Investigation of foraging habits and prey selection by humpback whales (Megaptera novaeangliae) using acoustic tags and concurrent fish surveys

Tags containing acoustic time-depth transmitters (ATDT) were attached to four humpback whales near Kodiak, Alaska. Tags allowed for whale dive depths to be recorded in real time. Acoustic and mid-water trawl surveys were conducted concurrent with tagging efforts within the study area to quantify available fish resources and describe potential prey selection by humpback whales. Recorded dives were grouped through visual assessment and t -tests. Dives that indicated likely foraging occurred at a mean maximum depth of 106.2 m with 62% of dives occurring between 92 m and 120 m. Acoustic backscatter from fish surveys was attributed to potential humpback prey based on known target strength values and 10 net tows. Capelin comprised 84% of the total potential prey abundance in the region followed by age 0 (12%) and juvenile pollock (2%), and eulachon (<1%). Although horizontally segregated in the region, both capelin and age 0 pollock were distributed at depths exceeding 92 m with maximum abundance between 107 m and 120 m. The four-tagged humpbacks were found to forage in areas with greatest capelin densities but bypassed areas of high age 0 pollock abundance. The location and diving behavior of tagged whales suggested that whales were favoring capelin over pollock as a prey source.     

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

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

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

Assessing changes in numbers and distribution of large whale entanglements in Newfoundland and Labrador,Canada1

Entanglements of large whales in commercial fisheries in Newfoundland and Labrador, Canada, have been consistently recorded since 1979, as part of a program aimed at releasing captured animals and reducing costs to fishermen. This data set represented an opportunity to identify fisheries posing particular entanglement risks to local whale populations. Data were assessed over the periods 1979–1992 and 1993–2008, corresponding to distinct phases in fisheries distribution and intensity. Between 1979 and 2008, 1,209 large whale entanglements were recorded in Newfoundland and Labrador. These were mostly humpback whales (Megaptera novaeangliae; 80%) and minke whales (Balaenoptera acutorostrata; 15%). Dramatic declines in reported inshore whale entanglement rates were observed following the 1992 moratorium on Atlantic cod (Gadus morhua) fisheries. Recently, more entanglements have been reported further offshore, largely due to expansion of fisheries targeting snow crab (Chionoecetes opilio). For all whale species, entanglement rates and associated mortality rates varied considerably in different fishing gear. Fractions of humpback and minke whales found dead in different fishing gear differed substantially, with minke whales far more likely to be found dead than humpback whales.     

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.     

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.     

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

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

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

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

Killer whales (Orcinus orca) feeding on lumpfish (Cyclopterus lumpus) in northern Norway

Killer whales (Orcinus orca) in Norwegian waters have long been known to rely on Atlantic herring (Clupea harengus) as a main prey resource. However, research almost exclusively conducted at seasonal herring grounds may have biased studies away from detecting other potentially significant prey species. Since 2013, dedicated research efforts have focused on monitoring killer whale occurrence and foraging ecology throughout the year in northern Norway. This study presents results on site-fidelity of photographically identified individuals, predation records and behavioral patterns from five spring seasons (March–April) in 2014–2018 in Andfjord, northern Norway. A minimum number of 75 adult and subadult killer whales (out of a catalog of 971 individuals) returned seasonally to the study area for foraging and residency for up to six weeks. Lumpfish (or lumpsucker, Cyclopterus lumpus) was the only type of prey identified (based on molecular or visual identification) on 22 predation events from 2016 (n = 4), 2017 (n = 2) and 2018 (n = 16). Spatial group cohesion observed when foraging was a potential adaptation for efficiently hunting this prey species. These whales were also encountered at herring wintering grounds the same years, but with different group sizes. Such behavioral adaptations suggested intraannual switching between prey resources and foraging strategies.     

Evidence of resource partitioning between humpback and minke whales around the western Antarctic Peninsula

For closely related sympatric species to coexist, they must differ to some degree in their ecological requirements or niches ( e.g. , diets) to avoid interspecific competition. Baleen whales in the Antarctic feed primarily on krill, and the large sympatric prewhaling community suggests resource partitioning among these species or a nonlimiting prey resource. In order to examine ecological differences between sympatric humpback and minke whales around the Western Antarctic Peninsula, we made measurements of the physical environment, observations of whale distribution, and concurrent acoustic measurements of krill aggregations. Mantel's tests and classification and regression tree models indicate both similarities and differences in the spatial associations between humpback and minke whales, environmental features, and prey. The data suggest (1) similarities (proximity to shore) and differences (prey abundance versus deep water temperatures) in horizontal spatial distribution patterns, (2) unambiguous vertical resource partitioning with minke whales associating with deeper krill aggregations across a range of spatial scales, and (3) that interference competition between these two species is unlikely. These results add to the paucity of ecological knowledge relating baleen whales and their prey in the Antarctic and should be considered in conservation and management efforts for Southern Ocean cetaceans and ecosystems.     

The timing of spawning in capelin (Mallotus villosus Müller) at a coastal location in eastern Newfoundland

We tested three hypotheses concerning the timing of spawning for a circumpolar species, capelin (Mallotus villosus), for which timing of larval emergence is known to be synchronized by physical conditions. The first hypothesis, developed from previous studies, was that spawning would be synchronized by upwelling events. Initial results from Middle Cove Beach in eastern Newfoundland indicated that spawning was not synchronized with upwelling. We next hypothesized that spawning was a function of several environmental variables. Results from logistic regression indicated that neither single-factor nor multi-factor models could explain the timing of spawning. Single variables could predict spawning in some years but no variable could reliably predict the time of spawning year after year. Finally, we hypothesized that the probability of spawning increased as a set of significant variables approached preferred levels. For capelin at Middle Cove, the set of variables that influence capelin spawning were identified as wave height, sea surface roughness and capelin abundance in the water. Thus only a combination of variables explained the timing of spawning for capelin. Preferred conditions for capelin spawning were wave heights less than 20 cm at the beach, a sea surface with a slight ripple, and an intermediate rank abundance of capelin in the water corresponding to hundreds to thousands of individuals. Capelin abundance alone was not a useful predictor. During the course of the study we observed a shift in the dates that capelin arrived and spawned at the beach. During 1987–1990 capelin spawned at Middle Cove Beach during June, but in more recent years (1991–1993) capelin did not arrive or spawn until July.     

Population spatial structuring on the feeding grounds in North Atlantic humpback whales (Megaptera novaeangliae)

Population spatial structuring among North Atlantic humpback whales Megaptera novaeangliae on the summer feeding grounds was investigated using movement patterns of identified individuals. We analysed the results from an intensive 2-year ocean-basin-scale investigation resulting in 1658 individuals identified by natural markings and 751 individuals by genetic markers supplemented with data from a long-term collaborative study with 3063 individuals identified by natural markings. Re-sighting distances ranged from <1 km to >2200 km. The frequencies ( F ) of re-sighting distances ( D ) observed in consecutive years were best modelled by an inverse allometric function ( F =6631 D ^−1.24, r ²=0.984), reflecting high levels of site fidelity (median re-sighting distance <40 km) with occasional long-distance movement (5% of re-sightings >550 km). The distribution of re-sighting distances differed east and west of 45°W, with more long-distance movement in the east. This difference is consistent with regional patterns of prey distribution and predictability. Four feeding aggregations were identified: the Gulf of Maine, eastern Canada, West Greenland and the eastern North Atlantic. There was an exchange rate of 0.98% between the western feeding aggregations. The prevalence of long-distance movement in the east made delineation of possible additional feeding aggregations less clear. Limited exchange between sites separated by as little as tens of kilometres produced lower-level structuring within all feeding aggregations. Regional and temporal differences in movement patterns reflected similar foraging responses to varying patterns of prey availability and predictability. A negative relationship was shown between relative abundance of herring and sand lance in the Gulf of Maine and humpback whale movement from the Gulf of Maine to eastern Canada.     

Signals from the south; humpback whales carry messages of Antarctic sea‐ice ecosystem variability

Southern hemisphere humpback whales (Megaptera novaeangliae) rely on summer prey abundance of Antarctic krill (Euphausia superba) to fuel one of the longest‐known mammalian migrations on the planet. It is hypothesized that this species, already adapted to endure metabolic extremes, will be one of the first Antarctic consumers to show measurable physiological change in response to fluctuating prey availability in a changing climate; and as such, a powerful sentinel candidate for the Antarctic sea‐ice ecosystem. Here, we targeted the sentinel parameters of humpback whale adiposity and diet, using novel, as well as established, chemical and biochemical markers, and assembled a time trend spanning 8 years. We show the synchronous, inter‐annual oscillation of two measures of humpback whale adiposity with Southern Ocean environmental variables and climate indices. Furthermore, bulk stable isotope signatures provide clear indication of dietary compensation strategies, or a lower trophic level isotopic change, following years indicated as leaner years for the whales. The observed synchronicity of humpback whale adiposity and dietary markers, with climate patterns in the Southern Ocean, lends strength to the role of humpback whales as powerful Antarctic sea‐ice ecosystem sentinels. The work carries significant potential to reform current ecosystem surveillance in the Antarctic region.     

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.     

Future recovery of baleen whales is imperiled by climate change

Historical harvesting pushed many whale species to the brink of extinction. Although most Southern Hemisphere populations are slowly recovering, the influence of future climate change on their recovery remains unknown. We investigate the impacts of two anthropogenic pressures—historical commercial whaling and future climate change—on populations of baleen whales (blue, fin, humpback, Antarctic minke, southern right) and their prey (krill and copepods) in the Southern Ocean. We use a climate–biological coupled “Model of Intermediate Complexity for Ecosystem Assessments” (MICE) that links krill and whale population dynamics with climate change drivers, including changes in ocean temperature, primary productivity and sea ice. Models predict negative future impacts of climate change on krill and all whale species, although the magnitude of impacts on whales differs among populations. Despite initial recovery from historical whaling, models predict concerning declines under climate change, even local extinctions by 2100, for Pacific populations of blue, fin and southern right whales, and Atlantic/Indian fin and humpback whales. Predicted declines were a consequence of reduced prey (copepods/krill) from warming and increasing interspecific competition between whale species. We model whale population recovery under an alternative scenario whereby whales adapt their migratory patterns to accommodate changing sea ice in the Antarctic and a shifting prey base. Plasticity in range size and migration was predicted to improve recovery for ice‐associated blue and minke whales. Our study highlights the need for ongoing protection to help depleted whale populations recover, as well as local management to ensure the krill prey base remains viable, but this may have limited success without immediate action to reduce emissions.     

Threshold foraging by gray whales in response to fine scale variations in mysid density

The gray whale (Eschrichtius robustus) is a coastal species whose nearshore summer foraging grounds off the coast of British Columbia offer an opportunity to study the fine scale foraging response of baleen whales. We explore the relationship between prey density and gray whale foraging starting with regional scale (10 km) assessments of whale density (per square kilometer) and foraging effort as a response to regional mysid density (per cubic meter), between 2006 and 2007. In addition we measure prey density at a local scale (100 m), while following foraging whales during focal surveys. We found regional mysid density had a significant positive relationship with both gray whale density and foraging effort. We identify a threshold response to regional mysid density for both whale density and foraging effort. In 2008 the lowest average local prey density measured beside a foraging whale was 2,300 mysids/m³. This level was maintained even when regional prey density was found to be substantially lower. Similar to other baleen whales, the foraging behavior of gray whales suggests a threshold response to prey density and a complex appreciation of prey availability across fine scales.     

Trophic niche partitioning and diet composition of sympatric fin (Balaenoptera physalus) and humpback whales (Megaptera novaeangliae) in the Gulf of Alaska revealed through stable isotope analysis

Fin and humpback whales are large consumers that are often sympatric, effectively sharing or partitioning their use of habitat and prey resources. Stable carbon and nitrogen isotopes in the skin of fin and humpback whales from two regions in the western Gulf of Alaska, Kodiak, and Shumagin Islands, were analyzed to test the hypothesis that these sympatric baleen whales exhibit trophic niche partitioning within these regions. Standard ellipse areas, estimated using Bayesian inference, suggested that niche partitioning between species is occurring in the Kodiak region but not in the Shumagin Islands. Isotopic mixing models based on stable isotopes from whales and local prey samples, were used to estimate possible diet solutions for whales in the Kodiak region. Comparison of isotopic niches and diet models support niche partitioning, with fin whales foraging primarily on zooplankton and humpback whales foraging on zooplankton and small forage fish. The results of this study show that niche partitioning between sympatric species can vary by region and may be the result of prey availability, prey preferences, or both.     

A citizen science approach to long-term monitoring of humpback whales (Megaptera novaeangliae) off Sydney,Australia

The Cape Solander Whale Migration Study is a citizen science project that annually counts northward migrating humpback whales (Megaptera novaeangliae) off Cape Solander, Sydney, Australia. Dedicated observers have compiled a 20-year data set (1997–2017) of shore-based observations from Cape Solander's high vantage point. Using this long-term data set collected by citizen scientists, we sought to estimate the humpback whale population trend as it continues to recover postexploitation. We estimated an exponential growth rate of 0.099 (95% CI = 0.079–0.119) using a generalized linear model, based on observer effort (number of observation days) and number of whales observed, equating to 10% per annum growth in sightings since 1997. We found that favorable weather conditions for spotting whales off Cape Solander consisted of winds <30 km/hr from a southerly through a north westerly direction. Incidental observations of other cetacean species included the endangered blue whale (Balaenoptera musculus) and data deficient species such as killer whales (Orcinus orca) and false killer whales (Pseudorca crassidens). Citizen science-based studies can provide a cost-effective approach to monitoring wildlife over the time necessary to detect change in a population. Information obtained from citizen science projects like this may help inform policy makers responsible for State and Federal protection of cetaceans in Australian waters and beyond.     

Underwater acrobatics by the world's largest predator: 360° rolling manoeuvres by lunge-feeding blue whales

The extreme body size of blue whales requires a high energy intake and therefore demands efficient foraging strategies. As an obligate lunge feeder on aggregations of small zooplankton, blue whales engulf a large volume of prey-laden water in a single, rapid gulp. The efficiency of this feeding mechanism is strongly dependent on the amount of prey that can be captured during each lunge, yet food resources tend to be patchily distributed in both space and time. Here, we measured the three-dimensional kinematics and foraging behaviour of blue whales feeding on krill, using suction-cup attached multi-sensor tags. Our analyses revealed 360° rolling lunge-feeding manoeuvres that reorient the body and position the lower jaws so that a krill patch can be engulfed with the whale''s body inverted. We also recorded these rolling behaviours when whales were in a searching mode in between lunges, suggesting that this behaviour also enables the whale to visually process the prey field and maximize foraging efficiency by surveying for the densest prey aggregations. These results reveal the complex manoeuvrability that is required for large rorqual whales to exploit prey patches and highlight the need to fully understand the three-dimensional interactions between predator and prey in the natural environment.