Amy R. Knowlton

Gross and histologic evidence of sharp and blunt trauma in North Atlantic right whales (Eubalaena glacialis) killed by vessels

Abstract Vessel-whale collision events represented the ultimate cause of death for 21 (52.5%) of the 40 North Atlantic right whales (Eubalaena glacialis) necropsied between 1970 and December 2006. Injuries seen in vessel-struck whales fall into two distinct categories: 1) sharp trauma, often resulting from contact with the propeller, and 2) blunt trauma, presumably resulting from contact with a vessels hull. This study analyzes four trauma cases that resulted from vessel-whale collisions, which together provide a framework for a more critical understanding of lethal blunt and sharp trauma resulting from vessel collisions with right whales. In case no. 1, contact with a propeller resulted in three deep lacerations. The animal survived acute trauma only to succumb nearly 14 years later when the lesions reopened and became infected. In case no. 2, anecdotal reports linked the laceration of large arteries of the peduncle and histologic evidence of perimortem trauma at a bone fracture site to vessel-whale collision trauma. Case no. 3 had a laceration of the oral rete and a fracture of the rostrum. Both of the areas displayed histologic evidence of perimortem blunt trauma. Finally, in case no. 4, an antemortem mandibular fracture, two additional skull fractures, and widespread hemorrhage were consistent with severe blunt trauma. Evidence from each case, including the timing of trauma relative to the time of death and identifying characteristics of both trauma types, are presented. Before this study, no detailed comparative analysis of trauma pathology that resulted from lethal interactions between vessels and right whales had been conducted. This study demonstrates the importance of detailed gross and histologic examination in determining the significance and timing of traumatic events. This work represents a new paradigm for the differential diagnosis of lethal sharp and blunt trauma in right whales hit by ships and will enhance the present understanding of the impact of anthropogenic mortality on this critically endangered species.

Population histories of right whales (Cetacea: Eubalaena) inferred from mitochondrial sequence diversities and divergences of their whale lice (Amphipoda: Cyamus)

Right whales carry large populations of three ‘whale lice’ (Cyamus ovalis, Cyamus gracilis, Cyamus erraticus) that have no other hosts. We used sequence variation in the mitochondrial COI gene to ask (i) whether cyamid population structures might reveal associations among right whale individuals and subpopulations, (ii) whether the divergences of the three nominally conspecific cyamid species on North Atlantic, North Pacific, and southern right whales (Eubalaena glacialis, Eubalaena japonica, Eubalaena australis) might indicate their times of separation, and (iii) whether the shapes of cyamid gene trees might contain information about changes in the population sizes of right whales. We found high levels of nucleotide diversity but almost no population structure within oceans, indicating large effective population sizes and high rates of transfer between whales and subpopulations. North Atlantic and Southern Ocean populations of all three species are reciprocally monophyletic, and North Pacific C. erraticus is well separated from North Atlantic and southern C. erraticus. Mitochondrial clock calibrations suggest that these divergences occurred around 6 million years ago (Ma), and that the Eubalaena mitochondrial clock is very slow. North Pacific C. ovalis forms a clade inside the southern C. ovalis gene tree, implying that at least one right whale has crossed the equator in the Pacific Ocean within the last 1–2 million years (Myr). Low‐frequency polymorphisms are more common than expected under neutrality for populations of constant size, but there is no obvious signal of rapid, interspecifically congruent expansion of the kind that would be expected if North Atlantic or southern right whales had experienced a prolonged population bottleneck within the last 0.5 Myr.

Patterns of male reproductive success in a highly promiscuous whale species: the endangered North Atlantic right whale

Parentage analyses of baleen whales are rare, and although mating systems have been hypothesized for some species, little data on realized male reproductive success are available and the patterns of male reproductive success have remained elusive for most species. Here we combine over 20 years of photo‐identification data with high‐resolution genetic data for the majority of individual North Atlantic right whales to assess paternity in this endangered species. There was significant skew in male reproductive success compared to what would be expected if mating was random (P < 0.001). The difference was due to an excess of males assigned zero paternities, a deficiency of males assigned one paternity, and an excess of males assigned as fathers for multiple calves. The variance in male reproductive success was high relative to other aquatically mating marine mammals, but was low relative to mammals where the mating system is based on resource‐ and/or mate‐defence polygyny. These results are consistent with previous data suggesting that the right whale mating system represents one of the most intense examples of sperm competition in mammals, but that sperm competition on its own does not allow for the same degree of polygyny as systems where males can control access to resources and/or mates. The age distribution of assigned fathers was significantly biased towards older males (P < 0.05), with males not obtaining their first paternity until ~15 years of age, which is almost twice the average age of first fertilization in females (8 years), suggesting that mate competition is preventing younger males from reproducing. The uneven distribution of paternities results in a lower effective population size in this species that already has one of the lowest reported levels of genetic diversity, which may further inhibit reproductive success through mate incompatibility of genetically similar individuals.

Fatally entangled right whales can die extremely slowly

Unlike smaller marine mammals that lack the mass and power to break free from serious entanglements in fixed fishing gear, right whales can do so, but they are not always rope free. The remaining rope can gradually constrict one or more body parts and the resulting debilitation and ultimate death can take many months. Thus the practices that lead to these mortalities need to be viewed not only as a conflict between the cultural and socioeconomic value of a fishery versus a potential species extinction process, but also in terms of an extreme animal welfare issue

Using hierarchical bayes to understand movement, health, and survival in the endangered north atlantic right whale.

Body condition is an indicator of health, and it plays a key role in many vital processes for mammalian species. While evidence of individual body condition can be obtained, these observations provide just brief glimpses into the health state of the animal. An analytical framework is needed for understanding how health of animals changes over space and time.Through knowledge of individual health we can better understand the status of populations. This is particularly important in endangered species, where the consequences of disruption of critical biological functions can push groups of animals rapidly toward extinction. Here we built a state-space model that provides estimates of movement, health, and survival. We assimilated 30+ years of photographic evidence of body condition and three additional visual health parameters in individual North Atlantic right whales, together with survey data, to infer the true health status as it changes over space and time. We also included the effect of reproductive status and entanglement status on health. At the population level, we estimated differential movement patterns in males and females. At the individual level, we estimated the likely animal locations each month. We estimated the relationship between observed and latent health status. Observations of body condition, skin condition, cyamid infestation on the blowholes, and rake marks all provided measures of the true underlying health. The resulting time series of individual health highlight both normal variations in health status and how anthropogenic stressors can affect the health and, ultimately, the survival of individuals. This modeling approach provides information for monitoring of health in right whales, as well as a framework for integrating observational data at the level of individuals up through the health status of the population. This framework can be broadly applied to a variety of systems – terrestrial and marine – where sporadic observations of individuals exist.

Effects of fishing rope strength on the severity of large whale entanglements

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.

Implications of fishing rope strength on the severity of large whale entanglements

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.

Sounds recorded in the presence of sei whale, Balaenoptera borealis

Opportunistic recordings were made on 12 days during the months of August and September in 1986–1989 between Browns and Baccari Banks on the Nova Scotian shelf. Sei whales, Balaenoptera borealis, were seen in the recording area on 16 of the 32 recording sessions. Subsequent signal analysis revealed that a distinctive sound type was recorded on all of the 16 recording sessions when sei whales were seen and on six sessions when no sei whales were seen. On some recordings two to three different sources (based on sound quality and received levels) are apparent. All these sounds share the same basic characteristics. The signal consists of two phrases, each lasting about 0.5–0.8 s, with an inter‐phrase interval of 0.4–1 s, and restricted to the 1.5–3.5 kHz band. Each phrase is composed of a series of 10–20 FM sweeps lasting approximately 30–40 ms/sweep. These signals are similar to the few sei whale sounds reported by Thompson et al. [in Behavior of Marine Animals, Vol. 3, Cetaceans, edited by H. E. Winn and B....

Characterizing the Duration and Severity of Fishing Gear Entanglement on a North Atlantic Right Whale (Eubalaena glacialis) Using Stable Isotopes, Steroid and Thyroid Hormones in Baleen

North Atlantic right whales (Eubalaena glacialis) are highly endangered and frequently exposed to a myriad of human activities and stressors in their industrialized habitat. Entanglements in fixed fishing gear represent a particularly pervasive and often drawn-out source of anthropogenic morbidity and mortality to the species. To better understand both the physiological response to entanglement, and to determine fundamental parameters such as acquisition, duration, and severity of entanglement, we measured a suite of biogeochemical markers in the baleen of an adult female that died from a well-documented chronic entanglement in 2005 (whale Eg2301). Steroid hormones (cortisol, corticosterone, estradiol, and progesterone), thyroid hormones (triiodothyronine (T3) and thyroxine (T4)), and stable isotopes (δ13C and δ15N) were all measured in a longitudinally sampled baleen plate. This yielded an 8-year profile of foraging and migration behavior, stress response, and reproduction. Stable isotopes cycled in annual patterns that reflect the animal’s north-south migration behavior and seasonally abundant zooplankton diet. A progesterone peak, lasting approximately 23 months, was associated with the single known calving event (in 2002) for this female. Estradiol, cortisol, corticosterone, T3, and T4 were also elevated, although variably so, during the progesterone peak. This whale was initially sighted with a fishing gear entanglement in September 2004, but the hormone panel suggests that the animal first interacted with the gear as early as June 2004. Elevated δ15N, T3, and T4 indicate that Eg2301 potentially experienced increased energy expenditure, significant lipid catabolism, and thermal stress approximately three months before the initial sighting with fishing gear. All hormones in the panel (except cortisol) were elevated above baseline by September 2004. This novel study illustrates the value of using baleen to reconstruct recent temporal profiles and as a comparative matrix in which key physiological indicators of individual whales can be used to understand the impacts of anthropogenic activity on threatened whale populations.

Predicting lethal entanglements as a consequence of drag from fishing gear

Large whales are frequently entangled in fishing gear and sometimes swim while carrying gear for days to years. Entangled whales are subject to additional drag forces requiring increased thrust power and energy expenditure over time. To classify entanglement cases and aid potential disentanglement efforts, it is useful to know how long an entangled whale might survive, given the unique configurations of the gear they are towing. This study establishes an approach to predict drag forces on fishing gear that entangles whales, and applies this method to ten North Atlantic right whale cases to estimate the resulting increase in energy expenditure and the critical entanglement duration that could lead to death. Estimated gear drag ranged 11-275N. Most entanglements were resolved before critical entanglement durations (mean±SD 216±260days) were reached. These estimates can assist real-time development of disentanglement action plans and U.S. Federal Serious Injury assessments required for protected species.