Social experiences shape song preference learning independently of developmental exposure to song.

Communication governs the formation and maintenance of social relationships. The interpretation of communication signals depends not only on the signal's content but also on a receiver's individual experience. Experiences throughout life may interact to affect behavioural plasticity, such that a lack of developmental sensory exposure could constrain adult learning, while salient adult social experiences could remedy developmental deficits. We investigated how experiences impact the formation and direction of female auditory preferences in the zebra finch. Zebra finches form long-lasting pair bonds and females learn preferences for their mate's vocalizations. We found that after 2 weeks of cohabitation with a male, females formed pair bonds and learned to prefer their partner's song regardless of whether they were reared with ('normally reared') or without ('song-naive') developmental exposure to song. In contrast, females that heard but did not physically interact with a male did not prefer his song. In addition, previous work has found that song-naive females do not show species-typical preferences for courtship song. We found that cohabitation with a male ameliorated this difference in preference. Thus, courtship and pair bonding, but not acoustic-only interactions, strongly influence preference learning regardless of rearing experience, and may dynamically drive auditory plasticity for recognition and preference.

Automated echolocation classifiers vary in accuracy for northeastern U.S. bat species.

Acoustic surveys of bat echolocation calls are an important management tool for determining presence and probable absence of threatened and endangered bat species. In the northeastern United States, software programs such as Bat Call Identification (BCID), Kaleidoscope Pro (KPro), and Sonobat can automatically classify ultrasonic detector sound files, yet the programs' accuracy in correctly classifying calls to species has not been independently assessed. We used 1,500 full-spectrum reference calls with known identities for nine northeastern United States bat species to test the accuracy of these programs using calculations of Positive Predictive Value (PPV), Negative Predictive Value (NPV), Sensitivity (SN), Specificity (SP), Overall Accuracy, and No Information Rate. We found that BCID performed less accurately than other programs, likely because it only operates on zero-crossing data and may be less accurate for recordings converted from full-spectrum to zero-crossing. NPV and SP values were high across all species categories for SonoBat and KPro, indicating these programs' success at avoiding false positives. However, PPV and SN values were relatively low, particularly for individual Myotis species, indicating these programs are prone to false negatives. SonoBat and KPro performed better when distinguishing Myotis species from non-Myotis species. We expect less accuracy from these programs for acoustic recordings collected under normal working conditions, and caution that a bat acoustic expert should verify automatically classified files when making species-specific regulatory or conservation decisions.

Machine learning with taxonomic family delimitation aids in the classification of ephemeral beaked whale events in passive acoustic monitoring.

Passive acoustic monitoring is an essential tool for studying beaked whale populations. This approach can monitor elusive and pelagic species, but the volume of data it generates has overwhelmed researchers' ability to quantify species occurrence for effective conservation and management efforts. Automation of data processing is crucial, and machine learning algorithms can rapidly identify species using their sounds. Beaked whale acoustic events, often infrequent and ephemeral, can be missed when co-occurring with signals of more abundant, and acoustically active species that dominate acoustic recordings. Prior efforts on large-scale classification of beaked whale signals with deep neural networks (DNNs) have approached the class as one of many classes, including other odontocete species and anthropogenic signals. That approach tends to miss ephemeral events in favor of more common and dominant classes. Here, we describe a DNN method for improved classification of beaked whale species using an extensive dataset from the western North Atlantic. We demonstrate that by training a DNN to focus on the taxonomic family of beaked whales, ephemeral events were correctly and efficiently identified to species, even with few echolocation clicks. By retrieving ephemeral events, this method can support improved estimation of beaked whale occurrence in regions of high odontocete acoustic activity.

Multimodal communication and audience directedness in the greeting behaviour of semi-captive African savannah elephants.

Many species communicate by combining signals into multimodal combinations. Elephants live in multi-level societies where individuals regularly separate and reunite. Upon reunion, elephants often engage in elaborate greeting rituals, where they use vocalisations and body acts produced with different body parts and of various sensory modalities (e.g., audible, tactile). However, whether these body acts represent communicative gestures and whether elephants combine vocalisations and gestures during greeting is still unknown. Here we use separation-reunion events to explore the greeting behaviour of semi-captive elephants (Loxodonta africana). We investigate whether elephants use silent-visual, audible, and tactile gestures directing them at their audience based on their state of visual attention and how they combine these gestures with vocalisations during greeting. We show that elephants select gesture modality appropriately according to their audience's visual attention, suggesting evidence of first-order intentional communicative use. We further show that elephants integrate vocalisations and gestures into different combinations and orders. The most frequent combination consists of rumble vocalisations with ear-flapping gestures, used most often between females. By showing that a species evolutionarily distant to our own primate lineage shows sensitivity to their audience's visual attention in their gesturing and combines gestures with vocalisations, our study advances our understanding of the emergence of first-order intentionality and multimodal communication across taxa.

Extensive data engineering to the rescue: building a multi-species katydid detector from unbalanced, atypical training datasets.

Passive acoustic monitoring (PAM) is a powerful tool for studying ecosystems. However, its effective application in tropical environments, particularly for insects, poses distinct challenges. Neotropical katydids produce complex species-specific calls, spanning mere milliseconds to seconds and spread across broad audible and ultrasonic frequencies. However, subtle differences in inter-pulse intervals or central frequencies are often the only discriminatory traits. These extremities, coupled with low source levels and susceptibility to masking by ambient noise, challenge species identification in PAM recordings. This study aimed to develop a deep learning-based solution to automate the recognition of 31 katydid species of interest in a biodiverse Panamanian forest with over 80 katydid species. Besides the innate challenges, our efforts were also encumbered by a limited and imbalanced initial training dataset comprising domain-mismatched recordings. To overcome these, we applied rigorous data engineering, improving input variance through controlled playback re-recordings and by employing physics-based data augmentation techniques, and tuning signal-processing, model and training parameters to produce a custom well-fit solution. Methods developed here are incorporated into Koogu, an open-source Python-based toolbox for developing deep learning-based bioacoustic analysis solutions. The parametric implementations offer a valuable resource, enhancing the capabilities of PAM for studying insects in tropical ecosystems. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Time and habitat structure shape insect acoustic activity in the Amazon.

Insects are the most diverse animal taxon on Earth and play a key role in ecosystem functioning. However, they are often neglected by ecological surveys owing to the difficulties involved in monitoring this small and hyper-diverse taxon. With technological advances in biomonitoring and analytical methods, these shortcomings may finally be addressed. Here, we performed passive acoustic monitoring at 141 sites (eight habitats) to investigate insect acoustic activity in the Viruá National Park, Brazil. We first describe the frequency range occupied by three soniferous insect groups (cicadas, crickets and katydids) to calculate the acoustic evenness index (AEI). Then, we assess how AEI varies spatially and temporally among habitat types, and finally we investigate the relationship between vegetation structure variables and AEI for each insect category. Overall, crickets occupied lower and narrower frequency bands than cicadas and katydids. AEI values varied among insect categories and across space and time. The highest acoustic activity occurred before sunrise and the lowest acoustic activity was recorded in pastures. Canopy cover was positively associated with cricket acoustic activity but not with katydids. Our findings contribute to a better understanding of the role of time, habitat and vegetation structure in shaping insect activity within diverse Amazonian ecosystems. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Towards acoustic monitoring of bees: wingbeat sounds are related to species and individual traits.

Global pollinator decline urgently requires effective methods to assess their trends, distribution and behaviour. Passive acoustics is a non-invasive and cost-efficient monitoring tool increasingly employed for monitoring animal communities. However, insect sounds remain highly unexplored, hindering the application of this technique for pollinators. To overcome this shortfall and support future developments, we recorded and characterized wingbeat sounds of a variety of Iberian domestic and wild bees and tested their relationship with taxonomic, morphological, behavioural and environmental traits at inter- and intra-specific levels. Using directional microphones and machine learning, we shed light on the acoustic signature of bee wingbeat sounds and their potential to be used for species identification and monitoring. Our results revealed that frequency of wingbeat sounds is negatively related with body size and environmental temperature (between-species analysis), while it is positively related with experimentally induced stress conditions (within-individual analysis). We also found a characteristic acoustic signature in the European honeybee that supported automated classification of this bee from a pool of wild bees, paving the way for passive acoustic monitoring of pollinators. Overall, these findings confirm that insect sounds during flight activity can provide insights on individual and species traits, and hence suggest novel and promising applications for this endangered animal group. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Multi-year soundscape recordings and automated call detection reveals varied impact of moonlight on calling activity of neotropical forest katydids.

Night-time light can have profound ecological effects, even when the source is natural moonlight. The impacts of light can, however, vary substantially by taxon, habitat and geographical region. We used a custom machine learning model built with the Python package Koogu to investigate the in situ effects of moonlight on the calling activity of neotropical forest katydids over multiple years. We prioritised species with calls that were commonly detected in human annotated data, enabling us to evaluate model performance. We focused on eight species of katydids that the model identified with high precision (generally greater than 0.90) and moderate-to-high recall (minimum 0.35), ensuring that detections were generally correct and that many calls were detected. These results suggest that moonlight has modest effects on the amount of calling, with the magnitude and direction of effect varying by species: half of the species showed positive effects of light and half showed negative. These findings emphasize the importance of understanding natural history for anticipating how biological communities respond to moonlight. The methods applied in this project highlight the emerging opportunities for evaluating large quantities of data with machine learning models to address ecological questions over space and time. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Hydroacoustic study of fin whales around the Southern Wake Island: Type, vocal behavior, and temporal evolution from 2010 to 2022.

The progress of fin whale study is hindered by the debate about whether the two typical type-A and type-B calls (characterized by central source frequencies of 17-20 Hz and 20-30 Hz, respectively) originate from a single fin whale or two individual fin whales. Here, hydroacoustic data is employed to study the type, vocal behavior, and temporal evolution of fin whale calls around the Southern Wake Island from 2010 to 2022. It is identified that (1) type-A and type-B calls come from two individuals based on the large source separation of the two calls through high-precision determination of source location; (2) type-A fin whales exhibit vocal influence on type-B fin whales, where type-B fin whales become paired with type-A calls and vocalize regularly when type-A fin whales appear, and type-A fin whales always lead the call sequences; and (3) some type-A fin whales stop calling when another type-A fin whale approaches at a distance of about 1.6 km. During 2010-2022, type-A calls occur every year, whereas type-B calls are prevalent only after November 2018. A culture transmission is proposed from type-A fin whales to type-B fin whales and/or a population increase of type-B fin whales in the region after November 2018.

Daily singing of adult songbirds functions to maintain song performance independently of auditory feedback and age.

Many songbirds learn to produce songs through vocal practice in early life and continue to sing daily throughout their lifetime. While it is well-known that adult songbirds sing as part of their mating rituals, the functions of singing behavior outside of reproductive contexts remain unclear. Here, we investigated this issue in adult male zebra finches by suppressing their daily singing for two weeks and examining the effects on song performance. We found that singing suppression decreased the pitch, amplitude, and duration of songs, and that those song features substantially recovered through subsequent free singing. These reversible song changes were not dependent on auditory feedback or the age of the birds, contrasting with the adult song plasticity that has been reported previously. These results demonstrate that adult song structure is not stable without daily singing, and suggest that adult songbirds maintain song performance by preventing song changes through physical act of daily singing throughout their life. Such daily singing likely functions as vocal training to maintain the song production system in optimal conditions for song performance in reproductive contexts, similar to how human singers and athletes practice daily to maintain their performance.

Vocal complexity in the long calls of Bornean orangutans.

Vocal complexity is central to many evolutionary hypotheses about animal communication. Yet, quantifying and comparing complexity remains a challenge, particularly when vocal types are highly graded. Male Bornean orangutans (Pongo pygmaeus wurmbii) produce complex and variable "long call" vocalizations comprising multiple sound types that vary within and among individuals. Previous studies described six distinct call (or pulse) types within these complex vocalizations, but none quantified their discreteness or the ability of human observers to reliably classify them. We studied the long calls of 13 individuals to: (1) evaluate and quantify the reliability of audio-visual classification by three well-trained observers, (2) distinguish among call types using supervised classification and unsupervised clustering, and (3) compare the performance of different feature sets. Using 46 acoustic features, we used machine learning (i.e., support vector machines, affinity propagation, and fuzzy c-means) to identify call types and assess their discreteness. We additionally used Uniform Manifold Approximation and Projection (UMAP) to visualize the separation of pulses using both extracted features and spectrogram representations. Supervised approaches showed low inter-observer reliability and poor classification accuracy, indicating that pulse types were not discrete. We propose an updated pulse classification approach that is highly reproducible across observers and exhibits strong classification accuracy using support vector machines. Although the low number of call types suggests long calls are fairly simple, the continuous gradation of sounds seems to greatly boost the complexity of this system. This work responds to calls for more quantitative research to define call types and quantify gradedness in animal vocal systems and highlights the need for a more comprehensive framework for studying vocal complexity vis-à-vis graded repertoires.

Exploring the role of vocalizations in regulating group dynamics.

Because of the diverging needs of individuals, group life can lead to disputes and competition, but it also has many advantages, such as reduced predation risk, information sharing and increased hunting success. Social animals have to maintain group cohesion and need to synchronize activities, such as foraging, resting, social interactions and movements, in order to thrive in groups. Acoustic signals are highly relevant for social dynamics, some because they are long-ranging and others because they are short-ranging, which may serve important within-group functions. However, although there has been an increase in studies concentrating on acoustic communication within groups in the past decade, many aspects of how vocalizations relate to group dynamics are still poorly understood. The aim of this review is to present an overview of our current knowledge on the role of vocalizations in regulating social group dynamics, identify knowledge gaps and recommend potential future research directions. We review the role that vocalizations play in (i) collective movement, (ii) separation risk and cohesion maintenance, (iii) fission-fusion dynamics, and (iv) social networks. We recommend that future studies aim to increase the diversity of studied species and strengthen the integration of state-of-the-art tools to study social dynamics and acoustic signals. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

The power of sound: unravelling how acoustic communication shapes group dynamics.

Acoustic signalling is a key mode of communication owing to its instantaneousness and rapid turnover, its saliency and flexibility and its ability to function strategically in both short- and long-range contexts. Acoustic communication is closely intertwined with both collective behaviour and social network structure, as it can facilitate the coordination of collective decisions and behaviour, and play an important role in establishing, maintaining and modifying social relationships. These research topics have each been studied separately and represent three well-established research areas. Yet, despite the close connection of acoustic communication with collective behaviour and social networks in natural systems, only few studies have focused on their interaction. The aim of this theme issue is therefore to build a foundation for understanding how acoustic communication is linked to collective behaviour, on the one hand, and social network structure on the other, in non-human animals. Through the building of such a foundation, our hope is that new questions in new avenues of research will arise. Understanding the links between acoustic communication and social behaviour seems crucial for gaining a comprehensive understanding of sociality and social evolution. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

Sex differences in the impact of social relationships on individual vocal signatures in grey mouse lemurs (Microcebus murinus).

Vocalizations coordinate social interactions between conspecifics by conveying information concerning the individual or group identity of the sender. Social accommodation is a form of vocal learning where social affinity is signalled by converging or diverging vocalizations with those of conspecifics. To investigate whether social accommodation is linked to the social lifestyle of the sender, we investigated sex-specific differences in social accommodation in a dispersed living primate, the grey mouse lemur (Microcebus murinus), where females form stable sleeping groups whereas males live solitarily. We used 482 trill calls of 36 individuals from our captive breeding colony to compare acoustic dissimilarity between individuals with genetic relatedness, social contact time and body weight. Our results showed that female trills become more similar the more time females spend with each other, independent of genetic relationship, suggesting vocal convergence. In contrast, male trills were affected more by genetic than social factors. However, focusing only on socialized males, increasing time as cage partners caused greater divergence in males' trills. Thus, grey mouse lemurs show the capacity for social accommodation, with females converging their trills to signal social closeness to sleeping group partners, whereas males do not adapt or diverge their trills to signal individual distinctiveness. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

In pop pursuit: social bond strength predicts vocal synchrony during cooperative mate guarding in bottlenose dolphins.

Vocal communication is an emblematic feature of group-living animals, used to share information and strengthen social bonds. Vocalizations are also used to coordinate group-level behaviours in many taxa, but little is known of the factors that may influence vocal behaviour during cooperative acts. Allied male Indo-Pacific bottlenose dolphins (Tursiops aduncus) use the 'pop' vocalization as a coercive signal when working together to herd single oestrous females. Using long-term association and acoustic data, we examined the influence of social and non-social factors on pop use by allied male dolphins in this context. Neither pop rate nor pop bout duration were influenced by any of the factors examined. However, allied males with stronger social bonds engaged in higher rates of vocal synchrony; whereby they actively matched the timing of their pop production. Hence, social bond strength influenced pop use in a cooperative context, suggesting dual functions of pop use: to induce the female to remain close, and to promote social bond maintenance and cooperation among males. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

Not your private tête-à-tête: leveraging the power of higher-order networks to study animal communication.

Animal communication is frequently studied with conventional network representations that link pairs of individuals who interact, for example, through vocalization. However, acoustic signals often have multiple simultaneous receivers, or receivers integrate information from multiple signallers, meaning these interactions are not dyadic. Additionally, non-dyadic social structures often shape an individual's behavioural response to vocal communication. Recently, major advances have been made in the study of these non-dyadic, higher-order networks (e.g. hypergraphs and simplicial complexes). Here, we show how these approaches can provide new insights into vocal communication through three case studies that illustrate how higher-order network models can: (i) alter predictions made about the outcome of vocally coordinated group departures; (ii) generate different patterns of song synchronization from models that only include dyadic interactions; and (iii) inform models of cultural evolution of vocal communication. Together, our examples highlight the potential power of higher-order networks to study animal vocal communication. We then build on our case studies to identify key challenges in applying higher-order network approaches in this context and outline important research questions that these techniques could help answer. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

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'.

Calling to the collective: contact calling rates within groups of disc-winged bats do not vary by kinship or association.

Many group-living animals coordinate social behaviours using contact calls, which can be produced for all group members or targeted at specific individuals. In the disc-winged bat, Thyroptera tricolor, group members use 'inquiry' and 'response' calls to coordinate daily movements into new roosts (furled leaves). Rates of both calls show consistent among-individual variation, but causes of within-individual variation remain unknown. Here, we tested whether disc-winged bats produce more contact calls towards group members with higher kinship or association. In 446 experimental trials, we recorded 139 random within-group pairs of one flying bat (producing inquiry calls for roost searching) and one roosting bat (producing response calls for roost advertising). Using generalized linear mixed-effect models (GLMM), we assessed how response and inquiry calling rates varied by sender, receiver, genetic kinship and co-roosting association rate. Calling rates varied consistently across senders but not by receiver. Response calling was influenced by inquiry calling rates, but neither calling rate was higher when the interacting pair had higher kinship or association. Rather than dyadic calling rates indicating within-group relationships, our findings are consistent with the hypothesis that bats produce contact calls to maintain contact with any or all individuals within a group while collectively searching for a new roost site. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

Hidden social complexity behind vocal and acoustic communication in non-avian reptiles.

Social interactions are inevitable in the lives of most animals, since most essential behaviours require interaction with conspecifics, such as mating and competing for resources. Non-avian reptiles are typically viewed as solitary animals that predominantly use their vision and olfaction to communicate with conspecifics. Nevertheless, in recent years, evidence is mounting that some reptiles can produce sounds and have the potential for acoustic communication. Reptiles that can produce sound have an additional communicative channel (in addition to visual/olfactory channels), which could suggest they have a higher communicative complexity, the evolution of which is assumed to be driven by the need of social interactions. Thus, acoustic reptiles may provide an opportunity to unveil the true social complexity of reptiles that are usually thought of as solitary. This review aims to reveal the hidden social interactions behind the use of sounds in non-avian reptiles. Our review suggests that the potential of vocal and acoustic communication and the complexity of social interactions may be underestimated in non-avian reptiles, and that acoustic reptiles may provide a great opportunity to uncover the coevolution between sociality and communication in non-avian reptiles. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

The relative contribution of acoustic signals versus movement cues in group coordination and collective decision-making.

To benefit from group living, individuals need to maintain cohesion and coordinate their activities. Effective communication thus becomes critical, facilitating rapid coordination of behaviours and reducing consensus costs when group members have differing needs and information. In many bird and mammal species, collective decisions rely on acoustic signals in some contexts but on movement cues in others. Yet, to date, there is no clear conceptual framework that predicts when decisions should evolve to be based on acoustic signals versus movement cues. Here, we first review how acoustic signals and movement cues are used for coordinating activities. We then outline how information masking, discrimination ability (Weber's Law) and encoding limitations, as well as trade-offs between these, can identify which types of collective behaviours likely rely on acoustic signals or movement cues. Specifically, our framework proposes that behaviours involving the timing of events or expression of specific actions should rely more on acoustic signals, whereas decisions involving complex choices with multiple options (e.g. direction and destination) should generally use movement cues because sounds are more vulnerable to information masking and Weber's Law effects. We then discuss potential future avenues of enquiry, including multimodal communication and collective decision-making by mixed-species animal groups. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamic'.