The response of female Mountain White-crowned Sparrows to songs from their natal dialect and an alien dialect

Summary Female Mountain White-crowned Sparrows from Colorado were tested in a laboratory playback experiment to determine their response to male songs from their own natal dialect and to those from an alien dialect of the same subspecies, Zonotrichia leucophrys oriantha (Fig. 1). Observations were made on locomotor activity and copulation postures produced during each experimental test session.The subjects gave copulation displays almost exclusively when hearing their natal dialect and new or no displays when hearing the alien dialect (Fig. 2). The birds also exhibited significantly greater locomotor activity when hearing the natal dialect than they did when hearing the alien dialect (Fig. 3).We conclude that females are sensitized by their early song learning experience to be responsive to songs from their natal dialect and virtually sexually unresponsive to songs from an alien dialect. We speculate that the copulation posture given in the experiment implies that females in natural populations would preferably only mate assortatively with males from their natal dialect region.     

Population biology of White-crowned Sparrows: Residence time and local movements of juveniles

Summary A field study of the ranging behavior of post-fledging White-crowned Sparrows (Zonotrichia leucophrys nuttalli) was carried out in undisturbed natural habitat in the Point Reyes National Seashore, Marin Co., California, USA. Forty-three first-brood juveniles, with tail feathers still growing, were banded at an estimated average age of 27 days and re-trapped an average of an additional 24 days for a total minimum time in residence in the immediate natal area of 51 days (Table 1).The distance from the site of first capture to subsequent recapture sites was about 60 m until age 35 days; then the distance increased to about 250 m by age 50 days (Fig. 2). These data indicate that the average juvenile White-crowned Sparrow completes at least the first 50 days of life in its natal area. Experimental work on song ontogeny has previously demonstrated that the sensitive period for song learning ends at about 50 days. Hence, young in our study population probably learn their natal dialect prior to any major dispersal movements. The correspondence between residence time and song learning would have the consequence of perpetuating song dialects as geographically structured populations.     

Female response to song reflects male developmental history in swamp sparrows

One explanation for why female songbirds attend to male song is that the quality of a male's song is associated with the quality of his developmental history. We tested this hypothesis by playing back to female swamp sparrows (Melospiza georgiana) songs recorded from males of either inferior or superior developmental histories, as assessed by their rates of mass gain during the first 18 days post-hatching. Females showed significantly higher levels of courtship display in response to songs of males with superior growth than to songs of males with inferior growth. Out of nine song traits measured, only song duration correlated with variation in female response; duration was also the only trait that differed significantly in univariate comparisons between the superior growth songs and the inferior growth songs. In a multivariate analysis, however, inferior growth songs were best discriminated from superior growth songs by combining three song traits: trill rate, stereotypy, and the number of notes per syllable. We suggest that early developmental stress degrades song in many small ways, and that it is the cumulative effect of the resulting deficits that explains lower female response.     

Response of territorial male song sparrows to song types and variations

Summary The song repertoires of male song sparrows (Melospiza melodia) are comprised of distinct song types and many overlapping variations within each type. This study examines the sensitivity of territorial male song sparrows to both multiple song types and variation within types. A playback experiment was conducted using stimulus tapes that had either single songs (but not variations), song variations (but not types) or a single song. Response during playback was similar for all three stimulus groups, but diminished sooner after playback for the single-song tape. Thus our results indicate that male song sparrows are sensitive to song variability both within and among song types.     

Song as an attractant for male and female European starlings,and the influence of song complexity on their response

Summary We tested the hypothesis that song attracts females and repels males in the European starling. We broadcast recorded song from speakers attached to nestboxes, while paired boxes with silent speakers served as controls. As predicted, females were attracted to the song boxes. Contrary to prediction, males were also attracted to the `song' boxes. Singing by male starlings may be costly because it attracts competitors for limited nesting sites, but the cost cannot be avoided due to the need to attract a mate. In a second experiment simple song (composed of 20 different phrase types) and complex song (40 phrase types) were played simultaneously. More males were captured at boxes where simple song was played. Song complexity may function as an indicator of male quality and be used by male starlings to assess potential competitors.     

Singing onstage: female and male common nightingales eavesdrop on song type matching

In bird communication, listening individuals may obtain information on the quality and motivation of a male not only from solo-singing, but also from song interactions and listeners base their future decisions in territorial and mating contexts on such public information. Eavesdropping on male interactions may thus have a strong influence on sexual selection. In singing interactions, temporal coordination (e.g. overlapping vs. alternating) of two singers as well as structural interaction patterns (e.g. song type matching or repertoire matching) have been described, but the latter is far less studied. By conducting dual-speaker playback experiments with common nightingales Luscinia megarhynchos, we simulated an interaction where one singer was repeatedly song–type matching his counterpart. Playbacks were broadcast to male and female nightingales, and their approach behaviour and singing responses (in the case of male focals) were analysed. We found that both, males and females, spent more time with the matched bird, whereas males additionally sang more songs towards the matching bird. This can be taken as strong hint that eavesdropping occurs in nightingale communication and that listening to male vocal contests might be an important strategy for both sexes to adjust their behavioural output. With regard to the function of song matching, we assume that song-matching is not an aggressive signal per se in nightingales. We rather conclude that vocal leaders within an interaction, here the matched bird, may elicit stronger responses in conspecifics than vocal followers, here the matching bird.     

Reproductive value and appropriateness of response to predators by White-Crowned Sparrows

Summary Twenty-five pair of breeding Whitecrowned Sparrows were presented with a live snake and models of a hawk, jay, and junco. Pairs were tested either when the female was brooding eggs, feeding nestlings, or fledglings. It was found that the snake was responded to the most when the pair had nestlings, very little when the female was brooding eggs, and at a moderate level with fledglings. The hawk and jay models were responded to the least with eggs, more with nestlings, and the most with fledglings. The junco model elicited little response. Both the pattern and level of response is influenced by whether or not the stimulus represents an effective predator at the particular stage of the breeding cycle. They also are influenced by the reproductive value of the progeny. A model is suggested that includes two factors: stimulus value and reproductive value. We propose that this combined model is more suitable than a single factor one.     

Soft song in song sparrows: response of males and females to an enigmatic signal

Low-amplitude “soft song” is used by a variety of songbirds; in some species during aggressive encounters, in others during courtship, and yet others in both these contexts. In song sparrows (Melospiza melodia), soft song has thus far been observed only in aggressive encounters, where its production is a more reliable predictor of attack than any other signaling behavior. We used song playback to test the response of both male and female song sparrows to soft song. The design of the playback experiments took into account the existence of two classes of soft song: crystallized soft song, which consists of song types also found in the broadcast repertoire, and warbled soft song, which consists of less-structured song types not found in the broadcast repertoire. Female song sparrows responded with significantly less courtship display to the playback of crystallized soft song than to that of normal broadcast song, and response to warbled soft song was if anything lower than to that of crystallized soft song. Male song sparrows responded equally aggressively to normal broadcast song as to crystallized soft song, and equally aggressively to warbled soft song as to crystallized soft song. The female results support the conclusion that neither form of soft song functions in courtship. The male results suggest that the reliability of soft song as a signal of aggressive intent is not maintained by a receiver retaliation rule.     

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

Female and male serins (Serinus serinus) respond differently to derived song traits

We tested if male or female behavior towards manipulated song indicates intra- or inter-sexual selection of two characteristics of serin song that are extreme and evolutionarily derived in this species: high frequency and fast syllable rate. In a first experiment, we monitored vocal responses and attendance to song playbacks. Female behavior indicated a preference for high-frequency song and suggested an aggressive function for fast syllable rates, as fast songs inhibited vocal response. Males did not show discrimination of frequency or syllable rate with this experimental design. The second experiment used a simple approach/no approach design, and in this experiment, males showed stronger discrimination between stimuli than did females. Therefore, sex differences in discrimination appear not to result from differences in perceptual abilities but from differences in the context of stimulus presentation. The second experiment also supported a role of song frequency in female choice, as the effect of frequency was limited to females: males did not respond differently to song frequency and approached high-frequency songs less than females did. Results of this experiment also supported an aggressive function for fast syllable rates, as the effect of fast songs did extend to male behavior. Taken together, our results indicate that the high frequency and fast syllable rate of serin song cannot result from a single selection process: while high frequency may have evolved by inter-sexual selection, syllable rate provokes a pattern of response that is more consistent with intra-sexual selection.     

Altered responses to female odors in parasitized male mice: neuromodulatory mechanisms and relations to female choice

There is accumulating evidence that females may preferentially select parasite-free or -resistant males. Minimal attention has, however, been paid to the mate preferences and responses of the parasitized male hosts themselves. Here, we considered the effects of parasitic infection on male host mate responses, the neuromodulatory correlates of these responses, and the relations of these responses to female mate choice. Using an odor “preference” test, we examined the effects of different stages of an acute, sub-clinical infection with the naturally occurring, enteric, single host, protozoan parasite, Eimeria vermiformis, on the responses of male mice, Mus musculus domesticus, to the odors of estrous females along with the responses of uninfected females to the parasitized males. At 4 days post-infection (non-infective, pre-patent stage) E. vermiformis-infected male mice showed a significantly decreased preference for the odors of estrous females, whereas at 10 days post-infection (infective, patent stage) infected males showed a significantly increased preference for the odors of estrous females. Parasitized males displayed no significant changes in their responses to the odors of non-estrous females, supporting effects on the reproductively related responses of the host. In parallel, estrous females displayed a reduced interest in the odors of infected males. Least interest was expressed in the odors of the patent, infective males, consistent with the avoidance of contagion. Using selective opioid peptide receptor agonists and antagonists we found evidence that enhanced kappa opioid peptide (e.g., dynorphin) activity was related to the decreased sexual interest of the pre-infective males, while augmented delta opioid peptide (e.g., enkephalin) activity was associated with the enhanced responses of the infective males to females. We further showed that acute kappa opiate administration reduced the responses of uninfected males to females and that uninfected females displayed modified responses to the odors of uninfected males subject to acute modifications of opioid activity. We suggest that these differential shifts in endogenous opioid activity in the parasitized males are associated with and, or related to alterations in neuro-immune and endocrine functions. These findings show that parasitic infection can have, depending on the stage of infection and associated neuromodulatory changes, either significant facilitatory or inhibitory effects on male host preferences for and responses to females. Received: 22 April 1996 / Accepted after revision: 15 March 1997     

Female mate choice in the zebra finch — the effect of male beak colour and male song

Studies of female mate preference in zebra finches (Taenopygia guttata) have shown that male beak colour and song rate are important. However, the two characters are correlated. Here the effect of beak colour and song rate on female choice are examined independently. In mate choice tests involving two males, beak colour was manipulated artificially using nail varnish. The results showed that females showed a significant preference for males with a high song rate, but not with a red beak. Females did not prefer males with a red beak if song rate was low and females preferred males with orange beaks who expressed a high song rate. Female preference for males with red beaks was not found when beak and song characters were no longer correlated.     

Use of the song repertoire in intersexual and intrasexual contexts by male red-winged blackbirds

Summary Live and stuffed male and female red-winged blackbirds (Agelaius phoeniceus) were presented to territorial males in order to study the repertoire organization of males during intersexual and intrasexual encounters. Territorial male red-winged blackbirds switched song types more frequently and sang more song types in response to presentation of a female stimulus than during control periods, but decreased their switching frequency when a male stimulus was presented. Switching frequency in response to the female stimulus was three times that in response to the male stimulus. These results support the hypothesis that song repertoires of male red-winged blackbirds evolved primarily in response to intersexual rather than intrasexual selection. They also suggest that male red-winged blackbirds have been selected to de-emphasize their song repertoires during encounters with conspecific males as a result of some as yet unidentified cost. Offprint requests to: W.A. Searcy     

Acoustic communication in phaneropterid bushcrickets: species-specific delay of female stridulatory response and matching male sensory time window

Summary The stridulatory sounds and movements produced by the females of various bushcricket species (Tettigonoidea: Phaneropteridae: Barbitistini) are compared with those of the males. Behavioral experiments are carried out to determine the significance of the female sounds in acoustic communication. Selection factors involved in the evolution of female stridulation are discussed. The morphological apparatus for sound production has evolved independently in males and females. Whereas males rub a toothed file on the underside of the left wing over the inner edge of the right wing, the plectrum, the females stridulate by rubbing a thickened vein on the underside of the left wing over modified spines on the upper surface of the right wing (Fig. 2). Similarly, the movements responsible for sound production are not homologous in males and females. In the male the audible closing movement is always preceded by wing opening, whereas the female in general initiates the closing movement when the wings are in the resting slightly opened position, and abruptly produces complete closure (Fig. 3, 4 and 5). The female responds to male singing by emitting one to several highly damped sound pulses each lasting less than one ms. The interval between the song of a conspecific male and this response is a very precise species-specific characteristic (Fig. 7). In species with male songs that are complicated in structure or continuous, the females respond only at specific time-after patticular markers in the song of the male. The time interval between male song and female response is an important criterion by which the male identifies conspecific female song (Fig. 8). Because the response delay of the female and corresponding neuronal time window in the male are distinct they may be important in species discrimination.     

A female evolutionary response when survival is at risk: male harassment mediates early reallocation of resources to increase egg number and size

One unexplored area in sexual conflict studies is the female physiological costs and possible resource reallocation that accompany evolutionary costs due to male harassment. Using females of the damselfly Hetaerina americana, I first investigated whether male harassment affected female mating rate and survival and explored whether such effects induced a resource allocation from immunity (in the form of phenoloxidase activity) and muscular fat reserves to egg number and size. Using two seasons that differed in male harassment, it was found that the higher the male harassment, the fewer are the female matings and the lower is the female survival. These results were corroborated using an experimental approach in which a situation of high male harassment was induced. It was also found that when the first mating takes place and at high male harassment, females had more reduced phenoloxidase activity and fat reserves and tended to lay most of the eggs they produce in their lifetime and these were considerably large. However, at low male harassment, egg number and size were more equally produced across matings. Females under high male harassment seemed to suffer the survival costs but may show a plastic evolutionary response of reallocating resources to egg traits to maximize fitness.     

Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the solo performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.Communicated by I. Hartley     

The effect of male calling on female locomotor activity of Teleogryllus commodus

Summary The calling song of the -cricket, Teleogryllus commodus, attracts for mating. Both calling and -locomotor activity are under circadian control and the behaviors occur approximately at the same time. Daily playback of calling songs for 3–12 h in constant light failed to entrain -running. Instead, calling acts as an external stimulus which can release -locomotion at any time of the 24-h period, while the clock-induced activity continues to run freely. The stimulatory effect of the calling song causes flexible extension of -locomotor activity, but a subsequent restorative period is necessary. The ecological consequences of acoustical stimulation are discussed.     

The scent of dominance: female field crickets use odour to predict the outcome of male competition

Pheromones may convey information about mate quality and social status. In the field cricket Gryllus integer, females mount the males for copulation, such that males cannot coerce females to mate. We examined whether virgin G. integer females preferred the scent of potentially dominant males to that of subordinate males. First, we collected pheromones by confining males on filter paper. Next, we offered filter paper from each of two size-matched males and control paper to females that had never been exposed to males, and measured the time spent by the female on each kind of paper. Finally, dominance status of the males in each size-matched pair was determined by pitting the two males against one another in agonistic contests. When offered filter paper from subsequently dominant versus subsequently subordinate males, females spent more time on the paper from the dominant male than the subordinate male, and much less time on control paper. Thus, pheromones may inform female G. integer about a male's potential to achieve dominant social status. Male pheromones were also associated with the female's tendency to mount a male. In contrast to cockroaches, where females prefer the scent of subordinate males (presumably to avoid risk of injury), female crickets prefer the scent of potentially dominant males and are more likely than males to wound their mating partners.