Reproductive strategy and singing activity: blue tit and great tit compared

The costs and benefits of bird song are likely to vary among species, and different singing patterns may reflect differences in reproductive strategies. We compared temporal patterns of singing activity in two songbird species, the blue tit (Cyanistes caeruleus) and the great tit (Parus major). The two species live side by side year round, and they have similar breeding ecology and similar rates of extra-pair paternity. However, they differ in two aspects of reproductive strategy that may have an influence on song output: blue tits are facultatively polygynous and have a fairly short breeding season with almost no second broods, whereas great tits are socially monogamous but more commonly raise second broods. We found that great tit males continued singing at high levels during the egg-laying and incubation periods, while monogamously paired blue tit males strongly reduced singing activity after the first days of egg-laying by their female. Since males of both species sang much more intensely shortly before sunrise than after sunrise, at midday or in the evening, this difference was most conspicuous at dawn. No differences in singing activity were found within species when testing for male age. We suggest that in contrast to blue tits, great tit males continued singing after egg-laying to defend the territory and to encourage the female for a possible second brood.     

Interactions between singing Hawaiian humpback whales and conspecifics nearby

Summary Interactions of singing humpback whales, Megaptera novaeangliae, with conspecifics nearly were studied during the breeding season off the west coast of Maui, Hawaii. On 35 occasions singing humpbacks were followed by boats (Table 1). The movement patterns of these singing whales and other conspecifics nearby were recorded by observers on land using a theodolite.Thirteen of 35 singers stopped singing and joined with nonsinging whales either simultaneously or within a few minutes after ceasing to sing. Another 15 also stopped singing while under observation and were not seen to join with another whale, but all singing whales that joined with other whales stopped singing. Singing whales often pursue nonsinging whales, while nonsinging whales usually turn away from singers (Figs. 4, 5).When a singer joined with a female and calf unaccompanied by another adult, behavior tentatively associated with courtship and mating was observed (Fig. 7). Such behavior also occurred during several interactions between singers and individuals of unknown sex. Aggressive behavior was observed during three interactions between singers and individuals of unknown sex (Fig. 4) and it predominated whenever more than one adult accompanied a cow and calf. During the other occasions when a singer joined another whale, we could not determine the nature of the interaction. Many times the singers and joiner would surface together only once and would then separate. However, on several occasions the singer and joiner would remain together for as long as we could follow them, up to 1.5 h.The roles of singer and joiner can be interchangeable. For instance, on two occasions a singer joined with a whale that either had been singing or started singing later in the day (Fig. 3). Furthermore, on several occasions, a nonsinging whale appeared to displace the singer. Individual singing humpbacks are not strictly territorial, although singers appear to avoid other singers.As the breeding season progressed, singers sang for longer periods of time (Fig. 2). In addition, the probability of a whale joining with the singer decreased by 42% from the first half of the observation period to the second half. Furthermore, this increase in duration of song bouts occurred during that section of the season when female reproductive activity as measured by rate of ovulation is reported to be decreasing in other areas.Our observations support the hypothesis that humpback song plays a reproductive role similar to that of bird song. Humpbacks sing only during the breeding season. If, as seems likely, most singing humpbacks are male, then singing humpbacks probably communicate their species, sex, location, readiness to mate with females, and readiness to engage in agonistic behavior with other whales.     

How a songbird with a continuous singing style modulates its song when territorially challenged

In songbirds of the temperate zone, often only males sing and their songs serve to attract females and to deter territorial rivals. In many species, males vary certain aspects of their singing behavior when engaged in territorial interactions. Such variation may be an honest signal of the traits of the signaler, such as fighting strength, condition, or aggressive motivation, and may be used by receivers in decisions on whether to retreat or to escalate a fight. This has been studied intensively in species that sing discontinuously, in which songs are alternating with silent pauses. We studied contextual variation in the song of skylarks (Alauda arvensis), a songbird with a large vocal repertoire and a continuous and versatile singing style. We exposed subjects to simulated territorial intrusions by broadcasting conspecific song and recorded their vocal responses. We found that males sing differently if they are singing spontaneously with no other conspecific around than if they are territorially challenged. In this last case, males produced lower-frequency syllables. Furthermore, they increased the sound density of their song: they increased the proportion of sound within song. They seem to do so by singing different elements of their repertoire when singing reactively. Furthermore, they increased the consistency of mean peak frequency: they repeated syllable types with less variability when singing reactively. Such contextual variation suggests that skylarks might use low frequencies, sound density, and song consistency to indicate their competitive potential, and thus, those song features might be important for mutual assessment of competitive abilities.     

Voices in the dark: predation risk by owls influences dusk singing in a diurnal passerine

Predation is an important cost of communication in animals and thus a potent selection pressure on the evolution of signaling behavior. Heterospecific eavesdropping by predators may increase the vulnerability of vocalizing prey, particularly during low light, such as at dusk when nocturnal predators are actively hunting. Despite the risk it entails, dawn and dusk chorusing is common in passerines. However, the dusk chorus has not been studied much, neglecting the opportunity for understanding how eavesdropping between predators and prey may shape communication in birds. Here, we report the first demonstration of simulated predation risk (playback of owl vocalizations) altering the dusk chorus of a diurnal passerine, the veery (Catharus fuscescens). Veeries have a pronounced dusk chorus, singing well after sunset and potentially exposing themselves to predation by owls. In response to brief playbacks of owl calls (~30 s of calls presented three times over 25 min), veeries sang fewer songs post-sunset and stopped singing earlier relative to control trials. These changes in singing remained evident 30 min after the last owl stimulus. Although the avian dusk chorus has received relatively little attention to date, our results suggest that the dusk chorus may pose a higher predation risk to singing males that may influence the evolution of singing behavior in diurnal birds.     

Changes in singing behavior of male black-capped chickadees (Parus atricapillus) following mate removal

We removed the mates of ten male black-capped chickadees (Pares atricapillus) during the nest-building period to determine the effect of female presence on dawn singing. During the first dawn chorus following mate removal, males sang significantly longer, increased movement within their territory, and increased the percentage of their territory covered while singing. After the female was returned, these parameters returned to the pre-removal values. Males did not alter the frequency range or modal frequency of their songs when the mate was removed, nor did they change the degree of frequency shifting in the fee-bee song. We conclude that dawn singing in the black-capped chickadee acts, in part, as an intersexual signal, and that the behavior of frequency shifting in the song may be directed more toward rival males than females. Correspondence to: K. Otter     

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.     

Vocal interactions in common nightingales (<Emphasis Type="Italic">Luscinia megarhynchos</Emphasis>): males take it easy after pairing

Seasonal patterns of bird song have been studied intensively with a focus on individual males. However, little is known about seasonal patterns of singing during vocal interactions between males. Vocal interactions have been shown to be important in sexual selection as males may signal aspects of motivation or quality. Here, we investigated in nightingales (Luscinia megarhynchos) whether a male’s behaviour in vocal interactions at different stages of the breeding season is influenced by its mating status. We examined how males that differ in their subsequent mating success respond to a non-interactive, nocturnal playback presented during the period of mate attraction and subsequently during the egg-laying period. We found that mated males overlapped fewer songs and had a lower song rate during the egg-laying period compared to their responses during the mate-attraction period, whereas unpaired males did not vary in their responses between the two periods. Our results suggest that mating status is a key factor affecting singing behaviour in vocal interactions and that a time-specific singing pattern like song overlapping is used flexibly during vocal interactions. Because song overlapping is thought to be a signal of aggression in male–male vocal interactions, it seems that males vary the level of aggression in vocal interactions according to their mating status and to the stage in the breeding season.     

Choice of singing sites by male bushcrickets (Tettigonia viridissima) in relation to signal propagation

Summary The spatial dispersion of singing male bushcrickets (Tettigonia viridissima) in a marshland habitat was found to be significantly clumped. Males clustered in patches of taller vegetation, buth within such clusters, males were regularly spaced with a mean distance of 6 m between nearest-neighbours. Males occupied perches on plants that were, on average, 0.3 m above the top of surrounding vegetation. Physical and acoustical interactions between males were observed more frequently when males were singing from higher sites. Excess attenuation of the male song was found to increase with frequency but decreased markedly with increasing elevation of the singing male above the ground. The maximum detection range of the song, realized when the insect was singing > 1 m above the surrounding vegetation, was estimated as 60 m for the fundamental frequency (10 kHz), 38 m for the 1st harmonic (20 kHz) and 14 m for the second harmonic (30 kHz). By contrast, when males sang from the middle of dense reed beds, the estimated detection distance was only 8 m, 6 m and 4 m for each frequency band, respectively. Males could have increased the detection range of their songs almost three fold by singing from higher positions than those usually observed in the field. This suggests that there may be a cost of singing at higher elevations such as an increased risk of predation and/or increased aggression from neighbours. We suggest the spacing strategy adopted by males reflects a compromise between maximizing the range over which their songs can be detected and accurately localized by females and minimizing interference from competing males. Offprint requests to: T. Eiriksson     

Sound degradation as a distance cue in great tit (Parus major) song

Summary We report an experiment designed to test the ideas that: 1. male songbirds can use cues from the distortion of song by environmental factors (degradation) to estimate the distance of another singing male; 2. song degradation is assessed by reference to an internal standard. Great tits respond more strongly to undegraded than to degraded songs when both are played at the same amplitude and from the same position in the territory. This difference in response is shown only if the playback song is familiar to the test bird; familiar songs being those sung either by the test bird or neighbours of the test bird. We interpret these results as evidence that cues from song degradation can be used to estimate the distance of a singing conspecific and that degradation assessment is only possible if the bird has an internal representation of the song (because either it and/or a neighbour sings the song). We discuss the implications of these results for Morton's (1982) ranging hypothesis, and for the distinction between learning and performance in bird song. Our results partially support the ranging hypothesis, but question the nature of unrangeable songs sensu Morton. The finding that birds can assess the degree of degradation of songs that they do not sing, supports the idea that birds learn more songs than they sing.     

A dusk chorus effect in a nocturnal bird: support for mate and rival assessment functions

The dawn chorus is a striking feature of spring mornings and a characteristic behaviour of many bird species, particularly the passerines. Dawn singing has been considered a reliable signal of male quality for mate and rival assessment. Singing is presumed to be relatively costly at dawn both because air temperatures are relatively low and because birds have not fed overnight. Models of optimal daily routine predict the existence of a “dusk chorus” in nocturnal birds, although this prediction has received little empirical attention. Nocturnal birds at dusk may be energy-limited because of a lack of daytime feeding, and singing at dusk may thus ensure signal reliability. Here, we used an observational and experimental approach to study vocal behaviour at dusk and dawn in a nocturnal raptor, the little owl Athene noctua. We assess whether male little owls adjust their vocal behaviour according to feeding stage (i.e. period of the night), ambient air temperature and territorial context (i.e. spontaneous calling behaviour vs elicited calling by intrusion). Across different temperatures, we find that both spontaneous vocal activity and inter-individual variability in call duration increased at dusk, clearly indicating a dusk chorus phenomenon. Results from playback presentations suggest that food, rather than air temperature, is likely to be more constraining at dusk. We discuss how comparing dusk and dawn choruses in nocturnal and diurnal species can provide insights into both mechanistic and functional aspects of signalling behaviour.     

Structure and evolution of song form in the wrens Thryothorus sinaloa and T. felix

Summary The songs of two species of wrens, Thryothorus sinaloa and T. felix, are less similar than previously believed. We have found that song structure, bout patterning, and other aspects of singing behavior are widely divergent between the two species, which are sympatric over much of their ranges. The nature of these differences supports the hypothesis of contrast reinforcement in bird song. Further evidence is provided by essentially parallel differences in another pair of sympatric species, T. pleurostictus and T. maculipectus. A cluster analysis of song form of the above four species plus one other, T. ludovicianus, suggests that those species which are allopatric are the most similar in song form, while sympatric species are the most dissimilar. We relate various features of singing behavior in the two species to some aspects to their ecology, and examine relationships among various song parameters within the genus Thryothorus. The parallel differences in pairs of sympatric Thryothorus in structure of song units, temporal organization of song, and duetting, along with the close correlation of groups of song parameters in the five species of Thryothorus leads us to propose that strategies or adaptive complexes of singing characteristics may exist in the genus.     

Bachelor and paired male rufous-and-white wrens use different singing strategies

To attract a breeding partner, males may behave differently when they are bachelors compared to when they are paired. Comparisons between groups of paired males versus groups of unpaired males in temperate-breeding animals have revealed such differences in signalling behaviour. Few studies, however, have explored how individual males alter their signalling behaviour with changes in pairing status, and very few investigations have explored paired versus unpaired male behaviour in tropical animals. During a 5-year study in Costa Rica, we analysed changes in the singing behaviour of male rufous-and-white wrens (Thryothorus rufalbus) when they were paired and when they were bachelors. We compared three aspects of male vocal behaviour: gross differences in song output, variation in repertoire use and differences in song structure. Males as bachelors had significantly higher song output and switched song types less frequently. Contrary to our expectation, bachelors sang significantly fewer song types from their repertoire compared to when those same males had a breeding partner. Songs sung by bachelor males were higher in syllabic diversity and had broader-bandwidth terminal syllables than the songs those males sang only when paired. Within song types, the fine structure of songs remained consistent across pairing status. Our results demonstrate that males change their singing behaviour with pairing status, delivering songs at a higher rate but with less variety when they are bachelors. Rufous-and-white wrens are renowned for their vocal duets, and we discuss the pattern of repertoire use in light of their duetting behaviour. These results enhance our understanding of how male behaviour varies with pairing status and the importance of vocal signalling behaviour in socially monogamous tropical animals.     

Effects of different levels of song overlapping on singing behaviour in male territorial nightingales (Luscinia megarhynchos)

In signalling interactions, animals can directly address information to a specific individual. Vocal overlapping is such a signalling strategy used in songbirds, anurans, and insects. In songbirds, numerous studies using high rates of song overlap to simulate an escalating situation have shown that song overlapping is perceived as a threatening signal by interacting and by listening (eavesdropping) individuals, indicating a high social relevance of song overlapping. Here we present a playback experiment on nocturnally singing male territorial nightingales (Luscinia megarhynchos). Using three different rates of song overlap (1, 25, or 50%), we tested whether or not lower levels of song overlapping act as a signal of aggressive intent and if birds would increase the intensity of their response with increasing level of song overlapping. Subjects did not vary song duration in response to the different playback treatments but increasingly interrupted their singing with increasing overlap by the three playback treatments. The effects persisted even after the playback ceased to overlap and switched to an alternating singing mode. These results expand on previous studies by showing that song overlapping is interpreted as an aggressive signal even when it is used at low or moderate levels. They suggest that, within the range tested here, increasing levels of song overlapping are perceived to be increasingly aggressive.     

Factors affecting aural detections of songbirds.

Many factors affect the number of birds detected on point count surveys of breeding songbirds. The magnitude and importance of these factors are not well understood. We used a bird song simulation system to quantify the effects of detection distance, singing rate, species differences, and observer differences on detection probabilities of birds detected by ear. We simulated 40 point counts consisting of 10 birds per count for five primary species (Black-and-white Warbler Mniotilta varia, Black-throated Blue Warbler Dendroica caerulescens, Black-throated Green Warbler Dendroica virens, Hooded Warbler Wilsonia citrina, and Ovenbird Seiurus aurocapillus) over a range of 15 distances (34-143 m). Songs were played at low (two songs per count) and high (13-21 songs per count) singing rates. Detection probabilities averaged across observers ranged from 0.60 (Black-and-white Warbler) to 0.83 (Hooded Warbler) at the high singing rate and 0.41 (Black-and-white Warbler) to 0.67 (Hooded Warbler) at the low singing rate. Logistic regression analyses indicated that species, singing rate, distance, and observer were all significant factors affecting detection probabilities. Singing rate x species and singing rate X distance interactions were also significant. Simulations of expected counts, based on the best logistic model, indicated that observers detected between 19% (for the worst observer, lowest singing rate, and least detectable species) and 65% (for the best observer, highest singing rate, and most detectable species) of the true population. Detection probabilities on actual point count surveys are likely to vary even more because many sources of variability were controlled in our experiments. These findings strongly support the importance of adjusting measures of avian diversity or abundance from auditory point counts with direct estimates of detection probability.     

Temporal performance roles during vocal interactions in nightingales (Luscinia megarhynchos B.)

Summary The temporal patterning of vocal interactions between territorial nightingales (Luscinia megarhynchos B). was investigated during nocturnal dyadic singing. We distinguished three forms of temporal performance roles (Table 1; Fig. 3): inserter (preferred song start 0.5–1.0 s after offset of a neighbour's song), overlapper (preferred song start 0.5–1.0 s after onset of a neighbour's song), and Autonomous bird (no evidence for temporal responses adjusted to the timing of a neighbour's singing; Fig. 4). With the beginning of the daily dawn chorus, mutual temporal adjustment of song performances could no longer be ascertained (Fig. 2). To test the flexibility of song onset timing, we presented subjects with three playback programs (I, II, III) simulating a non-flexible conspecific bird. Sequences of songs recorded from a stranger that shared no song types with the subjects were used; intersong pauses were unaltered in I, lengtened in II, and shortened in III. Results confirmed that the timing of song onset can be influenced by the temporal patterning of auditory stimuli (Table 1; Fig. 5): Inserters and overlappers sang at a slower rate during program II than during program I. During program III, which by accelerated stimulus succession caused frequent overlap of songs, inserters and overlappers interrupted their singing behaviour (Fig. 6). This suggests that overlapping, which results in signal interference, is of some functional significance in vocal interactions.     

Flocking behaviour of passerines: a dynamic model for the non-reproductive season

Summary The behaviour of a small male passerine bird over a typical winter day is studied by a dynamic programming model. The bird can be either unpaired or paired; an unpaired bird can forage in a flock, forage alone or sing to attract a mate. Foraging increases his reserves, while singing reduces them. The optimal policy and the expected behaviour of birds depend both on time and reserves. The model predicts that birds will flock, especially in the morning, if flocking birds find more food (foraging efficiency), and also more flocking can be expected when the predation risk is lower in a flock (antipredator benefit). Where flocking gives lower variance in food intake, with the same mean (reduced variance benefit), birds with low reserves at the end of the day choose to forage alone (high variance option), while otherwise they are risk-averse and forage in a flock. The cost of flocking increases with time in a day and with the probability of mate attraction through singing. Decisions inevitably involve trade-offs. Where flocking results in antipredator benefit, but also lower foraging gain, birds with low reserves forage alone, but birds with high reserves flock. Offprint requests to: A.I. Houston     

Daily patterns of energy storage in food-caching birds under variable daily predation risk: a dynamic state variable model

Accumulating and maintaining sufficient energy reserves is critical for winter survival of birds. Because high fat levels are assumed to be associated with higher risk of predation, birds have been hypothesized to regulate their body mass as a trade-off between risk of starvation and risk of predation. Theoretical models of energy management in birds typically assume that predation risk is constant throughout the day. However, this important assumption has little empirical support, and there is some evidence suggesting that it might not always be correct and that predation risk may vary during the day. Because predation risk is a critical component of the predation-starvation trade-off, any change in its value through the day might have a profound effect on birds' optimal daily tactics of energy accumulation. We used a dynamic optimization model to investigate how changes in predation risk might affect birds' energy management decisions. Daily patterns of fat accumulation and feeding activity were predicted to change with predation risk in a manner consistent with previous models (lower mass gain and less feeding when predation risk is high). Our more counterintuitive results concern daily patterns of food caching and cache retrieval. When predation risk was assumed to peak at midday, birds were predicted to cache primarily in the afternoon and not in the morning even though predation risk was identical in the morning and afternoon. With other temporal patterns of predation risk, caching intensity was highest when predation risk was lowest. Surprisingly, the daily pattern of cache retrieval was predicted to be unaffected by daily patterns of predation risk: birds were always predicted to retrieve their caches primarily during the late afternoon with a small peak in the morning. Highest mortality was predicted with predation risk decreasing from morning to evening whereas lowest mortality was predicted with predation risk increasing from morning to evening. Our model helps explain large variations in observed daily patterns of energy management in birds and provides testable predictions that could help us understand the daily dynamics of predation risk and how birds should respond to it.     

Patterns of Rarity in the Birds of the Atlantic Forest of Brazil

Patterns of rarity in species are generally explained by several factors: evolutionary history, spatial distribution, and genetic structure of each taxon. Human intervention also leads to or increases rarity in species. The discernment of causes of rarity is essential to the understanding of extinction patterns, and thus to devising conservation strategies. I examine patterns of rarity among bird species in the Atlantic forest region in Brazil, one of the most threatened ecosystems in the world. I assigned bird species to one of eight possible categories that differ in degree of vulnerability and that are based on three parameters of rarity: geographic distribution, habitat specificity, and population size. The Atlantic forest avifauna is a highly endangered group; 68% of the species are rare. Patterns of rarity among the birds in the region likely result from their specific ecologies or evolutionary histories. In addition, human alteration of natural habitats and hunting pressures have undoubtedly influenced rarity for a number of species.     

Persistent bimodal activity patterns in wild and captive black-tailed godwit <Emphasis Type="Italic">Limosa limosa</Emphasis> under different environmental conditions

There is a large literature dealing with daily foraging routines of wild birds during the non-breeding season. While different laboratory studies have showed that some bird activity patterns are a persistent property of the circadian system, most of field studies preclude the potential role of an endogenous circadian rhythm in controlling bird’s foraging routines. In this study we compared the patterns of diurnal foraging activity and intake rates of migrating black-tailed godwits, Limosa limosa (radio-tagged and non-tagged individuals) at two stopover sites (habitats) with different environmental characteristics, aiming at identifying proximate factors of bird activity routines. To gain insights into the role of food availability in control of such foraging routines, we also estimated foraging activity patterns in captive godwits subjected to constant food availability. Captive and wild black-tailed godwits showed a persistent bimodal activity pattern through daylight period. Food availability had a significant effect on the intake rates, but had a subtler effect on foraging and intake rate rhythms. Temperature and wind speed (combined in a weather index) showed non-significant effects on both rhythms. Although we could not discard a role for natural diurnal changes in light intensity, an important timing cue, our findings support the idea that an endogenous circadian rhythm could be an important proximate factor regulating foraging activity and food items taken per unit time of wild black-tailed godwits during migration.     

Living in a ghetto within a local population: an empirical example of an ideal despotic distribution

Merging patterns and processes about the way individuals should be distributed in a habitat is a key issue in the framework of spatial ecology. Here the despotic distribution of individuals in two distinct and neighboring patches within a local population of a long-lived colonial bird, the Yellow-legged Gull (Larus michahellis), was assessed. There was no density dependence for suitable habitat at the study population, but behavioral data suggested that birds from the good patch precluded birds from the bad patch from breeding in their patch. Younger breeders were almost exclusively found in the bad patch, where individuals were probably attracted by conspecific attraction from the good patch. Most breeding parameters were lower in the bad patch, resulting mainly from a higher vulnerability to environmental perturbations and a higher rate of intraspecific nest predation. Attempts at breeding dispersal between the two patches were only observed from the bad to the good patch. Strikingly, adult survival and large-scale dispersal, two life history parameters that are very conservative in long-lived organisms, were also more affected at the bad patch when catastrophic predation occurred. The study was consistent with an ideal despotic distribution at small spatial scale, and suggests that individual behavior can influence local population dynamics.