Spatial and temporal analysis of killer whale (Orcinus orca) strandings in the North Pacific Ocean and the benefits of a coordinated stranding response protocol

Killer whales (Orcinus orca) are widely distributed throughout the world's oceans, yet little has been documented about their stranding patterns. Knowledge of stranding patterns improves our ability to examine and sample carcasses and provides a foundation for understanding killer whale natural history, diet, reproduction, anthropogenic stressors, emerging diseases, and patterns of unusual mortality. We compiled published and unpublished killer whale stranding data to describe stranding patterns in the North Pacific Ocean. Between 1925 and 2011, 371 stranded killer whales were reported in Japan (20.4%), Russia (3.5%), Alaska (32.0%), British Columbia (27.4%), Washington (4.0%), Oregon (2.7%), California (5.1%), Mexico (3.8%), and Hawaii (0.8%). Strandings occurred at all times of year, but regionally specific seasonal differences were observed. Mortality and annual census data from Northern and Southern Resident populations were extrapolated to estimate that across the North Pacific, an average of 48 killer whales die annually. However, over the last two decades, an average of only 10 killer whale carcasses were recovered annually in this ocean, making each event a rare opportunity for study. Publication of a standardized killer whale necropsy protocol and dedicated funding facilitated the number of complete postmortem necropsies performed on stranded killer whales from 1.6% to 32.2% annually.     

Review of cetaceans stranded on the Irish coast, 1901–95

Published records of cetaceans stranded on the Irish coast during the period 1901–95 are reviewed. In this review the number of stranding events has been used in the analysis and includes both live strandings and those animals washed up dead. There were 529 records involving 21 species. The Harbour Porpoise (27%) was the most frequently reported species, followed by Common Dolphins (16%) and Pilot Whales (15%). Minke Whales (8%) were the most frequently reported mysticete. The number of reported strandings has increased since the 1960s which is thought to be mainly due to increased observer effort. Cetaceans have stranded on all Irish coasts but mainly along the south coast and along the western seaboard but with no apparent overall seasonal trend. There was a peak in the strandings of Common Dolphins during 1991–92 when 27% (28 records) of all strandings were reported and of White-sided Dolphins when 60% (28 records) were reported, both of which were attributed to possible interactions with fisheries. The number of Striped Dolphins stranded on the Irish coast has increased steadily since the 1980s and may reflect increasing water temperatures.
These stranding records are considered inadequate to determine the status of most species of cetaceans in Irish waters but are sufficient to identify unusual stranding events such as high mortalities due to fisheries interactions or epizootics. More observer coverage is required before the stranding data are adequate for monitoring the status of most species but a stranding scheme is considered the most effective and efficient method of long-term monitoring of cetaceans in Irish waters.     

Recovery rates of bottlenose dolphin (Tursiops truncatus) carcasses estimated from stranding and survival rate data

Recovery of cetacean carcasses provides data on levels of human‐caused mortality, but represents only a minimum count of impacts. Counts of stranded carcasses are negatively biased by factors that include at‐sea scavenging, sinking, drift away from land, stranding in locations where detection is unlikely, and natural removal from beaches due to wave and tidal action prior to detection. We estimate the fraction of carcasses recovered for a population of coastal bottlenose dolphins (Tursiops truncatus), using abundance and survival rate data to estimate annual deaths in the population. Observed stranding numbers are compared to expected deaths to estimate the fraction of carcasses recovered. For the California coastal population of bottlenose dolphins, we estimate the fraction of carcasses recovered to be 0.25 (95% CI = 0.20–0.33). During a 12 yr period, 327 animals (95% CI = 253–413) were expected to have died and been available for recovery, but only 83 carcasses attributed to this population were documented. Given the coastal habits of California coastal bottlenose dolphins, it is likely that carcass recovery rates of this population greatly exceed recovery rates of more pelagic dolphin species in the region.     

Were Multiple Stressors a ‘Perfect Storm’ for Northern Gulf of Mexico Bottlenose Dolphins (Tursiops truncatus) in 2011?

An unusual number of near term and neonatal bottlenose dolphin (Tursiops truncatus) mortalities occurred in the northern Gulf of Mexico (nGOM) in 2011, during the first calving season after two well documented environmental perturbations; sustained cold weather in 2010 and the Deepwater Horizon oil spill (DWHOS). Preceding the stranding event, large volumes of cold freshwater entered the nGOM due to unusually large snowmelt on the adjacent watershed, providing a third potential stressor. We consider the possibility that this extreme cold and freshwater event contributed to the pattern of perinatal dolphin strandings along the nGOM coast. During the 4-month period starting January 2011, 186 bottlenose dolphins, including 46% perinatal calves (nearly double the percentage for the same time period from 2003-2010) washed ashore from Louisiana to western Florida. Comparison of the frequency distribution of strandings to flow rates and water temperature at a monitoring buoy outside Mobile Bay, Alabama (the 4(th) largest freshwater drainage in the U.S.) and along the nGOM coast showed that dolphin strandings peaked in Julian weeks 5, 8, and 12 (February and March), following water temperature minima by 2-3 weeks. If dolphin condition was already poor due to depleted food resources, bacterial infection, or other factors, it is plausible that the spring freshet contributed to the timing and location of the unique stranding event in early 2011. These data provide strong observational evidence to assess links between the timing of the DWHOS, other local environmental stressors, and mortality of a top local predator. Targeted analyses of tissues from stranded dolphins will be essential to define a cause of death, and our findings highlight the importance of considering environmental data along with biological samples to interpret stranding patterns during and after an unusual mortality event.     

How much are stranding records affected by variation in reporting rates? A case study of small delphinids in the Bay of Biscay

Marine vertebrate strandings offer an opportunistic sampling scheme that can provide abundant data over long periods. Because the stranding process involves biological, physical and sociological parameters, confounding complicates the interpretation of results. The statistical analysis of these data relies on generalized linear or additive models in order to infer long-term trends, but does not easily account for drift or variation in reporting rates. Here, we capitalized on county-level (administrative) variation following the passing of a law for compulsory reporting of stranded marine mammals in France to investigate how variation in reporting rates may affect the observed trend in stranded small delphinids in the Bay of Biscay. Using a time-series spanning more than 30 years across eight administrative counties, we built variance partitioning models for the analysis of count data. We discussed the choice of an appropriate likelihood to conclude the Negative Binomial useful and interpretable in the context of small delphinid strandings. We expanded the model with a recent methodology to detect structural breaks in the time series, focusing on overdispersion. We performed statistical robustness checks with respect to variations in reporting rates and discuss their causal interpretation in the context of observational data. Stranding frequencies increased on average 7-fold over 30 years. We conclude that reporting rates to the French stranding network have been stable since the early 1990s, and the average 3-fold increase in stranded small delphinids observed in the Bay of Biscay since 1990 is due to other factors, including bycatch. Codes and data are available to replicate the analysis to other national stranding networks.     

Factors Affecting Harp Seal (Pagophilus groenlandicus) Strandings in the Northwest Atlantic

The effects of climate change on high latitude regions are becoming increasingly evident, particularly in the rapid decline of sea ice cover in the Arctic. Many high latitude species dependent on sea ice are being forced to adapt to changing habitats. Harp seals (Pagophilus groenlandicus) are an indicator species for changing high-latitude ecosystems. This study analyzed multiple factors including ice cover, demographics, and genetic diversity, which could affect harp seal stranding rates along the eastern coast of the United States. Ice cover assessments were conducted for the month of February in the Gulf of St. Lawrence whelping region from 1991–2010 using remote sensing data, and harp seal stranding data were collected over the same time period. Genetic diversity, which may affect how quickly species can adapt to changing climates, was assessed using ten microsatellite markers to determine mean d ² in a subset of stranded and by-caught (presumably healthy) seals sampled along the northeast U.S. coast. Our study found a strong negative correlation (R ² = 0.49) between ice cover in the Gulf of St. Lawrence and yearling harp seal strandings, but found no relationship between sea ice conditions and adult strandings. Our analysis revealed that male seals stranded more frequently than females during the study period and that this relationship was strongest during light ice years. In contrast, we found no significant difference in mean d ² between stranded and by-caught harp seals. The results demonstrate that sea ice cover and demographic factors have a greater influence on harp seal stranding rates than genetic diversity, with only a little of the variance in mean d ² among stranded seals explained by ice cover. Any changes in these factors could have major implications for harp seals, and these findings should be considered in the development of future management plans for the Arctic that incorporate climate variability.     

Hearing loss in stranded odontocete dolphins and whales

The causes of dolphin and whale stranding can often be difficult to determine. Because toothed whales rely on echolocation for orientation and feeding, hearing deficits could lead to stranding. We report on the results of auditory evoked potential measurements from eight species of odontocete cetaceans that were found stranded or severely entangled in fishing gear during the period 2004 through 2009. Approximately 57% of the bottlenose dolphins and 36% of the rough-toothed dolphins had significant hearing deficits with a reduction in sensitivity equivalent to severe (70-90 dB) or profound (>90 dB) hearing loss in humans. The only stranded short-finned pilot whale examined had profound hearing loss. No impairments were detected in seven Risso's dolphins from three different stranding events, two pygmy killer whales, one Atlantic spotted dolphin, one spinner dolphin, or a juvenile Gervais' beaked whale. Hearing impairment could play a significant role in some cetacean stranding events, and the hearing of all cetaceans in rehabilitation should be tested.     

RATES AND POTENTIAL CAUSES OF MORTALITY IN NORTH ATLANTIC RIGHT WHALES (EUBALAENA GLACIALIS)

North Atlantic right whale mortality rates range from 2% to 17% over the first four years of life. Sources of mortality in this population include ship collisions, entanglements, and natural causes. A combined analysis of stranding data, entanglement records, and photographic information indicates that approximately one third of all right whale mortality is caused by human activities. Anthropogenic sources of right whale mortality may be a significant factor inhibiting growth in the North Atlantic population.     

Risk analysis reveals global hotspots for marine debris ingestion by sea turtles

Plastic marine debris pollution is rapidly becoming one of the critical environmental concerns facing wildlife in the 21st century. Here we present a risk analysis for plastic ingestion by sea turtles on a global scale. We combined global marine plastic distributions based on ocean drifter data with sea turtle habitat maps to predict exposure levels to plastic pollution. Empirical data from necropsies of deceased animals were then utilised to assess the consequence of exposure to plastics. We modelled the risk (probability of debris ingestion) by incorporating exposure to debris and consequence of exposure, and included life history stage, species of sea turtle and date of stranding observation as possible additional explanatory factors. Life history stage is the best predictor of debris ingestion, but the best‐fit model also incorporates encounter rates within a limited distance from stranding location, marine debris predictions specific to the date of the stranding study and turtle species. There is no difference in ingestion rates between stranded turtles vs. those caught as bycatch from fishing activity, suggesting that stranded animals are not a biased representation of debris ingestion rates in the background population. Oceanic life‐stage sea turtles are at the highest risk of debris ingestion, and olive ridley turtles are the most at‐risk species. The regions of highest risk to global sea turtle populations are off of the east coasts of the USA, Australia and South Africa; the east Indian Ocean, and Southeast Asia. Model results can be used to predict the number of sea turtles globally at risk of debris ingestion. Based on currently available data, initial calculations indicate that up to 52% of sea turtles may have ingested debris.     

Social and demographic effects of anthropogenic mortality: a test of the compensatory mortality hypothesis in the red wolf

Whether anthropogenic mortality is additive or compensatory to natural mortality in animal populations has long been a question of theoretical and practical importance. Theoretically, under density-dependent conditions populations compensate for anthropogenic mortality through decreases in natural mortality and/or increases in productivity, but recent studies of large carnivores suggest that anthropogenic mortality can be fully additive to natural mortality and thereby constrain annual survival and population growth rate. Nevertheless, mechanisms underlying either compensatory or additive effects continue to be poorly understood. Using long-term data on a reintroduced population of the red wolf, we tested for evidence of additive vs. compensatory effects of anthropogenic mortality on annual survival and population growth rates, and the preservation and reproductive success of breeding pairs. We found that anthropogenic mortality had a strong additive effect on annual survival and population growth rate at low population density, though there was evidence for compensation in population growth at high density. When involving the death of a breeder, anthropogenic mortality was also additive to natural rates of breeding pair dissolution, resulting in a net decrease in the annual preservation of existing breeding pairs. However, though the disbanding of a pack following death of a breeder resulted in fewer recruits per litter relative to stable packs, there was no relationship between natural rates of pair dissolution and population growth rate at either high or low density. Thus we propose that short-term additive effects of anthropogenic mortality on population growth in the red wolf population at low density were primarily a result of direct mortality of adults rather than indirect socially-mediated effects resulting in reduced recruitment. Finally, we also demonstrate that per capita recruitment and the proportion of adults that became reproductive declined steeply with increasing population density, suggesting that there is potential for density-dependent compensation of anthropogenically-mediated population regulation.     

Managing multiple vital rates to maximize greater sage-grouse population growth

Despite decades of field research on greater sage-grouse, range-wide demographic data have yet to be synthesized into a sensitivity analysis to guide management actions. We reviewed range-wide demographic rates for greater sage-grouse from 1938 to 2011 and used data from 50 studies to parameterize a 2-stage, female-based population matrix model. We conducted life-stage simulation analyses to determine the proportion of variation in population growth rate (λ) accounted for by each vital rate, and we calculated analytical sensitivity, elasticity, and variance-stabilized sensitivity to identify the contribution of each vital rate to λ. As expected for an upland game bird, greater sage-grouse showed marked annual and geographic variation in several vital rates. Three rates were demonstrably important for population growth: female survival, chick survival, and nest success. Female survival and chick survival, in that order, had the most influence on λ per unit change in vital rates. However, nest success explained more of the variation in λ than did the survival rates. In lieu of quantitative data on specific mortality factors driving local populations, we recommend that management efforts for greater sage-grouse first focus on increasing female survival by restoring large, intact sagebrush-steppe landscapes, reducing persistent sources of human-caused mortality, and eliminating anthropogenic habitat features that subsidize species that prey on juvenile, yearling, and adult females. Our analysis also supports efforts to increase chick survival and nest success by eliminating anthropogenic habitat features that subsidize chick and nest predators, and by managing shrub, forb, and grass cover, height, and composition to meet local brood-rearing and nesting habitat guidelines. We caution that habitat management to increase chick survival and nest success should not reduce the cover or height of sagebrush below that required for female survival in other seasons (e.g., fall, winter). The success or failure of management actions for sage-grouse should be assessed by measuring changes in vital rates over long time periods to avoid confounding with natural, annual variation. © 2011 The Wildlife Society.     

Frequency,Magnitude, and Possible Causes of Stranding and Mass-Mortality Events of the Beach Clam Tivela mactroides (Bivalvia: Veneridae)

Stranding combined with mass-mortality events of sandy-beach organisms is a frequent but little-understood phenomenon, which is generally studied based on discrete episodes. The frequency, magnitude, and possible causes of stranding and mass-mortality events of the trigonal clam Tivela mactroides were assessed based on censuses of stranded individuals, every four days from September 2007 through December 2008, in Caraguatatuba Bay, southeastern Brazil. Stranded clams were classified as dying (closed valves did not open when forced) or dead (closed valves were easily opened). Information on wave parameters and the living intertidal clam population was used to assess possible causes of stranding. This fine-scale monitoring showed that stranding occurred widely along the shore and year-round, with peaks interspersed with periods of low or no mortality. Dead clams showed higher mean density than dying individuals, but a lower mean shell length, attributed to a higher tolerance to desiccation of larger individuals. Wave height had a significant negative relationship to the density of dying individuals, presumed to be due to the accretive nature of low-energy waves: when digging out, clams would be more prone to be carried upward and unable to return; while larger waves, breaking farther from the beach and with a stronger backwash, would prevent stranding in the uppermost areas. This ecological finding highlights the need for refined temporal studies on mortality events, in order to understand them more clearly. Last, the similar size structure of stranded clams and the living population indicated that the stranded individuals are from the intertidal or shallow subtidal zone, and reinforces the ecological and behavioral components of this process, which have important ecological and socioeconomic implications for the management of this population.     

The application of GIS and spatiotemporal analyses to investigations of unusual marine mammal strandings and mortality events

In 2006–2007, an unusually high number of harbor porpoises (Phocoena phocoena) stranded along the Washington and Oregon coastlines. Spatiotemporal analyses were used to examine their ability to detect clusters of porpoise strandings during an unusual mortality event (UME) in the Pacific Northwest using stranding location data. Strandings were evaluated as two separate populations, outer coast and inland waters. The presence of global clustering was evaluated using the Knox spatiotemporal test, and the presence of local clusters was investigated using a spatiotemporal scan statistic (space–time permutation). There was evidence of global clustering, but no local clustering, supporting the hypothesis that strandings were due to more varied etiologies instead of localized causes. Further analyses at subregional levels, and concurrently assessing environmental factors, might reveal additional geographic distribution patterns. This article describes the spatial analytical tools applied in this study and how they can help elucidate the spatiotemporal epidemiology of other UMEs and assist in determining their causes. More than one spatial analytical technique should be used if the study objective is to detect and describe clustering in time and space and to generate hypotheses regarding causation of marine mammal disease and stranding events.     

A MODEL LIFE TABLE FOR BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) FROM THE INDIAN RIVER LAGOON SYSTEM, FLORIDA, U.S.A.

Data gathered from 220 stranded bottlenose dolphins ( Tursiops truncatus ) in the Indian River Lagoon system, Florida, were used to derive a life table. Survivorship curves were fit to the data using Siler's competing-risk model and a maximum likelihood approach. Population growth was estimated to be between r = 0.0 and 0.046 based on the observed numbers of stranded dolphins. Variance in survival rates was estimated using an individual-based, age-structured population projection model. We estimate that the overall annual mortality rate for this population was 9.8% per year. Sex-specific differences in survivorship were apparent with females outliving males. The overall mortality curve resembles that of other large mammals, with high calf mortality and an exponentially increasing risk of senescent mortality. The inclusion of live-capture removals of individuals from this population did not significantly affect the estimation of survival parameters for most age classes.     

Predicted distributions and abundances of the sea turtle ‘lost years’ in the western North Atlantic Ocean

Oceanic dispersal characterizes the early juvenile life-stages of numerous marine species of conservation concern. This early stage may be a ‘critical period’ for many species, playing an overriding role in population dynamics. Often, relatively little information is available on their distribution during this period, limiting the effectiveness of efforts to understand environmental and anthropogenic impacts on these species. Here we present a simple model to predict annual variation in the distribution and abundance of oceanic-stage juvenile sea turtles based on species’ reproductive output, movement and mortality. We simulated dispersal of 25 cohorts (1993–2017) of oceanic-stage juveniles by tracking the movements of virtual hatchling sea turtles released in a hindcast ocean circulation model. We then used estimates of annual hatchling production from Kemp's ridley Lepidochelys kempii (n = 3), green Chelonia mydas (n = 8) and loggerhead Caretta caretta (n = 5) nesting areas in the northwestern Atlantic (inclusive of the Gulf of Mexico, Caribbean Sea and eastern seaboard of the U.S.) and their stage-specific mortality rates to weight dispersal predictions. The model's predictions indicate spatial heterogeneity in turtle distribution across their marine range, identify locations of increasing turtle abundance (notably along the U.S. coast), and provide valuable context for temporal variation in the stranding of young sea turtles across the Gulf of Mexico. Further effort to collect demographic, distribution and behavioral data that refine, complement and extend the utility of this modeling approach for sea turtles and other dispersive marine taxa is warranted. Finally, generating these spatially-explicit predictions of turtle abundance required extensive international collaboration among scientists; our findings indicate that continued conservation of these sea turtle populations and the management of the numerous anthropogenic activities that operate in the northwestern Atlantic Ocean will require similar international coordination.     

Patterns of mortality in endangered Cook Inlet beluga whales: Insights from pairing a long-term photo-identification study with stranding records

Mortality is a demographic metric crucial for understanding the dynamics of endangered populations such as Cook Inlet beluga whales (CIBWs, Delphinapterus leucas), but patterns of mortality are currently not well understood for CIBWs, making decisions about recovery actions challenging. We combined long-term photo-ID data from approximately 420 individual belugas identified during the period 2005–2017 with stranding data from 95 dead belugas to identify patterns of mortality with respect to age, sex, geographic range, cause of death, and to estimate minimum mortality rates. Reported mortality was greatest for adults of reproductive age, followed by calves, with fewer subadults and no adults older than 49 years in the stranding data set despite lifespans of 70+ years reported in other beluga populations. Dead females and males were evenly represented. Live stranding was the predominant assigned cause of death but represented only ~33% of deaths of known cause. Causal factors for the majority of deaths and live strandings are unknown. Annual mortality estimated from reported carcasses relative to total population size averaged 2.2%. Our analysis advances our current understanding of mortality patterns in CIBWs but linking a greater proportion of carcasses to photo-ID individuals would further improve our understanding; we conclude with recommendations for achieving this.     

Strandings provide insight into social group structure of Atlantic white-sided dolphins

Atlantic white-sided dolphins (Lagenorhynchus acutus) are highly social odontocetes with a poorly understood tendency to mass strand. With limited capacity to study social ecology in the open ocean, mass strandings provide an opportunity to improve our understanding of group structure. Our study of 32 mass stranding events that occurred on Cape Cod, Massachusetts, between 1999 and 2009 identifies aspects of social ecology that vary across the year. A greater number of mass stranding events occurred outside of the breeding season and there was evidence of age-structuring during the breeding season. We find generally low average intragroup pairwise relatedness assessed across eight microsatellite loci in a subset of 16 mass stranding events. Mass stranded groups do not show higher than expected relatedness when compared to a baseline estimate derived from single-stranded individuals. Overall, our integration of genetic estimates of relatedness with data on sex and maturity-class from stranded specimens suggests that Atlantic white-sided dolphins fall near the more fluid end of the continuum from short-term, highly fluid social associations to long-term, stable groups represented among the odontocetes. Despite their tendency to mass strand, stable, kin-based associations are not a defining feature of social group structure in this species.     

Estimating At-Sea Mortality of Marine Turtles from Stranding Frequencies and Drifter Experiments

Strandings of marine megafauna can provide valuable information on cause of death at sea. However, as stranding probabilities are usually very low and highly variable in space and time, interpreting the results can be challenging. We evaluated the magnitude and distribution of at-sea mortality of marine turtles along the Pacific coast of Baja California Sur, México during 2010–11, using a combination of counting stranded animals and drifter experiments. A total of 594 carcasses were found during the study period, with loggerhead (62%) and green turtles (31%) being the most common species. 87% of the strandings occurred in the southern Gulf of Ulloa, a known hotspot of loggerhead distribution in the Eastern Pacific. While only 1.8% of the deaths could be definitively attributed to bycatch (net marks, hooks), seasonal variation in stranding frequencies closely corresponded to the main fishing seasons. Estimated stranding probabilities from drifter experiments varied among sites and trials (0.05–0.8), implying that only a fraction of dead sea turtles can be observed at beaches. Total mortality estimates for 15-day periods around the floater trials were highest for PSL, a beach in the southern Gulf of Ulloa, ranging between 11 sea turtles in October 2011 to 107 in August 2010. Loggerhead turtles were the most numerous, followed by green and olive ridley turtles. Our study showed that drifter trials combined with beach monitoring can provide estimates for death at sea to measure the impact of small-scale fisheries that are notoriously difficult to monitor for by-catch. We also provided recommendations to improve the precision of the mortality estimates for future studies and highlight the importance of estimating impacts of small–scale fisheries on marine megafauna.     

What can cetacean stranding records tell us? A study of UK and Irish cetacean diversity over the past 100 years

There are many factors that may explain why cetaceans (whales, dolphins, and porpoises) strand. Around the UK and Ireland, over 20,000 stranding records have been collected since 1913, resulting in one of the longest, continuous, systematic stranding data sets in the world. We use this data set to investigate temporal and spatial trends in cetacean strandings and use generalized additive models (GAMs) to investigate correlates of strandings. We find a dramatic increase in strandings since the 1980s, most likely due to increases in recording effort, and the formation of formal strandings networks. We found no correlation between the numbers of cetaceans stranding each year and several potential environmental and anthropogenic predictors: storms, geomagnetic activity, North Atlantic Oscillations, sea‐surface temperature, and fishing catch. We suggest that this is because the scale of change in the variables is too coarse to detect any potential correlations. It may also highlight the idiosyncratic nature of species’ responses to external pressures, and further the need to investigate other potential correlates of strandings, such as bycatch and military sonar. Long‐term cetacean stranding data provide vital information on past and present diversity for common, rare, and inconspicuous species. This study underlines the importance of continued support for stranding networks.