Cooperative hunting behavior,prey selectivity and prey handling by pack ice killer whales (Orcinus orca), type B,in Antarctic Peninsula waters

Currently, there are three recognized ecotypes (or species) of killer whales (Orcinus orca) in Antarctic waters, including type B, a putative prey specialist on seals, which we refer to as “pack ice killer whale” (PI killer whale). During January 2009, we spent a total of 75.4 h observing three different groups of PI killer whales hunting off the western Antarctic Peninsula. Observed prey taken included 16 seals and 1 Antarctic minke whale (Balaenoptera bonaerensis). Weddell seals (Leptonychotes weddellii) were taken almost exclusively (14/15 identified seal kills), despite the fact that they represented only 15% of 365 seals identified on ice floes; the whales entirely avoided taking crabeater seals (Lobodon carcinophaga; 82% relative abundance) and leopard seals (Hydrurga leptonyx; 3%). Of the seals killed, the whales took 12/14 (86%) off ice floes using a cooperative wave‐washing behavior; they produced 120 waves during 22 separate attacks and successfully took 12/16 (75%) of the Weddell seals attacked. The mean number of waves produced per successful attack was 4.1 (range 1–10) and the mean attack duration was 30.4 min (range 15–62). Seal remains that we examined from one of the kills provided evidence of meticulous postmortem prey processing perhaps best termed “butchering.”     

Predation by killer whales (Orcinus orca) and the evolution of whistle loss and narrow-band high frequency clicks in odontocetes

A disparate selection of toothed whales (Odontoceti) share striking features of their acoustic repertoires including the absence of whistles and high frequency but weak (low peak-to-peak source level) clicks that have a relatively long duration and a narrow bandwidth. The non-whistling, high frequency click species include members of the family Phocoenidae, members of one genus of delphinids, Cephalorhynchus, the pygmy sperm whale, Kogia breviceps, and apparently the sole member of the family Pontoporiidae. Our review supports the 'acoustic crypsis' hypothesis that killer whale predation risk was the primary selective factor favouring an echolocation and communication system in cephalorhynchids, phocoenids and possibly Pontoporiidae and Kogiidae restricted to sounds that killer whales hear poorly or not at all (< 2 and > 100 kHz).     

KILLER WHALE PREDATION ON BELUGAS IN COOK INLET, ALASKA: IMPLICATIONS FOR A DEPLETED POPULATION

Killer whale predation on belugas in Cook Inlet, Alaska, has become a concern since the decline of these belugas was documented during the 1990s. Accordingly, killer whale sightings were compiled from systematic surveys, observer databases, and anecdotal accounts. Killer whales have been relatively common in lower Cook Inlet (at least 100 sightings from 1975 to 2002), but in the upper Inlet, north of Kalgin Island, sightings were infrequent (18 in 27 yr), especially prior to the 1990s. Beach cast beluga carcasses with teeth marks and missing flesh also provided evidence of killer whale predation. Most observed killer whale/beluga interactions were in the upper Inlet. During 11 of 15 observed interactions, belugas were obviously injured or killed, either through direct attacks or indirectly as a result of stranding. Assuming at least one beluga mortality occurred during the other four encounters, we can account for 21 belugas killed between 1985 and 2002. This would suggest a minimum estimate of roughly 1/yr and does not include at least three instances where beluga calves accompanied an adult that was attacked.     

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 (Orcinus orca) predation in a multi-prey system

Predation can regulate prey numbers but predator behaviour in multiple-prey systems can complicate understanding of control mechanisms. We investigate killer whale (Orcinus orca) predation in an ocean system where multiple marine mammal prey coexist. Using stochastic models with Monte-Carlo simulations, we test the most likely outcome of predator selection and compare scenarios where killer whales: (1) focus predation on larger prey which presumably offer more energy per effort, (2) generalize by feeding on prey as encountered during searches, or (3) follow a mixed foraging strategy based on a combination of encounter rate and prey size selection. We test alternative relationships within the Hudson Bay geographic region, where evidence suggests killer whales seasonally concentrate feeding activities on the large-bodied bowhead whale (Balaena mysticetus). However, model results indicate that killer whales do not show strong prey specialization and instead alternatively feed on narwhal (Monodon monoceros) and beluga (Delphinapterus leucas) whales early and late in the ice-free season. Evidence does support the conjecture that during the peak of the open water season, killer whale predation can differ regionally and feeding techniques can focus on bowhead whale prey. The mixed foraging strategy used by killer whales includes seasonal predator specialization and has management and conservation significance since killer whale predation may not be constrained by a regulatory functional response.     

Fight or flight: antipredator strategies of baleen whales

• 1 The significance of killer whale Orcinus orca predation on baleen whales (Mysticeti) has been a topic of considerable discussion and debate in recent years. Discourse has been constrained by poor understanding of predator‐prey dynamics, including the relative vulnerability of different mysticete species and age classes to killer whales and how these prey animals avoid predation. Here we provide an overview and analysis of predatory interactions between killer whales and mysticetes, with an emphasis on patterns of antipredator responses.
• 2 Responses of baleen whales to predatory advances and attacks by killer whales appear to fall into two distinct categories, which we term the fight and flight strategies. The fight strategy consists of active physical defence, including self‐defence by single individuals, defence of calves by their mothers and coordinated defence by groups of whales. It is documented for five mysticetes: southern right whale Eubalaena australis, North Atlantic right whale Eubalaena glacialis, bowhead whale Balaena mysticetus, humpback whale Megaptera novaeangliae and grey whale Eschrichtius robustus. The flight strategy consists of rapid (20–40 km/h) directional swimming away from killer whales and, if overtaken and attacked, individuals do little to defend themselves. This strategy is documented for six species in the genus Balaenoptera.
• 3 Many aspects of the life history, behaviour and morphology of mysticetes are consistent with their antipredator strategy, and we propose that evolution of these traits has been shaped by selection for reduced predation. Fight species tend to have robust body shapes and are slow but relatively manoeuvrable swimmers. They often calve or migrate in coastal areas where proximity to shallow water provides refuge and an advantage in defence. Most fight species have either callosities (rough and hardened patches of skin) or encrustations of barnacles on their bodies, which may serve (either primarily or secondarily) as weapons or armour for defence. Flight species have streamlined body shapes for high‐speed swimming and they can sustain speeds necessary to outrun pursuing killer whales (>15–20 km/h). These species tend to favour pelagic habitats and calving grounds where prolonged escape sprints from killer whales are possible.
• 4 The rarity of observed successful attacks by killer whales on baleen whales, especially adults, may be an indication of the effectiveness of these antipredator strategies. Baleen whales likely offer low profitability to killer whales, relative to some other marine mammal prey. High‐speed pursuit of flight species has a high energetic cost and a low probability of success while attacks on fight species can involve prolonged handling times and a risk of serious injury.

     

Interactions between killer whales (<Emphasis Type="Italic">Orcinus orca</Emphasis>) and Steller sea lions (<Emphasis Type="Italic">Eumetopias jubatus</Emphasis>) in the vicinity of Brat Chirpoev Island,Kuril Islands

The behaviors of breeding Steller sea lions in response to encounters with killer whales near the shore were observed on Brat Chirpoev Island, Kuril Islands between May and July 2002–2007. Approaches by killer whales and sea lion behavior was observed visually and recorded. Killer whales approached the rookery 104 times during the entire period of observations (289 days). In most cases (n = 95), beached sea lions did not show any apparent reactions to the presence of killer whales, and there were no observed interactions. Sea lions showed agitation during nine of the approaches; five of these events were considered to be predation attempts. The killer whales attacked the sea lions three times, however all the attacks were unsuccessful. We recorded two different types of responses towards the killer whales: (1) beaching on the shore (three times) and (2) mass exodus from the rookery with subsequent formation of a tight, actively swimming and vocalizing group (six times). The latter is the first recorded observation of this behavior for Steller sea lions. The observation suggests a low degree of interactions between these two species near the studied rookery. Despite the numerous observations of killer whales near the rookery, there were no observations of direct predation on sea lions. It is likely the killer whale predation has little or no direct impact on the Steller sea lion population on Brat Chirpoev Islands during the breeding period.     

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.     

KILLER WHALES AND MARINE MAMMAL TRENDS IN THE NORTH PACIFIC—A RE-EXAMINATION OF EVIDENCE FOR SEQUENTIAL MEGAFAUNA COLLAPSE AND THE PREY-SWITCHING HYPOTHESIS

Springer et al . (2003) contend that sequential declines occurred in North Pacific populations of harbor and fur seals, Steller sea lions, and sea otters. They hypothesize that these were due to increased predation by killer whales, when industrial whaling's removal of large whales as a supposed primary food source precipitated a prey switch. Using a regional approach, we reexamined whale catch data, killer whale predation observations, and the current biomass and trends of potential prey, and found little support for the prey-switching hypothesis. Large whale biomass in the Bering Sea did not decline as much as suggested by Springer et al ., and much of the reduction occurred 50–100 yr ago, well before the declines of pinnipeds and sea otters began; thus, the need to switch prey starting in the 1970s is doubtful. With the sole exception that the sea otter decline followed the decline of pinnipeds, the reported declines were not in fact sequential. Given this, it is unlikely that a sequential megafaunal collapse from whales to sea otters occurred. The spatial and temporal patterns of pinniped and sea otter population trends are more complex than Springer et al . suggest, and are often inconsistent with their hypothesis. Populations remained stable or increased in many areas, despite extensive historical whaling and high killer whale abundance. Furthermore, observed killer whale predation has largely involved pinnipeds and small cetaceans; there is little evidence that large whales were ever a major prey item in high latitudes. Small cetaceans (ignored by Springer et al .) were likely abundant throughout the period. Overall, we suggest that the Springer et al . hypothesis represents a misleading and simplistic view of events and trophic relationships within this complex marine ecosystem.     

Linking killer whale survival and prey abundance: food limitation in the oceans' apex predator?

Killer whales (Orcinus orca) are large predators that occupy the top trophic position in the world''s oceans and as such may have important roles in marine ecosystem dynamics. Although the possible top-down effects of killer whale predation on populations of their prey have received much recent attention, little is known of how the abundance of these predators may be limited by bottom-up processes. Here we show, using 25 years of demographic data from two populations of fish-eating killer whales in the northeastern Pacific Ocean, that population trends are driven largely by changes in survival, and that survival rates are strongly correlated with the availability of their principal prey species, Chinook salmon (Oncorhynchus tshawytscha). Our results suggest that, although these killer whales may consume a variety of fish species, they are highly specialized and dependent on this single salmonid species to an extent that it is a limiting factor in their population dynamics. Other ecologically specialized killer whale populations may be similarly constrained to a narrow range of prey species by culturally inherited foraging strategies, and thus are limited in their ability to adapt rapidly to changing prey availability.     

Analysis of a Blainville's beaked whale's movement response to playback of killer whale vocalizations

Increasing evidence links exposure to Navy sonar with certain mass stranding events of deep diving beaked whales. Although the cause of these strandings is unknown, one theory suggests that the animals confuse the sonar signals with vocalizations of killer whales, a known predator. Here we analyze the movement patterns of a tagged female Blainville's beaked whale in reaction to playback of killer whale predation calls. During a deep foraging dive, the whale was exposed to a playback of killer whale vocalizations with the source level slowly increased until the whale prematurely ceased foraging. The heading data from the tag were analyzed using a rotation test with a likelihood ratio calculated for a nonparametric kernel density estimate. We found a significant difference (P < 0.005) in the distribution of Δheading (the change in heading averaged over 200 s) after the cessation of the killer whale playback. A test of the angular standard deviation (SD) of the Δheading showed that after the playback, the SD was significantly reduced (P = 0.0064), which indicates that the animal maintained a straighter than normal course for an extended period of time. The prolonged directed avoidance response observed here suggests a behavioral reaction that could pose a risk factor for stranding.     

DIETS OF FIN, SEI, AND SPERM WHALES IN BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS, 1963–1967

Diets of fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), and sperm whales ( Physeter macrocephalus ) were estimated from the stomach contents of individuals killed along the British Columbia coast from 1963 to 1967. The dominant prey types of fin whales were euphausiids, with minor contributions from copepods and fish. Sei whale stomachs contained primarily copepods in three years, whereas euphausiids or a variety of fish dominated the diet in the other two years. Sperm whales consumed primarily North Pacific giant squid ( Moroteuthis robusta ), but secondary prey differed between males and females. Female sperm whales frequently consumed ragfish ( Icosteus spp.) and other fish, whereas the male diet also contained rockfish ( Sebastes spp.). The high abundance of euphausiids along the British Columbia coast likely contributed to the presence of a summer resident population of fin whales. The high abundance of large copepods farther north probably influenced the migration of sei whales through the offshore waters of British Columbia. Sperm whale stomach contents differed by sex reflecting location and possibly breeding behaviors.     

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.     

Spatio-temporal distribution of sperm whales (Physeter macrocephalus) off Kaikoura,New Zealand,in relation to bathymetric features

Using a shore-based station we monitored the position of sperm whales (Physeter macrocephalus) within the Kaikoura submarine canyon from 2010 to 2012. We tracked sperm whales using a theodolite station for a total of 290 days. We extracted the distance from the nearest coast, the depth and the bathymetric slope using ArcGIS 10.1. We estimated the seasonal spatial distribution of sperm whales using general additive models. The distribution varied significantly between seasons; individuals were found in deeper water and further offshore in the spring than in winter. This study improved our understanding of the variability of sperm whale distribution patterns off Kaikoura. We determined that the distribution was linked to the bathymetric features and we hypothesized that whales adapted their use of the submarine canyon in relation to food aggregation. We would encourage further studies to evaluate the sperm whale relationship with oceanographic variables off Kaikoura.     

Prey items and predation behavior of killer whales (Orcinus orca) in Nunavut, Canada based on Inuit hunter interviews

Background

Killer whales (Orcinus orca) are the most widely distributed cetacean, occurring in all oceans worldwide, and within ocean regions different ecotypes are defined based on prey preferences. Prey items are largely unknown in the eastern Canadian Arctic and therefore we conducted a survey of Inuit Traditional Ecological Knowledge (TEK) to provide information on the feeding ecology of killer whales. We compiled Inuit observations on killer whales and their prey items via 105 semi-directed interviews conducted in 11 eastern Nunavut communities (Kivalliq and Qikiqtaaluk regions) from 2007-2010.

Results

Results detail local knowledge of killer whale prey items, hunting behaviour, prey responses, distribution of predation events, and prey capture techniques. Inuit TEK and published literature agree that killer whales at times eat only certain parts of prey, particularly of large whales, that attacks on large whales entail relatively small groups of killer whales, and that they hunt cooperatively. Inuit observations suggest that there is little prey specialization beyond marine mammals and there are no definitive observations of fish in the diet. Inuit hunters and elders also documented the use of sea ice and shallow water as prey refugia.

Conclusions

By combining TEK and scientific approaches we provide a more holistic view of killer whale predation in the eastern Canadian Arctic relevant to management and policy. Continuing the long-term relationship between scientists and hunters will provide for successful knowledge integration and has resulted in considerable improvement in understanding of killer whale ecology relevant to management of prey species. Combining scientists and Inuit knowledge will assist in northerners adapting to the restructuring of the Arctic marine ecosystem associated with warming and loss of sea ice.     

Killer whale ecotypes: is there a global model?

Killer whales, Orcinus orca, are top predators occupying key ecological roles in a variety of ecosystems and are one of the most widely distributed mammals on the planet. In consequence, there has been significant interest in understanding their basic biology and ecology. Long‐term studies of Northern Hemisphere killer whales, particularly in the eastern North Pacific (ENP), have identified three ecologically distinct communities or ecotypes in that region. The success of these prominent ENP studies has led to similar efforts at clarifying the role of killer whale ecology in other regions, including Antarctica. In the Southern Hemisphere, killer whales present a range of behavioural, social and morphological characteristics to biologists, who often interpret this as evidence to categorize individuals or groups, and draw general ecological conclusions about these super‐predators. Morphologically distinct forms (Type A, B, C, and D) occur in the Southern Ocean and studies of these different forms are often presented in conjunction with evidence for specialised ecology and behaviours. Here we review current knowledge of killer whale ecology and ecotyping globally and present a synthesis of existing knowledge. In particular, we highlight the complexity of killer whale ecology in the Southern Hemisphere and examine this in the context of comparatively well‐studied Northern Hemisphere populations. We suggest that assigning erroneous or prefatory ecotypic status in the Southern Hemisphere could be detrimental to subsequent killer whale studies, because unsubstantiated characteristics may be assumed as a result of such classification. On this basis, we also recommend that ecotypic status classification for Southern Ocean killer whale morphotypes be reserved until more evidence‐based ecological and taxonomic data are obtained.     

Mortality rate acceleration and post-reproductive lifespan in matrilineal whale species

The strength of selection to increase the span of a life stage is dependent upon individuals at that stage being able to contribute towards individual fitness and the probability of their surviving to that stage. Complete reproductive cessation and a long post-reproductive female lifespan as found in humans are also found in killer whale (Orcinus orca) and short-finned pilot whale (Globicephala macrorhynchus), but not in the long-finned pilot whale (Globicephala melaena). Each species forms kin-based, stable matrilineal groups and exhibits kin-directed behaviours that could increase inclusive fitness. Here, the initial mortality rate and mortality rate-doubling time of females of these three closely related whale species are compared. The initial mortality rate shows little variation among pilot whale species; however mortality rate accelerates almost twice as fast in the long-finned pilot whale as it does in killer whale and short-finned pilot whale. Selection for a long post-reproductive female lifespan in matrilineal whales may therefore be determined by the proportion of females surviving past the point of reproductive cessation.     

WHY DO BALEEN WHALES MIGRATE?1

The annual migrations of baleen whales are a conspicuous but unexplained feature of their behavioral repertoire. Some hypotheses offered to explain whale migration focus on direct benefits to the calf (thermoregulation, calm water) and some do not (resource tracking, and the “evolutionary holdover” hypothesis). Here, we suggest that a major selective advantage to migrating pregnant female baleen whales is a reduced risk of killer whale (Orcinus orca) predation on their newborn calves in low-latitude waters. Killer whale abundance in high latitudes is substantially greater than that in lower latitudes, and most killer whales do not appear to migrate with baleen whales. We suggest that the distribution of killer whales is determined more by their primary marine mammal prey, pinnipeds, and that following the baleen whale migrations would remove them from their pinniped prey. There are problems with all current hypotheses, most of which stem from a lack of directed research. We explore variation in migratory habits between species, populations, and individuals that may provide a “natural laboratory” for discriminating among the competing hypotheses.     

OPEN-BOAT WHALING ON THE STRAITS OF GIBRALTAR GROUND AND ADJACENT WATERS

Logbooks ( n = 317) from whaling expeditions made in the North Atlantic during the 19th century were examined to investigate activity in the Gibraltar Straits grounds. At least forty expeditions of whaling vessels from European and American ports visited the area. In all cases the main target was the sperm whale, but pilot whales, dolphins, sea turtles, and even a blue whale were also taken. Whaling effort concentrated on the Atlantic side of the Straits; only two expeditions ventured into the Mediterranean Sea, obtaining negligible catches. The whaling season extended during spring and summer and peaked in June–July. This seasonality appeared not to be governed by changes in whale density but by the trade winds necessary to sail southward or westward to cross the Atlantic. Searching effort continued while trying out, but the rate of sighting cetaceans was about half that of searching periods. However, the rate of sighting or capturing a sperm whale remained unchanged during processing, probably because the gregarious habits of the species produced clumping of catches. For every whale secured, 1.31 whales were struck. After correcting for struck but lost whales and for "gammed" vessels, the minimum number of removals of sperm whales during 1862–1889 is estimated at 237.     

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.