Male Song Research Articles

Artificial Light at Night Advances the Onset of Vocal Activity in Both Male and Female Great Tits During the Breeding Season, While Noise Pollution Has Less Impact and Only in Females

Artificial light at night (ALAN) and noise pollution are two important stressors associated with urbanisation that can have a profound impact on animal behaviour and physiology, potentially disrupting biological rhythms. Although the influence of ALAN and noise pollution on daily activity patterns of songbirds has been clearly demonstrated, studies often focus on males, and the few that examined females have not included the potential influence of males on female activity patterns. Using free-living pairs of great tits (Parus major) as a model, we examined for the first time the effects of ALAN and noise pollution and their interaction on the onset of (vocal) activity in both members of a pair. We focused on the egg-laying phase, when both sexes are most vocally active. The onset of male dawn song, female emergence time from the nest box and the onset of female calling in the nest box were measured and used as a proxy for the chronotype. The repeatabilities for all chronotype proxies were high, with higher repeatabilities for males. Consistent with previous studies, ALAN advanced the onset of male dawn song, while it did not elicit a strong response in female emergence time. Additionally, our results suggest an indirect effect of ALAN on the onset of female vocal activity via acoustic interaction with the male. Noise pollution advanced the emergence time in females, while an interaction between ALAN and noise pollution was found for the onset of female calling. In agreement with previous studies, several covariables were shown to have an influence on the activity onset. Taking several proxies for chronotype into account, this study has provided robust evidence of effects of ALAN on male and female cavity-nesting songbirds during the egg-laying period.

Ancestral neural circuits potentiate the origin of a female sexual behavior in Drosophila

Courtship interactions are remarkably diverse in form and complexity among species. How neural circuits evolve to encode new behaviors that are functionally integrated into these dynamic social interactions is unknown. Here we report a recently originated female sexual behavior in the island endemic Drosophila species D. santomea, where females signal receptivity to male courtship songs by spreading their wings, which in turn promotes prolonged songs in courting males. Copulation success depends on this female signal and correlates with males’ ability to adjust his singing in such a social feedback loop. Functional comparison of sexual circuitry across species suggests that a pair of descending neurons, which integrates male song stimuli and female internal state to control a conserved female abdominal behavior, drives wing spreading in D. santomea. This co-option occurred through the refinement of a pre-existing, plastic circuit that can be optogenetically activated in an outgroup species. Combined, our results show that the ancestral potential of a socially-tuned key circuit node to engage the wing motor circuit facilitates the expression of a new female behavior in appropriate sensory and motivational contexts. More broadly, our work provides insights into the evolution of social behaviors, particularly female behaviors, and the underlying neural mechanisms.

Chickadees sing different songs in sympatry versus allopatry.

Character displacement theory predicts that closely-related co-occurring species should diverge in relevant traits to reduce costly interspecific interactions such as competition or hybridization. While many studies document character shifts in sympatry, few provide corresponding evidence that these shifts are driven by the costs of co-occurrence. Black-capped (Poecile atricapillus) and mountain chickadees (P. gambeli) are closely-related, ecologically similar, and broadly distributed songbirds with both allopatric and sympatric populations. In sympatry, both species appear to suffer costs of their co-occurrence: 1) both species are in worse body condition compared to allopatry and 2) hybridization sometimes yields sterile offspring. Here, we explored character displacement in the songs of black-capped and mountain chickadees by characterizing variation in male songs from sympatric and allopatric populations. We found that mountain chickadees sing differently in sympatry versus allopatry. Specifically, they produced more notes per song, were more likely to include an extra introductory note, and produced a smaller glissando in their first notes compared to all other populations. Combined with previous research on social dominance and maladaptive hybridization between black-capped and mountain chickadees, we posit that differences in sympatric mountain chickadee song are population-wide shifts to reduce aggression from dominant black-capped chickadees and/or prevent maladaptive hybridization.

Temperature effects on interspecific eavesdropping in the wild

Abstract Mating signals are targets of conspecific signal recognition and sexual selection, but are also subject to abiotic temperature effects and to biotic interspecific eavesdroppers. In crickets, the male calling song becomes faster at warmer temperatures, and female crickets’ recognition of male song tracks temperature in a coordinated manner, termed ‘temperature coupling.’ But female crickets are not the only ecologically relevant listeners: some cricket species are parasitized by Ormia ochracea, a parasitoid fly which finds its cricket hosts by eavesdropping on male cricket song. How temperature affects parasitoid fly phonotaxis to song is largely unexplored, with only one previous study conducted under field conditions. Here we explore six possible patterns of thermal effects on fly responses to cricket song, including temperature coupling, using field playbacks of synthetic Gryllus lineaticeps songs designed to be species-typical at various temperatures. We find that temperature does affect fly response, but that the temperature deviation of songs from ambient does not impact numbers of flies caught. We extend this finding by comparing the temperatures of the air (where flies search for their hosts) and the ground (where their host crickets signal) to show that temperature coupling is unlikely to be effective given microhabitat variation and differential rates of cooling in the evening hours when flies are most active. Our results can be interpreted more broadly to suggest (i) temperature effects on intraspecific communication systems may be more tightly coupled than are effects on interspecific eavesdropping, and (ii) variation in thermal microhabitats in the field make it difficult to translate laboratory physiological responses to natural selection in the wild.

Conspecific cues and breeding territory selection in Whinchat on abandoned fields

Many bird species use social information to guide territory selection. Different species in different habitats may use both pre-breeding and post-breeding conspecific cues. Abandoned agricultural fields are of particular interest for studies of conspecific attraction because the ephemeral nature of the habitat suggests territory search just before breeding, but high predation pressure suggests the use of performance-based conspecific cues. We investigated whether post-breeding or pre-breeding social cues affect Whinchat (Saxicola rubetra) abundance at the territorial scale during a two-year experiment in abandoned fields. We assigned 27 experimental plots (3.14 ha) to one of three treatments: post-breeding treatment, pre-breeding treatment and silent control. We conducted playback experiments with fledgling calls (evidence of past reproductive success) during the post-breeding period and male songs (evidence of conspecific presence) during the pre-breeding period. We estimated the difference in Whinchat abundance in two consecutive years and tested whether this value differed between treatment and control plots. We observed a slight increase (0.67 ± 0.29 pairs per plot) in Whinchat abundance in the post-breeding treatment plots and no significant changes in the pre-breeding (−0.22 ± 0.32 pairs per plot) and control (−0.22 ± 0.22 pairs per plot) plots. Our results suggest that Whinchats do not use conspecific acoustic cues during the pre-breeding period and provide limited evidence for the use of fledgling calls as conspecific cues in the post-breeding period.

Interacting phenotypic plasticities: Do male and female responses to the sociosexual environment interact to determine fitness?

Socially induced plasticity in reproductive effort is a widely documented phenomenon. However, few empirical studies have examined how male and female plastic responses to the social environment might interact in determining fitness outcomes. In field crickets, Teleogryllus oceanicus, males respond to rival song by increasing expenditure on seminal fluid proteins that enhance competitive fertilization success at the cost of reduced embryo survival. It remains unknown whether plastic responses in females could moderate the effects of male competitiveness on offspring performance. Here we used a fully factorial design to explore the interacting effects on fitness of male and female plasticity to the sociosexual environment. We found that female crickets exposed to male song increased the number of eggs produced during early life reproduction, which came at a cost of reduced offspring size. There was evidence, albeit weak, that interacting effects of male and female sociosexual environment contributed to variation in the hatching success of eggs laid by females. Lifetime offspring production was unaffected by the sociosexual environments to which upstream male and female plastic responses were made. Our data offer a rare test of the theoretical expectation that male and female plasticities should interact in their effects on female fitness.

Neighbors affect vocal behavior of tropical wrens: a multispeaker density-manipulation experiment.

For territorial animals, the behavior of conspecific neighbors sets the social context of communication. Despite numerous investigations of vocalizations related to territory defense and mate attraction, the effect of neighbor density on animal vocal behavior has received little attention, particularly in tropical animals and animals where both sexes produce complex acoustic signals. In this study, we used an innovative multispeaker playback experiment to manipulate the apparent density of neighbors in rufous-and-white wrens, Thryophilus rufalbus, living in Costa Rica's tropical dry forest. In this tropical songbird, both males and females defend year-round territories and sing complex, learned songs for territory defense. We recorded the singing behavior of 24 subjects (12 pairs), and then we used an array of 6 loudspeakers to simulate the presence of 6 new territorial neighbors (3 simulated pairs) outside each subject pair's breeding territory. The stimuli persisted for 3 consecutive days, with both male and female songs broadcast at a natural rate from dawn to dusk. We found that the mean male song rate increased by almost 50% in response to the simulated increase in local density. Females showed less frequent song-type switching rates following the simulated increase in local density, although it was a marginal increase. These findings reveal that male and female songbirds' vocal behavior varies with the local density of territorial neighbors. We conclude that birds are sensitive to acoustic signals of conspecific density arising from sounds beyond their territory boundaries, and that they use this public information to guide their vocal behavior.

Agonistic song rate positively correlates with male breeding success and avian malaria infection in Acrocephalus paludicola (Aquatic Warbler), a promiscuous songbird with female-only parental care

Abstract The link between male song and reproductive success has been explored at length in socially monogamous birds, but results were inconsistent and under-represented socially non-monogamous species with uniparental care. Here, we evaluated whether metrics of male song in Acrocephalus paludicola (Aquatic Warbler), a promiscuous songbird with female-only care and large repertoires, are associated with fitness- and quality-related traits. We showed that the number of 9-day-old nestlings fathered and probability of avian malaria infection increased with the rate of agonistic song, produced in male–male interactions, and that male-scaled mass index (proxy for energy reserves) positively correlated with the repertoire size. The male breeding success was not explained by repertoire size and song duty cycle. None of the song variables clearly explained the male return rate or change in the tarsus length (proxy for structural body size) and scaled mass index of the fathered chicks between days 2 and 9 post-hatch. No other relationships between the song characteristics and the inbreeding coefficient, scaled mass index, wing-length and Trypanosoma infection status of the males were supported, and Plasmodium infection was unrelated to the song duty cycle and repertoire size. We conclude that in male A. paludicola the repertoire size could be a signal of early developmental stress or current body condition, and the agonistic song rate could be shaped by sexual selection and signal to females a genetic makeup that enhances survival upon infection by Plasmodium.

Comparisons of acoustic structures between sexes in a duetting, montane bird

Female song in birds is more widespread than previously thought, and studies across different species are critical for better understanding the entire evolutionary process of bird song. In this study, we recorded the songs of males and females in a duetting, montane bird species, the Elliot's laughingthrush Trochalopteron elliotii, across consecutive breeding seasons. We specifically focused on identifying the number and structure of different song types by males and females, and compared these acoustic structures between the sexes. Our findings revealed that both males and females sang sex‐specific solos. More specifically, females sang a single type of solo that varied significantly in the number of notes, whereas males produced three different solos composed of 2, 3, or 4 notes, respectively. Female solos exhibited significantly more notes and longer song duration compared to males. Male solos typically had a significantly higher maximum frequency for the entire song. No significant differences were observed in the duration of the first note, song rate, and other frequency characteristics between male and female solos. Furthermore, paired males and females coordinated their sex‐specific solos to form duets when challenged by conspecific territory intruders, both within and outside the breeding season. Sex‐specific solos suggested that male and female songs play different roles and may be subject to different selective pressures. Further research is necessary for elucidating the functions of male song, female song, and duets in this montane bird species.

Mating behavior and three types of mating songs of the sandy beach-dwelling ground cricket Dianemobius csikii (Grylloidea, Trigonidiidae, Nemobiinae)

This study aimed to investigate the mating behavior of the sandy beach-dwelling ground cricket, Dianemobius csikii (Bolívar, 1901) (Orthoptera, Grylloidea, Trigonidiidae, Nemobiinae). The calling songs of males were recorded, and the temporal structures of the songs were analyzed. Subsequently, the courtship song and mating behavior of cricket pairs were observed. The calling song consisted only of monotonous chirps, while the courtship song consisted of similar chirps and ticks consisting of a single pulse. Dianemobius csikii exhibited a relatively longer courtship duration than other cricket species. The female stayed with the male for approximately 25 min while the male emitted the courtship song. The male then changed to the trill song, which is a continuous song, just before copulation. Copulation occurred within 10–40 s of the male emitting the trill song. The courtship behavior differs from that of other well-studied cricket species, such as Gryllinae. The findings of this study provide insight into the mating behaviors of crickets.

Response of the emerald‐spotted wood‐dove to the song of conspecific males and sympatric congeners

AbstractBird song contains crucial information that enables recognition of conspecific individuals from a distance, which facilitates subsequent complex behaviors related to mate attraction and rival deterrence. Loud broadcast signals of non‐learning bird species are usually treated as less complex than songs of Oscines, but several studies have revealed that song learning is not necessary for the evolution of complex signaling. Here, we focused on a species of African wood‐dove which produces superficially simple songs consisting of short notes organized into two parts of different amplitude. Our model was a population of emerald‐spotted wood‐doves (Turtur chalcospilos) that live in the same area as the physically and vocally similar blue‐spotted wood‐dove (Turtur afer). We tested the responses of male emerald‐spotted wood‐doves to different playbacks simulating territorial intrusion. We used songs of the focal species and of the congener, two types of artificially mixed songs with different parts belonging to the focal or congeneric species, and a control song. We aimed to assess (i) whether the focal species responds only to its own species' song or exhibits interspecific territoriality, and (ii) which part of the song is responsible for coding species identity. We found that male emerald‐spotted wood‐doves responded strongly to playback, but almost exclusively approached only the playback of their own species' song. Additionally, only conspecific song caused a decrease in song rate during playback and an increase in song output after playback. Our results suggest that emerald‐spotted wood‐doves are able to discriminate their own songs from those of congeners and do not exhibit interspecies territoriality. We were unable to identify a single part of the song that codes species‐specificity, as mixed songs of any kind did not substantially increase responsiveness to playback in comparison to the congener song or the control. We discuss these results in the context of current hypotheses regarding interspecific territoriality and the evolution of species‐identity coding.

Vocal differences in note, sequence and great call sequence among three closely related Nomascus gibbon species

AbstractThe vocal structure of gibbons has long been believed to be under strong genetic control and consistent with phylogeny. Gibbon vocalizations show hierarchical structures and song coordination behaviors, which may have different functions and convey distinct information compared to basic notes. However, previous studies have not compared vocalizations at multiple levels across species. We compared the vocal structures of three Nomascus species in the wild in South China, namely the western black gibbon (Nomascus concolor), Cao vit gibbon (N. nasutus), and Hainan gibbon (N. hainanus) at the levels of note (six features of four types of notes), male sequence (eight features), and song coordination (four features of the overlap pattern between male and female songs). We selected 15 recordings for each species: N. concolor (six groups, 2006 to 2021), N. nasutus (seven groups, 2008 to 2021), and N. hainanus (six groups, 2020 to 2021). We used permuted random forest analysis to test for species differences in vocal structure and the contribution of acoustic features to species differences at each level. We used acoustic features to construct a hierarchical clustering tree, and compared it with phylogenetic relationships. We found significant differences at all levels except the boom note. Acoustic similarity did not match genetic similarity in boom and pre-modulated notes, suggesting that genetic differences are not the only factor leading to species differences in vocal structures. We also found different contributions of acoustic features to differentiation in vocal structure at different levels. Our study suggests that we need to compare gibbon vocalizations at different levels to understand their differentiation and evolution.

Geographical variation in Isophya camptoxypha (Orthoptera, Tettigoniidae) male songs: Part 1—Solo performance across different microphones

Through the analysis of male-song characteristics in Isophya camptoxypha (Fieber), a Curve-tailed plump bush-cricket species prevalent in the highlands of the Alps and the Carpathians, this study establishes that acoustic signals not only serve for species identification among closely related counterparts but also contribute to biogeographical assumptions and constructions. The investigation delves into the formation of a relict locality on Stinka Mountain, situated at the junction of two Carpathian biogeographical regions. Additionally, the research explores the implications of using different types of microphones for bioacoustic measurements.

Courtship behavior: Resurrecting an undead song

The fly Drosophila yakuba has lost an ancestral component of the male courtship song: this is due to ontogenetic death of effector neurons in the ventral nerve cord, a result of the D.yakuba sex-determining gene dsx producing a male isoform, dsxM, with cell-death-promoting activity similar to that of the female isoform, dsxF, in D.melanogaster.

A new species of Ducetia (Tettigoniidae, Phaneropterinae) from Mindanao, Philippines based on bioacoustics, stridulatory file morphology and genetic data

The Ducetia japonica species group consists of 11 species widely distributed in Asia and Australia. It includes a single species from the Philippines: Ducetia adspersa Brunner von Wattenwyl, 1878. Previously synonymized under Ducetia japonica (Thunberg, 1815), this species was resurrected for its unique stridulatory file. Based on its distinct stridulatory file morphology, male calling song and cytochrome c oxidase subunit I signature, we discovered and describe here a new species from Mindanao in the south of the Philippines: Ducetia laniae sp. nov. We also describe the calling song of this new species. This discovery underscores the rich biodiversity of the region and highlights the necessity for continued exploration and study to unravel the complexities within this genus.

The taxonomic status of recently described Isophya taxa from Serbia (Tettigoniidae, Phaneropterinae)

During recent decades, increasing research of the taxonomy of the genus Isophya resulted in taxonomic descriptions of several new species. The delimitation of these species is mainly based on oscillographic song analysis of the male song in combination with morphological characters, such as the shape of male cerci, tegmina and ovipositor. In the present paper, we used an integrative taxonomic approach in order to resolve the status of four recently described Isophya taxa from Serbia. Based on our molecular and bioacoustic analyses, all analyzed taxa belong to a single species: Isophya modestior. The majority of the morphological characters used to differentiate the taxa showed strong intra- and interpopulational variability, proving that describing new species within the genus Isophya should not rely on morphological characters alone, but needs to consider bioacoustic analyses as the main tool and a larger series of specimens.

House cricket males reared at different perceived acoustic population densities differ in adult behaviour but not physiology

Abstract If environments stay relatively constant over an individual's lifetime, a juvenile that accurately perceives environmental conditions, like population density, may adjust adult traits to better match their environment, thereby increasing success. While previous studies have explored how adult exposure to population density affects physiological and behavioural plasticity, the influence of juvenile density experience on adult traits is less studied. Using the common house cricket, Acheta domesticus, we explored whether perceived acoustic population density during development affected adult physiology and behaviour. We simulated high‐ and low‐densities using live ambient male song. Upon maturation, we measured metabolic (resting respiration) rate, reproductive investment (testes and accessory gland masses), calling song characteristics and aggressive behaviours from pairwise contests between males from different densities. Male rearing density did not affect resting metabolic rates or reproductive organ masses. However, high‐density males had significantly faster, longer chirps, with more pulses—known to be preferred by females—and higher dominant frequency. Low‐density males won more aggressive contests and sang the aggressive song more. Initiation of aggressive behaviours or song and singing more aggressive song were the only other significant predictors of contest outcome. These results suggest that males may plastically adjust calling song characteristics and aggressive behaviour, but not physiology, based on perceived density during development. We hypothesize that alternative mating tactics—that is, territory guarding versus attractive song production—may underlie these observed patterns. Overall, our study highlights the significant influence of early‐life biotic environments on adult behavioural decisions to enhance success in diverse environments.

Translocated wild birds are predisposed to learn songs of their ancestral population

Population differences in socially learned mating signals like oscine birdsong are particularly vulnerable to breakdown through dispersal.1 Despite this challenge, geographic variation in learned signals is ubiquitous.2 A proposed explanation for this pattern is that birds express predispositions to selectively learn and produce population-typical songs.3,4,5 While experimental studies on lab-reared birds have shown the existence of within-species learning predispositions,6,7,8,9,10 it remains unclear whether and how learning predispositions influence song acquisition in the wild. Here, we investigated innate song learning predispositions in wild pied flycatchers (Ficedula hypoleuca) by measuring the songs of individuals translocated as eggs from a Dutch population to a breeding population in Sweden. We compared the songs of the adult males hatched from these translocated eggs with those from the ancestral and receiving populations. Songs of translocated males closely resemble the local Swedish songs to which they were exposed during development, supporting the importance of social learning. However, translocated males selectively learned those local Swedish song elements that sound the most "Dutch-like." As a result, their songs are significantly shifted toward those of the ancestral Dutch population. This suggests that innate learning predispositions track ongoing song evolution in wild populations of pied flycatchers. We propose that as songs continue to diverge over time, this coevolutionary relationship between song and learning predispositions may contribute to the emergence of incipient pre-mating barriers.

The ecology of zebra finch song and its implications for vocal communication in multi-level societies.

Acoustic signalling is crucial in affecting movements and in social interactions. In species with dynamic social structures, such as multi-level societies, acoustic signals can provide a key mechanism allowing individuals to identify and find or avoid each other and to exchange information. Yet, if the spacing between individuals regularly exceeds the maximum signalling range, the relation between movements and signals becomes more complex. As the best-studied songbird in captivity, the zebra finch (Taeniopygia castanotis) is a species with individually distinct songs that are audible over just a few metres and a widely ranging dynamic multi-level social organization in the wild, raising questions on the actual role of its song in social cohesion and coordination. Here, we provide an overview of birdsong in social organizations (networks) and use the ecology of the zebra finch and male song to discuss how singing can facilitate social cohesion and coordination in species where the signal range is very short. We raise the question of the extent to which zebra finches are a representative species to understand the function of song in communication, and we broaden current views on the function of birdsong and its individual signature. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

Clarification of taxonomic status of Pseudochorthippus geminus (Mistshenko, 1951) (Orthoptera: Acrididae: Gomphocerinae) based on male calling song analysis

Pseudochorthippus geminus is redescribed and illustrated based on specimens from southern Kazakhstan and northern Kyrgyzstan. Oscillograms of the male calling songs of P. geminus from these localities and of P. parallelus from southern Kazakhstan are given. Comparative analysis of the song temporal patterns of P. geminus and P. parallelus clearly indicates that they represent separate biological species, but the morphological differences between them are indistinct and do not always provide reliable characters for species diagnostics.