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Directional aggressive pulse sounds in the bottlenose dolphin (Tursiops Truncatus): technical aspects and social implications
Linköping University, Department of Physics, Chemistry and Biology. Linköping University, The Institute of Technology.
2004 (English)Doctoral thesis, comprehensive summary (Other academic)
Abstract [en]

The ability of dolphins to produce pulse sounds with a bandwidth of up to 150 kHz, has been intensively studied with focus on their sonar capabilities, i.e. the detection, ranging and discrimination of objects of various shapes, textures and material. The sonar pulses have been found to be directional, with a -3db beam width of about 10 degrees in both the vertical and the horizontal plane, and with the beam axis ea. 5 degrees above that of the rostrum. Pulse sounds also frequently occur in social situations (Paper I), but studies of these sounds traditionally have been limited to their audio band, omni-directional components. Hypothesising that social pulse sounds in bottlenose dolphins (Tursiops truncatus) also were directional, early recordings within this study were done across a net barrier, in a narrow channel connecting two pools, to increase the probability to record such sounds with a fixed hydrophone. These recordings confirmed the presence of ultrasonic frequencies in pulse sounds emitted in aggressive interactions, where the interacting animals were oriented face-to-face on either side of the net barrier. There were also indications that they were directional (Paper I). These sounds also had a pronounced energy peak <20 kHz, considerably louder than sonar clicks, making them sound loud to the human ear. Pulse trains of low pulse repetition rate and long duration (> 1 sec), labelled "Machine-gun" sound, occurred frequently in the initial phase of the aggressive interactions. While emitting this sound, the animal kept its rostrum, and possibly the sound beam axis aimed at the agonist, suggesting that it was deliberately used to expose the opponent to powerful broadband pulse sounds, probably to provoke a fight (Paper III).

To avoid the restrictions on the behaviour imposed on the dolphins by the net barrier setup, an acoustic tag system was developed (Paper II). The tag was attached to the receiving dolphin instead of the transmitter, and made it possible to study the use of the directionality of the pulse sounds in social contexts, while the interacting dolphins were fully free-swimming. The tag, based on click detectors, recorded the envelope of pulses within two narrow band pass frequency filters, centred on 70kHz and 120kHz, respectively (Paper II). The tag recordings verified the presence of energy within both filter frequency bands in social pulse sounds, in particular in aggressive interactions (Paper III). This thesis suggests that thedirectionality of such pulse sounds is used intentionally to address social signals to a specific individual, or group of individuals. The relatively loud omni-directional audio band component will facilitate for nearby conspecifics to monitor the progress of the interaction. It will also make the signal clearly audible to the opponents throughout an interaction, even if they may be tumbling around, being unable to keep the sound beam axis aimed at the other. Similar pulse bursts were recorded in association with what was interpreted as play-fight, between subadults and juveniles (Paper IV). However, unlike the real aggressive bursts, these bursts had a trailing FM whistle, and this combined signal is suggested to be an appeasement signal, analogous with the laugh and giggle observed in apes. It is supposed to prevent play-fights from escalating into physical and potentially dangerous real fights. It was never recorded in true aggressive interactions among adults.

There were individual differences between the animals in amount of received directional pulse sounds while carrying the tag (Paper III). The socially most active of the females, having an intermediate position in the dominance hierarchy, received significantly more directional pulse sounds than the other, socially less active, but higher ranked female. The breeding male received the least number of pulse sounds. This lends support to the presence of "echolocation manners", as suggested by Johnson and Norris (1994), making it socially unacceptable to aiming the sonar beam towards conspecifics unless the intention is to conveying a social message.

Dolphins lack the facial plasticity playing a dominant role in the social life of many terrestrial mammals, e.g. primates, canids and felids. This directional signal modality is used to express internal emotions and can be addressed to selected individuals, further stressed by a direct glare. Dolphins completely lack such a facial plasticity but most likely not the need for expressing emotions. This thesis suggests that in dolphins the facial expressions may have been replaced bydirectional pulse sounds. The facial expressions in e.g., wolves, is supposed to be the result of a mixture of dominance/aggression vs. submission/fear emotions. We propose that the pulse sounds, seen in the aggressive interactions, reflect a similar mix of feelings. Obvious parameters in such a graded social signal system are pulse repetition rate, amplitude, duration and power spectrum. To study this further a broadband acoustic tag, preferably attached to several animals simultaneously, is required.

Place, publisher, year, edition, pages
Linköping: Linköpings universitet , 2004. , 28 p.
Series
Linköping Studies in Science and Technology. Dissertations, ISSN 0345-7524 ; 905
National Category
Natural Sciences
Identifiers
URN: urn:nbn:se:liu:diva-29360Local ID: 14683ISBN: 91-85295-70-1 (print)OAI: oai:DiVA.org:liu-29360DiVA: diva2:250172
Public defence
2004-12-17, Forsknings- och Utbildningscentrum, Kolmårdens Djurpark, 13:15 (Swedish)
Opponent
Available from: 2009-10-09 Created: 2009-10-09 Last updated: 2013-01-08
List of papers
1. High-frequency burst-pulse sounds in agonistic/aggresive interactions in bottlenose dolphins, Tursiops truncatus
Open this publication in new window or tab >>High-frequency burst-pulse sounds in agonistic/aggresive interactions in bottlenose dolphins, Tursiops truncatus
2004 (English)In: Echolocation in bats and dolphins / [ed] Jeanette Thomas, Cynthia Moss, and Marianne Vater, Chicago: The University of Chicago Press , 2004, 425-431 p.Chapter in book (Other academic)
Abstract [en]

Although bats and dolphins live in very different environments, are vastly different in size, and hunt different kinds of prey, both groups have evolved similar sonar systems, known as echolocation, to locate food and navigate the skies and seas. While much research has been conducted over the past 30 years on echolocation in bats and dolphins, this volume is the first to compare what is known about echolocation in each group, to point out what information is missing and to identity future areas of research. "Echolocation in Bats and Dolphins" consists of six sections: mechanisms of echolocation signal production; the anatomy and physiology of signal reception and interpretation; performance and cognition; ecological and evolutionary aspects of echolocation mammals; theoretical and methodological topics; and possible echolocation capability in other mammals, including shrews, seals and baleen whales. Animal behaviourists, ecologists, physiologists and both scientists and engineers who work in the field of bioacoustics should benefit from this book.

The objectives of this study were to investigate whether burst-pulse sounds emitted in aggressive interactions contain ultrasonic frequencies similar to the sonar sounds and to describe their repetition rate patterns and concurrent visual behavior patterns.

Place, publisher, year, edition, pages
Chicago: The University of Chicago Press, 2004
Keyword
Bottlenose dolphins, interactions, sounds
National Category
Natural Sciences
Identifiers
urn:nbn:se:liu:diva-29361 (URN)14684 (Local ID)0-226-79599-3 (ISBN)0-226-68446-6 (ISBN)978-0-2267-9599-7 (ISBN)14684 (Archive number)14684 (OAI)
Available from: 2009-10-09 Created: 2009-10-09 Last updated: 2013-10-21Bibliographically approved
2. An acoustic tag for recording, directional, pulsed ultrasounds aimed at free-swimming Bottlenose dolphins (Tursiops truncatus) by conspecifics
Open this publication in new window or tab >>An acoustic tag for recording, directional, pulsed ultrasounds aimed at free-swimming Bottlenose dolphins (Tursiops truncatus) by conspecifics
2004 (English)In: Aquatic Mammals, ISSN 0167-5427, E-ISSN 1996-7292, Vol. 30, no 3, 345-356 p.Article in journal (Refereed) Published
Abstract [en]

We developed an acoustic tag, called MOSART (MObile Submersible Acoustic Recorder of Transients), for recording directional social pulses produced by a bottlenose dolphin (Tursiops truncatus). The tag was attached to the dorsal fin of two dolphins by means of suction cups. Two adult bottlenose dolphins at the Kolmårdens Djurpark, Sweden, were trained to carry the tag comfortably through a desensitising program. The tag included two envelope click-detectors, each with a narrow bandpass filter, centred at 120 and 70 kHz, respectively. The duration of the original pulses and their relative amplitude within the two filter frequency bands was retained. The amplitude differences between the two filter bands reflected changes in the source frequency spectrum and/or the position of the tag hydrophone in the incoming sound beam. The tag recorded "echolocation click trains," "slow and irregular pulses," and "pulse bursts" with varying amounts of energy in both frequency bands. The peak amplitude and duration of clicks in "echolocation click trains" and in "slow and irregular pulses" were logged correctly; however, the tag recorder had more difficulties in handling the complex pulses in the aggressive "pulse bursts," where the duration of the individual pulses could not be determined. Still, the amplitude and the pulse repetition rate could be measured. The possible impact of the tag was investigated by analysing the dolphin's behaviours (12 categories), sounds (3 categories), preferred location in the pool, and respiration intervals. Only four of the behaviours and one preferred location in the pool showed significant differences among pre-tag baselines, tag periods, and post-tag follow-ups, suggesting that the tag had only a minor impact on the dolphin. We describe and discuss the tag and its capacity to record different pulsed sounds.

Keyword
acoustic tag, bottlenose dolphin, Tursiops truncatus, click detector, suction cup tag, directional pulse, broadband pulse burst, ultrasound, aggressive sounds
National Category
Natural Sciences
Identifiers
urn:nbn:se:liu:diva-30352 (URN)10.1578/AM.30.3.2004.345 (DOI)15895 (Local ID)15895 (Archive number)15895 (OAI)
Available from: 2009-10-09 Created: 2009-10-09 Last updated: 2017-12-13Bibliographically approved
3. Acoustic tag recordings of social pulse sounds in bottlenose dolphins (Tursiops truncatus) in human care during intra-specific aggressive interactions
Open this publication in new window or tab >>Acoustic tag recordings of social pulse sounds in bottlenose dolphins (Tursiops truncatus) in human care during intra-specific aggressive interactions
(English)Manuscript (preprint) (Other academic)
Abstract [en]

The tag recorded 549 regular click trains, 1056 slow and irregular pulse trains and 255 pulse bursts in 70 and 120 kHz band-pass filter bands. Pulse bursts occurred in high intensity aggressive situations, with animals face-to-face at close distance. Plotting peak repetition rate (PRR) against sound duration, two clusters were found, with a mean PRR of 642 pulses per second (pps) and 76 pps, respectively, and a mean sound duration of 110 msec and 224 msec, respectively. In 1085 audio band pulse sounds, recorded with a stationary hydrophone during 25 aggressive interactions between two females, also two PRR vs. sound duration clusters were found, with mean values 148 pps/203 msec, and 550 pps/196 msec, respectively. Also > 1 sec long pulse trains with <200 pps repetition rate were found, including "Machine-gun" sounds, which often initiated an aggressive interaction, when one animal aimed its rostrum at another from a distance. If the targeted animal turned towards the aggressor, they approached each other face-to-face, displaying "Open mouth", "Head & pectoral fin jerk" and "Jaw clap" behaviours, concurrent with "Pulse burst" emissions. We propose that these directional pulse sounds are analogous to the facial expressions seen in e.g. aggressive primates and canids, offering the dolphin the possibility to express underlying emotions, and to address aggressive signals to selected con-specifics. The strong, omni-directional audio band component allows others to monitor the progress of the aggressive interaction.

National Category
Natural Sciences
Identifiers
urn:nbn:se:liu:diva-86958 (URN)
Available from: 2013-01-08 Created: 2013-01-08 Last updated: 2013-01-08
4. An acoustic play-fight signal in bottlenose dolphins (Tursiops truncatus) in human care
Open this publication in new window or tab >>An acoustic play-fight signal in bottlenose dolphins (Tursiops truncatus) in human care
2005 (English)In: Aquatic Mammals, ISSN 0167-5427, E-ISSN 1996-7292, Vol. 31, no 2, 187-194 p.Article in journal (Refereed) Published
Abstract [en]

Play-fighting is common in many mammals, especially among juveniles and subadults, providing a safe opportunity to practice behaviours important in adult life. To prevent escalation into a potentially dangerous real fight, play-fighting often is accompanied by acoustic and/or visual appeasement behaviours. We studied aggressive and play-fight behaviours in bottlenose dolphins (Tursiops truncatus) at the Kolmården Djurpark. The results showed that play-fighting subadult dolphins emitted a characteristic sound, which was never observed in aggressive interactions. This was a short pulse burst followed by an FM-whistle. By plotting pulse repetition rate (PRR) vs. duration of the bursts, two main clusters were found. The bottom cluster had a mean PRR of 59 pulses per second (pps), and a mean duration of 154 msec. The top cluster had a mean PRR of 502 pps and a mean duration of 149 msec. These play-fight clusters were compared separately to corresponding adult aggressive pulse burst clusters. Taking both PRR and duration into consideration, no significant difference was found between the top clusters, or between the bottom clusters, in the two age groups. The trailing whistles were divided into five different frequency contour categories. These did not resemble the signature whistles of any of the play-fighting dolphins. The average start and end frequencies were 13.0 kHz and 10.1 kHz, respectively, and the maximum and minimum frequencies were 13.7 kHz and 7.0 kHz, respectively. The mean duration was 410 msec. Based on the fact that this sound occurred only in play-fights, we propose that it helps prevent a play-fight from escalating into a real fight and, hence, is analogous to the "laugh" and "chuckle" seen in apes.

Keyword
Bottlenose dolphins, acoustic play-fight signal
National Category
Natural Sciences
Identifiers
urn:nbn:se:liu:diva-30351 (URN)10.1578/AM.31.2.2005.187 (DOI)15893 (Local ID)15893 (Archive number)15893 (OAI)
Available from: 2009-10-09 Created: 2009-10-09 Last updated: 2017-12-13
5. Pulsed sounds recorded with an acoustic tag on a free-ranging bottlenose dolphin (Tursiops truncatus) in the coastal waters of New Jersey, USA
Open this publication in new window or tab >>Pulsed sounds recorded with an acoustic tag on a free-ranging bottlenose dolphin (Tursiops truncatus) in the coastal waters of New Jersey, USA
(English)Manuscript (preprint) (Other academic)
Abstract [en]

No abstract available.

National Category
Natural Sciences
Identifiers
urn:nbn:se:liu:diva-86959 (URN)
Available from: 2013-01-08 Created: 2013-01-08 Last updated: 2013-01-08

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