiment, we compared the effects of CS to a control. Female CS caused a decrease in the time that males spent courting females and a decrease in the intensity of courtship compared with the control treatment. Male CS did not, however, affect the time that males spent displaying courtship or the intensity of that courtship. Neither male nor female CS had significant effects on male ritualized combat behaviour, including time that males spent in combat or the intensity of combat behaviours displayed. Furthermore, neither female nor male CS had an effect on female courtship versus controls. The inhibition of brown tree snake reproductive behaviours is specific to female CS inhibiting male courtship behaviour. This pheromone acts in concert with the female sex pheromone to regulate the events leading to copulation.Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

The importance of calling song and courtship song in female mate choice in the variable field cricket

Male field crickets produce calling songs, courtship songs, tactile signals and chemical signals. Although calling songs are known to play an important role in female mate choice, the importance of the other signals in mate choice is poorly understood. In the variable field cricket, Gryllus lineaticeps, females select mates, in part, based on variation in male calling song. Females prefer higher chirp rates, a trait which is partially dependent on male nutrient intake, and females prefer longer chirp durations, a trait which appears to be independent of male nutrient intake. We tested whether females also have preferences based on variation in male courtship song, and whether the structure of male courtship song varies with nutrient intake. First, we reexamined female preference for calling song chirp rate. Then, we examined: (1) female preference based on courtship song chirp rate; (2) the relative importance of calling song and courtship song chirp rate; (3) the nutrition dependence of courtship song chirp rate; and (4) the correlation between calling song and courtship song chirp rate. As reported previously, females preferred higher calling song chirp rates, and in addition, preferred higher courtship song chirp rates. Females were more likely to switch from a speaker broadcasting more attractive calling song to a speaker broadcasting less attractive calling song when the attractive calling song was associated with an unattractive courtship song than when it was associated with an attractive courtship song. Preferences based on courtship song may thus cause females to alter the choices that they made based on calling song. Males that received greater nutrients did not produce higher courtship song chirp rates. There was no correlation between calling song and courtship song chirp rate. As a result, the two traits may provide information to females about different aspects of male quality. Copyright 2000 The Association for the Study of Animal Behaviour.     

Triggers of the Postural Display of Courtship in <Emphasis Type="Italic">Drosophila persimilis</Emphasis> Flies

D. persimilis courtship shows some flexibility and courting males sometimes perform an elaborate postural display in addition to the standard courtship behaviours shared by most Drosophila species. This postural display includes the acrobatic contortion and tremulation of their abdomen, accompanied by the generation of substrate-borne vibrations, and they proffer a nutritional droplet to the female. Here, we use courtship and choice assays to ask what triggers this display and what advantages males may gain from it during courtship. In pair assays, we found no differences in the courtship duration and copulation success between displaying and non-displaying males. In trio assays, however, the female always mated with the male who performed the display. To investigate what promotes the male display, we varied the level of receptivity of the female and studied the impact of a second male. We found that rejection by the female does not induce the male to display, contrary to what was previously suggested. We present evidence that the male display is in fact promoted by the presence of an attentive and sexually receptive female and the absence of male competition, with the greatest exhibition rate obtained if the courted female is starved. These findings provide valuable information about the social ecology of flies, and how internal and external cues influence sexual behaviours and mate choice.     

Ritual behaviors associated with spermatophore transfer in Deuterosminthurus bicinctus (Collembola: Bourletiellidae)

The springtail Deuterosminthurus bicinctus, similarly to other members of Bourletiellidae, use their antennae, legs or heads to monopolize, stimulate, and direct female partners to spermatophores. The mating behavior of this species was examined by analysis of video recordings made on the leaves of its host plant. The characteristic stages of the behavioral sequence leading to sperm transfer were (1) preliminary courtship, highly variable in time and intensity—a male (rarely a female) endeavors to recruit a partner for further courtship, (2) push-and-retreat ritual—a rigid and rhythmical head-to-head dance of both partners, composed of some 180 phases and interspersed by several (3) free turns of a male and ended by (4) spermatophore deposition in front of a female, followed by stimulation of a female by a male to walk over the spermatophore, (5) spermatophore pick-up by a female with her gonopore, or alternately, (6) total spermatophore consumption by a female, as happened in 29% of the observed pairs. The mating ended always with (7) competition by both partners for the consumption of spermatophore residuals, usually won by the female. Comparison of mating elements and morphological features of partners engaged in mating sessions that ended either with spermatophore pick-up or consumption did not reveal any substantial differences. This suggests that changes in female motivation (hunger, state of impregnation) may be crucial for the male success in sperm transfer to a female. A digital video image is available at An erratum to this article can be found at     

Single gene mutations in Drosophila: what can they tell us about the evolution of sexual behaviour?

The molecular analysis of specific mutant genes that affect the courtship behaviours of Drosophila melanogaster males and females is discussed in the light of the possibility that they may contribute to mate choice. There is clear evidence that some genes can act as a reservoir of species-specific behaviour, particularly for the male actions during courtship. However, to date there has not been a single genetic locus that has been isolated at the molecular level and shown to be associated with a change in female preference. There are some promising avenues of exploration, in that recent genetic analyses suggest that a small number of genes may make major contributions to female preferences. Finally a candidate gene approach is advocated in which orthologous genes from other species of Drosophila are used as natural mutations, and transformed into D. melanogaster hosts to investigate whether they carry species-specific mating information of the donor.     

Escape response of the rotifer Polyarthra: a high-speed cinematographic analysis

Summary Cinefilms of unconstrained P. vulgaris at 17°C were taken at a low magnification (2x) and 120–200 fps to analyze body movements during swimming and escape responses mediated by movements of the 12 lateral, bladelike appendages or paddles. Cinefilms of partially constrained P. vulgaris and P. dolichoptera at 16°C were taken at a higher magnification (10x) and 300 fps, using interference contrast optics, to resolve paddle movements during escape responses. When swimming, P. vulgaris moved at a velocity of 0.348±0,025 (S.E.) mm·s^-1 (2.64 body lenghs·s^-1), having a Reynolds number of 0.05. During escape responses, P. vulgaris traveled 1.947±0.124 (S.E.) mm (15 body lengths) during 0.0564±0.0038 (S.E.) s, continuously moving at a velocity of 35.7±1.2 (S.E.) mm·s^-1 (270 body lengths·s^-1) and having a Reynolds number of 5. During these responses, P. vulgaris tumbled sinuously but mostly-88.9±2.3 (S.E.) %-in a constant direction; the angular change in direction from one frame to the next was 28±2 (S.E.) degrees, but the sign of the change in direction frequently alternated. Escape responses are caused by 1–3 cycles of paddle movements. In each cycle, the rigid paddles move up asynchronously until they are all directly overhead, and then they move downwards to their original resting positions, again asynchronously. Polyarthra's body moves along the flight path during all phases of this cycle. A single cycle may take as little as 26 ms, 13 ms for the paddles to elevate and 13 ms for them to descend. The asynchronous upward and downward movements of each of the 12 paddles explain why Polyarthra's body tumbles continuously through its low Reynolds number, viscous environment. Escape responses generally were initiated by contact with another rotifer. In one P. dolichoptera response, the time lag between such contact and the initiation of paddle elevation was about 7 ms. The very short lag time, great velocity, considerable displacement, and unpredictable directionality of Polyarthra's escape response make it a very effective defense against capture by some invertebrate predators.     

Androgen-induced alterations in vocalizations of femaleXenopus laevis: modifiability and constraints

Summary We examined effects of exogenous androgen (testosterone and dihydrotestosterone) on vocalizations of ovariectomized, adult female South African clawed frogs,Xenopus laevis. When paired with sexually active males, all ovariectomized females exhibited ticking, the unreceptive or release call. Ticking consists of low amplitude, regularly spaced clicks with a mean interclick interval of 154 ms. When androgen-treated and paired with sexually active males, these ovariectomized females also exhibited an aberrant call (atypical ticking) in which click multiples replaced the single clicks of ticking. Mean ICI's for atypical ticking were 37 ms for click doublets and 22 ms for click quadruplets. Androgen treatment decreased the total time spent vocalizing (typical and atypical ticking) by ovariectomized females.All androgen-treated females were then tested repeatedly with sexually receptive females in an attempt to elicit the male-typical vocalization, mate calling. Six of 17 females did not vocalize at all, even when gonadotropin injected. Eight females gave rapid (mean ICI, 36 ms) trains of clicks in an irregular temporal pattern (tick-like calls). Three females gave brief trills with alternating fast and slow components. Comparison of mate calllike vocalizations of androgen-treated females to mate calling of males reveals that calls in females are considerably shorter in duration (female: 0.32 min versus male: 45 min) and slower in tempo (ICI's; fast trill, female: 21 ms, male: 14 ms; slow trill, female: 36 ms, male: 28 ms). Incomplete masculinization of the vocal pattern of females by androgen treatment in adulthood may be due to developmental constraints on the modifiability of the neurons and muscles responsible for calling.Abbreviations C cholesterol - DHT dihydrotestosterone - HCG human chorionic gonadotropin - IBI interburst interval - ICI interclick interval - ovx ovariectomized - T testosterone