Two‐component calls in short‐finned pilot whales (Globicephala macrorhynchus)

Short‐finned pilot whales (Globicephala macrorhynchus) have complex vocal repertoires that include calls with two time‐frequency contours known as two‐component calls. We attached digital acoustic recording tags (DTAGs) to 23 short‐finned pilot whales off Cape Hatteras, North Carolina, and assessed the similarity of two‐component calls within and among tags. Two‐component calls made up <3% of the total number of calls on 19 of the 23 tag records. For the remaining four tags, two‐component calls comprised 9%, 23%, 24%, and 57% of the total calls recorded. Measurements of six acoustic parameters for both the low and high frequency components of all two‐component calls from the five tags were compared using a generalized linear model. There were significant differences in the acoustic parameters of two‐component calls between tags, verifying that acoustic parameters were more similar for two‐component calls recorded on the same tag than for calls between tags. Spectrograms of all two‐component calls from the five tags were visually graded and independently categorized by five observers. A test of inter‐rater reliability showed substantial agreement, suggesting that each tag contained a predominant two‐component call type that was not shared across tags.     

Survival and abundance of short‐finned pilot whales in the archipelago of Madeira,NE Atlantic

Estimates of population parameters for the short‐finned pilot whale, Globicephala macrorhynchus, are scarce in literature, contributing to an International Union for Conservation of Nature (IUCN) status of Data Deficient. In this study, photo‐identification data collected over 7 yr from Madeira were used to estimate for the first time survivorship, capture probability, and abundance in this species using mark‐recapture methodology. The Cormack‐Jolly‐Seber model estimated that the adult island‐associated (i.e., resident and regular visitor) whales had a constant survival rate of 0.960 (95% CI: 0.853–0.990) and an annual capture probability varying between 0.372 (CI: 0.178–0.619) and 0.843 (CI: 0.619–0.947). A parameterization of the Jolly‐Seber model estimated that 140 island‐associated whales (CI: 131–151) used the area throughout the course of the study. Based on a closed population model, the most precise (lower CV) annual estimate of the total number of pilot whales using the southern and eastern waters of Madeira (~900 km²) in a 3 mo period covering summer/autumn was 334 animals (CI: 260–437). No trend was observed. Despite including biases, the approach used in this study provided plausible estimates of population parameters, which can contribute to the regional conservation strategies.     

Stomach contents of mass-stranded short-finned pilot whales (Globicephala macrorhynchus) from North Carolina

We examined the stomach contents of 27 short‐finned pilot whales (Globicephala macrorhynchus) that mass stranded on the North Carolina coast on 15 January 2005. Eleven whales had prey parts in their forestomachs. We used frequency of occurrence and numerical abundance to assess the relative importance of prey. Brachioteuthis riisei (numerical abundance 28%), an oceanic species, was the most important cephalopod prey, but Taonius pavo (12%) and Histioteuthis reversa (9%) also represented a substantial part of the diet. A large number of otoliths belonging to the fish Scopelogadus beanii were present (25%). These results differ from reports of the stomach contents of short‐finned pilot whales from the Pacific coast in which neritic species dominate the diet. Our findings also suggest that there is a considerable difference between the diet of short‐ and long‐finned pilot whales (Globicephala melas) in the western North Atlantic. The latter feed predominantly on the long‐finned squid (Loligo pealei) whereas the former feed on deep‐water species. Our results indicate the whales fed primarily off the continental shelf prior to stranding.     

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

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 whale's 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.     

The density of odontocete integument depends on blubber lipid composition and temperature

Cetacean integument serves many functional roles, including contribution to whole body buoyancy. The blubber of the integument of different cetacean species contains varying concentrations of triacylglycerols (TAG) and wax esters (WE); generally, these lipid classes have different densities. Integument can also experience a wide range of temperatures during a dive, so its density may change with depth. The goals of this study were to measure integument density and isolated blubber lipid density in three deep‐diving odontocete species (n = 3–4)—short‐finned pilot whales (Globicephala macrorhynchus), pygmy sperm whales (Kogia breviceps), and Gervais' beaked whales (Mesoplodon europeaus)—at different temperatures (6°C–35°C), and to relate these densities to lipid content and composition. Kogia and Mesoplodon integument and isolated lipids had high WE content (78.7–99.5 wt%) and were less dense (by 1.7%–9.3%) than those of Globicephala, which were composed predominately of TAG. Generally, densities increased as temperature decreased. Changes in integument densities mirrored those of isolated lipid densities, suggesting that blubber lipids are largely responsible for the buoyant properties of cetacean integument. These data demonstrate that the contribution of the integument to whole body density depends on lipid class and temperature, and therefore may provide useful, species‐specific correction factors for diving energetics models.     

A simple photograph‐based approach for discriminating between free‐ranging long‐finned (Globicephala melas) and short‐finned (G. macrorhynchus) pilot whales off the east coast of the United States

Line transect based abundance estimation is complicated for long‐finned (LFPW, Globicephala melas) and short‐finned (SFPW, G. macrorhynchus) pilot whales because of their similarity in appearance and their overlapping summertime range in some areas. We developed a photograph‐based approach to distinguish between species of free‐ranging pilot whales in the northwest Atlantic. We collected skin samples and photographs during the summers of 2004–2007 and used skin samples to distinguish species based on mitochondrial DNA. Relative morphometric measurements from photographs were examined using mixed‐effect models and logistic regression. The best model among 94 candidate models had an overall classification error rate of 2.5%. We tested the presence/absence of pigmentation in four regions of the dorsal body (melon, eye, cape, and saddle) for differences. Pigmentation was present in all four regions in 100% of the SFPWs sampled. Melon patch, blaze, and saddle patch pigmentation were present in 6%, 68%, and 50%, respectively, of the LFPWs, but the cape was completely absent. Both types of analyses provided positive species discrimination of free‐ranging animals. We created a cost‐effective, simple tool which could ultimately assist in providing appropriate management, mitigation, and conservation strategies for both northwest Atlantic species of pilot whales.     

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.     

High site fidelity,strong associations,and long‐term bonds: Short‐finned pilot whales off the island of Hawai‘i

Studies of short‐finned pilot whales suggest they travel in stable mixed‐sex groups composed of strongly associated individuals; however, temporal analyses of social structure are lacking. To examine site fidelity, association patterns, and temporal relationships, we analyzed data from 267 encounters of this species off the island of Hawai‘i from 2003 through 2007, identifying 448 distinctive individuals (68.1% seen more than once). About 72% of the whales were linked by association into a single social network, suggesting the possibility of multiple populations using the area. Sighting histories suggested that only some individuals exhibit high site fidelity. Individuals demonstrated preferential associations and community division was strongly supported by average‐linkage hierarchical cluster analysis of the association data. Nine longitudinally stable social units composed of key individuals and their constant companions were identified. Qualitative assignment of age and sex classes of unit members indicated that some segregation between adult males and female/calf pairs may occur. Temporal analyses of individuals encountered on the same day indicate stable long‐term associations. Differential patterns of residency and site fidelity were unexpected and may be indicative of multiple populations around the main Hawaiian Islands. The presence of a resident population demonstrating strong, long‐term site fidelity and associations off Hawai‘i Island may warrant special management considerations.     

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.     

Notes on New Zealand mammals 6. Second report on the stomach contents of long‐finned pilot whales,Globicephala melas

Abstract

We report data on the stomach contents of the long‐finned pilot whale, Globicephala melas, recovered from a group of whales stranded on Ruakaka Beach, northeastern New Zealand, in November 2006. In nine whales for which identifiable stomach contents were recovered (three that stranded on 10 November and six that stranded on 11 November) prey remains comprised exclusively cephalopod beaks attributed to five squid species. The stomachs of a further two whales contained unidentifiable upper beaks only, while the stomachs of five whales were completely empty. No whale appeared to have been satiated immediately before stranding, given that the maximum biomass of prey recently consumed by any one whale was calculated to be <5 kg. All squids ingested represented oceanic species, found from 50 to 1000 m but more common towards the deeper end of this range. These data both complement and contrast with the only other dietary information available for this species in New Zealand waters, reported from stomach contents of whales stranded on Farewell Spit, South Island in December 2005.     

Assessment of wound healing of tagged gray (Eschrichtius robustus) and blue (Balaenoptera musculus) whales in the eastern North Pacific using long‐term series of photographs

Tags have been used to examine migration routes and habitat use of large whales for >40 yr, however, evaluation of tag wound healing has largely been short‐term, anecdotal or generalized. This study developed methods for systematic photographic assessment of long‐term external consequences of tag placement, to determine potential differences in wound healing between species and tag types and thus advise future tagging efforts to possibly minimize undesirable side effects. Tag site appearance and healing characteristics were evaluated by two reviewers and a time series evaluated by five veterinarians from photographs during 995 postdeployment encounters with 34 gray and 63 blue whales tagged in the North Pacific. Blue whale resightings were less frequent, but spanned a longer time period due to earlier tag deployments than the more frequent gray whale follow‐up observations. Swelling occurred in 74% of reencountered gray whales, with the highest frequency 6 mo postdeployment. Swellings were common in blue whales with early tag designs but rare with current models. Depressions occurred in 82% of gray and 71% of blue whales. This study demonstrates the value of follow‐up studies of tagged animals and systematic scoring of photographs to quantitatively compare tag response.     

Stomach contents of long‐finned pilot whales,Globicephala melas,mass‐stranded on Farewell Spit,Golden Bay in 2005 and 2008

Abstract

New data are reported on the diet of the long‐finned pilot whale, Globicephala melas, based on stomach contents recovered from whales involved in a mass stranding on Farewell Spit, Golden Bay, South Island, on 23 January 2008. The stomachs of 11 whales were examined, from which identifiable prey remains were recovered from six, four females and two males (3.1–5.4 m in length). Prey remains comprised exclusively cephalopod beaks (1–46 beaks per whale), attributed to two genera in two orders: arrow squid, Nototodarus spp. (Teuthoidea: Ommastrephidae), and common octopus, Pinnoctopus cordiformis (Octopoda: Octopodidae). The stomachs of eight whales were infested with parasitic nematodes, with two ulcerated; the stomachs of five whales did not contain any prey remains. These data complement and are comparable to the only other information available for this species from this region, reported from whales mass‐stranded at this same location in December 2005. Lower beak rostral length versus mantle length and biomass regression equations for Nototodarus spp. are reviewed, highlighting the importance of the use of species‐specific regression equations for reconstructing both cephalopod mantle length and biomass from lower beak remains in dietary studies.     

First report on the stomach contents of long‐finned pilot whales,Globicephala melas,stranded in New Zealand

Abstract

Stomach contents of the long‐finned pilot whale, Globicephala melas, are reported for the first time from New Zealand waters. Analyses based on two male and three female whales (2.5–5.3 m in length) that stranded on Farewell Spit, Golden Bay, South Island in December 2005 revealed a diet comprised exclusively of cephalopods (2-33 lower cephalopod beaks per stomach). Two genera of cephalopod from two orders; arrow squid, Nototodarus spp. (Teuthoidea: Ommastrephidae), and common octopus, Pinnoctopus cordiformis (Octopoda: Octopodidae) were represented. A further five pilot whale stomachs were examined and found to be empty.     

Depredation of Patagonian toothfish (Dissostichus eleginoides) by two sympatrically occurring killer whale (Orcinus orca) ecotypes: Insights on the behavior of the rarely observed type D killer whales

Sympatric forms of ecologically distinctive killer whales (Orcinus orca) have been documented worldwide. This study focused on a new case of such sympatric occurrence of the “Crozet” type and the recently described “type D” killer whales off the Crozet Islands. The two ecotypes are morphologically and genetically distinct, but they both depredate the same local longline fishery. We used observational, photo‐identification, and fishing data, collected between 2003 and 2015, to examine differences in their patterns of depredation. Of the 828 sets where ecotype could be confirmed, type D killer whales interacted with 82 (11%) of the sets, including 9 (1%) sets that were simultaneously depredated by both ecotypes. Associations between the two types were never observed. Type D killer whales typically occurred in larger groups and both ecotypes preferentially depredated Patagonian toothfish (Dissostichus eleginoides). GLMM modeling revealed that the probability of type D depredation significantly increased throughout the study period, especially in deep waters, and photo‐identification data suggested that a subset of all individuals were habituating to depredation. This study documents the partitioning of resources between two distinct ecotypes of killer whales and provides preliminary insight into the feeding ecology of the rare type D killer whale.     

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

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.     

Mark‐resight abundance and survival estimation of Indo‐Pacific bottlenose dolphins,Tursiops aduncus,in the Swatch‐of‐No‐Ground,Bangladesh

A mark‐resight analysis under Pollock's robust design was applied to Indo‐Pacific bottlenose dolphins Tursiops aduncus in the Swatch‐of‐No‐Ground (SoNG) submarine canyon, Bangladesh, during the winter seasons of 2005–2009. Information from sightings of photo‐identified individuals (1,144) and unmarked individuals generated abundance estimates of 1,701 (95% confidence interval [CI]= 1,533–1,888), 1,927 (95% CI = 1,851–2,006), 2,150 (95% CI = 1,906–2,425), and 2,239 (95% CI = 1,985–2,524) individuals for seasons 1–4, respectively. This makes the population among the largest assessed of the species. Overall apparent survival was estimated as 0.958 (95% CI = 0.802–0.992). Interseasonal probabilities of transitioning to an unobservable state were estimated as 0.045, 0.363, and 0.300 for years 1–2, 2–3, and 3–4, respectively, and the overall probability of remaining in an unobservable state was 0.688. These probabilities, together with an apparent increase in abundance during the study period, indicate that the identified dolphins are part of a larger superpopulation moving throughout a more extensive geographic area. Of the photo‐identified dolphins, 28.2% exhibited injuries related to entanglements with fishing gear. This implies a strong potential for fatal interactions that could jeopardize the conservation status of the population, which otherwise appears favorable.     

Structure and Dynamics of Minke Whale Surfacing Patterns in the Gulf of St. Lawrence,Canada

Animal behavioral patterns can help us understand physiological and ecological constraints on animals and its influence on fitness. The surfacing patterns of aquatic air-breathing mammals constitute a behavioral pattern that has evolved as a trade-off between the need to replenish oxygen stores at the surface and the need to conduct other activities underwater. This study aims to better understand the surfacing pattern of a marine top predator, the minke whale (Balaenoptera acutorostrata), by investigating how their dive duration and surfacing pattern changes across their activity range. Activities were classified into resting, traveling, surface feeding and foraging at depth. For each activity, we classified dives into short and long dives and then estimated the temporal dependence between dive types. We found that minke whales modified their surfacing pattern in an activity-specific manner, both by changing the expression of their dives (i.e. density distribution) and the temporal dependence (transition probability) between dive types. As the depth of the prey layer increased between activities, the surfacing pattern of foraging whales became increasingly structured, going from a pattern dominated by long dives, when feeding at the surface, to a pattern where isolated long dives were followed by an increasing number of breaths (i.e. short dives), when the whale was foraging at depth. A similar shift in surfacing pattern occurred when prey handling time (inferred from surface corralling maneuvers) increased for surface feeding whales. The surfacing pattern also differed between feeding and non-feeding whales. Resting whales did not structure their surfacing pattern, while traveling whales did, possibly as a way to minimize cost of transport. Our results also suggest that minke whales might balance their oxygen level over multiple, rather than single, dive cycles.     

Occurrence of false killer whales (Pseudorca crassidens) and their association with common bottlenose dolphins (Tursiops truncatus) off northeastern New Zealand

On a global scale, false killer whales (Pseudorca crassidens) remain one of the lesser‐known delphinids. The occurrence, site fidelity, association patterns, and presence/absence of foraging in waters off northeastern New Zealand are examined from records collected between 1995 and 2012. The species was rarely encountered; however, of the 61 distinctive, photo‐identified individuals, 88.5% were resighted, with resightings up to 7 yr after initial identification, and movements as far as 650 km documented. Group sizes ranged from 20 to ca. 150. Results indicate that all individuals are linked in a single social network. Most observations were recorded in shallow (<100 m) nearshore waters. Occurrence in these continental shelf waters is likely seasonal, coinciding with the shoreward flooding of a warm current. During 91.5% of encounters, close interspecific associations with common bottlenose dolphins (Tursiops truncatus) were observed. Photo‐identification reveals repeat inter‐ and intraspecific associations among individuals with 34.2% of common bottlenose dolphins resighted together with false killer whales over 1,832 d. While foraging was observed during 39.5% of mixed‐species encounters, results suggest that social and antipredatory factors may also play a role in the formation of these mixed‐species groups.     

APPARENT LATERALIZED BEHAVIOR IN GRAY WHALES FEEDING OFF THE CENTRAL BRITISH COLUMBIA COAST

A digital acoustic recording tag was used to examine the 3‐D orientation of gray whales feeding along the central British Columbia coast. A total of 96 feeding dives were recorded from six different whales. More than half (53.1%) of the whales' bottom time was spent rolled at an angle greater than 45°. Whales rolled an average of 2.9 times per feeding dive, and rolling behavior was often accompanied by a negative pitch angle. Out of 282 recorded rolls, 274 (97.2%) were to the right. Likewise, 98.5% of the total time spent rolled at an angle greater than 45° was spent rolled to the right. The gray whales in this study showed a significant right‐side bias on both an individual (P≤ 0.009) and group level (P < 0.001). Based on the findings of this study and previous reports of uneven baleen wear ( Kasuya and Rice 1970 ), it is proposed that gray whales exhibit a population‐wide right‐side rolling bias similar in character to the 90/10 split of right handedness in humans.     

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.