Neighbour-stranger discrimination in an African wood dove inhabiting equatorial rainforest.

We investigated within- and between-individual song variation and song-based neighbour-stranger discrimination in a non-learning bird species, the blue-headed wood-dove (Turtur brehmeri), which inhabits lowland rainforests of West and Central Africa. We found that songs of this species are individually specific and have a high potential for use in individual recognition based on the time-frequency pattern of note distribution within song phrases. To test whether these differences affect behaviour, we conducted playback experiments with 19 territorial males. Each male was tested twice, once with the songs of a familiar neighbour and once with the songs of an unfamiliar stranger. We observed that males responded more aggressively to playback of a stranger's songs: they quickly approached close to the speaker and spent more time near it. However, no significant differences between treatments were observed in the vocal responses. In addition, we explored whether responses differed based on the song frequency of the focal male and/or that of the simulated intruder (i.e., playback), as this song parameter is inversely related to body size and could potentially affect males' decisions to respond to other birds. Song frequency parameters (of either the focal male or the simulated intruder) had no effect on the approaching response during playback. However, we found that the pattern of response after playback was significantly affected by the song frequency of the focal male: males with lower-frequency songs stayed closer to the simulated intruder for a longer period of time without singing, while males with higher-frequency songs returned more quickly to their initial song posts and resumed singing. Together, these results depict a consistently strong response to strangers during and after playback that is dependent on a male's self-assessment rather than assessment of a rival's strength based on his song frequency. This work provides the first experimental evidence that doves (Columbidae) can use songs for neighbour-stranger discrimination and respond according to a "dear enemy" scheme that keeps the cost of territory defence at a reasonable level.

Detection ranges of forest bird vocalisations: guidelines for passive acoustic monitoring.

Passive acoustic monitoring has proven to have many advantages for monitoring efforts and research activities. However, there are considerations to be taken into account regarding the placement of autonomous sound recorders. Detection ranges differ among species and in response to variable conditions such as weather or the location of vocalising animals. It is thus important to the success of a research project to understand, with a certain degree of confidence, the distances at which birds might be detected. In two types of forests in Poland, we played back the vocalisations of 31 species of European forest birds exemplifying different singing characteristics. Based on recordings obtained along a 500-m transect, we estimated the probability of detection and maximum detection distance of each vocalisation. We broadcasted the recording at three heights of singing and repeated playbacks three times during the breeding season to evaluate the effect of vegetation growth. Our results revealed that environmental and meteorological factors had a significant influence on both detection probability and maximum detection distances. This work provides comprehensive measurements of detection distance for 31 bird species and can be used to plan passive acoustic monitoring research in Europe, taking into account species traits and individual characteristics of the study area.

Response of forest Turtur doves to conspecific and congeneric songs in sympatry and allopatry.

Birds have a diverse acoustic communication system, and the ability to recognise their own species' song from a distance facilitates complex behaviours related to mate attraction and rival deterrence. However, certain species, including doves, do not learn songs and their vocal repertoires are much simpler than those of better-studied songbirds. In these so-called non-learning birds, relatively little is known about the role that bird song plays in intra- and interspecific interactions, and how such behaviours might be acquired (inherited or learned from experience). To investigate this question, we focused on two species of African wood doves whose long-range songs are used in a territorial context. Specifically, we examined the responses of sympatric and allopatric populations of male blue-headed wood-doves (Turtur brehmeri) and tambourine doves (Turtur tympanistria) to different types of simulated territorial intrusions, i.e. playback of conspecific, congeneric, and control songs. We aimed to assess (i) whether these species, which have similar songs, respond only to their own species' song or exhibit interspecific territoriality, and (ii) if the response pattern is affected by the presence or absence of congeners in the general area. We found that both species responded strongly to playback of their own species in both sympatric and allopatric populations. In allopatry, though, male tambourine doves misdirected their response and also approached the playback of congeneric songs. Our results indicate that, in areas where the studied Turtur doves live in sympatry, they do not exhibit consistent interspecific territoriality. However, we cannot exclude the possibility that the smaller tambourine dove avoids its larger congener during the process of territory establishment. The difference in tambourine doves' response toward the song of present (sympatric) or absent (allopatric) congeners suggests that the ability to discriminate between songs of similarly singing potential competitors is acquired through earlier interactions and learning. This plasticity in response supports the misdirected aggression hypothesis, which argues that interspecific territorialism emerges as a maladaptive by-product of signal similarity. However, on an evolutionary timescale, such an ability could be considered an adaptive cognitive tool useful for resolving competing interests with congeners.

Male song of the Aquatic Warbler, a promiscuous bird without paternal care, is more complex than previously thought.

The Aquatic Warbler Acrocephalus paludicola is one of the rarest European passerines and is characterised by promiscuity, lack of pair-bonds and female-only parental care. This makes the species an important model for studying the function of avian courtship song. The song of the Aquatic Warbler consists of whistle and rattle phrases produced as discontinuous A-, B- and C-songs, which are built by a single rattle, a rattle and a whistle, and more than two phrases of both kinds, respectively. The A- and B-songs are thought to be aggressive signals in male-male interactions, while C-songs are thought to be important for female choice. Here, we analysed recordings of 40 individually marked males, and determined the phrase repertoire. The enumerated repertoire (males recorded for ≥ 10 min) ranged from 16 to 158 (mean 99), however, it did not capture the complete phrase repertoires. We then used models from species diversity ecology to estimate the actual phrase repertoire size, which ranged between 18 and 300 phrases (mean 155). The estimated repertoire was predicted by the number of C-songs. The rattle repertoire was larger than the whistle repertoire, and both positively correlated with the number of C-songs. Our study indicates that male Aquatic Warblers have highly complex phrase repertoires that vary widely in size. Their courtship song is flexible and efficient, enabling relative song complexity to be demonstrated in a short sample, thus facilitating both female attraction through the quick presentation of large phrase repertoires and rival deterrence through the production of many short and simple A- and B-songs.

Sperm morphology and performance in relation to postmating prezygotic isolation in two recently diverged passerine species.

Divergence in sperm phenotype and female reproductive environment may be a common source of postmating prezygotic (PMPZ) isolation between species. However, compared to other reproductive barriers it has received much less attention. In this study, we examined sperm morphology and velocity in two hybridizing passerine species, the common nightingale (Luscinia megarhynchos) and thrush nightingale (L. luscinia). In addition, we for the first time characterized a passerine female reproductive tract fluid proteome. We demonstrate that spermatozoa of the common nightingale have significantly longer and wider midpiece (proximal part of the flagellum containing mitochondria) and longer tail compared to spermatozoa of thrush nightingale. On the other hand, they have significantly shorter and narrower acrosome. Importantly, these differences did not have any effect on sperm velocity. Furthermore, the fluid from the reproductive tract of common nightingale females did not differentially affect velocity of conspecific and heterospecific sperm. Our results indicate that the observed changes in the flagellum and acrosome size are unlikely to contribute to PMPZ isolation through differential sperm velocity of conspecific and heterospecific sperm in the female reproductive tract. However, they could affect other postcopulatory processes, which might be involved in PMPZ isolation, such as sperm storage, longevity or sperm-egg interaction.

Male and female songs propagation in a duetting tropical bird species in its preferred and secondary habitat.

Acoustic signals produced by animals must transmit through the environment to reach potential receivers and change their behaviour. Both the environment (vegetation, air properties, other animals) and the form of the signal affect the propagation process. Here we investigated how the transmission of different song types of a duetting songbird species inhabiting an extreme environment within African montane forest, varies between males and females as well as different types of micro-habitats. We hypothesised that male and female songs would have different transmission properties, reflecting known differences in signal form and function. We analysed signal-to-noise ratio (SNR), excess attenuation (EA) and tail-to-signal ratio (TSR) of songs of male and female Yellow-breasted Boubous (Laniarius atroflavus) that were played and re-recorded in a range of sites representing the species-typical habitats. We found significant effects of distance, site (habitat) and sex reflected in all three measures of sound degradation. The clearest, primarily distance-dependent pattern was found for SNR of songs propagated in level forest site. EA was substantially higher in shrubs than in forest habitats, while TSR reflecting longer echoes appeared at longer distances in forest sites. Thus, Yellow-breasted Boubou songs are better propagated in forests than in disturbed sites covered with shrubs. We found that all male song types used for broadcast singing propagated farther than female songs, with significantly higher SNR at all distances. The different male song types which are known to have different functions, also demonstrated a differentiated pattern of propagation reflecting their functionality. All signals that were tested propagated the furthest in the ideal condition described as forest with a level terrain. Signals degraded much faster during transmission through shrubs regrowing after forest burning. On this site, the differences in the propagation of male and female songs, as well as the differences between male song types, were relatively least pronounced. Transmission in typical mountain forest among streams and with substantial terrain variation revealed that degradation pattern in such habitat could be perturbed in a non-linear way. Streams acting as a source of high noise level also negatively affected transmission and may strongly limit the perception of birds staying close to them. However, stream noise did not affect sex differences in song propagation as was found for the site located in shrubs. Male songs showed more efficient transmission through all habitats (least in the shrubs) than female song. These differences were the result of male songs having a whistle structure that is better adapted for long-range propagation than the atonal, wideband frequency female vocalisations. Results support the idea that signals of males of the Yellow-breasted Boubous evolved under the pressure of long-range communication both with rivals and females, while females of the species are much more focused on within-pair communication or signalling together with their partner. The consequence of deforestation resulting in pushing back territories to the forest remnants along streams may be a shortening of the song's active range, in particular, in females.

Comparison of Karyotypes in Two Hybridizing Passerine Species: Conserved Chromosomal Structure but Divergence in Centromeric Repeats.

Changes in chromosomal structure involving chromosomal rearrangements or copy number variation of specific sequences can play an important role in speciation. Here, we explored the chromosomal structure of two hybridizing passerine species; the common nightingale (Luscinia megarhynchos) and the thrush nightingale (Luscinia luscinia), using conventional cytogenetic approaches, immunostaining of meiotic chromosomes, fluorescence in situ hybridization as well as comparative genomic hybridization (CGH). We found that the two nightingale species show conserved karyotypes with the same diploid chromosome number of 2n = 84. In addition to standard chromosomes, both species possessed a small germline restricted chromosome of similar size as a microchromosome. Just a few subtle changes in chromosome morphology were observed between the species, suggesting that only a limited number of chromosomal rearrangements occurred after the species divergence. The interspecific CGH experiment suggested that the two nightingale species might have diverged in centromeric repetitive sequences in most macro- and microchromosomes. In addition, some chromosomes showed changes in copy number of centromeric repeats between the species. The observation of very similar karyotypes in the two nightingale species is consistent with a generally slow rate of karyotype evolution in birds. The divergence of centromeric sequences between the two species could theoretically cause meiotic drive or reduced fertility in interspecific hybrids. Nevertheless, further studies are needed to evaluate the potential role of chromosomal structural variations in nightingale speciation.

Song type and song type matching are important for joint territorial defense in a duetting songbird.

Birds have a diverse acoustic communication system, with species-specific repertoires facilitating more complex behaviors in terms of both within- and between-pair communications. Certain song types are produced for specific functions, such as aggressive encounters. In addition, song matching behaviors, whereby neighboring individuals match song types, can be used in aggressive interactions as a sophisticated acoustic behavior. In this study, we examined the functions of song types, in a duet context, of male yellow-breasted boubous (Laniarius atroflavus), an Afromontane bush-shrike with a vocal sexual dimorphism. We aimed at assessing whether, structurally, certain song types elicited a heightened reaction than others and also whether song matching affected response behavior. A dual speaker playback procedure was performed for 18 pairs of boubous, each pair being exposed to duets with three different male song types. We found differences in response toward the different duet types but these differences resulted from the amount at which males matched different song types. Pairs responded stronger when a focal male matched the playback type, and matching was significantly more often found in cases where the rarest type of male song was used. We found no sex differences in terms of response strength to playback type. Our results indicate a two-level way of coding aggression toward intruding pairs. The yellow-breasted boubous utilize their repertoires, linking matching with structure in order to show aggression in terms of territory defense and sexual conflict. This study also confirms joint territorial defense as a main function of duets in this species.

Seasonality and social factors, but not noise pollution, influence the song characteristics of two leaf warbler species.

Changes in the acoustic signalling of animals occupying urban ecosystems is often associated with the masking effects of noise pollution, but the way in which they respond to noise pollution is not straightforward. An increasing number of studies indicate that responses can be case specific, and some species have been found to respond differently to high levels of natural versus anthropogenic noise, as well as different levels of the latter. While the perception of noise between species may vary with its source, amplitude and temporal features, some species may possess broader environmental tolerance to noise pollution, as they use higher frequency vocalizations that are less masked by low-frequency urban noise. In this study, we explored the song variation of two closely related leaf warblers, the Common Chiffchaff Phylloscopus collybita and the Willow Warbler Phylloscopus trochilus, inhabiting urban green spaces and nonurban forests. The main goal of our study was to evaluate the impact of moderate levels of noise pollution on the songs of species which use higher frequency vocalizations and large frequency bandwidth. Previous studies found that the Common Chiffchaff modified their song in response to intense noise pollution, while no such data is available for the Willow Warbler. However, the majority of urban green spaces, which serve as wildlife hot spots in urban environments are usually polluted with moderate noise levels, which may not mask the acoustic signals of species that communicate with higher frequency. We analysed the spectral and temporal song parameters of both warblers and described the ambient noise present in males' territories. Additionally, we looked at the social and seasonal aspects of bird song, since there is more than just noise in urban ecosystems which may affect acoustic communication. We found no evidence for noise-related bird song divergence in either species, however, we showed that social factors, time of day and season influence certain Common Chiffchaff and Willow Warbler song characteristics. Lack of noise-related bird song divergence may be due to the relatively low variation in its amplitude or other noise features present within the song frequency range of the studied species. Similar results have previously been shown for a few songbird species inhabiting urban ecosystems. Although in many cases such results remain in the shadow of the positive ones, they all contribute to a better understanding of animal communication in urban ecosystems.

Gut microbiota in two recently diverged passerine species: evaluating the effects of species identity, habitat use and geographic distance.

BACKGROUND: It has been proposed that divergence in the gut microbiota composition between incipient species could contribute to their reproductive isolation. Nevertheless, empirical evidence for the role of gut microbiota in speciation is scarce. Moreover, it is still largely unknown to what extent closely related species in the early stages of speciation differ in their gut microbiota composition, especially in non-mammalian taxa, and which factors drive the divergence. Here we analysed the gut microbiota in two closely related passerine species, the common nightingale (Luscinia megarhynchos) and the thrush nightingale (Luscinia luscinia). The ranges of these two species overlap in a secondary contact zone, where both species occasionally hybridize and where interspecific competition has resulted in habitat use differentiation. RESULTS: We analysed the gut microbiota from the proximal, middle and distal part of the small intestine in both sympatric and allopatric populations of the two nightingale species using sequencing of bacterial 16S rRNA. We found small but significant differences in the microbiota composition among the three gut sections. However, the gut microbiota composition in the two nightingale species did not differ significantly between either sympatric or allopatric populations. Most of the observed variation in the gut microbiota composition was explained by inter-individual differences. CONCLUSIONS: To our knowledge, this is the first attempt to assess the potential role of the gut microbiota in bird speciation. Our results suggest that neither habitat use, nor geographical distance, nor species identity have strong influence on the nightingale gut microbiota composition. This suggests that changes in the gut microbiota composition are unlikely to contribute to reproductive isolation in these passerine birds.

Structure and functions of Yellow-breasted Boubou (Laniarius atroflavus) solos and duets.

BACKGROUND: Birds have extremely well-developed acoustic communication and have become popular in bioacoustics research. The majority of studies on bird song have been conducted in the temperate zones where usually males of birds sing to attract females and defend territories. In over 360 bird species mostly inhabiting the tropics both males and females sing together in duets. Avian duets are usually formed when a male and female coordinate their songs. We focused on a species with relatively weakly coordinated duets, with male solo as the prevailing vocalisation type. METHODS: Instead of analysing a set of recordings spread over a long time, we analysed whole day microphone-array recordings of the Yellow-breasted Boubou (Laniarius atroflavus), a species endemic to West African montane rainforests. We described the structure of the solo and duet vocalisations and temporal characteristics of daily activity based on 5,934 vocal bouts of 18 focal pairs and their neighbours. RESULTS: Birds had small, sex specific repertoires. All males shared three types of loud whistles functioning as song type repertoires in both solos and duets. Females vocalised with five types of harsh, atonal notes with a more variable and usually lower amplitude. Three of them were produced both as solos and in duets, while two seem to function as alarm and excitement calls given almost exclusively as a solo. Solos were the most common vocalisation mode (75.4%), with males being more vocally active than females. Duets accounted for 24.6% of all vocalisations and in most cases were initiated by males (81%). The majority of duets were simple (85.1%) consisting of a single male and female song type, but altogether 38 unique duet combinations were described. Males usually initiated singing at dawn and for this used one particular song type more often than expected by chance. Male solo and duet activities peaked around dawn, while female solos were produced evenly throughout the day. DISCUSSION: Yellow-breasted Boubou is a duetting species in which males are much more vocal than females and duetting is not a dominating type of vocal activity. Duet structure, context and timing of daily production support the joint resource defence hypothesis and mate guarding/prevention hypotheses, however maintaining pair contact also seems to be important. This study provides for the first time the basic quantitative data describing calls, solos and duet songs in the Yellow-breasted Boubou.

Habitat-related differences in song structure and complexity in a songbird with a large repertoire.

BACKGROUND: Urbanisation has been shown to influence many aspects of animal vocal communication. Much attention has been paid to anthropogenic noise, which is often described as one of the most challenging disturbances for urban dwellers. While a large body of literature describes associations between vocal behavior of avian populations and background noise level, most of these studies were conducted on species with relatively simple songs and small repertoire sizes. This study focuses on the song thrush, Turdus philomelos, a common Eurasian songbird with a complex singing style and large syllable repertoire. Our objective was to determine whether frequency, repertoire and temporal organisation of song parameters vary between birds inhabiting urban and adjacent forest habitats in which ambient noise levels differ. RESULTS: Songs of urban males were found to be more complex than in conspecifics from natural forest populations. Urban dwellers possessed greater syllable repertoires and repeated syllable sequences more often. In addition, they used a smaller proportion of whistles and a higher proportion of twitter syllables when singing compared to the nonurban males. Moreover, we found significant differences in the minimum and peak frequency of the whistle syllable between studied populations. CONCLUSIONS: These findings may be an example of adaptation of acoustic communication in noisy urban environments, but we also discuss other possible explanations. We emphasize the need for further investigation into the relationships between birdsong and habitat characteristics, male quality, population density and ambient noise level in populations occupying urban and nonurban habitats.

Sing softly to evoke a response only from a recent intruder.

Low-amplitude soft songs have been described in many birds, but in most species, research has addressed only broadcast songs. Soft songs may have a similar or distinctive structure in comparison to broadcast songs produced in order to defend a territory and attract females. In some species, such soft songs were found to be produced in an aggressive context and were the best predictors of conflict escalation and later physical attack. However, such observations are not consistent across all species studied. Several hypotheses have been put forward to explain the function(s) of soft songs and why they are so quiet. Studies on the ortolan bunting (Emberiza hortulana) show that males produce soft songs similar in structure to broadcast songs during close interactions with conspecific intruders. However, experiments with the playback of loud and soft songs as well as taxidermic mount presentation revealed that soft song in this species does not fulfil aggressive signal criteria. Here we performed an experiment with two loudspeakers simulating movements of the intruder in order to test whether the soft songs are used to evoking a response from a nearby but not localised rival. We found that males responded with similar strength to songs played back from single and double speaker systems. Simultaneously, males produced more soft songs during and just after the phase of the experiment in which we simulated short flights of the intruder. Our results indicate that soft songs might be used during close interactions with rivals without being an aggressive signal and are used as short-range signals to check if the rival is still around. Our results also provide an alternative explanation of soft song behaviour in comparison to the hypotheses of eavesdropping avoidance and readiness to fight.

Functional significance of cuckoo Cuculus canorus calls: responses of conspecifics, hosts and non-hosts.

Male cuckoos Cuculus canorus produce calls that differ in number of syllables depending on environmental conditions and presence of male and female conspecifics. Why different males produce so repeatable calls that vary greatly in duration among males remains an open question. We used playback of cuckoo calls with few or many syllables (hereafter short and long calls), and woodpigeon calls (a control that also produces few or many syllables), predicting that playback of longer cuckoo calls should attract more male cuckoos (if males with such calls are dominant and successfully out-compete other males due to intraspecific competition), and attract more hosts mobbing male cuckoos (cuckoos with such calls and their females attract more hosts because of an increased risk of parasitism). Because cuckoos differentially parasitize hosts away from human habitation, we also tested whether the number of syllables in cuckoo calls differed with distance from buildings. Playback showed significant effects of number of syllables in cuckoo calls, but not woodpigeon Columba palumbus calls, with an additional effect of distance from human habitation decreasing the response to playback. These findings are consistent with the hypothesis that longer cuckoo calls, especially played back near human habitation, attract more conspecifics and hosts than shorter calls. To the best of knowledge this is the first study showing that cuckoo call response modified both other cuckoo individuals, as well as hosts response.

Dawn chorus interpretation differs when using songs or calls: the Dupont's Lark Chersophilus duponti case.

BACKGROUND: Territorial songbirds vocalise intensively before sunrise and then decrease their vocal activity. This creates a communication network that disseminates essential information for both males and females. The function of dawn chorus in birds has been frequently interpreted according to seasonal variation of singing as the breeding season advances, but potential differences in seasonal variation of song and calls for the same species have not been taken into account. METHODS: We chose Dupont's Lark as a model species to study whether the seasonal pattern of dawn chorus differs between singing and calling activity, because in this species most daily songs and calls are uttered at dawn. We registered vocal activity of Dupont's Lark before and around dawn in three different populations, through repeated sampling over the entire breeding season of two consecutive years. RESULTS: We found that dawn singing parameters remained constant or presented an increasing trend while dawn calling activity decreased as breeding season advanced. We also found different daily patterns for singing and calling, with birds calling mostly during the first 30 minutes of dawn choruses and singing peaking afterwards. DISCUSSION: The different time patterns of songs and calls may indicate that they serve diverse functions at dawn in the Dupont's Lark. Relaxation of dawn calling activity after the first month of the breeding season would suggest that dawn calling may be mainly related to mate attraction, while constant dawn singing throughout the breeding period would support a relationship of dawn singing to territorial defence. Our study highlights that the type of vocalisation used is an important factor to consider in further research on dawn choruses, since results may differ depending on whether calls or songs are analysed.

Shell colour, temperature, (micro)habitat structure and predator pressure affect the behaviour of Cepaea nemoralis.

Although shell colour polymorphism of the land snail Cepaea nemoralis is a well-known phenomenon, proximate and ultimate factors driving its evolution remain uncertain. Polymorphic species show variation in behavioural responses to selective forces. Therefore, we estimated effects of various environmental factors (temperature, humidity, food availability, (micro)habitat structure and predatory pressure) on behavioural response (frequency of locomotion, climbing and hiding) of C. nemoralis morphs, in experimental and natural conditions. In the experimental part of study, the frequency of locomotion was negatively affected by temperature and the presence of food and positively influenced by the presence of light. Morphs significantly differed in behavioural responses to environmental variability. Pink mid-banded and yellow five-banded morphs climbed less often and hide in shelter more often than yellow and pink unbanded individuals when temperature was low and food was absent. Snails fed most often at moderate temperature compared to low and high temperatures. Field investigations partially confirmed differences among morphs in frequency of climbing, but not in terms of probability of hiding in sheltered sites. In natural colonies, temperature and (micro)habitat structure significantly affected frequency of climbing as well as hiding in shelter. Snails more often hid in sheltered sites where thrushes preyed on Cepaea. Tendency of unbanded morphs to climb trees may have evolved under avian predatory pressure as thrushes forage on a ground. Tendency of banded morphs to hide in sheltered sites may reflect prey preferences for cryptic background. The results implicate that differential behaviour of C. nemoralis morphs compensate for their morphological and physiological limitations of adaptation to habitat.

Song duration mediates responses of territory owner in a songbird species with a small song repertoire.

Song is a sexually selected trait that is involved in mate attraction and territory defence in birds. Songs may convey information about different male quality components. They are flexible in terms of frequency, amplitude, and duration. Although changes in song duration are common, the function of this behaviour has been studied less strongly. It is known that song duration changes within a singing bout and may provide information about aggressive motivation. We tested whether the elongation and shortening of songs affects the responses of territorial ortolan bunting males to neighbour song playback. If changing song duration signals level of aggressiveness, then songs differing solely in duration may elicit behavioural responses of different strength. We performed two tests with different males assigned to two experimental groups and measured approaching and vocal response. In Experiment 1, we tested 18 males, which responded to the playbacks of elongated and normal (control) neighbour songs. In Experiment 2, we tested 17 males, which responded to the playbacks of shortened and normal (control) neighbour songs. Males responded significantly stronger to longer songs in both experiments as measured by the approach variable. Vocal response was not affected by treatment, but it was affected by the order of playback presentation. Our results indicate that song duration might be used for signalling current aggressive motivation during close interactions with rivals.

Individually specific call feature is not used to neighbour-stranger discrimination: the corncrake case.

In various contexts, animals rely on acoustic signals to differentiate between conspecifics. Currently, studies examining vocal signatures use two main approaches. In the first approach, researchers search for acoustic characteristics that have the potential to be individual specific. This approach yields information on variation in signal parameters both within and between individuals and generates practical tools that can be used in population monitoring. In the second approach, playback experiments with natural calls are conducted to discern whether animals are capable of discriminating among the vocal signatures of different individuals. However, both approaches do not reveal the exact signal characteristics that are being used in the discrimination process. In this study, we tested whether an individual-specific call characteristic--namely the length of the intervals between successive maximal amplitude peaks within syllables (PPD)--is crucial in neighbour-stranger discrimination by males of the nocturnal and highly secretive bird species, the corncrake (Crex crex). We conducted paired playback experiments in which corncrakes (n = 47) were exposed to artificial calls with PPD characteristics of neighbour and stranger birds. These artificial calls differed only in PPD structure. The calls were broadcast from a speaker, and we recorded the birds' behavioural responses. Although corncrakes have previously been experimentally shown to discriminate between neighbours and strangers, we found no difference in the responses to the artificial calls representing neighbours versus strangers. This finding demonstrates that even if vocal signatures are individual specific within a species, it does not automatically mean that said signatures are being crucial in discrimination among individuals. At the same time, the birds' aggressive responses to the artificial calls indicated that the information transmitted by PPDs is important in species-specific call recognition and may be used by males and/or females to evaluate sender quality, similarly like sound frequency in some insect species.

Functionally similar acoustic signals in the corncrake (Crex crex) transmit information about different states of the sender during aggressive interactions.

We combined playback experiments with hormonal manipulations to study the information content of acoustic signals during aggressive interactions between male corncrakes. During territorial conflicts, fights are uncommon, but the intensity of signaling usually increases. Such signals can be temporally and contextually associated with many aggressive behaviors and most likely function as threats or as indicators of the sender's quality or motivation. However, such correlational data are unsatisfactory for the proper interpretation of the function and information content of signals. Experimental tests are required to determine whether signals and aggressive behaviors are controlled by common or independent mechanisms. In our experiment, we assigned subjects to four groups: testosterone-implanted birds, flutamide-implanted birds, birds with empty implants, and non-captured control birds. Males produced two types of calls (quiet soft calls and loud broadcast calls), both of which are known to be reliable predictors of aggressive escalation. When testosterone action was blocked with flutamide, males significantly limited the amount of time spent close to the playback speaker and stopped responding to playback with soft calls. Broadcast calling was unaffected by the blockage of testosterone. Conversely, increased levels of testosterone neither affected calling nor the time spent near the speaker, indicating a permissive, rather than a graded effect of androgens. We concluded that, despite the seemingly similar function, both signals may transmit information about different states of the sender; soft calls seem to imply a threat of force, while broadcast calls appear to be more similar to an announcement, which is only indirectly associated with a male's aggressive behavior.

No male identity information loss during call propagation through dense vegetation: the case of the corncrake.

Individually specific acoustic signals in birds are used in territorial defence. These signals enable a reduction of energy expenditure due to individual recognition between rivals and the associated threat levels. Mechanisms and acoustic cues used for individual recognition seem to be versatile among birds. However, most studies so far have been conducted on oscine species. Few studies have focused on exactly how the potential for individual recognition changes with distance between the signaller and receiver. We studied a nocturnally active rail species, the corncrake, which utters a seemingly simple disyllabic call. The inner call structure, however, is quite complex and expressed as intervals between maximal amplitude peaks, called pulse-to-pulse durations (PPD). The inner call is characterized by very low within-individual variation and high between-individuals difference. These variations and differences enable recognition of individuals. We conducted our propagation experiments in a natural corncrake habitat. We found that PPD was not affected by transmission. Correct individual identification was possible regardless of the distance and position of the microphone which was above the ground. The results for sounds from the extreme distance propagated through the vegetation compared to those transmitted above the vegetation were even better. These results support the idea that PPD structure has evolved under selection favouring individual recognition in a species signalling at night, in a dense environment and close to the ground.