Long-term influence of simulated territorial intrusions on dawn and dusk singing in the Winter Wren: spring versus autumn

Males of many songbird species have peaks of singing activity at dawn and dusk. Singing during those twilight periods can function in territory proclamation, and males are suggested to adjust song output to the level of intruder pressure. We used song playback during the breeding season to simulate intrusions into territories of male Winter Wrens (Troglodytes troglodytes) shortly after dawn. We then compared male singing behaviour during the dawn and dusk chorus before and 1 day after the simulated intrusion. One day after the playback, male Wrens increased their song output before sunrise, which confirms our results from a previous study on dawn singing in autumn territories. At dusk, on the evening following the playback, males slightly increased song output after sunset, but singing activity at dusk was generally very low. We found no significant changes of song output after sunrise, before sunset, and between 2 days of control without playback. These results are consistent with the hypothesis that dawn and dusk singing is important for territory defence in spring. Unlike in autumn, however, increased singing in spring at dawn and dusk could also serve to defend other resources such as fertile mates or to strengthen the pair bond after a territorial challenge. In comparison with the results on autumnal singing, male Wrens started singing earlier at dawn during the breeding season, and they generally sang more songs at dawn and immediately after playback. The increase in absolute numbers of songs sung in the morning after playback seemed greater in spring than in autumn; however, the proportional increase relative to overall song output was similar in both seasons.     

Conveying information with one song type: changes in dawn song performance correspond to different female breeding stages

Many bird species participate in dawn singing, a behaviour categorized by intensive singing at dawn; however, many of these species deliver only one song type at dawn. While there are many proximate and ultimate hypotheses for why birds sing at dawn, little is known about whether males are able to vary one simple song to convey different information. We used autonomous acoustic recorders to record dawn songs of field sparrows and quantified three parameters of singing performance: 1) bout length, 2) song rate, and 3) song complexity. We found that males sang the longest dawn bouts during their mate's fertile period, the highest song rates during the post-fertile period, and the most complex songs during the pre-fertile period. The change in dawn singing behaviour with their mate's breeding stage suggest the purpose of dawn song may be context dependent. Our results demonstrate that male field sparrows, while only having a single song type sung at dawn, may convey information for both intra- and intersexual purposes. While it is generally assumed that dawn song has a specific function, the variability in the duration, rate, and complexity of dawn song in field sparrows suggests that they are conveying different information and that dawn song likely has multiple functions.     

Honest advertisement and song output during the dawn chorus of black-capped chickadees

Costly signals can evolve under sexual selection, as only thosesignals that are difficult to produce and reflect the relativequality of individuals should be important in mate choice. Onesuch signal may be dawn singing behavior in birds. We assessedwhether the song output at dawn of breeding male black-cappedchickadees Parus atricapilhis honestly reflects quality, whererelative quality is assessed by relative dominance rank in winterflocks. Dawn choruses were recorded from 20 male chickadeesfrom 10 flocks during the fertile period of their mates in 1992,1994, and 1995. Dominance ranks of males were assessed by tabulatinginteractions at winter feeders from 1993 to 1995. A comparisonof the dawn singing behavior of the high-ranking and the low-rankingmales from each of the 10 flocks showed that high-ranking malesbegan singing earlier, sang longer, and sang at higher averageand maximum rates than low-ranking flockmates. Age of the maleshad less effect on song output at dawn than rank; older malestended to sing longer dawn choruses, but there was no differencein onset of singing, average song rate, or maximum song rateat dawn between hatch year and after-hatch year males. Our findingssuggest the dawn chorus can provide an accurate signal to femalesof the relative quality of their mate compared to neighboringmales     

Singing the Popular Songs? Nightingales Share More Song Types with Their Breeding Population in Their Second Season than in Their First

Signals used in communication often change throughout an individual’s life course. For example, in many song bird species, males modify their song especially between their first and second breeding season. To address one possible reason of such modification, we investigated whether common nightingales Luscinia megarhynchos adjust their song type repertoires to sing the song types commonly occurring in their breeding population. We analysed nocturnal singing of six nightingales in their first and second breeding season and compared their repertoire composition and usage to the ‘typical’ repertoire and usage on the breeding ground (represented by seven reference birds). Songs that were maintained between the first and second season by the six focal birds occurred in most of the repertoires of the seven reference birds and were sung often. In contrast, song types that were dropped from the repertoires occurred less often in the reference birds’ repertoires and were sung less often. Furthermore, in the first year, each focal nightingale’s repertoire was less similar to the reference birds’ repertoires than in the second year. Thus, nightingales adjusted their singing towards the songs popular in the breeding grounds by keeping song types that were common and frequently sung by other individuals in their breeding area and by disposing of infrequently performed ones. This resulted in increased similarity with the population’s repertoire from the first to the second year. We discuss possible ontogenetic processes that may lead to such an adjustment and suggest an improved ability to match song types as possible adaptive value.     

The effect of variability in the food supply on the daily singing routines of European robins: a test of a stochastic dynamic programming model

A stochastic dynamic programming (SDP) model offers a general explanation of daily singing routines in birds, but remains almost untested empirically. I examined a central prediction of the SDP model, that a more variable food supply decreases the bird's song output at dawn, relative to its song output at dusk. I provided supplementary food to make the food supply more or less variable over 2-week periods in the territories of free-living European robins Erithacus rubecula. Robins sang relatively less at dawn than at dusk after weeks in which their supplementary food supply was variable, and more at dawn than at dusk after weeks in which their food supplementation was constant. These results provide strong support for the prediction of the SDP model. Copyright 1999 The Association for the Study of Animal Behaviour.     

Daily energy expenditure of singing great reed warblers Acrocephalus arundinaceus

According to honest signalling theory, signals must be costly to produce to retain information about the signaller's quality. The song produced by male birds during breeding is a vocal "ornament" used for intra- and inter-sexual purposes. The energetic cost of this vocal signal remains a contentious issue. We used the doubly labelled water method to measure field metabolic rate by estimating CO₂ production and then convert this to daily energy expenditure (DEE) in great reed warbler males singing under natural conditions (10 at low to moderate intensity and 7 at very high intensity from dawn to dusk). There was a significant positive relationship between singing intensity and DEE. From this relationship we extrapolated the average DEE for intensely singing males (i.e., males producing song sounds 50% of the time and hence sitting at their elevated song post in the top of a reed stem more or less continuously throughout the ∼20 h of daylight) to 3.3×BMR (basal metabolic rate) and for non-singing males to 2.2×BMR. The mean DEE measured for the seven males singing with very high intensity was 3.1×BMR. The maximum measured DEE for a single male was 3.9×BMR, i.e. close to the maximum sustainable DEE (4×BMR), and the minimum DEE was 2.1×BMR for a male singing at very low intensity. These results imply that producing intensive advertising song in birds may incur a substantial cost in terms of increased energy expenditure.     

Experimentally Elevated Plasma Testosterone Levels Do Not Influence Singing Behaviour of Male Blue Tits (Parus caeruleus) During the Early Breeding Season

Sexual selection theory suggests that females might prefer males on the basis of testosterone (T)‐dependent secondary sexual traits such as song. Correlational studies have linked high plasma T‐levels to high diurnal song output. This has been confirmed in experiments where T‐levels were kept high at times when natural T‐levels have decreased. However, surprisingly little is known about the relation between T‐levels during the early breeding season and song. In many passerine birds males sing at a high rate at dawn early in the breeding season, referred to as the dawn chorus. In blue tits (Parus caeruleus), the dawn chorus coincides with the fertile period of the female, whereas diurnal song occurs throughout the breeding season. Previous studies on blue tits showed that characteristics of the dawn chorus correlate with male reproductive success. We experimentally elevated plasma T‐levels in male blue tits during the pre‐fertile and fertile period. Our aim was to test whether increased plasma T‐levels affect dawn song characteristics and increase the amount of diurnal song. Although T‐implants successfully raised circulating T‐levels, we did not find any difference between T‐ and control males in temporal performance measures of dawn song or in diurnal song output. Our results suggest that either there is no direct causal link between song output or quality and individual T‐levels, or experimental manipulations of T‐levels using implants do not permit detection of such effects during the early breeding season. Although we cannot exclude that individual T‐levels are causally linked to other (e.g. structural) song parameters, our results cast doubt on T‐dependence as the mechanisms that enforces honesty on song as a sexually selected trait.     

Variation in Song Rate during the Breeding Cycle of the Chaffinch,Fringilla coelebs

The variation in song rate during the breeding season was studied in two individually marked chaffinch Fringilla coelebs populations. We gathered data to investigate especially the recently presented mate-guarding hypothesis. The active singing has been supposed to function as a form of mate guarding during the female's fertile period by announcing the high status of the male and preventing extra-pair copulations by neighbouring males. There was no clear dawn chorus in the chaffinch, i.e. a peak in the song rate before sunrise. Male chaffinches continued to sing after mating, but the song rate dropped significantly. In contrast to the mate-guarding hypothesis the song rate was lower during the fertile period of the female than during pre-mating and incubation. Thus, the males do not announce the fertility status of their mates or their own quality and status by active singing. The song does not function as a form of mate guarding in the chaffinch. One function of the song of the chaffinch is mate attraction: singing activity was highest before pair formation in early spring and decreased after mating but increased again if the male lost his mate later in the breeding season.     

Interspecific Response to Playback of Bird Song

Responses to bird song have usually only been studied at the intraspecific level. I experimentally tested whether playback of the song of the black wheatear Oenanthe leucura in an area in S Spain resulted in responses from conspecifics as well as heterospecific birds by comparing the numbers of individuals singing before and after playback. The number of singing male black wheatears increased considerably, but also the number of singing males of five other passerine species increased significantly. The heterospecific response to playback may be due (1) to interspecific territoriality, (2) to black wheatear song signalling the absence of predators, or (3) to heterospecifics confusing the species-identity of the singer. The second alternative is considered more likely, since an ecologically wide array of species increased their song rate following playback. The conspicuous dawn (and dusk) chorus of bird song may be augmented by social facilitation due to the singing of conspecifics as well as heterospecifics.     

Understanding the function of nocturnal song in ovenbirds: males do not countersing at night

Many diurnal bird species vocalize at night, however the function of nocturnal song is, generally, still poorly understood. Previous research has suggested that nocturnal song may serve a social function and is influenced by environmental factors. To test whether males attend to the nocturnal song of conspecifics, we experimentally exposed ovenbirds Seiurus aurocapilla to nocturnal flight songs, and recorded their response both during the night and during the following dawn chorus. We compared latency to song and vocal output before and after playback exposure to determine if males altered their vocalizations in response to exposure to flight songs from an unknown male. We found no evidence of counter singing or change in nocturnal song output, nor a change in vocal output during the dawn chorus following playback exposure. Our results suggest that, in ovenbirds, nocturnal song does not serve as an intraspecific social function. Nocturnal song, through rare, may be significant in the mating systems of some diurnal bird species, and requires additional study.     

Two tests of a stochastic dynamic programming model of daily singing routines in birds

Many hypotheses have been put forward to account for the dawn chorus in birds. Few of these, however, are able to account for variation in song output over the whole day, or for differences in daily singing routines between species, individuals, seasons and environmental conditions. One hypothesis that does offer a more general explanation is based on a stochastic dynamic programming (SDP) model of daily singing routines. This model relates the relative costs and benefits of feeding and singing at different times of day to the size of a bird's fat reserves and calculates the optimal daily routines of singing and foraging that will maximize the amount that the bird can sing while avoiding starvation. The use of SDP models in behavioural ecology has become well established, but they remain largely untested empirically. I tested two predictions of the SDP model of daily routines of singing, using free-living European robins Erithacus rubecula. The results supported both predictions: (1) food supplementation causing unpredictable short-term increases in foraging success increased subsequent song output; and (2) changes in ambient temperature were positively associated with changes in subsequent song output. Copyright 1999 The Association for the Study of Animal Behaviour.     

The costs of singing in nightingales

The costs of singing in birds are poorly understood. One potential type of cost is a metabolic cost of singing. Previous studies have measured short-term changes in oxygen consumption associated with bouts of vocalizations, with equivocal results. In this study, I used an alternative approach to measuring the metabolic cost of singing, by measuring overnight loss of body mass, in male common nightingales, Luscinia megarhynchos, singing at night at different rates. Nightingales were shown not to forage at night. They reached a higher mass at dusk prior to singing more at night, and lost more mass overnight when dusk mass and overnight song rate were high. These results show that singing at night is associated with increased overnight consumption of body reserves, which represents a significant metabolic cost of singing at night. However, the correlation between dusk mass and overnight song rate makes it impossible to determine whether these costs arise from the energetic costs of the singing itself, or from the metabolic costs of the additional body reserves laid down at dusk on nights when song rate was high. There are also likely to be costs associated with accumulating and carrying these extra body reserves during daylight, as well as other potential costs of singing such as an increased risk of predation. These results are consistent with those models of signalling in biology that predict or assume that honest signals are costly. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sex-specific timing of mate searching and territory prospecting in the nightingale: nocturnal life of females

Formal models have shown that diel variation in female mate searching is likely to have profound influence on daily signalling routines of males. In studies on acoustic communication, the temporal patterns of the receivers'' signal evaluation should thus be taken into account when investigating the functions of signalling. In bird species in which diel patterns of signalling differ between males singing to defend a territory or to attract a mate, the diel patterns of mate and territory prospecting are suggested to depend on the sex of the prospector. We simulated newly arriving female nightingales (Luscinia megarhynchos) by translocating radio-tagged females to our study site. The mate-searching females prospected the area mostly at night, visiting several singing males. The timing of female prospecting corresponded to the period of the night when the singing activity of unpaired males was higher than that of paired males. In contrast to females, territory searching males have been shown to prospect territories almost exclusively during the dawn chorus. At dawn, both paired and unpaired males sang at high rates, suggesting that in contrast to nocturnal singing, dawn singing is important to announce territory occupancy to prospecting males. In the nightingale, the sex-specific timing of prospecting corresponded to the differential signalling routines of paired and unpaired males. The temporal patterns in the behaviour of signallers and receivers thus appear to be mutually adapted.     

Social factors affect the singing rates of female northern cardinals Cardinalis cardinalis

Social factors, such as the duration of territorial occupancy or of time paired with a mate can affect the rate at which individual birds sing. This study examined the influence of duration of pair-bond and territory occupancy as well as date on the rate of singing by male and female northern cardinals Cardinalis cardinalis . When differences in breeding status were controlled, female cardinals sang at higher rates earlier in the season. Females in newly formed pairs sang at significantly higher rates than those in pairs that had previously bred together. In contrast, male cardinals did not show significant variation in the rate of singing throughout the season. The song rates of males in newly formed and established pairs did not differ significantly. Song rates for males and females in mated pairs were not significantly correlated. This study suggests that social factors have a strong effect on the rate at which female cardinals sing. It is possible that increased intra-sexual aggression by females when they are establishing a new territory with a new mate leads to this higher level of song output. Once females have established a territory and acquired a mate, dear-enemy effects might diminish the need for acoustic territorial defence. Differences in the constraints of nesting or the attraction of extra-pair mates might explain the sexual differences in male and female singing behaviour.     

The Return of the Intruder: Immediate and Later Effects of Different Approach Distances in a Territorial Songbird

Male songbirds often maintain territories throughout the breeding season, and one of the main functions of song is to deter invaders. Therefore, the distance of an unknown singing rival should play a crucial role within territorial singing interactions of males. This distance is expected to be assessed as more threatening the closer the rival approaches. Here, we tested this assumption by conducting nocturnal playbacks from two different distances in territorial Common nightingales (Luscinia megarhynchos). Immediate vocal responses of birds were examined by analysing changes in song structure as well as temporal response features. The next morning, follow‐up playbacks from an intermediate distance allowed us to investigate longer‐lasting effects of nocturnal playbacks. We found that the distance of a simulated rival had an effect on both immediate and later vocal responses of territorial male nightingales with different song parameters being affected during nocturnal and diurnal singing. This indicates that birds perceive intruder distances and adjust their response behaviour both immediately and in later interactions.     

Patterns of dawn singing by Buff-breasted Flycatchers

ABSTRACT.   Many passerine species exhibit a "dawn chorus"—a bout of intense singing activity before or at dawn, but our understanding of this phenomenon is poor. Tyrant flycatchers (Tyrannidae) exhibit pronounced daily bouts of dawn singing. I documented this behavior in several populations of Buff-breasted Flycatchers ( Empidonax fulvifrons ) in Arizona, tape recording >30,000 songs of 23 individuals during dawn singing. Individual males sang a dawn bout each morning, even during breeding phases when daytime song was almost completely absent. Dawn bouts began 5–10 min before local civil twilight and continued for 25–30 min. Each male possessed two song types delivered at high rates during dawn singing (four times the rate during sustained daytime singing). Song rate varied significantly over the course of dawn bouts, increasing to 55 songs / min at mid-bout, then declining to the end of the bout. Type 2 songs comprised about 30% of songs during dawn singing, and decreased significantly in proportion during the final 10 min of the bout. Songs of the two types were delivered in a nonrandom fashion. Males sang at locations near territory boundaries and pairs of neighbors engaged in counter-singing from the same locations each morning. A number of dawn singing bouts ended with attempted or successful copulations. These observations are consistent with the social dynamics hypothesis for the functional significance of dawn singing in this species.     

Social factors affect the singing rates of female northern cardinals Cardinalis cardinalis

Social factors, such as the duration of territorial occupancy or of time paired with a mate can affect the rate at which individual birds sing. This study examined the influence of duration of pair‐bond and territory occupancy as well as date on the rate of singing by male and female northern cardinals Cardinalis cardinalis. When differences in breeding status were controlled, female cardinals sang at higher rates earlier in the season. Females in newly formed pairs sang at significantly higher rates than those in pairs that had previously bred together. In contrast, male cardinals did not show significant variation in the rate of singing throughout the season. The song rates of males in newly formed and established pairs did not differ significantly. Song rates for males and females in mated pairs were not significantly correlated. This study suggests that social factors have a strong effect on the rate at which female cardinals sing. It is possible that increased intra‐sexual aggression by females when they are establishing a new territory with a new mate leads to this higher level of song output. Once females have established a territory and acquired a mate, dear‐enemy effects might diminish the need for acoustic territorial defence. Differences in the constraints of nesting or the attraction of extra‐pair mates might explain the sexual differences in male and female singing behaviour.     

Dawn song of male blue tits as a predictor of competitiveness in midmorning singing interactions

During the dawn chorus, territorial male songbirds vocalise intensively within signalling range of several conspecific males and can therefore be considered members of a busy communication network. The more or less continuous singing over a long period of time under standardised stimulus conditions makes the dawn song a potentially important information source both for simple receivers and for eavesdroppers. Male blue tits (Parus caeruleus) vary in features of their dawn song, e.g. older males sing longer strophes, and females choose males that sing longer strophes as extra-pair partners. However, so far, dawn song in the blue tit has been investigated separately from other singing behaviour of the same males. In this study, we investigate aspects of blue tit male quality, reflected in dawn song characteristics, and their predictive value for how males behave during singing interactions later in the morning. We acted as simple receivers by recording the singing activity of one male at a time at dawn and compared features of its dawn song, such as onset before sunrise, repertoire size, mean bout length, strophe length and percentage performance time to responses of the same male to a territory intrusion simulated by playback of synthesised songs later during the same morning. We assume that an aggressive response towards an intruder will involve a fast approach to the loudspeaker broadcasting strophes of blue tit song, searching for the intruder (flying around), and a high amount of counter singing and overlapping of the intruders songs. Aspects of vigour of response to the simulated intrusion could be predicted from all five investigated dawn song parameters as well as male age. This is, to our knowledge, the first indication that a simple receiver could extract reliable information from a males dawn singing behaviour about its competitiveness later in the day.Communicated by P.K. McGregor     

Two explanations of the dawn chorus compared: how monotonically changing light levels favour a short break from singing

I used optimality modelling to compare two of the most plausible and general explanations for the dawn and dusk peaks in bird song output. Kacelnik's explanation is that foraging is inefficient in poor light, but that social interactions are less affected, making singing more worthwhile than foraging. McNamara et al.'s explanation is based on stochasticity in foraging success and overnight energy requirements; it has been extensively analysed with stochastic dynamic programming models. Both explanations are now incorporated into this sort of model. I used various functions to link success of foraging and singing to time of day, but assumed that above some light level there is no further effect. Kacelnik's explanation has as strong an effect as stochasticity in generating dawn and dusk choruses. It also predicts short pauses in the singing output just after the dawn chorus and before the dusk chorus. The former arises because birds delay foraging when it will become more profitable later, until foraging success reaches a plateau, when the energetic debt accumulated makes them forage. The principle of this see-saw double switch in behaviours may apply to other explanations for the dawn chorus, and to other shifts in behaviour when conditions change gradually. The model predicts that from day to day cloud cover determines when a dawn chorus starts, but that overnight temperature and wind strength have more effect on chorus intensity and duration. I discuss what sort of observational and experimental data on singing routines would better test this model. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Light pollution alters the phenology of dawn and dusk singing in common European songbirds

Artificial night lighting is expanding globally, but its ecological consequences remain little understood. Animals often use changes in day length as a cue to time seasonal behaviour. Artificial night lighting may influence the perception of day length, and may thus affect both circadian and circannual rhythms. Over a 3.5 month period, from winter to breeding, we recorded daily singing activity of six common songbird species in 12 woodland sites, half of which were affected by street lighting. We previously reported on analyses suggesting that artificial night lighting affects the daily timing of singing in five species. The main aim of this study was to investigate whether the presence of artificial night lighting is also associated with the seasonal occurrence of dawn and dusk singing. We found that in four species dawn and dusk singing developed earlier in the year at sites exposed to light pollution. We also examined the effects of weather conditions and found that rain and low temperatures negatively affected the occurrence of dawn and dusk singing. Our results support the hypothesis that artificial night lighting alters natural seasonal rhythms, independently of other effects of urbanization. The fitness consequences of the observed changes in seasonal timing of behaviour remain unknown.