Turnover of territorial Scops Owls Otus scops as estimated by spectrographic analyses of male hoots

In this study we assessed territory turnover of the European Scops Owl Otus scops, a migratory nocturnal raptor, using sonagrams of hoots recorded in the same territories in the 1993 and 1994 breeding seasons. Three structural parameters of the hoot (duration, inter-note interval and frequency) varied considerably, and a Discriminant Function Analysis classified >90% of individual calls correctly. In order to investigate yearly turnover and site-fidelity of male Scops Owls, the discriminant functions derived from the hoot sample recorded in 1993 were used to classify hoots of individuals recorded in the same territories in 1994. Apparently, turnover rate of territory owners was high and rapid, because 55–78% of territories censused in 1993 were defended by a different male in 1994. The fast decline of our Scops Owl population, probably due to winter mortality, could be responsible for such a high turnover of territorial owners.     

Vicariance and dispersal in the differentiation of vocalization in the Ryukyu Scops Owl Otus elegans

The distribution of species and species diversity can be affected by vicariance or dispersal. To understand their role in shaping species distribution and population structure these two processes must be estimated within and among populations. I analysed large‐scale variation in the call structure of the Ryukyu Scops Owl Otus elegans. This owl is distributed over a 1200‐km range, and only inhabits islands. Within this range, I studied this species across 22 continental islands of the Ryukyu Archipelago and two oceanic islands. The study aimed to assess whether there is variation in the acoustic structure of Owl hoot calls within islands, among major groups of islands and across a large area comprising a major biogeographical barrier (the Kerama Gap). The acoustic structure of calls was homogeneous within islands and among major island‐groups. Acoustic differentiation, however, increased over longer geographical distances of up to about 1200 km. The acoustic structure of hoots of the Ryukyu Scops Owl populations was clearly divided into two groups, north and south of the Kerama Gap. It is suggested that the Kerama Gap acted as a biogeographical barrier and contributed to the differentiation between the two major island‐groups. It is likely that this difference developed during the fragmentation of a widespread ancestral population by vicariant isolating events. There was also evidence of an effect of dispersal on vocal differentiation in subspecies inhabiting the two oceanic islands.     

Communication of male quality in owl hoots

The evolution of communication through intrasexual selection is expected to lead signalers to transmit honest information on their fighting ability. Here we studied the information encoded in the acoustic structure of the territorial calls of a nocturnal raptor. During territorial contests, male scops owls give hoots composed of a downward frequency shift followed by a stable plateau. We found that the frequency of the hoot was negatively correlated with the body weight of the vocalizer. We shifted the frequency contour of natural hoots in order to create resynthesized calls corresponding to individuals of varying body weight and used these stimuli in playback experiments simulating an intrusion into the territory of established breeders. Territory owners responded less intensely when they heard hoots simulating heavier intruders, and males with heavier apparent weight tended to give hoots with a lower plateau in response to playbacks simulating heavier intruders.     

Phrase structure of wild chimpanzee pant hoots: Patterns of production and interpopulation variability

Recordings and behavioral observations of wild chimpanzees were made over a 2-year period in the Kibale National Park, Uganda (Kanyawara and Ngogo communities) to investigate patterns of acoustic variability in long-distance calls. The phrase structures of pant hoots, the species-typical loud call given predominantly by adult males, were analyzed. Analysis revealed that the build-up phrase was frequently absent from the pant hoots of Kibale chimpanzees. By contrast, analysis of published data on Gombe and Mahale chimpanzees (Tanzania) indicated that these animals typically included the build-up in their calls. These results were interpreted as evidence for phrase-level differences between populations in the acoustic morphology of this compound call. Data on age and sex differences in the context of production of pant hoots were analyzed, and their relevance to the possibility that aspects of pant hoot acoustic morphology are learned is discussed. Adult males initiated pant hoots more than subadult males, and adult males joined other pant hooting individuals with pant hoots more than subadult males did. It is suggested that younger males may pant hoot with specific adult males preferentially and that this may affect the development of their pant hoot acoustic morphologies. A peculiar pant hoot variant previously reported from Gombe, the whimper hoot, is described from Kibale. The production of this call by low-ranking individuals suggests that there are social constraints on pant hooting in the chimpanzee community. Ideas concerning the effect of social relationships on interpopulation variability in vocal signals are briefly discussed. © 1996 Wiley-Liss, Inc.     

Conservation of Scops Owl Otus scops in the Alps: relationships with grassland management, predation risk and wider biodiversity

The Scops Owl Otus scops is the most endangered and least studied owl in Europe, with widespread declines reported throughout Europe. The species is qualitatively associated with two threats: changes in agricultural practices and predation by Tawny Owls Strix aluco . We investigated these two threats to a population in the Alps, where land abandonment is causing widespread woodland expansion with unknown consequences. Predation risk and environmental indicators of the degree of agricultural change both predicted Scops Owl distribution. Furthermore, agricultural change also affected broader biodiversity, as estimated by the richness and diversity of bird and diurnal butterfly species. This resulted in a strong association between Scops Owl presence and wider biodiversity, which held at different spatial scales and justified a conservation focus on a single, threatened species. The persistence of Scops Owls in the Alps could be promoted through subsidies halting land abandonment and promoting extensive grassland management practices. Such interventions would have the additional advantage of yielding broader biodiversity benefits.     

Wild Chimpanzees Produce Group‐Specific Calls: a Case for Vocal Learning?

Vocal learning, where animals can modify the structure of their vocalizations as a result of experience, has been found in a range of birds and mammals. Although vocal learning is a fundamental aspect of developing spoken language, there is as yet little evidence that vocal learning occurs in primates. Here we examine whether vocal learning may occur in chimpanzees. We analysed whether wild male chimpanzees, Pan troglodytes verus, of four communities living in a similar habitat in the Taï Forest, Côte d'Ivoire, developed community specific pant hoots. If so, we expected males of three contiguous communities to have distinct pant hoots, while pant hoots of males from a fourth, distant community, located 70 km away, should only differ from those of the contiguous communities by chance. Our analysis confirmed these expectations. In addition, the acoustic distances between the pant hoots of pairs of individuals did not correlate with the genetic relatedness of those pairs, where genetic relatedness was determined using nuclear DNA analysis. Thus, neither habitat nor genetic differences accounted for the observation that there were acoustic differences in the pant hoot structure of males living in neighbouring communities, but not in those of males from a distant community. This suggests that chimpanzees may actively modify pant hoots to be different from their neighbours, providing support for the vocal learning hypothesis.     

Male collared lizards,Crotaphytus collaris (Sauria: Crotaphytidae), signal females by broadcasting visual displays

The signalling function of displays broadcast when animals are distant from conspecifics can be difficult to determine. I tested the extent to which visually transmitted broadcast displays given by free‐ranging territorial male collared lizards signalled same‐sex rivals or females. One test involved recording the frequency of broadcast displays, aggressive contests with rivals, and courtship encounters with females during ten reproductive seasons when local sex ratios varied markedly. The frequency of broadcast displays decreased as the ratio of male competitors to females increased. The frequency with which males initiated contests with rivals was not related to the ratio of competitors to females, whereas the frequency of courtship interactions decreased with sex ratio because there were fewer females to court. The behaviour of males that defended territories during two successive seasons showed a similar pattern. Broadcast display frequency was positively correlated with courtship frequency, but not with the frequency of contests with rivals. Lastly, individual males gave more broadcast displays during focal observations when they also engaged in courtship encounters with females than other observations when they engaged in aggressive conflicts with rival males. Although these results do not reject the possibility that broadcast displays may also signal male rivals, they support a major role of these displays in advertisement to females. © 2013 The Linnean Society of London     

Ready to Fight: Reliable Predictors of Attack in a Cooperatively Breeding,Non‐Passerine Bird

Signal reliability is a major focus of animal communication research. Aggressive signals are ideal for measuring signal reliability because the signal referent – attack or no attack – can be measured unambiguously. Signals of aggressive intent occur at elevated rates in aggressive contexts, predict subsequent aggression by the signaler, and elicit appropriate responses from receivers. We tested the ‘predictive criterion’ in smooth‐billed anis, Crotophaga ani, by broadcasting one of two playback types (‘ahnee’ calls only or ‘ahnee + hoot’ calls), presenting a taxidermic mount, and observing the animals’ behavior. Based on the hypotheses that ‘hoot’ calls and ‘throat‐inflation’ displays signal aggressive intent, we predicted that they would be associated with attack, and that signaling rate would increase over the time period leading up to an attack. Indeed, both hoots and throat‐inflation displays reliably predicted attack. The second prediction, that signaling rate increases in the time leading up to attack, was strongly supported for throat‐inflation displays, which increased over the pre‐attack period in both treatments. Hoots increased over the pre‐attack period in ahnee playbacks but not in ahnee + hoot playbacks. Hierarchical signaling systems are characterized by early, less‐reliable predictors of attack, and later, more reliable predictors of attack. During both natural and simulated interactions, the more‐reliable throat‐inflation display tended to precede the less‐reliable hoot call, suggesting that this signaling system is not hierarchical. In a comparison of 11 putative signals of aggressive intent in birds, the throat‐inflation display had the second highest mutual information (reduction in uncertainty) among visual signals and non‐passerine signals while hoots had below‐average mutual information. Natural observations indicate that both hoots and throat‐inflation displays occur in the context of aggressive between‐group encounters, and hoots also occur during within‐group interactions. Throat‐inflation displays appear to be reliable indicators of aggressive intent, but the function of hoot calls is less clear.     

Importance of Frequency and Temporal Song Matching in Black-Capped Chickadees: Evidence from Interactive Playback

Song-type matching during territorial contests may allow males of some bird species to direct their signal to a particular receiver. By matching the song-type and also responding immediately to a rival (temporal matching), a signalling male may indicate his willingness to escalate the contest. Black-capped chickadees ( Poecile atricapilla ) sing a single song-type, but are able to sing this song over a wide range of absolute frequencies. By using interactive playback to instigate and control the level of matching during trials, we investigated whether matching the frequency and the temporal patterning of song escalates contests. Males that were matched for both the frequency and the temporal pattern of their songs during trials escalated contests more than males that were not matched, while males that were only matched temporally had an intermediate response. During trials that consisted of temporal matching only, focal males often shifted frequency to match the playback. Our results confirm that frequency matching and temporal matching using a single song-type allows graded signalling during aggressive interactions in chickadees.     

Overlapping and matching in the song contests of black-capped chickadees

During vocal contests, animals alter both the timing and the patterning of responses to their opponents. Time-specific responses (such as overlapping an opponent's song) and pattern-specific responses (such as matching the type of an opponent's song) may reveal information about interacting animals. Here we explore the consequences of overlapping and frequency matching during song contests of male black-capped chickadees, Poecile atricapillus. Using interactive playback, we engaged birds of high and low dominance status in vocal interactions with a simulated territorial intruder. The playback intruder either overlapped or avoided overlapping the subjects' songs and either matched or avoided matching the frequency of the subjects' songs. Individuals who were overlapped by the playback intruder showed higher variability in their song length and song timing than individuals who were not overlapped. Individuals who were frequency-matched by the playback intruder responded with more agitated responses (more flights and passes over the speaker and closer distances of closest approach) and spent more time farther away from the loudspeaker. We argue that the timing of song delivery and the choice of song type are distinct functional components of vocal interactions, where overlapping and matching songs are threatening signals that have separate consequences for opponent behaviour and song performance. High-ranking males responded at greater distances from the loudspeaker in all treatments and responded with lower agitation levels than low-ranking males. We demonstrate that males of different quality show different behavioural responses to territorial intruders, where males of high status appear reluctant to engage an intruder as intensely as males of low status.     

Variation in the male territorial hoot of the Tawny Owl Strix aluco in three English populations

Little is known about owl song. We made sonagrams of the territorial calls of 50 male Tawny Owls Strix aluco from three different areas. Six temporal and four frequency measures of the calls were recorded from the sonagrams. The measures of the calls were then subjected to analysis to try to separate between individual owls and between owls from different areas. We also looked for similarities between calls of neighbouring owls and for any effect of habitat on owl hoots. Individual owls were separated on the basis of their hoots with a high degree of success (98.6% overall), and there were significant differences between areas. Differences were found between calls in woodland and farmland habitats, but these differences were not in the direction expected to increase sound transmission. Calls of neighbouring owls did not resemble each other more than calls from owls that were not in vocal contact, implying that if calls are learned by Tawny Owls, they are learned before dispersal.     

RECENT BIOACOUSTIC PUBLICATIONS—from 1991 and a few from 1990 and 1989

ABSTRACT

The focus of this study was to determine whether individual vocal identification of Scops Owls Otus scops was possible and if there was a stability of the hoot-calls over a short time period in the same individuals. Spontaneous vocalizations of 13 owls were recorded in 2004 in Southern Tuscany, Italy. Visual analysis of spectrograms and quantitative multivariate analysis of six vocal features showed marked individual differences. In some owls a repertoire of two different hoot types was found. In 2005, 10 Scops owls were recorded three times in the same breeding season (2 hours and 10 days after the first session). Statistical analysis of data showed that 60% of owls did not change call features over time. However a slight but significant variability between successive vocal performances of the same owl was found in 40% of cases. This variability may decrease the recognition power by acoustic analysis. To overcome this obstacle I suggest a multi step qualitative/quantitative approach. A Difference Index (DI) was calculated to set a threshold between the slight intra-individual and the very high inter-individual variability. This method allowed the recognition of calls of each owl recorded over time in 2005.     

Food and habitats requirements of the Eurasian Scops Owl (Otus scops) in Switzerland revealed by very high-resolution multi-scale models

In Europe, agricultural practices have progressively evolved towards high productivity leading either to the intensification of productive and accessible areas or to the abandonment of less profitable sites. Both processes have led to the degradation of semi-natural habitats like extensive grasslands, threatening species such as the Eurasian Scops Owl Otus scops that rely on extensively managed agricultural landscapes. In this work, we aimed to assess the habitat preferences of the Scops Owl using habitat suitability models combined with a multi-scale approach. We generated a set of multi-scale predictors, considering both biotic and abiotic variables, built on two newly developed vegetation management and orthopteran abundance models. To select the variables to incorporate in a ‘best multi-scale model’, we chose the best spatial scale for each variable using univariate models and by calculating their relative importance through multi-model inference. Next, we built ensembles of small models (ESMs) at 10 different scales from 50 to 1000 m, and an additional model with each variable at its best scale (‘best multi-scale model’). The latter performed better than most of the other ESMs and allowed the creation of a high-resolution habitat suitability map for the species. Scops Owls showed a preference for dry sites with extensive and well-structured habitats with 30–40% bush cover, and relied strongly on semi-extensive grasslands covering at least 30% of the surface within 300 m of the territory centre and with high orthopteran availability near the centre (50-m radius), revealing a need for good foraging grounds near the nest. At a larger spatial scale within a radius of 1000 m, the habitat suitability of Scops Owls was negatively related to forest cover. The resulting ESM predictions provide valuable tools for conservation planning, highlighting sites in need of particular conservation efforts together with offering estimates of the percentage of habitat types and necessary prey abundance that could be used as targets in future management plans to ensure the persistence of the population.     

Ultraviolet signals fighting ability in a lizard

Ultraviolet (UV) signals are used in female mate choice in numerous taxa; however, the role of UV signals in male contests remains relatively unexplored. We experimentally reduced throat UV of free-ranging lizards (Platysaurus broadleyi) to test whether UV acts as a signal of fighting ability during male contests. We found that UV-reduced males were more likely to be challenged than control males. However, contest outcome was not influenced by UV-reduction, and this was despite other obvious asymmetries between opponents, such as body size and residency. Throat UV was confirmed as a signal of fighting ability because contests were more likely to escalate when one contestant had reduced UV. Therefore, throat UV, not body size or residency, was used during the initial stage of opponent assessment, but this did not influence contest outcome. The results suggest that UV overrides other traits that could function as signals during rival assessment.     

Factors affecting spontaneous vocal activity of Tawny Owls Strix aluco and implications for surveying large areas

To use vocalizations properly for the estimation of owl population size, it is important to identify how environmental factors affect owl calling behaviour. Here, we analyse how intrinsic and extrinsic factors modify the vocal activity of Tawny Owls Strix aluco in two areas of northern Spain. From March 2013 to May 2015, we radiotracked 20 Tawny Owls and also undertook a systematic survey in which we collected data on spontaneous vocal activity (hoot/call) of the tagged owls, plus their mates and neighbours (36 untagged owls). After 223 nights in Valle de Mena and 224 in Duranguesado we obtained a total of 8791 records of vocal activity. The annual proportion of surveys on which an owl called was 6.3% and did not differ between the study areas. Vocal activity of Tawny Owls varied with sex, annual cycle stage and weather. Male owls were significantly more vocal than females year‐round, and vocal activity peaked during the incubation and post‐breeding periods. Wind and rain adversely affected vocal activity of both sexes throughout the year. Tagged owls were detected more often than untagged owls, which we interpret as an observer effect because the systematic survey ensured that short distances to tagged owls increased the probability of detecting vocal activity. In fact, 2.8% of variation in vocal activity was due to detectability differences between tagged and untagged owls. We conclude that if fieldwork is carried out during the optimum period of the year for vocal detection (i.e. incubation, which in our case was around mid‐April), and under good weather conditions (dry and calm nights), censuses based on spontaneous vocal activity would detect only approximately 12% of the true Tawny Owl population.     

Acoustic analysis of wild chimpanzee pant hoots: Do Kibale Forest chimpanzees have an acoustically distinct food arrival pant hoot?

Recordings were made of 193 pant hoots given by 6 adult males during ad lib sampling over a 16-month period in 1988 and 1989. The presence or absence of a let-down phase, and acoustic measures of the let-down and climax phases of the calls, were compared for different call contexts to determine if an acoustically distinct pant hoot was given uniquely upon arrival at fruiting trees. The greatest proportion of pant hoots with a let-down (LD pant hoots) occurred immediately upon arrival at fruiting trees. However, LD pant hoots also occurred at other times during feeding bouts. The frequency and duration of the first exhaled element of 19 letdown phases were measured, and four measures were made on the highest pitched element of 49 climax phases: duration, maximum and minimum frequencies, and average frequency. No differences were found in these acoustic measures that distinguished calls given immediately upon arrival at a food tree from calls given later during feeding bouts. Thus no evidence was found that an acoustically distinct pant hoot was given uniquely upon arrival at fruiting trees. However, the analyses did suggest that identifiable pant hoot variants are given in different social contexts. The proportion of LD pant hoots decreased in more interactive social contexts, and other acoustic features may be available to distinguish this pant hoot variant at long distances. We suggest that different pant hoot variants might broadcast information specific to social, as opposed to ecological, context. © 1993 Wiley-Liss, Inc.     

Vocal rate as an assessment process during fallow deer contests

Two types of model propose that strategic decisions during contests are determined either by (i) a mutual-assessment process or (ii) a self-assessment process. Vocal signals are thought to convey information about the competitive abilities of individuals, the ultimate function of which is a reduction in costs associated with fighting consistent with the principle of mutual assessment. Nevertheless, the limited evidence that male ungulates engage in mutual assessment of vocal rates during dyadic contests has been questioned. Therefore, we examined the vocal rates of winners and losers during escalated dyadic contests between male fallow deer in order to further inform on this issue. Our results showed that winners and losers did not differ in vocal rate. The best model fit that accounted for individual vocal rates included a preponderance of factors related to the opponent indicating that contestants were attending to their opponent during fights. Vocal rate was, therefore, dependent on estimates of opponent quality without reference to self, supporting an 'opponent-only' rather than a mutual assessment process.     

Information transfer during cricket frog contests

We investigated information transfer during vocal interactions between cricket frogs, Acris crepitans, with a specific focus on information about size and intention. In response to opponents, cricket frogs alter both temporal and spectral (frequency) aspects of their calls. Previous work suggests that males use dominant frequency, which is correlated with size, to provide information about fighting ability, and use temporal call characters, which are independent of size, to provide information about aggressive intent. We tested this hypothesis by investigating the relationship between call characters and contest behaviour. We presented a focal male with a simulated opponent, and categorized his behavioural response as attack, abandon or tolerate. We found that information about opponent size does not appear to influence a male's decision to fight, flee or tolerate an intruder. Whether or not males use the size information that is encoded in call frequency remains unclear. In contrast, changes in call frequency, which are not correlated with size, predicted the outcome of simulated contests, suggesting that male cricket frogs signal information about agonistic intent. Temporal call characters indicated whether or not a resident tolerated an opponent, but they did not predict contest outcome (attack versus abandon). Furthermore, the difference between the temporal call characters of a focal male and the simulated opponent predicted whether the resident tolerated the opponent. We suggest, therefore, that temporal call changes may be a cooperative signal designed to facilitate assessment of size through physical contact in wrestling bouts. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Selection for acoustic individuality within the vocal repertoire of wild chimpanzees

Individual primates typically produce acoustically distinct calls. To investigate the factors that facilitate the evolution of individual vocal signatures, we examined two components of the call repertoire of chimpanzees: the pant hoot and pant grunt. Pant hoots are long-distance signals whose recipients can be several hundred meters away, while pant grunts are short-range calls given to conspecifics within close visual range. Given their markedly different contexts of emission, we predicted that natural selection would favor the elaboration of individually distinctive acoustic features in pant hoots compared with pant grunts. Analyses of nine acoustic features revealed that pant hoots are more stereotyped within-individuals and variable between-individuals than pant grunts. These data are consistent with the hypothesis that selection may act to encode varying degrees of individuality in different components of the vocal repertoire of a single species.     

Effect of Willow Tit Poecile montanus alarm calls on attack rates by Pygmy Owls Glaucidium passerinum

One suggested anti‐predator function of alarm calls is to deliver a message to a predator that it has been detected. Moreover, giving the alarm call could provide a signal to the predator that capturing the individual giving the alarm is more difficult than capturing its silent group members, as the caller is probably the most aware of the predator's location. In an aviary experiment using stuffed dummy Willow Tits Poecile montanus, we assessed whether an authentic alarm call given by Willow Tit affected Pygmy Owl Glaucidium passerinum prey preference. In the experiment, the Owls attacked only the ‘silent’ dummy individuals, suggesting that alarm calling could offer direct fitness benefits to the caller by decreasing the attack risk of the caller relative to its group members.