Philip K. Stoddard

Recognition of individual neighbors by song in the song sparrow, a species with song repertoires

SummaryPrevious theory and research have suggested that bird species with song repertoires in general, and song sparrows (Melospiza melodia) in particular, cannot readily discriminate between the songs of neighbors and strangers. In a recent study (Stoddard et al. 1991) we showed that song sparrows can in fact discriminate neighbors from strangers on the basis of song. In this study we sought to demonstrate that song sparrows can make the finer discrimination between individual neighbors and that they can do so on the basis of a single song type. We compared the response of territorial males to song playback of neighbors and strangers at three locations: the neighbors regular boundary, the opposite boundary, and the center of the territory. The birds showed strong neighbor-stranger discrimination at the regular boundary but not at the opposite boundary, nor in the center of the territory. The differences in song discrimination between different boundary locations indicate that song sparrows associate particular songs with particular territories, effectively discriminating between individual neighbors on the basis of song. Song repertoires themselves do not interfere with neighbor recognition to the extent originally postulated. As speakers are moved inside the territory from the border, however, the degree of discrimination diminishes. We believe that differences in speaker placement may have contributed to the variability in neighbor-stranger discrimination observed in previous studies of the song sparrow and perhaps other repertoire species as well. This interpretation is consistent with data from another song sparrow population showing that half the territory takeovers are by immediate neighbors.

Predation enhances complexity in the evolution of electric fish signals

Theories of sexual selection assume that predation is a restrictive, simplifying force in the evolution of animal display characters and many empirical studies have shown that predation opposes excessive elaboration of sexually selected traits. In an unexpected turnaround, I show here that predation pressure on neotropical, weakly electric fish (order Gymnotiformes) seems to have selected for greater signal complexity, by favouring characters that have enabled further signal elaboration by sexual selection. Most gymnotiform fish demonstrate adaptations that lower detectability of their electrolocation/communication signals by key predators. A second wave phase added to the ancestral monophasic signal shifts the emitted spectrum above the most sensitive frequencies of electroreceptive predators. By using playback trials with the predatory electric eel (Electrophorus electricus), I show that these biphasic signals are less detectable than the primitive monophasic signals. But sexually mature males of many species in the family Hypopomidae extend the duration of the second phase of their electric signal pulses and further amplify this sexual dimorphism nightly during the peak hours of reproduction. Thus a signal element that evolved for crypsis has itself been modified by sexual selection.

Song repertoire size predicts initial mating success in male song sparrows, Melospiza melodia

Male song sparrows sing repertoires of 4–13 distinct song types and have proved a valuable model for testing hypotheses concerning the function and evolution of song complexity. Captive female song sparrows solicit more copulations in response to playback of larger repertoires, yet it remains unclear whether male repertoire size influences female mate choice in natural situations. We used long-term data from free-living song sparrows inhabiting Mandarte Island, British Columbia, Canada, to investigate whether male song repertoire size predicted three components of reproductive performance during the first year: territory acquisition, mating success and laying date. Across males whose song was recorded, males with larger repertoires were not more likely to acquire a territory, to acquire a larger territory or to settle sooner. However, after we controlled for territory size and between-year variation in the population sex ratio, first-year males with larger repertoires were more likely to mate. This was because they were more likely to pair with newly settled females, not because they were more likely to acquire territories where older females were already resident. After we controlled for territory size and between-year variation in breeding date, newly settled females laid earlier when mated with males with larger repertoires. Together with the results of previous mate choice experiments, these patterns are consistent with the hypothesis that male song repertoire size is a sexually selected trait that influences female mate choice in song sparrows.

PLASTICITY OF THE ELECTRIC ORGAN DISCHARGE WAVEFORM OF THE ELECTRIC FISH BRACHYHYPOPOMUS PINNICAUDATUS I. QUANTIFICATION OF DAY-NIGHT CHANGES

Abstract The electric organ discharge of the gymnotiform fish Brachyhypopomus pinnicaudatus is a biphasic waveform. The females electric organ discharge is nearly symmetric but males produce a longer second phase than first phase. In this study, infrared-sensitive video cameras monitored the position of unrestrained fish, facilitating precise measurement of electric organ discharge duration and amplitude every 2 h for 24 h. Males (n=27) increased electric organ discharge duration by 37 ± 12% and amplitude by 24 ± 9% at night and decreased it during the day. In contrast, females (n=8) exhibited only minor electric organ discharge variation over time. Most of a males increase occurred rapidly within the first 2–3 h of darkness. Electric organ discharge values gradually diminished during the second half of the dark period and into the next morning. Modulation of the second phase of the biphasic electric organ discharge produced most of the duration change in males, but both phases changed amplitude by similar amounts. Turning the lights off at mid-day triggered an immediate increase in electric organ discharge, suggesting modification of existing ion channels in the electric organ, rather than altered genomic expression. Exaggeration of electric organ discharge sex differences implies a social function. Daily reduction of duration and amplitude may reduce predation risk or energy expenditure.

Fitness Correlates Of Song Repertoire Size In Free-Living Song Sparrows (Melospiza Melodia)

Models of sexual selection propose that exaggerated secondary sexual ornaments indicate a male’s own fitness and the fitness of his offspring. These hypotheses have rarely been thoroughly tested in free‐living individuals because overall fitness, as opposed to fitness components, is difficult to measure. We used 20 years of data from song sparrows (Melospiza melodia) inhabiting Mandarte Island, British Columbia, Canada, to test whether a male’s song repertoire size, a secondary sexual trait, predicted overall measures of male or offspring fitness. Males with larger song repertoires contributed more independent and recruited offspring, and independent and recruited grandoffspring, to Mandarte’s population. This was because these males lived longer and reared a greater proportion of hatched chicks to independence from parental care, not because females mated to males with larger repertoires laid or hatched more eggs. Furthermore, independent offspring of males with larger repertoires were more likely to recruit and then to leave more grandoffspring than were offspring of males with small repertoires. Although we cannot distinguish whether observed fitness variation reflected genetic or environmental effects on males or their offspring, these data suggest that female song sparrows would gain immediate and intergenerational fitness benefits by pairing with males with large song repertoires.

Plasticity of the electric organ discharge waveform of male Brachyhypopomus pinnicaudatus. II. Social effects.

Abstract. Many electric fish produce sexually dimorphic electric organ discharges. Although electric organ discharges are comprised of action potentials, those of the Gymnotiform family Hypopomidae show significant plasticity in response to stress and time of day. We show here that male Brachyhypopomus pinnicaudatus (Hopkins 1991), adjusts the degree of sexual dimorphism in its electric organ discharge depending on immediate social conditions. Three to five days of isolation resulted in gradual decrease of two sexually dimorphic waveform characters: duration and amplitude. Introduction of a second fish to the experimental tank restored electric organ discharge duration and amplitude. Duration recovered quicker than amplitude, and both recovered faster in the presence of males than females. In studies of other electric fish species, treatment with steroid sex hormones have taken several days to increase sexual dimorphism in the electric organ discharge. The socially induced changes seen in this study are initiated too quickly to involve classic steroid action of genomic transcription and thus may depend on another mechanism. Socially induced regulation of the males electric organ discharge waveform is consistent with the compromises in signaling strategy shown by other taxa with costly sexual advertisement signals.

Regulation and modulation of electric waveforms in gymnotiform electric fish

Weakly electric gymnotiform fish specialize in the regulation and modulation of the action potentials that make up their multi-purpose electric signals. To produce communication signals, gymnotiform fish modulate the waveforms of their electric organ discharges (EODs) over timescales spanning ten orders of magnitude within the animal’s life cycle: developmental, reproductive, circadian, and behavioral. Rapid changes lasting milliseconds to seconds are the result of direct neural control of action potential firing in the electric organ. Intermediate-term changes taking minutes to hours result from the action of melanocortin peptides, the pituitary hormones that induce skin darkening and cortisol release in many vertebrates. Long-term changes in the EOD waveform taking days to weeks result from the action of sex steroids on the electrocytes in the electric organ as well as changes in the neural control structures in the brain. These long-term changes in the electric organ seem to be associated with changes in the expression of voltage-gated ion channels in two gene families. Electric organs express multiple voltage-gated sodium channel genes, at least one of which seems to be regulated by androgens. Electric organs also express multiple subunits of the shaker (Kv1) family of voltage-gated potassium channels. Expression of the Kv1 subtype has been found to vary with the duration of the waveform in the electric signal. Our increasing understanding of the mechanisms underlying precise control of electric communication signals may yield significant insights into the diversity of natural mechanisms available for modifying the performance of ion channels in excitable membranes. These mechanisms may lead to better understanding of normal function in a wide range of physiological systems and future application in treatment of disease states involving pathology of excitable membranes.

Sex differences in energetic costs explain sexual dimorphism in the circadian rhythm modulation of the electrocommunication signal of the gymnotiform fish Brachyhypopomus pinnicaudatus.

SUMMARY To understand the evolution of sexually dimorphic communication signals, we must quantify their costs, including their energetic costs, the regulation of these costs, and the difference between the costs for the sexes. Here, we provide the first direct measurements of the relative energy expended on electric signals and show for the focal species Brachyhypopomus pinnicaudatus that males spend a significantly greater proportion of their total energy budget on signal generation (11–22%) compared with females (3%). Both sexes significantly reduce the energy spent on electric signals during daylight hours through circadian modulation of the amplitude, duration and repetition rate of the electric signal, but this effect is more marked in males. Male body condition predicted the energy spent on electric signals (R2=0.75). The oxygen consumed by males for signal production closely paralleled the product of the electric signals waveform area (R2=0.99) and the discharge rate (R2=0.59), two signal parameters that can be assessed directly by conspecifics. Thus the electric communication signal of males carries the information to reveal their body condition to prospective mates and competing males. Because the electric signal constitutes a significant fraction of the energy budget, energy savings, along with predation avoidance, provides an adaptive basis for the production of circadian rhythms in electric signals.

Accuracy of auditory distance and azimuth perception by a passerine bird in natural habitat

Small birds should localize sound poorly because small head size limits azimuth resolution and because the loose correlation of acoustic degradation with distance limits accurate estimation of auditory distance. We determined the accuracy of sound localization by a passerine bird in the field using an open-loop phonotaxis experiment. After hearing a playback of a conspecific contact call, eastern towhees, Pipilo erythrophthalmus, approached the silenced source. Mean auditory distance resolution was 7% of total speaker distance and mean azimuth resolution was +/-5 degrees. In a second experiment, we played birds the same calls rerecorded previously over the 10- or 20-m distance beyond each playback location. In 13 of 30 trials, the birds over-flew the speaker by a distance propotional to rerecording; but in 15 trials, approach distances were comparable to speaker distance despite the addition of distance simulated by attenuating and rerecording the calls. Signal-specific and location-specific distance cues are derived to explain the bimodal distribution of flight distances we observed. Copyright 1998 The Association for the Study of Animal Behaviour

Mate preference in female electric fish, Brachyhypopomus pinnicaudatus

Weakly electric fish communicate with brief electrostatic field pulses called electric organ discharges (EODs). EOD waveforms are sexually dimorphic in most genera, a condition thought to result from mate choice acting to shape the electric signal’s constituent action potentials. We have no direct behavioural evidence that sexual selection by either mate choice or intrasexual competition is responsible for sex differences in the EOD waveforms of electric fish. We explored sexual selection in electric fish by conducting two-choice unforced preference tests with live, unaltered gymnotiform electric fish, Brachyhypopomus pinnicaudatus, which are sexually dimorphic. In the initial test, gravid females selected males over females only when the males were larger than average. Gravid females in later tests preferred larger males to smaller males in a significant majority of those trials in which they showed a preference. In about one-third of those trials, females spawned with their preferred male, confirming their preference. We concluded that passage through the choice apparatus was related to mate choice. The signals of chosen males had larger EOD amplitudes and longer EOD durations. These findings show that female B. pinnicaudatus do have a preference for a certain male phenotype. The system requires additional study to dissociate correlated male phenotypic characters to identify which male traits the female prefers.  2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.