Formant Frequencies are Acoustic Cues to Caller Discrimination and are a Weak Indicator of the Body Size of Corncrake Males

Source–filter theory assumes that calls are generated by a vocal source and are subsequently filtered by the vocal tract. The air in the vocal tract vibrates preferentially at certain resonant frequencies, called formants. Formant frequencies can be a good indicator of the caller's characteristics, such as sex, age, body size or individual identity. Although source–filter theory was originally proposed for mammals, formants are also observed in birds, and some bird species have been shown to perceive formants. In this study, we evaluated the hypotheses that formant frequencies (1) are an indicator of body size and (2) can be used for individual discrimination by a nocturnal bird species, the corncrake (Crex crex). We analysed calls of 104 males from Poland and the Czech Republic. Linear regression models showed that the males with a longer head (including the bill length) had a significantly lower formant dispersion and lower fourth and fifth formant frequencies. However, we found no significant relationships between body weight and any filter‐related acoustic measurement. The formant frequencies had smaller within‐ than between‐individual coefficients of variation. This characteristic of the formant frequencies implies a high potential for individual coding. A discriminant function analysis correctly assigned 94.8% of the calls to the caller based on formants from second to fifth. Our results indicated that the formant frequencies are a weak indicator of the body size of the sender in the corncrake. However, even weak dependence between body size and acoustic properties of signal may be important in natural selection process. Alternatively, such a weak dependence may be observed, because receivers ignore the acoustical, formant‐based cues of body size. Simultaneously, the formants might potentially provide acoustic cues to individual discrimination and could be used to census and monitoring tasks.     

Acoustic cues to caller identity in lemurs: a case study

This study investigated the acoustic structure of grunt vocalizations in red-bellied lemurs (Eulemur rubriventer) and its potential for individual discrimination. Acoustic analyses were performed on 1,605 grunts recorded from seven lemurs belonging to two captive groups. From the perspective of sound-filter theory, we described the acoustic structure of grunts, measuring two sets of parameters: fundamental frequency characteristics as larynx-related variables and four formant frequencies as filter-related features. Formants were effective in assigning 80.5% of the vocalizations to the correct emitter against 24.9% scored by the model based on larynx-related variables. We concluded that vocal tract resonances might potentially provide conspecifics with individual cues.     

Can Nocturnal Flight Calls of the Migrating Songbird,American Redstart,Encode Sexual Dimorphism and Individual Identity?

Bird species often use flight calls to engage in social behavior, for instance maintain group cohesion and to signal individual identity, kin or social associations, or breeding status of the caller. Additional uses also exist, in particular among migrating songbirds for communication during nocturnal migration. However, our understanding of the information that these vocalizations convey is incomplete, especially in nocturnal scenarios. To examine whether information about signaler traits could be encoded in flight calls we quantified several acoustic characteristics from calls of a nocturnally migrating songbird, the American Redstart. We recorded calls from temporarily captured wild specimens during mist-netting at the Powdermill Avian Research Center in Rector, PA. We measured call similarity among and within individuals, genders, and age groups. Calls from the same individual were significantly more similar to one another than to the calls of other individuals, and calls were significantly more similar among individuals of the same sex than between sexes. Flight calls from hatching-year and after hatching-year individuals were not significantly different. Our results suggest that American Redstart flight calls may carry identifiers of gender and individual identity. To our knowledge, this is the first evidence of individuality or sexual dimorphism in the flight calls of a migratory songbird. Furthermore, our results suggest that flight calls may have more explicit functions beyond simple group contact and cohesion. Nocturnal migration may require coordination among numerous individuals, and the use of flight calls to transmit information among intra- and conspecifics could be advantageous. Applying approaches that account for such individual and gender information may enable more advanced research using acoustic monitoring.     

Otter scent signals age, sex, and reproductive status

Scent is used across taxa to communicate information about signaler identity. Eurasian otters Lutra lutra are mainly solitary and thought to use scent as their primary means of communication. Little is known, however, about what information otters communicate through scent or what social function this performs. Headspace solid-phase microextraction and gas chromatography-mass spectrometry were used to sample and analyze volatile organic compounds from anal scent gland secretion from 158 otters of differing sex, age, and female reproductive status. Univariate and multivariate differences were clear between adult and juvenile otters. Complex sex differences were apparent in adult otters but not in younger individuals, suggesting the use of this scent secretion in mate attraction. The scent of pregnant and lactating females was highly differentiated from male and juvenile scent, but anecdotal reports suggest females avoid communication during these times.     

The spatial organization of otters (Lutra lutra) in Shetland

The dispersion of otters Lutra lutra was studied along the coasts of an area of Shetland. The population was relatively stable, with about one adult otter per kilometre of coast. They fished diurnally in a strip of water usually within 100 m of the shore. Females lived in exclusive group ranges, occupied by up to four females, with well-defined boundaries which were respected by successive occupants. For two groups, where relatively complete information was available, range size was 4.7 and 6.4 km, occupied by two and four females, respectively; a third range was about 14 km, with possibly 4–5 females. Within the group ranges individual females moved on their own or with their cubs. Each had a core area, where they were observed > 50% of the time; these core areas were separate from each other, although each individual used the entire group range. Several or all females in each range bred in any one summer. Male ranges were larger than those of females, and overlapped with two or more female group ranges; there were several resident males along each section of coast, but the information on males was incomplete. The habitats of the sexes were different; males spent more time on exposed coasts than did females. There were frequent visits to all coasts from non-resident otters of both sexes.
Behavioural interactions related to territoriality are described; otters showed little overt aggression, except between adult males, but there was some individual avoidance. It is argued that the survival value of this spatial organization can be understood in terms of resource dispersion, but several types of resources have to be considered (including prey and fresh water), and detailed predictions of spatial organization from resource dispersion are unwarranted.     

Statistical discrimination of sex in Melanoides tuberculata (Gastropoda: Thiaridae)

Systematists may rely on morphometric differences among samples of specimens for the recognition of living and fossil species, even though morphometric differentiation may be caused by non-genetic factors, such as ecophenotypy, differential growth rates and taphonomic mixing. When genetic differences between sexes or among closely related species are expressed as differences in the morphology of the individual or population, potentially valuable information becomes available to the systematist for a variety of genetic and ecological investigations. We have studied the morphology of the freshwater snail Melanoides tuberculata (Muller, 1774) in Israel, where males occur in what would otherwise be normally parthenogenetic (all female) populations. In modern M. tuberculata, sex may be determined by observation of gonadal tissue; in fossil specimens, any classification according to sex must be accomplished using only preservable features of the mineralized shell. Previous research confirmed that in large samples, mean shell shape of male and female snails differed significantly, but the degree of difference was too small to identify the sex of any individual specimen. We apply a three stage process that results with a high degree of accuracy in the discrimination of individual M. tuberculata specimens by sex on the basis of continuous morphological characters: (1) measurement of many aspects of shell morphology of individuals of known sex, and stepwise discrimination to discover which of the variables, if any, contribute to the morphometric differentiation of males from females (one time only, for the species); (2) use of these selected variables in a clustering procedure to make a preliminary assignment of each specimen to sex; (3) use of cluster assignments in a discrimination procedure to optimally predict sex. For species that exhibit morphometric differences between two groups, and for which continuous morphometric variation precludes the a priori recognition of discrete clusters, this sequential procedure may be of broad applicability. These objective methods may be applied to the discrimination within any set of specimens for which the hypothesis of two, and only two, constituent groups may be entertained.     

Lessons from the use of non-invasive genetic sampling as a way to estimate Eurasian otter population size and sex ratio

Given the difficulties in establishing population parameters of elusive animals in the wild by traditional methods, such as trapping, much attention has been given in recent years to non-invasive genetic sampling. Our work compared estimates of population size and sex ratio derived from genetic sampling with the known number and sex of animals released during an otter reintroduction and reports on the pitfalls and opportunities that may be encountered in studies of this kind. This study makes use of 121 samples of otter spraints (faeces) collected over 7 months during a reintroduction in the Upper Thames (UK) where a total of 17 otters was released in two consecutive phases. Spraints were processed with a multiple tubes approach and seven microsatellites were used. Of all collected samples, 19 % were complete for at least five loci, the minimum required for discrimination between individuals. Six out of nine of the otters that were released in the first phase were detected, four males and two females, while none of the otters released in the second phase was detected probably due to a combination of sampling pitfalls and otter behaviour. In particular, the specific sex (mostly females) and dominance composition (lower) of the individuals in the second release group may explain our failure to detect individuals in this group. Taken together, our results add further evidence that genetic sampling approaches represent a potentially accurate and non-invasive route to census populations of otters but that the sampling design should take into account factors like the sex ratio and dominance composition of the population in order to maximise detection and minimise error.     

Why Do Large Dogs Sound More Aggressive to Human Listeners: Acoustic Bases of Motivational Misattributions

Previous research has highlighted that while human listeners are capable of estimating the body size of dogs using the acoustic components of their growls, they also rate growls from larger dogs as more being aggressive than growls from smaller dogs. The aim of this study was to investigate the relative contributions of two cues to body size, fundamental frequency (F0) and formant frequency dispersion (Δf) to perceived levels of aggression. We found that participants that had just made an accurate assessment of caller size based on these characteristics then misattributed aggressiveness levels on the basis of these same size‐related acoustic cues. More specifically, stimuli in which F0 and/or Δf were typical of larger dogs were rated as being more aggressive than stimuli typical of smaller dogs. Although both F0 and Δf influenced aggressiveness ratings independently, their interaction also had a significant affect. These results are discussed with respects to the human tendency to generalise reliable between‐class acoustic cues to within‐class stimuli and the resulting potential for making perceptual misattributions.     

Vocal-auditory functions in the chimpanzee: Vowel perception

The perception of vowels was studied in chimpanzees and humans, using a reaction time task in which reaction times for discrimination of vowels were taken as an index of similarity between vowels. Vowels used were five synthetic and natural Japanese vowels and eight natural French vowels. The chimpanzees required long reaction times for discrimination of synthetic [i] from [u] and [e] from [o], that is, they need long latencies for discrimination between vowels based on differences in frequency of the second formant. A similar tendency was observed for discrimination of natural [i] from [u]. The human subject required long reaction times for discrimination between vowels along the first formant axis. These differences can be explained by differences in auditory sensitivity between the two species and the motor theory of speech perception. A vowel, which is pronounced by different speakers, has different acoustic properties. However, humans can perceive these speech sounds as the same vowel. The phenomenon of perceptual constancy in speech perception was studied in chimpanzees using natural vowels and a synthetic [o]- [a] continuum. The chimpanzees ignored the difference in the sex of the speakers and showed a capacity for vocal tract normalization.     

Species and sex differences in the screams of chimpanzees and bonobos

We examined screams of chimpanzees and bonobos to investigate interspecific and intraspecific variability in call structure. Measurement of 11 acoustic features of screams revealed differences between and within species. One-way analyses of variance and discriminant function analyses show that the calls of chimpanzees and bonobos differ primarily in spectral characteristics. Spectral features also account for acoustic differences between the sexes. These acoustic variations may be attributable to differences in body size and social dispersion between the two species and sexes. The effectiveness with which an acoustic feature could be used to discriminate the two species and female bonobos from male bonobos is negatively associated with its relative variability. These data are consistent with the hypothesis that optimal signals for group identification vary little within groups but differ widely between groups.     

Breeding durations as estimators of adult sex ratios and population size

Adult sex ratios (ASRs) and population size are two of the most fundamental parameters in population biology, as they are the main determinants of genetic and demographic viability, and vulnerability of a population to stochastic events. Underpinning the application of population viability analysis for predicting the extinction risk of populations is the need to accurately estimate parameters that determine the viability of populations (i.e. the ASR and population size). Here we demonstrate that a lack of temporal information can confound estimation of both parameters. Using acoustic telemetry, we compared differences in breeding durations of both sexes for a giant Australian cuttlefish Sepia apama breeding aggregation to the strongly male-biased operational sex ratio (4:1), in order to estimate the population ASR. The ratio of breeding durations between sexes was equal to the operational sex ratio, suggesting that the ASR is not strongly male-biased, but balanced. Furthermore, the short residence times of individuals at the breeding aggregation suggests that previous density-based abundance estimates have significantly underestimated population size. With the current wide application of population viability analysis for predicting the extinction risk of populations, tools to improve the accuracy of such predictions are vital. Here we provide a new approach to estimating the fundamental ASR parameter, and call for temporal considerations when estimating population size.     

Geographic variation in sex‐chromosome differentiation in the common frog (Rana temporaria)

In sharp contrast with birds and mammals, sex‐determination systems in ectothermic vertebrates are often highly dynamic and sometimes multifactorial. Both environmental and genetic effects have been documented in common frogs (Rana temporaria). One genetic linkage group, mapping to the largest pair of chromosomes and harbouring the candidate sex‐determining gene Dmrt1, associates with sex in several populations throughout Europe, but association varies both within and among populations. Here, we show that sex association at this linkage group differs among populations along a 1500‐km transect across Sweden. Genetic differentiation between sexes is strongest (F_ST = 0.152) in a northern‐boreal population, where male‐specific alleles and heterozygote excesses (F_IS = ?0.418 in males, +0.025 in females) testify to a male‐heterogametic system and lack of X‐Y recombination. In the southernmost population (nemoral climate), in contrast, sexes share the same alleles at the same frequencies (F_ST = 0.007 between sexes), suggesting unrestricted recombination. Other populations show intermediate levels of sex differentiation, with males falling in two categories: some cluster with females, while others display male‐specific Y haplotypes. This polymorphism may result from differences between populations in the patterns of X‐Y recombination, co‐option of an alternative sex‐chromosome pair, or a mixed sex‐determination system where maleness is controlled either by genes or by environment depending on populations or families. We propose approaches to test among these alternative models, to disentangle the effects of climate and phylogeography on the latitudinal trend, and to sort out how this polymorphism relates to the ‘sexual races’ described in common frogs in the 1930s.     

Production and perception of sex differences in vocalizations of Wied's black‐tufted‐ear marmosets (Callithrix kuhlii)

Males and females from many species produce distinct acoustic variations of functionally identical call types. Social behavior may be primed by sex‐specific variation in acoustic features of calls. We present a series of acoustic analyses and playback experiments as methods for investigating this subject. Acoustic parameters of phee calls produced by Wied's black‐tufted‐ear marmosets (Callithrix kuhlii) were analyzed for sex differences. Discriminant function analyses showed that calls contained sufficient acoustic variation to predict the sex of the caller. Several frequency variables differed significantly between the sexes. Natural and synthesized calls were presented to male–female pairs. Calls elicited differential behavioral responses based on the sex of the caller. Marmosets became significantly more vigilant following the playback of male phee calls (both natural and synthetic) than following female phee calls. In a second playback experiment, synthesized calls were modified by independently manipulating three parameters that were known to differ between the sexes (low‐, peak‐, and end‐frequency). When end‐frequency‐modified calls were presented, responsiveness was differentiable by sex of caller but did not differ from responses to natural calls. This suggests that marmosets did not use end‐frequency to determine the sex of the caller. Manipulation of peak‐and low‐frequency parameters eliminated the discrete behavioral responses to male and female calls. Together, these parameters may be important features that encode for the sex‐specific signal. Recognition of sex by acoustic cues seems to be a multivariate process that depends on the congruency of acoustic features. Am. J. Primatol. 71:324–332, 2009. © 2008 Wiley‐Liss, Inc.     

Variable asymmetry and resonance in the avian vocal tract: a structural basis for individually distinct vocalizations

The social vocalizations of the oilbird (Steatornis caripensis) frequently have their acoustic energy concentrated into 3 prominent formants which appear to arise from the filter properties of their asymmetrical vocal tract with its bronchial syrinx. The frequency of the second and third formants approximate the predicted fundamental resonances of the unequal left and right cranial portions of each primary bronchus, respectively. Reversibly plugging either bronchus eliminates the corresponding formant. The first formant may arise in the trachea. The degree of vocal tract asymmetry varies between individuals, endowing them with different formant frequencies and providing potential acoustic cues by which individuals of this nocturnal, cave dwelling species may recognize each other in their dark, crowded colonies.     

Low frequency groans indicate larger and more dominant fallow deer (Dama dama) males

Background

Models of honest advertisement predict that sexually selected calls should signal male quality. In most vertebrates, high quality males have larger body sizes that determine higher social status and in turn higher reproductive success. Previous research has emphasised the importance of vocal tract resonances or formant frequencies of calls as cues to body size in mammals. However, the role of the acoustic features of vocalisations as cues to other quality-related phenotypic characteristics of callers has rarely been investigated.

Methodology/Principal Findings

We examined whether the acoustic structure of fallow deer groans provides reliable information on the quality of the caller, by exploring the relationships between male quality (body size, dominance rank, and mating success) and the frequency components of calls (fundamental frequency, formant frequencies, and formant dispersion). We found that body size was not related to the fundamental frequency of groans, whereas larger males produced groans with lower formant frequencies and lower formant dispersion. Groans of high-ranking males were characterised by lower minimum fundamental frequencies and to a lesser extent, by lower formant dispersions. Dominance rank was the factor most strongly related to mating success, with higher-ranking males having higher mating success. The minimum fundamental frequency and the minimum formant dispersion were indirectly related to male mating success (through dominance rank).

Conclusion/Significance

Our study is the first to show that sexually selected vocalisations can signal social dominance in mammals other than primates, and reveals that independent acoustic components encode accurate information on different phenotypic aspects of male quality.     

Sounds produced by the cichlid fish Metriaclima zebra allow reliable estimation of size and provide information on individual identity

Sounds produced by male cichlids Metriaclima zebra during aggressive interactions were recorded to conduct a detailed analysis and to search for potential individual acoustic signatures. Fish from two different size groups (small and large individuals) were analysed. The two groups were significantly different for all acoustic variables considered; six of seven features demonstrated a significant interindividual variability and most of them were correlated with the size of the emitter. A cross-validated and permuted discriminant function analysis (pDFA) separated the two groups and correctly classified around 50% of the sounds to the correct individuals. Acoustic features that best distinguished among males were the instantaneous frequency of sounds and the modulation of pulse amplitude. These results suggest that acoustic signals could bear information about individual identity. The long-term stability of this signature is likely to be weak since the signature of a growing individual may change over time.     

Differentiation of subspecies and sexes of Beringian Dunlins using morphometric measures

Five subspecies of Dunlins (Calidris alpina) that breed in Beringia are potentially sympatric during the non‐breeding season. Studying their ecology during this period requires techniques to distinguish individuals by subspecies. Our objectives were to determine (1) if five morphometric measures (body mass, culmen, head, tarsus, and wing chord) differed between sexes and among subspecies (C. a. actites, arcticola, kistchinski, pacifica, and sakhalina), and (2) if these differences were sufficient to allow for correct classification of individuals using equations derived from discriminant function analyses. We conducted analyses using morphometric data from 10 Dunlin populations breeding in northern Russia and Alaska, USA. Univariate tests revealed significant differences between sexes in most morphometric traits of all subspecies, and discriminant function equations predicted the sex of individuals with an accuracy of 83–100% for each subspecies. We provide equations to determine sex and subspecies of individuals in mixed subspecies groups, including the (1) Western Alaska group of arcticola and pacifica (known to stage together in western Alaska) and (2) East Asia group of arcticola, actites, kistchinski, and sakhalina (known to winter together in East Asia). Equations that predict the sex of individuals in mixed groups had classification accuracies between 75% and 87%, yielding reliable classification equations. We also provide equations that predict the subspecies of individuals with an accuracy of 22–96% for different mixed subspecies groups. When the sex of individuals can be predetermined, the accuracy of these equations is increased substantially. Investigators are cautioned to consider limitations due to age and feather wear when using these equations during the non‐breeding season. These equations will allow determination of sexual and subspecies segregation in non‐breeding areas, allowing implementation of taxonomic‐specific conservation actions.     

The influence of age,sex, population group,and dentition on the mandibular angle as measured on a South African sample

The mandibular angle is measured in physical anthropological assessments of human remains to possibly assist with the determination of sex and population affinity. The purpose of this investigation was to establish how the mandibular angle changes with age and loss of teeth among the sexes in South African population groups. The angles of 653 dried adult mandibles from the Pretoria Bone Collection were measured with a mandibulometer. Males and females of both South African whites and blacks were included. To compensate for imbalances in numbers among subgroups, type IV ANOVA testing was applied. No association was found between age and angle within either of the populations, within sexes, or within dentition groups. The angle was the most obtuse in individuals without molars and with an uneven distribution of molars, and most acute in the group with an even distribution of molars on both sides. Statistically significant differences (P < 0.001) were found in the angle between the two population groups and sexes in the overall sample as well as in the subgroup with absent molar teeth (P = 0.003 for sex, males more acute angle, and P = 0.001 for population group, blacks more acute angle), although a very large overlap existed. No significant differences could be demonstrated between the sexes or populations within the subgroups with molars. We concluded that the loss of molars, especially if complete or uneven, has a considerable effect on the mandibular angle. In the assessment of human remains, the mandibular angle is not very usable in determining sex. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Conflict between Fishermen and Giant Otters Pteronura brasiliensis in Western Brazilian Amazon

The recovery of giant otter populations after the hunting prohibition and restriction of the pelt trade resulted in more frequent conflicts with fishermen. In this study, fisherman–giant otter conflicts were analyzed in the Uacari Sustainable Development Reserve, where giant otters are accused of interfering with fisheries by eating the fish (predation), frightening the fish away (local interference), and damaging fishing equipment (direct interference). Interference by predation was analyzed by evaluating overlap in fish species consumption between humans (measured by subsistence and commercial catches) and giant otters. The giant otter diet was assessed from fecal samples, and the human diet through questionnaires. Local and direct interferences were analyzed through fish samples using gillnets and comparing capture efficiency with and without giant otters’ presence. The overlap between human and giant otter diets was low (0.37), varied seasonally, and was smaller in the low water (0.24) than in the high water period (0.60), when both species tend to be more generalists. Overlap between fish species consumed by giant otters and those exploited by commercial fisheries was small (0.34). Giant otter presence during the experimental fishing was low (9.5%), restricted to the high water period, and did not significantly reduce the captures (U = 13, P = 0.61). The low overlap in diet may be a result of differences in preferences and fishing strategies. The conflict between giant otters and fishermen is greater in the high water period, when the income of the fisheries decreases; however, the conflict seems to be mainly motivated by the resident's prejudice against giant otters.