Papers by Petter Kvadsheim
ABSTRACT LONG-TERM GOALS The DTAG was originally developed to record an animal's response... more ABSTRACT LONG-TERM GOALS The DTAG was originally developed to record an animal's responses to acoustic stimuli, such as naval sonar signals, and the traditional method for deploying these non invasive (suction cup) tags on large and mid-sized cetaceans has been to use long carbon fiber poles (Moore et al. 2001, Johnson and Tyack 2003). This technique works well with some of the large whales, but some mid-sized cetaceans (e.g., beaked whales, minke whales) tend to be quicker, more maneuverable, and elusive, making pole tagging rather inefficient (Johnson and Tyack 2003). The goal of this project was to develop a system that launches the DTAG through the air using a pneumatic launcher (Aerial Rocket Tag System, or ARTS), in order to extend the tagging range and thus increase tag deployment rate. A preliminary version of the ARTS-DTAG system has already proven to have potential in improving tagging efficiency, particularly with "difficult" whale species (Kvadsheim et al. 2009). However, this system needed improvement in several areas to become operational. This project included redesign of the ARTS-DTAG system compared to the 2009 version and an extensive test program, including ballistic testing in the lab, at sea testing on a floating dummy whale and a field trial with tag deployments on minke and pilot whales. Public reporting burden for the collection of information is estimated to average 1 hour per response, including the time for reviewing instructions, searching existing data sources, gathering and maintaining the data needed, and completing and reviewing the collection of information. Send comments regarding this burden estimate or any other aspect of this collection of information, including suggestions for reducing this burden, to Washington Headquarters Services, Directorate for Information Operations and Reports, 1215 Jefferson Davis Highway, Suite 1204, Arlington VA 22202-4302. Respondents should be aware that notwithstanding any other provision of law, no person shall be subject to a penalty for failing to comply with a collection of information if it does not display a currently valid OMB control number.
Ecological applications : a publication of the Ecological Society of America, 2016
The time and energetic costs of behavioral responses to incidental and experimental sonar exposur... more The time and energetic costs of behavioral responses to incidental and experimental sonar exposures, as well as control stimuli, were quantified using hidden state analysis of time series of acoustic and movement data recorded by tags (DTAG) attached to 12 sperm whales (Physeter macrocephalus) using suction cups. Behavioral state transition modeling showed that tagged whales switched to a non-foraging, non-resting state during both experimental transmissions of low-frequency active sonar from an approaching vessel (LFAS; 1-2 kHz, source level 214 dB re 1 µPa m, four tag records) and playbacks of potential predator (killer whale, Orcinus orca) sounds broadcast at naturally occurring sound levels as a positive control from a drifting boat (five tag records). Time spent in foraging states and the probability of prey capture attempts were reduced during these two types of exposures with little change in overall locomotion activity, suggesting an effect on energy intake with no immediate...
Acta Physiologica Scandinavica, Sep 30, 1997
The trunk of marine mammals is encased in a blubber layer which provides thermal insulation that ... more The trunk of marine mammals is encased in a blubber layer which provides thermal insulation that can be changed by circulatory adjustments. The extremities, on the other hand, are poorly insulated but have vascular arrangements constructed for prevention or promotion of heat loss, depending on the thermal state of the animal. We have studied the importance of different body parts as sites for heat dissipation and also assessed the effect of circulatory adjustments on heat transfer through blubber, by combining direct measurements of heat flux from the flippers and trunk with simultaneous recordings of temperature gradients through the blubber and metabolic rates of harp seals (Phoca groenlandica) subjected to water temperatures between 1 and 24 °C. We also determined the thermal conductivity of blubber samples from the same animals after death, and compared this with the insulative properties of live blubber. At the lowest water temperatures, the insulative properties of live blubber were similar to those of dead blubber, and heat loss from the ¯ippers only accounted for 2±6% of the metabolic heat production. As heat load increased with increasing water temperatures, the fraction of heat lost from the ¯ippers increased, to 19±48% at 24 °C, while the fraction lost from the trunk decreased, despite an increase in the convective (circulatory) heat transfer through the blubber layer.
Marine Mammal Science, 2002
In phocid seals, blubber serves as the main thermal insulation instead of fur. The thermal functi... more In phocid seals, blubber serves as the main thermal insulation instead of fur. The thermal function of fur, at least in adult phocid seals, has therefore been questioned. We measured the relative contribution of fur to the combined thermal resistance (insulation) offered by blubber, skin, and fur in newborn and adult harp (Pagophilus gvoenlandiczls) and hooded (Cystophora cristutu) seals, in air and water, to elucidate the role of fur as insulation in phocid seals. In air the fur contributed 70% of the combined thermal resistance of blubber, skin, and fur in newborn harp seal pups and 27% in adults, whereas in hooded seals the fur contributed 73% in newborn pups and 34% in adults. When submerged the thermal resistance of the fur was reduced by 84%-92%, and contributed 65% to the total insulation in newborn harp seal pups and 3% in adults, and 26% in newborn hooded seal pups and 5% in adults. We conclude that in air the fur of phocid seals makes an important contribution to the insulation of pups, and also contributes considerably to the insulation of adult animals. In water, even though the thermal resistance of the fur is dramatically reduced, the fur still contributes substantially to the insulation of pups, but its contribution in adults is negligible.
Journal of Theoretical Biology, 2000
Heat flux models have been used to predict metabolic rates of marine mammals, generally by estima... more Heat flux models have been used to predict metabolic rates of marine mammals, generally by estimating conductive heat transfer through their blubber layer. Recently, Kvadsheim et al. (1997) found that such models tend to overestimate metabolic rates, and that such errors probably result from the asymmetrical distribution of blubber. This problem may be avoided if reliable estimates of heat flux through the skin of the animals are obtained by using models that combine calculations of conductive heat flux through the skin and fur, and convective heat flux from the surface of the animal to the environment. We evaluated this approach based on simultaneous measurements of metabolic rates and of input parameters necessary for heat flux calculations, as obtained from four harp seals (Phoca groenlandica) resting in cold water. Heat flux estimates were made using two free convection models (double-flat-plate and cylindrical geometry) and one forced convection model (single-flat-plate geometry). We found that heat flux estimates generally underestimated metabolic rates, on average by 26-58%, and that small variations in input parameters caused large variations in these estimates. We conclude that cutaneous heat flux models are too inaccurate and sensitive to small errors in input parameters to provide reliable estimates of metabolic rates of marine mammals.
ICES Journal of Marine Science, 2013
In this study, retrospective data on naval sonar activity and prey abundance were correlated with... more In this study, retrospective data on naval sonar activity and prey abundance were correlated with killer whale sightings within a fjord basin in northern Norway. In addition, passive acoustic and visual marine mammal surveys were conducted before, during, and after a specific navy exercise in 2006. Herring abundance was the main factor affecting killer whale presence. Naval sonar, either operational navy sonar exercises (Flotex) or experimental sonar activity (CEE) alone, did not explain killer whale occurrence. However, naval sonar activity during a period of low prey availability seemed to have had a negative effect on killer whale presence. We conclude that the level of reaction to sonar can be influenced by multiple factors, including availability of prey.
American Institute of Physics Conference Series, Nov 18, 2004
A study of the effects of active sonar transmissions on fish and marine mammals in Norwegian wate... more A study of the effects of active sonar transmissions on fish and marine mammals in Norwegian waters has been launched following the ordering of new frigates by the Royal Norwegian Navy (RNoN). The frigates will be equipped with active sonars operating at lower frequencies than those of the sonars currently in operation in the RNoN. Lower frequency sonar transmissions are believed to be potentially more harmful to marine life than higher frequency transmissions. The objective of the study is to acquire knowledge about the effects of active sonar on marine life, and produce a set of recommended rules for naval sonar operations in Norwegian waters based on scientific grounds.
The Journal of the Acoustical Society of America, 2009
Military antisubmarine sonars produce intense sounds within the hearing range of most clupeid fis... more Military antisubmarine sonars produce intense sounds within the hearing range of most clupeid fish. The behavioral reactions of overwintering herring ͑Clupea harengus͒ to sonar signals of two different frequency ranges ͑1-2 and 6 -7 kHz͒, and to playback of killer whale feeding sounds, were tested in controlled exposure experiments in Vestfjorden, Norway, November 2006. The behavior of free ranging herring was monitored by two upward-looking echosounders. A vessel towing an operational naval sonar source approached and passed over one of them in a block design setup. No significant escape reactions, either vertically or horizontally, were detected in response to sonar transmissions. Killer whale feeding sounds induced vertical and horizontal movements of herring.
Denne rapporten oppsummerer resultatet fra et internasjonalt forskningstokt i Vestfjorden i novem... more Denne rapporten oppsummerer resultatet fra et internasjonalt forskningstokt i Vestfjorden i november 2006. Bakgrunnen for undersøkelsen er påstander om at forsvarets bruk av sonarer skremmer bort sild og spekkhoggere fra området. Målsetningen for dette toktet var derfor å studere effekten av militaere lavfrekvente (LFAS 1-2 kHz) og mellomfrekvente (MFAS 6-7 kHz) aktive sonarer på spekkhogger og sild. I tillegg testet man bruk av både passive og aktive sonarer til deteksjon av marine pattedyr, slik at man kan begrense eventuelt negative effekter av militaere sonarer eller seismiske kilder som opereres i naerheten av pattedyr.
Advances in Experimental Medicine and Biology, 2016
Ramp-up procedures are used to mitigate the impact of sound on marine mammals. Sound exposure mod... more Ramp-up procedures are used to mitigate the impact of sound on marine mammals. Sound exposure models combined with observations of marine mammals responding to sound can be used to assess the effectiveness of ramp-up procedures. We found that ramp-up procedures before full-level sonar operations can reduce the risk of hearing threshold shifts with marine mammals, but their effectiveness depends strongly on the responsiveness of the animals. In this paper, we investigated the effect of sonar parameters (source level, pulse-repetition time, ship speed) on sound exposure by using a simple analytical model and highlight the mechanisms that limit the effectiveness of ramp-up procedures.
Advances in Experimental Medicine and Biology, 2016
Effects of noise on fish populations may be predicted by the population consequence of acoustic d... more Effects of noise on fish populations may be predicted by the population consequence of acoustic disturbance (PCAD) model. We have predicted the potential risk of population disturbance when the highest sound exposure level (SEL) at which adult herring do not respond to naval sonar (SEL0) is exceeded. When the population density is low (feeding), the risk is low even at high sonar source levels and long-duration exercises (>24 h). With densely packed populations (overwintering), a sonar exercise might expose the entire population to levels >SEL0 within a 24-h exercise period. However, the disturbance will be short and the response threshold used here is highly conservative. It is therefore unlikely that naval sonar will significantly impact the herring population.
Advances in Experimental Medicine and Biology, 2016
In mitigating the risk of sonar operations, the behavioral response of cetaceans is one of the ma... more In mitigating the risk of sonar operations, the behavioral response of cetaceans is one of the major knowledge gaps that needs to be addressed. The 3S-Project has conducted a number of controlled exposure experiments with a realistic sonar source in Norwegian waters from 2006 to 2013. In total, the following six target species have been studied: killer, long-finned pilot, sperm, humpback, minke, and northern bottlenose whales. A total of 38 controlled sonar exposures have been conducted on these species. Responses from controlled and repeated exposure runs have been recorded using acoustic and visual observations as well as with electronic tags on the target animal. So far, the first dose-response curves as well as an overview of the scored severity of responses have been revealed. In this paper, an overview is presented of the approach for the study, including the results so far as well as the current status of the ongoing analysis.
Frontiers in Physiology, May 10, 2012
Naval sonar has been accused of causing whale stranding by a mechanism which increases formation ... more Naval sonar has been accused of causing whale stranding by a mechanism which increases formation of tissue N 2 gas bubbles. Increased tissue and blood N 2 levels, and thereby increased risk of decompression sickness (DCS), is thought to result from changes in behavior or physiological responses during diving. Previous theoretical studies have used hypothetical sonar-induced changes in both behavior and physiology to model blood and tissue N 2 tension P N 2 , but this is the first attempt to estimate the changes during actual behavioral responses to sonar. We used an existing mathematical model to estimate blood and tissue N 2 tension P N 2 from dive data recorded from sperm, killer, long-finned pilot, Blainville's beaked, and Cuvier's beaked whales before and during exposure to Low-(1-2 kHz) and Mid-(2-7 kHz) frequency active sonar. Our objectives were: (1) to determine if differences in dive behavior affects risk of bubble formation, and if (2) behavioral-or (3) physiological responses to sonar are plausible risk factors. Our results suggest that all species have natural high N 2 levels, with deep diving generally resulting in higher end-dive P N 2 as compared with shallow diving. Sonar exposure caused some changes in dive behavior in both killer whales, pilot whales and beaked whales, but this did not lead to any increased risk of DCS. However, in three of eight exposure session with sperm whales, the animal changed to shallower diving, and in all these cases this seem to result in an increased risk of DCS, although risk was still within the normal risk range of this species. When a hypothetical removal of the normal dive response (bradycardia and peripheral vasoconstriction), was added to the behavioral response during model simulations, this led to an increased variance in the estimated end-dive N 2 levels, but no consistent change of risk. In conclusion, we cannot rule out the possibility that a combination of behavioral and physiological responses to sonar have the potential to alter the blood and tissue end-dive N 2 tension to levels which could cause DCS and formation of in vivo bubbles, but the actually observed behavioral responses of cetaceans to sonar in our study, do not imply any significantly increased risk of DCS.
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Papers by Petter Kvadsheim