Gray whale habitat use and reproductive success during seismic surveys near their feeding grounds: comparing state-dependent life history models and field data.

We used a stochastic dynamic programming (SDP) model to quantify the consequences of disturbance on pregnant western gray whales during one foraging season. The SDP model has a firm basis in bioenergetics, but detailed knowledge of minimum reproductive length of females (Lmin) and the relationship between length and reproductive success (Rfit) was lacking. We varied model assumptions to determine their effects on predictions of habitat use, proportion of animals disturbed, reproductive success, and the effects of disturbance. Smaller Lmin values led to higher predicted nearshore habitat use. Changes in Lmin and Rfit had little effect on predictions of the effect of disturbance. Reproductive success increased with increased Lmin and with higher probability of reproductive success by length. Multiple seismic surveys were conducted in 2015 off the northeast coast of Sakhalin Island, with concomitant benthic prey surveys, photo-identification studies, and whale distribution sampling, thus providing a unique opportunity to compare output from SDP models with empirical observations. SDP model predictions of reproductive success and habitat use were similar with and without acoustic disturbance, and SDP predictions of reproductive success and large-scale habitat use were generally similar to values and trends in the data. However, empirical estimates of the proportion of pregnant females nearshore were much higher than SDP model predictions (a large effect, measured by Cohen's d) during the first week, and the SDP model overestimated whale density in the south and underestimated density around the mouth of Piltun Bay. Such differences in nearshore habitat use would not affect SDP predictions of reproductive success or survival under the current seismic air gun disturbance scenario.

Western gray whale behavioral response to seismic surveys during their foraging season.

Gray whales utilizing their foraging grounds off northeastern Sakhalin Island, Russia, have been increasingly exposed to anthropogenic activities related to oil and gas development over the past two decades. In 2015, four seismic vessels, contracted by two operators, conducted surveys near and within the gray whale feeding grounds. Mitigation and monitoring plans were developed prior to the survey and implemented in the field, with real-time data transfers to assist the implementation of measures aimed at minimizing impacts of acoustic exposure. This study examined the behavioral response of gray whales relative to vessel proximities and sounds generated during seismic exploration. Five shore-based teams monitored gray whale behavior from 1 June to 30 September using theodolite tracking and focal follow methodologies. Behavioral data were combined with acoustic and benthic information from studies conducted during the same period. A total of 1270 tracks (mean duration = 0.9 h) and 401 focal follows (1.1 h) were collected with gray whales exposed to sounds ranging from 59 to 172 dB re 1 µPa2 SPL. Mixed models were used to examine 13 movement and 10 respiration response variables relative to "natural," acoustic, and non-acoustic explanatory variables. Water depth and behavioral state were the largest predictors of gray whale movement and respiration patterns. As vessels approached whales with increasing seismic/vessel sound exposure levels and decreasing distances, several gray whale movement and respiration response variables significantly changed (increasing speed, directionality, surface time, respiration intervals, etc.). Although the mitigation measures employed could have reduced larger/long-term responses and sensitization to the seismic activities, this study illustrates that mitigation measures did not eliminate behavioral responses, at least in the short-term, of feeding gray whales to the activities.

Gray whale density during seismic surveys near their Sakhalin feeding ground.

Oil and gas development off northeastern Sakhalin Island, Russia, has exposed the western gray whale population on their summer-fall foraging grounds to a range of anthropogenic activities, such as pile driving, dredging, pipeline installation, and seismic surveys. In 2015, the number of seismic surveys within a feeding season surpassed the level of the number and duration of previous seismic survey activities known to have occurred close to the gray whales' feeding ground, with the potential to cause disturbance to their feeding activity. To examine the extent that gray whales were potentially avoiding areas when exposed to seismic and vessel sounds, shore-based teams monitored the abundance and distribution of gray whales from 13 stations that encompassed the known nearshore feeding area. Gray whale density was examined in relation to natural (spatial, temporal, and prey energy) and anthropogenic (cumulative sound exposure from vessel and seismic sounds) explanatory variables using Generalized Additive Models (GAM). Distance from shore, water depth, date, and northing explained a significant amount of variation in gray whale densities. Prey energy from crustaceans, specifically amphipods, isopods, and cumaceans also significantly influenced gray whale densities in the nearshore feeding area. Increasing cumulative exposure to vessel and seismic sounds resulted in both a short- and longer-term decline in gray whale density in an area. This study provides further insights about western gray whale responses to anthropogenic activity in proximity to and within the nearshore feeding area. As the frequency of seismic surveys and other non-oil and gas anthropogenic activity are expected to increase off Sakhalin Island, it is critical to continue to monitor and assess potential impacts on this endangered population of gray whales.

Western gray whales on their summer feeding ground off Sakhalin Island in 2015: who is foraging where?

In the face of cumulative effects of oil and gas activities on the endangered western gray whale, informed management decisions rely on knowledge of gray whale spatial use patterns as a function of demographic group and prey energy. In particular, the gray whale foraging ground off Sakhalin Island consists of two distinct areas (nearshore and offshore) with the offshore feeding area exhibiting markedly high prey energy content. Based on photo-identification data collected from 2002 to 2015, we determined that gray whale use of the offshore feeding area increased with age. Pregnant females were more likely to be sighted only nearshore when nearshore prey energy and the proportion of nearshore energy from amphipods were higher. Likewise, females arriving with calves were less likely to be sighted offshore when the proportion of nearshore energy from amphipods was higher. Photo-identification effort in 2015 was increased substantially, with the intent of maximizing resighting data of individual whales to determine the relative proportion of different demographic groups utilizing the nearshore and offshore feeding areas. Comparing sighting data collected in 2015 with data from all previous years combined, mothers arriving with calves were sighted in the offshore feeding area earlier in 2015, with no evidence that they returned to forage nearshore later in the season. Other reproductive females constituted a higher proportion of the animals foraging nearshore prior to 2015, while juveniles were a higher proportion during 2015. Thus, the offshore feeding area is an important component of the gray whales' annual life cycle, particularly if nearshore prey energy continues to decline, and offshore anthropogenic activities need to be monitored and addressed.

Predicting the population consequences of acoustic disturbance, with application to an endangered gray whale population.

Acoustic disturbance is a growing conservation concern for wildlife populations because it can elicit physiological and behavioral responses that can have cascading impacts on population dynamics. State-dependent behavioral and life history models implemented via Stochastic Dynamic Programming (SDP) provide a natural framework for quantifying biologically meaningful population changes resulting from disturbance by linking environment, physiology, and metrics of fitness. We developed an SDP model using the endangered western gray whale (Eschrichtius robustus) as a case study because they experience acoustic disturbance on their summer foraging grounds. We modeled the behavior and physiological dynamics of pregnant females as they arrived on the feeding grounds and predicted the probability of female and offspring survival, with and without acoustic disturbance and in the presence/absence of high prey availability. Upon arrival in mid-May, pregnant females initially exhibited relatively random behavior before they transitioned to intensive feeding that resulted in continual fat mass gain until departure. This shift in behavior co-occurred with a change in spatial distribution; early in the season, whales were more equally distributed among foraging areas with moderate to high energy availability, whereas by mid-July whales transitioned to predominate use of the location that had the highest energy availability. Exclusion from energy-rich offshore areas led to reproductive failure and in extreme cases, mortality of adult females that had lasting impacts on population dynamics. Simulated disturbances in nearshore foraging areas had little to no impact on female survival or reproductive success at the population level. At the individual level, the impact of disturbance was unequally distributed across females of different lengths, both with respect to the number of times an individual was disturbed and the impact of disturbance on vital rates. Our results highlight the susceptibility of large capital breeders to reductions in prey availability, and indicate that who, where, and when individuals are disturbed are likely to be important considerations when assessing the impacts of acoustic activities. This model provides a framework to inform planned acoustic disturbances and assess the effectiveness of mitigation strategies for large capital breeders.

Site-fidelity and spatial movements of western North Pacific gray whales on their summer range off Sakhalin, Russia.

The Western North-Pacific (WNP) gray whale feeding grounds are off the northeastern coast of Sakhalin Island, Russia and is comprised of a nearshore and offshore component that can be distinguished by both depth and location. Spatial movements of gray whales within their foraging grounds were examined based on 13 years of opportunistic vessel and shore-based photo-identification surveys. Site fidelity was assessed by examining annual return and resighting rates. Lagged Identification Rates (LIR) analyses were conducted to estimate the residency and transitional movement patterns within the two components of their feeding grounds. In total 243 individuals were identified from 2002-2014, among these were 94 calves. The annual return rate over the period 2002-2014 was 72%, excluding 35 calves only seen one year. Approximately 20% of the individuals identified from 2002-2010 were seen every year after their initial sighting (including eight individuals that returned for 13 consecutive years). The majority (239) of the WNP whales were observed in the nearshore area while only half (122) were found in the deeper offshore area. Within a foraging season, there was a significantly higher probability of gray whales moving from the nearshore to the offshore area. No mother-calf pairs, calves or yearlings were observed in the offshore area, which was increasingly used by mature animals. The annual return rates, and population growth rates that are primarily a result of calf production with little evidence of immigration, suggest that this population is demographically self-contained and that both the nearshore and offshore Sakhalin feeding grounds are critically important areas for their summer annual foraging activities. The nearshore habitat is also important for mother-calf pairs, younger individuals, and recently weaned calves. Nearshore feeding could also be energetically less costly compared to foraging in the deeper offshore habitat and provide more protection from predators, such as killer whales.