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  • 1
    Publikationsdatum: 2022-05-25
    Beschreibung: Author Posting. © The Author(s), 2018. This is the author's version of the work. It is posted here under a nonexclusive, irrevocable, paid-up, worldwide license granted to WHOI. It is made available for personal use, not for redistribution. The definitive version was published in Current Biology 28 (2018): 3878-3885.e3, doi:10.1016/j.cub.2018.10.037.
    Beschreibung: Toothed whales are apex predators varying in size from 40-kg porpoises to 50-ton sperm whales that all forage by emitting high-amplitude ultrasonic clicks and listening for weak returning echoes [1, 2]. The sensory field of view of these echolocating animals depends on the characteristics of the biosonar signals and the morphology of the sound generator, yet it is poorly understood how these biophysical relationships have shaped evolution of biosonar parameters as toothed whales adapted to different foraging niches. Here we test how biosonar output, frequency, and directivity vary with body size to understand the co-evolution of biosonar signals and sound-generating structures. We show that the radiated power increases twice as steeply with body mass (P ∝ M1.47±0.25) than expected from typical scaling laws of call intensity [3], indicating hyperallometric investment into sound production structures. This is likely driven by a strong selective pressure for long-range biosonar in larger oceanic or deep-diving species to search efficiently for patchy prey. We find that biosonar frequency scales inversely with body size (F∝ M-0.19±0.03), resulting in remarkably stable biosonar beamwidth that is independent of body size. We discuss how frequency scaling in toothed whales cannot be explained by the three main hypotheses for inverse scaling of frequency in animal communication [3-5]. We propose that a narrow acoustic field of view, analogous to the fovea of many visual predators, is the primary evolutionary driver of biosonar frequency in toothed whales, serving as a spatial filter to reduce clutter levels and facilitate long-range prey detection.
    Beschreibung: FHJ received support from a Carlsberg Foundation travel grant and an AIAS-COFUND fellowship from Aarhus Institute of Advanced Studies. ML was funded by a PhD stipend from the Faculty of Science and Technology, Aarhus University, and National Research Council grants to PTM. DMW was supported by the Danish National Research Foundation and Carlsberg Foundation grants to PTM. MJ was partly supported by an Aarhus University visiting professorship.
    Schlagwort(e): Echolocation ; Toothed whales ; Evolution ; Phylogenetic comparative methods ; Foraging ; Ecology ; Biosonar directivity ; Field of view ; Frequency scaling
    Repository-Name: Woods Hole Open Access Server
    Materialart: Preprint
    Standort Signatur Erwartet Verfügbarkeit
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  • 2
    Publikationsdatum: 2022-05-25
    Beschreibung: Author Posting. © The Company of Biologists, 2015. This article is posted here by permission of The Company of Biologists for personal use, not for redistribution. The definitive version was published in Journal of Experimental Biology 218 (2015): 3091-3101, doi:10.1242/jeb.120501.
    Beschreibung: Toothed whales produce echolocation clicks with source parameters related to body size; however, it may be equally important to consider the influence of habitat, as suggested by studies on echolocating bats. A few toothed whale species have fully adapted to river systems, where sonar operation is likely to result in higher clutter and reverberation levels than those experienced by most toothed whales at sea because of the shallow water and dense vegetation. To test the hypothesis that habitat shapes the evolution of toothed whale biosonar parameters by promoting simpler auditory scenes to interpret in acoustically complex habitats, echolocation clicks of wild Amazon river dolphins were recorded using a vertical seven-hydrophone array. We identified 404 on-axis biosonar clicks having a mean SLpp of 190.3±6.1 dB re. 1 µPa, mean SLEFD of 132.1±6.0 dB re. 1 µPa2s, mean Fc of 101.2±10.5 kHz, mean BWRMS of 29.3±4.3 kHz and mean ICI of 35.1±17.9 ms. Piston fit modelling resulted in an estimated half-power beamwidth of 10.2 deg (95% CI: 9.6–10.5 deg) and directivity index of 25.2 dB (95% CI: 24.9–25.7 dB). These results support the hypothesis that river-dwelling toothed whales operate their biosonars at lower amplitude and higher sampling rates than similar-sized marine species without sacrificing high directivity, in order to provide high update rates in acoustically complex habitats and simplify auditory scenes through reduced clutter and reverberation levels. We conclude that habitat, along with body size, is an important evolutionary driver of source parameters in toothed whale biosonars.
    Beschreibung: Field work was funded by Danish National Research Council grants to P.T.M., Associação Amigos do Peixe Boi da Amazônia (AMPA) and Petrobras Ambiental grants to V.M.F.d.S., Augustinus Fonden grants to M.L. and a travelling fellowship awarded to M.d.F. by Journal of Experimental Biology. M.L. was funded by a PhD stipend from the Faculty of Science and Technology, Aarhus University, and National Research Council grants to P.T.M. F.H.J. was funded by a Carlsberg Foundation travel grant.
    Beschreibung: 2016-10-07
    Schlagwort(e): Beamwidth ; Clutter ; Directionality ; Echolocation ; Habitat ; Toothed whale
    Repository-Name: Woods Hole Open Access Server
    Materialart: Article
    Format: application/pdf
    Standort Signatur Erwartet Verfügbarkeit
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  • 3
    Publikationsdatum: 2022-05-25
    Beschreibung: Author Posting. © The Author(s), 2018. This is the author's version of the work. It is posted here under a nonexclusive, irrevocable, paid-up, worldwide license granted to WHOI. It is made available for personal use, not for redistribution. The definitive version was published in Proceedings of the Royal Society B. Biological Sciences 285 (2018): 20181178, doi:10.1098/rspb.2018.1178.
    Beschreibung: The costs of predation may exert significant pressure on the mode of communication used by an animal, and many species balance the benefits of communication (e.g. mate attraction) against the potential risk of predation. Four groups of toothed whales have independently evolved narrowband high-frequency (NBHF) echolocation signals. These signals help NBHF species avoid predation through acoustic crypsis by echolocating and communicating at frequencies inaudible to predators such as mammal-eating killer whales. Heaviside’s dolphins (Cephalorhynchus heavisidii) are thought to exclusively produce NBHF echolocation clicks with a centroid frequency around 125 kHz and little to no energy below 100 kHz. To test this, we recorded wild Heaviside’s dolphins in a sheltered bay in Namibia. We demonstrate that Heaviside’s dolphins produce a second type of click with lower frequency and broader bandwidth in a frequency range that is audible to killer whales. These clicks are used in burst-pulses and occasional click series but not foraging buzzes. We evaluate three different hypotheses and conclude that the most likely benefit of these clicks is to decrease transmission directivity and increase conspecific communication range. The expected increase in active space depends on background noise but ranges from 2.5 (Wenz Sea State 6) to 5 times (Wenz Sea State 1) the active space of NBHF signals. This dual click strategy therefore allows these social dolphins to maintain acoustic crypsis during navigation and foraging, and to selectively relax their crypsis to facilitate communication with conspecifics.
    Beschreibung: This research was supported by a Fulbright U.S. Research Fellowship, the National Geographic Society’s Emerging Explorers Grant in conjunction with the Waitt Foundation (38115) and the University of Pretoria’s Zoology Department. TG was funded by the Claude Leon Foundation, and SE was funded by the South African National Research Foundation. FHJ acknowledges funding from the Office of Naval Research (N00014-1410410) and an AIAS-COFUND fellowship from Aarhus Institute of Advanced Studies.
    Schlagwort(e): Acoustic crypsis ; Active space ; Communication ; Echolocation ; Heaviside’s dolphin ; Narrowband high-frequency clicks
    Repository-Name: Woods Hole Open Access Server
    Materialart: Preprint
    Standort Signatur Erwartet Verfügbarkeit
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  • 4
    Publikationsdatum: 2022-05-26
    Beschreibung: Author Posting. © The Company of Biologists, 2015. This is the author's version of the work. It is posted here by permission of Company of Biologists for personal use, not for redistribution. The definitive version was published in Journal of Experimental Biology (2015), doi:10.1242/​jeb.116285.
    Beschreibung: Echolocating animals exercise an extensive control over the spectral and temporal properties of their biosonar signals to facilitate perception of their actively generated auditory scene when homing in on prey. The intensity and directionality of the biosonar beam defines the field of view of echolocating animals by affecting the acoustic detection range and angular coverage. However, the spatial relationship between an echolocating predator and its prey changes rapidly, resulting in different biosonar requirements throughout prey pursuit and capture. Here we measured single click beam patterns using a parametric fit procedure to test whether free-ranging Atlantic spotted dolphins (Stenella frontalis) modify their biosonar beamwidth. We recorded echolocation clicks using a linear array of receivers and estimated the beamwidth of individual clicks using a parametric spectral fit, cross-validated with well-established composite beam pattern estimates. The dolphins apparently increased the biosonar beamwidth, to a large degree without changing the signal frequency, when they approached the recording array. This is comparable to bats that also expand their field of view during prey capture, but achieve this by decreasing biosonar frequency. This behaviour may serve to decrease the risk that rapid escape movements of prey take them outside the biosonar beam of the predator. It is likely that shared sensory requirements have resulted in bats and toothed whales expanding their acoustic field of view at close range to increase the likelihood of successfully acquiring prey using echolocation, representing a case of convergent evolution of echolocation behaviour between these two taxa.
    Beschreibung: The study was funded by frame grants from the Danish Natural Science Foundation to PTM and MW, and by the National Oceanographic Partnership Programme via a research agreement between La Laguna University (NAS) and the Woods Hole Oceanographic Institution (MJ). FHJ was supported by the Danish Council for Independent Research | Natural Sciences, and is currently funded by a postdoctoral fellowship from the Carlsberg Foundation.
    Beschreibung: 2016-03-12
    Schlagwort(e): Echolocation ; Directionality ; Field of view ; Perception ; Dolphin ; Prey capture
    Repository-Name: Woods Hole Open Access Server
    Materialart: Preprint
    Format: application/pdf
    Standort Signatur Erwartet Verfügbarkeit
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  • 5
    Publikationsdatum: 2022-05-26
    Beschreibung: Author Posting. © Company of Biologists, 2019. This article is posted here by permission of Company of Biologists for personal use, not for redistribution. The definitive version was published in Journal of Experimental Biology 223 (2019): jeb.216283, doi: 10.1242/jeb.216283.
    Beschreibung: Toothed whales have evolved flexible biosonar systems to find, track and capture prey in diverse habitats. Delphinids, phocoenids and iniids adjust inter-click intervals and source levels gradually while approaching prey. In contrast, deep-diving beaked and sperm whales maintain relatively constant inter-click intervals and apparent output levels during the approach followed by a rapid transition into the foraging buzz, presumably to maintain a long-range acoustic scene in a multi-target environment. However, it remains unknown whether this rapid biosonar adjustment strategy is shared by delphinids foraging in deep waters. To test this, we investigated biosonar adjustments of a deep-diving delphinid, the Risso's dolphin (Grampus griseus). We analyzed inter-click interval and apparent output level adjustments recorded from sound recording tags to quantify in situ sensory adjustment during prey capture attempts. Risso's dolphins did not follow typical (20logR) biosonar adjustment patterns seen in shallow-water species, but instead maintained stable repetition rates and output levels up to the foraging buzz. Our results suggest that maintaining a long-range acoustic scene to exploit complex, multi-target prey layers is a common strategy amongst deep-diving toothed whales. Risso's dolphins transitioned rapidly into the foraging buzz just like beaked whales during most foraging attempts, but employed a more gradual biosonar adjustment in a subset (19%) of prey approaches. These were characterized by higher speeds and minimum specific acceleration, indicating higher prey capture efforts associated with evasive prey. Thus, tracking and capturing evasive prey using biosonar may require a more gradual switch between multi-target echolocation and single-target tracking.
    Beschreibung: F.H.J. was supported by an AIAS-COFUND fellowship from Aarhus Institute of Advanced Studies under the FP7 program of the EU Seventh Framework Programme (agreement no. 609033). F.V. was supported by the Office of Naval Research (ONR) (grants N00014-15-1-2341 and N00014-17-1-2715) and the Dutch Research Council (award number 016.Veni.181.086). P.L.T. was supported by ONR (grants N00014-15-1-2553 and N00014-18-1-2062) and acknowledges the support of the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) in the completion of this study. MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.
    Beschreibung: 2020-12-10
    Schlagwort(e): Echolocation ; Sensory ecology ; Mesopelagic foraging ; Deep-water environment ; Biosonar strategies ; Gain control
    Repository-Name: Woods Hole Open Access Server
    Materialart: Article
    Standort Signatur Erwartet Verfügbarkeit
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