Daniel M. Palacios

Southern Hemisphere Humpback Whales Wintering Off Central America: Insights from Water Temperature into the Longest Mammalian Migration

We report on a wintering area off the Pacific coast of Central America for humpback whales (Megaptera novaeangliae) migrating from feeding areas off Antarctica. We document seven individuals, including a mother/calf pair, that made this migration (approx. 8300 km), the longest movement undertaken by any mammal. Whales were observed as far north as 11° N off Costa Rica, in an area also used by a boreal population during the opposite winter season, resulting in unique spatial overlap between Northern and Southern Hemisphere populations. The occurrence of such a northerly wintering area is coincident with the development of an equatorial tongue of cold water in the eastern South Pacific, a pattern that is repeated in the eastern South Atlantic. A survey of location and water temperature at the wintering areas worldwide indicates that they are found in warm waters (21.1–28.3°C), irrespective of latitude. We contend that while availability of suitable reproductive habitat in the wintering areas is important at the fine scale, water temperature influences whale distribution at the basin scale. Calf development in warm water may lead to larger adult size and increased reproductive success, a strategy that supports the energy conservation hypothesis as a reason for migration.

Spatial and Temporal Occurrence of Blue Whales off the U.S. West Coast, with Implications for Management

Mortality and injuries caused by ship strikes in U.S. waters are a cause of concern for the endangered population of blue whales (Balaenoptera musculus) occupying the eastern North Pacific. We sought to determine which areas along the U.S. West Coast are most important to blue whales and whether those areas change inter-annually. Argos-monitored satellite tags were attached to 171 blue whales off California during summer/early fall from 1993 to 2008. We analyzed portions of the tracks that occurred within U.S. Exclusive Economic Zone waters and defined the ‘home range’ (HR) and ‘core areas’ (CAU) as the 90% and 50% fixed kernel density distributions, respectively, for each whale. We used the number of overlapping individual HRs and CAUs to identify areas of highest use. Individual HR and CAU sizes varied dramatically, but without significant inter-annual variation despite covering years with El Niño and La Niña conditions. Observed within-year differences in HR size may represent different foraging strategies for individuals. The main areas of HR and CAU overlap among whales were near highly productive, strong upwelling centers that were crossed by commercial shipping lanes. Tagged whales generally departed U.S. Exclusive Economic Zone waters from mid-October to mid-November, with high variability among individuals. One 504-d track allowed HR and CAU comparisons for the same individual across two years, showing similar seasonal timing, and strong site fidelity. Our analysis showed how satellite-tagged blue whales seasonally used waters off the U.S. West Coast, including high-risk areas. We suggest possible modifications to existing shipping lanes to reduce the likelihood of collisions with vessels.

WhaleWatch: a dynamic management tool for predicting blue whale density in the California Current

Summary Management of highly migratory species is reliant on spatially and temporally explicit information on their distribution and abundance. Satellite telemetry provides time-series data on individual movements. However, these data are underutilized in management applications in part because they provide presence-only information rather than abundance information such as density. Eastern North Pacific blue whales are listed as threatened, and ship strikes have been suggested as a key factor limiting their recovery. Here, we developed a satellite-telemetry-based habitat model in a case–control design for Eastern North Pacific blue whales Balaenoptera musculus that was combined with previously published abundance estimates to predict habitat preference and densities. Further, we operationalize an automated, near-real-time whale density prediction tool based on up-to-date environmental data for use by managers and other stakeholders. A switching state-space movement model was applied to 104 blue whale satellite tracks from 1994 to 2008 to account for errors in the location estimates and provide daily positions (case points). We simulated positions using a correlated random walk model (control points) and sampled the environment at each case and control point. Generalized additive mixed models and boosted regression trees were applied to determine the probability of occurrence based on environmental covariates. Models were used to predict 8-day and monthly resolution, year-round density estimates scaled by population abundance estimates that provide a critical tool for understanding seasonal and interannual changes in habitat use. The telemetry-based habitat model predicted known blue whale hot spots and had seasonal agreement with sightings data, highlighting the skill of the model for predicting blue whale habitat preference and density. We identified high interannual variability in occurrence emphasizing the benefit of dynamic models compared to multiyear averages. Synthesis and applications. This near-real-time tool allows a more accurate examination of the year-round spatio-temporal overlap of blue whales with potentially harmful human activities, such as shipping. This approach should also be applicable to other species for which sufficient telemetry data are available. The dynamic predictive product developed here is an important tool that allows managers to consider finer-scale management areas that are more economically feasible and socially acceptable.

Oceanographic and climatic variation drive top‐down/bottom‐up coupling in the Galápagos intertidal meta‐ecosystem

The impact of herbivores on primary producers in differing oceanographic regimes is a matter of intense ecological interest due to ongoing changes in their abundance, that of their predators, and anthropomorphic alteration of nutrient cycles and climatic patterns. Interactions between productivity and herbivory in marine habitats have been studied on temperate rocky shores, coral reefs, mangroves, and salt marshes, but less so at tropical latitudes. To determine how herbivore-alga dynamics varied with oceanographic regime, we used the comparative-experimental approach in rocky intertidal communities on the Galapagos Islands from January 2006 to January 2009. This setting was selected because strongly contrasting oceanographic conditions occurred within a range of ;181 km, with significant differences in temperature, nutrients, phytoplankton productivity, and intertidal communities, and in abundance of macro-herbivores, including marine iguanas. Experiments and measurements were conducted at two sites in each of three oceanographic regimes characterized by low, intermediate, and high bottom-up inputs. At sites of low inputs, macro- herbivores (fish, crabs, iguanas) had a consistent top-down effect, reducing algal abundance, and leaving a few grazer-resistant varieties. At sites of intermediate and high inputs, consumer impacts were stronger during La Ni˜ na (cool phase) than during El Ni˜ no (warm phase). At sites of high inputs, algal biomass was naturally relatively high and was dominated by the edible algae Ulva spp. Macro-grazers reduced algal biomass, but their primary effect was indirect, as articulated corallines displaced other species of algae in their absence. Prior results from the tropics had revealed dominant effects of top-down interactions and recruitment in structuring intertidal communities. Our results suggest that, when a broader oceanographic scenario is taken into account, the relative importance of top-down and bottom-up forces are context dependent, varying with oceanographic regime and climatic variability.

Pycnocline Variations in the Eastern Tropical and North Pacific, 1958–2008

AbstractClimatic variability of the pycnocline in the eastern tropical and North Pacific has oceanographic and ecological implications. Gridded monthly profiles of temperature and salinity from the Simple Ocean Data Assimilation (SODA) reanalysis, 1958–2008, were used to derive estimates of four variables related to the density structure of the upper-ocean water column: surface temperature, pycnocline depth, mixed layer depth, and stratification (potential energy anomaly). The pycnocline is primarily a thermal gradient in this region, except in subarctic waters at the northern extreme of the study area, where salinity becomes more important than temperature in determining stratification. Spatial patterns of mean and standard deviation of the four pycnocline variables are presented. Partitioning of variance between seasonal and interannual scales shows the predominance of interannual variability in the tropics and seasonal variability at higher latitudes. Low-frequency variations (trends) in the pycnocline...

Inferring cetacean population densities from the absolute dynamic topography of the ocean in a hierarchical Bayesian framework

We inferred the population densities of blue whales (Balaenoptera musculus) and short-beaked common dolphins (Delphinus delphis) in the Northeast Pacific Ocean as functions of the water-column’s physical structure by implementing hierarchical models in a Bayesian framework. This approach allowed us to propagate the uncertainty of the field observations into the inference of species-habitat relationships and to generate spatially explicit population density predictions with reduced effects of sampling heterogeneity. Our hypothesis was that the large-scale spatial distributions of these two cetacean species respond primarily to ecological processes resulting from shoaling and outcropping of the pycnocline in regions of wind-forced upwelling and eddy-like circulation. Physically, these processes affect the thermodynamic balance of the water column, decreasing its volume and thus the height of the absolute dynamic topography (ADT). Biologically, they lead to elevated primary productivity and persistent aggregation of low-trophic-level prey. Unlike other remotely sensed variables, ADT provides information about the structure of the entire water column and it is also routinely measured at high spatial-temporal resolution by satellite altimeters with uniform global coverage. Our models provide spatially explicit population density predictions for both species, even in areas where the pycnocline shoals but does not outcrop (e.g. the Costa Rica Dome and the North Equatorial Countercurrent thermocline ridge). Interannual variations in distribution during El Niño anomalies suggest that the population density of both species decreases dramatically in the Equatorial Cold Tongue and the Costa Rica Dome, and that their distributions retract to particular areas that remain productive, such as the more oceanic waters in the central California Current System, the northern Gulf of California, the North Equatorial Countercurrent thermocline ridge, and the more southern portion of the Humboldt Current System. We posit that such reductions in available foraging habitats during climatic disturbances could incur high energetic costs on these populations, ultimately affecting individual fitness and survival.

A Dynamic State Model of Migratory Behavior and Physiology to Assess the Consequences of Environmental Variation and Anthropogenic Disturbance on Marine Vertebrates

Integrating behavior and physiology is critical to formulating new hypotheses on the evolution of animal life-history strategies. Migratory capital breeders acquire most of the energy they need to sustain migration, gestation, and lactation before parturition. Therefore, when predicting the impact of environmental variation on such species, a mechanistic understanding of the physiology of their migratory behavior is required. Using baleen whales as a model system, we developed a dynamic state variable model that captures the interplay among behavioral decisions, energy, reproductive needs, and the environment. We applied the framework to blue whales (Balaenoptera musculus) in the eastern North Pacific Ocean and explored the effects of environmental and anthropogenic perturbations on female reproductive success. We demonstrate the emergence of migration to track prey resources, enabling us to quantify the trade-offs among capital breeding, body condition, and metabolic expenses. We predict that periodic climatic oscillations affect reproductive success less than unprecedented environmental changes do. The effect of localized, acute anthropogenic impacts depended on whales’ behavioral response to the disturbance; chronic, but weaker, disturbances had little effect on reproductive success. Because we link behavior and vital rates by modeling individuals’ energetic budgets, we provide a general framework to investigate the ecology of migration and assess the population consequences of disturbance, while identifying critical knowledge gaps.

Reproductive constraints influence habitat accessibility, segregation, and preference of sympatric albatross species

BackgroundThe spatiotemporal distribution of animals is dependent on a suite of factors, including the distribution of resources, interactions within and between species, physiological limitations, and requirements for reproduction, dispersal, or migration. During breeding, reproductive constraints play a major role in the distribution and behavior of central place foragers, such as pelagic seabirds. We examined the foraging behavior and marine habitat selection of Laysan (Phoebastria immutabilis) and black-footed (P. nigripes) albatrosses throughout their eight month breeding cycle at Tern Island, Northwest Hawaiian Islands to evaluate how variable constraints of breeding influenced habitat availability and foraging decisions. We used satellite tracking and light-based geolocation to determine foraging locations of individuals, and applied a biologically realistic null usage model to generate control locations and model habitat preference under a case–control design. Remotely sensed oceanographic data were used to characterize albatross habitats in the North Pacific.ResultsIndividuals of both species ranged significantly farther and for longer durations during incubation and chick-rearing compared to the brooding period. Interspecific segregation of core foraging areas was observed during incubation and chick-rearing, but not during brooding. At-sea activity patterns were most similar between species during brooding; neither species altered foraging effort to compensate for presumed low prey availability and high energy demands during this stage. Habitat selection during long-ranging movements was most strongly associated with sea surface temperature for both species, with a preference for cooler ocean temperatures compared to overall availability. During brooding, lower explanatory power of habitat models was likely related to the narrow range of ocean temperatures available for selection.ConclusionsLaysan and black-footed albatrosses differ from other albatross species in that they breed in an oligotrophic marine environment. During incubation and chick-rearing, they travel to cooler, more productive waters, but are restricted to the low-productivity environment near the colony during brooding, when energy requirements are greatest. Compared to other albatross species, Laysan and black-footed albatrosses spend a greater proportion of time in flight when foraging, especially during the brooding period; this strategy may be adaptive for locating dispersed prey in an oligotrophic environment.

The development of an intermediate-duration tag to characterize the diving behavior of large whales

Abstract The development of high‐resolution archival tag technologies has revolutionized our understanding of diving behavior in marine taxa such as sharks, turtles, and seals during their wide‐ranging movements. However, similar applications for large whales have lagged behind due to the difficulty of keeping tags on the animals for extended periods of time. Here, we present a novel configuration of a transdermally attached biologging device called the Advanced Dive Behavior (ADB) tag. The ADB tag contains sensors that record hydrostatic pressure, three‐axis accelerometers, magnetometers, water temperature, and light level, all sampled at 1 Hz. The ADB tag also collects Fastloc GPS locations and can send dive summary data through Service Argos, while staying attached to a whale for typical periods of 3–7 weeks before releasing for recovery and subsequent data download. ADB tags were deployed on sperm whales (Physeter macrocephalus; N = 46), blue whales (Balaenoptera musculus; N = 8), and fin whales (B. physalus; N = 5) from 2007 to 2015, resulting in attachment durations from 0 to 49.6 days, and recording 31 to 2,539 GPS locations and 27 to 2,918 dives per deployment. Archived dive profiles matched well with published dive shapes of each species from short‐term records. For blue and fin whales, feeding lunges were detected using peaks in accelerometer data and matched corresponding vertical excursions in the depth record. In sperm whales, rapid orientation changes in the accelerometer data, often during the bottom phase of dives, were likely related to prey pursuit, representing a relative measure of foraging effort. Sperm whales were documented repeatedly diving to, and likely foraging along, the seafloor. Data from the temperature sensor described the vertical structure of the water column in all three species, extending from the surface to depths >1,600 m. In addition to providing information needed to construct multiweek time budgets, the ADB tag is well suited to studying the effects of anthropogenic sound on whales by allowing for pre‐ and post‐exposure monitoring of the whales dive behavior. This tag begins to bridge the gap between existing long‐duration but low‐data throughput tags, and short‐duration, high‐resolution data loggers.

Sperm whale dive behavior characteristics derived from intermediate-duration archival tag data

Abstract Here, we describe the diving behavior of sperm whales (Physeter macrocephalus) using the Advanced Dive Behavior (ADB) tag, which records depth data at 1‐Hz resolution and GPS‐quality locations for over 1 month, before releasing from the whale for recovery. A total of 27 ADB tags were deployed on sperm whales in the central Gulf of California, Mexico, during spring 2007 and 2008, of which 10 were recovered for data download. Tracking durations of all tags ranged from 0 to 34.5 days (median = 2.3 days), and 0.6 to 26.6 days (median = 5.0 days) for recovered tags. Recovered tags recorded a median of 50.8 GPS‐quality locations and 42.6 dives per day. Dive summary metrics were generated for archived dives and were subsequently classified into six categories using hierarchical cluster analysis. A mean of 77% of archived dives per individual were one of four dive categories with median Maximum Dive Depth >290 m (V‐shaped, Mid‐water, Benthic, or Variable), likely associated with foraging. Median Maximum Dive Depth was <30 m for the other two categories (Short‐ and Long‐duration shallow dives), likely representing socializing or resting behavior. Most tagged whales remained near the tagging area during the tracking period, but one moved north of Isla Tiburón, where it appeared to regularly dive to, and travel along the seafloor. Three whales were tagged on the same day in 2007 and subsequently traveled in close proximity (<1 km) for 2 days. During this period, the depth and timing of their dives were not coordinated, suggesting they were foraging on a vertically heterogeneous prey field. The multiweek dive records produced by ADB tags enabled us to generate a robust characterization of the diving behavior, activity budget, and individual variation for an important predator of the mesopelagos over temporal and spatial scales not previously possible.