A path reconstruction method integrating dead-reckoning and position fixes applied to humpback whales

PW received a PhD studentship with matched funding from The Netherlands Ministry of Defence (administered by TNO) and the UK Natural Environment Research Council (NE/J500276/1). The 3S2 project was funded by the US Office of Naval Research (N00014-10-1-0355), the Norwegian Ministry of Defence, and The Netherlands Ministry of Defence. Part of this work was supported by the Multi-study Ocean acoustics Human effects Analysis (MOCHA) project funded by the US Office of Naval Research (N00014-12-1-0204).BACKGROUND: Detailed information about animal location and movement is often crucial in studies of natural behaviour and how animals respond to anthropogenic activities. Dead-reckoning can be used to infer such detailed information, but without additional positional data this method results in uncertainty that grows with time. Combining dead-reckoning with new Fastloc-GPS technology should provide good opportunities for reconstructing georeferenced fine-scale tracks, and should be particularly useful for marine animals that spend most of their time under water. We developed a computationally efficient, Bayesian state-space modelling technique to estimate humpback whale locations through time, integrating dead-reckoning using on-animal sensors with measurements of whale locations using on-animal Fastloc-GPS and visual observations. Positional observation models were based upon error measurements made during calibrations. RESULTS: High-resolution 3-dimensional movement tracks were produced for 13 whales using a simple process model in which errors caused by water current movements, non-location sensor errors, and other dead-reckoning errors were accumulated into a combined error term. Positional uncertainty quantified by the track reconstruction model was much greater for tracks with visual positions and few or no GPS positions, indicating a strong benefit to using Fastloc-GPS for track reconstruction. Compared to tracks derived only from position fixes, the inclusion of dead-reckoning data greatly improved the level of detail in the reconstructed tracks of humpback whales. Using cross-validation, a clear improvement in the predictability of out-of-set Fastloc-GPS data was observed compared to more conventional track reconstruction methods. Fastloc-GPS observation errors during calibrations were found to vary by number of GPS satellites received and by orthogonal dimension analysed; visual observation errors varied most by distance to the whale. CONCLUSIONS: By systematically accounting for the observation errors in the position fixes, our model provides a quantitative estimate of location uncertainty that can be appropriately incorporated into analyses of animal movement. This generic method has potential application for a wide range of marine animal species and data recording systems.Publisher PDFPeer reviewe

Vocal foragers and silent crowds: context-dependent vocal variation in Northeast Atlantic long-finned pilot whales

13 pages, 6 figures, 3 tables.-- Fleur Visser ... et al.-- This article is an open access publicationVocalisations form a key component of the social interactions and foraging behaviour of toothed whales. We investigated changes in calling and echolocation behaviour of long-finned pilot whales between foraging and non-foraging periods, by combining acoustic recordings and diving depth data from tagged individuals with concurrent surface observations on social behaviour of their group. The pilot whales showed marked vocal variation, specific to foraging and social context. During periods of foraging, pilot whales showed more vocal activity than during non-foraging periods (rest, travel). In addition to the expected increase in echolocation activity, call rates also increased, suggesting that pilot whales communicate more during foraging. Furthermore, calls with multiple inflections occurred more often immediately before and after foraging dives and during the early descent and late ascent phases of foraging dives. However, these calls were almost never detected at diving depths of the tagged whale beyond 350 m. Calls with no or few inflections were produced at all times, irrespective of diving depth of the tagged whale. We discuss possible explanations for the distinct vocal variation associated with foraging periods. In addition, during non-foraging periods, the pilot whales were found to be more silent (no calling or echolocation) in larger, more closely spaced groups. This indicates that increased levels of social cohesion may release the need to stay in touch acoustically.This study was financially supported by the US Office of Naval Research, The Netherlands Ministry of Defence, the Norwegian Research Council and the Norwegian Ministry of DefencePeer reviewe

Experimental field studies to measure behavioral responses of cetaceans to sonar

Funding was provided by a variety of military and governmental funding sources from several nations acknowledged within referenced publications, notably the US Office of Naval Research, US Navy Living Marine Resources Program, and the navies of the USA, Norway, and the Netherlands. P.L.T. 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.Substantial recent progress has been made in directly measuring behavioral responses of free-ranging marine mammals to sound using controlled exposure experiments. Many studies were motivated by concerns about observed and potential negative effects of military sonar, including stranding events. Well-established experimental methods and increasingly sophisticated technologies have enabled fine-resolution measurement of many aspects of baseline behavior and responses to sonar. Studies have considered increasingly diverse taxa, but primarily odontocete and mysticete cetaceans that are endangered, particularly sensitive, or frequently exposed to sonar. This review focuses on recent field experiments studying cetacean responses to simulated or actual active military sonars in the 1 to 8 kHz band. Overall results demonstrate that some individuals of different species display clear yet varied responses, some of which have negative implications, while others appear to tolerate relatively high levels, although such exposures may have other consequences not measured. Responses were highly variable and may not be fully predictable with simple acoustic exposure metrics (e.g. received sound level). Rather, differences among species and individuals along with contextual aspects of exposure (e.g. behavioral state) appear to affect response probability. These controlled experiments provide critically needed documentation of identified behavioral responses occurring upon known sonar exposures, and they directly inform regulatory assessments of potential effects. They also inform more targeted opportunistic monitoring of potential responses of animals during sonar operations and have stimulated adaptations of field methods to consider increasingly realistic exposure scenarios and how contextual factors such as behavioral state and source proximity influence response type and probability.Publisher PDFPeer reviewe

Gait switches in deep-diving beaked whales : biomechanical strategies for long-duration dives

Work in the Canary Islands was funded by the Office of Naval Research and the National Oceanographic Partnership Program (NOPP), by a consortium consisting of the Canary Islands Government, the Spanish Ministry of Environment and the Spanish Ministry of Defense and by the European environmental funding LIFE-INDEMARES program for the inventory and designation of the Natura 2000 network in marine areas of the Spanish territory, headed by Fundacion Biodiversidad, with additional support from the Cabildo Insular of El Hierro. Fieldwork in the Ligurian Sea was funded by NOPP. Research in the Gully was funded by the Strategic Environmental Research and Development Program (SERDP) programs RC-2113 and RC-2337. L.M.M.L. was funded by a ‘la Caixa’ Fellowship within the 2010 UK Framework Programme. N.A.d.S. was funded for this study by ONR and by the EU FP7 Marie Curie project SOUNDMAR. M.J. was funded by a Marie Curie Career Integration Grant, and from the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) and their support is gratefully acknowledged. MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.Diving animals modulate their swimming gaits to promote locomotor efficiency and so enable longer, more productive dives. Beaked whales perform extremely long and deep foraging dives that probably exceed aerobic capacities for some species. Here, we use biomechanical data from suction-cup tags attached to three species of beaked whales (Mesoplodon densirostris, N = 10; Ziphius cavirostris, N = 9; and Hyperoodon ampullatus, N = 2) to characterize their swimming gaits. In addition to continuous stroking and stroke and-glide gaits described for other diving mammals, all whales produced occasional fluke-strokes with distinctly larger dorsoventral acceleration, which we termed 'type-B' strokes. These high-power strokes occurred almost exclusively during deep dive ascents as part of a novel mixed gait. To quantify body rotations and specific acceleration generated during strokes we adapted a kinematic method combining data from two sensors in the tag. Body rotations estimated with high-rate magnetometer data were subtracted from accelerometer data to estimate the resulting surge and heave accelerations. Using this method, we show that stroke duration, rotation angle and acceleration were bi-modal for these species, with B-strokes having 76% of the duration, 52% larger body rotation and four times more surge than normal strokes. The additional acceleration of B-strokes did not lead to faster ascents, but rather enabled brief glides, which may improve the overall efficiency of this gait. Their occurrence towards the end of long dives leads us to propose that B-strokes may recruit fast-twitch fibres that comprise similar to 80% of swimming muscles in Blainville's beaked whales, thus prolonging foraging time at depth.PostprintPeer reviewe

Call combination patterns in Icelandic killer whales (Orcinus orca)

Funding: Funding for data collection was provided by the Fundação para a Ciência e a Tecnologia (grant number SFRH/BD/30303/2006), the Icelandic Research Fund (grant numbers 120248042 and 217519), the National Geographic Global Exploration Fund (grant number GEFNE65-12), and a Russell Trust Award from the University of St. Andrews to FIPS. This project was funded in part by the generous support of Earthwatch. Additionally, funding was provided by the US Office of Naval Research (grant number N00014-08-1-0984), US Living Marine Resources (project 57), UK Defence Science and Technology Laboratory, and French Direction Générale de l’Armement to PJOM. A RANNÍS Infrastructure Fund (grant number 200229) provided funding to JS and PJW for CATS tags and tracking equipment. AS was supported by Doctoral Student Grants (grant number 206808 and 239641) from the Icelandic Research Fund.Acoustic sequences have been described in a range of species and in varying complexity. Cetaceans are known to produce complex song displays but these are generally limited to mysticetes; little is known about call combinations in odontocetes. Here we investigate call combinations produced by killer whales (Orcinus orca), a highly social and vocal species. Using acoustic recordings from 22 multisensor tags, we use a first order Markov model to show that transitions between call types or subtypes were significantly different from random, with repetitions and specific call combinations occurring more often than expected by chance. The mixed call combinations were composed of two or three calls and were part of three call combination clusters. Call combinations were recorded over several years, from different individuals, and several social clusters. The most common call combination cluster consisted of six call (sub-)types. Although different combinations were generated, there were clear rules regarding which were the first and last call types produced, and combinations were highly stereotyped. Two of the three call combination clusters were produced outside of feeding contexts, but their function remains unclear and further research is required to determine possible functions and whether these combinations could be behaviour- or group-specific.Publisher PDFPeer reviewe

Behavioral responses to predatory sounds predict sensitivity of cetaceans to anthropogenic noise within a soundscape of fear

Funding was provided by the US Navy Living Marine Resources and Office of Naval Research programs, Netherlands Ministry of Defence, Norwegian Ministry of Defence, UK Defence Science and Technology Laboratory, and DGA French Ministry of Defence.Acoustic signals travel efficiently in the marine environment, allowing soniferous predators and prey to eavesdrop on each other. Our results with four cetacean species indicate that they use acoustic information to assess predation risk and have evolved mechanisms to reduce predation risk by ceasing foraging. Species that more readily gave up foraging in response to predatory sounds of killer whales also decreased foraging more during 1- to 4-kHz sonar exposures, indicating that species exhibiting costly antipredator responses also have stronger behavioral reactions to anthropogenic noise. This advance in our understanding of the drivers of disturbance helps us to predict what species and habitats are likely to be most severely impacted by underwater noise pollution in oceans undergoing increasing anthropogenic activities. As human activities impact virtually every animal habitat on the planet, identifying species at-risk from disturbance is a priority. Cetaceans are an example taxon where responsiveness to anthropogenic noise can be severe but highly species and context specific, with source–receiver characteristics such as hearing sensitivity only partially explaining this variability. Here, we predicted that ecoevolutionary factors that increase species responsiveness to predation risk also increase responsiveness to anthropogenic noise. We found that reductions in intense-foraging time during exposure to 1- to 4-kHz naval sonar and predatory killer whale sounds were highly correlated (r = 0.92) across four cetacean species. Northern bottlenose whales ceased foraging completely during killer whale and sonar exposures, followed by humpback, long-finned pilot, and sperm whales, which reduced intense foraging by 48 to 97level responses to killer whale playbacks, implying a similar level of perceived risk. The correlation cannot be solely explained by hearing sensitivity, indicating that species- and context-specific antipredator adaptations also shape cetacean responses to human-made noise. Species that are more responsive to predator presence are predicted to be more disturbance sensitive, implying a looming double whammy for Arctic cetaceans facing increased anthropogenic and predator activity with reduced ice cover.Publisher PDFPeer reviewe

Disturbance-specific social responses in long-finned pilot whales, Globicephala melas

The study was funded by the US Office of Naval Research, The Netherlands Ministry of Defence, Norwegian Ministry of Defence and French Ministry of Defence. F.V., C.C., P.K., F.P.L. and P.M. were supported by one or two of these funders. P.T. received funding from the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland). MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.Social interactions among animals can influence their response to disturbance. We investigated responses of long-finned pilot whales to killer whale sound playbacks and two anthropogenic sources of disturbance: Tagging effort and naval sonar exposure. The acoustic scene and diving behaviour of tagged individuals were recorded along with the social behaviour of their groups. All three disturbance types resulted in larger group sizes, increasing social cohesion during disturbance. However, the nature and magnitude of other responses differed between disturbance types. Tagging effort resulted in a clear increase in synchrony and a tendency to reduce surface logging and to become silent (21% of cases), whereas pilot whales increased surface resting during sonar exposure. Killer whale sounds elicited increased calling rates and the aggregation of multiple groups, which approached the sound source together. This behaviour appears to represent a mobbing response, a likely adaptive social defence against predators or competitors. All observed response-Tactics would reduce risk of loss of group coordination, suggesting that, in social pilot whales, this could drive behavioural responses to disturbance. However, the behavioural means used to achieve social coordination depends upon other considerations, which are disturbance-specific.Publisher PDFPeer reviewe

Separating underwater ambient noise from flow noise recorded on stereo acoustic tags attached to marine mammals

A.M.v.B.B. and P.B. were funded by The Netherlands Ministry of Defence. Fieldwork efforts and support for P.M. and F.S. was provided by the US Office of Naval Research [award numbers N00014-08-1-0984 and N00014-10-1-0355]. P.W. received a PhD studentship with matched funding from The Netherlands Ministry of Defence (administered by The Netherlands Organisation for Applied Scientific Research, TNO) and UK Natural Environment Research Council [NE/J500276/1].Sound-recording acoustic tags attached to marine animals are commonly used in behavioural studies. Measuring ambient noise is of interest to efforts to understand responses of marine mammals to anthropogenic underwater sound, or to assess their communication space. Noise of water flowing around the tag reflects the speed of the animal, but hinders ambient noise measurement. Here, we describe a correlation-based method for stereo acoustic tags to separate the relative contributions of flow and ambient noise. The uncorrelated part of the noise measured in digital acoustic recording tag (DTAG) recordings related well to swim speed of a humpback whale (Megaptera novaeangliae), thus providing a robust measure of flow noise over a wide frequency bandwidth. By removing measurements affected by flow noise, consistent ambient noise estimates were made for two killer whales (Orcinus orca) with DTAGs attached simultaneously. The method is applicable to any multi-channel acoustic tag, enabling application to a wide range of marine species.Publisher PDFPeer reviewe

Killer whales (Orcinus orca) produce ultrasonic whistles

This study reports that killer whales, the largest dolphin, produce whistles with the highest fundamental frequencies ever reported in a delphinid. Using wide-band acoustic sampling from both animal-attached (Dtag) and remotely deployed hydrophone arrays, ultrasonic whistles were detected in three Northeast Atlantic populations but not in two Northeast Pacific populations. These results are inconsistent with analyses suggesting a correlation of maximum frequency of whistles with body size in delphinids, indicate substantial intraspecific variation in whistle production in killer whales, and highlight the importance of appropriate acoustic sampling techniques when conducting comparative analyses of sound repertoires

Predictive model of sperm whale prey capture attempts from time-depth data

Funding: Research was supported by the Portuguese Science & Technology Foundation (FCT), the Azorean Science & Technology Fund (FRCT), and the EU through research projects WATCH IT-Acores-01-0145-FEDER-000057, FCT-IF/00943/2013/CP1199/CT0001, META-FA_06_2017_017, and SUMMER-H2020 GA 817806, co-funded by FEDER, COMPETE, QREN, POPH, ESF, PO AZORES 2020, Portuguese Ministry for Science and Education, and individual contracts/grants to CO (WATCH IT-Acores-01-0145-FEDER-000057, 3/SRMCT/DRAM/2019 under RAGES-SUB/ENV.C.2-GA 110661, and INTERTAGUA-MAC2/1.1a/385), SPJ (SUMMER-H2020 GA 817806), IC (FCT-IP Project UIDP/05634/2020). PJW is funded by RANNÍS Icelandic Research Fund grant 207081. RP and MAS are co-financed by AZORES2020, through the EU Fund 01-0145-FEDER-000140 “MarAZ Researchers: Consolidate a body of researchers in Marine Sciences in the Azores”. Okeanos is funded by FCT (UIDB/05634/2020) and by the Regional Government of the Azores through the initiative to support the Research Centers of the University of the Azores (M1.1.A/REEQ.CIENTÍFICO UI&D/2021/010).Background High-resolution sound and movement recording tags offer unprecedented insights into the fine-scale foraging behaviour of cetaceans, especially echolocating odontocetes, enabling the estimation of a series of foraging metrics. However, these tags are expensive, making them inaccessible to most researchers. Time-Depth Recorders (TDRs), which have been widely used to study diving and foraging behaviour of marine mammals, offer a more affordable alternative. Unfortunately, data collected by TDRs are bi-dimensional (time and depth only), so quantifying foraging effort from those data is challenging. Methods A predictive model of the foraging effort of sperm whales (Physeter macrocephalus) was developed to identify prey capture attempts (PCAs) from time-depth data. Data from high-resolution acoustic and movement recording tags deployed on 12 sperm whales were downsampled to 1 Hz to match the typical TDR sampling resolution and used to predict the number of buzzes (i.e., rapid series of echolocation clicks indicative of PCAs). Generalized linear mixed models were built for dive segments of different durations (30, 60, 180 and 300 s) using multiple dive metrics as potential predictors of PCAs. Results Average depth, variance of depth and variance of vertical velocity were the best predictors of the number of buzzes. Sensitivity analysis showed that models with segments of 180 s had the best overall predictive performance, with a good area under the curve value (0.78 ± 0.05), high sensitivity (0.93 ± 0.06) and high specificity (0.64 ± 0.14). Models using 180 s segments had a small difference between observed and predicted number of buzzes per dive, with a median of 4 buzzes, representing a difference in predicted buzzes of 30%. Conclusions These results demonstrate that it is possible to obtain a fine-scale, accurate index of sperm whale PCAs from time-depth data alone. This work helps leveraging the potential of time-depth data for studying the foraging ecology of sperm whales and the possibility of applying this approach to a wide range of echolocating cetaceans. The development of accurate foraging indices from low-cost, easily accessible TDR data would contribute to democratize this type of research, promote long-term studies of various species in several locations, and enable analyses of historical datasets to investigate changes in cetacean foraging activity.Publisher PDFPeer reviewe