Remote measurement of the physiology, behaviour and energetic status of free-living animals is made possible by a variety of techniques that we refer to collectively as 'biotelemetry'. This set of tools ranges from transmitters that send their signals to receivers up to a few kilometers away to those that send data to orbiting satellites and, more frequently, to devices that log data. They enable researchers to document, for long uninterrupted periods, how undisturbed organisms interact with each other and their environment in real time. In spite of advances enabling the monitoring of many physiological and behavioural variables across a range of taxa of various sizes, these devices have yet to be embraced widely by the ecological community. Our review suggests that this technology has immense potential for research in basic and applied animal ecology. Efforts to incorporate biotelemetry into broader ecological research programs should yield novel information that has been challenging to collect historically from free-ranging animals in their natural environments. Examples of research that would benefit from biotelemetry include the assessment of animal responses to different anthropogenic perturbations and the development of life-time energy budgets.

/pdf/biotelemetry-a-mechanistic-approach-to-ecology-1pkho6xgij.pdf

Biotelemetry: a mechanistic approach to ecology.

(2008). MARINE MAMMAL NOISE-EXPOSURE CRITERIA: INITIAL SCIENTIFIC RECOMMENDATIONS. Bioacoustics: Vol. 17, No. 1-3, pp. 273-275.

Marine mammal noise-exposure criteria: initial scientific recommendations

1
Since the last thorough review of the effects of anthropogenic noise on cetaceans in 1995, a substantial number of research reports has been published and our ability to document response(s), or the lack thereof, has improved. While rigorous measurement of responses remains important, there is an increased need to interpret observed actions in the context of population-level consequences and acceptable exposure levels. There has been little change in the sources of noise, with the notable addition of noise from wind farms and novel acoustic deterrent and harassment devices (ADDs/AHDs). Overall, the noise sources of primary concern are ships, seismic exploration, sonars of all types and some AHDs.

2
Responses to noise fall into three main categories: behavioural, acoustic and physiological. We reviewed reports of the first two exhaustively, reviewing all peer-reviewed literature since 1995 with exceptions only for emerging subjects. Furthermore, we fully review only those studies for which received sound characteristics (amplitude and frequency) are reported, because interpreting what elicits responses or lack of responses is impossible without this exposure information. Behavioural responses include changes in surfacing, diving and heading patterns. Acoustic responses include changes in type or timing of vocalizations relative to the noise source. For physiological responses we address the issues of auditory threshold shifts and ‘stress’, albeit in a more limited capacity; a thorough review of physiological consequences is beyond the scope of this paper.

3
Overall, we found significant progress in the documentation of responses of cetaceans to various noise sources. However, we are concerned about the lack of investigation into the potential effects of prevalent noise sources such as commercial sonars, depth finders and fisheries acoustics gear. Furthermore, we were surprised at the number of experiments that failed to report any information about the sound exposure experienced by their experimental subjects. Conducting experiments with cetaceans is challenging and opportunities are limited, so use of the latter should be maximized and include rigorous measurements and or modelling of exposure.

Responses of cetaceans to anthropogenic noise

Foundations of modern auditory theory

Global increases in environmental noise levels – arising from expansion of human populations, transportation networks, and resource extraction – have catalysed a recent surge of research into the effects of noise on wildlife. Synthesising a coherent understanding of the biological consequences of noise from this literature is challenging. Taxonomic groups vary in auditory capabilities. A wide range of noise sources and exposure levels occur, and many kinds of biological responses have been observed, ranging from individual behaviours to changes in ecological communities. Also, noise is one of several environmental effects generated by human activities, so researchers must contend with potentially confounding explanations for biological responses. Nonetheless, it is clear that noise presents diverse threats to species and ecosystems and salient patterns are emerging to help inform future natural resource-management decisions. We conducted a systematic and standardised review of the scientific literature published from 1990 to 2013 on the effects of anthropogenic noise on wildlife, including both terrestrial and aquatic studies. Research to date has concentrated predominantly on European and North American species that rely on vocal communication, with approximately two-thirds of the data set focussing on songbirds and marine mammals. The majority of studies documented effects from noise, including altered vocal behaviour to mitigate masking, reduced abundance in noisy habitats, changes in vigilance and foraging behaviour, and impacts on individual fitness and the structure of ecological communities. This literature survey shows that terrestrial wildlife responses begin at noise levels of approximately 40 dBA, and 20% of papers documented impacts below 50 dBA. Our analysis highlights the utility of existing scientific information concerning the effects of anthropogenic noise on wildlife for predicting potential outcomes of noise exposure and implementing meaningful mitigation measures. Future research directions that would support more comprehensive predictions regarding the magnitude and severity of noise impacts include: broadening taxonomic and geographical scope, exploring interacting stressors, conducting larger-scale studies, testing mitigation approaches, standardising reporting of acoustic metrics, and assessing the biological response to noise-source removal or mitigation. The broad volume of existing information concerning the effects of anthropogenic noise on wildlife offers a valuable resource to assist scientists, industry, and natural-resource managers in predicting potential outcomes of noise exposure.

https://www.wildliferesearch.co.uk/Wildlife_Research/Publications_files/A%20synthesis%20of%20two%20decades%20of%20research%20documenting%20the%20effects%20of%20noise%20on%20wildlife%20Shannon.pdf

A synthesis of two decades of research documenting the effects of noise on wildlife

Traditionally, sperm whale clicks have been described as multipulsed, long duration, nondirectional signals of moderate intensity and with a spectrum peaking below 10 kHz. Such properties are counterindicative of a sonar function, and quite different from the properties of dolphin sonar clicks. Here, data are presented suggesting that the traditional view of sperm whale clicks is incomplete and derived from off-axis recordings of a highly directional source. A limited number of assumed on-axis clicks were recorded and found to be essentially monopulsed clicks, with durations of 100 micros, with a composite directionality index of 27 dB, with source levels up to 236 dB re: 1 microPa (rms), and with centroid frequencies of 15 kHz. Such clicks meet the requirements for long-range biosonar purposes. Data were obtained with a large-aperture, GPS-synchronized array in July 2000 in the Bleik Canyon off Vesteralen, Norway (69 degrees 28' N, 15 degrees 40' E). A total of 14 h of sound recordings was collected from five to ten independent, simultaneously operating recording units. The sound levels measured make sperm whale clicks by far the loudest of sounds recorded from any biological source. On-axis click properties support previous work proposing the nose of sperm whales to operate as a generator of sound.

/pdf/the-monopulsed-nature-of-sperm-whale-clicks-56fyifcszu.pdf

The monopulsed nature of sperm whale clicks

In sperm whales (Physeter catodon L. 1758) the nose is vastly hypertrophied, accounting for about one-third of the length or weight of an adult male. Norris and Harvey [in Animal Orientation and Navigation, NASA SP-262 (1972), pp. 397–417] ascribed a sound-generating function to this organ complex. A sound generator weighing upward of 10 tons and with a cross-section of 1 m is expected to generate high-intensity, directional sounds. This prediction from the Norris and Harvey theory is not supported by published data for sperm whale clicks (source levels of 180 dB re 1 μPa and little, if any, directionality). Either the theory is not borne out or the data is not representative for the capabilities of the sound-generating mechanism. To increase the amount of relevant data, a five-hydrophone array, suspended from three platforms separated by 1 km and linked by radio, was deployed at the slope of the continental shelf off Andenes, Norway, in the summers of 1997 and 1998. With this system, source levels up to 223 dB re 1 μPa peRMS were recorded. Also, source level differences of 35 dB for the same click at different directions were seen, which are interpreted as evidence for high directionality. This implicates sonar as a possible function of the clicks. Thus, previously published properties of sperm whale clicks underestimate the capabilities of the sound generator and therefore cannot falsify the Norris and Harvey theory.

Sperm whale clicks: Directionality and source level revisited

Delphinoids (Delphinidae, Odontoceti) produce tonal sounds and clicks by forcing pressurized air past phonic lips in the nasal complex. It has been proposed that homologous, hypertrophied nasal structures in the deep-diving sperm whale (Physeter macrocephalus) (Physeteridae, Odontoceti) are dedicated to the production of clicks. However, air volumes in diving mammals are reduced with increasing ambient pressure, which seems likely to influence pneumatic sound production at depth. To study sperm whale sound production at depth, we attached ultrasound time/depth-recording tags to sperm whales by means of a pole and suction cup. We demonstrate that sperm whale click production in terms of output and frequency content is unaffected by hydrostatic reduction in available air volume down to less than 2 % of the initial air volume in the nasal complex. We present evidence suggesting that the sound-generating mechanism has a bimodal function, allowing for the production of clicks suited for biosonar and clicks more suited for communication. Shared click features suggest that sound production in sperm whales is based on the same fundamental biomechanics as in smaller odontocetes and that the nasal complexes are therefore not only anatomically but also functionally homologous in generating the initial sound pulse.

/pdf/sperm-whale-sound-production-studied-with-ultrasound-time-4wdsyyaiqs.pdf

Sperm whale sound production studied with ultrasound time/depth-recording tags.

Sperm whales (Physeter macrocephalus) are deep-diving predators foraging in meso- and bathypelagic ecosystems off the continental shelves. To investigate the ecophysiological and communicative function of various click types from male sperm whales in a high-latitude habitat, we deployed a large-aperture array of calibrated hydrophones off northern Norway (N69, E15). Data show that sperm whales in this habitat produce three click types: usual clicks, creak clicks and, occasionally, slow clicks. Usual clicks and creak clicks exhibit short duration, profound directionality and a frequency content suited for echolocation on meso- and bathypelagic fish and squids. The acoustic properties and low repetition rate of usual clicks are suited for long-range echolocation, whereas creak clicks have properties equivalent to signals in buzzes, the terminating pulse trains known from echolocating bats during prey capture. From these source parameters and the high acoustic activity during foraging dives, it is concluded that echolocation is an important sensory cue in prey location. Sound pressure levels of creak clicks and usual clicks measured off the acoustic axis suggest that sperm whales may be subjected to eavesdropping by conspecifics, thereby conveying information about food aggregations at estimated ranges of 6 km for creak clicks and 16 km for usual clicks. Slow clicks exhibit low directionality, low-frequency emphasis and a reduced repetition rate, suggesting that this click type is more suited for communication than for echolocation. Slow clicks can be detected by submerged conspecifics at ranges up to 60 km. Thus, sperm whales producing slow clicks may represent an odontocete species that utilizes long-range sound communication.

Male sperm whale (Physeter macrocephalus) acoustics in a high-latitude habitat: implications for echolocation and communication

A linear error propagation analysis was applied to a hydrophone array used to locate sperm whales [see Mohl et al., J. Acoust. Soc. Am. 107, 638–648 (2000)]. The accuracy of two-dimensional (2D) and three-dimensional (3D) array configurations was investigated. The precision in source location was estimated as a function of inaccuracies in measurements of sound velocity, time-of-arrival differences (TOADs), and receiver positions. The magnitude of additional errors caused by geometric simplification was also assessed. The receiver position uncertainty had the largest impact on the precision of source location. A supplementary vertical linear array consisting of three receivers gave information on the vertical bearing and distance to the sound sources. The TOAD data from an additional receiver as well as from surface reflections were used to form an overdetermined location system. This system rendered positions within two standard deviations of the estimated errors from the original 3D array.

Bertel Møhl

Papers

The monopulsed nature of sperm whale clicks

Sperm whale clicks: Directionality and source level revisited

Sperm whale sound production studied with ultrasound time/depth-recording tags.

Male sperm whale (Physeter macrocephalus) acoustics in a high-latitude habitat: implications for echolocation and communication

Estimating source position accuracy of a large-aperture hydrophone array for bioacoustics