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Fifty-four right whale mortalities have been reported from between Florida, USA and the Canadian Maritimes from 1970 to 2002. Thirty of those animals were examined: 18 adults and juveniles, and 12 calves. Morphometric data are presented such that prediction of body weight is possible if the age, or one or more measurements are known. Calves grew approximately linearly in their first year. Total length and fluke width increased asymptotically to a plateau with age, weight increased linearly with age, weight and snout to blowhole distance increased exponentially with total length, whereas total length was linearly related to fluke width and flipper length. Among the adults and juveniles examined in this study, human interaction appeared to be a major cause of mortality, where in 14/18 necropsies, trauma was a significant finding. In 10/14 of these, the cause of the trauma was presumed to be vessel collision. Entanglement in fishing gear accounted for the remaining four cases. Trauma was also present in 4/12 calves. In the majority of calf mortalities (8/12) the cause of death was not determined. Sharp ship trauma included propeller lacerations inducing multiple, deep lacerations that often incised vital organs including the brain, spinal cord, major airways, vessels and musculature. Blunt ship trauma resulted in major internal bruising and fractures often without any obvious external damage. In at least two cases fatal gear entanglements were extremely protracted: where the entanglements took at least 100 and 163 days respectively to be finally lethal. The sum of these findings show two major needs: (1) that extinction avoidance management strategies focused on reducing trauma to right whales from ship collisions and fishing gear entanglement are highly appropriate and need to be continued and; (2) that as mitigation measures continue to be introduced into shipping and fishing industry practices, there is a strong effort to maximise the diagnostic quality of post-mortem examination of right whale mortalities, to ensure an optimal understanding of resultant trends.

Conservation and management of many cetaceans is hindered by the difficulty of acquiring samples from free-swimming individuals to obtain essential data on health, diet, reproduction and physiological impacts of environmental and anthropogenic stressors. This is particularly true for large whales, which are logistically difficult to live-capture for sampling. In North Atlantic right whales (Eubalaena glacialis), a significant decline in reproduction and health in the 1990s led to the application of faecal-based analyses to study stress and reproductive endocrinology, marine biotoxin exposure and prevalence of disease-causing protozoa. However, this approach was limited by low sample acquisition rates with opportunistic faecal (scat) collection methods. The work presented here evaluates the relative sampling efficiency of scent detection dogs trained to locate North Atlantic right whale scat versus opportunistic scat collection during photoidentification surveys. Three years of sample collection using both detection dogs and opportunistic methods are summarised. Faecal sample collection rates using detection dogs were over four times higher than opportunistic methods. The use of detection dogs for scat collection from free-swimming right whales has for the first time provided adequate numbers of samples for statistical analyses. The endocrine, disease, genetic and biotoxin studies currently being performed on these samples markedly improve the ability to address fundamental questions vital to effective conservation and management of highly endangered right whales.

Because of the increase in usage of remote sensing tags to track the migration and movements of cetaceans, it has become increasingly important to assess the impact of such techniques on the target species. Between 1988 and 1997, 55 tags (41 satellite telemetry, 14 VHF or acoustic radio transmitter) were attached to 49 North Atlantic right whales (Eubalaena glacialis). All tags had implantable barbs or were fully implanted below the dermis. Right whales are photographically identifiable and the New England Aquarium curates the North Atlantic catalog, which currently numbers 374 individuals. The photo catalog has made it possible for tagged individuals to be tracked after the tag falls off the whale. Photo documentation during and after tagging provides an opportunity to monitor physiological effects from tags and healing responses to tags.

Enclosed is a report on a workshop held at the New England Aquarium on October 23rd, 1999, to review the effects of implantable satellite and radio tags on right whales in the North Atlantic. Transcripts from the meeting served as the basis for a draft of this report, which was circulated on April 12th . Comments were received from several participants and have been incorporated. Although no consensus was reached on the workshop topic, most participants agreeing on the need for better follow-up of animals subject to tagging activities. Additional recommendations were made by participants, but they are not consensus statements. The results of the workshop do not provide support for a moratorium on tagging, but they also do not suggest that researchers can be complacent about the potential effects on animals. The need to identify missing right whale habitats means that tagging will continue to be an essential part of the biologist's toolkit. This report indicates that while tag technology is rapidly improving, a commensurate improvement in monitoring effects is needed. It seems likely that both will lead to better tagging efforts in the future.

Funding: Office of Naval Research - N000142012697, N000142112096; Strategic Environmental Research and Development Program - RC20-1097, RC20-7188, RC21-3091. Assessing cumulative effects of human activities on ecosystems is required by many jurisdictions, but current science cannot meet regulatory demands. Regulations define them as effect(s) of one human action combined with other actions. Here we argue for an approach that evaluates the cumulative risk of multiple stressors for protected wildlife populations within their ecosystems. Monitoring effects of each stressor is necessary but not sufficient to estimate how multiple stressors interact to affect wildlife populations. Examining the mechanistic pathways, from cellular to ecological, by which stressors affect individuals can help prioritize stressors and interpret how they interact. Our approach uses health indicators to accumulate the effects of stressors on individuals and to estimate changes in vital rates, driving population status. We advocate using methods well-established in human health and integrating them into ecosystem-based management to protect the health of commercially and culturally important wildlife populations and to protect against risk of extinction for threatened species. Our approach will improve abilities to conserve and manage ecosystems but will also demand significant increases in research and monitoring effort. We advocate for increased investment proportional to the economic scale of human activities in the Anthropocene and their pervasive effects on ecology and biodiversity. Publisher PDF

Offshore wind energy development is growing quickly around the world. In southern New England, USA, one of the largest commercial offshore wind energy farms in the USA will be established in the waters off Massachusetts and Rhode Island, an area used by the Critically Endangered North Atlantic right whale Eubalaena glacialis. Prior to 2011, little was known about the use of this area by right whales. We examined aerial survey data collected between 2011-2015 and 2017-2019 to quantify right whale distribution, residency, demography, and movements in the region. Right whale occurrence increased during the study period. Since 2017, whales have been sighted in the area nearly every month, with peak sighting rates between late winter and spring. Model outputs suggest that 23% of the species’ population is present from December through May, and the mean residence time has tripled to an average of 13 d during these months. Age and sex ratios of the individuals present in the area are similar to those of the species as a whole, with adult males the most common demographic group. Movement models showed that southern New England is an important destination for right whales, including conceptive and reproductive females, and qualitative observations included animals feeding and socializing. Implementing mitigation procedures in coordination with these findings will be crucial in lessening the potential impacts on right whales from construction noise, increased vessel traffic, and habitat disruption in this region.

Funding information: Office of Naval Research, Grant/Award Numbers: N000141126207, N000141210286, N000141210389, N000141712817; Strategic Environmental Research and Development Program, Grant/Award Number: RC20-C2-1097; National Oceanic and Atmospheric Administration. North Atlantic right whales frequently become entangled in fishing gear, which can negatively affect their reproductive output and probability of survival. We estimated individual whale health from a hierarchical Bayesian model fit to photographic indices of health. We reviewed 696 whales sighted from 1980 to 2011 and assigned 1196 entanglement events to 573 individuals in six categories of increasing injury severity and estimated monthly median health scores (0–100 scale) for the duration of their life within the study period. We then quantified the relationship between entanglement injury events and their severity with survival, reproduction, and population health. Severe entanglements resulted in worse health for all whales—males and females with severe injuries were eight times more likely to die than males with minor injuries. Females with severe injuries that survived had the lowest birth rates. Though the relationship between entanglement and fecundity was complex, we found that as the health of reproductively active females declined, their calving intervals increased. Unimpacted whale health scores declined significantly over three decades, 1980s, 1990s, and 2000s, suggesting food limitations may be contributing to population-wide health declines. Decadal health scores of entangled whales showed a more notable reduction in health suggesting a clear and perhaps synergistic effect. Publisher PDF

The population of western North Atlantic right whales (Eubalaena glacialis) is distributed primarily between Florida, USA and NovaScotia, Canada, aggregating seasonally in five geographically distinct, high-use areas. To test the effectiveness of monitoring alldemographic classes (juvenile and adult males and females) of the population in these five habitat areas, an evaluation was carried out ofthe identification records of catalogued right whales collected between 1980 and 1992, for which the age, sex and reproductive status (foradult females) were known. The mean annual identification frequency of adult females was significantly lower than that of adult males,juvenile females and juvenile males. Among adult females, reproductively active females were seen significantly more often than expectedwhen lactating (with a calf) than during their pregnancy or resting years. These data suggest that, while research efforts in the five high-usehabitat areas have had relatively equal success at monitoring juvenile males and females and adult males, many adult females are segregatedat times from the rest of the population. Lower variability in annual identification frequencies of adult females indicates that they may bemore site specific in their distribution than males, particularly during the years when they are pregnant or resting from a previous pregnancy.Re-running these analyses using sighting records updated through 2000 will help determine if the trends continue to be documentedregardless of changes in survey effort and patterns of habitat use of some animals.

North Atlantic right whale reproduction was assessed for the period 1980 through 1998. At the end of this period, we estimated there werebetween 299 and 437 right whales alive, including 70 mature females. Using maximum and minimum population estimates for the entireperiod, mean values for gross annual reproductive rate were 0.36 and 0.49 respectively, and the mean value for calves per mature femaleper year was 0.25. There is a significant decreasing trend in calves per mature female per year over the entire study period. The mean ageat first calving is 9.53 years. The mean number of cows recruited annually since 1985 is 3.8. Mean annual calving intervals have increasedsignificantly during the study period from 3.67 years (1980-1992) to over 5 years (1993-1998). Although the North Atlantic population isaffected by significant anthropogenic mortality, diminishing reproductive rates are probably also responsible for the plight of thisspecies.

Northern right whales in the western North Atlantic number about 300 animals and have shown little sign of recovery in recent d ecades. Mortality and serious injury due to human activities, particularly commercial fishing and shipping, are thought to be significa nt factors limiting their recovery. From 1970-1999, 45 right whale deaths were reliably documented. Sixteen of these fatalities (35.5%) we re due to ship collisions, and three (6.7%) were due to entanglement in fishing gear. The remainder were neonates (13; 28.9%) and ‘unknown cause’ mortalities (13; 28.9%). Criteria for defining serious injuries and mortalities from entanglement or ship strikes were develope d and include any animal carrying fishing gear, cuts from entanglement or ship strike deeper than 8cm, swelling or necrosis, evidence of poor health from such interactions, and, in carcasses, evidence of haematoma, haemorrhaging or broken bones. A total of 56 animals fitting the d efined criteria were documented from 1970-1999: 31 (55.4%) from entanglement and 25 (44.6%) from ship strikes. Nineteen were fatal (16 ship strikes, 3 entanglements), 10 were possibly fatal (2 ship strikes, 8 entanglements) and 27 were non-fatal (7 ship strikes, 20 e ntanglements). The breakdown of potentially serious injuries by age and sex reveals no difference in levels between sexes but shows a 3.3:1 hi gher level of interaction in juveniles and calves versus adults. The data show that ship strikes are more immediately lethal, but entanglements can result in long term deterioration of an animal and may be responsible for higher levels of mortality than previously thought. Consider ing that some animals become entangled, drown and never return to the surface, even these levels may be underestimated. Between 1986 and 1999 , 84 animals were presumed dead based on a lack of resightings for six years. There were 32 confirmed deaths during this time period suggesting that at least as many unreported deaths occurred as carcasses were reported. Definitive actions need to be taken to reduce the level and severity of anthropogenic injuries and deaths. Actions could include continued disentanglement efforts, gear modifications, sea sonal closures for fisheries, mandatory ship reporting, ships’ routing measures and speed restrictions for commercial shipping.

One of today’s greatest conservation challenges is balancing policies, laws, and management strategies established to achieve economic goals that depend on extracting ocean resources with those established to conserve marine biodiversity. We use the Northeast Canyons and Seamounts Marine National Monument (NCSM) as a case study to explore the consequences of changing from a policy that prevents fishing to a fisheries management policy. We found that opening the NCSM to commercial fishing reduces species protections. Fishing with pelagic and bottom tending fixed gear exposes species inhabiting the sea surface and midwater regions to entanglement and bycatch risk. Fishing with bottom tending fixed gear also exposes deep-sea coral habitat to gear known to have detrimental impacts. The NCSM was designated as a marine national monument due to the area’s unique ecological resources that are a subject of scientific interest. Our case study demonstrates that a fisheries management policy is insufficient to protect these ecological resources.

Evaluations of the conservation status of the endangered North Atlantic right whale as well as many other wildlife species often rely extensively on counts and cause‐of‐death determinations of carcasses found accidentally or during dedicated surveys. Even when survey effort dedicated to a population is extensive, many deaths may go unseen. We used an abundance estimation model to derive estimates of cryptic mortality for North Atlantic right whales and found that observed carcasses accounted for only 36% of all estimated death during 1990–2017. We found strong evidence that total mortality varied over time, and that observed carcass counts were poor predictors of estimated annual numbers of whales dying. Importantly, there were substantial differences between fractions of deaths determined to be entanglement related during necropsy (49%) and the fraction of cryptic deaths suffering serious injuries related to entanglement (87%). Although we concluded that a single year's observations produced poor estimates of carcass detection rates due to the volatility of ratios of small counts, ratio estimates of data pooled over periods of consistent survey may offer better information on detection rates. Additionally, it appears unwise to consider cause of death determinations from detected carcasses as representative of cause‐specific mortality rates in right whales given the large number of seriously injured whales from entanglement that are likely part of the unseen mortality.

The Northeast Canyons and Seamounts Marine National Monument (NECSMNM) was designated by President Barack Obama in 2016, using his authority under the Antiquities Act of 1906. The Act allows a President to proclaim as national monuments “historic landmarks, historic and prehistoric structures, and other objects of historic or scientific interest” that are “upon the lands owned or controlled” by the United States but to reserve each designation to “the smallest area compatible with the proper care and management of the objects to be protected.” Protection in general excludes commercial scale extraction and is in perpetuity. Here we present analyses of physiographic and ecological datasets that facilitated assessment of the conservation benefits of protections for a new monument. We also review and synthesize the ecological literature to describe processes that operate in continental margin and deep-sea settings, in order to demonstrate the monument area is bounded for proper management and is an object of scientific interest. Results indicate that the current monument designation is an area of high diversity and ecological connectivity across depths and along the continental margin. The monument boundaries contain hot spots (areas of high abundance and species richness) for seafloor communities (inclusive of benthic invertebrate and demersal fish) as well as marine mammals in the epipelagic. Many species are sensitive to disturbance and vulnerable to human activities (e.g., deep-sea corals and sponges) with very long recovery times and extremely low resilience. The monument contains at least nine exemplars of offshore northwest Atlantic marine wildlife communities and habitats (e.g., deep shelf invertebrates, shelf fish, deep sea corals and sponges in canyons and on seamounts, deep sea fish, chemosynthetic communities, deep sea soft sediment, shelf edge cetaceans, and seabirds). The region is relatively undisturbed and can serve as a reference site to focus future research on ecological processes in an increasingly industrialized ocean and one subject to the synergies of regional climate effects. These results suggest that there is great potential for discovery and novel research in this first Atlantic Ocean Marine National Monument.

Six baleen whale species are found in the temperate western North Atlantic Ocean, with limited information existing on the distribution and movement patterns for most. There is mounting evidence of distributional shifts in many species, including marine mammals, likely because of climate‐driven changes in ocean temperature and circulation. Previous acoustic studies examined the occurrence of minke (Balaenoptera acutorostrata) and North Atlantic right whales (NARW; Eubalaena glacialis). This study assesses the acoustic presence of humpback (Megaptera novaeangliae), sei (B. borealis), fin (B. physalus), and blue whales (B. musculus) over a decade, based on daily detections of their vocalizations. Data collected from 2004 to 2014 on 281 bottom‐mounted recorders, totaling 35,033 days, were processed using automated detection software and screened for each species' presence. A published study on NARW acoustics revealed significant changes in occurrence patterns between the periods of 2004–2010 and 2011–2014; therefore, these same time periods were examined here. All four species were present from the Southeast United States to Greenland; humpback whales were also present in the Caribbean. All species occurred throughout all regions in the winter, suggesting that baleen whales are widely distributed during these months. Each of the species showed significant changes in acoustic occurrence after 2010. Similar to NARWs, sei whales had higher acoustic occurrence in mid‐Atlantic regions after 2010. Fin, blue, and sei whales were more frequently detected in the northern latitudes of the study area after 2010. Despite this general northward shift, all four species were detected less on the Scotian Shelf area after 2010, matching documented shifts in prey availability in this region. A decade of acoustic observations have shown important distributional changes over the range of baleen whales, mirroring known climatic shifts and identifying new habitats that will require further protection from anthropogenic threats like fixed fishing gear, shipping, and noise pollution., Limited information exists on the distribution and movement patterns for most baleen whale species in the western North Atlantic Ocean. Previously, we conducted a broad‐scale passive acoustic monitoring study to analyze North Atlantic right whale seasonal distribution and shifts therein, based on identified changes in occurrence patterns starting in 2010. This study uses similar acoustic datasets and protocols to understand the seasonal distribution of humpback, sei, fin, and blue whales within the western North Atlantic, and to determine whether any of these species exhibited similar shifts in distribution patterns across time.

Cetorhinus maximus aggregations recorded during extensive aerial survey efforts off the north-eastern United States between 1980 and 2013 included aggregations centring on sightings with group sizes of at least 30 individuals. These aggregations occurred in summer and autumn months and included aggregation sizes of up to 1398 individuals, the largest aggregation ever reported for this species. The aggregations were associated with sea surface temperatures of 13-24° C and chlorophyll-a concentrations of 0·4-2·6 mg m-3 and during one aggregation, a high abundance of zooplankton prey was present. Photogrammetric tools allowed for the estimation of total body lengths ranging between 4 and 8 m. Characterization of these events provides new insight into the potential biological function of large aggregations in this species.

Until now, physiological stress assessment of large whales has predominantly focused on adrenal glucocorticoid (GC) measures. Elevated GC concentrations in feces (fGC) are known to reflect stressful disturbances, such as fishing gear entanglement and human-generated underwater noise, in North Atlantic right whales (Eubalaena glacialis). However, there can be considerable variation in GC production as a function of sex and life history stage, which may confound the interpretation of fGC levels. Additionally, GC antibodies used in immunoassays can cross-react with other fecal metabolites (i.e., non-target steroids), potentially influencing fGC data. Here, aldosterone concentrations (fALD; aldosterone and related metabolites) were measured in fecal samples from right whales (total n=315 samples), including samples from identified individuals of known life history (n=82 individual whales), to evaluate its utility as a complementary biomarker to fGC for identifying adrenal activation. Concentrations of fALD were positively correlated with fGCs in right whales (r=0.59, P0.001), suggesting concurrent secretion of these hormones by the adrenal gland. However, fALD levels were less influenced by concentrations of reproductive steroids in feces, minimizing the potential confounder of assay cross-reactivity in samples with highly skewed hormone ratios. Across different life history states for right whales, fALD concentrations showed similar patterns to those reported for fGC, with higher levels in pregnant females (35.9±7.6ng/g) followed by reproductively mature males (9.5±0.9ng/g) (P0.05), providing further evidence of elevated adrenal activation in these groups of whales. The addition of fALD measurement as a biomarker of adrenal activation may help distinguish between intrinsic and external causes of stress hormone elevations in large whales, as well as other free-living wildlife species, providing a more comprehensive approach for associating adrenal activation with specific natural and anthropogenic stressors.

Given new distribution patterns of the endangered North Atlantic right whale (NARW; Eubalaena glacialis) population in recent years, an improved understanding of spatio-temporal movements are imperative for the conservation of this species. While so far visual data have provided most information on NARW movements, passive acoustic monitoring (PAM) was used in this study in order to better capture year-round NARW presence. This project used PAM data from 2004 to 2014 collected by 19 organizations throughout the western North Atlantic Ocean. Overall, data from 324 recorders (35,600 days) were processed and analyzed using a classification and detection system. Results highlight almost year-round habitat use of the western North Atlantic Ocean, with a decrease in detections in waters off Cape Hatteras, North Carolina in summer and fall. Data collected post 2010 showed an increased NARW presence in the mid-Atlantic region and a simultaneous decrease in the northern Gulf of Maine. In addition, NARWs were widely distributed across most regions throughout winter months. This study demonstrates that a large-scale analysis of PAM data provides significant value to understanding and tracking shifts in large whale movements over long time scales.

As the need for aquaculture continues to grow, expansion of marine aquaculture into the coastal ocean and beyond will require an understanding of ocean use and attention to spatial planning with en...

Recent surveys of wind energy areas offshore of Massachusetts and Rhode Island (USA) have demonstrated that they encompass habitat used by the Endangered North Atlantic right whale Eubalaena glacialis. Prior to 2011, little systematic survey effort had been conducted in the area. The Bureau of Ocean Energy Management and the state of Massachusetts supported 3.5 yr of twice-monthly aerial surveys by the Northeast Large Pelagic Survey Collaborative (NLPSC). Additional survey teams including the Northeast Fisheries Science Center and the Center for Coastal Studies have collected sightings data in the region. Data systematically collected by the NLPSC allowed analyses of monthly sightings rates, sightings per unit effort, and hot spots which provided information on current temporal and spatial use patterns. Abundance estimates for each season-year (i.e. a 3 mo period within a given survey year) were calculated. Behaviors observed included feeding and surface active groups. Photo-identification of whales since 2010 yielded a minimum count of 196 unique individuals (annual average = 35), or over one-third of the current population estimate. Analyses of demographics of these individuals revealed that 34 known calving females (30% of the total currently presumed alive) visited the study area. These results demonstrate consistent annual use of this area by a significant portion of the E. glacialis population, with a strong correlation between season and presence. These findings can inform management activities and development planning, and be used as a baseline dataset for assessing long-term impacts to the species.

Two wind energy areas (WEAs), designated by the Bureau of Ocean Energy Management, offshore of Massachusetts and Rhode Island were surveyed between October 2011 and June 2015. A total of 969 cetacean sightings of over 10,000 animals were documented during systematic line-transect aerial surveys (67,525 km flown). Twelve cetacean species were documented, including seven odontocete and five mysticete species. Cetaceans were recorded in all seasons demonstrating inter-annual, seasonal, and spatial use of the WEAs. Peak presence occurred in the spring and summer for most species, with the exception of North Atlantic right whales, which occurred primarily in the winter and spring. Right and sei whales were documented regularly throughout the study and in higher numbers than expected. Seasonal estimates of abundance were calculated for seven species, including five baleen whales (fin, minke, humpback, right, and sei) and two dolphins (common and bottlenose). The results suggest the area is a previously underestimated seasonal habitat for cetaceans including several endangered baleen whale species. These data will help inform mitigation strategies for future wind energy development. Climate change effects, increased coastal urbanization, and other factors argue for continued monitoring of cetaceans in the WEAs offshore of New England.

Body condition has been correlated with survival and reproductive success in both terrestrial and marine mammals, including North Atlantic right whales Eubalaena glacialis. We used photographs of individually identified right whales to assess visual changes in body condition in reproductive females, adult males, juveniles, and entangled whales. Images from sightings of individual whales were grouped sequentially by habitat region, and each group of images was assigned a body condition score of good, fair, or poor based on the dorsal profile posterior to the blowholes. Temporally consecutive groups of images (n = 1496) of 340 individual whales were compared to investigate the frequency, direction, and minimum timeframe between changes in body condition. Changes in body condition scores of right whales were significantly influenced by group category. Lactating females and severely entangled right whales were more likely to exhibit declining body condition than other groups. Resting females were significantly more likely to improve in condition than other groups but exhibited the longest timeframe for improving condition. Young juveniles were less likely to improve in condition compared to adult males, but remained in compromised condition less frequently than older juveniles and adult males. The shortest timeframes between changing body condition scores were 11 d for declining condition and 12 d for improving condition. This study demonstrates that photographic analysis can detect rapid body condition changes and identifies groups of right whales that are particularly vulnerable to declining condition and delayed recovery from energetically taxing events.

All three extant right whales [Eubalaena australis (Southern; SRW), glacialis (North Atlantic; NARW), and japonica (North Pacific; NPRW)] were heavily exploited, and the status of the two northern hemisphere species remains precarious. Recently, limited gains made by the NARW have been reversed and urgent changes to management approaches are needed if extinction is to be averted. By contrast, some SRW populations are recovering. Given their close phylogenetic relationship, morphological, demographic, and ecological similarities, the contrasting recovery rates between populations and species provide an opportunity to apply a comparative approach to inform the differences in recovery as follows. (1) Recovery: All right whale species were internationally protected in 1931, but NARW, eastern NPRW and some SRW populations have barely recovered from whaling, while others are doing so at maximal rates. Are these differences a legacy of extreme depletion (e.g., loss of genetic diversity and cultural knowledge) or primarily due to anthropogenic factors (e.g., high mortality from ship strike and fisheries entanglement)? If modern anthropogenic threats are not affecting remote SRW populations, can these serve as baseline populations for comparison with NARW and NPRW? (2) Linking individuals to population-level responses: In wild mammals, strong links exist between reproductive indices and environmental conditions within the context of life-history strategies. Individual identification of whales provides the ability to track survival, reproduction and other demographic parameters, and their population-level consequences, providing the tools with which to uncover these links. Robust life-history analyses are now available for NARW and several SRW populations, linking demography with environmental conditions, providing the potential for teasing out important influencing factors. (3) Adapting to shifting resources: Recent reproductive declines in NARW appear linked to changing food resources. While we know some large-scale movement patterns for NARW and a few SRW populations, we know little of mesoscale movements. For NPRW and some SRW populations, even broad-scale movements are poorly understood. In the face of climate change, can methodological advances help identify Eubalaena distributional and migratory responses? (4) Emergent diseases and the vulnerability of populations under stress: Marine mammals are vulnerable to infectious diseases, particularly when subjected to stressors such as fishing gear entanglements, acoustic disturbance, and prey shortages. New tools to assess large whale health include body condition imaging, viromes, microbiomes, as well as metabolic and stress hormones. Comparative analysis of the three Eubalaena spp. could identify causes of varying recovery. (5) Comparative synthesis and cumulative effects: The lack of a good analytical approach for cumulative effects is an urgent bio-statistical problem in conservation biology. Without such a framework every stressor is managed in isolation, limiting efficacy. We propose a comparative synthesis to inform future cumulative effect analyses and outline future research priorities to achieve these goals. Publisher PDF

We provide guidance for monitoring whether human activities affect the physiology or behavior of marine mammals and, if so, whether those effects may lead to changes in survival and reproduction at the population level. We suggest that four elements be included in designing and implementing such a monitoring program. The first is development of a theory of change: a set of mechanistic hypotheses that outline why a given activity might be expected to have one or more measurable effects on individuals and populations, and ideally the magnitude, timing, and duration of the effects. The second element, definition of biologically meaningful effect sizes, ultimately facilitates the development of a monitoring program that can detect those magnitudes of effect with the desired levels of precision. The third element, selection of response variables for monitoring, allows inference to whether observed changes in the status of individuals or populations are attributable to a given activity. Visual observations, passive acoustic and tagging instruments, and direct physical measurements all can provide data that facilitate quantitative hypothesis testing. The fourth element is specification of the temporal sequence of monitoring. These elements also can be used to inform monitoring of the responses of other taxonomic groups to human activities.

Entanglement in fixed fishing gear affects whales worldwide. In the United States, deaths of North Atlantic right (Eubalaena glacialis) and humpback whales (Megaptera novaeangliae) have exceeded management limits for decades. We examined live and dead whales entangled in fishing gear along the U.S. East Coast and the Canadian Maritimes from 1994 to 2010. We recorded whale species, age, and injury severity and determined rope polymer type, breaking strength, and diameter of the fishing gear. For the 132 retrieved ropes from 70 cases, tested breaking strength range was 0.80-39.63 kN (kiloNewtons) and the mean was 11.64 kN (SD 8.29), which is 26% lower than strength at manufacture (range 2.89-53.38 kN, mean = 15.70 kN [9.89]). Median rope diameter was 9.5 mm. Right and humpback whales were found in ropes with significantly stronger breaking strengths at time of manufacture than minke whales (Balaenoptera acuturostrata) (19.30, 17.13, and 10.47 mean kN, respectively). Adult right whales were found in stronger ropes (mean 34.09 kN) than juvenile right whales (mean 15.33 kN) and than all humpback whale age classes (mean 17.37 kN). For right whales, severity of injuries increased since the mid 1980s, possibly due to changes in rope manufacturing in the mid 1990s that resulted in production of stronger ropes at the same diameter. Our results suggest that broad adoption of ropes with breaking strengths of ≤ 7.56 kN (≤ 1700 lbsf) could reduce the number of life-threatening entanglements for large whales by at least 72%, and yet could provide sufficient strength to withstand the routine forces involved in many fishing operations. A reduction of this magnitude would achieve nearly all the mitigation legally required for U.S. stocks of North Atlantic right and humpback whales. Ropes with reduced breaking strength should be developed and tested to determine the feasibility of their use in a variety of fisheries.

Managing the nonlethal effects of disturbance on wildlife populations has been a long‐term goal for decision makers, managers, and ecologists, and assessment of these effects is currently required by European Union and United States legislation. However, robust assessment of these effects is challenging. The management of human activities that have nonlethal effects on wildlife is a specific example of a fundamental ecological problem: how to understand the population‐level consequences of changes in the behavior or physiology of individual animals that are caused by external stressors. In this study, we review recent applications of a conceptual framework for assessing and predicting these consequences for marine mammal populations. We explore the range of models that can be used to formalize the approach and we identify critical research gaps. We also provide a decision tree that can be used to select the most appropriate model structure given the available data. Synthesis and applications: The implementation of this framework has moved the focus of discussion of the management of nonlethal disturbances on marine mammal populations away from a rhetorical debate about defining negligible impact and toward a quantitative understanding of long‐term population‐level effects. Here we demonstrate the framework's general applicability to other marine and terrestrial systems and show how it can support integrated modeling of the proximate and ultimate mechanisms that regulate trait‐mediated, indirect interactions in ecological communities, that is, the nonconsumptive effects of a predator or stressor on a species' behavior, physiology, or life history.

Exhaled breath analysis is a non-invasive assessment tool that has shown promise in human diagnostics, and could greatly benefit research, management, and conservation of large whales. However, hormone assessment of whale respiratory vapor (blow) has been challenged by variable water content and unknown total volume of collected samples. To advance this technique, we investigated urea (a compound present in narrow range in circulation) as a normalizing factor to correct for blow sample concentration. Normalized progesterone, testosterone, and cortisol concentrations of 100 blow samples from 46 photo-identified North Atlantic right whales (Eubalaena glacialis) were more biologically relevant compared to absolute estimates, varying by sex, age class, or individual. Progesterone was elevated in adult females compared with other cohorts and highest in one independently confirmed pregnant female. For both sexes, testosterone was two-fold higher in reproductively mature whales but studied adult females showed the widest variation. Cortisol was present in relatively low concentrations in blow and demonstrated variation between individual whales, suggesting potential for studies of individual differences in adrenal activity. Incorporation of methodologies that normalize sample concentration are essential for blow hormone analysis of free-swimming whales, and measurement of urea could be used to optimize non-invasive physiological assessment of whales.

Summary Fishing gear entanglement is a problem for sea birds, marine turtles, dolphins, and other marine mammals. Here, large whale entanglements are highlighted.

North Atlantic right whales (Eubalaena glacialis Müller 1776) present an interesting problem for abundance and trend estimation in marine wildlife conservation. They are long lived, individually identifiable, highly mobile, and one of the rarest of cetaceans. Individuals are annually resighted at different rates, primarily due to varying stay durations among several principal habitats within a large geographic range. To date, characterizations of abundance have been produced that use simple accounting procedures with differing assumptions about mortality. To better characterize changing abundance of North Atlantic right whales between 1990 and 2015, we adapted a state–space formulation with Jolly‐Seber assumptions about population entry (birth and immigration) to individual resighting histories and fit it using empirical Bayes methodology. This hierarchical model included accommodation for the effect of the substantial individual capture heterogeneity. Estimates from this approach were only slightly higher than published accounting procedures, except for the most recent years (when recapture rates had declined substantially). North Atlantic right whales' abundance increased at about 2.8% per annum from median point estimates of 270 individuals in 1990 to 483 in 2010, and then declined to 2015, when the final estimate was 458 individuals (95% credible intervals 444–471). The probability that the population's trajectory post‐2010 was a decline was estimated at 99.99%. Of special concern was the finding that reduced survival rates of adult females relative to adult males have produced diverging abundance trends between sexes. Despite constraints in recent years, both biological (whales' distribution changing) and logistical (fewer resources available to collect individual photo‐identifications), it is still possible to detect this relatively recent, small change in the population's trajectory. This is thanks to the massive dataset of individual North Atlantic right whale identifications accrued over the past three decades. Photo‐identification data provide biological information that allows more informed inference on the status of this species.

Immunoassay of hormone metabolites extracted from faecal samples of free-ranging large whales can provide biologically relevant information on reproductive state and stress responses. North Atlantic right whales (Eubalaena glacialis Muller 1776) are an ideal model for testing the conservation value of faecal metabolites. Almost all North Atlantic right whales are individually identified, most of the population is sighted each year, and systematic survey effort extends back to 1986. North Atlantic right whales number

To enhance recovery of the Endangered North Atlantic right whale Eubalaena glacialis, mitigation strategies are needed to reduce the leading causes of injury and mortality of these animals, which include ship strikes and entanglement in fishing gear. Such efforts require information on the spatial and temporal distribution of right whales that can be analyzed against risk factors to identify solutions to protect whales while minimizing economic impacts to industry. Currently, no methods adequately make use of all available data to characterize right whale distribution, often leaving data gaps or misrepresenting whale activity in areas where we have limited systematic survey data. In response to this, an areal co-kriging interpolation technique was developed using ArcGIS 10.1 Geostatistical Analyst, utilizing all available right whale location data including systematic survey data, opportunistic sightings data, and satellite tag data. With many management plans shaped around specific geographic zones, this methodology was developed with the option to summarize information within user-defined polygons depending on the management regime. For this paper, whale distribution predictions were summarized to 524 irregularly shaped polygons off coastal Maine, representing lobster fishing zones at a 2 mo temporal scale. Results indicate that the predicted values fall within reasonable ranges, appropriately represent seasonal differences, and better represent right whale distribution patterns compared to other methods.

This work is partially supported by The Marine Alliance for Science and Technology for Scotland pooling initiative (funded by the Scottish Funding Council, grant reference HR09011, and contributing institutions) Environmental changes (a type of disturbance) are altering the habitat of southern elephant seals Mirounga leonina, an apex marine predator in the Southern Ocean. As a result, individuals may shift their behaviour, spending more time in transit and less time foraging. The effects of these sublethal changes in behaviour can accumulate, indirectly impacting lifetime fitness through changes in individual survival and reproduction. If a sufficient proportion of the population is affected, the probability of population persistence will be altered. We used data from long-term telemetry studies of female elephant seals at Macquarie Island, Australia, to model the effect of behaviour on the seals’ health (i.e. all internal factors that affect homeostasis). Through simulation, we investigated the effect of increasing periods of behavioural shifts, quantifying how the exclusion of maternal southern elephant seals from foraging habitat may affect their health, offspring survival, individual fitness and population growth rate. A long period of altered behaviour (>50% of an average foraging trip at sea) in 1 yr resulted in a small (0.4%) decline in population size the following year. However, a persistent disruption (e.g. 30 yr), caused for example by the long-term effects of climate change, could result in a 0.3% decline in individual fitness and a 10% decline in population size. Our approach to estimating the long-term population effects of short-term changes in individual behaviour can be generalised to include physiological effects and other causes of behavioural and physiological disruption, such as anthropogenic disturbance, for any species. Publisher PDF

Marine aerial surveys are designed to maximize the potential for detecting target species. Collecting data on different taxa from the same platform is economically advantageous but normally comes at the cost of compromising optimal taxon-specific scanning patterns and survey parameters, in particular altitude. Here, we describe simultaneous visual and photographic sampling methods as a proof of concept for detecting large whales and turtles from a single aircraft, despite very different sighting cues. Data were collected for fishing gear, fish, sharks, turtles, seals, dolphins, and whales using two observers and automated vertical photography. The photographic method documented an area directly beneath the aircraft that would otherwise have been obscured from observers. Preliminary density estimates were calculated for five species for which there were sufficient sample sizes from both methods after an initial year of data collection. The photographic method yielded significantly higher mean density estimates for loggerhead turtles, ocean sunfish, and blue sharks (p < 0.01), despite sampling a substantially smaller area than visual scanning (less than 11%). Density estimates from these two methods were not significantly different for leatherback turtles or basking sharks (p > 0.05), two of the largest species included in the analysis, which are relatively easy to detect by both methods. Although postflight manual processing of photographic data was extensive, this sampling method comes at no additional in-flight effort and obtains high-quality digital documentation of sightings on the trackline. Future directions for this project include automating photographic sighting detections, expanding the area covered by photography, and performing morphometric measurement assessments.

1 New England Aquarium, Boston, MA, USA, 2 Graduate School of Oceanography, University of Rhode Island, Narragansett, RI, USA, 3 Right Whale Ecology Program, Provincetown Center for Coastal Studies, Provincetown, MA, USA, 4 Biology and Marine Biology, University of North Carolina, Wilmington, NC, USA, 5 Biology Department, Woods Hole Oceanographic Institution, Woods Hole, MA, USA, 6 Nicholas School of the Environment and Pratt School of Engineering, Duke University

Right whales are vulnerable to many sources of anthropogenic disturbance including ship strikes, entanglement with fishing gear, and anthropogenic noise. The effect of these factors on individual health is unclear. A statistical model using photographic evidence of health was recently built to infer the true or hidden health of individual right whales. However, two important prior assumptions about the role of missing data and unexplained variance on the estimates were not previously assessed. Here we tested these factors by varying prior assumptions and model formulation. We found sensitivity to each assumption and used the output to make guidelines on future model formulation.

Expert elicitation is a rigorous method for synthesizing expert knowledge to inform decision making and is reliable and practical when field data are limited. We evaluated the feasibility of applying expert elicitation to estimate population-level effects of disturbance on marine mammals. Diverse experts estimated parameters related to mortality and sublethal injury of North Atlantic right whales (Eubalaena glacialis). We are now eliciting expert knowledge on the movement of right whales among geographic regions to parameterize a spatial model of health. Expert elicitation complements methods such as simulation models or extrapolations from other species, sometimes with greater accuracy and less uncertainty.

Entanglement in non-mobile fishing gear has been identified as one of the leading causes of mortality in North Atlantic right whales Eubalaena glacialis. To investigate this issue further, all available photographs of right whales taken from 1980 to 2009 were examined for evidence of entanglement with gear used in fisheries based on presence of rope or netting on the whale or scars inferred to have been caused by an encounter with rope. Photographs of 626 individual whales were assessed and 1032 unique entanglement events were documented. Of the 626 animals, 519 (82.9%) had been entangled at least once and 306 of the 519 (59.0%) had been entangled more than once. Males and females were entangled at similar rates. Juveniles were entangled at a higher rate than adults. On average, 25.9% of adequately photographed animals acquired new wounds or scars from fishing gear annually with no significant trend over time detected. However, the annual percentage of animals observed with rope on the body increased significantly during the study period, suggesting that it is becoming more difficult for whales to free themselves completely from fishing gear. Such high annual rates of entanglement remain a serious conservation concern for right whales because entanglements can have both lethal and sub-lethal effects. Federally required changes to fixed-gear fisheries in US waters have not reduced serious injuries and mortality to legally required levels. Here we show how documenting various annual rates of entanglement can monitor progress and impacts that fishing gear regula- tions may have on right whale recovery.

Baleen whales ( Mysticeti ) communicate using low-frequency acoustic signals. These long-wavelength sounds can be detected over hundreds of kilometres, potentially allowing contact over large distances. Low-frequency noise from large ships (20–200 Hz) overlaps acoustic signals used by baleen whales, and increased levels of underwater noise have been documented in areas with high shipping traffic. Reported responses of whales to increased noise include: habitat displacement, behavioural changes and alterations in the intensity, frequency and intervals of calls. However, it has been unclear whether exposure to noise results in physiological responses that may lead to significant consequences for individuals or populations. Here, we show that reduced ship traffic in the Bay of Fundy, Canada, following the events of 11 September 2001, resulted in a 6 dB decrease in underwater noise with a significant reduction below 150 Hz. This noise reduction was associated with decreased baseline levels of stress-related faecal hormone metabolites (glucocorticoids) in North Atlantic right whales ( Eubalaena glacialis ). This is the first evidence that exposure to low-frequency ship noise may be associated with chronic stress in whales, and has implications for all baleen whales in heavy ship traffic areas, and for recovery of this endangered right whale population.

The abundance of New Zealand subantarctic southern right whales (Eubalaena australis) was estimated for the first time using mark-recapture methods based on photo-identification and microsatellite genotyping (13 loci). Individual identification photographs of 383 whales and microsatellite genotypes of 235 whales were collected during annual austral winter field surveys from 1995 to 1998. Given the 4-year survey period and lack of geographic and demographic closure, we estimated super-population abundance using the POPAN Jolly-Seber model implemented in the software programme MARK. Models with constant survivorship but time-varying capture probability and probability of entry into the population were the most suitable due to the survey design. This provided estimates of abundance in 1998 of 908 non-calf whales (95% C.L. = 755, 1,123) for the photo-identification and 910 non-calf whales (95% C.L. = 641, 1,354) for the microsatellite genotype data sets. The current estimate of 900 whales may represent less than 5% of the pre-whaling abundance in New Zealand waters.

Vessel strikes are the leading cause of mortality for the endangered North Atlantic right whale Eubalaena glacialis. Prior to a December 2008 implementation of a mandatory, seasonally based vessel-speed rule (10 knots, 18.5 km h–1) along the eastern US seaboard, voluntary recommended speeds and routes were established. We used Automatic Identification System (AIS) data to evaluate and compare the compliance rates between the mandatory and voluntary measures to protect right whales in the southeast US critical habitat area off Florida and Georgia during the winters from 2005–06 to 2008–09. Vessel compliance was significantly higher under mandatory versus voluntary recommended speed restrictions, with compliance rates of 75 and 16%, respectively. Average vessel speeds were slower under mandatory speed restrictions (10.5 knots, 19.6 km h–1) compared to voluntary recommended speed restrictions (14.5 knots, 26.9 km h–1). Although vessels only slow down when mandated, they change their routing voluntarily. Compliance rates with voluntary recommended routes steadily increased during this period, from 43% prior to rulemaking, to 52% during the first year, 84% in the second year, and 96% in the final year of the study. Combining reduced speeds with recommended routes reduces the probability of right whale mortality from ships by 71.9% from the pre-implementation period. These results support long-term implementation of both vessel-speed reduction and restricted vessel routes for the survival and recovery of the North Atlantic right whale.

Evaluating impacts of human activities on marine ecosystems is difficult when effects occur out of plain sight. Oil spill severity is often measured by the number of marine birds and mammals killed, but only a small fraction of carcasses are recovered. The Deepwater Horizon/BP oil spill in the Gulf of Mexico was the largest in the U.S. history, but some reports implied modest environmental impacts, in part because of a relatively low number (101) of observed marine mammal mortalities. We estimate historical carcass-detection rates for 14 cetacean species in the northern Gulf of Mexico that have estimates of abundance, survival rates, and stranding records. This preliminary analysis suggests that carcasses are recovered, on an average, from only 2% (range: 0–6.2%) of cetacean deaths. Thus, the true death toll could be 50 times the number of carcasses recovered, given no additional information. We discuss caveats to this estimate, but present it as a counterpoint to illustrate the magnitude of misrepresentation implicit in presenting observed carcass counts without similar qualification. We urge methodological development to develop appropriate multipliers. Analytical methods are required to account explicitly for low probability of carcass recovery from cryptic mortality events (e.g., oil spills, ship strikes, bycatch in unmonitored fisheries and acoustic trauma).