Dawn P. Noren

Speaking up: Killer whales (Orcinus orca) increase their call amplitude in response to vessel noise

This study investigated the effects of anthropogenic sound exposure on the vocal behavior of free-ranging killer whales. Endangered Southern Resident killer whales inhabit areas including the urban coastal waters of Puget Sound near Seattle, WA, where anthropogenic sounds are ubiquitous, particularly those from motorized vessels. A calibrated recording system was used to measure killer whale call source levels and background noise levels (1-40 kHz). Results show that whales increased their call amplitude by 1 dB for every 1 dB increase in background noise levels. Furthermore, nearby vessel counts were positively correlated with these observed background noise levels.

Competing conservation objectives for predators and prey: estimating killer whale prey requirements for Chinook salmon.

Ecosystem-based management (EBM) of marine resources attempts to conserve interacting species. In contrast to single-species fisheries management, EBM aims to identify and resolve conflicting objectives for different species. Such a conflict may be emerging in the northeastern Pacific for southern resident killer whales (Orcinus orca) and their primary prey, Chinook salmon (Oncorhynchus tshawytscha). Both species have at-risk conservation status and transboundary (Canada–US) ranges. We modeled individual killer whale prey requirements from feeding and growth records of captive killer whales and morphometric data from historic live-capture fishery and whaling records worldwide. The models, combined with caloric value of salmon, and demographic and diet data for wild killer whales, allow us to predict salmon quantities needed to maintain and recover this killer whale population, which numbered 87 individuals in 2009. Our analyses provide new information on cost of lactation and new parameter estimates for other killer whale populations globally. Prey requirements of southern resident killer whales are difficult to reconcile with fisheries and conservation objectives for Chinook salmon, because the number of fish required is large relative to annual returns and fishery catches. For instance, a U.S. recovery goal (2.3% annual population growth of killer whales over 28 years) implies a 75% increase in energetic requirements. Reducing salmon fisheries may serve as a temporary mitigation measure to allow time for management actions to improve salmon productivity to take effect. As ecosystem-based fishery management becomes more prevalent, trade-offs between conservation objectives for predators and prey will become increasingly necessary. Our approach offers scenarios to compare relative influence of various sources of uncertainty on the resulting consumption estimates to prioritise future research efforts, and a general approach for assessing the extent of conflict between conservation objectives for threatened or protected wildlife where the interaction between affected species can be quantified.

Thermoregulation of Weaned Northern Elephant Seal (Mirounga angustirostris) Pups in Air and Water

Fasting weaned northern elephant seal pups (Mirounga angustirostris) experience diverse environmental conditions on land and in water on a daily basis. Each environment undoubtedly induces distinct energetic costs that may vary for pups of differing body condition. To determine the energetic costs associated with different environmental conditions and whether costs vary between individuals, body mass, surface area, volume, body composition, resting metabolic rate, and core body temperature were determined for 17 weaned northern elephant seal pups from Año Nuevo, California. Metabolic rate and body temperature were measured for pups resting in air (20.9° ± 0.8°C), cold water (3.8° ± 0.4°C), and warm water (14.5° ± 0.2°C). Resting metabolic rate increased with body mass (range: 62.0–108.0 kg) and was also correlated with lean mass and lipid mass. Metabolic rates ranged from 293.6 to 512.7 mL O2 min−1 and were lowest for pups resting in cold water. Thermal conductance, calculated from metabolic rate and core body temperature, ranged from 3.1 to 15.2 W °C−1, with the highest values in air and the lowest values in cold water. Metabolic responses to the three environmental conditions did not differ with individual variation in body condition. For all elephant seal pups, a consequence of high lipid content is that thermoregulatory costs are greatest on land and lowest in cold water, a pattern that contrasts markedly with terrestrial mammals.

Effects of noise levels and call types on the source levels of killer whale calls

Accurate parameter estimates relevant to the vocal behavior of marine mammals are needed to assess potential effects of anthropogenic sound exposure including how masking noise reduces the active space of sounds used for communication. Information about how these animals modify their vocal behavior in response to noise exposure is also needed for such assessment. Prior studies have reported variations in the source levels of killer whale sounds, and a more recent study reported that killer whales compensate for vessel masking noise by increasing their call amplitude. The objectives of the current study were to investigate the source levels of a variety of call types in southern resident killer whales while also considering background noise level as a likely factor related to call source level variability. The source levels of 763 discrete calls along with corresponding background noise were measured over three summer field seasons in the waters surrounding the San Juan Islands, WA. Both noise level and call type were significant factors on call source levels (1-40 kHz band, range of 135.0-175.7 dB(rms) re 1 [micro sign]Pa at 1 m). These factors should be considered in models that predict how anthropogenic masking noise reduces vocal communication space in marine mammals.

Vocal performance affects metabolic rate in dolphins: implications for animals communicating in noisy environments

ABSTRACT Many animals produce louder, longer or more repetitious vocalizations to compensate for increases in environmental noise. Biological costs of increased vocal effort in response to noise, including energetic costs, remain empirically undefined in many taxa, particularly in marine mammals that rely on sound for fundamental biological functions in increasingly noisy habitats. For this investigation, we tested the hypothesis that an increase in vocal effort would result in an energetic cost to the signaler by experimentally measuring oxygen consumption during rest and a 2 min vocal period in dolphins that were trained to vary vocal loudness across trials. Vocal effort was quantified as the total acoustic energy of sounds produced. Metabolic rates during the vocal period were, on average, 1.2 and 1.5 times resting metabolic rate (RMR) in dolphin A and B, respectively. As vocal effort increased, we found that there was a significant increase in metabolic rate over RMR during the 2 min following sound production in both dolphins, and in total oxygen consumption (metabolic cost of sound production plus recovery costs) in the dolphin that showed a wider range of vocal effort across trials. Increases in vocal effort, as a consequence of increases in vocal amplitude, repetition rate and/or duration, are consistent with behavioral responses to noise in free-ranging animals. Here, we empirically demonstrate for the first time in a marine mammal, that these vocal modifications can have an energetic impact at the individual level and, importantly, these data provide a mechanistic foundation for evaluating biological consequences of vocal modification in noise-polluted habitats. Summary: Increased vocal effort by bottlenose dolphins has metabolic costs at the individual level, indicating energetic consequences for wild populations of marine mammals communicating in noise-polluted environments.

The metabolic cost of communicative sound production in bottlenose dolphins (Tursiops truncatus)

SUMMARY Bottlenose dolphins (Tursiops truncatus) produce various communicative sounds that are important for social behavior, maintaining group cohesion and coordinating foraging. For example, whistle production increases during disturbances, such as separations of mother–calf pairs and vessel approaches. It is clear that acoustic communication is important to the survival of these marine mammals, yet the metabolic cost of producing whistles and other socials sounds and the energetic consequences of modifying these sounds in response to both natural and anthropogenic disturbance are unknown. We used flow-through respirometry to determine whether the metabolic cost of sound production could be quantified in two captive dolphins producing social sounds (whistles and squawks). On average, we found that metabolic rates measured during 2 min periods of sound production were 1.2 times resting values. Up to 7 min were required for metabolism to return to resting values following vocal periods. The total metabolic cost (over resting values) of the 2 min vocal period plus the required recovery period (163.3 to 2995.9 ml O2 or 3279.6 to 60,166.7 J) varied by individual as well as by mean duration of sounds produced within the vocal period. Observed variation in received cumulative sound energy levels of vocalizations was not related to total metabolic costs. Furthermore, our empirical findings did not agree with previous theoretical estimates of the metabolic cost of whistles. This study provides the first empirical data on the metabolic cost of sound production in dolphins, which can be used to estimate metabolic costs of vocal responses to environmental perturbations in wild dolphins.

Competing tradeoffs between increasing marine mammal predation and fisheries harvest of Chinook salmon

Many marine mammal predators, particularly pinnipeds, have increased in abundance in recent decades, generating new challenges for balancing human uses with recovery goals via ecosystem-based management. We used a spatio-temporal bioenergetics model of the Northeast Pacific Ocean to quantify how predation by three species of pinnipeds and killer whales (Orcinus orca) on Chinook salmon (Oncorhynchus tshawytscha) has changed since the 1970s along the west coast of North America, and compare these estimates to salmon fisheries. We find that from 1975 to 2015, biomass of Chinook salmon consumed by pinnipeds and killer whales increased from 6,100 to 15,200 metric tons (from 5 to 31.5 million individual salmon). Though there is variation across the regions in our model, overall, killer whales consume the largest biomass of Chinook salmon, but harbor seals (Phoca vitulina) consume the largest number of individuals. The decrease in adult Chinook salmon harvest from 1975–2015 was 16,400 to 9,600 metric tons. Thus, Chinook salmon removals (harvest + consumption) increased in the past 40 years despite catch reductions by fisheries, due to consumption by recovering pinnipeds and endangered killer whales. Long-term management strategies for Chinook salmon will need to consider potential conflicts between rebounding predators or endangered predators and prey.

Energetic Cost of Behaviors Performed in Response to Vessel Disturbance: One Link in the Population Consequences of Acoustic Disturbance Model

Several studies have shown that cetaceans respond to the physical presence and/or acoustic emissions from marine vessels. For example, cetaceans perform surface-active behaviors (SABs) in response to an increase in the number of and/or close approaches by vessels (Lusseau 2006; Noren et al. 2009; Williams et al. 2002, 2009). SABs are often performed in bouts of one or more behaviors performed sequentially, and the majority of SABs provide both visual and acoustic signals that are important to social marine mammals. Indeed, the use of sound is essential to the survival and reproduction of cetaceans (National Research Council 2003), and because of this, anthropogenic sound exposure in marine mammals is a concern. Individuals may compensate for increased vessel noise by changing the amplitude (Holt et al. 2009; Scheifele et al. 2005), duration (Foote et al. 2004), repetition rate, and/or frequency of the sounds they produce.

Assessing long‐term impacts of vocal compensation to ambient noise by measuring the metabolic cost of sound production in bottlenose dolphins.

The use of sound is essential to the survival of cetaceans. Therefore, anthropogenic sound exposure is a concern. Cetaceans can change the amplitude, duration, repetition rate, and/or frequency of sounds they produce to compensate for masking noise. Potential costs of such compensation are unknown, and no empirical data on the metabolic cost of sound production in marine mammals exist. This study aims to determine the metabolic cost of cetacean vocalizations to assess the biological significance of vocal compensation. Oxygen consumption, respiration rates, and vocalizations of two captive bottlenose dolphins were recorded during sound production at moderate levels. One dolphin produced his signature whistle while the other produced a pulsed squawk or squeaklike sound. Both types of vocalizations increased oxygen consumption while respiration rates did not change. This increased oxygen consumption is likely due to increased metabolic demand related to sound production, rather than changes in breathing patt...

Comparing the metabolic costs of different sound types in bottlenose dolphins

Cetaceans produce different types of sounds that vary according to behavioral context. They also modify their acoustic signals in response to noise. The metabolic costs of producing social sounds and clicks were recently measured in two bottlenose dolphins using flow-through respirometry methods. For both sound types, metabolic rates significantly increased as vocal effort increased, illustrating a modest cost of vocal modification. Using these data, metabolic costs can be extrapolated to more typical (higher) values of free-ranging dolphins and compared among sound types. However, cost comparisons are complicated by important differences in methodology and the acoustic energy output between sound type trials. In this investigation, existing data are analyzed specifically to scale and compare costs with these differences taken into account. Total metabolic cost of sound production was calculated above baseline values in ml O2 and related to vocal effort in dB re 1 μPa2s (adjusted to on-axis source levels)...