SWIMMING SPEEDS OF SINGING AND NON-SINGING HUMPBACK WHALES DURING MIGRATION

Limited data exist on swimming speeds of humpback whales ( Megaptera novaeangliae ) and none on swimming speeds of singing whales during migration. We tracked humpback whales visually and acoustically during migration from the breeding grounds past our study site on the east coast of Australia (latitude 26°28'S). The mean swimming speed for whales while singing was 2.5 km/h, significantly less than for non-singing whales with a mean of 4.0 km/h but significantly greater than the mean of 1.6 km/h observed for singing whales on the Hawaiian breeding grounds. Between song sessions, there was no significant difference in speeds between whales that had been singing and other whales. Migration speeds were less for whales while singing but increased during the season. Although humpback whales can swim rapidly while singing (maximum observed 15.6 km/h), they generally do not do so, even during migration. Slower migration by singers would delay their return to the polar feeding areas and may be costly, but may be a strategy to provide access to more females.     

INTERACTIONS OF SINGING HUMPBACK WHALES WITH OTHER MALES

Two non-mutually exclusive hypotheses on the function of the humpback whale song are: (1) it attracts females to the male singer; (2) it is a male-male display, that may order status. To evaluate these, from 24 January-13 April 1997 off Maui, Hawaii, 42 singers were located, audio-recorded, photo-identified and monitored for interactions with other whales. Whales that joined singers were biopsy sampled for molecular determination of sex. In 76% (32 of 42) of the interactions, a lone non-singing adult joined the singer. In the remainder, singers stopped singing and joined a nearby group or accompanied other whales. In 81% (26 of 32) instances where a lone adult joined a singer, the pair split again within minutes; otherwise a group formed. In one such group the pair became a trio and eventually joined a competitive group. Behavior in joining/splitting interactions ranged from a single pass-by, to surface activity such as tail lobs and breaches. The sex of 22 joiners was determined: 14 genetically and eight behaviorally. All were males. Humpback whale song preceded, and at times followed, male-male interactions of variable duration and agonistic level in and around the breeding season. If considered within the context of a proposed dominance polygyny mating system, these observations appear to support speculation that the song may function in male social ordering.     

Humpback Whale Song and Foraging Behavior on an Antarctic Feeding Ground

Reports of humpback whale (Megaptera novaeangliae) song chorusing occurring outside the breeding grounds are becoming more common, but song structure and underwater behavior of individual singers on feeding grounds and migration routes remain unknown. Here, ten humpback whales in the Western Antarctic Peninsula were tagged in May 2010 with non-invasive, suction-cup attached tags to study foraging ecology and acoustic behavior. Background song was identified on all ten records, but additionally, acoustic records of two whales showed intense and continuous singing, with a level of organization and structure approaching that of typical breeding ground song. The songs, produced either by the tagged animals or close associates, shared phrase types and theme structure with one another, and some song bouts lasted close to an hour. Dive behavior of tagged animals during the time of sound production showed song occurring during periods of active diving, sometimes to depths greater than 100 m. One tag record also contained song in the presence of feeding lunges identified from the behavioral sensors, indicating that mating displays occur in areas worthy of foraging. These data show behavioral flexibility as the humpbacks manage competing needs to continue to feed and to prepare for the breeding season during late fall. This may also signify an ability to engage in breeding activities outside of the traditional, warm water breeding ground locations.     

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.     

Detecting diel patterns in the songs of Chipping Sparrows using citizen-science data

Previous studies have revealed diel patterns in the songs of Chipping Sparrows (Spizella passerina), with songs shorter in duration before dawn than after. However, the extent to which this phenomenon generalizes to the full geographic range of these sparrows is unclear, as is the question of whether citizen-science data can be used to detect diel patterns in song. We analyzed all available songs of Chipping Sparrows from the Macaulay Library and xeno-canto databases and compared the distributions of song features of recordings made at different times of day. We show that, across their entire geographic range in North and Central America, Chipping Sparrows sing shorter songs before sunrise (dawn song) than after sunrise (day song). Furthermore, we show that Chipping Sparrows shorten their songs by singing fewer syllables, not by singing faster: the number of syllables per song accounts for the observed difference in duration, not the syllable nor the intersyllable duration. Our results demonstrate that recordings from public repositories can be used to determine whether daily song patterns exist in species even in the absence of prior fieldwork, and we further propose that citizen-science recordings can be used to inform cross-species hypotheses and facilitate future studies to determine whether diel patterns in song are associated with differences in social behavior.     

The multiple functions of male song within the humpback whale (Megaptera novaeangliae) mating system: review,evaluation, and synthesis

Humpback whales (Megaptera novaeangliae) are seasonal breeders, annually migrating from high‐latitude summer feeding grounds to low‐latitude winter breeding grounds. The social matrix on the winter grounds is a loose network of interacting individuals and groups and notably includes lone males that produce long bouts of complex song that collectively yield an asynchronous chorus. Occasionally, a male will sing while accompanying other whales. Despite a wealth of knowledge about the social matrix, the full characterization of the mating system remains unresolved, without any firm consensus, as does the function of song within that system. Here, I consider and critically analyse three proposed functions of song that have received the most attention in the literature: female attraction to individual singers, determining or facilitating male–male interactions, and attracting females to a male aggregation within the context of a lekking system. Female attraction suggests that humpback song is an advertisement and invitation to females, but field observations and song playback studies reveal that female visits to individual singers are virtually absent. Other observations suggest instead that females might convey their presence to singers (or to other males) through the percussive sounds of flipper or tail slapping or possibly through vocalizations. There is some evidence for male–male interactions, both dominance and affiliative: visits to singers are almost always other lone males not singing at that time. The joiner may be seeking a coalition with the singer to engage cooperatively in attempts to obtain females, or may be seeking to disrupt the song or to affirm his dominance. Some observations support one or the other intent. However, other observations, in part based on the brevity of most pairings, suggest that the joiner is prospecting, seeking to determine whether the singer is accompanying a female, and if not soon departs. In the lekking hypothesis, the aggregation of vocalizing males on a winter ground and the visits there by non‐maternal females apparently for mating meet the fundamental definition of a lekking system and its role though communal display in attracting females to the aggregation, although not to an individual singer. Communal singing is viewed as a form of by‐product mutualism in which individuals benefit one another as incidental consequences of their own selfish actions. Possibly, communal singing may also act to stimulate female receptivity. Thus, there are both limitations and merit in all three proposals. Full consideration of song as serving multiple functions is therefore necessary to understand its role in the mating system and the forces acting on the evolution of song. I suggest that song may be the prime vector recruiting colonists to new winter grounds pioneered by vagrant males as population pressures increase or as former winter grounds become unavailable or undesirable, with such instances documented relatively recently. Speculatively, song may have evolved historically as an aggregating call during the dynamic ocean conditions and resulting habitat uncertainties in the late Miocene–early Pliocene epochs when Megaptera began to proliferate. Early song may have been comprised of simpler precursor sounds that through natural selection and ritualization evolved into complex song.     

Humpback whale songs during winter in subarctic waters

The songs of the male humpback whales (Megaptera novaeangliae) have traditionally been associated with mating at tropical and subtropical mating grounds during winter. However, songs also occur out of mating season, both on feeding grounds in spring, late summer and fall. This study provides the first report of humpback whale singing behaviour in the subarctic waters of Northeast Iceland (Skjálfandi Bay) using long-term bottom-moored acoustic recorders during September 2008–February 2009 and from April to September 2009. Singing started in late November and peaked in February, within the breeding season. No songs were detected from spring to fall, despite visual detections of humpback whales. Non-song sound signals from humpback whales were detected during all recording months. Songs were partly composed of fundamental units common with other known mating grounds, and partly of song units likely unique to the study area. The variety of song unit types in the songs increased at the end of the winter recordings, indicating a gradual change in the songs throughout the winter season; as has been shown on traditional mating grounds. The relative proportion of songs compared with non-song signals was higher during dark hours than daylight hours. The short light periods of the winter, and where food is available, likely influence the daily occurrence of humpback whales’ songs in the subarctic.     

Acoustic monitoring on a humpback whale (Megaptera novaeangliae) feeding ground shows continual singing into late Spring

Singing by males is a major feature of the mating system of humpback whales, Megaptera novaeangliae (Borowski). Although a few songs have been opportunistically recorded on the whales' high-latitude feeding grounds, singing in these regions was thought to be only sporadic. We report results from the first continuous acoustic monitoring of a humpback whale feeding ground (off Cape Cod, MA, USA) in spring. Using autonomous sea-floor recording systems, we found singing on a daily basis over the entire 25 day monitoring period, from 14 May to 7 June 2000. For much of the period, song was recorded 24 h per day. These results, combined with evidence for aseasonal conceptions in whaling catch data, suggest that the humpback whale breeding season should no longer be considered as confined to lower-latitude regions in winter. Rather, we suggest breeding extends geographically and temporally onto feeding grounds into at least spring and early summer. Singing at these times represents either low-cost opportunistic advertising by (perhaps relatively few) males to court females that failed to conceive during the winter, and/or possibly an intrasexual display.     

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.     

Long-term influence of simulated territorial intrusions on dawn and dusk singing in the Winter Wren: spring versus autumn

Males of many songbird species have peaks of singing activity at dawn and dusk. Singing during those twilight periods can function in territory proclamation, and males are suggested to adjust song output to the level of intruder pressure. We used song playback during the breeding season to simulate intrusions into territories of male Winter Wrens (Troglodytes troglodytes) shortly after dawn. We then compared male singing behaviour during the dawn and dusk chorus before and 1 day after the simulated intrusion. One day after the playback, male Wrens increased their song output before sunrise, which confirms our results from a previous study on dawn singing in autumn territories. At dusk, on the evening following the playback, males slightly increased song output after sunset, but singing activity at dusk was generally very low. We found no significant changes of song output after sunrise, before sunset, and between 2 days of control without playback. These results are consistent with the hypothesis that dawn and dusk singing is important for territory defence in spring. Unlike in autumn, however, increased singing in spring at dawn and dusk could also serve to defend other resources such as fertile mates or to strengthen the pair bond after a territorial challenge. In comparison with the results on autumnal singing, male Wrens started singing earlier at dawn during the breeding season, and they generally sang more songs at dawn and immediately after playback. The increase in absolute numbers of songs sung in the morning after playback seemed greater in spring than in autumn; however, the proportional increase relative to overall song output was similar in both seasons.     

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.     

Modelling heterogeneity in detection probabilities in land and aerial abundance surveys in humpback whales (<Emphasis Type="Italic">Megaptera novaeangliae</Emphasis>)

The effective management and conservation of animal populations relies on statistically-sound and replicable surveys to obtain estimates of abundance and assess trends. Surveys of cetaceans, such as humpback whales Megaptera novaeangliae, are difficult to conduct and are particularly affected by bias in detection probability. For example, the probability of detection of whales from land decreases substantially with increased distance from the platform. This distance effect is also true for aerial surveys, combined with the problem that animals are unavailable for detection (underwater) whilst in the field of view. We present a novel approach that combines corrected double-platform land surveys with corrected aerial surveys to obtain a robust estimate of g(0), the probability of detection on the survey line, for aerial surveys of migrating humpback whales. Several sources of heterogeneity in detection probabilities were identified within the land and aerial surveys (including group composition, bearing of first sighting, number of groups being tracked simultaneously and cloud cover). After including these into our estimate of ĝ(0), we found that only 29% of available whales are being detected on the survey line (ĝ(0) = 0.288), which is a considerably smaller estimate than many available for humpback whales using other methods. Incorporating heterogeneity into the population surveys shows that we are likely to be underestimating the population size of whales on the east coast of Australia. The implications of this result for their conservation and management in light of increased whale-human conflict is discussed.     

Minimal similarity in songs suggests limited exchange between humpback whales (Megaptera novaeangliae) in the southern Indian Ocean

Comparing humpback whale song from different breeding assemblages can reveal similarities in song due to acoustically interacting males, and therefore indirectly test whether males from different breeding sites are mixing. Northern Hemisphere song comparisons illustrated that whales within ocean basins share similar songs and are subpopulations within a larger population, whereas whales in different ocean basins are isolated populations and therefore do not share songs. During the 2006 breeding season, recordings were collected in Madagascar and Western Australia, and were compared visually plus aurally. Both regions shared one theme, whereas each region had four and six private themes, respectively. This study had a substantially low number of shared themes. The co‐occurrence of one theme was interpreted as an indication of limited exchange between these breeding assemblages, and we speculate that limited song similarity is due to inter‐oceanic interactions. Male(s) from an Indian Ocean breeding group could be exposed to novel song when they geographically overlap, and acoustically interact, with males from a different ocean basin. Novel song could induce rapid temporal changes as new song content is incorporated, thereby minimizing song similarities between that breeding group and other Indian Ocean breeding groups that were not exposed to the novel song.     

The potential beginning of a postwhaling recovery in New Zealand humpback whales (Megaptera novaeangliae)

Between the 1940s and 1970s Southern Hemisphere populations of humpback whales (including eastern Australia and Oceania populations) were hunted to near extinction by extensive commercial whaling activities in Antarctica, with fewer whales taken in shore whaling operations in New Zealand, Australia (including Norfolk Island) and Tonga. Variable rates of recovery of these populations have been documented, ranging from recovery to prewhaling numbers in eastern Australian humpbacks to very little sign of recovery in many Oceania populations. Here we analyze recent sighting data collected over 12 yr, from annual surveys in Cook Strait, New Zealand. The data show an increase in sightings, at an estimated rate of 13% (95% CI of 4.9% and 21.7%) in 2015, of humpback whales migrating through Cook Strait. The wide confidence intervals preclude substantive conclusions about the rate of increase but suggest humpback whales are returning to this region in increasing numbers, indicating an influx of immigrants from neighboring areas, namely eastern Australia.     

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.     

ESTIMATES OF THE NUMBERS OF HUMPBACK WHALES OBSERVED MIGRATING PAST CAPE VIDAL, SOUTH AFRICA, 1988–1991

A recovery-monitoring program of the African east coast humpback whale population was carried out through shore-based visual surveys from Cape Vidal, northern Natal. Surveys of the northward migration were undertaken each winter from 1988 to 1991, and a survey of the southward migration was undertaken in 1990. Independent observer surveys were undertaken during June 1990 and during the entire 1991 survey. Hourly densities of groups sighted each day were adjusted for groups missed by observers with distance from the shore and under different sighting conditions. Densities were multiplied by 24 h and the mean group size of the survey year to give resulting daily densities of individuals, which were summed to provide totals of whales sighted during each year's survey. The best estimate of population size was 1,711 (made during the northward migration of 1990), although this is likely to be biased downwards by a proportion of the population passing outside of observers'view. Bootstrapping of the 1991 daily data resulted in CVs between 11.4% and 12.2%. The numbers sighted show the population to have undergone considerable recovery since protection in October 1963.     

BLUE WHALE VISUAL AND ACOUSTIC ENCOUNTER RATES IN THE SOUTHERN CALIFORNIA BIGHT

The relationship between blue whale ( Balaenoptera musculus ) visual and acoustic encounter rates was quantitatively evaluated using hourly counts of detected whales during shipboard surveys off southern California. Encounter rates were estimated using temporal, geographic, and weather variables within a generalized additive model framework. Visual encounters (2.06 animals/h, CV = 0.10) varied with subregion, Julian day, time of day, and year. Acoustic encounters of whales producing pulsed A and tonal B call sequences (song; 0.65 animals/h, CV = 0.06) varied by Julian day, survey mode (transit or stationary), and subregion, and encounters of whales producing downswept (D) calls (0.41 animals/h, CV = 0.09) varied by Julian day and the number of animals seen. Inclusion of Julian day in all models reflects the seasonal occurrence of blue whales off southern California; however, the seasonal peak in visual encounters and acoustic encounters of D calling whales (July–August) was offset from the peak in acoustic encounters of singing whales (August–September). The relationship between visual and acoustic encounter rates varied regionally, with significant differences in several northern regions. The number of whales heard D calling was positively related to the number of animals seen, whereas the number of singing whales was not related to visual encounter rate.     

HISTORICAL OBSERVATIONS OF HUMPBACK AND BLUE WHALES IN THE NORTH ATLANTIC OCEAN: CLUES TO MIGRATORY ROUTES AND POSSIBLY ADDITIONAL FEEDING GROUNDS

The seasonal distributions of humpback and blue whales ( Megaptera novaeangliae and Balaenoptera musculus , respectively) in the North Atlantic Ocean are not fully understood. Although humpbacks have been studied intensively in nearshore or coastal feeding and breeding areas, their migratory movements between these areas have been largely inferred. Blue whales have only been studied intensively along the north shore of the Gulf of St. Lawrence, and their seasonal occurrence and movements elsewhere in the North Atlantic are poorly known. We investigated the historical seasonal distributions of these two species using sighting and catch data extracted from American 18th and 19th century whaling logbooks. These data suggest that humpback whales migrated seasonally from low-latitude calving/ breeding grounds over a protracted period, and that some of them traveled far offshore rather than following coastal routes. Also, at least some humpbacks apparently fed early in the summer west of the Mid-Atlantic Ridge, well south of their known present-day feeding grounds. In assessing the present status of the North Atlantic humpback population, it will be important to determine whether such offshore feeding does in fact occur. Blue whales were present across the southern half of the North Atlantic during the autumn and winter months, and farther north in spring and summer, but we had too few data points to support inferences about these whales' migratory timing and routes.