INCOMPLETE SONG STROPHES IN THE CHAFFINCH FRINGILLA COELEBS L.: GENERAL INFLUENCES ON A SPECIFIC BEHAVIOURAL OUTPUT

ABSTRACT

Incomplete song strophes in free-living territorial Chaffinch males can be induced by different experimental as well as natural stimulus situations including replay of species- specific song, approaching of human beings, and aggressive encounters with con-specific males. While the first post-stimulus song strophe is shortest the following ones gradually attain their full number of elements again. The strength of this reaction differs with regard to different stimuli.     

INDIVIDUAL AND SPECIES-SPECIFIC SONG PATTERNS OF ROCK AND MEADOW PIPITS: PHYSICAL CHARACTERISTICS AND EXPERIMENTS

ABSTRACT

The songs of Anthus spinoletta littoralis and A. pratensis, in sympatric populations in SW Sweden were recorded, analysed in sound spectrographs and tested in the field. The purpose was to investigate: a) if the strophes are species specifically and individually distinct and, if so, whether pipits are able to make such distinctions; b) if the species-specific and individual-specific patterns are concentrated in different sections of the song and, if so, whether the species-specific section transmits individual-specific information and the individual-specific section transmits species-specific information as well.

The basic structure of the syllables in the terminating section of the strophe was found to be intraspecifically stereotyped, but varied between the two species. Replay experiments showed that territorial males could discriminate between con- and heterospecific strophes and this ability persisted when the terminating section of the strophe was replayed alone. The terminating part of the strophe did not seem to transmit individual-specific information to conspecific males. On the other hand, the basic structure of the syllables in the first section of the strophe was intraindividually stereotyped but varied between most conspecific males. Playback experiments showed that a territorial male could discriminate his neighbours' song patterns and this ability remained intact when the introductory phrase of the strophe was replayed alone. Both song pattern and position were necessary properties of the identifying signal in these pipits. But the introductory phrase of a total stranger did not seem to transmit unambiguous species-specific information to conspecific males.     

Attraction of Hoopoe Upupa epops females and males by means of song playback in the field: influence of strophe length

Experimental Hoopoe Upupa epops songs prepared with synthetic sound to differ in strophe length were used to test whether Hoopoe females prefer long strophes. The songs were broadcast simultaneously in the field from two loudspeakers situated 100 m apart, in early spring, when females actively search for mates. The playbacks attracted a total of 87 individuals, both males and females, with a maximum of five individuals (three males and two females) per trial. A female was considered to have been attracted by the song broadcast from a loudspeaker when she approached unaccompanied by a male, and when there was no male nearby. A total of 15 females chose one of the songs and significantly more were attracted by the one with long strophes. These results suggest that in the Hoopoe, male song attracts females, and that strophe length is a sexually selected song cue. There were no differences in the number or kind of males (classified according to strophe length) attracted by each playback. Frequently the experiment attracted more than one male simultaneously. These aggregations cannot be explained as territorial responses, and their significance is discussed together with that of natural spontaneous groupings of displaying males. Apparently males aggregate where they expect to find females.     

Strophe Length in Spontaneous Songs Predicts Male Response to Playback in the Hoopoe Upupa epops

Hoopoe (Upupa epops, Coraciformes) males produce a very simple song during the breeding season in order to attract females and repel intruders. Strophes vary in length (i.e. number of elements) both within and between males, and previous studies have shown that this song cue is positively correlated with male condition and breeding success. In the present study we tested whether strophe length of males influences male behaviour during intra‐sexual contests, in a colour‐ringed population in southeast Spain. Paired males were presented with a recorded song with long strophes during the pre‐laying period, while they were near their mates, in order to provoke male mate‐defence behaviour. Most males responded to the playback, but the strategy of defence adopted depended on their own strophe length in spontaneous songs recorded before the experiments. While singing responses were common to most of the males, only those using long strophes adopted the most risky strategy of approaching the loudspeaker. However, the males that approached produced abnormal songs during playback, that were shorter and with fewer strophes than those of males that did not approach, and used shorter strophes in comparison with spontaneous songs before the experiment. These differences in quality of the song produced in response to the playback suggest that long‐strophe males were basing their response mainly on attacking rather than singing, while short‐strophe males tried to resolve the contest at a distance by means of their song. These results show that strophe length reflects some component of the competitive ability of males (either physical strength or aggressiveness) in the hoopoe, which together with previous results regarding its role for female choice, show that it is a sexual signal with dual function.     

Song structure in the Hoopoe (Upupa epops) — Strophe length reflects male condition

Summary The structure of Hoopoe (Upupa epops) song is analysed in a colour ringed population in southern Spain. The song of males in this species is very simple, with a repertoire size of one. The strophes of a male only differ in the number of elements that they include (strophe length), and strophe length is the main song feature differing between males. During the prelaying period each individual used mainly strophes of only two lengths, between 2 and 6 elements per strophe, and mean strophe length of males during this period was highly repeatable. However, some males changed the range of strophe types produced and decreased their mean strophe length after unsuccessful breeding or spending long periods of time singing (unpaired males). These changes show that strophe length is phenotypically plastic, and suggest that singing long strophes is energetically more costly than singing short ones. The significant relationship between strophe length and body condition, and the fact that long strophes were associated with longer previous pauses than short strophes, also suggest that increasing strophe length is costly. All these findings are in accordance with the hypothesis that strophe length reflects male condition in the Hoopoe, although it is not clear what the actual cost of singing long strophes is.

---

Struktur des Gesangs des Wiedehopfs (Upupa epops) — Strophenlänge reflektiert Männchen-Qualität

Zusammenfassung Die Struktur des Gesangs des Wiedehopfs wurde in einer farbberingten südspanischen Population untersucht. Der Gesang des Männchens ist sehr einfach und umfaßt nur ein Repertoire. Die Strophen eines Männchens differieren nur in der Anzahl Elemente (Strophenlänge), und die Männchen unterscheiden sich vor allem in der Strophenlänge voneinander. In der Vorbrutphase verwendeten die Männchen meist nur zwei verschieden lange Strophen, bestehend aus 2 und 6 Elementen je Strophe, und die durchschnittliche Strophenlänge war in dieser Phase sehr konstant. Nach Brutverlust oder bei kontinuierlich singenden, ledigen Männchen kam es zu einer Verkürzung der mittleren Strophenlänge. Die Strophenlänge ist phänotypisch plastisch, und lange Strophen zu singen scheint energetisch aufwendiger als kurze Strophen. Der enge Zusammenhang zwischen Strophenlänge und männlicher Konstitution einerseits und die Beobachtung, daß lange Strophen mit langen Pausen korrelieren andererseits, zeigen ebenfalls, daß längere Strophen energetisch aufwendiger sind. Die wirklichen Kosten dafür sind jedoch noch nicht bekannt.

     

Song complexity and song structure in the Moustached Warbler Acrocephalus melanopogon

We examined different song parameters leading to a complex song configuration, and song output (production), and their relation in male Moustached Warblers and discuss them with reference to female choice. With more than 120 different syllables per two minutes of song, male Moustached Warblers can be regarded as one of the most complex singers within the genus Acrocephalus . We found significant differences between males for almost all song parameters investigated (repertoire size, switching and repetition rate, song speed and strophe length) but not for time spent singing per 20 minutes. This individual variation probably is an important cue for female mate choice. Male Moustached Warblers achieve song complexity mainly via frequent syllable switching. Song complexity seems to be consistent throughout the song and our results suggest that females could assess the overall song complexity of a singer by examining even very short song bouts. Furthermore, a high proportion of motifs – clusters of syllables with the same sequence – occurs within the song of each male which would also facilitate assessment of song complexity by females. However, repertoire size and strophe length were positively related, thus it remains to be investigated which song feature, if any, females actually use.     

AUDITORY EXPERIENCE AND SONG DEVELOPMENT IN THE CHAFFINCH FRINGILLA COELEBS

The present paper describes the results of experiments designed to evaluate the importance of auditory-motor experience in the development of song in the Chaffinch. For this purpose a number of Chaffinches collected as nestlings or adults were deafened by extirpating the cochlea at different ages. Recordings of their song made subsequently provide the material on which the paper is based. Since the results of the experiments can only be interpreted in the light of knowledge of normal song development, the latter is described from the observation of previous workers and from some new material. Of the 20 males deafened, only three came into full song of their own accord; song was induced in the rest by the administration of testosterone phenylacetate or testosterone propionate. Songs of abnormal length were common among first-year males though males deafened when two or more years old produced song of the normal length. The song of male Chaffinches deafened as adults, that is after they had produced full song for one or more breeding seasons, was infistinguishable from that produced by intact adult males. They even went through a “plastic” song stage in which some “intermediate” themes occurred. A first-year male deafened when it had almost stabilized its song produced song that gradually deviated more and more from normal song. This indicates that the bird had not yet laid down a long-term memory of its song independent of auditory feedback. First-year male Chaffinches deafened during early spring, mid-winter and late sumer, respectively, show that the more they were deprived of the normal period of auditory-motor experience, the simpler was the song they produced in their first spring. Thus, at the time of deafening they had acquired a particular auditory-motor experience, and it is the type of element established up to that time that is incorporated into full song. Male chaffinches brought into the laboratory as nestlings and deafened when three months old produced a virtually sfyctureless song, in which the only recognizable element was a sound reminiscent of the juvenile “chirrup”. The subsong and other calls of these birds were also highly abnormal. Song development in the Chaffinch clearly starts before song as such has made its appearance. The available evidence suggests that the kind of song a male Chaffinch produces depends directly on its auditory-motor experience before deafening. There is no indication that deafening achieves its effects on the resultant song by interfering with the hormonal substrate. Relatively stable song end patterns develop in the absence of auditory feedback, so their occurrence does not necessitate the postulation of templates or preconceived patterns. Song development differs markedly from species to species.     

Effects of an Immune Challenge on Multiple Components of Song Display in Barn Swallows Hirundo rustica: Implications for Sexual Selection

Bird song is believed to honestly reveal male quality including the ability of singers to face parasitism. In a natural population of barn swallows Hirundo rustica, we experimentally imposed a cost on song production by an immune challenge. We therefore vaccinated a group of reproducing males with an antigen (Newcastle disease virus), and injected phosphate‐buffered saline to a control group. Immune challenge significantly reduced one song feature, rattle duration. This decrease was related to male quality, as measured by tail length, because males with short tails reduced the duration of their rattle significantly more than males with long tails. In addition, another song feature, strophe duration, decreased in the control group, while it remained constant in the challenged group. Duration of the rattle has previously been found to be positively related to testosterone level, and it may hence reflect male competitive ability. Thus, male barn swallows may not have the potential to produce long rattles when their immune system is challenged. By maintaining their strophe duration after an immune challenge, males may compensate for the decrease in rattle duration. Our results suggest that different song features may convey different types of information, and that barn swallow song, in particular their rattle, may reliably reveal information about activation of the immune system.     

The Dawn Song of the Blue Tit Parus caeruleus and its Role in Sexual Selection

Sexual selection arises when variance in male reproductive success is non-randomly related to phenotypic characters of males. Song can be considered as such a phenotypic character and several studies have shown that song complexity and/or song output are important in competition among males or in partner choice. In the blue tit Parus caeruleus a peak in male singing activity occurs at dawn during the female fertile period, i.e. after pair formation. The function of this dawn chorus is not well understood. In this study 20 male blue tits were recorded at dawn and song complexity and output were expressed as versatility, mean strophe length, mean percentage performance time and bouts with or without drift, i.e. with or without a systematic decline in percentage performance time. Females mated to males with a higher mean percentage performance time (output) and a higher versatility (complexity) started to lay eggs earlier, but the latter was not significant. Females mated to males that showed no drift in their song bouts laid significantly larger clutches. Our results thus suggest that in the blue tit, song output at dawn, rather than song complexity, might be a trait under sexual selection.     

ANNUAL CHANGES OF LEARNED BEHAVIOUR—VARIATION OF SONG PATTERN IN FREE-LIVING CHAFFINCHES,FRINGILLA COELEBS,DURING THE BREEDING SEASON

ABSTRACT

Seasonal changes of parameters of full song were studied in a free-living population of chaffinches Fringilla coelebs during one entire reproductive period. Approximately 7000 strophes sung by 14 male chaffinches were recorded and analysed by sonography and a particular oscillographic method. While the general pattern of song strophes, i.e. characteristics of elements, number and arrangement of phrases, and final flourish, remained constant throughout the reproductive period, full song varied with respect to the repetition rate of strophes, number of strophe types used, intensity of singing, duration of strophes, and percentage of incomplete strophes sung. These changes are discussed as results of learning processes, social interactions in the population, and endogenous mechanisms activating memorized information.     

Male quality and territory quality in the great tit parus major

In this study there was no positive significant relationship between the quality of male great tits, as measured by average strophe length (a strophe is a short burst of song), and territory quality. (1) Males singing longer strophes did not defend territories in areas of higher recruitment, and (2) naturally settling males were better singers than replacement birds were. The second result was probably caused by a difference in age composition of the two groups, because naturally settling yearlings and replacement yearlings did not differ significantly in average strophe length. These observations on a non-migratory species are in contrast to those of several other studies on passerine birds, which were mostly concerned with migratory species. The relationship between male quality and territory quality is predicted to be closer in species that migrate than in non-migratory species.     

Extrapair paternity in the Hoopoe Upupa epops: an exploration of the influence of interactions between breeding pairs, non-pair males and strophe length

Previous studies of the Hoopoe Upupa epops have shown that the strophe length of male songs influences female mate choice, and is correlated with female reproductive rates and male production of fledglings in the male’s own brood. However, frequent interactions between breeding pairs and non‐pair males suggests that extrapair copulations could occur and could affect the real number of fledglings sired by males, and therefore the relationship between strophe length and breeding success. Here we analyse the incidence of interactions between breeding pairs and non‐pair males, and of extrapair paternity, the interrelation of these parameters, the influence of male strophe length on them, and whether extrapair fertilizations affect the correlation between strophe length and breeding success of males, in a colour‐ringed population of Hoopoes in south‐eastern Spain. Multilocus DNA‐fingerprinting revealed that 10% of the broods contained offspring sired by extrapair males, representing 7.7% of the chicks. However, the interactions between pairs and non‐pair males were more frequent, with more than 25% of broods being visited by non‐pair males, and about 10% being helped (fed or defended) by males other than the nest owner. Most of these relationships were apparently attempts by visitor males to obtain copulations with paired females, or to obtain access to such females or nests in future breeding attempts. However, there was no significant link between the detection of interactions with alien males in a nest and the occurrence of extrapair paternity in it, indeed extrapair paternity was found in only 30% of the nests with interactions, and therefore the detection of visits or helping by non‐pair males cannot be considered evidence of extrapair paternity in visited or helped broods. Males that sang with long strophes never suffered losses of paternity within their broods, while 25% of males that sang with short strophes did. However, these differences were not significant. Nevertheless, strophe length of males was significantly positively correlated with per brood and seasonal production of fledglings after accounting for losses of paternity within their own broods.     

SONG POPULATIONS AND DISPERSAL IN STEELBLUE AND PURPLE WIDOWFINCHES

Payne, R. B. 1985. Song populations and dispersal in Steelblue and Purple Widowfinches. Ostrich 56:135-146.

Songs of Steelblue Widowfinches Vidua chalybeata were recorded over eight years at Lochinvar National Park, Zambia. Neighbouring males share their song-type repertoires. Individual birds changed the structural details of all song types in their repertoire from year to year, and over 4–5 years the songs accumulated changes so that a song type could scarcely be recognized as the same. A few males in addition switched their song repertoires when they dispersed from one song population to another, or when their old neighbours disappeared and new neighbouring males sang a different song repertoire. Individual marked adult birds moved from one song neighbourhood to another. Purple Widowfinches had a similar song behaviour but only one song dialect was found in the study area.

An estimate of the proportion of immigrants was made from the proportion of birds with songs unlike their neighbours, and from birds that had a mixed song repertoire or that switched their song repertoires. Song populations were sampled at several localities in the Transvaal, Botswana, Zambia, Kenya, Tanzania, Cameroon, and Nigeria. An overall minimal estimate of dispersal between song neighbourhoods was 18 %. The incidence of observed dispersal and the movements indicated by the song differences among local widowfinches were considerable and more than sufficient to prevent genetic isoloation and differentiation of local song populations.     

NATURAL SELECTION ON BEAK AND BODY SIZE IN THE SONG SPARROW

We documented temporal patterns of natural selection on beak and body traits in a song sparrow population. We looked for evidence of selection in association with reproduction and overwinter survival in order to identify the conditions under which size in beak and body traits is adaptive. We also attempted to identify the specific traits most closely associated with fitness under these conditions. Selection was observed in association with both survival and reproduction. Patterns of selection differed between the sexes. Selection on males was weak and stabilizing in association with overwinter survival. Selection on females was strong, was both stabilizing and directional, and was associated with both survival and reproduction. In females, traits that enhanced juvenile survival also reduced reproductive success; i.e., there was a trade-off between survival and reproduction. Patterns of selection in the song sparrow parallel those reported for the Galápagos finch, Geospiza fortis. However, in song sparrows, selection occurred mainly on tarsus length and beak length, and not on beak depth or width as in G. fortis. This difference may occur because most North American sparrows partition food resources by habitat, while most Galápagos ground finches partition food by seed size.     

The anti-exhaustion hypothesis: a new hypothesis to explain song performance and song switching in the great tit

This paper describes the results of a detailed analysis of 52 song bouts, recorded from 22 great tits, Parus major, during the dawn chorus. A song bout consists of a number of song bursts (called strophes) separated by periods of silence. High quality males, as measured by average strophe length, sang their bouts with a higher percentage performance time (i.e. the percentage of time spent singing in a bout), but the average number of strophes per bout was not related to male quality. In 31 of 52 bouts there was a systematic decrease in the percentage performance time throughout the bout. This was mainly caused by a prolongation of the pauses between the strophes, and sometimes by a shortening of the strophes. Both high and low quality males sang bouts with and without this decrease in the percentage performance time. Bouts that started with longer strophes and/or shorter inter-strophe pauses showed on average a more rapid decrease in the percentage performance time, and contained fewer strophes, than bouts that started with shorter strophes and/or longer inter-strophe pauses. After switching to another song type the males again used longer strophes and/or shorter inter-strophe pauses. An ‘anti-exhaustion’ hypothesis is proposed and discussed. This hypothesis gives a mainly causal explanation for the existence of song switching and song repertoires in passerine birds.     

青海门源赭红尾鸲鸣唱方言的时间稳定性

鸟类的鸣唱方言是研究非人类文化进化及其与种群遗传分化相互作用的理想素材。该研究通过比较跨度8年的2001年和2009年青海门源赭红尾鸲(Phoenicurus ochruros rufiventris)鸣唱,研究结果显示,青海门源赭红尾鸲的种特异性音节、经典唱段中的部分音节、典型唱段结构和鸣唱方言的基本形式都具有较高的时间稳定性。2009年赭红尾鸲唱段曲目多于2001年,共享唱段出现了一定的分化,唱段的共享程度和鸣唱的相似性与个体领域间的距离相关,相邻个体的鸣唱相似性程度高于较远距离的个体。     

Mate choice in Pied Flycatchers Ficedula hypoleuca: can females use song to find high‐quality males and territories?

High-quality male Pied Flycatchers Ficedula hypoleuca (defined by brighter plumage, better condition and more experience) have previously been shown to have larger syllable repertoires and greater song versatility than males of inferior quality. Thus, by preferring more complex songs, females could choose a high-quality male. Females may also use song as a cue to find a high-quality territory since early arriving males may obtain the best territories and these males have more complex songs than late-arriving males. We found that males with more complex songs had a greater chance of becoming paired and stayed unpaired for a shorter period than males with less elaborate songs. When controlling for arrival order, however, only strophe versatility was still correlated with pairing order. Males defending popular territories had more complex and longer songs and were also in better body condition than males in less popular territories. A multiple logistic regression showed that song length was important in explaining whether a male defended a popular nestbox or not. Thus, male arrival time seems to be important in deciding the quality of a male's territory, which in turn explains female choice. However, song quality seems to add important information. Thus, females could find both high-quality males and high-quality territories by using song cues during mate choice.     

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     

SONG REPERTOIRES AND REPERTOIRE SHARING IN A LOCAL GROUP OF BLACKBIRDS

ABSTRACT

The song repertoires of a local group of blackbird Turdus merula males were determined and quantified with respect to individuality and sharing between neighbours. We focused on the long-ranging, introductory whistle part of the song and its motifs of fixed sequences of sounds. These motifs, whether they are used to start the song or placed centrally in the song, constitute the largest units of song that are repeated in the same way every time they are sung. Blackbird motifs therefore are equivalents of song types in other species. The result shows that one needs to analyse more than 200 songs to estimate a male's repertoire, which averaged 44 different motifs. Relative to other species, this is a medium to large song repertoire. The size of the repertoire of start motifs (on average 32) varied only a little between the individuals, whereas that of central motifs (on average 12) varied rather much between the same individuals, suggesting that they form a potential cue for assessment of male quality. The males within the neighbourhood showed a high degree of start motif sharing, which, together with the relatively large repertoires, could be constraining neighbour recognition. However, in most males the possession of a few individually distinctive and frequently repeated motifs could counter such an effect.