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.     

False killer whales (Pseudorca crassidens) around the main Hawaiian Islands: Long-term site fidelity, inter-island movements, and association patterns

Despite their world-wide distribution throughout the tropics and subtropics, false killer whales ( Pseudorca crassidens ) are one of the lesser-known large odontocetes. Genetic evidence indicates a demographically isolated population around the main Hawaiian Islands. We examine site fidelity, movements and association patterns in this population using data from directed surveys and opportunistic photographs from 1986 to 2007. This species was only infrequently encountered, and while found in depths from 38 to 4,331 m, sighting rates were greatest in depths >3,000 m. We photo-identified 152 distinctive individuals. Resighting rates were high, with an average of 76.8% of distinctive individuals within groups documented on more than one occasion. Most (86.6%) were linked by association into a single social network; only one large group (16 distinctive individuals), documented the farthest offshore (42–70 km), did not link by association to that large network, and may be part of an offshore population. Individual movements of up to 283 km were documented, with a large proportion of individuals moving among islands. Individuals were resighted up to 20.1 yr after first being documented, showing long-term fidelity to the islands. Repeated associations among individuals were documented for up to 15 yr, and association analyses indicate preferred associations and strong bonds among individuals.     

Spatial and temporal analysis of killer whale (Orcinus orca) strandings in the North Pacific Ocean and the benefits of a coordinated stranding response protocol

Killer whales (Orcinus orca) are widely distributed throughout the world's oceans, yet little has been documented about their stranding patterns. Knowledge of stranding patterns improves our ability to examine and sample carcasses and provides a foundation for understanding killer whale natural history, diet, reproduction, anthropogenic stressors, emerging diseases, and patterns of unusual mortality. We compiled published and unpublished killer whale stranding data to describe stranding patterns in the North Pacific Ocean. Between 1925 and 2011, 371 stranded killer whales were reported in Japan (20.4%), Russia (3.5%), Alaska (32.0%), British Columbia (27.4%), Washington (4.0%), Oregon (2.7%), California (5.1%), Mexico (3.8%), and Hawaii (0.8%). Strandings occurred at all times of year, but regionally specific seasonal differences were observed. Mortality and annual census data from Northern and Southern Resident populations were extrapolated to estimate that across the North Pacific, an average of 48 killer whales die annually. However, over the last two decades, an average of only 10 killer whale carcasses were recovered annually in this ocean, making each event a rare opportunity for study. Publication of a standardized killer whale necropsy protocol and dedicated funding facilitated the number of complete postmortem necropsies performed on stranded killer whales from 1.6% to 32.2% annually.     

Life history and population dynamics of southern Alaska resident killer whales (Orcinus orca)

Resident (fish eating) killer whales (Orcinus orca) in the North Pacific have been the subject of long‐term studies in several geographical regions. The current study examines population parameters in the southern Alaska resident population from 1984 to 2010 and develops a population model. The southern Alaska resident population ranges from southeastern Alaska through the Kodiak archipelago and contains over 700 individuals. We follow the life histories of 343 identifiable whales in 10 pods from two clans born before and during the study. Population parameters were comparable to those of the British Columbia northern resident population during the 1970s and 1980s, except that age of maturity was approximately one year earlier. The average annual rate of increase was slightly higher in Alaska (3.5%) than for the British Columbia northern residents (2.9%) and probably represents a population at r‐max (maximum rate of growth). Reasons for the high growth rate in Alaska could be a recovery following past anthropogenic mortalities, or more likely, a response to increasing salmon returns in recent decades, resulting in an increase in carrying capacity. The slow maturation and low rate of reproductive response makes these whales slow to recover from natural or anthropogenic catastrophes.     

False killer whales and fisheries interactions in Hawaiian waters: Evidence for sex bias and variation among populations and social groups

We assessed scarring patterns as evidence of fisheries interactions for three populations of false killer whales in Hawai‘i. Bycatch of the pelagic population in the tuna longline fishery exceeds their Potential Biological Removal level. Scarring was assessed by seven evaluators as consistent, possibly consistent, or not consistent with fisheries interactions, and average scores computed. Scores were highest for scarred main Hawaiian Island (MHI) false killer whales, followed by pelagic and Northwestern Hawaiian Island (NWHI) individuals. Considering only whales for which the majority of evaluators scored scarring as consistent revealed significant differences among populations in the percentage of individuals scarred; MHI: 7.5%, pelagic: 0%, NWHI: 0%. Assessment by social cluster for the MHI population showed that 4.2% of Cluster 1, 7.1% of Cluster 2, and 12.8% of Cluster 3 individuals had such scarring, although differences between clusters were not statistically significant. There was a significant sex bias; all sexed individuals (n = 7) with injuries consistent with fisheries interactions were female. The higher proportion of MHI individuals with fisheries‐related scarring suggests that fisheries interactions are occurring at a higher rate in this population. The bias towards females suggests that fisheries‐related mortality has a disproportionate impact on population dynamics.     

Sex‐related site fidelity of humpback whales (Megaptera novaeangliae) to the Fueguian Archipelago feeding area,Chile

We investigated sex‐related site fidelity by humpback whales to the Fueguian Archipelago, a new feeding area in the eastern South Pacific, by examining the resighting histories of 45 males and 39 females recorded from 2003 to 2012. Results indicated an overall annual return to the feeding area of 74.8%, and annual sex ratio is roughly equal in the population. The probability of an individual being resighted across years and in subsequent years was not significantly different for both males and females, however, the proportion of resighting within a year was significantly higher for individual males compared to females. Potential sources of sex‐related bias were analyzed, but none were found to be significant. Greater intraannual resighting frequency for males may reflect sex‐based differences in spatial occupation and short‐range movements due to potential differences in energy budgets.     

Mirror image processing in three marine mammal species: killer whales (Orcinus orca), false killer whales (Pseudorca crassidens) and California sea lions (Zalophus californianus)

Dolphins (Tursiops truncatus) and their relatives might be expected to show mirror-induced contingency checking, a prerequisite to self-recognition, because of their high brain development, their complex social life and their demonstrated abilities in bodily imitation. A study of killer whales'(Orcinus orca) behaviour in front of a mirror is presented, including a mark test. Shorter investigations of mirror behaviour are also described in false killer whales (Pseudorca crassidens) and California sea lions (Zalophus californianus). Contingency checking was present in killer whales and possibly also in false killer whales, but no clear contingency checking was observed in sea lions. The mark test on killer whales suggested that the marked animal anticipated that its image would look different. This study shows that killer whales and false killer whales, like bottlenose dolphins, appear to possess the cognitive abilities required for self-recognition.     

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.     

The diets of humpback whales (Megaptera novaeangliae) on the shelf and oceanic feeding grounds in the western North Pacific inferred from stable isotope analysis

Humpback whales feed on a variety of prey, but significant differences likely occur between regional feeding grounds. In this study, the diets of humpback whales were analyzed by comparing stable isotope ratios in animal tissues at three humpback whale feeding grounds in the Russian Far East: Karaginsky Gulf, Anadyr Gulf, and the Commander Islands. Anadyr Gulf is a neritic zone far from a shelf break, Karaginsky Gulf is a neritic zone close to a shelf break, and the Commander Islands represent an open oceanic ecosystem where whales feed off the shelf break. Samples from the Commander Islands had the lowest mean δ¹³C and δ¹⁵N values (mean ± SE: δ¹³C = ?18.7 ± 0.1, δ¹⁵N = 10.4 ± 0.1) compared to the samples from Karaginsky Gulf (δ¹³C = ?17.2 ± 0.1, δ¹⁵N = 12.7 ± 0.2) and Anadyr Gulf (δ¹³C= ?17.8 ± 0.1, δ¹⁵N = 14.0 ± 0.4). The samples from Anadyr Gulf had the highest δ¹⁵N values, while the samples from Karaginsky Gulf had the highest δ¹³C values. Both δ¹³C and δ¹⁵N values differed significantly among all three areas. Our data support the hypothesis that humpback whales tend to feed on fish in neritic areas and on plankton in deep oceanic waters.     

Combined effects of temperature and food concentration on growth and reproduction of Eodiaptomus japonicus (Copepoda: Calanoida) from Lake Biwa (Japan)

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Geographic variation in the time‐frequency characteristics of high‐frequency whistles produced by killer whales (Orcinus orca)

Investigating intraspecific variation in acoustic signals can indicate the extent of isolation and divergence between populations and adaptations to local environments. Here we analyze the variation in killer whale high‐frequency (>17 kHz) whistles recorded off Norway, Iceland, and in the North Pacific. We used a combination of methods including multivariate comparisons of spectral and temporal parameters and categorization of contours to types. Our results show that spectral and temporal characteristics of high‐frequency whistles recorded in the North Pacific show significant differences from whistles recorded in the Northeast Atlantic, being generally stereotyped, lower in frequency, and slightly longer in duration. Most high‐frequency whistles from the North Pacific were downsweeps, whereas this was one of the least common types recorded in the Northeast Atlantic. The repertoire of whistles recorded in Norway was similar to Iceland, but whistles produced in Norway had significantly lower maximum frequency and frequency range. Most methods were able to discriminate between whistles of the North Pacific and the Northeast Atlantic, but were unable to consistently distinguish whistles from Iceland and Norway. This suggests that macro‐ and microgeographic differences in high‐frequency whistles of killer whales may reflect historical geographic isolation between ocean basins and more recent divergence between adjacent populations.     

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.     

Seasonal acoustic behavior of blue whales (Balaenoptera musculus) in the Gulf of California,Mexico

Six years of passive acoustic monitoring data from the Gulf of California reveal seasonality and movements for the northeastern Pacific blue whales. Three sites were studied, one from the southern (Punta Pescadero) and two from the northern (Isla Tiburon and Canal de Ballenas) regions. A total of 4,953 h were analyzed, and 15,539 blue whale calls were detected, of which 2,723 (18%) were A calls, 11,249 were B calls (72%), and 1,567 were D calls (10%). A and B calls were produced both as song units (2,524) or AB singular calls (2,026). The high rate of songs and their seasonality suggest that the GC is a winter‐breeding ground. A shift from AB call predominance in winter, to D calls in spring and early summer, especially at the entrance of the GC, suggests the importance of this area for reproduction and foraging. Analysis of calling frequency suggests a clear movement of blue whales from the southern region (Punta Pescadero) to the northern regions (Canal de Ballenas and Isla Tiburon), with subsequent southern movement in March. The seasonality and mobility of blue whales in the Gulf of California, inferred from their calling, contributes to the ecological understanding of this population.     

Ecotype and geographical variation in carbon and nitrogen stable isotope values in western North Pacific killer whales (Orcinus orca)

Killer whales are top predators in marine trophic chains, and therefore their feeding preferences can substantially affect the abundance of species on the lower trophic levels. Killer whales are known to feed on many different types of prey from small fish to large whales, but a given killer whale population usually focuses on a specific type of prey. Stable isotope analysis is widely used to study whale diets, because direct observations are often impossible. Killer whale feeding habits in the western North Pacific are poorly studied, and the large-scale stable isotope analysis provides a unique opportunity to gain insights into the trophic links of this top predator. In this study, we compare the δ¹³C and δ¹⁵N stable isotope values from killer whale skin samples obtained in different areas of the western North Pacific from fish-eating (R-type) and mammal-eating (T-type) killer whale ecotypes. The effect of ecotype was highly significant: both carbon and nitrogen stable isotope values were lower in R-type whales than in T-type whales. The geographical variation also affected killer whale stable isotope values due to both the differences in killer whale diet and the variation in baseline stable isotope values across the study areas.     

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.     

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.     

Male‐killer dynamics in Danaus chrysippus (L.) (Lepidoptera: Nymphalidae) in East Africa

Within certain regions in East Africa, the butterfly Danaus chrysippus (L.) shows female‐biased population sex ratio, because of the production by some females of all‐female broods, as a result of infection by maternally inherited, male‐killing bacterium of the genus Spiroplasma. In this study, we describe a 3‐year field survey for the population dynamics of the male‐killing Spiroplasma in D. chrysippus in four independent localities, namely Uganda, Ghana, Sudan and Madagascar. The prevalence of the bacterium was found to show extensive variations at multiple scales among different sites, in various countries, seasons and years. A novel, selection‐based hypothesis was suggested to explain the high variability of male‐killer prevalence over space and time, based on the existence of an adaptive link between larval food‐plant density and the magnitude of resource reallocation fitness advantage for the male‐killer.     

Sexual dimorphism in striped dolphin (Stenella coeruleoalba) crania from South Africa

This study assessed the extent of sexual dimorphism in striped dolphin (Stenella coeruleoalba) cranial size and shape off the South African coast. Dorsal and ventral features of 60 striped dolphin crania from both the western and eastern coasts of South Africa were analyzed using landmark‐based geometric morphometrics. Although there was no evidence of dimorphism in cranial size, evidence for small, but significant, variation in both dorsal and ventral cranial shape was found between the sexes. The observed dimorphism was partly associated with changes in shape around the temporal fossa, occipital condyle and supraoccipital bone, the nasal bone, and the paraoccipital process and basiocciptal. The temporal fossa serves as an attachment point for the temporal muscle, which functions to close the lower jaw, and the occipital area serves as the anterior insertion of the epaxial muscles, which power the upstroke of the flukes during swimming. Both the paraoccipital process and basiocciptal are associated with the functioning of the hyoid apparatus, which serves as an attachment point for muscles and ligaments involved in feeding and sound production. These findings suggest the possibility of differences in diet, foraging behavior, vocalizations, and locomotion between the sexes of this species.     

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.     

Genetic differentiation between humpback whales (Megaptera novaeangliae) from Atlantic and Pacific breeding grounds of South America

Humpback whales wintering in tropical waters along the Atlantic and Pacific coasts of the South American continent are thought to represent distinct populations or “stocks.” Here we present the first analysis of genetic differentiation and estimates of gene flow between these breeding stocks, based on both mitochondrial DNA (mtDNA) control region sequences (465 bp) and 16 microsatellite loci from samples collected off Brazil (n = 277) and Colombia (n = 148), as well as feeding areas near the western Antarctic Peninsula (n = 86). We found significant differentiation between Brazilian and Colombian breeding grounds at both mtDNA (F_ST = 0.058) and microsatellite (F_ST = 0.011) markers and corroborated previous studies showing genetic similarity between humpbacks from Colombia and those from Antarctic Peninsula feeding areas. Estimates of long‐term gene flow between Brazil and Colombia were low to moderate, asymmetrical, and mostly mediated by males. Assignment procedures detected some cases of interchange and individuals of admixed ancestry between breeding grounds, indicating limited mixing of individuals between these stocks. Overall, results highlight the differentiation of humpback whale breeding populations with adjacent feeding grounds. This appears to be a remarkable example of fidelity to seasonal habitat in the absence of any contemporary barriers.