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North Atlantic right whales (Eubalaena glacialis) ignore ships but respond to alerting stimuli.

North Atlantic right whales were extensively hunted during the whaling era and have not recovered. One of the primary factors inhibiting their recovery is anthropogenic mortality caused by ship strikes. To assess risk factors involved in ship strikes, we used a multi-sensor acoustic recording tag to measure the responses of whales to passing ships and experimentally tested their responses to controlled sound exposures, which included recordings of ship noise, the social sounds of conspecifics and a signal designed to alert the whales. The whales reacted strongly to the alert signal, they reacted mildly to the social sounds of conspecifics, but they showed no such responses to the sounds of approaching vessels as well as actual vessels. Whales responded to the alert by swimming strongly to the surface, a response likely to increase rather than decrease the risk of collision.

Acoustics↗

Motor mechanisms of a vocal mimic: implications for birdsong production.

The diverse vocal performances of oscine songbirds are produced by the independent but coordinated patterns of activity in muscles controlling separate sound generators on the left and right sides of their duplex vocal organ, the syrinx. Species with different song styles use the two sides of their syrinx in different ways to produce their species-typical songs. Understanding how a vocal mimic copies another species' song may provide an insight into whether there are alternative motor mechanisms for generating the model's song and what parts of his song are most difficult to produce. We show here that when a vocal mimic, the northern mockingbird, accurately copies the song of another species it also uses the vocal motor pattern employed by the model species. Deviations from the model's production mechanism result in predictable differences in the mockingbird's song. Species-specific acoustic features of the model seem most difficult to copy, suggesting that they have been exposed to the strongest selective pressure to maximize their performance.

Acoustics↗

Signature-whistle production in undisturbed free-ranging bottlenose dolphins (Tursiops truncatus).

Data from behavioural observations and acoustic recordings of free-ranging bottlenose dolphins (Tursiops truncatus) were analysed to determine whether signature whistles are produced by wild undisturbed dolphins, and how whistle production varies with activity and group size. The study animals were part of a resident community of bottlenose dolphins near Sarasota, Florida, USA. This community of dolphins provides a unique opportunity for the study of signature-whistle production, since most animals have been recorded during capture-release events since 1975. Three mother-calf pairs and their associates were recorded for a total of 141.25 h between May and August of 1994 and 1995. Whistles of undisturbed dolphins were compared with those recorded from the same individuals during capture-release events. Whistles were conservatively classified into one of four categories: signature, probable signature, upsweep or other. For statistical analyses, signature and probable signature whistles were combined into a 'signature' category; upsweep and other whistles were combined into a 'non-signature' category. Both 'signature' and 'non-signature' whistle frequencies significantly increased as group size increased. There were significant differences in whistle frequencies across activity types: both 'signature' and 'non-signature' whistles were most likely to occur during socializing and least likely to occur during travelling. There were no significant interactions between group size and activity type. Signature and probable signature whistles made up ca. 52% of all whistles produced by these free-ranging bottlenose dolphins.

Animals↗

Hornbills can distinguish between primate alarm calls.

Some mammals distinguish between and respond appropriately to the alarm calls of other mammal and bird species. However, the ability of birds to distinguish between mammal alarm calls has not been investigated. Diana monkeys (Cercopithecus diana) produce different alarm calls to two predators: crowned eagles (Stephanoaetus coronatus) and leopards (Panthera pardus). Yellow-casqued hornbills (Ceratogymna elata) are vulnerable to predation by crowned eagles but are not preyed on by leopards and might therefore be expected to respond to the Diana monkey eagle alarm call but not to the leopard alarm call. We compared responses of hornbills to playback of eagle shrieks, leopard growls, Diana monkey eagle alarm calls and Diana monkey leopard alarm calls and found that they distinguished appropriately between the two predator vocalizations as well as between the two Diana monkey alarm calls. We discuss possible mechanisms leading to these responses.

Acoustic Stimulation↗

Acoustic monitoring on a humpback whale (Megaptera novaeangliae) feeding ground shows continual singing into late Spring.

Singing by males is a major feature of the mating system of humpback whales, Megaptera novaeangliae (Borowski). Although a few songs have been opportunistically recorded on the whales' high-latitude feeding grounds, singing in these regions was thought to be only sporadic. We report results from the first continuous acoustic monitoring of a humpback whale feeding ground (off Cape Cod, MA, USA) in spring. Using autonomous sea-floor recording systems, we found singing on a daily basis over the entire 25 day monitoring period, from 14 May to 7 June 2000. For much of the period, song was recorded 24 h per day. These results, combined with evidence for aseasonal conceptions in whaling catch data, suggest that the humpback whale breeding season should no longer be considered as confined to lower-latitude regions in winter. Rather, we suggest breeding extends geographically and temporally onto feeding grounds into at least spring and early summer. Singing at these times represents either low-cost opportunistic advertising by (perhaps relatively few) males to court females that failed to conceive during the winter, and/or possibly an intrasexual display.

Animals↗

Dynamics of jamming avoidance in echolocating bats.

Animals using active sensing systems such as echolocation or electrolocation may experience interference from the signals of neighbouring conspecifics, which can be offset by a jamming avoidance response (JAR). Here, we report JAR in one echolocating bat (Tadarida teniotis: Molossidae) but not in another (Taphozous perforatus: Emballonuridae) when both flew and foraged with conspecifics. In T. teniotis, JAR consisted of shifts in the dominant frequencies of echolocation calls, enhancing differences among individuals. Larger spectral overlap of signals elicited stronger JAR. Tadarida teniotis showed two types of JAR: (i) for distant conspecifics: a symmetric JAR, with lower- and higher-frequency bats shifting their frequencies downwards and upwards, respectively, on average by the same amount; and (ii) for closer conspecifics: an asymmetric JAR, with only the upper-frequency bat shifting its frequency upwards. In comparison, 'wave-type' weakly electric fishes also shift frequencies of discharges in a JAR, but unlike T. teniotis, the shifts are either symmetric in some species or asymmetric in others. We hypothesize that symmetric JAR in T. teniotis serves to avoid jamming and improve echolocation, whereas asymmetric JAR may aid communication by helping to identify and locate conspecifics, thus minimizing chances of mid-air collisions.

Animals↗

Sperm whale behaviour indicates the use of echolocation click buzzes "creaks" in prey capture.

During foraging dives, sperm whales (Physeter macrocephalus) produce long series of regular clicks at 0.5-2 s intervals interspersed with rapid-click buzzes called "creaks". Sound, depth and orientation recording Dtags were attached to 23 whales in the Ligurian Sea and Gulf of Mexico to test whether the behaviour of diving sperm whales supports the hypothesis that creaks are produced during prey capture. Sperm whales spent most of their bottom time within one or two depth bands, apparently feeding in vertically stratified prey layers. Creak rates were highest during the bottom phase: 99.8% of creaks were produced in the deepest 50% of dives, 57% in the deepest 15% of dives. Whales swam actively during the bottom phase, producing a mean of 12.5 depth inflections per dive. A mean of 32% of creaks produced during the bottom phase occurred within 10 s of an inflection (13x more than chance). Sperm whales actively altered their body orientation throughout the bottom phase with significantly increased rates of change during creaks, reflecting increased manoeuvring. Sperm whales increased their bottom foraging time when creak rates were higher. These results all strongly support the hypothesis that creaks are an echolocation signal adapted for foraging, analogous to terminal buzzes in taxonomically diverse echolocating species.

Acoustics↗

Red deer stags use formants as assessment cues during intrasexual agonistic interactions.

While vocal tract resonances or formants are key acoustic parameters that define differences between phonemes in human speech, little is known about their function in animal communication. Here, we used playback experiments to present red deer stags with re-synthesized vocalizations in which formant frequencies were systematically altered to simulate callers of different body sizes. In response to stimuli where lower formants indicated callers with longer vocal tracts, stags were more attentive, replied with more roars and extended their vocal tracts further in these replies. Our results indicate that mammals other than humans use formants in vital vocal exchanges and can adjust their own formant frequencies in relation to those that they hear.

Acoustic Stimulation↗

Flexibility in assessment of prey cues: frog-eating bats and frog calls.

Predators use cues associated with their prey to assess prey quality and to avoid consuming poisonous prey. Considerable attention has been given to predators' use of aposematic cues to assess prey quality, but little is known about predators that eavesdrop on prey cues that are not intended for them. Here we investigate the prey-cue/prey-quality associations of a predator that eavesdrops on the sexual advertisement signals of its prey. Stability is expected in prey-cue/prey-quality associations when mistakes in prey assessment are lethal. Conversely, flexibility is possible when mistakes are less costly. Predators that must respond to temporal and spatial fluctuations in prey availability should be more flexible in their assessment of prey quality. Given these predictions, we examined flexibility in the ability of wild-caught bats to reverse prey-cue/prey-quality associations for a preferred prey and a poisonous one. We found that the predatory bat, Trachops cirrhosus, has a heretofore undescribed ability to reverse its evaluations of the cues that signal preferred prey.

Adaptation, Biological↗

Ambient noise and the design of begging signals.

The apparent extravagance of begging displays is usually attributed to selection for features, such as loud calls, that make the signal costly and hence reliable. An alternative explanation, however, is that these design features are needed for effective signal transmission and reception. Here, we test the latter hypothesis by examining how the begging calls of tree swallow (Tachycineta bicolor) nestlings and the response to these calls by parents are affected by ambient noise. In a field study, we found that call length, amplitude and frequency range all increased with increasing noise levels at nests. In the laboratory, however, only call amplitude increased in response to the playback of noise to nestlings. In field playbacks to parents, similar levels of noise abolished parental preferences for higher call rates, but the preference was restored when call amplitude was increased to the level that nestlings had used in the laboratory study. Our results show that nestling birds, like other acoustic signallers, consistently increase call amplitude in response to ambient noise and this response appears to enhance discrimination by receivers. Thus, selection for signal efficacy may explain some of the seemingly extravagant features of begging displays.

Acoustics↗

Biosonar behaviour of free-ranging porpoises.

Detecting objects in their paths is a fundamental perceptional function of moving organisms. Potential risks and rewards, such as prey, predators, conspecifics or non-biological obstacles, must be detected so that an animal can modify its behaviour accordingly. However, to date few studies have considered how animals in the wild focus their attention. Dolphins and porpoises are known to actively use sonar or echolocation. A newly developed miniature data logger attached to a porpoise allows for individual recording of acoustical search efforts and inspection distance based on echolocation. In this study, we analysed the biosonar behaviour of eight free-ranging finless porpoises (Neophocaena phocaenoides) and demonstrated that these animals inspect the area ahead of them before swimming silently into it. The porpoises inspected distances up to 77 m, whereas their swimming distance without using sonar was less than 20 m. The inspection distance was long enough to ensure a wide safety margin before facing real risks or rewards. Once a potential prey item was detected, porpoises adjusted their inspection distance from the remote target throughout their approach.

Animals↗

Vibrational communication facilitates cooperative foraging in a phloem-feeding insect.

Insects are the dominant herbivores in tropical forests, with a range of mechanisms for exploiting plant resources. For group-living species, such mechanisms may involve communication. The Neotropical treehopper Calloconophora pinguis (Hemiptera: Membracidae) is a sap-feeding species in which groups of siblings feed on new leaves during the brief period of leaf expansion. Using an experimental approach, a process of cooperative foraging among siblings was documented, in which a few individuals in a group behave as scouts, locating a new feeding site and advertising it using plant-borne vibrational signals. Signalling leads to a period of positive feedback in which newly recruited individuals signal in concert with those already there. The food signalling system of C. pinguis is unique in its use of synchronized group displays and in the tight coordination of receiver responses with collective signals. Examples from a number of taxonomic groups show that vibrational communication can allow group-living insects to solve the challenges of feeding on plants, such as remaining in a foraging group or avoiding predation. While most research has focused on leaf-feeding species, sap-feeding species may remove just as much biomass. This study shows that cooperative vibrational communication underlies the ability of a sap-feeding species to exploit plant resources during a narrow window of availability.

Analysis of Variance↗

Geographical variation in sound production in the anemonefish Amphiprion akallopisos.

Because of pelagic-larval dispersal, coral-reef fishes are distributed widely with minimal genetic differentiation between populations. Amphiprion akallopisos, a clownfish that uses sound production to defend its anemone territory, has a wide but disjunct distribution in the Indian Ocean. We compared sounds produced by these fishes from populations in Madagascar and Indonesia, a distance of 6500 km. Differentiation of agonistic calls into distinct types indicates a complexity not previously recorded in fishes' acoustic communication. Moreover, various acoustic parameters, including peak frequency, pulse duration, number of peaks per pulse, differed between the two populations. The geographic comparison is the first to demonstrate 'dialects' in a marine fish species, and these differences in sound parameters suggest genetic divergence between these two populations. These results highlight the possible approach for investigating the role of sounds in fish behaviour in reproductive divergence and speciation.

Agonistic Behavior↗

Parasites affect song complexity and neural development in a songbird.

There is now considerable evidence that female choice drives the evolution of song complexity in many songbird species. However, the underlying basis for such choice remains controversial. The developmental stress hypothesis suggests that early developmental conditions can mediate adult song complexity by perturbing investment in the underlying brain nuclei during their initial growth. Here, we show that adult male canaries (Serinus canaria), infected with malaria (Plasmodium relictum) as juveniles, develop simpler songs as adults compared to uninfected individuals, and exhibit reduced development of the high vocal centre (HVC) song nucleus in the brain. Our results show how developmental stress not only affects the expression of a sexually selected male trait, but also the structure of the underlying song control pathway in the brain, providing a direct link between brain and behaviour. This novel experimental evidence tests both proximate and ultimate reasons for the evolution of complex songs and supports the Hamilton-Zuk hypothesis of parasite-mediated sexual selection. Together, these results propose how developmental costs may help to explain the evolution of honest advertising in the complex songs of birds.

Animals↗

Vocal performance influences male receiver response in the banded wren.

In a variety of songbirds the production of trilled song elements is constrained by a performance tradeoff between how fast a bird can repeat trill units (trill rate) and the range of frequencies each unit can span (frequency bandwidth). High-performance trills serve as an assessment signal for females, but little is known about the signal value of vocal performance for male receivers. We investigated the relationship between trill rate and frequency bandwidth in banded wren (Thryothorus pleurostictus) songs. Trilled song elements showed the same performance tradeoff found in other passerines and individuals differed in performance of some trill types. We tested the hypothesis that males of this species assess each other based on trill performance with a two-speaker experiment, in which territory owners were presented with alternating renditions of the same song type manipulated to differ in trill rate. Subjects were significantly more likely to approach the faster trill stimulus first. However, subjects that received trill types closer to the performance limit spent less time close to the fast speaker. Our results show that male banded wrens discriminate and respond differently to songs based on their vocal performance. Thus, performance of physically challenging songs may be important in intra- as well as inter-sexual assessment.

Acoustic Stimulation↗

Experimental support for a model of birdsong production.

In this work we present an experimental validation of a recently proposed model for the production of birdsongs. We have previously observed that driving the model with simple functions of time, which represent tensions in vocal muscles, produces a wide variety of sounds resembling birdsongs. In this work we drive the model with functions whose time dependence comes from recordings of muscle activities and air sac pressure. We simultaneously recorded the birds' songs and compared them with the synthetic songs. The model produces recognizable songs. Beyond finding a qualitative agreement, we also test some predictions of the model concerning the relative levels of activity in the gating muscles at the beginning and end of a syllable.

Air Sacs↗

Recurrence plot analysis of nonstationary data: the understanding of curved patterns.

Recurrence plots of the calls of the Nomascus concolor (Western black crested gibbon) and Hylobates lar (White-handed gibbon) show characteristic circular, curved, and hyperbolic patterns superimposed to the main temporal scale of the signal. It is shown that these patterns are related to particular nonstationarities in the signal. Some of them can be reproduced by artificial signals like frequency modulated sinusoids and sinusoids with time divergent frequency. These modulations are too faint to be resolved by conventional time-frequency analysis with similar precision. Therefore, recurrence plots act as a magnifying glass for the detection of multiple temporal scales in slightly modulated signals. The detected phenomena in these acoustic signals can be explained in the biomechanical context by taking in account the role of the muscles controlling the vocal folds.

Algorithms↗

Simple neural substrate predicts complex rhythmic structure in duetting birds.

Horneros (Furnarius Rufus) are South American birds well known for their oven-looking nests and their ability to sing in couples. Previous work has analyzed the rhythmic organization of the duets, unveiling a mathematical structure behind the songs. In this work we analyze in detail an extended database of duets. The rhythms of the songs are compatible with the dynamics presented by a wide class of dynamical systems: forced excitable systems. Compatible with this nonlinear rule, we build a biologically inspired model for how the neural and the anatomical elements may interact to produce the observed rhythmic patterns. This model allows us to synthesize songs presenting the acoustic and rhythmic features observed in real songs. We also make testable predictions in order to support our hypothesis.

Action Potentials↗