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'.

A Relationship between the Characteristics of the Oval Nucleus of the Mesopallium and Parrot Vocal Response to Playback.

Correlations between differences in animal behavior and brain structures have been used to infer function of those structures. Brain region size has especially been suggested to be important for an animal's behavioral capability, controlled by specific brain regions. The oval nucleus of the mesopallium (MO) is part of the anterior forebrain vocal learning pathway in the parrot brain. Here, we compare brain volume and total number of neurons in MO of three parrot species (the peach-fronted conure, Eupsittula aurea, the peach-faced lovebird, Agapornis roseicollis, and the budgerigar, Melopsittacus undulatus), relating the total neuron numbers with the vocal response to playbacks of each species. We find that individuals with the highest number of neurons in MO had the shortest vocal latency. The peach-fronted conures showed the shortest vocal latency and largest number of MO neurons, the peach-faced lovebird had intermediary levels of both, and the budgerigar had the longest latency and least number of neurons. These findings indicate the MO nucleus as one candidate region that may be part of what controls the vocal capacity of parrots.

Follow the leader? Orange-fronted conures eavesdrop on conspecific vocal performance and utilise it in social decisions.

Animals regularly use social information to make fitness-relevant decisions. Particularly in social interactions, social information can reduce uncertainty about the relative quality of conspecifics, thus optimising decisions on with whom and how to interact. One important resource for individuals living in social environments is the production of information by signalling conspecifics. Recent research has suggested that some species of parrots engage in affiliative contact call matching and that these interactions may be available to conspecific unintended receivers. However, it remains unclear what information third parties may gain from contact call matching and how it can be utilised during flock decisions. Here, using a combined choice and playback experiment, we investigated the flock fusion choices and vocal behaviour of a social parrot species, the orange-fronted conure (Eupsittula canicularis), to a contact call matching interaction between two individuals of different sexes and with different vocal roles. Our results revealed that orange-fronted conures chose to follow vocal leaders more often than vocal followers during fusions. Furthermore, flocks responded with higher call rates and matched the stimulus calls closer when subsequently choosing a vocal leader. Interestingly, orange-fronted conures also showed higher contact call rates and closer matches when choosing males over females. These results suggest that paying attention to conspecific contact call interactions can provide individuals with social information that can be utilised during fission and fusion events, significantly influencing the social dynamics of orange-fronted conures.

Contingency and determinism in the evolution of bird song sound frequency.

Sexual signals are archetypes of contingent evolution: hyper-diverse across species, often evolving fast and in unpredictable directions. It is unclear to which extent their evolutionary unpredictability weakens deterministic evolution, or takes place bounded by deterministic patterns of trait evolution. We compared the evolution of sound frequency in sexual signals (advertisement songs) and non-sexual social signals (calls) across > 500 genera of the crown songbird families. Contrary to the acoustic adaptation hypothesis, we found no evidence that forest species used lower sound frequencies in songs or calls. Consistent with contingent evolution in song, we found lower phylogenetic signal for the sound frequency of songs than calls, which suggests faster and less predictable evolution, and found unpredictable direction of evolution in lineages with longer songs, which presumably experience stronger sexual selection on song. Nonetheless, the most important deterministic pattern of sound frequency evolution-its negative association with body size-was stronger in songs than calls. This can be explained by songs being longer-range signals than most calls, and thus using sound frequencies that animals of a given size produce best at high amplitude. Results indicate that sexual selection can increase aspects of evolutionary contingency while strengthening, rather than weakening, deterministic patterns of evolution.

An exploratory investigation of glucocorticoids, personality and survival rates in wild and rehabilitated hedgehogs (Erinaceus europaeus) in Denmark.

BACKGROUND: The European population of hedgehogs (Erinaceus europaeus) is declining. It is therefore essential to optimise conservation initiatives such as the rehabilitation of sick, injured and orphaned hedgehogs. Wild animals placed in captivity may be prone to chronic stress, potentially causing negative health effects. Therefore, the effects of these rehabilitation efforts should consequently be evaluated. Furthermore, hand-raising orphaned hedgehogs is a laborious and costly task, and it is therefore relevant to document whether they have equal post release survival rates compared to their wild conspecifics. The objectives of this research were therefore to conduct an exploratory study of glucocorticoid levels in hedgehogs from different backgrounds and compare the post release survival of translocated, rehabilitated and wild, juvenile hedgehogs as well as the possible effect on survival of differences in shy or bold behaviour (personality) exhibited by individuals. RESULTS: We measured glucocorticoid levels in 43 wild-caught (n = 18) and rehabilitated (n = 25) hedgehogs and compared the post release survival and spatial behaviour of 18 translocated juvenile hedgehogs (eight hand-raised and ten wild) until hibernation. The possible effect on survival of differences in shy or bold behaviour (personality) exhibited by 17 juvenile individuals (seven hand-raised and ten wild) was also examined. Rehabilitated individuals and females had higher levels of faecal corticosterone metabolites compared to wild individuals and males, respectively. Rehabilitated individuals showed higher levels of saliva corticosterone than wild. The personality tests labelled 13 individuals as shy and 11 as bold. Post release survival was 57% for rehabilitated and 50% for wild individuals. Neither background nor personality affected post release survival. Home range measures were 3.54 and 4.85 ha. Mean dispersal length from the release sites was 217 ± 100 m. CONCLUSION: The higher levels of corticosterone observed in rehabilitated compared to wild hedgehogs calls for consideration of the duration of admission to wildlife rehabilitation centres to reduce stress levels in the patients. Hand-raised juveniles appear to have the same prospects as wild, and personality does not seem to affect post release survival in hedgehogs, indicating that hand-raising of orphaned juvenile hedgehogs is a relevant contribution to the conservation of this species.

The Allometry of Sound Frequency Bandwidth in Songbirds.

AbstractTheory predicts that allometric constraints on sound production should be stronger for the lower frequencies of vocalizations than for the higher frequencies, which could originate from an allometry for sound frequency bandwidth. Using song recordings of approximately 1,000 passerine species (from >75% passerine genera), we show a significantly steeper allometry for the lower song frequencies than for the higher song frequencies, resulting in a positive allometry of frequency bandwidth: larger species can use wider bandwidths than smaller species. The bandwidth allometry exists in songbirds (oscines) but not in nonoscine passerines, indicating that it emerges from a combination of constraints to sound frequency production or transmission and the evolved behavior of oscines: unlike the narrow bandwidths of most nonoscine songs, the learned songs of oscines often use wide bandwidths that can be limited by both lower and upper constraints to sound frequency. This bandwidth allometry has implications for several research topics in acoustic communication.

Habitat suitability analysis reveals high ecological flexibility in a "strict" forest primate.

BACKGROUND: Research of many mammal species tends to focus on single habitats, reducing knowledge of ecological flexibility. The Javan lutung (Trachypithecus auratus) is considered a strict forest primate, and little is known about populations living in savannah. In 2017-2018, we investigated the density and distribution of Javan lutung in Baluran National Park, Indonesia. We conducted ad libitum follows and line transect distance sampling with habitat suitability analysis of Javan lutung. RESULTS: Estimated density was 14.91 individuals km- 2 (95% CI 7.91-28.08), and estimated population size was 3727 individuals (95% CI 1979 - 7019). Long-tailed macaque (Macaca fascicularis) habitat suitability was the main driver of lutung habitat suitability as the probability of lutung occurrence increased greatly with macaque habitat suitability. Distance to roads, and distance to secondary forest had a negative relationship with lutung occurrence. Lutung habitat suitability decreased with increasing elevation, however, Mt Baluran and the primary forest on Mt Baluran was under-sampled due to treacherous conditions. Follows of six focus groups revealed considerable use of savannah, with terrestrial travel. The follows also revealed polyspecific associations with long-tailed macaques through shared sleeping sites and inter-specific vocalisations. CONCLUSIONS: Our study provides new knowledge on the general ecology of Javan lutung, such as use of savannah habitats, underlining our need to branch out in our study sites to understand the flexibility and adaptability of our study species. Another undocumented behaviour is the polyspecific association with long-tailed macaques. We encourage more research on this subject.

The ecology of suburban juvenile European hedgehogs (Erinaceus europaeus) in Denmark.

European hedgehog (Erinaceus europaeus) populations are widespread across diverse habitats but are declining in Western Europe. Drastic declines have been described in the UK, with the most severe declines occurring in rural areas. Hedgehogs are widely distributed in Denmark, but their status remains unknown.Fieldwork on hedgehogs has tended to focus on rural areas, leaving their ecology in suburban habitats largely unexplored, with clear implications for conservation initiatives. Here, we study the ecology of 35 juvenile hedgehogs using radio tracking during their first year of life in the suburbs of western Copenhagen.We use radio-tracking data to estimate (a) home range sizes in autumn and spring/summer, (b) survival during their first year of life, (c) the body mass changes before, during, and after hibernation, and (d) the hibernation behavior of the juvenile hedgehogs.We show that males and females have small home ranges compared with previous studies. The 95% MCP home range sizes in autumn were 1.33 ha (95% CI = 0.88-2.00) for males and 1.40 ha (95% CI = 0.84-2.32) for females; for spring/summer they were 6.54 ha (95% CI = 3.76-11.38) for males and 1.51 ha (95% CI = 0.63-3.63) for females. The juvenile survival probabilities during the study period from September 2014 to July 2015 were .56 for females and .79 for males. All healthy individuals gained body mass during the autumn and survived hibernation with little body mass loss thus demonstrating that the juveniles in the study were capable of gaining sufficient weight in the wild to survive their first hibernation.The climate is changing, but there is a lack of knowledge on how this affects mammal ecology. The exceptionally mild autumn of 2014 caused the juvenile hedgehogs to delay hibernation for up to a month compared with previous studies in Denmark.

Evolution of non-kin cooperation: social assortment by cooperative phenotype in guppies.

Cooperation among non-kin constitutes a conundrum for evolutionary biology. Theory suggests that non-kin cooperation can evolve if individuals differ consistently in their cooperative phenotypes and assort socially by these, such that cooperative individuals interact predominantly with one another. However, our knowledge of the role of cooperative phenotypes in the social structuring of real-world animal populations is minimal. In this study, we investigated cooperative phenotypes and their link to social structure in wild Trinidadian guppies (Poecilia reticulata). We first investigated whether wild guppies are repeatable in their individual levels of cooperativeness (i.e. have cooperative phenotypes) and found evidence for this in seven out of eight populations, a result which was mostly driven by females. We then examined the social network structure of one of these populations where the expected fitness impact of cooperative contexts is relatively high, and found assortment by cooperativeness, but not by genetic relatedness. By contrast, and in accordance with our expectations, we did not find assortment by cooperativeness in a population where the expected fitness impact of cooperative contexts is lower. Our results provide empirical support for current theory and suggest that assortment by cooperativeness is important for the evolution and persistence of non-kin cooperation in real-world populations.

The number of neurons in specific amygdala regions is associated with boldness in mink: a study in animal personality.

Conspecifics vary consistently in their behavioural responses towards environment stimuli such as exposure to novel objects; ethologists often refer to this variability as animal personality. The neurological mechanisms underlying animal personality traits remain largely unknown, but linking the individual variation in emotional expression to brain structural and neurochemical factors is attracting renewed interest. While considerable research has focused on hormonal and neurotransmitter effects on behavioural responses, less is known about how individual variation in the number of specific neuron populations contributes to individual variation in behaviour. The basolateral amygdala (BLA) and the central nuclei of the amygdala (CeA) mediate emotional processing by regulating behavioural responses of animals in a potentially threatening situation. As such, these structures are good candidates for evaluating the relationship between neuronal populations and behavioural traits. We now show that individual American mink (Neovison vison) reacting more boldly towards novelty have more neurons in the BLA than do their more timid conspecifics, suggesting that a developmental pattern of the number of amygdala neurons can influence behavioural traits of an adult animal. Furthermore, post hoc correlations revealed that individuals performing with higher arousal, as reflected by their frequency of startle behaviour, have more CeA neurons. Our results support a direct link between the number of neurons in amygdala regions and aspects of animal personality.

Mate choice screening in captive solitary carnivores: The role of male behavior and cues on mate preference and paternity in females of a model species, American mink (Neovison vison).

Mate choice studies suggest that choosy females benefit from increased fecundity, litter size, and offspring survival. Thus, providing females with the opportunity to choose among potential mates, deemed genetically suitable based on studbook data, might improve breeding management in production and zoo animals and thereby the sustainability of captive populations. Investigating mate preference via odor from potential mates before animal transfer is a proposed strategy for incorporating mate choice into breeding management. In this study, we test whether olfactory cues and signals from males can be used to assess and measure female mate preference in American mink. Eighteen females were subjected to a 4-day stimulus test in which females showed a preference for one of two males' urine and feces. Subsequently, each female was subjected to a 10-day mate preference test involving the same two males of the first test. Paternity tests revealed that 13 females had offspring, which could be assigned to only one male, suggesting that these females performed a mate choice. In nine of these females preference during the stimulus test was directed toward the male that fathered their offspring. Our results suggest that even though there was a preference difference in scent stimulus trials from potential mates this preference was not predictive of eventual mate preference or paternity. Other factors such as aspects of male behavior seem to play a role, when the mates are introduced. Our study supports that mate preference and mate choice are complex matters influenced by multiple cues and signals.

Context Matters: Multiple Novelty Tests Reveal Different Aspects of Shyness-Boldness in Farmed American Mink (Neovison vison).

Animal personality research is receiving increasing interest from related fields, such as evolutionary personality psychology. By merging the conceptual understanding of personality, the contributions to both fields of research may be enhanced. In this study, we investigate animal personality based on the definition of personality traits as underlying dispositional factors, which are not directly measurable, but which predispose individuals to react through different behavioural patterns. We investigated the shyness-boldness continuum reflected in the consistency of inter-individual variation in behavioural responses towards novelty in 47 farmed American mink (Neovison vison), which were raised in identical housing conditions. Different stages of approach behaviour towards novelty, and how these related within and across contexts, were explored. Our experimental design contained four tests: two novel object tests (non-social contexts) and two novel animated stimuli tests (social contexts). Our results showed consistency in shyness measures across multiple tests, indicating the existence of personality in farmed American mink. It was found that consistency in shyness measures differs across non-social and social contexts, as well as across the various stages in the approach towards novel objects, revealing that different aspects of shyness exist in the farmed American mink. To our knowledge this is the first study to reveal aspects of the shyness-boldness continuum in the American mink. Since the mink were raised in identical housing conditions, inherited factors may have been important in shaping the consistent inter-individual variation. Body weight and sex had no effect on the personality of the mink. Altogether, our results suggest that the shyness-boldness continuum cannot be explained by a simple underlying dispositional factor, but instead encompasses a broader term of hesitating behaviour that might comprise several different personality traits.

Core and Shell Song Systems Unique to the Parrot Brain.

The ability to imitate complex sounds is rare, and among birds has been found only in parrots, songbirds, and hummingbirds. Parrots exhibit the most advanced vocal mimicry among non-human animals. A few studies have noted differences in connectivity, brain position and shape in the vocal learning systems of parrots relative to songbirds and hummingbirds. However, only one parrot species, the budgerigar, has been examined and no differences in the presence of song system structures were found with other avian vocal learners. Motivated by questions of whether there are important differences in the vocal systems of parrots relative to other vocal learners, we used specialized constitutive gene expression, singing-driven gene expression, and neural connectivity tracing experiments to further characterize the song system of budgerigars and/or other parrots. We found that the parrot brain uniquely contains a song system within a song system. The parrot "core" song system is similar to the song systems of songbirds and hummingbirds, whereas the "shell" song system is unique to parrots. The core with only rudimentary shell regions were found in the New Zealand kea, representing one of the only living species at a basal divergence with all other parrots, implying that parrots evolved vocal learning systems at least 29 million years ago. Relative size differences in the core and shell regions occur among species, which we suggest could be related to species differences in vocal and cognitive abilities.

Evidence for varying social strategies across the day in chacma baboons.

Strong social bonds can make an important contribution to individual fitness, but we still have only a limited understanding of the temporal period relevant to the adjustment of social relationships. While there is growing recognition of the importance of strong bonds that persist for years, social relationships can also vary over weeks and months, suggesting that social strategies may be optimized over shorter timescales. Using biological market theory as a framework, we explore whether temporal variation in the benefits of social relationships might be sufficient to generate daily adjustments of social strategies in wild baboons. Data on grooming, one measure of social relationships, were collected from 60 chacma baboons (Papio ursinus) across two troops over a six month period. Our analyses suggest that social strategies can show diurnal variation, with subordinates preferentially grooming more dominant individuals earlier in the day compared with later in the day. These findings indicate that group-living animals may optimize certain elements of their social strategies over relatively short time periods.

Knowing your audience affects male-male interactions in Siamese fighting fish (Betta splendens).

Aggressive interactions between animals often occur in the presence of third parties. By observing aggressive signalling interactions, bystanders may eavesdrop and gain relevant information about conspecifics without the costs of interacting. On the other hand, interactants may also adjust their behaviour when an audience is present. This study aimed to test how knowledge about fighting ability of an audience affects aggressive interactions in male Siamese fighting fish. Subjects were positioned between two dyads of non-interacting males and allowed to observe both dyads shortly before the view to one of the dyads was blocked, and the dyads were allowed to interact. Subjects were subsequently exposed to an unknown opponent in the presence of either the winner or the loser of the seen or unseen interaction. The results suggest a complex role of the characteristic of an audience in the agonistic behaviours of a subject engaged in an interaction. The presence of a seen audience elicited more aggressive displays towards the opponent if the audience was a loser. This response was different in the presence of an unseen audience. Subjects then directed a higher aggressiveness against their opponent if the audience was a winner. These results also suggest a potentially more complex and interesting process allowing individuals to gain information about the quality and threat level of an unknown audience while it is interacting with a third party. The importance of information acquisition for an individual to adapt its behaviour and the role of communication networks in shaping social interactions are discussed.

Vocal imitation in parrots allows addressing of specific individuals in a dynamic communication network.

Parrots in captivity are known for their ability to vocally imitate humans and recently it has been shown that wild-living orange-fronted conures are able to immediately imitate other individuals' contact calls. The function of this exceptional ability to imitate remains unclear. However, orange-fronted conures live in fission-fusion flocks where they encounter many different individuals every day, and it is possible that their vocal imitation ability is a flexible means to address a specific individual within a flock. We tested this via playback to short-term captive wild conures. Test birds were placed together in pairs in outdoor aviaries to form simple flocks. To simulate imitation of a specific individual these pairs received playback of contact calls that primarily imitate one of the two birds. Overall, individuals that received simulated vocal imitations of its calls responded more frequently and faster than the other individual. This suggests that orange-fronted conures can use imitations of contact calls to address specific individuals of a flock. In the discussion we argue that the fission-fusion flock dynamics of many parrot species has been an important factor in evolving conures' and other parrots' exceptional ability to imitate.

Social preferences based on sexual attractiveness: a female strategy to reduce male sexual attention.

Male sexual harassment of females is common across sexually reproducing species and can result in fitness costs to females. We hypothesized that females can reduce unwanted male attention by constructing a social niche where their female associates are more sexually attractive than themselves, thus influencing the decision-making of males to their advantage. We tested this hypothesis in the Trinidadian guppy (Poecilia reticulata), a species with high levels of male sexual harassment. First, we confirmed that non-receptive females were harassed less when they were paired with a more sexually attractive (receptive) female than with another non-receptive female. We then found that, indeed, females exploit this as a strategy to reduce sexual harassment; non-receptive females actively preferred to associate with receptive over non-receptive females. Importantly, when given access only to chemosensory cues, non-receptive females still showed this preference, suggesting that they use information from chemical cues to assess the sexual attractiveness of potential female partners. Receptive females in contrast showed no such preferences. Our results demonstrate that females can decrease male harassment by associating with females that are more sexually attractive than themselves and that they perform active partner choices based on this relative attractiveness. We propose that this strategy is likely to represent an important pathway by which females can construct social niches that influence the decision-making of others to their advantage; in this case, to reduce the sexual harassment they experience.

Degradation of rural and urban great tit song: testing transmission efficiency.

Acoustic signals play a fundamental role in avian territory defence and mate attraction. Several studies have now shown that spectral properties of bird song differ between urban and rural environments. Previously this has been attributed to competition for acoustic space as a result of low-frequency noise present in cities. However, the physical structure of urban areas may have a contributory effect. Here we investigate the sound degradation properties of woodland and city environments using both urban and rural great tit song. We show that although urban surroundings caused significantly less degradation to both songs, the transmission efficiency of rural song compared to urban song was significantly lower in the city. While differences between the two songs in woodland were generally minimal, some measures of the transmission efficiency of rural song were significantly lower than those of urban song, suggesting additional benefits to singing rural songs in this setting. In an attempt to create artificial urban song, we mimicked the increase in minimum frequency found several times previously in urban song. However, this did not replicate the same transmission properties as true urban song, suggesting changes in other song characteristics, such as temporal adjustments, are needed to further increase transmission of an avian signal in the city. We suggest that the structure of the acoustic environment, in addition to the background noise, plays an important role in signal adaptation.

A neurological comparative study of the harp seal (Pagophilus groenlandicus) and harbor porpoise (Phocoena phocoena) brain.

The cetacean brain is well studied. However, few comparisons have been done with other marine mammals. In this study, we compared the harp seal (Pagophilus groenlandicus) and the harbor porpoise brain (Phocoena phocoena). Stereological methods were applied to compare three areas of interest: the entire neocortex and two subdivisions of the neocortex, the auditory and visual cortices. The total number of neurons and glial cells in the three regions was estimated. The main results showed that the harbor porpoise have an estimated 14.9 × 10(9) neocortical neurons and 34.8 × 10(9) neocortical glial cells, whereas the harp seal have 6.1 × 10(9) neocortical neurons and 17.5 × 10(9) neocortical glial cells. The harbor porpoise have significantly more neurons and glial cells in the auditory cortex than in the visual cortex, whereas the pattern was opposite for the harp seal. These results are the first to provide estimates of the number of neurons and glial cells in the neocortex of the harp seal and harbor porpoise brain and offer new data to the comparative field of mammalian brain evolution.

Knee-clicks and visual traits indicate fighting ability in eland antelopes: multiple messages and back-up signals.

BACKGROUND: Given the costs of signalling, why do males often advertise their fighting ability to rivals using several signals rather than just one? Multiple signalling theories have developed largely in studies of sexual signals, and less is known about their applicability to intra-sexual communication. We here investigate the evolutionary basis for the intricate agonistic signalling system in eland antelopes, paying particular attention to the evolutionary phenomenon of loud knee-clicking. RESULTS: A principal components analysis separated seven male traits into three groups. The dominant frequency of the knee-clicking sound honestly indicated body size, a main determinant of fighting ability. In contrast, the dewlap size increased with estimated age rather than body size, suggesting that, by magnifying the silhouette of older bulls disproportionately, the dewlap acts as an indicator of age-related traits such as fighting experience. Facemask darkness, frontal hairbrush size and body greyness aligned with a third underlying variable, presumed to be androgen-related aggression. A longitudinal study provided independent support of these findings. CONCLUSION: The results show that the multiple agonistic signals in eland reflect three separate components of fighting ability: (1) body size, (2) age and (3) presumably androgen-related aggression, which is reflected in three backup signals. The study highlights how complex agonistic signalling systems can evolve through the simultaneous action of several selective forces, each of which favours multiple signals. Specifically, loud knee-clicking is discovered to be an honest signal of body size, providing an exceptional example of the potential for non-vocal acoustic communication in mammals.