Echolocation click parameters and biosonar behaviour of the dwarf sperm whale (Kogia sima)

PhD and fieldwork funding were provided by the Danmarks Grundforskningsfond (27125 to P.T.M.), the Oticon Fonden (18-0340 to C.E.M.) the Dansk Akustisk Selskab (to C.E.M.), the South Africa National Research Foundation (research career advancement fellowship to S.E.) and the Claude Leon Foundation (postdoctoral fellowship to T.G.).Dwarf sperm whales (Kogia sima) are small toothed whales that produce narrow-band high-frequency (NBHF) echolocation clicks. Such NBHF clicks, subject to high levels of acoustic absorption, are usually produced by small, shallow-diving odontocetes, such as porpoises, in keeping with their short-range echolocation and fast click rates. Here, we sought to address the problem of how the little-studied and deep-diving Kogia can hunt with NBHF clicks in the deep sea. Specifically, we tested the hypotheses that Kogia produce NBHF clicks with longer inter-click intervals (ICIs), higher directionality and higher source levels (SLs) compared with other NBHF species. We did this by deploying an autonomous deep-water vertical hydrophone array in the Bahamas, where no other NBHF species are present, and by taking opportunistic recordings of a close-range Kogia sima in a South African harbour. Parameters from on-axis clicks (n=46) in the deep revealed very narrow-band clicks (root mean squared bandwidth, BWRMS, of 3±1 kHz), with SLs of up to 197 dB re. 1 µPa peak-to-peak (μPapp) at 1 m, and a half-power beamwidth of 8.8 deg. Their ICIs (mode of 245 ms) were much longer than those of porpoises (<100 ms), suggesting an inspection range that is longer than detection ranges of single prey, perhaps to facilitate auditory streaming of a complex echo scene. On-axis clicks in the shallow harbour (n=870) had ICIs and SLs in keeping with source parameters of other NBHF cetaceans. Thus, in the deep, dwarf sperm whales use a directional, but short-range echolocation system with moderate SLs, suggesting a reliable mesopelagic prey habitat.Publisher PDFPeer reviewe

Behavioral responses of satellite tracked Blainville's beaked whales (Mesoplodon densirostris) to mid-frequency active sonar

Funding support for tagging was provided by the US Navy's Office of Naval Research and Living Marine Resources program, the Chief of Naval Operations' Energy and Environmental Readiness Division and the NOAA Fisheries Ocean Acoustics Program. Trevor Joyce was supported by a National Research Council postdoctoral research associateship, hosted by NOAA's Southwest Fisheries Science Center.The vulnerability of beaked whales (Family: Ziphiidae) to intense sound exposure has led to interest in their behavioral responses to mid‐frequency active sonar (MFAS, 3–8 kHz). Here we present satellite‐transmitting tag movement and dive behavior records from Blainville's beaked whales (Mesoplodon densirostris) tagged in advance of naval sonar exercises at the Atlantic Undersea Test and Evaluation Center (AUTEC) in the Bahamas. This represents one of the largest samples of beaked whales individually tracked during sonar operations (n = 7). The majority of individuals (five of seven) were displaced 28–68 km after the onset of sonar exposure and returned to the AUTEC range 2–4 days after exercises ended. Modeled sound pressure received levels were available during the tracking of four individuals and three of those individuals showed declines from initial maxima of 145–172 dB re 1 μPa to maxima of 70–150 dB re 1 μPa following displacements. Dive behavior data from tags showed a continuation of deep diving activity consistent with foraging during MFAS exposure periods, but also suggested reductions in time spent on deep dives during initial exposure periods. These data provide new insights into behavioral responses to MFAS and have important implications for modeling the population consequences of disturbance.Publisher PDFPeer reviewe

Using individual-based bioenergetic models to predict the aggregate effects of disturbance on populations : a case study with beaked whales and Navy sonar

Funding: This research was supported by the Office of Naval Research (https://www.onr.navy.mil/) grant N0001419WX00431 and N000142012045: “Integrating information on displacement caused by mid-frequency active sonar and measurements of prey field into a population consequences of disturbance model for beaked whales” awarded to Dave Moretti, ND, SW, JH, LT, KB-B, AdR & VH. Funding support for tagging was provided by the US Navy's Office of Naval Research and Living Marine Resources program, the Chief of Naval Operations' Energy and Environmental Readiness Division and the NOAA Fisheries Ocean Acoustics Program.Anthropogenic activities can lead to changes in animal behavior. Predicting population consequences of these behavioral changes requires integrating short-term individual responses into models that forecast population dynamics across multiple generations. This is especially challenging for long-lived animals, because of the different time scales involved. Beaked whales are a group of deep-diving odontocete whales that respond behaviorally when exposed to military mid-frequency active sonar (MFAS), but the effect of these nonlethal responses on beaked whale populations is unknown. Population consequences of aggregate exposure to MFAS was assessed for two beaked whale populations that are regularly present on U.S. Navy training ranges where MFAS is frequently used. Our approach integrates a wide range of data sources, including telemetry data, information on spatial variation in habitat quality, passive acoustic data on the temporal pattern of sonar use and its relationship to beaked whale foraging activity, into an individual-based model with a dynamic bioenergetic module that governs individual life history. The predicted effect of disturbance from MFAS on population abundance ranged between population extinction to a slight increase in population abundance. These effects were driven by the interaction between the temporal pattern of MFAS use, baseline movement patterns, the spatial distribution of prey, the nature of beaked whale behavioral response to MFAS and the top-down impact of whale foraging on prey abundance. Based on these findings, we provide recommendations for monitoring of marine mammal populations and highlight key uncertainties to help guide future directions for assessing population impacts of nonlethal disturbance for these and other long-lived animals.Publisher PDFPeer reviewe

Beaked whales respond to simulated and actual navy sonar

This article is distributed under the terms of the Creative Commons Public Domain declaration. The definitive version was published in PLoS One 6 (2011): e17009, doi:10.1371/journal.pone.0017009.Beaked whales have mass stranded during some naval sonar exercises, but the cause is unknown. They are difficult to sight but can reliably be detected by listening for echolocation clicks produced during deep foraging dives. Listening for these clicks, we documented Blainville's beaked whales, Mesoplodon densirostris, in a naval underwater range where sonars are in regular use near Andros Island, Bahamas. An array of bottom-mounted hydrophones can detect beaked whales when they click anywhere within the range. We used two complementary methods to investigate behavioral responses of beaked whales to sonar: an opportunistic approach that monitored whale responses to multi-day naval exercises involving tactical mid-frequency sonars, and an experimental approach using playbacks of simulated sonar and control sounds to whales tagged with a device that records sound, movement, and orientation. Here we show that in both exposure conditions beaked whales stopped echolocating during deep foraging dives and moved away. During actual sonar exercises, beaked whales were primarily detected near the periphery of the range, on average 16 km away from the sonar transmissions. Once the exercise stopped, beaked whales gradually filled in the center of the range over 2–3 days. A satellite tagged whale moved outside the range during an exercise, returning over 2–3 days post-exercise. The experimental approach used tags to measure acoustic exposure and behavioral reactions of beaked whales to one controlled exposure each of simulated military sonar, killer whale calls, and band-limited noise. The beaked whales reacted to these three sound playbacks at sound pressure levels below 142 dB re 1 µPa by stopping echolocation followed by unusually long and slow ascents from their foraging dives. The combined results indicate similar disruption of foraging behavior and avoidance by beaked whales in the two different contexts, at exposures well below those used by regulators to define disturbance.The research reported here was financially supported by the United States (U.S.) Office of Naval Research (www.onr.navy.mil) Grants N00014-07-10988, N00014-07-11023, N00014-08-10990; the U.S. Strategic Environmental Research and Development Program (www.serdp.org) Grant SI-1539, the Environmental Readiness Division of the U.S. Navy (http://www.navy.mil/local/n45/), the U.S. Chief of Naval Operations Submarine Warfare Division (Undersea Surveillance), the U.S. National Oceanic and Atmospheric Administration (National Marine Fisheries Service, Office of Science and Technology) (http://www.st.nmfs.noaa.gov/), U.S. National Oceanic and Atmospheric Administration Ocean Acoustics Program (http://www.nmfs.noaa.gov/pr/acoustics/), and the Joint Industry Program on Sound and Marine Life of the International Association of Oil and Gas Producers (www.soundandmarinelife.org)

Biogeography in the deep : hierarchical population genomic structure of two beaked whale species

Funding for this research was provided by the Office of Naval Research, Award numbers N000141613017 and N000142112712. ABO was supported by a partial studentship from the University of St Andrews, School of Biology; OEG by the Marine Alliance for Science and Technology for Scotland (Scottish Funding Council grant HR09011); ELC by a Rutherford Discovery Fellowship from the Royal Society of New Zealand Te Aparangi; NAS by a Ramon y Cajal Fellowship from the Spanish Ministry of Innovation; MLM by the European Union’s Horizon 2020 Research and Innovation Programme (Marie Skłodowska-Curie grant 801199); CR by the Marine Institute (Cetaceans on the Frontier) and the Irish Research Council; and MTO by the Hartmann Foundation.The deep sea is the largest ecosystem on Earth, yet little is known about the processes driving patterns of genetic diversity in its inhabitants. Here, we investigated the macro- and microevolutionary processes shaping genomic population structure and diversity in two poorly understood, globally distributed, deep-sea predators: Cuvier’s beaked whale (Ziphius cavirostris) and Blainville’s beaked whale (Mesoplodon densirostris). We used double-digest restriction associated DNA (ddRAD) and whole mitochondrial genome (mitogenome) sequencing to characterise genetic patterns using phylogenetic trees, cluster analysis, isolation-by-distance, genetic diversity and differentiation statistics. Single nucleotide polymorphisms (SNPs; Blainville’s n = 43 samples, SNPs=13988; Cuvier’s n = 123, SNPs= 30479) and mitogenomes (Blainville’s n = 27; Cuvier’s n = 35) revealed substantial hierarchical structure at a global scale. Both species display significant genetic structure between the Atlantic, Indo-Pacific and in Cuvier’s, the Mediterranean Sea. Within major ocean basins, clear differentiation is found between genetic clusters on the east and west sides of the North Atlantic, and some distinct patterns of structure in the Indo-Pacific and Southern Hemisphere. We infer that macroevolutionary processes shaping patterns of genetic diversity include biogeographical barriers, highlighting the importance of such barriers even to highly mobile, deep-diving taxa. The barriers likely differ between the species due to their thermal tolerances and evolutionary histories. On a microevolutionary scale, it seems likely that the balance between resident populations displaying site fidelity, and transient individuals facilitating gene flow, shapes patterns of connectivity and genetic drift in beaked whales. Based on these results, we propose management units to facilitate improved conservation measures for these elusive species.Publisher PDFPeer reviewe

Biogeography in the deep: hierarchical population genomic structure of two beaked whale species

The deep sea is the largest ecosystem on Earth, yet little is known about the processes driving patterns of genetic diversity in its inhabitants. Here, we investigated the macro- and microevolutionary processes shaping genomic population structure and diversity in two poorly understood, globally distributed, deep-sea predators: Cuvier’s beaked whale (Ziphius cavirostris) and Blainville’s beaked whale (Mesoplodon densirostris). We used double-digest restriction associated DNA (ddRAD) and whole mitochondrial genome (mitogenome) sequencing to characterise genetic patterns using phylogenetic trees, cluster analysis, isolation-by-distance, genetic diversity and differentiation statistics. Single nucleotide polymorphisms (SNPs; Blainville’s n=43 samples, SNPs=13988; Cuvier’s n=123, SNPs= 30479) and mitogenomes (Blainville’s n=27; Cuvier’s n=35) revealed substantial hierarchical structure at a global scale. Both species display significant genetic structure between the Atlantic, Indo-Pacific and in Cuvier’s, the Mediterranean Sea. Within major ocean basins, clear differentiation is found between genetic clusters on the east and west sides of the North Atlantic, and some distinct patterns of structure in the Indo-Pacific and Southern Hemisphere. We infer that macroevolutionary processes shaping patterns of genetic diversity include biogeographical barriers, highlighting the importance of such barriers even to highly mobile, deep-diving taxa. The barriers likely differ between the species due to their thermal tolerances and evolutionary histories. On a microevolutionary scale, it seems likely that the balance between resident populations displaying site fidelity, and transient individuals facilitating gene flow, shapes patterns of connectivity and genetic drift. Based on these results, we propose management units to facilitate improved conservation measures for these elusive species

Population ecology of Blainville's beaked whales (Mesoplodon densirostris)

Quantifying population demographics is necessary to analyse the status of wildlife populations and to support effective conservation and management. Such a need exists for beaked whales which are vulnerable to anthropogenic noise, including navy sonar. Here, population demographics were estimated for Blainville’s beaked whales (Mesoplodon densirostris) in The Bahamas and the potential population-level effects of sonar investigated. Mark-recapture models were fitted to photo-identification data collected at the US Navy’s Atlantic Test and Evaluation Centre (AUTEC) where sonars were used regularly and 170 km away at Abaco where sonar use was limited, with the exception of a navy exercise correlated with a stranding of beaked whales in 2000. Life history data collected from 1997-2011 revealed that onset of sexual maturity occurred at age nine for both males and females and minimum longevities were 23 years. The annual turnover of individuals at Abaco was supported by the estimation of a larger parent population. However, adult females showed high site fidelity and survival, while adult males’ occupancy patterns were different, making survival of males difficult to separate from permanent emigration. Average annual abundance was lower at AUTEC when compared to a same-sized area at Abaco. Despite a similar number of adult females at both sites, a higher female:calf ratio was found at AUTEC, suggesting lower recruitment through births may have contributed to lower abundance. Population demographics in Abaco changed after the 2000 stranding; abundance and temporary emigration increased then returned to pre-2000 levels remaining stable thereafter. Two stranded whales re-floated in 2000 were later re-sighted having survived exposure to sonar and the physiological stresses related to stranding. This work provides evidence of a possible population-level effect of sonar use at a navy range and during a multi-ship exercise, emphasising the valuable role that longitudinal studies play in monitoring impacts of anthropogenic activities

Aquatic Mammals

Abstract Intraspeci c aggression constitutes a signi cant proportion of conspeci c behaviours among many mammal species. Agonistic interactions among male bottlenose dolphins (Tursiops truncatus) have been documented in several diVerent regions, including escalated aggression involving coalitions comprised of several males. However, despite many hours of direct observation most of these encounters are evidenced by minor injuries and often, physical wounds are not visible. Herein, we describe a unique encounter involving a long-term male alliance competing with a &apos;solo&apos; male that resulted in the temporary loss of consciousness of the lone male following repeated physical blows to his head region. This observation supports the increased tness experienced by males in alliances and illustrates the potential severity of aggressive interactions among adult bottlenose dolphins

Kinship as a basis for alliance formation between male bottlenose dolphins, Tursiops truncates, in the Bahamas

Alliances between reproductive males have been described in both cetacean and terrestrial mammal societies, and kin selection theory has often been invoked to explain these stable male coalitions. However, recent studies of bottlenose dolphins, Tursiops aduncus, in Australia suggest that mechanisms other than kinship are fundamental in the formation of male alliances. We supplemented skin biopsy sampling with the collection of faecal samples from individually recognized wild bottlenose dolphins, T. truncatus, to assess the role of kin selection in alliance membership in the northeast Bahamas. Stable alliances between pairs of males were identified based on association analyses of individual photoidentification data collected over 4 years. Molecular genetic analyses of tissue samples revealed highly significant correlations between patterns of association and both mitochondrial DNA haplotype identity and microsatellite relatedness, indicating that males within long-term alliances are more closely related than expected by chance. These data reinforce the high degree of plasticity previously reported within the genus Tursiops, and suggest that social and ecological differences between populations can markedly affect the role of kin selection in determining social alliances between male bottlenose dolphins