Fanny Rybak

How to identify dear enemies: the group signature in the complex song of the skylark Alauda arvensis.

SUMMARY Song geographic variation and Neighbour–Stranger (N–S) discrimination have been intensively but separately studied in bird species, especially in those with small- to medium-sized repertoires. Here, we establish a link between the two phenomena by showing that dialect features are used for N–S recognition in a territorial species with a large repertoire, the skylark Alauda arvensis. In this species, during the breeding season, many pairs settle in stable and adjoining territories gathered in locations spaced by a few kilometres. In a first step, songs produced by males established in different locations were recorded, analyzed and compared to identify possible microgeographic variation at the syntax level. Particular common sequences of syllables (phrases) were found in the songs of all males established in the same location (neighbours), whereas males of different locations (strangers) shared only few syllables and no sequences. In a second step, playback experiments were conducted and provided evidence for N–S discrimination consistent with the dear-enemy effect, i.e. reduced aggression from territorial birds towards neighbours than towards strangers. In addition, a similar response was observed when a `chimeric signal (shared phrases of the location artificially inserted in the song of a stranger) and a neighbour song were broadcast, indicating that shared sequences were recognized and identified as markers of the group identity. We thus show experimentally that the shared phrases found in the songs of neighbouring birds constitute a group signature used by birds for N–S discrimination, and serve as a basis for the dear-enemy effect.

Are bird song complexity and song sharing shaped by habitat structure? An information theory and statistical approach

In songbirds, song complexity and song sharing are features of prime importance for territorial defence and mate attraction. These aspects of song may be strongly influenced by changes in social environment caused by habitat fragmentation. We tested the hypothesis that habitat fragmentation induced by human activities influences song complexity and song sharing in the skylark, a songbird with a very large repertoire and whose population recently underwent a large decline. We applied powerful mathematical and statistical tools to assess and compare song complexity and song sharing patterns of syllables and sequences of syllables in two populations: a declining population in a fragmented habitat, in which breeding areas are separated from each other by unsuitable surroundings, and a stable population in a continuous habitat. Our results show that the structure of the habitat influences song sharing, but not song complexity. Neighbouring birds shared more syllables and sequences of syllables in the fragmented habitat than in the continuous one. Habitat fragmentation seems thus to have an effect on the composition of elements in songs, but not on the number and complexity of these elements, which may be a fixed feature of song peculiar to skylarks.

When to be a dear enemy: flexible acoustic relationships of neighbouring skylarks, Alauda arvensis

Numerous territorial species are less aggressive towards neighbours than strangers. This tolerance towards neighbouring conspecifics, termed the ‘dear enemy’ effect, seems to be a flexible feature of the relationship between neighbours, and has been shown to disappear in some species after experimental or natural modifications of the context. However, the maintenance over time of this singular relationship has been poorly studied. In this study, we followed the change of dear enemy relationships during the breeding season in a territorial songbird with a complex song, the skylark. We examined in the field the response of territory owners to playbacks of neighbour and stranger songs at three periods of the breeding season, corresponding to three ecological and social situations. Results showed that neighbours were dear enemies in the middle of the season, when territories were stable, but not at the beginning of the breeding season, during settlement and pair formation, nor at the end, when bird density increased owing to the presence of young birds becoming independent. Thus, the dear enemy relationship is not a fixed pattern but a flexible one likely to evolve with social and ecological circumstances.

Stridulations reveal cryptic speciation in neotropical sympatric ants.

The taxonomic challenge posed by cryptic species underlines the importance of using multiple criteria in species delimitation. In the current paper we tested the use of acoustic analysis as a tool to assess the real diversity in a cryptic species complex of Neotropical ants. In order to understand the potential of acoustics and to improve consistency in the conclusions by comparing different approaches, phylogenetic relationships of all the morphs considered were assessed by the analysis of a fragment of the mitochondrial DNA cytochrome b. We observed that each of the cryptic morph studied presents a morphologically distinct stridulatory organ and that all sympatric morphs produce distinctive stridulations. This is the first evidence of such a degree of specialization in the acoustic organ and signals in ants, which suggests that stridulations may be among the cues used by these ants during inter-specific interactions. Mitochondrial DNA variation corroborated the acoustic differences observed, confirming acoustics as a helpful tool to determine cryptic species in this group of ants, and possibly in stridulating ants in general. Congruent morphological, acoustic and genetic results constitute sufficient evidence to propose each morph studied here as a valid new species, suggesting that P. apicalis is a complex of at least 6 to 9 species, even if they present different levels of divergence. Finally, our results highlight that ant stridulations may be much more informative than hitherto thought, as much for ant communication as for integrative taxonomists.

Compared Ontogenesis of Courtship Song Components of Males from the Sibling Species, D. melanogaster and D. simulans

The courtship song of Drosophila is known to be an important signal involved in sex and species recognition. It consists of pulse song and sine song, which have been studied in a quantitative way with different parameters. For the first time the setting of both components of the acoustic signaling is described and compared for males belonging to the sibling species D. melanogaster and D. simulans. At early ages, these two species share similar interpulse interval values but maturation establishes the species specificity of this character. For D. melanogaster the variations of several acoustic parameters take place in two successive periods, whereas for D. simulans the majority of the acoustic features does not change much with age. In D. melanogaster, copulation success seems to be linked to the maturation of the acoustic performance, which is not the case for D. simulans.

Response to displaced neighbours in a territorial songbird with a large repertoire

Neighbour recognition allows territory owners to modulate their territorial response according to the threat posed by each neighbour and thus to reduce the costs associated with territorial defence. Individual acoustic recognition of neighbours has been shown in numerous bird species, but few of them had a large repertoire. Here, we tested individual vocal recognition in a songbird with a large repertoire, the skylark Alauda arvensis. We first examined the physical basis for recognition in the song, and we then experimentally tested recognition by playing back songs of adjacent neighbours and strangers. Males showed a lower territorial response to adjacent neighbours than to strangers when we broadcast songs from the shared boundary. However, when we broadcast songs from the opposite boundary, males showed a similar response to neighbours and strangers, indicating a spatial categorisation of adjacent neighbours’ songs. Acoustic analyses revealed that males could potentially use the syntactical arrangement of syllables in sequences to identify the songs of their neighbours. Neighbour interactions in skylarks are thus subtle relationships that can be modulated according to the spatial position of each neighbour.

First description of underwater acoustic diversity in three temperate ponds

The past decade has produced an increased ecological interest in sonic environments, or soundscapes. However, despite this rise in interest and technological improvements that allow for long-term acoustic surveys in various environments, some habitats’ soundscapes remain to be explored. Ponds, and more generally freshwater habitats, are one of these acoustically unexplored environments. Here we undertook the first long term acoustic monitoring of three temperate ponds in France. By aural and visual inspection of a selection of recordings, we identified 48 different sound types, and according to the rarefaction curves we calculated, more sound types are likely present in one of the three ponds. The richness of sound types varied significantly across ponds. Surprisingly, there was no pond-to-pond daily consistency of sound type richness variation; each pond had its own daily patterns of activity. We also explored the possibility of using six acoustic diversity indices to conduct rapid biodiversity assessments in temperate ponds. We found that all indices were sensitive to the background noise as estimated through correlations with the signal-to-noise ratio (SNR). However, we determined that the AR index could be a good candidate to measure acoustic diversities using partial correlations with the SNR as a control variable. Yet, research is still required to automatically compute the SNR in order to apply this index on a large data set of recordings. The results showed that these three temperate ponds host a high level of acoustic diversity in which the soundscapes were variable not only between but also within the ponds. The sources producing this diversity of sounds and the drivers of difference in daily song type richness variation both require further investigation. Such research would yield insights into the biodiversity and ecology of temperate ponds.

Does true syntax or simple auditory object support the role of skylark song dialect

Parallels between birdsong and human language are numerous and include particular temporal arrangements of acoustic units and the existence of dialects. In animal communication, modifications of the temporal ordering of existing acoustic units have rarely been clearly linked with changes in information content, particularly in a natural environment. Here, we show that the organization of birdsong units (‘syllables’) in sequences supports interindividual relationships within skylark communities. We manipulated the temporal arrangement of song dialect variants (‘shared phrases’) in the skylark, Alauda arvensis, a songbird with a very large repertoire of syllables and complex song. When tested with playback experiments performed in the field, skylarks were able to perceive subtle differences in the ordering of syllables. Modifications of the syllable ordering within shared phrases changed the information content from ‘group member’ to ‘unfamiliar individual’ and induced more aggressive reactions than shared phrases with a preserved syllable arrangement. Shared phrases often varied between individuals in the number of successive repetitions of similar syllable types, but were very consistent in terms of syllable type ordering. Our results indicate that skylarks rely not simply on the composition in syllable types of shared phrases to recognize group members, but on syllable type ordering. Shared phrases could be perceived by birds as ‘auditory objects’ embedded within songs. Alternatively, birds might identify incorrect phrases using grammatical rules governing the succession of syllables composing the phrases shared by their group. The presence of between-individual variation in phrase length, associated with consistent syllable type ordering revealed by our analysis, suggests that the latter hypothesis is more likely. Our results show that birds perceive disruptions in the natural temporal pattern of song units, and that this temporal pattern is behaviourally salient and carries information.

A SIMPLE METHOD FOR RECORDING LOW-AMPLITUDE SOUNDS. APPLICATION TO THE STUDY OF THE COURTSHIP SONG OF THE FRUIT FLY DROSOPHILA MELANOGASTER

ABSTRACT Some acoustic signals produced by small insects are very low in amplitude and attenuate rapidly with distance. To achieve high quality recordings with such signals, the use of specialised microphones or of sound insulation chambers is necessary. This paper presents a simple and efficient method for the recording of acoustic signals emitted by small sources. Its principle is based upon the use of two simultaneous digital recordings from two microphones: one records the ambient noise while the other records the ambient noise plus the signal to analyse. Both these recordings are converted into digital files and then a simple subtraction between the two isolates the signal with a good signal-to-noise ratio. With this method of background noise removal, the recording of low amplitude sounds in an uninsulated room with common microphones becomes possible. We have applied this method to the study of 12 complete courtships of Drosophila melanogaster and particularly to the analysis of pulse sounds produced by the male in presence of a female. The study focuses mainly on the rhythm of production of pulse trains over the course of the courtship.

Are Unfamiliar Neighbours Considered to Be Dear-Enemies?

Background Discriminating threatening individuals from non-threatening ones allow territory owners to modulate their territorial responses according to the threat posed by each intruder. This ability reduces costs associated with territorial defence. Reduced aggression towards familiar adjacent neighbours, termed the dear-enemy effect, has been shown in numerous species. An important question that has never been investigated is whether territory owners perceive distant neighbours established in the same group as strangers because of their unfamiliarity, or as dear-enemies because of their group membership. Methodology/Principal Findings To investigate this question, we played back to male skylarks (Alauda arvensis) songs of adjacent neighbours, distant neighbours established a few territories away in the same microdialect area and strangers. Additionally, we carried out a propagation experiment to investigate how far skylark songs are propagated in their natural habitat and we estimated repertoire similarity between adjacent neighbours, distant neighbours and strangers. We show that skylarks, in the field, respond less aggressively to songs of their distant and likely unfamiliar neighbours, as shown by the propagation experiment, compared to stranger songs. The song analysis revealed that individuals share a high amount of syllables and sequences with both their adjacent and distant neighbours, but only few syllables and no sequences with strangers. Conclusions The observed reduction of aggression between distant neighbours thus probably results from their familiarity with the vocal group signature shared by all members of the neighbourhood. Therefore, in skylarks, dear-enemy-like relationships can be established between unfamiliar individuals who share a common acoustic code.