Attaching bio-telemetry or -logging devices ('tags') to marine animals for research and monitoring adds drag to streamlined bodies, thus affecting posture, swimming gaits and energy balance. These ...costs have never been measured in free-swimming cetaceans. To examine the effect of drag from a tag on metabolic rate, cost of transport and swimming behavior, four captive male dolphins (Tursiops truncatus) were trained to swim a set course, either non-tagged (n=7) or fitted with a tag (DTAG2; n=12), and surface exclusively in a flow-through respirometer in which oxygen consumption VO₂ and carbon dioxide production (VO₂; ml kg(-1) min(-1)) rates were measured and respiratory exchange ratio (VO₂/resting VO₂) was calculated. Tags did not significantly affect individual mass-specific oxygen consumption, physical activity ratios (exercise /resting ), total or net cost of transport (COT; J m(-1) kg(-1)) or locomotor costs during swimming or two-minute recovery phases. However, individuals swam significantly slower when tagged (by ~11%; mean ± s.d., 3.31±0.35 m s(-1)) than when non-tagged (3.73±0.41 m s(-1)). A combined theoretical and computational fluid dynamics model estimating drag forces and power exertion during swimming suggests that drag loading and energy consumption are reduced at lower swimming speeds. Bottlenose dolphins in the specific swimming task in this experiment slowed to the point where the tag yielded no increases in drag or power, while showing no difference in metabolic parameters when instrumented with a DTAG2. These results, and our observations, suggest that animals modify their behavior to maintain metabolic output and energy expenditure when faced with tag-induced drag.
Drag force acting on swimming marine mammals is difficult to measure directly. Researchers often use simple modeling and kinematic measurements from animals, or computational fluid dynamics (CFD) ...simulations to estimate drag. However, studies that compare these methods are lacking. Here, computational simulation and physical experiments were used to estimate drag forces on gliding bottlenose dolphins (Tursiops truncatus). To facilitate comparison, variable drag loading (no‐tag, tag, tag + 4, tag + 8) was used to increase force in both simulations and experiments. During the experiments, two dolphins were trained to perform controlled glides with variable loading. CFD simulations of dolphin/tag geometry in steady flow (1–6 m/s) were used to model drag forces. We expect both techniques will capture relative changes created by experimental conditions, but absolute forces predicted by the methods will differ. CFD estimates were within a calculated 90% confidence interval of the experimental results for all but the tag condition. Relative drag increase predicted by the simulation vs. experiment, respectively, differed by between 21% and 31%: tag, 4% vs. 33%; tag + 4, 47% vs. 68%; and tag + 8, 108% vs. 77%. The results from this work provide a direct comparison of computational and experimental estimates of drag, and provide a framework to quantify uncertainty.
Lethal and sublethal fishing gear entanglement is pervasive in North Atlantic right whales (Eubalaena glacialis). Entanglement can lead to direct injury and is likely to incur substantial energetic ...costs. This study (1) evaluates drag characteristics of entangled right whales, (2) contextualizes gear drag measurements for individual whales, and (3) quantifies the benefits of partial disentanglement. A load cell measured drag forces on 15 sets of fishing gear removed from entangled right whales, a towed satellite telemetry buoy, and 200 m of polypropylene line as it was shortened to 25 m, as they were towed behind a vessel at ~0.77, 1.3, and 2.1 m/s (~1.5, 2.5, and 4 knots) and ~0, 3, and 6 m depth. Mean drag ranges from 8.5 N to 315 N, and can be predicted from the dry weight or length of the gear. Combining gear drag measurements with theoretical estimates of drag on whales' bodies suggests that on average, entanglement increases drag and propulsive power by 1.47 fold. Reducing trailing line length by 75% can reduce parasitic gear drag by 85%, reinforcing current disentanglement response practices. These drag measurements can be incorporated into disentanglement response, serious injury determination, and evaluation of sublethal effects on population dynamics.
To determine effectiveness of Seasonal Management Areas (SMAs), introduced in 2008 on the U.S. East Coast to reduce lethal vessel strikes to North Atlantic right whales, we analyzed observed large ...whale mortality events from 1990–2012 in the geographic region of the “Ship Strike Rule” to identify changes in frequency, spatial distribution, and spatiotemporal interaction since implementation. Though not directly coincident with SMA implementation, right whale vessel‐strike mortalities significantly declined from 2.0 (2000–2006) to 0.33 per year (2007–2012). Large whale vessel‐strike mortalities have decreased inside active SMAs, and increased outside inactive SMAs. We detected no significant spatiotemporal interaction in the 4‐year pre‐ or post‐Rule periods, although a longer time series is needed to detect these changes. As designed, SMAs encompass only 36% of historical right whale vessel‐strike mortalities, and 32% are outside managed space but within managed timeframes. We suggest increasing spatial coverage to improve the Rule's effectiveness.
Empirical metabolic rate and oxygen consumption estimates for free-ranging whales have been limited to counting respiratory events at the surface. Because these observations were limited and ...generally viewed from afar, variability in respiratory properties was unknown and oxygen consumption estimates assumed constant breath-to-breath tidal volume and oxygen uptake. However, evidence suggests that cetaceans in human care vary tidal volume and breathing frequency to meet aerobic demand, which would significantly impact energetic estimates if the findings held in free-ranging species. In this study, we used suction cup-attached video tags positioned posterior to the nares of two humpback whales (
) and four Antarctic minke whales (
) to measure inhalation duration, relative nares expansion, and maximum nares expansion. Inhalation duration and nares expansion varied between and within initial, middle, and terminal breaths of surface sequences between dives. The initial and middle breaths exhibited the least variability and had the shortest durations and smallest nares expansions. In contrast, terminal breaths were highly variable, with the longest inhalation durations and the largest nares expansions. Our results demonstrate breath-to-breath variability in duration and nares expansion, suggesting differential oxygen exchange in each breath during the surface interval. With future validation, inhalation duration or nares area could be used alongside respiratory frequency to improve oxygen consumption estimates by accounting for breath-to-breath variation in wild whales.
United States and Canadian governments have responded to legal requirements to reduce human-induced whale mortality via vessel strikes and entanglement in fishing gear by implementing a suite of ...regulatory actions. We analyzed the spatial and temporal patterns of mortality of large whales in the Northwest Atlantic (23.5° N to 48.0° N), 1970 through 2009, in the context of management changes. We used a multinomial logistic model fitted by maximum likelihood to detect trends in cause-specific mortalities with time. We compared the number of human-caused mortalities with U.S. federally established levels of potential biological removal (i.e., species-specific sustainable human-caused mortality). From 1970 through 2009, 1762 mortalities (all known) and serious injuries (likely fatal) involved 8 species of large whales. We determined cause of death for 43% of all mortalities; of those, 67% (502) resulted from human interactions. Entanglement in fishing gear was the primary cause of death across all species (n = 323), followed by natural causes (n = 248) and vessel strikes (n = 171). Established sustainable levels of mortality were consistently exceeded in 2 species by up to 650%. Probabilities of entanglement and vessel-strike mortality increased significantly from 1990 through 2009. There was no significant change in the local intensity of all or vessel-strike mortalities before and after 2003, the year after which numerous mitigation efforts were enacted. So far, regulatory efforts have not reduced the lethal effects of human activities to large whales on a population-range basis, although we do not exclude the possibility of success of targeted measures for specific local habitats that were not within the resolution of our analyses. It is unclear how shortfalls in management design or compliance relate to our findings. Analyses such as the one we conducted are crucial in critically evaluating wildlife-management decisions. The results of these analyses can provide managers with direction for modifying regulated measures and can be applied globally to mortality-driven conservation issues. Los gobiernos de Estados Unidos y Canadá han respondido a requerimientos legales para reducir la mortalidad de ballenas inducida por humanos por medio de impacto con embarcaciones y enmarañamiento en artes de pesca mediante la implementación de un conjunto de acciones reguladoras. Analizamos los patrones espaciales y temporales de la mortalidad de ballenas mayores en el Atlántico Noroccidental (23.5° N a 48.0° N), de 1970 a 2009, en el contexto de cambios de manejo. Utilizamos un modelo logístico multinomial ajustado por la máxima probabilidad de detección de tendencias en mortalidades por causa específica en el tiempo. Comparamos el número de muertes provocadas por humanos con los niveles de remoción biológica potencial (i.e., mortalidad específica provocada por humanos sustentable). De 1970 a 2009, hubo 1762 muertes (conocidas) y lesiones serias (casi fatales) involucrando 8 especies de ballenas mayores. Determinamos la causa de 43% de todas las muertes; de ellas, 67% (502) resultaron de interacciones humanas. El enmarañamiento en artes de pesca fue la causa principal de muerte en todas las especies (n = 323), seguida de causas naturales (n = 248) e impacto de embarcaciones (n = 171). Los niveles sustentables de mortalidad establecidos fueron excedidos consistentemente hasta en 650% en 2 especies. Las probabilidades de muerte por enmarañamiento y por impacto de embarcaciones incrementaron significativamente de 1990 a 2009. No hubo cambio significativo en la intensidad local de mortalidad por todas las causas o por impacto de embarcaciones antes y después de 2003, año en el que se implementaron numerosos esfuerzos de mitigación. Hasta ahora, los esfuerzos regulatorios no han reducido los efectos letales de las actividades humanas sobre las ballenas a nivel de población, aunque no excluimos la posibilidad de éxito de medidas enfocadas a hábitats locales específicos que no estuvieron dentro de la resolución de nuestro análisis. No es claro como se relacionan con nuestros resultados las deficiencias en el diseño o implementación del manejo. Análisis como el que realizamos son cruciales para la evaluación crítica de decisiones para el manejo de vida silvestre, y los resultados de estos análisis pueden proporcionar directrices a los manejadores para que modifiquen medidas regulatorias y puedan ser aplicadas globalmente en temas de conservación relacionadas con mortalidad.
Vessel strikes are the primary source of known mortality for the endangered North Atlantic right whale (
Eubalaena glacialis
). Multi-institutional efforts to reduce mortality associated with vessel ...strikes include vessel-routing amendments such as the International Maritime Organization voluntary "area to be avoided" (ATBA) in the Roseway Basin right whale feeding habitat on the southwestern Scotian Shelf. Though relative probabilities of lethal vessel strikes have been estimated and published, absolute probabilities remain unknown. We used a modeling approach to determine the regional effect of the ATBA, by estimating reductions in the expected number of lethal vessel strikes. This analysis differs from others in that it explicitly includes a spatiotemporal analysis of real-time transits of vessels through a population of simulated, swimming right whales. Combining automatic identification system (AIS) vessel navigation data and an observationally based whale movement model allowed us to determine the spatial and temporal intersection of vessels and whales, from which various probability estimates of lethal vessel strikes are derived. We estimate one lethal vessel strike every 0.775-2.07 years prior to ATBA implementation, consistent with and more constrained than previous estimates of every 2-16 years. Following implementation, a lethal vessel strike is expected every 41 years. When whale abundance is held constant across years, we estimate that voluntary vessel compliance with the ATBA results in an 82% reduction in the per capita rate of lethal strikes; very similar to a previously published estimate of 82% reduction in the relative risk of a lethal vessel strike. The models we developed can inform decision-making and policy design, based on their ability to provide absolute, population-corrected, time-varying estimates of lethal vessel strikes, and they are easily transported to other regions and situations.
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.
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•Large whales are often entangled in fishing gear for months to years.•Drag and energy burden from entangling gear can be estimated at the time of a whale's detection.•We develop tools for prognosis of specific entanglement cases to assist disentanglement action and inform stock assessment.
Marine mammals are streamlined for efficient movement in their relatively viscous fluid environment and are able to alter their kinematics (i.e. fluke stroke frequency, amplitude, or both) in ...response to changes in force balance. Entanglement in fishing gear adds significant drag and buoyant forces that can impact swimming behaviors across a range of timescales. We deployed biologging tags during the disentanglement of 2 North Atlantic right whales Eubalaena glacialis to (1) examine how their kinematics changed in response to drag and buoyancy from entanglement in fishing gear, and (2) calculate resultant changes in swimming efficiency for one individual. We observed variable responses in dive behavior, but neither whale appeared to exploit added buoyancy to reduce energy expenditure. While some of the observed changes in behavior were individually specific, some swimming kinematics were consistently modulated in response to high drag and buoyancy associated with entangling gear, affecting thrust production. In high drag and buoyancy conditions, fluke strokes were significantly shorter and more variable in shape, and gliding was less frequent. Thrust and efficiency significantly differed among dive phases. Disentanglement reduced thrust coefficients ~4-fold, leading to 1.2 to 1.8-fold lower power (W). Ideal propulsive efficiency was significantly lower when entangled, though we detected no difference in observed propulsive efficiency between the conditions. Similar to carrying heavy objects or changing shoes, we present another condition where animals perceive unique movement constraints over seconds to minutes and develop compensatory strategies, altering their movement accordingly.
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