Non-song acoustic communication in migrating humpback whales (Megaptera novaeangliae)

Humpback whales are renowned for the complex structured songs produced by males. A second, relatively understudied area of humpback acoustic communication concerns un-patterned sounds known as "social sounds," produced by both males and females. These include vocalizations as well as sounds produced at the surface of the water as a result of surface behaviors ( e.g. , breaching, pectoral slapping). This study describes a portion of the non-song social sound repertoire of southward migrating humpbacks in Australian waters, and explores the social relevance of these sounds. On migration, humpback whales travel in social groups of varying compositions. These social groups are not stable in that humpback whales continually change group composition by splitting from, or joining with, other groups. The results of this study suggest that "breaching" and "slapping" have a communicative function. Other sounds such as "underwater blows" and "cries" were heard mainly in competitive groups while other low-frequency sounds such as "grumbles,""snorts,""thwops," and "wops" may function in intra- or inter-group communication. Particular sounds ("grunts,""groans," and "barks") were almost exclusive to joining pods suggesting a role in social integration. Social sounds in humpbacks may have specific social and behavioral functions relating to social group composition, and the mediation of interactions between these social groups.     

The effect of close approaches for tagging activities by small research vessels on the behavior of humpback whales (Megaptera novaeangliae)

Small research vessels are often used as platforms for tagging activities to collect behavioral data on cetaceans and they have the potential to disturb that group or individual. If this disturbance is ignored, results and conclusions produced by that study could be inaccurate. Here land‐based behavioral data of migrating humpback whales (Megaptera novaeangliae) (n = 29) were used to determine the effect of close approaches for tagging by research vessels on their diving, movement and surface behaviors. Groups of whales were tagged, using digital recording tags, by small research vessels, as part of a behavioral response study. In groups that were approached for tagging, temporary changes in movement behaviors during close approaches were found, with subsequent recovery to “pre‐approach” levels. In female‐calf groups more long‐term changes in travel speed were found. Results suggest that, although close approaches for tagging by small vessels may cause behavioral changes in humpback whales, this change may be small and temporary. However, in female‐calf groups, the behavioral change may be greater and longer lasting. This study shows that when using small vessels for behavioral research, disturbance, and recovery should be measured to ensure integrity of data used for other analyses.     

Factors driving the variability in diving and movement behavior of migrating humpback whales (Megaptera novaeangliae): Implications for anthropogenic disturbance studies

Humpback whales (Megaptera novaeangliae) undertake one of the longest migrations of any animal and while on a broad‐scale this journey appears direct, on a fine‐scale, behaviors associated with socializing and breeding are regularly observed. However, little is known about which social and environmental factors influence behavior during this time. Here we examined the effect of multiple factors on the movement (speed and course) and diving behavior (dive and surfacing duration) of humpback whales during migration off the eastern coast of Australia. Focal data (202 h) were collected on 94 different whale groups with simultaneous social and environmental context data. The environmental factors water depth and wind speed were found to be important predictors of dive and movement behavior, whereas social factors were less influential at this site. Groups tended to dive for longer with increased water depth but traveled more slowly in increasing wind speeds. These baseline studies are crucial when examining the effect of anthropogenic disturbance. Determining which natural factors significantly affect behavior ensures any observed behavioral changes are correctly attributed to the disturbance and are not a result of other factors. In addition, any responses observed can be put into biological context and their relative magnitude determined.     

Whale killers: Prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia

Reports of killer whales (Orcinus orca) preying on large whales have been relatively rare, and the ecological significance of these attacks is controversial. Here we report on numerous observations of killer whales preying on neonate humpback whales (Megaptera novaeangliae) off Western Australia (WA) based on reports we compiled and our own observations. Attacking killer whales included at least 19 individuals from three stable social groupings in a highly connected local population; 22 separate attacks with known outcomes resulted in at least 14 (64%) kills of humpback calves. We satellite‐tagged an adult female killer whale and followed her group on the water for 20.3 h over six separate days. During that time, they attacked eight humpback calves, and from the seven known outcomes, at least three calves (43%) were killed. Overall, our observations suggest that humpback calves are a predictable, plentiful, and readily taken prey source for killer whales and scavenging sharks off WA for at least 5 mo/yr. Humpback “escorts” vigorously assisted mothers in protecting their calves from attacking killer whales (and a white shark, Carcharodon carcharias). This expands the purported role of escorts in humpback whale social interactions, although it is not clear how this behavior is adaptive for the escorts.     

Identifying behavioral states and habitat use of acoustically tracked humpback whales in Hawaii

Although humpback whales have been well‐studied on their Hawaiian breeding grounds, it is difficult to track individual animals over long distances without tags, particularly when they move offshore. Here, singing humpback whales were localized in three dimensions on the Pacific Missile Range Facility off Kauai, Hawaii, located between 20 km and 80 km offshore, from January 2011 through June 2014. Detailed behavioral analyses were conducted on the resulting tracks. One hundred and eight individual tracks were identified and metrics of these tracks were examined. Using these metrics, the tracks were classified into four behavior categories, described herein as Directed Travel, Repeated Stationary Dives, Mill, and tracks with Combinations of behavioral states. Some diel and seasonal patterns were identified, with Mill tracks occurring more at night than the other behaviors, Repeated Stationary Dive tracks occurring more during the day, and Directed Travel occurring only at the start and end of the breeding season. These results provide detailed insights into the movement of singing humpback whales, particularly in offshore waters where they may be migrating into or out of breeding grounds. This also contributes valuable information on the baseline behavior of humpback whales on a US Navy training range.     

Social structure in migrating humpback whales (Megaptera novaeangliae)

Although largely solitary, humpback whales exhibit a number of behaviours where individuals co-operate with one another, for example during bubble net feeding. Such cases could be due to reciprocal altruism brought on by exceptional circumstances, for example the presence of abundant shoaling fish. An alternative explanation is that these behaviours have evolved through kin selection. With little restriction to either communication or movement, diffuse groups of relatives could maintain some form of social organization without the need to travel in tight-nit units. To try to distinguish between these hypotheses, we took advantage of the fact that migrating humpback whales often swim together in small groups. If kin selection is important in humpback whale biology, these groups should be enriched for relatives. Consequently, we analysed biopsy samples from 57 groups of humpback whales migrating off Eastern Australia in 1992. A total of 142 whales were screened for eight microsatellite markers. Mitochondrial DNA sequences (371 bp) were also used to verify and assist kinship identification. Our data add support to the notion that mothers travel with their offspring for the first year of the calf's life. However, beyond the presence of mother-calf/yearling pairs, no obvious relatedness pattern was found among whales sampled either in the same pod or on the same day. Levels of relatedness did not vary between migratory phases (towards or away from the breeding ground), nor between the two sexes considered either overall or in the north or south migrations separately. These findings suggest that, if any social organization does exist, it is formed transiently when needed rather than being a constant feature of the population, and hence is more likely based on reciprocal altruism than kin selection.     

Delphinid whistle production and call matching during playback of simulated military sonar

In 2007 and 2008, controlled exposure experiments were performed in the Bahamas to study behavioral responses to simulated mid‐frequency active sonar (MFA) by three groups of odontocetes: false killer whales, Pseudorca crassidens; short‐finned pilot whales, Globicephala macrorhynchus; and melon‐headed whales, Peponocephala electra. An individual in each group was tagged with a Dtag to record acoustic and movement data. During exposures, some individuals produced whistles that seemed similar to the experimental MFA stimulus. Statistical tests were thus applied to investigate whistle‐MFA similarity and the relationship between whistle production rate and MFA reception time. For the false killer whale group, overall whistle rate and production rate of the most MFA‐like whistles decreased with time since last MFA reception. Despite quite low whistle rates overall by the melon‐headed whales, statistical results indicated minor transient silencing after each signal reception. There were no apparent relationships between pilot whale whistle rates and MFA sounds within the exposure period. This variability of responses suggests that changes in whistle production in response to acoustic stimuli depend not only on species and sound source, but also on the social, behavioral, or environmental contexts of exposure.     

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.     

OCCURRENCE, COMPOSITION, AND POTENTIAL FUNCTIONS OF NORTH ATLANTIC RIGHT WHALE (EUBALAENA GLACIALIS) SURFACE ACTIVE GROUPS

Marine mammal social interactions are poorly understood primarily because of the difficulty of observing these behaviors in the wild. Observations of copulations in North Atlantic right whale surface active groups (SAGs) have led to the hypothesis that the function of these groups is for conception. The occurrence and composition of SAGs sighted from 1992 to 2001 were investigated on the basis of age, sex, and reproductive status of individuals. A total of 918 SAG events were documented. SAGs were observed in all habitat areas and in all months in which right whales were sighted. Group compositions revealed both potentially conceptive groups and nonconceptive groups. Of right whales whose age was known, 93% were first sighted in SAGs as juveniles, and more than half of all observed groups contained at least one juvenile whale. The group composition and timing of occurrence of SAGs do not support the hypothesis that all groups serve a purely conceptive function. Their functional role is likely much broader. Other potential functions include play, mating practice, or maintenance of social bonds.     

Rorqual whale (Balaenopteridae) surface lunge‐feeding behaviors: Standardized classification,repertoire diversity,and evolutionary analyses

Rorqual whales (Family: Balaenopteridae) are the world's largest predators and sometimes feed near or at the sea surface on small schooling prey. Most rorquals capture prey using a behavioral process known as lunge‐feeding that, when occurring at the surface, often exposes the mouth and head above the water. New technology has recently improved historical misconceptions about the natural variation in rorqual lunge‐feeding behavior yet missing from the literature is a dedicated study of the identification, use, and evolution of these behaviors when used to capture prey at the surface. Here we present results from a long‐term investigation of three rorqual whale species (minke whale, Balaenoptera acutorostrata; fin whale, B. physalus; and blue whale, B. musculus) that helped us develop a standardized classification system of surface lunge‐feeding (SLF) behaviors. We then tested for differences in frequency of these behaviors among the three species and across all rorqual species. Our results: (1) propose a unified classification system of six homologous SLF behaviors used by all living rorqual whale species; (2) demonstrate statistically significant differences in the frequency of each behavior by minke, fin, and blue whales; and (3) provide new information regarding the evolution of lunge‐feeding behaviors among rorqual whales.     

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.     

Southeastern Pacific humpback whales (Megaptera novaeangliae) and their breeding grounds: Distribution and habitat preference of singers and social groups off the coast of Ecuador

Understanding the distribution, habitat preference, and social structure of highly migratory species at important life history stages (e.g., breeding and calving) is essential for conservation efforts. We investigated the spatial distribution and habitat preference of humpback whale social groups and singers, in relation to depth categories (<20 m, 20–50 m, and >50 m) and substrate type (muddy and mixed) on a coastal southeastern Pacific breeding ground. One hundred and forty‐three acoustic stations and 304 visual sightings were made at the breeding ground off the coast of Esmeraldas, Ecuador. Spatial autocorrelation analysis suggested singers were not randomly distributed, and Neu's method and Monte Carlo simulations indicated that singers frequented depths of <20 m and mixed substrate. Singletons, and groups with a calf displayed a preference for shallower waters (0–20 m), while pairs and groups with a calf primarily inhabited mixed bottom substrates. In contrast, competitive groups showed no clear habitat preference and exhibited social segregation from other whales. Understanding the habitat preference and distribution of humpback whales on breeding and calving grounds vulnerable to anthropogenic disturbance provides important baseline information that should be incorporated into conservation efforts at a regional scale.     

Singing through winter nights: Seasonal and diel occurrence of humpback whale (Megaptera novaeangliae) calls in and around the Gully MPA,offshore eastern Canada

Humpback whale use of areas off eastern Canada is poorly understood, a knowledge gap that could impact future conservation efforts. We describe the acoustic occurrence of humpback whales in and around the Gully Marine Protected Area (MPA), an eastern Scotian Shelf submarine canyon. Near‐continuous acoustic recordings sampling at 16 kHz were collected from the MPA and nearby slope areas from October 2012 to September 2014 using near‐bottom recorders. In an offshore region where humpbacks were thought to be rare, we observed calls from October to June with a peak in song and nonsong calls in December and January. This suggests that some individuals occur in Canadian waters in winter and the Gully region may be a North Atlantic humpback whale migratory corridor. Calls were predominantly songs indicating potential mating activities. Song and nonsong calls occurred more at sunset and during hours of darkness than during daylight. This study improves our understanding of the seasonal occurrence of humpback whales on the Scotian Slope and, more specifically, their use of an offshore protected area.     

Potential vessel collisions with Southern Hemisphere humpback whales wintering off Pacific Panama

Vessel collision is a threat to many whale species, and the risk has increased with expanding maritime traffic. This compromises international conservation efforts and requires urgent attention from the world's maritime industry. Humpback whales (Megaptera novaeangliae) are at the top of the death toll, and although Central America is a wintering area for populations from both the Northern and Southern Hemispheres, existing efforts to reduce ship‐whale collisions are meager. Herein, we evaluated the potential collisions between vessels and humpback whales wintering off Pacific Panama by following the movements of 15 whales tagged with satellite transmitters and comparing these data with tracks plotted using AIS real‐time latitude‐longitude points from nearly 1,000 commercial vessels. Movements of whales (adults and calves) in the Gulf of Panama coincide with major commercial maritime routes. AIS vessel data analyzed for individual whale satellite tracks showed that 53% (8 whales) of whales had 98 encounters within 200 m with 81 different vessels in just 11 d. We suggest implementing a 65 nmi Traffic Separation Scheme and a 10 kn maximum speed for vessel routing into the Gulf of Panama during the wintering season. In so doing, the area for potential whale‐vessel collisions could be reduced by 93%.     

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.     

THE GUNSHOT SOUND PRODUCED BY MALE NORTH ATLANTIC RIGHT WHALES (EUBALAENA GLACIALIS) AND ITS POTENTIAL FUNCTION IN REPRODUCTIVE ADVERTISEMENT

North Atlantic right whales ( Eubalaena glacialis ) make a short, distinctive broadband sound that is produced internally called a Gunshot sound. This sound has been recorded in the Bay of Fundy, Canada from both single whales ( n = 9) and social surface active groups ( n =49). Those single whales producing Gunshot sounds whose sex could be determined ( n = 9) were all mature males. Gunshot sounds were produced as part of a stereotyped behavioral sequence by these individuals, including frequent head-lifts and flipper slapping at the surface. In surface active groups, Gunshot sounds were commonly recorded when males were present in the group. The rate of production of Gunshot sounds was weakly correlated with the total number of males present in the group. Given the behavioral context of Gunshot sound production, and production of the sound only by male whales, Gunshots may function in a reproductive context as an advertisement signal to attract females, an agonistic signal directed toward other males, or a combination of the two functions.     

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.     

From the southern right whale hunting decline to the humpback whaling expansion: a review of whale catch records in the tropical western South Atlantic Ocean

相似文献   

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.     

Observations of cetaceans in the south-east Atlantic sector of the Southern Ocean,during summer 2008

Knowledge of cetacean species composition and their distribution in the south-east Atlantic sector of the Southern Ocean is scarce. During a survey in February–March 2008, systematic whale sightings were carried out along transect lines following the 5° and 15° E meridians between 35° and 67° S. In total, 67 toothed whales and 126 baleen whales were observed. Both fin whales (four animals) and Antarctic minke whales Balaenoptera bonaerenses (three animals) in addition to 16 individuals of unidentified species were among the observed baleen whales. The dominating baleen whale species in our study was humpback whales Megaptera novaeangliae with 108 individuals observed. They occurred single or in groups up to seven individuals (N _mean = 2.5 ind) and eight of the counts were of calves. The relationship between humpback whale occurrence and environmental variables including Antarctic krill (Euphausia superba) abundance from acoustic recordings, hydrography, bathymetry and production was tested using general additive models. Only temperature increased the predictive power of the model with whale occurrence increasing with the decreasing temperature in more southern areas.