Shallow and deep lunge feeding of humpback whales in fjords of the West Antarctic Peninsula

Humpback whales (Megaptera novaeangliae) belong to the class of marine mammals known as rorquals that feed through extraordinarily energetic lunges during which they engulf large volumes of water equal to as much as 70% of their body mass. To understand the kinematics of humpback lunge feeding, we attached high‐resolution digital recording tags incorporating accelerometers, magnetometers, pressure and sound recording to whales feeding on euphausiids in fjords of the West Antarctic Peninsula. Instances of near vertical lunges gave us the unique opportunity to use the signal from the accelerometer to obtain a fine scale record of the body accelerations involved in lunging. We found that lunges contain extreme accelerations reaching 2.5 m/s² in certain instances, which are then followed by decelerations. When animals are intensively feeding the inter‐lunge interval is similar for both deep and shallow lunges suggesting a biomechanical constraint on lunges. However, the number of lunges per dive varies from one for shallow feeding (<25 m) to a median of six for deeper dives. Different feeding patterns were evident in the kinematic record, for deep and shallow feeding bouts with the much greater mean turn rates occurring in shallow feeding.     

Behavioral ontogeny in humpback whale (Megaptera novaeangliae) calves during their residence in Hawaiian waters

The ontogeny of behavior in young offspring is a key component of a species life history, influencing short-term survival and life-long future fitness. In this study, we examine the behavioral ontogeny of humpback whale calves during their natal season on the winter breeding grounds. Behavioral data were collected during focal follows for 69 calves. Their relative age was estimated based on the degree of unfurling of the dorsal fin, and analysis of their time budgets revealed that behavior changed as the calves matured. Among the youngest group of calves ( n = 27), persistent travel accounted for median 85% (interquartile range 34%) of the calves' time budget, they spent little time alone at the surface and the breathing regime included frequent intermittent breaths. Within the oldest group of calves ( n = 26), time spent traveling dropped to a median 47% (IQR 42%) of the time budget, they spent more time at rest, surfaced alone more frequently, and the breathing regime more closely resembled that of adult humpbacks. We suggest potential functions for these and other components of calf behavior during this period, and review our findings in light of recent discussions on the role of predation pressure as a driving force in mysticete migration.     

Spatial patterns of humpback whale (Megaptera novaeangliae) sightings and survey effort: Insight into North Atlantic population structure

Understanding the population structure of a species is critical to its effective management and conservation. The humpback whale ( Megaptera novaeangliae ) has been the target of numerous research projects in several ocean basins, but no clear picture of its population structure has emerged. In the North Atlantic Ocean, genetic analyses and photo-identification movements have shown significant heterogeneity among the summer feeding grounds. Building on this knowledge, we test the hypothesis that the feeding grounds represent distinct populations by analyzing the spatial pattern of summer humpback whale sightings and survey effort. Controlling for the spatial pattern of effort, sightings are clustered, with peaks at radial distances of 300 km, 600 km, and 1,500 km. These results provide insight into the spatial extent of the summer population structure of humpback whales in the North Atlantic Ocean. Fine-scale clustering at distances of 300 km and 600 km is compatible with multiple populations consisting of the Gulf of Maine, eastern Canada, western Greenland, and Iceland. Broad-scale clustering at distances of 1,500 km may represent divisions between the western and eastern North Atlantic populations. These results provide spatial bounds to the feeding grounds of humpback whales and emphasize their distinct nature as management units.     

Incorporating tides into the acoustic ecology of humpback whales

North Pacific humpback whales (Megaptera novaeangliae) migrate annually to foraging grounds in Southeast Alaska that are characterized by semidiurnal tidal cycles. Tidal activity is an important driver of marine mammal behavior on foraging grounds, but is often omitted in studies of acoustic ecology. To better understand the role of sound in this vocal species we investigated the influence of tidal height and direction on humpback whale nonsong calling behavior in Frederick Sound and described new call types for this population. The likelihood of detecting a call from the low‐frequency‐harmonic, pulsed, or noisy‐complex call classes was independent of tidal activity. The likelihood of detecting a call from the tonal call class, and a feeding call in particular, was 2.1 times higher during flood tides than during ebb tides (95% CI 1.1–4.4). This likely reflects an indirect relationship between humpback whale foraging and tides.     

First record of humpback whale songs in Southern Chile: Analysis of seasonal and diel variation

Male humpback whales (Megaptera novaeangliae) produce complex, patterned songs that are traditionally recorded on their breeding grounds. In this work, we report results from the first continuous acoustic monitoring of a humpback whale feeding ground off southern Chile, Corcovado Gulf. Using an autonomous continuously recording system anchored to the seafloor and an automatic signal detector, we used the units within a song to analyze the temporal distribution and diel patterns of humpback whales. Acoustic recordings were made at the end of the austral summer and autumn of 2012. Songs occurred over the entire 130 d monitoring period, from 1 February to 15 June 2012. The percentage of units detected increased throughout the monitored period with the highest detections in the last recorded month (June), despite recording for fewer days that month. Furthermore, songs were detected during all light regimes studied, but most frequently during darkness. This study provides further evidence that, far from being rare or sporadic, humpback whale songs occur commonly at a feeding ground in high latitudes over different light conditions and in all months, with a peak in autumn.     

Habitat preference reflects social organization of humpback whales (Megaptera novaeangliae) on a wintering ground

Habitat preference is driven by a complex interaction among behavioural patterns, biological requirements, and environmental conditions. These variables are difficult to determine for any species but are further complicated for migratory marine mammals, such as humpback whales Megaptera novaeangliae . Patterns of habitat use in relation to social organization potentially exist for this species on their wintering grounds. Using an integrated GIS approach, we examined the degree to which spatial patterns of habitat stratification are correlated within different humpback whale group types from 6 years of sighting data (1996–2001) collected on the Antongil Bay, Madagascar, wintering ground. Stratification of humpback whale sightings by behavioural classification showed significant variation in depth and distance from shore. Distribution by depth could not be described as a function of group size but could be described as a function of social organization, with mother–calf pairs showing a strong preference for shallower water compared to all other group types. Group size and social organization seem to be factors in distribution by distance from shore. Significant diurnal patterns in distribution by depth and distance from shore also exist, where mother–calf groups maintain a relatively stable distribution and pairs and competitive groups are the most variable. Patterns of habitat preference on this wintering ground appear to be guided by social organization, where distribution by depth and distance from shore highlight areas critical to conservation.     

Distribution and migratory destinations of humpback whales off the Pacific coast of Central America during the boreal winters of 1996–2003

Here, we examine the distribution, habitat use, and migratory destinations of North Pacific humpback whales wintering off Central America. Coastal boat surveys were conducted off Costa Rica and Panama between 1996 and 2003. In 1999, a broader survey was conducted along most of Central America. Over 23,000 km were surveyed, with the greatest effort off southern Costa Rica. We made 191 sightings of 320 individual humpback whales. Whales were seen between 14°N and 8°N, making this the most southerly of the North Pacific wintering areas. Encounters included singles, adult pairs, singers, and mother/calf pairs. Mother/calf pairs accounted for 27% of all groups sighted, which is one of the highest sighting rates reported among North Pacific wintering areas. Sixty percent of sightings occurred in depths <50 m. Average sea surface temperature was 28.6°C (±1.0 SD). Ninety percent of the 77 unique whales photo‐identified were also seen in the California–Oregon–Washington feeding aggregation. The 1999 survey showed that humpback whales were widely distributed along the Central American coast at relatively low densities. The extensive distribution of animals, the higher proportion of calves, and the almost exclusive migration to a single feeding area contrast with observations in other regions.     

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.     

Distributional patterns of humpback whales (Megaptera novaeangliae) along the Newfoundland East Coast reflect their main prey,capelin (Mallotus villosus)

On the Newfoundland foraging ground, humpback whales (Megaptera novaeangliae) primarily consume capelin (Mallotus villosus), which experienced a population collapse in the early 1990s, associated with altered timing of spawning and spawning migration. We examined whether humpback whale movement and distribution match these prey changes. Combining tour company whale sighting reports and photographs, citizen science reports of capelin spawning and scientific monitoring, whales were found to move northward along the east coast and whale aggregation presence within bays was associated with spawning capelin presence, being later in northerly bays. Whale aggregations arrived 8–20 days later than spawning capelin in northern bays, however, suggesting inconsistent tendencies to track high abundance spawning capelin aggregations during migration. Repeated scientific surveys during July–August 2009, 2010, 2012, 2014–2017, within a biological hotspot associated with capelin spawning sites in Notre Dame Bay, revealed that whale presence was influenced by the date of capelin spawning rather than capelin abundance metrics (i.e., biomass, number of shoals, shoal density, shoal area). A photo-identification catalog compiled during July–August, 2003–2017, revealed a 22% return rate of whales to the hotspot. Overall, findings suggest that capelin spawning sites are important foraging areas for humpback whales in coastal Newfoundland under these altered prey conditions.     

Humpback whale song variations in La Reunion Island from 2016 to 2018 and in Madagascar in 2018

Male Megaptera novaeangliae produce complex and structured songs which are shared at the population level. Song patterns are culturally transmitted and evolve progressively through time, both over the breeding season and among years. The songs also undergo periods of relatively rapid change, termed “revolutions.” Acoustic monitoring was conducted from 2016 to 2018 in Reunion and throughout 2018 in Madagascar to assess spatiotemporal variation in song structures and population connectivity. A total of 46 high-quality song samples were selected, representing 2,760 min of recordings in Reunion. In Madagascar, 12 samples representing 240 min of recordings were analyzed. Analysis of songs revealed 11 phrases and their variants. Low levels of temporal variations were observed over the breeding season. Songs recorded in June were very similar to those recorded in September. Greater variation was observed between years, and some phrases identified in 2018 may have evolved from phrases recorded in 2017. More variants were described for each phrase type in 2018 compared to 2016. All themes recorded in Reunion were shared with Madagascar, suggesting a high degree of population connectivity during the breeding season.     

Modeling whale entanglement injuries: An experimental study of tissue compliance, line tension, and draw-length

Two test systems were developed to evaluate the influence of draw‐length and tissue compliance on entanglement‐induced epidermal abrasion in humpback (Megaptera novaeangliae) and right whale (Eubalaena glacialis) tissue samples. Under straight pull abrasion tests, an adult right whale fluke required 3.7 times the load and 15 times the draw‐length of a right whale calf flipper to induce epidermal failure whereas a humpback fluke was intermediate between these extremes. A load applied tangentially to the leading edge of the fluke or flipper resulted in a substantial, but reversible, deflection of the leading edge in the direction of the applied load. The maximum possible deformation of the leading edge under shear load, prior to the line slipping relative to the skin, was defined as the tissue compliance limit. Oscillatory abrasion tests revealed that line draw‐lengths exceeding the tissue compliance limit resulted in substantially increased tissue abrasion. In actual entanglements, line draw‐length relative to the tissue compliance may determine if the line will cut into the body or simply press against the skin. Increasing the entangling line's ability to stretch in response to a load could potentially minimize sliding of the line relative to the skin and help mitigate entanglement injuries.     

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.     

Migratory movements and surfacing rates of humpback whales (Megaptera novaeangliae) satellite tagged at Socorro Island,Mexico

Humpback whales wintering in the Revillagigedo Archipelago, Mexico, have been considered a different subpopulation from those found off mainland Mexico and Baja California. The primary feeding grounds for Revillagigedo humpbacks remain unknown. In February 2003, we deployed 11 Argos satellite‐monitored radio tags to track movements and surfacings of humpback whales (five adults without calves, five mothers with calves, one calf) off Socorro Island in the Revillagigedo Archipelago. Tracking ranged from 222 to 10,481 km over 4.9–149.1 d. Eight whales left Socorro Island: five visited other Mexican wintering destinations, seven moved north of these areas. Migration routes were primarily offshore (average 444 km). Two whales were tracked to feeding grounds: one to British Columbia (46 d migration), and one to Alaska (49 d migration). Mean travel speeds were 1.2 km/h in wintering areas, 4.0 km/h during migration, and 2.2 km/h in feeding areas. Overall surfacing rates ranged from 21 to 88 surfacings/h. Surfacing rates differed between the calf and all other whales, and between feeding areas and migratory/wintering areas for the calf and an adult without a calf. The calf also showed diel variation in surfacing rates. The offshore habits of tagged whales may explain scarce resightings of Revillagigedo humpbacks outside the Revillagigedo Archipelago.     

A quarter of a world away: female humpback whale moves 10,000 km between breeding areas

Fidelity of individual animals to breeding sites is a primary determinant of population structure. The degree and scale of philopatry in a population reflect the fitness effects of social facilitation, ecological adaptation and optimal inbreeding. Patterns of breeding-site movement and fidelity are functions of social structure and are frequently sex biased. We report on a female humpback whale (Megaptera novaeangliae) first identified by natural markings off Brazil that subsequently was photographed off Madagascar. The minimum travel distance between these locations is greater than 9800 km, approximately 4000 km longer than any previously reported movement between breeding grounds, more than twice the species' typical seasonal migratory distance and the longest documented movement by a mammal. It is unexpected to find this exceptional long-distance movement between breeding groups by a female, as models of philopatry suggest that male mammals move more frequently or over longer distances in search of mating opportunities. While such movement may be advantageous, especially in changeable or unpredictable circumstances, it is not possible to unambiguously ascribe causality to this rare observation. This finding illustrates the behavioural flexibility in movement patterns that may be demonstrated within a typically philopatric species.     

Seasonal changes in pod characteristics of eastern Australian humpback whales (Megaptera novaeangliae), Hervey Bay 1992–2005

We investigated the characteristics and composition of 4,506 humpback whale pods observed in Hervey Bay between 1992 and 2005. We use these data to analyze and model the variability of pod size and composition, and to assess the importance of Hervey Bay for particular classes of humpback whales. Pods ranged in size from one to nine individuals. Pairs were the most frequent pod type (1,344, 29.8%), followed by mother‐calf alone (1,249, 27.7%), trios (759, 16.8%), singletons (717, 15.9%), and 4+ whales (437, 9.7%). Of the 4,506 pods, calves were present in 40%, and 10.8% of all pods had one or more escorts present. Of the 1,804 pods observed with calves present, 1,251 (69.4%) were mothers alone with their calves. The size and composition of pods in the study area varied significantly as the season progressed. Pods with calves present were rarely recorded early in the season but dominated later in the season. A significant increase over years in larger groups may be related to social and behavioral changes as the population expands. The data indicate that Hervey Bay is important to immature males and females early in the season, to mature males and females in mid‐season, and to mother‐calf pairs (either alone or with escorts) in mid‐to‐late season.     

Non-lethal entanglement of humpback whales (Megaptera novaeangliae) in fishing gear in northern Southeast Alaska

Aim Entanglement in fishing gear is recognized as a potentially significant source of serious injury and mortality for humpback whales (Megaptera novaeangliae) in some parts of their range. In recent years, the number of humpback whales reported to have been entangled in Alaska has increased. In 2003–04 we quantified the prevalence of non‐lethal entanglements of humpback whales in northern Southeast Alaska (SEAK) with the ultimate goal of informing management discussions of the entanglement issue for the Central North Pacific stock of humpback whales. Location The near‐shore waters of northern Southeast Alaska. Methods We photographed individual humpback whales’ caudal peduncles as they dived and then examined the photographs for scars indicative of a previous entanglement. Results The percentage of whales assessed to have been non‐lethally entangled at some time in their lives ranged from 52% (minimal estimate) to 71% (conditional estimate) to 78% (maximal estimate). Of these, the conditional estimate is recommended because it is based solely on unambiguous scars. Eight per cent of the whales in one portion of the study area (Glacier Bay/Icy Strait) acquired new entanglement scars between 2003 and 2004, although the sample size was small. Calves were less likely than older whales to have entanglement scars, and males may be at higher risk than females. Whales with more photographs and/or photographic coverage may be more likely to be assessed as having been entangled than whales with fewer photographs and/or coverage. Main conclusions Caudal peduncle scars reveal that the majority of humpback whales in northern SEAK have been entangled. Comparison with statistics on reported entanglements suggests that most whales apparently shed the gear on their own, unless humans are disentangling whales much more often than is reported. While cumulative estimates of the percentage of whales with entanglement scars (e.g. the conditional estimate) provide useful baseline information, future efforts should focus on monitoring the annual rate of entanglement scar acquisition as a more powerful measure of contemporary entanglement rates. Our findings indicate that entanglement of humpback whales in fishing gear in SEAK is a management issue warranting increased attention. A proactive approach is needed to address the problem and to identify and implement preventive measures.     

Site fidelity and movements of humpback whales (Megaptera novaeangliae) on the Brazilian breeding ground,southwestern Atlantic

Site fidelity and movements were studied for humpback whales photo-identified from 1989 to 2006 in the Abrolhos Bank, southwestern Atlantic, Brazil. A total of 2,612 individuals were identified, 374 of which were observed on more than one occasion. The cumulative number of identified whales has increased since 1989. Recapture rate was low and varied among different years. A total of 33 whales was observed using the Abrolhos Bank for longer than 10 yr, up to a maximum of 16 yr. Our data suggest that different whales show distinct movement rates. Some whales used a large extent of the Abrolhos Bank region. Opportunistic photo-identification data (on the scale of the Brazilian coast from 4° to 23°S) revealed important information about stock identity. The longest distance between within-season resightings was over 600 km, while one whale was observed in two locations separated by more than 1,400 km in different years. Long-range movements within and between seasons support the single stock hypothesis for humpback whales wintering off the Brazilian coast.     

Immature euphausiids do not appear to be prey for humpback whales (Megaptera novaeangliae) during spring and summer in Southeast Alaska

Humpbacks whales (Megaptera novaeangliae) have shown a remarkable recovery in the North Pacific, raising concerns regarding their impact on marine communities. In Southeast Alaska, humpbacks feed heavily on euphausiids; however, it remains unclear whether they target immature individuals despite evidence that they do so elsewhere. I evaluate the hypothesis that humpbacks target immature euphausiids in late spring‐summer in Southeast Alaska. Plankton samples were collected at random sites (n = 44) and near whales (n = 53) between 8 June and 9 September 2008 in Frederick Sound and Stephens Passage. The proportion of samples containing immature euphausiids, and immature euphausiid abundance within those samples, were compared between the two sample types. Similar analyses were conducted for adult euphausiids (prey) and calanoid copepods (nonprey) for comparison. I found no statistical difference between the whale and random samples with respect to the occurrence or numerical density of immature euphausiids, which is consistent with the hypothesis that whales did not target them in 2008. Smaller size, insufficient numerical densities and lower energy density of immature euphausiids are suggested as possible reasons. These findings can assist in resolving regional humpback abundance and distribution patterns, and can contribute to an understanding of the trophic interactions characterizing the local ecosystem.