Male sperm whale (Physeter macrocephalus) acoustics in a high-latitude habitat: implications for echolocation and communication

Sperm whales (Physeter macrocephalus) are deep-diving predators foraging in meso- and bathypelagic ecosystems off the continental shelves. To investigate the ecophysiological and communicative function of various click types from male sperm whales in a high-latitude habitat, we deployed a large-aperture array of calibrated hydrophones off northern Norway (N69, E15). Data show that sperm whales in this habitat produce three click types: usual clicks, creak clicks and, occasionally, slow clicks. Usual clicks and creak clicks exhibit short duration, profound directionality and a frequency content suited for echolocation on meso- and bathypelagic fish and squids. The acoustic properties and low repetition rate of usual clicks are suited for long-range echolocation, whereas creak clicks have properties equivalent to signals in buzzes, the terminating pulse trains known from echolocating bats during prey capture. From these source parameters and the high acoustic activity during foraging dives, it is concluded that echolocation is an important sensory cue in prey location. Sound pressure levels of creak clicks and usual clicks measured off the acoustic axis suggest that sperm whales may be subjected to eavesdropping by conspecifics, thereby conveying information about food aggregations at estimated ranges of 6 km for creak clicks and 16 km for usual clicks. Slow clicks exhibit low directionality, low-frequency emphasis and a reduced repetition rate, suggesting that this click type is more suited for communication than for echolocation. Slow clicks can be detected by submerged conspecifics at ranges up to 60 km. Thus, sperm whales producing slow clicks may represent an odontocete species that utilizes long-range sound communication.     

Song repertoires and density assessment in red-winged blackbirds: further tests of the Beau Geste hypothesis

Summary The Beau Geste hypothesis proposes that song repertoires are advantageous in territory defense because nonterritorial males, who are prospecting for territories, use the number of song types they hear to assess the density of territorial males, and then avoid densely-settled areas. A territorial male can then inflate the apparent density of singers on his territory by singing several distinct song type. This hypothesis assumes (1) a positive correlation between the density of song types and the density of territorial males, and (2) a negative correlation between the rate of trespassing and the densities of both song types and territorial males. We studied the behavior of male red-winged blackbirds (Agelaius phoeniceus) and found support for the first, but not the second, assumption. Results showing a positive correlation between density of song types and density of territorial males show that the proposed method of density assessment is feasible in this species. However, we could find no support for the assumption that nonterritorial male red-winged blackbirds avoid densely settled areas. In contrast to the assumed negative correlation, rate of trespass per territory was not consistently correlated with male density, and trespass rate per unit area was positively correlated with male density. Further, these relationships were retained when we controlled statistically for the effects of territory quality. We conclude that prospecting male red-winged blackbirds to not attempt to avoid densely settled areas, and that, although they do avoid territories defended by many song types, they do not use song type density to assess the density of territorial males. It is thus unlikely that the Beau Geste hypothesis adequately accounts for the evolution of song repertoires in the red-winged blackbird.     

Are vocal signals used to recognize individuals during male–male competition in greater prairie-chickens (Tympanuchus cupido)?

Vocal signaling can be an important component of vertebrate communication during social interactions. If vocalizations vary among individuals but are consistent within a given individual, they may be used to discriminate among individuals. In many species, territorial males use vocalizations to discriminate between neighbors and strangers and either respond more aggressively toward strangers relative to neighbors (“dear enemy” effect) or they respond more aggressively toward neighbors relative to strangers (“nasty neighbor” effect). In the greater prairie-chicken (Tympanuchus cupido), male vocalizations are an integral part of the display males produce on leks. We investigated whether male greater prairie-chickens discriminate among familiar individuals on their own territory, familiar individuals outside their normal territory and strangers from a nearby lek. Vocal characteristics varied among males, suggesting that vocalizations may potentially be used by prairie-chickens to identify individuals. Males responded to playback of prairie-chicken calls by vocalizing at a faster rate and approaching the playback speaker, but did not vary in their response to the vocalizations based on the identity of the caller. Our results suggest that variation is present among the vocalizations of individual male greater prairie-chickens, but males do not appear to discriminate among familiar individuals and strangers based solely on their “boom” vocalizations. Greater prairie-chicken vocalization likely functions as a way of announcing that a territory is occupied and defended, but it may also serve as a way of advertising to conspecifics or as a signal that is secondary to other forms of communication.     

Patterns of egg yolk antioxidant co-variation in an avian brood parasite–host system

Despite their importance to offspring fitness outcomes, there has been little previous work on egg-mediated maternal effects in avian brood parasites which lay their eggs in the nests of other species. Here, we examine patterns of egg yolk antioxidant deposition in an avian host (red-winged blackbird, Agelaius phoeniceus) and their brood parasite (brown-headed cowbird, Molothrus ater). Cowbird nestlings compete directly for food with host blackbird chicks and succeed, despite hatching from a smaller egg, by begging more intensely and growing as or more rapidly than their hosts. Dietary-derived antioxidants, such as carotenoids and vitamins E and A, deposited in egg yolk, may enhance growth rate and reduce oxidative stress, and therefore provide a potential mechanism by which female cowbirds could enhance the competitiveness of their young in host nests. However, provision of antioxidants to eggs is costly so we predicted that female cowbirds should adjust the amount of antioxidants in response to variation in host quality. We found that whilst red-winged blackbird clutches parasitised by cowbirds did not differ in the pattern of antioxidant deposition compared to non-parasitised clutches, levels of vitamin E deposited in cowbird eggs were closely matched to that of the individual host clutch in which cowbirds laid their egg. This supports the prediction that female cowbirds adjust the antioxidant concentration of their eggs to increase the likely competitiveness of their offspring in the host nest into which they are laid and may help explain the success of cowbirds in competing with larger host young.     

Polygyny in the red-winged blackbird: do females prefer monogamy or polygamy?

Five hypotheses have been proposed to explain polygyny in the red-winged blackbird (Agelaius phoeniceus). We categorized the hypotheses into three groups based on female preference for unmated versus monogamously mated males: (1) the “polygyny threshold” model, “sexy son” hypothesis and the “asynchronous settlement” model, which assume that females prefer unmated males to mated males on breeding situations of homogeneous quality; (2) the “neutral mate choice” hypothesis, which assumes that females have no preference; and (3) the “cooperative female choice” model, which assumes that females prefer monogamously mated males to unmated males. We tested the direction of female preference in two field experiments. In both experiments, newly settling females were given a choice of two adjacent territories, one defended by an unmated male and the other by a monogamously mated male. Male mating status was randomized with respect to the variation in territory quality and male quality. Early in the breeding season, significantly more females settled with the unmated males than with the mated males. Although more females settled with the unmated males than with the mated males late in the breeding season, the difference was no longer significant. Female settlement late in the season appeared to be related to the tenure of the resident females: the new females avoided territories where the resident females were in early stages of their nesting, but settled on territories where the resident females were in late stages. The pattern of female settlement shows that females prefer unmated males to mated males. The preference is consistent with the polygyny threshold model, sexy son hypothesis and the asynchronous settlement model, and inconsistent with the neutral mate choice hypothesis and the cooperative female choice model. For this reason, the latter two hypotheses are unlikely to explain the occurrence of polygyny in our population of red-winged blackbirds. Received: 1 December 1994 / Accepted after revision: 28 October 1995     

Male Scudderia pistillata katydids defend their acoustic duet against eavesdroppers

Acoustic duets—in which the female reveals herself to the advertising male—are a common means of establishing a temporary, pre-mating pair bond within Phaneropterinae katydids. Such duets, however, are especially susceptible to eavesdropping males that orient to signaling females and interrupt the established duet. It would therefore be advantageous for an advertising male to protect his investment in a duet from eavesdroppers. Male Scudderia pistillata katydids produce acoustic advertisement calls after which a conspecific female enters the duet by responding during a specific time-window with her own call, one or more acoustic ticks. We demonstrate here that when a duet is occurring in the presence of other calling males, the focal male produces a single acoustic tick that has a spectrum similar to that of the female’s tick and occurs during the time-window in which the female would respond to his advertisement call within the duet. The male acoustic tick may confound eavesdropping males if they have difficulty performing accurate phonotaxis to a female sound source when sounds arrive from two locations. We tested this hypothesis in laboratory arenas and determined that silent males eavesdrop on advertising male’s duets and accurately locate his female. Moreover, we found that the added acoustic tick produced by advertising males serves to mimic the female response and reduces the accuracy of eavesdropping males to localize the female. Therefore, male ticks appear to act as a form of pre-copulatory acoustic mate guarding of the calling male’s temporary pair bond with the female.     

Variation in the temporal and spatial use of signals and its implications for multimodal communication

The use of signals across multiple sensory modalities in communication is common in animals and plants. Determining the information that each signal component conveys has provided unique insights into why multimodal signals evolve. However, how these complex signals are assessed by receivers will also influence their evolution, a hypothesis that has received less attention. Here, we explore multimodal signal assessment in a closely related complex of island flycatchers that have diverged in visual and acoustic signals. Using field experiments that manipulated song and plumage colour, we tested if song, a possible long-range signal, is assessed before plumage colour in conspecific recognition. We find that divergent song and colour are assessed in sequence, and this pattern of sequential assessment is likely mediated by habitat structure and the extent of differences in signal characteristics. A broad survey of the literature suggests that many organisms from a wide range of taxa sequentially assess multimodal signals, with long-range signals attracting conspecifics for further assessment of close-range signals. Our results highlight the need to consider how signals are assessed when understanding multimodal signal evolution. Finally, given the results of our field experiments indicating sequential assessment of divergent song and colour in the recognition of conspecifics, we discuss the consequences of multimodal signal divergence for the origin of species, as changes in signals across different sensory modalities may influence the evolution of premating reproductive isolation.     

Voices in the dark: predation risk by owls influences dusk singing in a diurnal passerine

Predation is an important cost of communication in animals and thus a potent selection pressure on the evolution of signaling behavior. Heterospecific eavesdropping by predators may increase the vulnerability of vocalizing prey, particularly during low light, such as at dusk when nocturnal predators are actively hunting. Despite the risk it entails, dawn and dusk chorusing is common in passerines. However, the dusk chorus has not been studied much, neglecting the opportunity for understanding how eavesdropping between predators and prey may shape communication in birds. Here, we report the first demonstration of simulated predation risk (playback of owl vocalizations) altering the dusk chorus of a diurnal passerine, the veery (Catharus fuscescens). Veeries have a pronounced dusk chorus, singing well after sunset and potentially exposing themselves to predation by owls. In response to brief playbacks of owl calls (~30 s of calls presented three times over 25 min), veeries sang fewer songs post-sunset and stopped singing earlier relative to control trials. These changes in singing remained evident 30 min after the last owl stimulus. Although the avian dusk chorus has received relatively little attention to date, our results suggest that the dusk chorus may pose a higher predation risk to singing males that may influence the evolution of singing behavior in diurnal birds.     

Female preferences for multiple attributes in the acoustic signals of the Italian treefrog, <Emphasis Type="Italic">Hyla intermedia</Emphasis>

The advertisement call of frogs and toads is an example of multiple message signal because different acoustic properties encode different kinds of biologically significant information. In the Italian treefrog, Hyla intermedia, pulse rate and frequency have been found to be under stabilizing female preferences and to encode information important for mate recognition, whereas the number of calls per call group have been found to be under directional preferences and, thus, to be important for mate quality assessment. In this study, we investigate preferences for calls that differ simultaneously in frequency, pulse rate, and number of calls per call group, and we ask how these properties interact with each other in influencing female mating decisions. Results of two-choice phonotaxis experiments provide no evidence to support the hypothesis that females process multi-attribute signals in a hierarchical way. In contrast, the pattern of preferences is consistent with the ‘preference function’ hypothesis, that is, with the hypothesis that females rank signals along an ordinal scale of values and choose accordingly. Pulse rate and frequency influence mating preferences more than does the number of calls per call group. The interaction between pulse rate and frequency is not additive but multiplicative: small differences in either pulse rate or frequency that, alone, have no effects on female choice, interact synergistically so that their combination has strong influence on female preferences. A preference repeatability test shows strong among-female differences in preference for multi-attribute signals. We suggest that this result reveals not only a variation in attribute values among females, but also a variation in the way females weight and combine attribute values into a single preference score.     

Song length variation serves multiple functions in the white-crowned sparrow

A wide variety of hypotheses has been proposed to explain the structural diversity in bird song repertoires. Song diversity is frequently described in terms of song “types” or within-type “variants.” Male Puget Sound white-crowned sparrows, Zonotrichia leucophrys pugetensis, produce variants of their single adult song type by altering the number of repetitions of syllables in the terminal trill. We tested whether variation in trill length correlated with distance to the receiver and with signaling context as predicted by the eavesdropping avoidance hypothesis and the strategic signaling hypothesis. In accordance with the eavesdropping avoidance hypothesis, males sang variants with shorter trills, and sang quieter and less frequently when near their mate during the incubation phase than during spontaneous singing while unpaired. Males also sang variants with short trills, but at a high rate and variable amplitude when within 10 m of an opponent during close male–male territorial interactions. In agreement with the strategic signaling hypothesis, males decreased trill length immediately before chasing an opponent, but did not change length consistently prior to flight. We conclude that the occurrence of short quiet songs sung near the mate agrees with predictions of the eavesdropping avoidance hypothesis, while short songs sung near other males are best explained by the strategic signaling hypothesis. Trill length variation may be a conventional signal of aggressive intentions in male–male contests stabilized by receiver-imposed retaliation costs.     

Communication by changing signals: call switching in red-winged blackbirds

Summary Territorial male red-winged blackbirds use many different alert calls which overlap broadly in the context of their use and which are often given in continuous, repetitive fashion. Males give one call type many times before switching to another type, and call during all daily activities. In this field study we demonstrate that males tend to give the same call types as their neighbors and change types to match the changes of their neighbors. Individual males change call types in response to the appearance of a mounted hawk and also match call type changes broadcast from a loudspeaker. The various call types are not associated with particular behavioral contexts. We suggest that red-winged blackbirds operate a general acoustic alert system by calling repetitiously and changing call types when detecting environmental changes such as appearances of predators. Evidence is presented that communication is acheived primarily during switching among different call signals, and not by specific calls that refer to particular stimuli or states of alertness.     

Singing onstage: female and male common nightingales eavesdrop on song type matching

In bird communication, listening individuals may obtain information on the quality and motivation of a male not only from solo-singing, but also from song interactions and listeners base their future decisions in territorial and mating contexts on such public information. Eavesdropping on male interactions may thus have a strong influence on sexual selection. In singing interactions, temporal coordination (e.g. overlapping vs. alternating) of two singers as well as structural interaction patterns (e.g. song type matching or repertoire matching) have been described, but the latter is far less studied. By conducting dual-speaker playback experiments with common nightingales Luscinia megarhynchos, we simulated an interaction where one singer was repeatedly song–type matching his counterpart. Playbacks were broadcast to male and female nightingales, and their approach behaviour and singing responses (in the case of male focals) were analysed. We found that both, males and females, spent more time with the matched bird, whereas males additionally sang more songs towards the matching bird. This can be taken as strong hint that eavesdropping occurs in nightingale communication and that listening to male vocal contests might be an important strategy for both sexes to adjust their behavioural output. With regard to the function of song matching, we assume that song-matching is not an aggressive signal per se in nightingales. We rather conclude that vocal leaders within an interaction, here the matched bird, may elicit stronger responses in conspecifics than vocal followers, here the matching bird.     

Bayesian estimation of age-specific bird nest survival rates with categorical covariates

The populations of many North American landbirds are showing signs of declining. Gathering information on breeding productivity allows critical assessment of population performance and helps identify good habitat-management practices. He (Biometrics (2003) 59 962–973) proposed a Bayesian model to estimate the age-specific nest survival rates. The model allows irregular visiting schedule under the assumption that the observed nests have homogeneous nest survival. Because nest survival studies are often conducted in different sites and time periods, it is not realistic to assume homogeneous nest survival. In this paper, we extend He’s model by incorporating these factors as categorical covariates. The simulation results show that the Bayesian hierarchical model can produce satisfactory estimates on nest survival and capture different factor effects. Finally the model is applied to a Missouri red-winged blackbird data set.     

Use of the song repertoire in intersexual and intrasexual contexts by male red-winged blackbirds

Summary Live and stuffed male and female red-winged blackbirds (Agelaius phoeniceus) were presented to territorial males in order to study the repertoire organization of males during intersexual and intrasexual encounters. Territorial male red-winged blackbirds switched song types more frequently and sang more song types in response to presentation of a female stimulus than during control periods, but decreased their switching frequency when a male stimulus was presented. Switching frequency in response to the female stimulus was three times that in response to the male stimulus. These results support the hypothesis that song repertoires of male red-winged blackbirds evolved primarily in response to intersexual rather than intrasexual selection. They also suggest that male red-winged blackbirds have been selected to de-emphasize their song repertoires during encounters with conspecific males as a result of some as yet unidentified cost. Offprint requests to: W.A. Searcy     

Multisensory vocal communication in primates and the evolution of rhythmic speech

The integration of the visual and auditory modalities during human speech perception is the default mode of speech processing. That is, visual speech perception is not a capacity that is “piggybacked” on to auditory-only speech perception. Visual information from the mouth and other parts of the face is used by all perceivers to enhance auditory speech. This integration is ubiquitous and automatic and is similar across all individuals across all cultures. The two modalities seem to be integrated even at the earliest stages of human cognitive development. If multisensory speech is the default mode of perception, then this should be reflected in the evolution of vocal communication. The purpose of this review is to describe the data that reveal that human speech is not uniquely multisensory. In fact, the default mode of communication is multisensory in nonhuman primates as well but perhaps emerging with a different developmental trajectory. Speech production, however, exhibits a unique bimodal rhythmic structure in that both the acoustic output and the movements of the mouth are rhythmic and tightly correlated. This structure is absent in most monkey vocalizations. One hypothesis is that the bimodal speech rhythm may have evolved through the rhythmic facial expressions of ancestral primates, as indicated by mounting comparative evidence focusing on the lip-smacking gesture.     

Parent-absent signalling of need and its consequences for sibling competition in the barn swallow

In species with parental care, competition among siblings for access to limited parental resources is common. Sibling competition can be mediated by begging behaviour, a suite of different visual and acoustic displays by which offspring solicit parental care. These are mostly addressed to the parents upon food provisioning, but can also be performed in the absence of the attending parents. This so-called parent-absent begging (PAB) may function as an intrabrood communication signal and potentially affect intrabrood competition dynamics for access to food. We investigated the role of PAB in moulding sibling interactions and its effect on food intake among altricial barn swallow (Hirundo rustica) nestlings, both under normal and experimentally reduced food intake. Frequency of PAB increased after food deprivation. Nestlings that had performed PAB increased their begging intensity upon the subsequent parental feeding visit, while siblings reduced their own begging level, but only when they had not been food-deprived. As a consequence, nestlings which had performed PAB before parental arrival had larger chances of receiving food. However, nestlings did not benefit from displaying PAB when competing with food-deprived siblings. Our findings show that PAB reliably reflects need of food, indicating that a nestling will vigorously compete for the subsequent food item. By eavesdropping siblings' PAB displays, nestlings may optimally balance the costs of scrambling competition, the direct fitness gains of being fed and the indirect fitness costs of subtracting food to needy kin. However, large asymmetries in satiation between competitors may lead individual offspring to monopolize parental resources, irrespective of PAB displays.     

A novel method for quantifying the glossiness of animals

The glossy sheen of healthy hair is an ideal of human beauty; however, glossiness has never been quantified in the context of non-human animal signaling. Glossiness, the specular reflectance characteristic of polished surfaces, has the potential to act as a signal of quality because it depends upon material integrity and cleanliness. Here, we undertook two studies of glossiness in avian plumage to determine (a) the repeatability of a recently developed measure of glossiness, (b) the relationship between glossiness and conventional measures of coloration, and (c) how glossiness is associated with quality signaling. Using museum specimens of three North American bird species with glossy plumage (red-winged blackbird, Agelaius phoeniceus; great-tailed grackle, Quiscalus mexicanus; Chihuahuan raven, Corvus cryptoleucus), we found that the glossiness measure was highly repeatable for all species and was significantly correlated with plumage coloration (e.g., chroma, brightness) in male great-tailed grackles. We then used wild-caught grackles to examine sexual dimorphism in plumage glossiness and its correlation to a potentially sexually selected trait in this species, male tail length. We found that males were significantly glossier than females and that male, but not female, glossiness correlated positively with tail length. This study provides a repeatable method to measure glossiness and highlights its potential as a signal of individual quality in animals.     

Seasonal variation in ejaculate traits of male red-winged blackbirds (Agelaius phoeniceus)

Many reproductive traits, including ejaculate characteristics, usually show remarkable seasonal variation, but the potential for such dynamics in sperm morphology has been overlooked. Several studies have revealed high within-male repeatability in sperm morphology, but samples have typically been collected within a few hours or days, and the consistency of sperm morphology over longer periods remains unexplored. Here, we tested whether ejaculate traits, including sperm morphology, sperm number, and sperm velocity, exhibit seasonal phenotypic plasticity in a long-lived seasonal breeder, the red-winged blackbird (Agelaius phoeniceus). We found absolute and/or relative flagellum length and sperm velocity to increase across the season, whereas sperm numbers within ejaculates declined. Sperm morphological traits were further positively associated with harem size or the total number of offspring that fledged in each male’s territory, suggesting that sperm morphology is likely to be linked to male reproductive quality. The underlying mechanisms of these patterns of seasonal variation remain unresolved, but we discuss our results in the context of dynamics of reproductive hormones, testicular structures and function, and reproductive behavior.     

Automated detection of frog calls and choruses by pulse repetition rate

Anurans (frogs and toads) are among the most globally threatened taxonomic groups. Successful conservation of anurans will rely on improved data on the status and changes in local populations, particularly for rare and threatened species. Automated sensors, such as acoustic recorders, have the potential to provide such data by massively increasing the spatial and temporal scale of population sampling efforts. Analyzing such data sets will require robust and efficient tools that can automatically identify the presence of a species in audio recordings. Like bats and birds, many anuran species produce distinct vocalizations that can be captured by autonomous acoustic recorders and represent excellent candidates for automated recognition. However, in contrast to birds and bats, effective automated acoustic recognition tools for anurans are not yet widely available. An effective automated call-recognition method for anurans must be robust to the challenges of real-world field data and should not require extensive labeled data sets. We devised a vocalization identification tool that classifies anuran vocalizations in audio recordings based on their periodic structure: the repeat interval-based bioacoustic identification tool (RIBBIT). We applied RIBBIT to field recordings to study the boreal chorus frog (Pseudacris maculata) of temperate North American grasslands and the critically endangered variable harlequin frog (Atelopus varius) of tropical Central American rainforests. The tool accurately identified boreal chorus frogs, even when they vocalized in heavily overlapping choruses and identified variable harlequin frog vocalizations at a field site where it had been very rarely encountered in visual surveys. Using a few simple parameters, RIBBIT can detect any vocalization with a periodic structure, including those of many anurans, insects, birds, and mammals. We provide open-source implementations of RIBBIT in Python and R to support its use for other taxa and communities.     

Echolocation beam shape in emballonurid bats, Saccopteryx bilineata and Cormura brevirostris

The shape of the sonar beam plays a crucial role in how echolocating bats perceive their surroundings. Signal design may thus be adapted to optimize beam shape to a given context. Studies suggest that this is indeed true for vespertilionid bats, but little is known from the remaining 16 families of echolocating bats. We investigated the echolocation beam shape of two species of emballonurid bats, Cormura brevirostris and Saccopteryx bilineata, while they navigated a large outdoor flight cage on Barro Colorado Island, Panama. C. brevirostris emitted more directional signals than did S. bilineata. The difference in directionality was due to a markedly different energy distribution in the calls. C. brevirostris emitted two call types, a multiharmonic shallowly frequency-modulated call and a multiharmonic sweep, both with most energy in the fifth harmonic around 68?kHz. S. bilineata emitted only one call type, multiharmonic shallowly frequency-modulated calls with most energy in the second harmonic (~46?kHz). When comparing same harmonic number, the directionality of the calls of the two bat species was nearly identical. However, the difference in energy distribution in the calls made the signals emitted by C. brevirostris more directional overall than those emitted by S. bilineata. We hypothesize that the upward shift in frequency exhibited by C. brevirostris serves to increase directionality, in order to generate a less cluttered auditory scene. The study indicates that emballonurid bats are forced to adjust their relative harmonic energy instead of adjusting the fundamental frequency, as the vespertilionids do, presumably due to a less flexible sound production.