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.

Seasonal patterns of singing activity vary with time of day in the nightingale (Luscinia megarhynchos)

Seasonal patterns of singing activity of male birds have been thoroughly studied, but little is known about how those patterns vary with time of day. Here, we censused mated and unmated male Nightingales (Luscinia megarhynchos) at four different hours of the day throughout the breeding cycle. In unmated males, singing activity increased until the young hatched in their neighborhood, and the seasonal variation was similar at each of the four hours of the day. In mated males, however, the seasonal patterns of singing activity differed between hours of the day. In morning (about the hour of egg-laying) and during the dusk chorus, the singing activity of mated males was strongly influenced by the females' reproductive state: singing activity was low before egg-laying and during incubation, but high during the egg-laying period. In the dawn chorus, however, singing activity showed a similar seasonal pattern in mated and unmated males and was high until late stages of the breeding cycle. Our results suggest that the social context influences singing behavior to a varying degree across the season, and that this variation also depends on time of day. The hour of data collection thus is an important but often neglected factor when seasonal changes of singing activity are studied.

On the ability to metaphorize: Talking trees and singing tigers in the English classroom

This article reviews and discusses how metaphor as a trope has been regarded as an essential element in rhetorical approaches to reading and to writing. In addition it considers the extent to which, while metaphor-making is a fundamental cognitive capacity, a metaphorizing habit of mind may be especially pertinent to some aspects of aesthetic activity in English and it has salience also in a multimodal environment. There is exploration of how contemporary practice in the English classroom could accommodate and consolidate the ability to metaphorize.

Experimentally increased noise levels change spatial and singing behaviour

The reasons why animal populations decline in response to anthropogenic noise are still poorly understood. To understand how populations are affected by noise, we must understand how individuals are affected by noise. By modifying the acoustic environment experimentally, we studied the potential relationship between noise levels and both spatial and singing behaviour in the European robin (Erithacus rubecula). We found that with increasing noise levels, males were more likely to move away from the noise source and changed their singing behaviour. Our results provide the first experimental evidence in a free ranging species, that not merely the presence of noise causes changes in behaviour and distribution, but that the level of noise pollution plays a crucial role as well. Our results have important implications for estimating the impact of infrastructure which differs in the level of noise produced. Thus, governmental planning bodies should not only consider the physical effect on the landscape when assessing the impact of new infrastructure, but also the noise levels emitted, which may reduce the loss of suitable habitats available for animals. © 2012 The Author(s) Published by the Royal Society. All rights reserved.

Temporal patterns of territory settlement and detectability in mated and unmated Nightingales Luscinia megarhynchos

In many bird species the sex ratio of adults is male-biased, which is likely to have consequences for the ecology as well as for the conservation of a species. For example, when some males remain unpaired in a population, there should be strong selection on behavioural traits that enhance pairing success. A surplus of males is also likely to have important implications for the interpretation of breeding bird survey data. In our study population of Nightingales Luscinia megarhynchos, about half of the males stayed unpaired, suggesting that the number of males encountered singing was greater than the number of breeding pairs. Furthermore, the detectability (the probability of encountering a male singing) of mated males was only two-thirds that of unmated males when censused in the morning or late in the breeding season. The relative detectability was more similar early in the season and during the twilight periods before sunrise and after sunset. Males that arrived earlier on the breeding grounds were more successful in attracting a mate than males arriving later. Some of the unmated males deserted their territories and prospected areas up to 4000 m distant, whereas others settled on the study site only late in the season and may actually have changed territories. We suggest that adult sex ratios and the time of the census should be taken into account when interpreting the results of breeding bird surveys.

Responses to interactive playback predict future pairing success in nightingales

Birdsong is a sexually selected trait that serves in territory defence and mate choice. Individual song traits can be affected by the body condition of the male and thus may reflect his quality. Such relations between male quality and general singing performance raise the question whether differences in male quality also affect response strategies used in dyadic interactions. To address this question, we studied the relation between pairing success of male common nightingales, Luscinia megarhynchos, and their responses to rivals posing different levels of threat. Using interactive playback, we exposed males prior to mating to either aggressively or moderately singing rivals (by song overlapping and song alternating, respectively). Males that remained unpaired throughout the season (bachelors) interrupted their singing significantly more often after being overlapped than after alternating playback, whereas subsequently mated males kept the number of singing interruptions more constant across playback treatment. This suggests that subsequently paired males are less discriminative than are bachelors when challenged by rivals varying in aggressiveness. Regardless of playback treatment, males that later became paired responded significantly more strongly than did bachelor males. Thus, an increase in singing after a vocal interaction prior to mating predicted future mating success. (c) 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Early birds are sexy: male age, dawn song and extrapair paternity in blue tits, Cyanistes (formerly Parus) caeruleus

Sexual selection theory predicts that signals reflecting the relative quality of individuals should be used in mate choice. Females could base their choice of copulation partners on male secondary sexual traits that honestly signal male age, as predicted by the age-based indicator mechanism. Studies have shown that female blue tits prefer older males and that aspects of dawn song reflect male quality, but it remains unknown whether dawn song characteristics correlate with male age. We compared dawn song characteristics of second-year (SY) and older (ASY) male blue tits (cross-sectional analysis), and tested for age-related changes within individuals (longitudinal analysis) and differential overwinter survival of SY males. We further investigated the relation between dawn song and paternity gain and loss. We found that ASY male blue tits began to sing earlier relative to sunrise than did SY males. This difference in the onset of dawn singing was due to age-related changes in individual performance rather than differential survival of individuals with varying expression of the trait. Males that began to sing earlier at dawn had more mating partners, and were more likely to gain extrapair paternity. Our findings suggest that the onset of dawn song can provide a simple mechanism for females to assess the relative quality of their mate and of neighbouring males. We propose that females use the onset of singing as a cue for their choice of extrapair partners. (c) 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Seasonal variation in dawn song characteristics in the common nightingale

Many male temperate zone passerines show a marked peak of singing activity before sunrise. The two main functions of this so-called dawn chorus are mate attraction and territory defence. We examined how seasonal patterns of different dawn song characteristics were related to mating status and to the breeding cycle of females in the common nightingale, Luscinia megarhynchos. We investigated two measures of song output: song rate and percentage performance time. We also calculated the proportion of 'whistle songs', a song category that is thought to be important in female choice. We predicted that if the main function of dawn singing in nightingales is to attract a social mate, then mated males should change their dawn singing behaviour after pair formation. In contrast, if dawn singing is mainly used in territory defence, we expected no difference in song traits between mated and unmated males throughout the season. We found that song rate and the proportion of whistle songs were low at the beginning of the season and did not predict future mating status. After arrival of females, all measures of dawn song performance remained largely constant throughout the breeding season, and we did not find significant differences in the seasonal variation between mated and unmated males. These findings are consistent with the hypothesis that song at dawn is important to defend a territory throughout the breeding season. (c) 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales

Animal communication often occurs in communication networks in which multiple signalers and receivers are within signaling range of each other. In such networks, individuals can obtain information on the quality and motivation of territorial neighbors by eavesdropping on their signaling interactions. In songbirds, extracting information from interactions involving neighbors is thought to be an important factor in the evolution of strategies of territory defense. In a playback experiment with radio-tagged nightingales Luscinia megarhynchos we here demonstrate that territorial males use their familiar neighbors' performance in a vocal interaction with an unfamiliar intruder as a standard for their own response. Males were attracted by a vocal interaction between their neighbor and a simulated stranger and intruded into the neighbor's territory. The more intensely the neighbor had interacted with playback, the earlier the intrusions were made, indicating that males eavesdropped on the vocal contest involving a neighbor. However, males never intruded when we had simulated by a second playback that the intruder had retreated and sang outside the neighbor's territory. These results suggest that territorial males use their neighbors' singing behavior as an early warning system when territorial integrity is threatened. Simultaneous responses by neighboring males towards unfamiliar rivals are likely to be beneficial to the individuals in maintaining territorial integrity.

The natural plasma testosterone profile of male blue tits during the breeding season and its relation to song output

In male birds, the gonadal hormone testosterone (T) is known to influence territorial and mating behaviour. Plasma levels of T show seasonal fluctuations which vary in relation to mating system and social instability. First, we determined the natural T profile of male blue tits Parus caeruleus during the breeding season. We found that plasma levels of T increased at the onset of nest building. Thus, the increase in circulating T was not associated with territory establishment, nor with the fertile period of the males' mates. In most individuals, T levels dropped to values close to zero during the period of chick feeding. Second, we investigated the relationship between plasma levels of T and male age, size, and singing behaviour. During the mating period, T levels did not differ between 1 yr old and older males and did not correlate with body size or condition. However, song output during the dawn chorus tended to be positively correlated with T levels. Therefore, if high T levels are costly, song output might be an honest indicator of male quality in blue tits. Finally, we show that plasma levels of T are significantly higher during the night than during the day. This pattern has also been observed in captive non-passerine birds, but its functional significance remains unknown.