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.     

Owl dusk chorus is related to the quality of individuals and nest‐sites

Dawn and dusk choruses represent one of the most investigated topics in avian vocal behaviour, but their underlying basis remains unclear. As with the dawn chorus in passerines, dusk chorus in owls seems to support the mate and rival assessment hypothesis and happens during the most constraining period, as individuals have not yet fed and, under the handicap principle, dusk chorus is likely to reveal inter‐individual differences in competitive ability, body condition and/or habitat quality. Here, a study of vocal displays at dusk of 14 Eurasian Eagle Owls Bubo bubo revealed a temporal succession in the order in which males began their vocalizations. The vocalization order appeared to be related to both the quality of the nesting territory (based upon mean number of fledged young and proportion of rats in the diet) and the male's individual quality, as revealed by haematocrit values and the brightness of the white throat patch.     

Lizards speed up visual displays in noisy motion habitats

Extensive research over the last few decades has revealed that many acoustically communicating animals compensate for the masking effect of background noise by changing the structure of their signals. Familiar examples include birds using acoustic properties that enhance the transmission of vocalizations in noisy habitats. Here, we show that the effects of background noise on communication signals are not limited to the acoustic modality, and that visual noise from windblown vegetation has an equally important influence on the production of dynamic visual displays. We found that two species of Puerto Rican lizard, Anolis cristatellus and A. gundlachi, increase the speed of body movements used in territorial signalling to apparently improve communication in visually 'noisy' environments of rapidly moving vegetation. This is the first evidence that animals change how they produce dynamic visual signals when communicating in noisy motion habitats. Taken together with previous work on acoustic communication, our results show that animals with very different sensory ecologies can face similar environmental constraints and adopt remarkably similar strategies to overcome these constraints.     

Seasonal changes in the nocturnal singing routines of Common Nightingales Luscinia megarhynchos

Male Common Nightingales Luscinia megarhynchos famously sing at night. There are two distinct types of nocturnal singing routine (the dusk-to-dawn pattern of variation in song rate): (1) dusk and dawn choruses, with little or no song during the middle of the night; (2) a rapid increase in song rate after dusk, reaching a broad peak in the middle of the night, declining towards dawn, and followed by a dawn chorus. Males sing different nocturnal singing routines at different times in the breeding season. Earlier in the breeding season, most males sing Type 2 routines. Later in the breeding season, most males sing Type 1 routines, as do birds on the wintering grounds. At least some individuals may also sing Type 1 routines during the first few days after their arrival on their breeding territories, before the arrival of females. The main function of nocturnal song appears to be mate attraction. The patterns of variation in song rate over the course of the night are qualitatively similar to those predicted by stochastic dynamic programming (SDP) models of daily singing and foraging routines, for birds that do not forage at night, in circumstances when birds can pair at night (Type 2 routines), and when they cannot (Type 1 routines). The observed seasonal changes in the types of routine sung are also consistent with the predictions of these models.     

Diurnal periodicity of spawning activity of permanently territorial damselfishes (Teleostei: Pomacentridae)

Synopsis Like other congeneric damselfishes, the herbivorous Stegastes altus defends individual feeding territories from heterospecific food competitors, regardless of sex. Females spawned demersal eggs for 31.4 min (n = 25) at the nest in territories of males 0–75 m away from theirs. Throughout the spawning season (June to September), spawning occurred only at dawn, mainly just after sunrise. Daily activity of the fish community showed that potential diurnal food competitors were few or inactive only at dawn and dusk. The frequency of intrusions into the female's territory by heterospecific competitors were as low when she left her territory to spawn at dawn, as they were in the daytime when she defended it. Removal of the female in the daytime resulted in a significant increase in intrusion frequency. An ‘anti-competitor hypothesis’, whereby dawn spawning ensures the food resource in the female's territory seems to explain not only the spawning periodicity in S. altus but also the timing of spawning of other permanently territorial damselfishes. Contribution from the Laboratory of Animal Ecology, Kyoto University, No. 492.     

Cicada acoustic communication: potential sound partitioning in a multispecies community from Mexico (Hemiptera: Cicadomorpha: Cicadidae)

Multispecies cicada communities in neotropical rainforests produce a complex and intense acoustic environment. In a fragment of a Mexican rainforest (Veracruz, Mexico), a cicada community at the end of the dry season consisted of nine species ( Daza montezuma; Pacarina schumanni; Miranha imbellis; Dorisiana sutori; Fidicinoides picea; Fidicinoides pronoe; Quesada gigas; one species of the genus Neocicada and one uncaught canopy species). Seven of the nine species formed dense choruses at dawn and at dusk. Each species showed preferences in the height of calling sites. Males of the species were solitary or gregarious, and followed a 'call-fly' or a 'call-stay' calling strategy. Acoustic signals of each species had particular time and frequency patterns. All these specific features appear to separate the nine species acoustically and lead to a partitioning of the acoustic environment. The acoustic partitioning might decrease the risk of heterospecific courting and mating.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 379–394.     

Duet function in the yellow-naped amazon, Amazona auropalliata: evidence from playbacks of duets and solos

The question of why animals participate in duets is an intriguing one, as many such displays appear to be more costly to produce than individual signals. Mated pairs of yellow-naped amazons, Amazona auropalliata, give duets on their nesting territories. We investigated the function of those duets with a playback experiment. We tested two hypotheses for the function of those duets: the joint territory defense hypothesis and the mate-guarding hypothesis, by presenting territorial pairs with three types of playback treatments: duets, male solos, and female solos. The joint territory defense hypothesis suggests that individuals engage in duets because they appear more threatening than solos and are thus more effective for the establishment, maintenance and/or defense of territories. It predicts that pairs will be coordinated in their response (pair members approach speakers and vocalize together) and will either respond more strongly (more calls and/or more movement) to duet treatments than to solo treatments, or respond equally to all treatments. Alternatively, the mate-guarding hypothesis suggests that individuals participate in duets because they allow them to acoustically guard their mate, and predicts uncoordinated responses by pairs, with weak responses to duet treatments and stronger responses by individuals to solos produced by the same sex. Yellow-naped amazon pairs responded to all treatments in an equivalently aggressive and coordinated manner by rapidly approaching speakers and vocalizing more. These responses generally support the joint territory defense hypothesis and further suggest that all intruders are viewed as a threat by resident pairs.     

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.     

A rain forest dusk chorus: cacophony or sounds of silence?

A rain forest dusk chorus consists of a large number of individuals of acoustically communicating species signaling at the same time. How different species achieve effective intra-specific communication in this complex and noisy acoustic environment is not well understood. In this study we examined acoustic masking interference in an assemblage of rain forest crickets and katydids. We used signal structures and spacing of signalers to estimate temporal, spectral and active space overlap between species. We then examined these overlaps for evidence of strategies of masking avoidance in the assemblage: we asked whether species whose signals have high temporal or spectral overlap avoid calling together. Whereas we found evidence that species with high temporal overlap may avoid calling together, there was no relation between spectral overlap and calling activity. There was also no correlation between the spectral and temporal overlaps of the signals of different species. In addition, we found little evidence that species calling in the understorey actively use spacing to minimize acoustic overlap. Increasing call intensity and tuning receivers however emerged as powerful strategies to minimize acoustic overlap. Effective acoustic overlaps were on average close to zero for most individuals in natural, multispecies choruses, even in the absence of behavioral avoidance mechanisms such as inhibition of calling or active spacing. Thus, call temporal structure, intensity and frequency together provide sufficient parameter space for several species to call together yet communicate effectively with little interference in the apparent cacophony of a rain forest dusk chorus.     

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.     

Differential effects of moving versus stationary territorial intruders on territory defence in a songbird

1.  In territorial contests, not only acoustic or other signals, but also the movements of a territorial intruder are likely to influence the response of a resident.
2.  We tested this movement hypothesis by simulating moving vs. stationary intruders into the territories of winter wrens Troglodytes troglodytes , using the same non-interactive song playbacks in both treatments.
3.  Male winter wrens showed a different long-term singing reaction in response to a moving than to a stationary intruder.
4.  One day after experiencing an intruder that was switching between three locations, residents started to sing earlier before sunrise, and they sang more and longer songs at dawn than before the intrusion.
5.  Residents receiving the same playback from one location only reacted by starting to sing later relative to sunrise, and by singing fewer and shorter songs than before the intrusion.
6.  We could not discriminate between the treatments when examining the short-term singing reactions during and immediately after the playbacks. However, our results clearly demonstrate an effect of the spatial behaviour of territorial intruders on the long-term territory defence of residents at dawn, about 24 h after an intrusion.
7.  We argue that spatial behaviour of territorial intruders should be an integral part of the study of animal territory defence behaviour. Investigating long-term changes in territory defence at dawn is a sensitive tool for discriminating between different types of intruders.     

The evolution of alternative adaptive strategies for effective communication in noisy environments

Animals communicating socially are expected to produce signals that are conspicuous within the habitats in which they live. The particular way in which a species adapts to its environment will depend on its ancestral condition and evolutionary history. At this point, it is unclear how properties of the environment and historical factors interact to shape communication. Tropical Anolis lizards advertise territorial ownership using visual displays in habitats where visual motion or "noise" from windblown vegetation poses an acute problem for the detection of display movements. We studied eight Anolis species that live in similar noise environments but belong to separate island radiations with divergent evolutionary histories. We found that species on Puerto Rico displayed at times when their signals were more likely to be detected by neighboring males and females (during periods of low noise). In contrast, species on Jamaica displayed irrespective of the level of environmental motion, apparently because these species have a display that is effective in a range of viewing conditions. Our findings appear to reflect a case of species originating from different evolutionary starting points evolving different signal strategies for effective communication in noisy environments.     

Roaring rhythms in black howler monkeys (Alouatta pigra) of Belize

Roaring was recorded during 15-min intervals for 55 days from April 29 to July 14, 1981 in the area around Bermuda Landing, Belize. Roaring profiles were compiled to represent four three-week periods in which roaring was graphed against the time of day. Rates of roaring were also calculated for three troops. Results indicate a gradual change in the daily profile of roaring from dry to rainy season. The marked bimodality of roaring at dawn and dusk in the dry season changed to a more dampened bimodality with reduced dawn and dusk peaks and more roaring during the midday in the rainy season. This seems related to the decreased hours of sun and the amount of rain during the wet season. Additionally, literature on monkeys with long range vocalizations suggests that bimodal peaks in calling may be more prevalent in species which display territorial defense.     

Natural entrainment without dawn and dusk: the case of the European ground squirrel (Spermophilus citellus).

Observational data collected in the field and in enclosures show that diurnal, burrow-dwelling European ground squirrels (Spermophilus citellus) never were above ground during twilight at dawn or at dusk. The animals emerged on average 4.02 h (SD = 0.45) after civil twilight at dawn and retreated in their burrows on average 2.87 h (SD = 0.47) before civil twilight at dusk. Daily patterns of light perceived by these burrowing mammals were measured with light-sensitive radio collar transmitters in an enclosure (the Netherlands) and in the field (Hungary). The observational data are corroborated by the telemetry data, which show clear daily patterns of timing of light perception including light perceived from the burrow entrances. The first light was observed by the animals on average 3.54 h (enclosure, SD = 0.45) and 3.60 h (field, SD = 0.31) after civil twilight at dawn, whereas the final observed light was on average 3.04 h (enclosure, SD = 0.64) and 2.02 h (field, SD = 0.72) before civil twilight at dusk. Thus, the animals do not perceive the rapid natural light-dark (LD) transitions that occur at civil twilight. Instead, they generate their own pattern of exposure to light within the natural LD cycle. The classical phase response model for entrainment by light or dark pulses cannot explain how the circadian system of this species remains entrained to the external, natural LD cycle while the major LD transitions are created by its own behavior.     

Group chorusing as an intragroup signal in the greater ani,a communally breeding bird

Group living often requires maintaining dynamic and varied relationships with fellow group members, while simultaneously monitoring and interacting with external competitors. Group members in many social species vocalize together to produce duets or choruses—coordinated, often conspicuous vocal displays—that may play a role in these interactions. Compared with male–female duets, however, relatively little research exists on the function and adaptive significance of group choruses, which involve three or more individuals. Here we investigate chorusing behavior in the greater ani (Crotophaga major), a communally breeding cuckoo that nests in stable social groups of four to eight unrelated individuals. Groups may remain together for several years on the same nesting territory, and groups occasionally destroy each other's clutches in conflicts over high-quality territories. We asked whether the raucous, highly stereotyped choruses performed by ani groups are primarily used for intra- or intergroup communication, and whether they contain information about the identity of the social group and the number of birds vocalizing. Behavioral observations and acoustic recordings from three breeding seasons revealed that choruses typically occurred during social interactions within the group (78% of choruses) or in response to a predator or extra-group individual (17%) and only rarely in intergroup interactions (4%). Consistent with this finding, choruses did not reliably reflect the number of birds vocalizing, and we found only limited evidence for group-specific acoustic signatures (driven by a single group whose choruses were highly distinct). These results suggest that group choruses play an important role in intra-group signaling, potentially in contexts such as group formation, reinforcement of social bonds within the group, and/or collective decision-making, and they motivate new research questions about the role of collective signaling in social evolution.     

Context Dependent Territory Defense: The Importance of Habitat Structure in Anolis sagrei

Territoriality is a potentially costly endeavor, and several mechanisms for mitigating the costs of territoriality have been investigated in the wild. For example, territory owners can reduce the costs of defending territory boundaries by prioritizing defense of the most valuable areas within territories, investing less energy in low quality areas. We staged pairwise encounters between adult male lizards on natural territories in the wild, to test whether male brown anoles, Anolis sagrei, would differentially defend certain regions of their territories over others. Based on our observations that male A. sagrei spend most of their time on elevated perches on tree trunks or branches compared with sites on the ground, we predicted that territory residents would respond more aggressively to territory invasions that took place on elevated perches than to invasions on the ground. We measured significant differences in the behavior of residents following invasion on the ground vs. on the elevated perches, and results partially supported our hypothesis. Males performed more displays and approached intruders more often when territory invasion took place on the ground, but were quicker to attack intruders that entered territories on elevated perches. Our hypothesis was only partially supported, potentially indicating that elevated perches are preferred as outposts to monitor valuable sites on the ground. Our study provides evidence that territory defense varies not just among individuals, but also within individuals at different locations in a territory.     

Matched Counter-singing Signals Escalation of Aggression in Black-capped Chickadees (Parus atricapillus)

Black-capped chickadees (Parus atricapillus) have the ability to shift their songs up and down over a wide range of absolute frequencies. Males can shift their songs over 465 ± 52.9 (SE) Hz. During the dawn chorus, males shift their songs by 80 Hz or more every 41 ± 8.8 (SE) songs, but it appears that males can sing at any frequency within their range. Frequency shifting may allow males to match counter-sing with rival males; that is, to switch song output to match that of a rival. During simultaneously recorded dawn choruses, however, there was no correlation over time in the frequency of neighbouring males' songs, nor was there a correlation over time in the size of shifts between their songs. Moreover, males did not match the frequencies of songs presented on a played-back tape at the edge of their territories during the dawn chorus. Matching was observed during some bouts of counter-singing between males. In these cases, matched counter-singing was highly associated with escalation of the conflict. We suggest that frequency matching in this species may be a graded signal that allows the singer to direct aggression towards a particular rival.     

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