REACTIONS OF HUMPBACK WHALES TO SKIN BIOPSY SAMPLING ON A WEST INDIES BREEDING GROUND

Abstract: Reactions of humpback whales, Megaptera novaeangliae , to the taking of skin biopsies and to associated activities were studied on one of their principal West Indies breeding grounds on Silver Bank (Dominican Republic). Results were in some cases different from those reported from a similar study of this species in a high-latitude feeding area. Almost half (44.1%) of 565 biopsied whales showed no immediate reaction to a hit, while a further 22.5% showed only low-level reactions. A total of 375 (87.8%) of 427 misses involved no reaction. Only one strong reaction was recorded. Behavior changes were recorded following 31 (5.5%) of 569 hits, and 18 (4.5%) of 404 misses. Evasive behavior related to vessel approach was exhibited prior to 72 (12.0%) of 598 hits and 100 (24.1%) of 415 misses. Mothers showed significantly fewer reactions to hits than other whales, and a similar frequency and type of behavior changes, although they tended to be more evasive before a shot was made. Presumed males in competitive groups also showed significantly fewer reactions to shots, and very few behavior changes. Overall, this study supports the belief that the biopsy itself has little effect on a whale and that, if the associated vessel approach is conducted with care, samples can usually be taken with minimal disturbance to the target animal. However, approaches may affect the probability of obtaining fluke photographs for individual identification.     

OBTAINING SKIN SAMPLES FROM LIVING SPERM WHALES

Samples of sperm whale skin, useful for modern molecular analyses of DNA, can be obtained from living animals either by collecting skin sloughed naturally by the whales, or by using biopsy darts fired from crossbows or compound bows. Sloughed skin was found frequently in warm waters, and particular samples could often be linked to photographs which enabled individuals to be identified. However, sloughed skin seemed less available at higher latitudes. Two types of darts were found to collect skin but collected samples were very small (<4 mm²) and insufficient for repeated DNA fingerprinting analyses. Sperm whales always reacted to darting by "startling" and showing changes of behavior over the next few minutes, but we found no indications of longer-term effects. In warm water studies, collection of sloughed skin seems to be generally effective, but for samples of sperm whale tissue at high latitudes modifications could probably be made to either of the darts in order to obtain larger-sized samples.     

KILLER WHALE ATTACKS ON MINKE WHALES: PREY CAPTURE AND ANTIPREDATOR TACTICS

We describe nine incidents of predation or attempted predation of minke whales ( Balaenoptera acutorostrata ) by mammal-hunting "transient" killer whales ( Orcinus orca ) in coastal waters of British Columbia, Washington, and southeastern Alaska. Pursuits of minke whales were characterized by prolonged chases on a straight heading at velocities of 15–30 km/h. In four of the nine cases the adultsized minke whale gradually outdistanced the killer whales, which abandoned the high-speed pursuit after 0.5–1 h. In one case the minke beached itself and died. Four attacks were successful. In one instance a subadult minke was killed in open water following a chase. In two cases the fleeing minke entered a confined bay and was killed by the killer whales. One adult minke was taken after apparently attempting to seek cover beside a large sailboat. Minke whales made no attempt to physically defend themselves and were killed by repeated ramming or by asphyxiation. Although killer whales are capable of sprinting speeds greater than those of minke whales, it appears that adult minkes can maintain higher sustained speeds and evade capture if sufficient space for an extended escape trajectory is available. Successful predation of minke whales in coastal waters is rare compared to pinnipeds and small cetaceans, the main prey of transient killer whales.     

KILLER WHALE PREDATION ON SPERM WHALES: OBSERVATIONS AND IMPLICATIONS

In October 1997 we observed a herd of approximately 35 killer whales ( Orcinus orca ) attack a pod of nine sperm whales ( Physeter macrocephalus ) 130 km off the coast of central California. During the four hours we watched, adult female killer whales, including some with calves, attacked in waves of four to five animals in what was apparently a "wound and withdraw" strategy. Adult male killer whales stood by until the very end when one charged in and quickly killed a seriously wounded sperm whale that had been separated from the group. The sperm whales appeared largely helpless: their main defensive behavior was the formation of a rosette ("marguerite"-heads together, tails out). When the killer whales were successful in pulling an individual out of the rosette, one or two sperm whales exposed themselves to increased attack by leaving the rosette, flanking the isolated individual, and leading it back into the formation. Despite these efforts, one sperm whale was killed and eaten and the rest were seriously, perhaps mortally, wounded. We also present details of two other encounters between sperm whales and killer whales that we observed. Although sperm whales, because of various behavioral and morphological adaptations, were previously thought to be immune to predation, our observations clearly establish their vulnerability to killer whales. We suggest that killer whale predation has potentially been an important, and underrated, selective factor in the evolution of sperm whale ecology, influencing perhaps the development of their complex social behavior and at-sea distribution patterns.     

Humpback whale (Megaptera novaeangliae) and killer whale (Orcinus orca) feeding aggregations for foraging on herring (Clupea harengus) in Northern Norway

Aggregations of predators on food patches have been documented for both terrestrial and marine animals. Here, we documented for the first time, and investigated, non-predatory aggregations occurring between humpback whales (Megaptera novaeangliae) and killer whales (Orcinus orca) while feeding on wintering Norwegian spring spawning herring (Clupea harengus) in Andfjord, northern Norway. Observational data were collected during 109 opportunistic surveys through three seasons 2013–2016. Killer whales were observed feeding on 59 occasions, with one to three humpback whales involved in 47 of these feeding events (79.7%), and there was an increased probability of finding feeding humpback whales when feeding killer whales also were observed. With killer whales identified as the initiating species in 94.4% of the feeding aggregations for which the first species was known, and with humpback whales joining and feeding on the fish ball afterwards, we suggest that humpback whales may benefit more from these aggregations than the opposite.     

The Relationship between Vessel Traffic and Noise Levels Received by Killer Whales (Orcinus orca)

Whale watching has become increasingly popular as an ecotourism activity around the globe and is beneficial for environmental education and local economies. Southern Resident killer whales (Orcinus orca) comprise an endangered population that is frequently observed by a large whale watching fleet in the inland waters of Washington state and British Columbia. One of the factors identified as a risk to recovery for the population is the effect of vessels and associated noise. An examination of the effects of vessels and associated noise on whale behavior utilized novel equipment to address limitations of previous studies. Digital acoustic recording tags (DTAGs) measured the noise levels the tagged whales received while laser positioning systems allowed collection of geo-referenced data for tagged whales and all vessels within 1000 m of the tagged whale. The objective of the current study was to compare vessel data and DTAG recordings to relate vessel traffic to the ambient noise received by tagged whales. Two analyses were conducted, one including all recording intervals, and one that excluded intervals when only the research vessel was present. For all data, significant predictors of noise levels were length (inverse relationship), number of propellers, and vessel speed, but only 15% of the variation in noise was explained by this model. When research-vessel-only intervals were excluded, vessel speed was the only significant predictor of noise levels, and explained 42% of the variation. Simple linear regressions (ignoring covariates) found that average vessel speed and number of propellers were the only significant correlates with noise levels. We conclude that vessel speed is the most important predictor of noise levels received by whales in this study. Thus, measures that reduce vessel speed in the vicinity of killer whales would reduce noise exposure in this population.     

Causes and consequences of marine mammal population declines in southwest Alaska: a food-web perspective

Populations of sea otters, seals and sea lions have collapsed across much of southwest Alaska over the past several decades. The sea otter decline set off a trophic cascade in which the coastal marine ecosystem underwent a phase shift from kelp forests to deforested sea urchin barrens. This interaction in turn affected the distribution, abundance and productivity of numerous other species. Ecological consequences of the pinniped declines are largely unknown. Increased predation by transient (marine mammal-eating) killer whales probably caused the sea otter declines and may have caused the pinniped declines as well. Springer et al. proposed that killer whales, which purportedly fed extensively on great whales, expanded their diets to include a higher percentage of sea otters and pinnipeds following a sharp reduction in great whale numbers from post World War II industrial whaling. Critics of this hypothesis claim that great whales are not now and probably never were an important nutritional resource for killer whales. We used demographic/energetic analyses to evaluate whether or not a predator–prey system involving killer whales and the smaller marine mammals would be sustainable without some nutritional contribution from the great whales. Our results indicate that while such a system is possible, it could only exist under a narrow range of extreme conditions and is therefore highly unlikely.     

Competing conservation objectives for predators and prey: estimating killer whale prey requirements for Chinook salmon

Ecosystem-based management (EBM) of marine resources attempts to conserve interacting species. In contrast to single-species fisheries management, EBM aims to identify and resolve conflicting objectives for different species. Such a conflict may be emerging in the northeastern Pacific for southern resident killer whales (Orcinus orca) and their primary prey, Chinook salmon (Oncorhynchus tshawytscha). Both species have at-risk conservation status and transboundary (Canada-US) ranges. We modeled individual killer whale prey requirements from feeding and growth records of captive killer whales and morphometric data from historic live-capture fishery and whaling records worldwide. The models, combined with caloric value of salmon, and demographic and diet data for wild killer whales, allow us to predict salmon quantities needed to maintain and recover this killer whale population, which numbered 87 individuals in 2009. Our analyses provide new information on cost of lactation and new parameter estimates for other killer whale populations globally. Prey requirements of southern resident killer whales are difficult to reconcile with fisheries and conservation objectives for Chinook salmon, because the number of fish required is large relative to annual returns and fishery catches. For instance, a U.S. recovery goal (2.3% annual population growth of killer whales over 28 years) implies a 75% increase in energetic requirements. Reducing salmon fisheries may serve as a temporary mitigation measure to allow time for management actions to improve salmon productivity to take effect. As ecosystem-based fishery management becomes more prevalent, trade-offs between conservation objectives for predators and prey will become increasingly necessary. Our approach offers scenarios to compare relative influence of various sources of uncertainty on the resulting consumption estimates to prioritise future research efforts, and a general approach for assessing the extent of conflict between conservation objectives for threatened or protected wildlife where the interaction between affected species can be quantified.     

The relationship between the acoustic behaviour and surface activity of killer whales (<Emphasis Type="Italic">Orcinus orca</Emphasis>) that feed on herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>)

We describe the acoustic behaviour of piscivorous killer whales in Norwegian and Icelandic waters. Whales were assigned to one of three activities (feeding, travelling or other), and sound recordings were made in their proximity with a single hydrophone and a digital audiotape (DAT) recorder. A quantitative analysis of the production of pulsed calls, whistles and echolocation clicks in the three activities revealed that there was a significant effect of activity on the production of these sound types. Both killer whales in Icelandic and Norwegian waters produced high rates of clicks and calls during feeding and low rates of click, calls and whistles during travelling. The differences can be used as acoustical markers and provides new possibilities for acoustic monitoring of killer whales in these areas. Based on the similarity between their prey choice, hunting strategies, phenotype and acoustic behaviour, we suggest that the killer whales in Icelandic and Norwegian waters belong to the same ecotype: Scandinavian herring-eating killer whales.     

Vocal activity of tropical dolphins is inhibited by the presence of killer whales,Orcinus orca

Research has suggested killer whale (Orcinus orca) predation may affect cetacean vocal behavior; however, few data exist to test this hypothesis. Data collected during 40,976 km of visual and acoustic shipboard surveys in the tropical Pacific Ocean, including 1,232 detections of 13 species, were examined to determine if changes in dolphin vocal activity could be attributed to the presence of killer whales. Generalized linear models and Random Forest analyses were used to test the hypothesis that dolphin vocal activity was related to the distance and time to the nearest killer whale sighting. Both results show that dolphin vocalizations were inversely correlated with the temporal proximity of killer whales (P < 0.05). Despite the relative rarity of killer whales in the tropics, they appear to influence vocal behavior of nearby dolphin schools. This disruption in communication may not significantly impact interactions necessary for survival in tropical waters where killer whale density is low. However, in temperate climates, where increased productivity supports a greater abundance of killer whales, this interruption in communication may have a greater impact. The lower incidence of whistling dolphins in temperate waters may be related to the greater abundance of killer whales in these areas.     

Killer whales (Orcinus orca) hunting for walruses (Odobenus rosmarus divergens) near Retkyn Spit, Chukotka

Group hunting by killer whales for walruses was observed in August 18, 2008, in the littoral area (3 km from the haulout of walruses, Retkyn Spit, Chukotka). The group of killer whales consisted of seven adults (one adult male did not participate in attacks) and two calves. Based on prey type, these killer whales were mammal-eating. The total duration of their hunt activity was not less than 95 min. The hunt consisted of three phases. The first phase was an attack on the group of walruses and choice of individual prey; the second phase was attacks on the chosen walrus; and the third (final) phase was a decrease in activity of killer whales and leaving group with walrus from sea shore. The main behavioral patterns of killer whales during the hunt were discerned. Two killer whales tried to kill walruses by chasing them and jumping out of the water on the shore. The video analysis of the ??attack phase?? showed that killer whales made 55 attacks on the walrus during 17.3 min. On average, each killer whale attacked the walrus seven times. The attack tactics of killer whales, the number of movements, and the location of killer whales (adults and calves) relative to each other and to the walrus were described. Well coordination of their movements and group actions was observed.     

Occurrence of killer whale Orcinus orca rake marks on Eastern Canada-West Greenland bowhead whales Balaena mysticetus

Killer whales (Orcinus orca) are increasing in occurrence and residence time in the eastern Canadian Arctic (ECA) in part due to a decrease in sea ice associated with global climate change. Killer whales prey on bowhead whales (Balaena mysticetus) of the Eastern Canada-West Greenland (EC-WG) population, but their patterns of predation pressure and effect on the EC-WG population’s ability to recover from historical whaling remain unknown. We analyzed photographs of individual bowhead whale flukes from five regions within the EC-WG population’s geographic range (Cumberland Sound, Foxe Basin, Isabella Bay, Repulse Bay and Disko Bay), taken during 1986 and from 2007 to 2012, to estimate the occurrence of rake marks (parallel scars caused by killer whale teeth). Of 598 identified whales, 10.2 % bore rake marks from killer whales. A higher occurrence of rake marks was found in Repulse and Disko Bays, where primarily adult bowhead whales occur seasonally, than in Foxe Basin, where juveniles and females with calves occur. Older bowheads, which have had greater exposure time to killer whales due to their age, had higher occurrences of rake marks than juveniles and calves, which may indicate that younger whales do not survive killer whale attacks. A high proportion of adult females also had rake marks, perhaps due to protecting their calves from killer whale predation. In order to quantify the effect of killer whales on EC-WG population recovery, further research is needed on the relationship between the occurrence of rake marks and bowhead adult, calf, and juvenile mortality in the ECA, as well as more information about Arctic killer whale ecology.     

The function of multi-pod aggregations of fish-eating killer whales (<Emphasis Type="Italic">Orcinus orca</Emphasis>) in Kamchatka,Far East Russia

In fish-eating North Pacific killer whales, large multi-pod aggregations of up to 100 animals often occur. These aggregations are thought to be reproductive gatherings where mating between members of different pods takes place. However, killer whales are social animals, and the role of these aggregations might also be establishing and maintaining social bonds between pods. Alternatively, it is also possible that multi-pod aggregations are in some way connected with foraging or searching for fish. In this study of killer whales in the western North Pacific, we describe multi-pod aggregations quantitatively and suggest their functional role in the life of fish-eating killer whales. We show that foraging is rare in multi-pod aggregations, whether inter-clan or intra-clan, and thus they are unlikely to play an important role in cooperative foraging. Socialising occurs more frequently in inter-clan rather than in intra-clan aggregations, which suggests the higher arousal level and possible mating during inter-clan aggregations. In summary, multi-pod aggregations of Kamchatka killer whales might be both reproductive assemblages and “clubs” of some kind in which whales gather to establish and maintain social bonds.     

The snail's love-dart delivers mucus to increase paternity

Many of the seemingly bizarre animal behaviours can be understood only by acknowledging the power of sex to shape evolution. A case in point is the so-called love-dart that some terrestrial molluscs shoot at their prospective sexual partners. Given that the likelihood of copulation is not different after solid hits than after complete misses, why do these suitors act so violently towards their chosen mates? Previously, it was shown that successful dart shooting enhances paternity. We conducted an experiment to determine whether the dart achieves its effect by a purely mechanical action or by transferring a bioactive substance. We found that injections of mucus from a gland associated with the dart more than doubled paternity relative to injections of saline. These results support the hypothesis that the dart transfers a substance capable of reconfiguring the spermatophore-receiving organs. While dart shooting probably evolved as the result of sperm competition, a role for cryptic female choice cannot be excluded. Our results imply that if cryptic female choice is operating in this system, it is likely to be based on the properties of the mucus and not on properties of the dart itself. Since we also found evidence of early-male sperm precedence, we conclude that snails can optimize their reproductive success by mating with virgins and shooting their darts accurately.     

UNDERWATER NOISE OF WHALE-WATCHING BOATS AND POTENTIAL EFFECTS ON KILLER WHALES (ORCINUS ORCA), BASED ON AN ACOUSTIC IMPACT MODEL

Underwater noise of whale-watching boats was recorded in the popular killer whale-watching region of southern British Columbia and northwestern Washington State. A software sound propagation and impact assessment model was applied to estimate zones around whale-watching boats where boat noise was audible to killer whales, where it interfered with their communication, where it caused behavioral avoidance, and where it possibly caused hearing loss. Boat source levels ranged from 145 to 169 dB re 1 μPa @ 1 m, increasing with speed. The noise of fast boats was modeled to be audible to killer whales over 16 km, to mask killer whale calls over 14 km, to elicit a behavioral response over 200 m, and to cause a temporary threshold shift (TTS) in hearing of 5 dB after 30–50 min within 450 m. For boats cruising at slow speeds, the predicted ranges were 1 km for audibility and masking, 50 m for behavioral responses, and 20 m for TTS. Superposed noise levels of a number of boats circulating around or following the whales were close to the critical level assumed to cause a permanent hearing loss over prolonged exposure. These data should be useful in developing whale-watching regulations. This study also gave lower estimates of killer whale call source levels of 105–124 dB re 1 μPa.     

Observations of killer whales (Orcinus orca) in the fjords of Chilean Patagonia

Killer whales occur in Chilean waters, but their seasonality, diets, and overall distribution are poorly known. Here, we present data on group composition, site fidelity, and prey species of individual killer whales recorded in 63 sightings between 2004 and 2012 in the Chilean Patagonian fjords. Group sizes were small (mean = 5, SD = 2.5 for calf groups; mean = 3, SD = 1.5 for non-calf groups), and occurrence was significantly lower in summer months. Photographs enabled identification of 55 individuals from natural markings, and all resembled Southern Ocean type A killer whales. The species was transient in the area; the average presence was 1.7 days with 60 % of individuals seen only once. Occupancy was 3–44 days, and low levels of site fidelity were recorded (64 % of individuals were seen in only 1 year). Group composition at short time scales (3 months) remained stable, but we detected changes at longer time scales. Prey included fish, otariids, and seabirds. Twelve individual killer whales showed a broad dietary spectrum: 3 ate otariids and fish, 2 ate birds and otariids, and 7 ate otariids, birds, and possibly fish. Further research is needed to increase basic biological knowledge of these killer whales and to determine the relationship with type A killer whales from the Southern Ocean.     

Site fidelity and association patterns of a rare species: Pygmy killer whales (Feresa attenuata) in the main Hawaiian Islands

Most of what we know about cetacean biology and ecology comes from studies of relatively common species. Despite their distribution throughout the tropics and sub-tropics, pygmy killer whales ( Feresa attenuata ) are rare throughout their range and are one of the most poorly-known species of odontocetes. During a 22-yr study of short-finned pilot whales ( Globicephala macrorhynchus ) off the island of Hawai'i, we opportunistically photo-identified pygmy killer whales whenever encountered. As part of a directed multi-species study throughout the main Hawaiian Islands from 2000 through 2007, we also photo-identified individuals and obtained information on habitat use and behavior. This species was extremely uncommon (representing only 1.2% of odontocete sightings in directed efforts). Given the low encounter rates, assessing trends of this population cannot be feasibly done with line-transect surveys. Despite their rarity, 80% of the distinctive individuals within groups documented off the island of Hawai'i were seen on multiple occasions, individuals were resighted over periods of up to 21 yr, and there was evidence of year-round use of the area. Association analyses indicate stable long-term associations in mixed-sex groups. High resighting rates indicate a small population of island-associated individuals that may be at risk from anthropogenic impacts.     

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.     

FRIENDS OF WILDLIFE SOUND AT NSA

ABSTRACT

The problem of categorization arises in any classification system because classes should be discrete while the characteristics of most natural objects and aspects of nature are more or less gradual. In systematics, this problem usually is solved by creating several levels of categories, such as class, order, family, genus and species. In the existing killer whale discrete call classification, only two levels occur—call type and call subtype. In this paper we describe structural categories at a broader level than call type in the discrete sounds of killer whales and compare these categories between and within vocal clans in a community of resident killer whales from Southeast Kamchatka, Russian Far East, and also with killer whales outside this community. We found four main classes of discrete calls in the repertoire of resident killer whales from Southeast Kamchatka. The calls of Southeast Kamchatka transient killer whales and Sakhalin killer whales do not fall into these classes. This suggests that the resident killer whale community from Southeast Kamchatka has some rules defining the structure of calls which are typical for this community. Consequently, all resident killer whales from Southeast Kamchatka can be said to share the same vocal tradition.     

Use of chemical tracers to assess diet and persistent organic pollutants in Antarctic Type C killer whales

Measuring chemical tracers in tissues of marine predators provides insight into the prey consumed and the predator's contaminant exposure. In this study, samples from Type C killer whales ( Orcinus orca ) biopsied in Antarctica were analyzed for chemical tracers ( i.e. , stable isotopes of carbon and nitrogen, fatty acids, and persistent organic pollutants [POPs]). Profiles of these individual tracers were very different from those of killer whale populations that have been studied in the eastern North and eastern Tropical Pacific. For example, δ¹³C and δ¹⁵N stable isotope values and most POP concentrations were significantly lower in the Antarctic population. In addition, multivariate statistical analyses of both fatty acid and POP profiles found distinctly different patterns for Antarctic Type C whales compared to those from whales in the other populations. Similar assays were conducted on four species of Antarctic marine fish considered potential prey for Type C killer whales. Results were consistent with a diet of fish for Type C whales, but other species ( e.g. , low trophic-level marine mammals or penguins) could not be eliminated as supplemental prey.