Singing Behavior Research Articles

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.

The hidden fitness of the male zebra finch courtship song.

Vocal learning in songbirds is thought to have evolved through sexual selection, with female preference driving males to develop large and varied song repertoires1-3. However, many songbird species learn only a single song in their lifetime4. How sexual selection drives the evolution of single-song repertoires is not known. Here, by applying dimensionality-reduction techniques to the singing behaviour of zebra finches (Taeniopygia guttata), we show that syllable spread in low-dimensional feature space explains how single songs function as honest indicators of fitness. We find that this Gestalt measure of behaviour captures the spectrotemporal distinctiveness of song syllables in zebra finches; that females strongly prefer songs that occupy more latent space; and that matching path lengths in low-dimensional space is difficult for young males. Our findings clarify how simple vocal repertoires may have evolved in songbirds and indicate divergent strategies for how sexual selection can shape vocal learning.

Music production and its role in coalition signaling during foraging contexts in a hunter-gatherer society.

Music is a cultural activity universally present in all human societies. Several hypotheses have been formulated to understand the possible origins of music and the reasons for its emergence. Here, we test two hypotheses: (1) the coalition signaling hypothesis which posits that music could have emerged as a tool to signal cooperative intent and signal strength of alliances and (2) music as a strategy to deter potential predators. In addition, we further explore the link between tactile cues and the propensity of mothers to sing toward infants. For this, we investigated the singing behaviors of hunter-gatherer mothers during daily foraging trips among the Mbendjele BaYaka in the Republic of the Congo. Although singing is a significant component of their daily activities, such as when walking in the forest or collecting food sources, studies on human music production in hunter-gatherer societies are mostly conducted during their ritual ceremonies. In this study, we collected foraging and singing behavioral data of mothers by using focal follows of five BaYaka women during their foraging trips in the forest. In accordance with our predictions for the coalition signaling hypothesis, women were more likely to sing when present in large groups, especially when group members were less familiar. However, predictions of the predation deterrence hypothesis were not supported as the interaction between group size and distance from the village did not have a significant effect on the likelihood of singing. The latter may be due to limited variation in predation risk in the foraging areas, because of the intense bush meat trade, and hence, future studies should include foraging areas with higher densities of wild animals. Lastly, we found that mothers were more likely to sing when they were carrying infants compared to when infants were close, but carried by others, supporting the prediction that touch plays an important prerequisite role in musical interaction between the mother and child. Our study provides important insight into the role of music as a tool in displaying the intent between or within groups to strengthen potentially conflict-free alliances during joint foraging activities.

Humpback whales increase the length of their songs during nearby airgun operations

There is concern that noise from “airguns” used during oil and gas exploration may cause behavioral changes in marine mammals. One important behavior of some mysticete whales is the production of songs which are likely used as reproductive displays. Humpback whales have conspicuous songs but there are few studies on the impact of noise on their singing behavior. Here, we test the hypothesis that airgun noise causes individual humpback whales to reduce the duration of their songs. In a series of experiments off the east coast of Australia, we exposed migrating humpback whales to airguns. We tracked and recorded singing males when we towed airguns through the study area as well as when there were no vessels or airguns present. We also noted interactions between singers and nearby conspecifics. Humpbacks usually stop singing when they join with others, and so joining is both a predictor of song duration and a biologically relevant outcome. Contrary to expectations, whales that were already singing prior to airgun exposure produced significantly longer songs than unexposed singers. However, the small number of whales that started singing during airgun exposure produced significantly shorter songs.

Distinct patterns of gene expression in the medial preoptic area are related to gregarious singing behavior in European starlings (Sturnus vulgaris)

BackgroundSong performed in flocks by European starlings (Sturnus vulgaris), referred to here as gregarious song, is a non-sexual, social behavior performed by adult birds. Gregarious song is thought to be an intrinsically reinforced behavior facilitated by a low-stress, positive affective state that increases social cohesion within a flock. The medial preoptic area (mPOA) is a region known to have a role in the production of gregarious song. However, the neurochemical systems that potentially act within this region to regulate song remain largely unexplored. In this study, we used RNA sequencing to characterize patterns of gene expression in the mPOA of male and female starlings singing gregarious song to identify possibly novel neurotransmitter, neuromodulator, and hormonal pathways that may be involved in the production of gregarious song.ResultsDifferential gene expression analysis and rank rank hypergeometric analysis indicated that dopaminergic, cholinergic, and GABAergic systems were associated with the production of gregarious song, with multiple receptor genes (e.g., DRD2, DRD5, CHRM4, GABRD) upregulated in the mPOA of starlings who sang at high rates. Additionally, co-expression network analyses identified co-expressing gene clusters of glutamate signaling-related genes associated with song. One of these clusters contained five glutamate receptor genes and two glutamate scaffolding genes and was significantly enriched for genetic pathways involved in neurodevelopmental disorders associated with social deficits in humans. Two of these genes, GRIN1 and SHANK2, were positively correlated with performance of gregarious song.ConclusionsThis work provides new insights into the role of the mPOA in non-sexual, gregarious song in starlings and highlights candidate genes that may play a role in gregarious social interactions across vertebrates. The provided data will also allow other researchers to compare across species to identify conserved systems that regulate social behavior.

Females occasionally create duets with males but they never sing solo-year-round singing behaviour in an Afrotropical songbird

Our knowledge of birdsong mainly comes from studies focused on male songs produced in a short breeding period, even though we know that sedentary species sing year-round, female song is quite widespread and many species sing collectively creating duets and choruses. In this study we focused on daily and seasonal changes in singing activity of an endemic, sedentary, duetting, Afrotropical songbird—the Bangwa forest warbler. We collected soundscape recordings in six recording locations and used singing activity index to examine how vocal activity of males and females varies daily and seasonally and how it correlates with the rainfall. We found that Bangwa forest warblers sing year-round, yet they do it more in wet than in dry season. The rapid increase of singing activity occurs after first rain, at the beginning of the rainy season. Males sing significantly more than females. Females never sing solo, however, in 13% of songs they create duets by joining male solos. The pattern of daily singing activity is sex-specific and seasonally variable, with two peaks (dawn and dusk) observed in males and only one in females (dawn). In Bangwa forest warbler male singing behaviour is similar to that of many songbirds, suggesting that territory defence and female attraction as main functions of singing. Females, which create duets and never sing solo may use songs in mate guarding, signalling commitment, resource defence or intersex territory defence. Duets observed year-round may suggest cooperative resource defence. Results of the study show that examining year-round singing behaviour is crucial to fully understand the evolution and functions of male and female songs.

Interspecific avoidance of song overlap in tropical songbirds: species-specific responses to acoustically similar and different intruders

To ensure effective acoustic communication, signals should reach receivers in the least distorted form possible. Animals use various short- and long-term strategies to avoid signal degradation and masking. However, we still have an insufficient understanding of how animals’ vocal behaviour is impacted by the vocalisations of other animals in their acoustic communities. We experimentally examined how two tropical, sedentary, territorial songbirds in Western Uganda—the scaly-breasted illadopsis (Illadopsis albipectus) and the green-backed camaroptera (Camaroptera brachyura)—modify their singing behaviour after the simulated appearance of new, unfamiliar acoustic competitors, whose songs vary in similarity to those of the species studied. We found that scaly-breasted illadopsis sang significantly less during the playback of songs of acoustically similar species than of acoustically different species or silence and avoided song overlapping with acoustically similar species but not with acoustically different species. Green-backed camaroptera sang significantly more during the playback of both acoustically similar and different simulated intruders than during the control containing silence, and patterns of overlap with the songs of both the acoustically similar and different species were random. Our results show that even a single-point noise source present within a territory can modify a bird’s singing behaviour. The new sound may affect species differently, depending in part on the level of acoustic similarity with the species’ song. To mitigate the effect of song masking, different species may use different strategies, such as temporal avoidance or signal redundancy. Studies examining the adaptive abilities of species in natural and modified habitats are needed to predict the consequences of changes in acoustic community structure.Significance statementTo ensure effective communication, birds may use different strategies to avoid signal masking in common acoustic space, particularly in the complex acoustic environment of a tropical forest. While multiple studies have focused on responses to interference caused by anthropogenic noise, the effect of new individual species on the acoustic community structure has received little attention. We simulated intrusions by unfamiliar species with different levels of song similarity into the territories of two tropical songbird species. The appearance of new simulated acoustic intruders modified the birds’ singing behaviour, but the two study species responded differently. These results suggest that the level of acoustic similarity, as well as the species ecology, may affect the species response, which may be particularly important when predicting the effects of new species appearance as a result of changes in habitat and climate.

The relationships of breeding stage to daytime singing behaviour and song perch height in Bermuda white‐eyed vireos Vireo griseus bermudianus

Bird song is crucial for attracting mates and defending territories, but different types of song or different singing behaviours may be involved in acquiring or maintaining each resource. Furthermore, male songbirds may adjust when and where they sing throughout the breeding season, depending on their breeding stage. However, such relationships remain untested in several avian taxa. Here, we studied male Bermuda white‐eyed vireos Vireo griseus bermudianus, a passerine with two distinct song types (discrete and rambling), to test the mate attraction and territory defence hypotheses. We compare song production and song perch height among different stages of the breeding season and during the non‐breeding season. We show that male vireos produce both song types during the breeding and non‐breeding seasons, suggesting dual roles in mate choice and territorial defence. Song production did not differ significantly between the breeding and non‐breeding seasons, but, within the breeding season, males without nesting duties sang significantly more songs than males with nesting duties. Song perch height was higher during the breeding season versus non‐breeding season, among males without nesting duties compared to males with nesting duties, and when males produced discrete versus rambling songs. Our findings suggest that male vireos increase their conspicuousness to prospecting females by increasing song production and song perch height, and that they sing during the breeding and non‐breeding seasons to defend year‐round territories. Collectively, our study supports the mate attraction and territory defence hypotheses of bird song.

Dehydroepiandrosterone (DHEA) increases undirected singing behavior and alters dopaminergic regulation of undirected song in non-breeding male European starlings (Sturnus vulgaris).

It has been proposed that in species that defend territories across multiple life history stages, brain metabolism of adrenal dehydroepiandrosterone (DHEA) regulates aggressive behavior at times when gonadal androgen synthesis is low (i.e. the non-breeding season). To date, a role for DHEA in the regulation of other forms of social behavior that are expressed outside of the context of breeding remains unknown. In this experiment, we used the European starling (Sturnus vulgaris) model system to investigate a role for DHEA in the neuroendocrine regulation of singing behavior by males in non-breeding condition. Starling song in a non-breeding context is spontaneous, not directed towards conspecifics, and functions to maintain cohesion of overwintering flocks. Using within-subjects design, we found that DHEA implants significantly increase undirected singing behavior by non-breeding condition male starlings. Given that DHEA is known to modulate multiple neurotransmitter systems including dopamine (DA) and DA regulates undirected song, we subsequently used immunohistochemistry for phosphorylated tyrosine hydroxylase (pTH, the active form of the rate-limiting enzyme in DA synthesis) to investigate the effect of DHEA on dopaminergic regulation of singing behavior in a non-breeding context. Pearson correlation analysis revealed a positive linear association between undirected singing behavior and pTH immunoreactivity in the ventral tegmental area and midbrain central gray of DHEA-implanted, but not control-implanted, males. Taken together, these data suggest that undirected singing behavior by non-breeding starlings is modulated by effects of DHEA on dopaminergic neurotransmission. More broadly, these data expand the social behavior functions of DHEA beyond territorial aggression to include undirected, affiliative social communication.

Frequent duets, rare choruses, and extremely rare solos: year-round singing behavior in Chubb’s Cisticola

Our current understanding of the function of coordinated acoustic displays usually comes from studies conducted over a short period of the breeding season. However, the function of particular types of vocalizations may vary according to sex and context, and such displays can extend beyond the time of reproduction. To fully understand this phenomenon, analyses of year-round singing behavior are required. In the current study, we focused on a small, year-round territorial Afrotropical songbird, Chubb’s Cisticola (Cisticola chubbi). We analyzed the structure of songs during the breeding season as well as year-round changes in the proportion of solos, duets, and choruses to investigate the potential function(s) of each type of vocalization. We found that: (1) females produced whistling notes, while males generated trilling ones; (2) up to five individuals formed coordinated choruses, and (3) individuals were always near to each other during cooperative singing. Over the course of a year, the majority of syllables recorded were duets (82%), with rarer choruses (16%) and extremely rare solos (2%). Outside of the breeding season, males produced the most solos, while females produced more at the beginning of the breeding season. The proportion of choruses was highest at the end of breeding season. Frequent year-round production of duets and choruses strongly supports territory defense as the main function of joint singing, while the highest proportion of choruses at the end of the breeding season suggests that offspring take part in the chorus. To better understand cooperative singing, it is essential to extend our looking beyond the breeding season.

Estradiol decreases the excitability of RA projection neurons in adult male zebra finches.

Zebra finches are essential animal models for studying learned vocal signals. The robust nucleus of the arcopallium (RA) plays an important role in regulating singing behavior. Our previous study showed that castration inhibited the electrophysiological activity of RA projection neurons (PNs) in male zebra finches, demonstrating that testosterone modulates the excitability of RA PNs. Testosterone can be converted into estradiol (E2) in the brain through aromatase; however, the physiological functions of E2 in RA are still unknown. This study aimed to investigate the electrophysiological activities of E2 on the RA PNs of male zebra finches through patch-clamp recording. E2 rapidly decreased the rate of evoked and spontaneous action potentials (APs) of RA PNs, hyperpolarized the resting membrane potential, and decreased the membrane input resistance. Moreover, the G-protein-coupled membrane-bound estrogen receptor (GPER) agonist G1 decreased both the evoked and spontaneous APs of RA PNs. Furthermore, the GPER antagonist G15 had no effect on the evoked and spontaneous APs of RA PNs; E2 and G15 together also had no effect on the evoked and spontaneous APs of RA PNs. These findings suggested that E2 rapidly decreased the excitability of RA PNs and its binding to GPER suppressed the excitability of RA PNs. These pieces of evidence helped us fully understand the principle of E2 signal mediation via its receptors to modulate the excitability of RA PNs in songbirds.

Changes in humpback whale song structure and complexity reveal a rapid evolution on a feeding ground in Northern Norway

Singing behaviour by male humpback whales (Megaptera novaeangliae) has traditionally been associated with low-latitude breeding grounds. However, in recent years, this vocal behaviour has been increasingly reported outside these areas. All singers in a given population sing the same version of a song and this song is continually evolving over time with modifications on different levels within the song structure. Tracing changes in whale song will help to undercover the drivers underlying this vocal display and contribute to the understanding of animal culture and its evolution. To determine the progressive changes in songs found on a subarctic feeding ground and migratory stopover, a detailed analysis of humpback whale song recordings from Northern Norway was conducted. Passive acoustic data from the Lofoten-Vesterålen Ocean Observatory (LoVe), collected using a bottom-moored underwater hydrophone, were used from January - April 2018 and January 2019. Two measures of the song structure were examined: (1) sequence similarities using the Levenshtein distance and (2) song complexity using a principal component analysis (PCA). In total, 21 distinct themes were identified which presented highly directional, structural changes over time. Two themes from 2018 reoccurred in 2019, whereas all other themes in 2019 appeared to be evolved versions of 2018 themes. All songs grouped into three general clusters, reflecting the rapid evolution over the study period. With all sampled animals singing the same version of the song, this might indicate that the singers are either from the same breeding population or that song learning occurred before the study period. Song complexity appeared to follow the trend of song progression; songs became more complex as they evolved over the months in 2018 and decreased in complexity between the years, returning to a more simplified song in 2019. The results confirm that humpback whale song exhibits a rapid progression on a shared subarctic feeding ground, with strong potential for song exchange and opportunities for cultural transmission between populations in the North Atlantic.

Mu opioid receptor stimulation in the medial preoptic area or nucleus accumbens facilitates song and reward in flocking European starlings.

It has been proposed that social cohesion in gregarious animals is reinforced both by a positive affective state induced by social interactions and by the prevention of a negative state that would be caused by social separation. Opioids that bind to mu opioid receptors (MORs) act in numerous brain regions to induce positive and to reduce negative affective states. Here we explored a potential role for MORs in affective states that may impact flocking behavior in mixed-sex flocks of nonbreeding European starlings, Sturnus vulgaris. Singing behavior, which is considered central to flock cohesion, and other social behaviors were quantified after infusions of the MOR agonist D-Ala2, N-Me-Phe4, glycinol5-ENK (DAMGO) into either the medial preoptic area (POM) or the nucleus accumbens (NAC), regions previously implicated in affective state and flock cohesion. We focused on beak wiping, a potential sign of stress or redirected aggression in this species, to provide insight into a presumed negative state. We also used conditioned place preference (CPP) tests to provide insight into the extent to which infusions of DAMGO into POM or NAC that stimulated song might be rewarding. We found that MOR stimulation in either POM or NAC dose-dependently promoted singing behavior, reduced beak wiping, and induced a CPP. Subtle differences in responses to MOR stimulation between NAC and POM also suggest potential functional differences in the roles of these two regions. Finally, because the location of NAC has only recently been identified in songbirds, we additionally performed a tract tracing study that confirmed the presence of dopaminergic projections from the ventral tegmental area to NAC, suggesting homology with mammalian NAC. These findings support the possibility that MORs in POM and NAC play a dual role in reinforcing social cohesion in flocks by facilitating positive and reducing negative affective states.

Driving singing behaviour in songbirds using a multi-modal, multi-agent virtual environment

Interactive biorobotics provides unique experimental potential to study the mechanisms underlying social communication but is limited by our ability to build expressive robots that exhibit the complex behaviours of birds and small mammals. An alternative to physical robots is to use virtual environments. Here, we designed and built a modular, audio-visual 2D virtual environment that allows multi-modal, multi-agent interaction to study mechanisms underlying social communication. The strength of the system is an implementation based on event processing that allows for complex computation. We tested this system in songbirds, which provide an exceptionally powerful and tractable model system to study social communication. We show that pair-bonded zebra finches (Taeniopygia guttata) communicating through the virtual environment exhibit normal call timing behaviour, males sing female directed song and both males and females display high-intensity courtship behaviours to their mates. These results suggest that the environment provided is sufficiently natural to elicit these behavioral responses. Furthermore, as an example of complex behavioral annotation, we developed a fully unsupervised song motif detector and used it to manipulate the virtual social environment of male zebra finches based on the number of motifs sung. Our virtual environment represents a first step in real-time automatic behaviour annotation and animal–computer interaction using higher level behaviours such as song. Our unsupervised acoustic analysis eliminates the need for annotated training data thus reducing labour investment and experimenter bias.

Effects of Choir Singing on Mental Health: Results of an Online Cross-sectional Study

The study aims to quantify the impact of sociodemographic, personal, and choir-related characteristics on perceived singing well-being and mental health changes due to singing. 847 German adult choristers (233m, 614f, age 18-86 years) were interviewed in a cross-sectional online questionnaire study that included questions on singing well-being, vocal and choral characteristics, the adapted versions of the Bochum change questionnaire (BCQ2000), the Utrecht Work Engagement Scale (UWES-09), and the WHO-5 well-being index. Multiple regression models were calculated with singing related well-being or mental health changes as dependent variables and individual and choir-related characteristics as factors. Predictors of mental health benefits were analyzed in gender subgroups and in persons with a low or normal/high score on the WHO-5. On average, participants rated the impact of singing on well-being and mental health changes positively. Subgroup analysis revealed smaller but significant positive mental health effects in men compared to women and in persons with a low WHO-5 score compared to those with a normal/high score. Education level and relationship status were not significantly related to subjects' perceptions of mental health benefits, whereas singing well-being increased with age in women and in participants with a low WHO-5 score. Larger improvements in mental health came along with longer choir membership, more singing hours per week, and a high engagement in choral activity. Significant positive associations of well-being with optimal singing behavior and vocal warm-ups were observed. Results suggest that singers of all ages, genders, and educational backgrounds perceive the choral experience as beneficial to their well-being and mental health. Positive effects are related to WHO-5 scores, engagement in choral activity, and optimal singing conditions. On average, women rate singing benefits higher than men and singers with preexisting vocal pathologies or low WHO-5 benefit slightly less.

Birdsong and the Neural Regulation of Positive Emotion.

Birds are not commonly admired for emotional expression, and when they are, the focus is typically on negative states; yet vocal behavior is considered a direct reflection of an individual’s emotional state. Given that over 4000 species of songbird produce learned, complex, context-specific vocalizations, we make the case that songbirds are conspicuously broadcasting distinct positive emotional states and that hearing songs can also induce positive states in other birds. Studies are reviewed that demonstrate that that the production of sexually motivated song reflects an emotional state of anticipatory reward-seeking (i.e., mate-seeking), while outside the mating context song in gregarious flocks reflects a state of intrinsic reward. Studies are also reviewed that demonstrate that hearing song induces states of positive anticipation and reward. This review brings together numerous studies that highlight a potentially important role for the songbird nucleus accumbens, a region nearly synonymous with reward in mammals, in positive emotional states that underlie singing behavior and responses to song. It is proposed that the nucleus accumbens is part of an evolutionarily conserved circuitry that contributes context-dependently to positive emotional states that motivate and reward singing behavior and responses to song. Neural mechanisms that underlie basic emotions appear to be conserved and similar across vertebrates. Thus, these findings in songbirds have the potential to provide insights into interventions that can restore positive social interactions disrupted by mental health disorders in humans.

Effect of Darkness on Intrinsic Motivation for Undirected Singing in Bengalese Finch (Lonchura striata Domestica): A Comparative Study With Zebra Finch (Taeniopygia guttata).

The zebra finch (ZF) and the Bengalese finch (BF) are animal models that have been commonly used for neurobiological studies on vocal learning. Although they largely share the brain structure for vocal learning and production, BFs produce more complex and variable songs than ZFs, providing a great opportunity for comparative studies to understand how animals learn and control complex motor behaviors. Here, we performed a comparative study between the two species by focusing on intrinsic motivation for non-courtship singing (“undirected singing”), which is critical for the development and maintenance of song structure. A previous study has demonstrated that ZFs dramatically increase intrinsic motivation for undirected singing when singing is temporarily suppressed by a dark environment. We found that the same procedure in BFs induced the enhancement of intrinsic singing motivation to much smaller degrees than that in ZFs. Moreover, unlike ZFs that rarely sing in dark conditions, substantial portion of BFs exhibited frequent singing in darkness, implying that such “dark singing” may attenuate the enhancement of intrinsic singing motivation during dark periods. In addition, measurements of blood corticosterone levels in dark and light conditions provided evidence that although BFs have lower stress levels than ZFs in dark conditions, such lower stress levels in BFs are not the major factor responsible for their frequent dark singing. Our findings highlight behavioral and physiological differences in spontaneous singing behaviors of BFs and ZFs and provide new insights into the interactions between singing motivation, ambient light, and environmental stress.

Singing strategies are linked to perch use on foraging territories in heart-nosed bats.

Acoustic communication allows animals to coordinate and optimize resource utilization in space. Cardioderma cor, the heart‐nosed bat, is one of the few species of bats known to sing during nighttime foraging. Previous research found that heart‐nosed bats react aggressively to song playback, supporting the territorial defense hypothesis of singing in this species. We further investigated the territorial defense hypothesis from an ecological standpoint, which predicts that singing should be associated with exclusive areas containing a resource, by tracking 14 individuals nightly during the dry seasons in Tanzania. We quantified the singing behavior of individuals at all perches used throughout the night. Using home range analysis tools, we quantified overall use, night ranges and singing ranges, as well as areas used in early and later time periods at night. Males sang back and forth from small (x¯ = 3.48 ± 2.71 ha), largely exclusive areas that overlapped with overall night ranges used for gleaning prey. Individuals varied in singing effort; however, all sang significantly more as night progressed. Subsequently, areas used earlier at night and overall use areas were both larger than singing areas. Individuals varied in singing strategies. Some males sang for long periods in particular trees and had smaller core areas, while others moved frequently among singing trees. The most prolific singers used more perches overall. Our results support the hypothesis that acoustic communication repertoires evolved in support of stable foraging territory advertisement and defense in some bats.

Singing Fin Whale Swimming Behavior in the Central North Pacific

Male fin whales sing using 20 Hz pulses produced in regular patterns of inter-note intervals, but little is known about fin whale swimming behavior while they are singing. Even less is known about fin whales in Hawaiian waters because they have rarely been sighted during surveys and passive acoustic monitoring has been limited to sparse hydrophone systems that do not have localization capabilities. We hypothesized that fin whale kinematics may be related to their singing behavior, or external variables such as time and sea state. To investigate this hypothesis, we analyzed 115 tracks containing 50,034 unique notes generated from passive acoustic recordings on an array of 14 hydrophones from 2011 to 2017 at the U.S. Navy Pacific Missile Range Facility off Kauai, Hawaii. Fin whales swam at an average speed of 1.1 m/s over relatively direct paths. We incorporated the whales' speed and turning angle into hidden Markov models to identify different behavioral states based on the whales' movements. We found that fin whale kinematic behavioral state was related to the vocalization rate (also known as cue rate) and time of day. When cue rate was higher, fin whales were more likely to swim slower and turn more than when cue rate was lower. During the night, fin whales were also more likely to swim slower and turn more than during the day. In addition, we examined whether the presence of singing fin whales was related to time and sea state using generalized additive models. Fin whale track presence was affected by day of the year and song season, and possibly also wind speed and wave height. Although the track kinematics from the fin whale tracks presented here are limited to a subset of whales that are acoustically active, they provide some of the only detailed movements of fin whales in the region and can be compared against fin whale swim speeds in other regions. Understanding how fin whale swimming behavior varies based on their vocalization patterns, time, and environmental factors will help us to contextualize potential changes in whale behavior during Navy training and testing on the range.

μ-Opioid Receptor Stimulation in the Nucleus Accumbens Increases Vocal-Social Interactions in Flocking European Starlings, Sturnus Vulgaris.

Social connections in gregarious species are vital for safety and survival. For these reasons, many bird species form large flocks outside the breeding season. It has been proposed that such large social groups may be maintained via reward induced by positive interactions with conspecifics and via the reduction of a negative affective state caused by social separation. Moreover, within a flock optimal social spacing between conspecifics is important, indicating that individuals may optimize spacing to be close but not too close to conspecifics. The μ-opioid receptors (MORs) in the nucleus accumbens (NAc) are well known for their role in both reward and the reduction of negative affective states, suggesting that MOR stimulation in NAc may play a critical role in flock cohesion. To begin to test this hypothesis, social and nonsocial behaviors were examined in male and female European starlings (Sturnus vulgaris) in nonbreeding flocks after intra-NAc infusion of saline and three doses of the selective MOR agonist d-Ala2-N-Me-Phe4-Glycol5-enkephalin (DAMGO). DAMGO in NAc dose-dependently increased singing behavior and facilitated social approaches while at the same time promoting displacements potentially used to maintain social spacing. These findings support the hypothesis that MORs in NAc promote social interactions important for group cohesion in nonsexual contexts and suggest the possibility that MORs in the NAc play a role in optimizing the pull of joining a flock with the push of potential agonistic encounters.