Factors driving the variability in diving and movement behavior of migrating humpback whales (Megaptera novaeangliae): Implications for anthropogenic disturbance studies

Humpback whales (Megaptera novaeangliae) undertake one of the longest migrations of any animal and while on a broad‐scale this journey appears direct, on a fine‐scale, behaviors associated with socializing and breeding are regularly observed. However, little is known about which social and environmental factors influence behavior during this time. Here we examined the effect of multiple factors on the movement (speed and course) and diving behavior (dive and surfacing duration) of humpback whales during migration off the eastern coast of Australia. Focal data (202 h) were collected on 94 different whale groups with simultaneous social and environmental context data. The environmental factors water depth and wind speed were found to be important predictors of dive and movement behavior, whereas social factors were less influential at this site. Groups tended to dive for longer with increased water depth but traveled more slowly in increasing wind speeds. These baseline studies are crucial when examining the effect of anthropogenic disturbance. Determining which natural factors significantly affect behavior ensures any observed behavioral changes are correctly attributed to the disturbance and are not a result of other factors. In addition, any responses observed can be put into biological context and their relative magnitude determined.     

Evidence for the functions of surface‐active behaviors in humpback whales (Megaptera novaeangliae)

As part of their social sound repertoire, migrating humpback whales (Megaptera novaeangliae) perform a large variety of surface‐active behaviors, such as breaching and repetitive slapping of the pectoral fins and tail flukes; however, little is known about what factors influence these behaviors and what their functions might be. We investigated the potential functions of surface‐active behaviors in humpback whale groups by examining the social and environmental contexts in which they occurred. Focal observations on 94 different groups of whales were collected in conjunction with continuous acoustic monitoring, and data on the social and environmental context of each group. We propose that breaching may play a role in communication between distant groups as the probability of observing this behavior decreased significantly when the nearest whale group was within 4,000 m compared to beyond 4,000 m. Involvement in group interactions, such as the splitting of a group or a group joining with other whales, was an important factor in predicting the occurrence of pectoral, fluke, and peduncle slapping, and we suggest that they play a role in close‐range or within‐group communication. This study highlights the potentially important and diverse roles of surface‐active behaviors in the communication of migrating humpback whales.     

Identifying behavioral states and habitat use of acoustically tracked humpback whales in Hawaii

Although humpback whales have been well‐studied on their Hawaiian breeding grounds, it is difficult to track individual animals over long distances without tags, particularly when they move offshore. Here, singing humpback whales were localized in three dimensions on the Pacific Missile Range Facility off Kauai, Hawaii, located between 20 km and 80 km offshore, from January 2011 through June 2014. Detailed behavioral analyses were conducted on the resulting tracks. One hundred and eight individual tracks were identified and metrics of these tracks were examined. Using these metrics, the tracks were classified into four behavior categories, described herein as Directed Travel, Repeated Stationary Dives, Mill, and tracks with Combinations of behavioral states. Some diel and seasonal patterns were identified, with Mill tracks occurring more at night than the other behaviors, Repeated Stationary Dive tracks occurring more during the day, and Directed Travel occurring only at the start and end of the breeding season. These results provide detailed insights into the movement of singing humpback whales, particularly in offshore waters where they may be migrating into or out of breeding grounds. This also contributes valuable information on the baseline behavior of humpback whales on a US Navy training range.     

Southeastern Pacific humpback whales (Megaptera novaeangliae) and their breeding grounds: Distribution and habitat preference of singers and social groups off the coast of Ecuador

Understanding the distribution, habitat preference, and social structure of highly migratory species at important life history stages (e.g., breeding and calving) is essential for conservation efforts. We investigated the spatial distribution and habitat preference of humpback whale social groups and singers, in relation to depth categories (<20 m, 20–50 m, and >50 m) and substrate type (muddy and mixed) on a coastal southeastern Pacific breeding ground. One hundred and forty‐three acoustic stations and 304 visual sightings were made at the breeding ground off the coast of Esmeraldas, Ecuador. Spatial autocorrelation analysis suggested singers were not randomly distributed, and Neu's method and Monte Carlo simulations indicated that singers frequented depths of <20 m and mixed substrate. Singletons, and groups with a calf displayed a preference for shallower waters (0–20 m), while pairs and groups with a calf primarily inhabited mixed bottom substrates. In contrast, competitive groups showed no clear habitat preference and exhibited social segregation from other whales. Understanding the habitat preference and distribution of humpback whales on breeding and calving grounds vulnerable to anthropogenic disturbance provides important baseline information that should be incorporated into conservation efforts at a regional scale.     

Prey preferences of sympatric fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales revealed by stable isotope mixing models

Over‐exploitation of top predators and fish stocks has altered ecosystems towards less productive systems with fewer trophic levels. In the Celtic Sea (CS), discards and bycatch levels have prompted concern about some fisheries, while fin and humpback whales are recovering from centuries of over‐exploitation. A lack of empirical evidence on the preferred diet of some predators such as whales in the CS has hindered the implementation of effective conservation measures using an ecosystem‐based approach to fisheries management. Using a Bayesian framework (SIAR), stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope mixing models were used to assign proportionate diet solutions to fin and humpback whales (skin biopsies) and putative prey items: herring (Clupea harengus), sprat (Sprattus sprattus), and krill (Meganyctiphanes norvegica and Nyctiphanes couchii) in the CS. Krill was the single most important prey item in the diet of fin whales, but one of the least important for humpback whales (albeit based on a small sample of humpback whale samples). Age 0 sprat and herring comprised a large proportion of the diet of both species, followed by older sprat (age 1–2) and older herring (age 2–4). An ecosystem based approach to fisheries management will be required in the CS if we seek effective conservation of both fin and humpback whales, and sustainable fisheries.     

Whale killers: Prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia

Reports of killer whales (Orcinus orca) preying on large whales have been relatively rare, and the ecological significance of these attacks is controversial. Here we report on numerous observations of killer whales preying on neonate humpback whales (Megaptera novaeangliae) off Western Australia (WA) based on reports we compiled and our own observations. Attacking killer whales included at least 19 individuals from three stable social groupings in a highly connected local population; 22 separate attacks with known outcomes resulted in at least 14 (64%) kills of humpback calves. We satellite‐tagged an adult female killer whale and followed her group on the water for 20.3 h over six separate days. During that time, they attacked eight humpback calves, and from the seven known outcomes, at least three calves (43%) were killed. Overall, our observations suggest that humpback calves are a predictable, plentiful, and readily taken prey source for killer whales and scavenging sharks off WA for at least 5 mo/yr. Humpback “escorts” vigorously assisted mothers in protecting their calves from attacking killer whales (and a white shark, Carcharodon carcharias). This expands the purported role of escorts in humpback whale social interactions, although it is not clear how this behavior is adaptive for the escorts.     

Delphinid whistle production and call matching during playback of simulated military sonar

In 2007 and 2008, controlled exposure experiments were performed in the Bahamas to study behavioral responses to simulated mid‐frequency active sonar (MFA) by three groups of odontocetes: false killer whales, Pseudorca crassidens; short‐finned pilot whales, Globicephala macrorhynchus; and melon‐headed whales, Peponocephala electra. An individual in each group was tagged with a Dtag to record acoustic and movement data. During exposures, some individuals produced whistles that seemed similar to the experimental MFA stimulus. Statistical tests were thus applied to investigate whistle‐MFA similarity and the relationship between whistle production rate and MFA reception time. For the false killer whale group, overall whistle rate and production rate of the most MFA‐like whistles decreased with time since last MFA reception. Despite quite low whistle rates overall by the melon‐headed whales, statistical results indicated minor transient silencing after each signal reception. There were no apparent relationships between pilot whale whistle rates and MFA sounds within the exposure period. This variability of responses suggests that changes in whistle production in response to acoustic stimuli depend not only on species and sound source, but also on the social, behavioral, or environmental contexts of exposure.     

How does social behavior differ among sperm whale clans?

When individuals primarily associate with and learn from those who behave similarly, society and culture become closely tied. Sperm whales (Physeter macrocephalus) exhibit multilevel social structure, the levels of which are differentiated in part by characteristic cultural behaviors. Sperm whales are organized into sympatric clans, with distinctive vocal repertoires that are socially learned. Other behaviors, such as movement patterns and foraging, also differ among clans. Here we ask whether the clan partition also includes divergences in social behavior. Off the Galápagos Islands, members of two clans differed consistently in diving synchrony, heterogeneity, and temporal stability of social relationships. While number of associates (indicated by social unit, group, and cluster sizes) were similar between clans, Regular clan members dived more synchronously and had more homogeneous relationships than the Plus‐One clan members. Plus‐One social units had generally longer associations than those of the Regular clan. Differences in surface‐time coordination and quality of social relationships are likely byproducts of the clan segregation, which could affect alloparental care giving, therefore scaling up to differential calf survival rates between clans. This new dimension of behavioral divergence between sperm whale clans indicates that sympatric, socio‐cultural entities of nonhumans can also display characteristic social behavior.     

Resource partitioning facilitates coexistence in sympatric cetaceans in the California Current

Resource partitioning is an important process driving habitat use and foraging strategies in sympatric species that potentially compete. Differences in foraging behavior are hypothesized to contribute to species coexistence by facilitating resource partitioning, but little is known on the multiple mechanisms for partitioning that may occur simultaneously. Studies are further limited in the marine environment, where the spatial and temporal distribution of resources is highly dynamic and subsequently difficult to quantify. We investigated potential pathways by which foraging behavior may facilitate resource partitioning in two of the largest co‐occurring and closely related species on Earth, blue (Balaenoptera musculus) and humpback (Megaptera novaeangliae) whales. We integrated multiple long‐term datasets (line‐transect surveys, whale‐watching records, net sampling, stable isotope analysis, and remote‐sensing of oceanographic parameters) to compare the diet, phenology, and distribution of the two species during their foraging periods in the highly productive waters of Monterey Bay, California, USA within the California Current Ecosystem. Our long‐term study reveals that blue and humpback whales likely facilitate sympatry by partitioning their foraging along three axes: trophic, temporal, and spatial. Blue whales were specialists foraging on krill, predictably targeting a seasonal peak in krill abundance, were present in the bay for an average of 4.7 months, and were spatially restricted at the continental shelf break. In contrast, humpback whales were generalists apparently feeding on a mixed diet of krill and fishes depending on relative abundances, were present in the bay for a more extended period (average of 6.6 months), and had a broader spatial distribution at the shelf break and inshore. Ultimately, competition for common resources can lead to behavioral, morphological, and physiological character displacement between sympatric species. Understanding the mechanisms for species coexistence is both fundamental to maintaining biodiverse ecosystems, and provides insight into the evolutionary drivers of morphological differences in closely related species.     

Using dive behavior and active acoustics to assess prey use and partitioning by fin and humpback whales near Kodiak Island,Alaska

Near the Kodiak Archipelago, fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales frequently overlap spatially and temporally. The Gulf Apex Predator‐prey study (GAP) investigated the prey use and potential prey partitioning between these sympatric species by combining concurrent analysis of vertical whale distribution with acoustic assessment of pelagic prey. Acoustic backscatter was classified as consistent with either fish or zooplankton. Whale dive depths were determined through suction cup tags. Tagged humpback whales (n = 10) were most often associated with distribution of fish, except when zooplankton density was very high. Associations between the dive depths of tagged fin whales (n = 4) and the vertical distribution of either prey type were less conclusive. However, prey assessment methods did not adequately describe the distribution of copepods, a potentially significant resource for fin whales. Mean dive parameters showed no significant difference between species when compared across all surveys. However, fin whales spent a greater proportion of dive time in the foraging phase than humpbacks, suggesting a possible difference in foraging efficiency between the two. These results suggest that humpback and fin whales may target different prey, with the greatest potential for diet overlap occurring when the density of zooplankton is very high.     

Response of southern right whales to simulated swim‐with‐whale tourism at Península Valdés,Argentina

Guidelines for sustainable tourism involving swimming with large whales are not well‐developed compared to those focused on programs of swimming with delphinids. From September to November 2005 and August to September 2006, we collected behavioral and movement data for southern right whales (Eubalaena australis) exposed to interactions with boats and swimmers at Península Valdés, Argentina. Whales were tracked from shore using a theodolite before, during, and after a series of directed interactions with swimmers and a boat. Resting, socializing, and surface active behavior decreased, traveling increased, and whales swam faster and reoriented more often during interactions. Responses were variable by age/sex class, with mother/calf pairs showing strongest responses. Increased levels of tourism activity are a concern, as reduction in resting time and disruption of socialization among adults, juveniles, and mother/calf pairs have unknown long‐term consequences. Additional data should be collected for whale behavior in proposed tourism and nontourism areas to build a long‐term database which can be used to determine if reactions of whales change over time. Our data suggest that swimming with whales in Chubut Province should not be legalized until further investigations are completed, especially in light of the recent southern right whale die‐offs recorded in Península Valdés.     

Evidence of two subaggregations of humpback whales on the Kodiak,Alaska, feeding ground revealed from stable isotope analysis

Knowledge of humpback whale (Megaptera novaeangliae) foraging on feeding grounds is becoming increasingly important as the growing North Pacific population recovers from commercial whaling and consumes more prey, including economically important fishes. We explored spatial and temporal (interannual, within‐season) variability in summer foraging by humpback whales along the eastern side of the Kodiak Archipelago as described by stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin (n = 118; 2004–2013). The trophic level (TL) of individual whales was calculated using basal food web δ¹⁵N values collected within the study area. We found evidence for the existence of two subaggregations of humpback whales (“North,” “South”) on the feeding ground that fed at different TLs throughout the study period. Linear mixed models suggest that within an average year, Kodiak humpback whales forage at a consistent TL during the feeding season. TL estimates support mixed consumption of fish and zooplankton species in the “North” (mean ± SE; 3.3 ± 0.1) and predominant foraging on zooplankton in the “South” (3.0 ± 0.1). This trend appears to reflect spatial differences in prey availability, and thus, our results suggest North Pacific humpback whales may segregate on feeding aggregations and target discrete prey species.     

Potential vessel collisions with Southern Hemisphere humpback whales wintering off Pacific Panama

Vessel collision is a threat to many whale species, and the risk has increased with expanding maritime traffic. This compromises international conservation efforts and requires urgent attention from the world's maritime industry. Humpback whales (Megaptera novaeangliae) are at the top of the death toll, and although Central America is a wintering area for populations from both the Northern and Southern Hemispheres, existing efforts to reduce ship‐whale collisions are meager. Herein, we evaluated the potential collisions between vessels and humpback whales wintering off Pacific Panama by following the movements of 15 whales tagged with satellite transmitters and comparing these data with tracks plotted using AIS real‐time latitude‐longitude points from nearly 1,000 commercial vessels. Movements of whales (adults and calves) in the Gulf of Panama coincide with major commercial maritime routes. AIS vessel data analyzed for individual whale satellite tracks showed that 53% (8 whales) of whales had 98 encounters within 200 m with 81 different vessels in just 11 d. We suggest implementing a 65 nmi Traffic Separation Scheme and a 10 kn maximum speed for vessel routing into the Gulf of Panama during the wintering season. In so doing, the area for potential whale‐vessel collisions could be reduced by 93%.     

Whales from space: Four mysticete species described using new VHR satellite imagery

Large‐bodied animals such as baleen whales can now be detected with very high resolution (VHR) satellite imagery, allowing for scientific studies of whales in remote and inaccessible areas where traditional survey methods are limited or impractical. Here we present the first study of baleen whales using the WorldView‐3 satellite, which has a maximum spatial resolution of 31 cm in the panchromatic band, the highest currently available to nonmilitary professionals. We manually detected, described, and counted four different mysticete species: fin whales (Balaenoptera physalus) in the Ligurian Sea, humpback whales (Megaptera novaeangliae) off Hawaii, southern right whales (Eubalaena australis) off Península Valdés, and gray whales (Eschrichtius robustus) in Laguna San Ignacio. Visual and spectral analyses were conducted for each species, their surrounding waters, and nonwhale objects (e.g., boats). We found that behavioral and morphological differences made some species more distinguishable than others. Fin and gray whales were the easiest to discern due to their contrasting body coloration with surrounding water, and their prone body position, which is proximal to the sea surface (i.e., body parallel to the sea surface). These results demonstrate the feasibility of using VHR satellite technology for monitoring the great whales.     

From the southern right whale hunting decline to the humpback whaling expansion: a review of whale catch records in the tropical western South Atlantic Ocean

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Migratory preferences of humpback whales between feeding and breeding grounds in the eastern South Pacific

Latitudinal preferences within the breeding range have been suggested for Breeding Stock G humpback whales that summer in different feeding areas of the eastern South Pacific. To address this hypothesis, humpback whales photo‐identified from the Antarctic Peninsula and the Fueguian Archipelago (southern Chile) were compared with whales photo‐identified from lower latitudes extending from northern Peru to Costa Rica. This comparison was performed over a time span that includes 18 austral seasons. A total of 238 whales identified from the Antarctic Peninsula and 25 whales from the Fueguian Archipelago were among those photo‐identified at the breeding grounds. Our findings showed that humpback whales from each feeding area were resighted unevenly across the breeding grounds, which suggests a degree of spatial structuring in the migratory pathway. Humpback whales that feed at the Antarctic Peninsula were more likely to migrate to the southern breeding range between northern Peru and Colombia, whereas whales that feed at the Fueguian Archipelago were more likely to be found in the northern range of the breeding ground off Panama. Further photo‐identification efforts and genetic sampling from poorly sampled or unsampled areas are recommended to confirm these reported connectivity patterns.     

Insights into the use of an unmanned aerial vehicle (UAV) to investigate the behavior of humpback whales (Megaptera novaeangliae) in Vava'u,Kingdom of Tonga

Vertical take-off and landing (VTOL) unmanned aerial vehicles (UAVs) are becoming invaluable data collection platforms for cetacean research. In particular, multi-rotors can be used to measure whales and investigate their behavior. Moreover, VTOL UAVs are increasingly accessible for recreational and commercial pilots, and close encounters with whales are widely documented. Unfortunately, quantitative assessments of potential disturbance for the targeted animals are not yet published and guidelines for responsible use of UAVs around cetaceans are still under development. We conducted VTOL UAV surveys on humpback whales in Vava'u, Kingdom of Tonga. Interestingly, whale behaviors such as socializing and nurturing were not detected by trained observers on board the research vessel, but were evident from the UAV. Nevertheless, no significant differences were detected in diving and swim parameters between absence and presence of UAV flying at 30 m altitude. These results suggest that VTOL UAVs can be a noninvasive tool to gather morphometric and behavioral data on baleen whales. However, further research is necessary to establish whether applications that require flight altitudes lower than 30 m and targeting different species may elicit behavioral responses.     

Development of an automated method of detecting stereotyped feeding events in multisensor data from tagged rorqual whales

The introduction of animal‐borne, multisensor tags has opened up many opportunities for ecological research, making previously inaccessible species and behaviors observable. The advancement of tag technology and the increasingly widespread use of bio‐logging tags are leading to large volumes of sometimes extremely detailed data. With the increasing quantity and duration of tag deployments, a set of tools needs to be developed to aid in facilitating and standardizing the analysis of movement sensor data. Here, we developed an observation‐based decision tree method to detect feeding events in data from multisensor movement tags attached to fin whales (Balaenoptera physalus). Fin whales exhibit an energetically costly and kinematically complex foraging behavior called lunge feeding, an intermittent ram filtration mechanism. Using this automated system, we identified feeding lunges in 19 fin whales tagged with multisensor tags, during a total of over 100 h of continuously sampled data. Using movement sensor and hydrophone data, the automated lunge detector correctly identified an average of 92.8% of all lunges, with a false‐positive rate of 9.5%. The strong performance of our automated feeding detector demonstrates an effective, straightforward method of activity identification in animal‐borne movement tag data. Our method employs a detection algorithm that utilizes a hierarchy of simple thresholds based on knowledge of observed features of feeding behavior, a technique that is readily modifiable to fit a variety of species and behaviors. Using automated methods to detect behavioral events in tag records will significantly decrease data analysis time and aid in standardizing analysis methods, crucial objectives with the rapidly increasing quantity and variety of on‐animal tag data. Furthermore, our results have implications for next‐generation tag design, especially long‐term tags that can be outfitted with on‐board processing algorithms that automatically detect kinematic events and transmit ethograms via acoustic or satellite telemetry.     

Fine‐scale spatial differences in humpback whale diet composition near Kodiak,Alaska

On the North Pacific feeding grounds, humpback whales (Megaptera novaeangliae) are recovering from commercial whaling at a rapid rate (6.8%). The potential effect that this recovery will have on trophic dynamics involving these predators is currently unknown. To better elucidate complex trophic dynamics, variability in diet composition of apex predators on their respective feeding grounds needs to be understood. Thus, we explored the diet composition of two defined subaggregations of humpback whales of the Kodiak Archipelago population (“North,” “South”) using stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin and regional prey samples in Bayesian dietary mixing models. Humpback whales in the “North” region consumed proportionally more fish, dominated by capelin (Mallotus villosus), whereas, whales in the “South” region consumed predominantly krill. The difference in diet composition appears to reflect regional differences in prey availability. Thus, regional variability in diet composition by humpback whales may have disproportionate impacts on prey resources of sympatric predators depending on available prey biomass. As a result, we suggest fine‐scale studies of apex predator diets are needed to better model trophic dynamics with accuracy.     

BEHAVIORAL RESPONSE OF FOUR SPECIES OF BALAENOPTERID WHALES TO BIOPSY SAMPLING

Behavioral responses to biopsy sampling of four species of northwestern Atlantic balaenopterid whales summering in the estuary and Gulf of St. Lawrence, Quebec, from 1990 to 1995 were studied to determine if this technique was an important disturbance to the whales. A total of 447 biopsy samples were taken using a small punch-type biopsy tip fired from a crossbow. Biopsies were successfully taken from 91.2% of the whales approached. Whales displayed no reaction to 45.2% of the successful biopsy attempts. Whales that responded to biopsy sampling typically resumed their normal behavior immediately or within a few minutes. Most humpback whales displayed a hard tail flick, and the majority of fin and blue whales submerged following biopsy sampling. Significantly different frequencies and intensities of responses were found between whale species. Minke and humpback whales were found to be more sensitive to biopsy sampling than fin and blue whales. Response frequencies were similar between females and males for all species, with the exception of fin whales where females had a higher response frequency than males. Biopsy sample length, i. e., penetration depth, did not explain variations in response intensity but may influence response frequency to biopsy sampling. Group size, geographical region, and number of biopsies taken per whale were not factors that explained variation in behavioral responses. The biopsy technique was found to be an efficient method for obtaining high-quality whale skin and blubber samples with limited behavioral disturbance to balaenopterid whales.