Maneuverability in cetaceans is facilitated by pectoral flippers, flukes and spinal flexibility, features that are pronounced in humpback whales (Megaptera novaeangliae). Humpback whales exhibit several foraging tactics requiring high maneuverability not seen in other baleen whales, including bubble-net feeding. We hypothesized that the significant lift force produced by the humpback whale's uniquely large pectoral flippers will result in them being the only species observed executing the tight, high-speed, sustained turns characteristic of solitary bubble-net feeding. To test this hypothesis, we used a combination of inertial sensor tag data and unoccupied aerial systems (UAS; drone) photogrammetry to quantify the turning performance of solitary bubble-net feeding humpback whales, and compared this to similar data from six other mysticete species. We found that solitary bubble-net feeding humpback whales exhibited centripetal accelerations (0.46 m s-2) that exceeded the upper limit quantified in comparable turns by all six other mysticetes. This enhanced turning performance can be attributed to a substantial lift force generated by the humpback whale's pectoral flippers (7800±85 N), which contributes to centripetal acceleration and facilitates faster roll rates, allowing humpback whales to more quickly bank inwards and utilize their spinal flexibility to decrease their turning radius. Our findings demonstrate how humpback whales are uniquely adapted to exploit prey patches that might otherwise be insufficient for capture by animals of such a large size.

Given recent declines in North Pacific humpback whale (Megaptera novaeangliae) reproductive output and calf survival, there is additional urgency to better understand how mother-calf pairs allocate energy resources across their migratory cycle. Here, unoccupied aerial system (UAS; or drone) photogrammetry was used to quantify the body size and condition (BC) of humpback whales on their Hawai'i (HI) breeding and Southeast Alaska (SEAK) feeding grounds. Between 2018 and 2022, we collected 2410 measurements of 1659 individuals. Rates of change in body volume (BV) and length (BL) were quantified using 803 repeat measurements of 275 individuals. On average, HI mothers lost 0.106 m3 or 96.84 kg day-1 while fasting, equivalent to 2641 MJ day-1 or 830 kg of krill and 424 kg of Pacific herring daily. HI calf BV and BL increased by 0.035 m3 and 2.6 cm day-1, respectively. In SEAK, maternal BV increased by 0.015 m3 or 14.54 kg day-1 (367 MJ day-1), while calf BV and BL increased by 0.039 m3 and 0.93 cm day-1, respectively. Maternal investment in calf growth correlated with both female BL and BC, with larger females producing larger, faster-growing calves. Finally, using 330 measurements from 156 females, we quantified differences in BC increase over four feeding seasons. Lactating females exhibited an average BC increase of 6.10%, half that of unclassified females (13.51%) and six times lower than pregnant females (37%). These findings represent novel insights into the life history of humpback whales across their migratory cycle, providing key baseline data for bioenergetic models elucidating the effects of anthropogenic disturbance and rapidly changing ocean ecosystems. KEY POINTS: On average, Hawai'i (HI) mothers lost 0.106 m3 or 96.84 kg day-1, equivalent to 2641 MJ day-1. Over a 60 day period, this corresponded to an estimated mean energetic cost of 158 GJ, or ≈50 tons of krill or ≈25 tons of Pacific herring, surpassing the total energetic cost of gestation estimated for humpback whales of similar length. In Southeast Alaska (SEAK), maternal body volume (BV) increased by just 0.015 m3 or 14.54 kg day-1 (367 MJ day-1). Further, SEAK lactating females showed the slowest rates of growth in body width and condition over a 150 day period compared to non-lactating females. Maternal investment in calf growth correlated with both maternal length and body condition, with larger females producing larger, faster-growing calves. In HI, however, the ratio between maternal BV lost and calf BV gained (conversion efficiency) was relatively low compared to other mammals.

Isolation by distance and biogeographical boundaries define patterns of population genetic structure for harbour porpoise along the Pacific coast from California to British Columbia. Until recently, inadequate sample sizes in many regions constrained efforts to characterise population genetic structure throughout the coastal waters of Alaska. Here, tissue samples from beachcast strandings and fisheries bycatch were supplemented with targeted environmental DNA (eDNA) samples in key regions of Alaska coastal and inland waters. Using a geographically explicit, hierarchical approach, we examined the genetic structure of Alaska harbour porpoises, using both mitochondrial DNA (mtDNA) sequence data and multilocus SNP genotypes. Despite a lack of evidence of genetic differentiation from nuclear SNP loci, patterns of relatedness and genetic differentiation from mtDNA suggest natal philopatry at multiple geographic scales, with limited gene flow among sites possibly mediated by male dispersal. A priori clustering of sampled areas at an intermediate scale (eastern and western Bering Sea, Gulf of Alaska and Southeast Alaska) best explained the genetic variance (12.37%) among regions. In addition, mtDNA differentiation between the Gulf of Alaska and eastern Bering Sea, and among regions within the Gulf of Alaska, indicated significant genetic structuring of harbour porpoise populations in Southeast Alaska. The targeted collection of eDNA samples from strata within Southeast Alaska was key for elevating the statistical power of our mtDNA dataset, and findings indicate limited dispersal between neighbouring strata within coastal and inland waters. These results provide evidence supporting a population boundary within the currently recognised Southeast Alaska Stock. Together, these findings will prove useful for ongoing management efforts to reduce fisheries conflict and conserve genetic diversity in this iconic coastal species.

For the 40 years after the end of commercial whaling in 1976, humpback whale populations in the North Pacific Ocean exhibited a prolonged period of recovery. Using mark–recapture methods on the largest individual photo-identification dataset ever assembled for a cetacean, we estimated annual ocean-basin-wide abundance for the species from 2002 through 2021. Trends in annual estimates describe strong post-whaling era population recovery from 16 875 (± 5955) in 2002 to a peak abundance estimate of 33 488 (± 4455) in 2012. An apparent 20% decline from 2012 to 2021, 33 488 (± 4455) to 26 662 (± 4192), suggests the population abruptly reached carrying capacity due to loss of prey resources. This was particularly evident for humpback whales wintering in Hawai‘i, where, by 2021, estimated abundance had declined by 34% from a peak in 2013, down to abundance levels previously seen in 2006, and contrasted to an absence of decline in Mainland Mexico breeding humpbacks. The strongest marine heatwave recorded globally to date during the 2014–2016 period appeared to have altered the course of species recovery, with enduring effects. Extending this time series will allow humpback whales to serve as an indicator species for the ecosystem in the face of a changing climate.

Abstract Heteroplasmy in the mitochondrial genome offers a rare opportunity to track the evolution of a newly arising maternal lineage in populations of non-model species. Here, we identified a previously unreported mitochondrial DNA haplotype while assembling an integrated database of DNA profiles and photo-identification records from humpback whales in southeastern Alaska (SEAK). The haplotype, referred to as A8, was shared by only 2 individuals, a mature female with her female calf, and differed by only a single base pair from a common haplotype in the North Pacific, referred to as A−. To investigate the origins of the A8 haplotype, we reviewed n = 1,089 electropherograms (including replicate samples) of n = 710 individuals with A− haplotypes from an existing collection. From this review, we found 20 individuals with clear evidence of heteroplasmy for A−/A8 (parental/derived) haplotypes. Of these, 15 were encountered in SEAK, 4 were encountered on the Hawaiian breeding ground (the primary migratory destination for whales in SEAK), and 1 was encountered in the northern Gulf of Alaska. We used genotype exclusion and likelihood to identify one of the heteroplasmic females as the likely mother of the A8 cow and grandmother of the A8 calf, establishing the inheritance and germ-line fixation of the new haplotype from the parental heteroplasmy. The mutation leading to this heteroplasmy and the fixation of the A8 haplotype provide an opportunity to document the population dynamics and regional fidelity of a newly arising maternal lineage in a population recovering from exploitation.

We delineated and scored Biologically Important Areas (BIAs) for cetacean species in the Gulf of Alaska region. BIAs represent areas and times in which cetaceans are known to concentrate for activities related to reproduction, feeding, and migration, and also the known ranges of small and resident populations. This National Oceanic and Atmospheric Association (NOAA)-led effort uses structured expert elicitation principles to build upon the first version of NOAA’s BIAs for cetaceans. Supporting evidence for these BIAs came from aerial-, land-, and vessel-based surveys; satellite-tagging data; passive acoustic monitoring; Indigenous knowledge; photo-identification data; and/or prey studies. A total of 20 BIAs were identified, delineated, and scored for six species: beluga whale ( Delphinapterus leucas ), fin whale ( Balaenoptera physalus ), gray whale ( Eschrichtius robustus ), humpback whale ( Megaptera novaeangliae ), North Pacific right whale ( Eubalaena japonica ), and sperm whale ( Physeter macrocephalus ). Of the 20 total BIAs, there were two small and resident populations, one migratory, and 17 feeding areas; no reproductive BIAs were identified. An additional five watch list areas were identified, a new feature to the second version of BIAs. In addition to more comprehensive narratives and maps, the BIA II products improve upon the first version by creating metadata tables and incorporating a scoring and labeling system which improves quantification and standardization of BIAs within and across regions. BIAs are compilations of the best available science and have no inherent regulatory authority. They have been used by NOAA, other federal agencies, and the public to support planning and marine mammal impact assessments, and to inform the development of conservation measures for cetaceans.

To better understand reproductive physiology of humpback whales Megaptera novaeangliae that reside in Hawai'i and Alaska, enzyme immunoassays were validated for both progesterone and testosterone in free-ranging and stranded animals (n = 185 biopsies). Concentrations were analyzed between different depths of large segments of blubber taken from skin to muscle layers of stranded female (n = 2, 1 pregnant, 1 non-pregnant) and male (n = 1) whales. Additionally, progesterone metabolites were identified between pregnant (n = 1) and non-pregnant (n = 3) females using high pressure liquid chromatography (HPLC). Progesterone concentrations were compared between juvenile (i.e., sexually immature), lactating, and pregnant females, and male whales, and pregnancy rates of sexually mature females were calculated. Based on replicate samples from ship struck animals collected at 7 depth locations, blubber containing the highest concentration of progesterone was located 1 cm below the skin for females, and the highest concentration of testosterone was in the skin layer of one male whale. HPLC of blubber samples of pregnant and non-pregnant females contain different immunoreactive progesterone metabolites, with the non-pregnant female eluate comprised of a more polar, and possibly conjugated, form of progesterone than the pregnant female. In females, concentrations of progesterone were highest in the blubber of pregnant (n = 28, 28.6 +/- 6.9 ng/g), followed by lactating (n = 16, 0.9 +/- 0.1 ng/g), and female juvenile (n = 5, 1.0 +/- 0.2 ng/g) whales. Progesterone concentrations in male (n = 24, 0.6 ng/g +/- 0.1 ng/g) tissues were the lowest all groups, and not different from lactating or juvenile females. Estimated summer season pregnancy rate among sexually mature females from the Hawai'i stock of humpback whales was 0.562 (95 % confidence interval 0.528-0.605). For lactating females, the year-round pregnancy rate was 0.243 (0.09-0.59), and varies depending on the threshold of progesterone assumed for pregnancy in the range between 3.1 and 28.5 ng/g. Our results demonstrate the synergistic value added when combining immunoreactive assays, HPLC, and long-term sighting histories to further knowledge of humpback whale reproductive physiology.

Understanding the ecosystem effects of ocean warming is increasingly important as marine heatwaves become more common and increase in severity. Here, we used Glacier Bay National Park long-term monitoring data (1985–2020) to investigate a sudden, sharp decline in humpback whale reproductive success and survival following the onset of the 2014–2016 Northeast Pacific marine heatwave (PMH). Oceanographic data confirm a persistent warm-water anomaly in 2015–2016 in Glacier Bay, months later than the PMH was documented in the North Pacific. We assessed changes in demographic parameters pre- and post-PMH using whale and calf counts and multi-state closed population capture–recapture models. Non-calf abundance decreased by 56

Extended acoustic interactions with a humpback whale ( Megaptera novaeangliae ) were captured via playbacks of the purported “throp” social call and hydrophone recordings of the animal’s vocalized responses during August 2021 in Frederick Sound, Southeast Alaska. Fluke photographs identified the animal as a female named Twain ( HappyWhale.com identity SEAK-0401) first observed some 38 years ago. We document Twain’s life history via sightings made over almost four decades. The observational history gives illuminating snapshots of the long history of the individual behind the acoustic interactions.

Despite their enormous size, whales make their living as voracious predators. To catch their much smaller, more maneuverable prey, they have developed several unique locomotor strategies that require high energetic input, high mechanical power output and a surprising degree of agility. To better understand how body size affects maneuverability at the largest scale, we used bio-logging data, aerial photogrammetry and a high-throughput approach to quantify the maneuvering performance of seven species of free-swimming baleen whale. We found that as body size increases, absolute maneuvering performance decreases: larger whales use lower accelerations and perform slower pitch-changes, rolls and turns than smaller species. We also found that baleen whales exhibit positive allometry of maneuvering performance: relative to their body size, larger whales use higher accelerations, and perform faster pitch-changes, rolls and certain types of turns than smaller species. However, not all maneuvers were impacted by body size in the same way, and we found that larger whales behaviorally adjust for their decreased agility by using turns that they can perform more effectively. The positive allometry of maneuvering performance suggests that large whales have compensated for their increased body size by evolving more effective control surfaces and by preferentially selecting maneuvers that play to their strengths.

Some of the longest and most comprehensive marine ecosystem monitoring programs were established in the Gulf of Alaska following the environmental disaster of the Exxon Valdez oil spill over 30 years ago. These monitoring programs have been successful in assessing recovery from oil spill impacts, and their continuation decades later has now provided an unparalleled assessment of ecosystem responses to another newly emerging global threat, marine heatwaves. The 2014–2016 northeast Pacific marine heatwave (PMH) in the Gulf of Alaska was the longest lasting heatwave globally over the past decade, with some cooling, but also continued warm conditions through 2019. Our analysis of 187 time series from primary production to commercial fisheries and nearshore intertidal to offshore oceanic domains demonstrate abrupt changes across trophic levels, with many responses persisting up to at least 5 years after the onset of the heatwave. Furthermore, our suite of metrics showed novel community-level groupings relative to at least a decade prior to the heatwave. Given anticipated increases in marine heatwaves under current climate projections, it remains uncertain when or if the Gulf of Alaska ecosystem will return to a pre-PMH state.

Sperm whales interact with commercially important groundfish fisheries offshore in the Gulf of Alaska (GOA). This study aims to use stable isotope analysis to better understand the trophic variability of sperm whales and their potential prey, and to use dietary mixing models to estimate the importance of prey species to sperm whale diets. We analysed tissue samples from sperm whales and seven potential prey (five groundfish and two squid species). Samples were analysed for stable carbon and nitrogen isotope ratios, and diet composition was estimated using Bayesian isotopic mixing models. Mixing model results suggest that an isotopically combined sablefish/dogfish group, skates and rockfish make up the largest proportion of sperm whale diets (35%, 28% and 12%) in the GOA. The top prey items of whales that interact more frequently with fishing vessels consisted of skates (49%) and the sablefish/dogfish group (24%). This is the first known study to provide an isotopic baseline of adult male sperm whales and these adult groundfish and offshore squid species, and to assign contributions of prey to whale diets in the GOA. This study provides information to commercial fishermen and fisheries managers to better understand trophic connections of important commercial species.

Developing a better understanding of the stress response is critical to ensuring the health and sustainability of marine mammal populations. However, accurately measuring and interpreting a stress response in free-ranging, large cetaceans is a nascent field. Here, an enzyme immunoassay for corticosterone was validated for use in biopsy samples from male humpback whales (Megaptera novaeangliae). Analyses were conducted on 247 male North Pacific humpback whale blubber samples, including 238 non-calves and 9 calves that were collected on the Hawaiian breeding and Southeast Alaskan feeding grounds from 2004 to 2006. Significant relationships were found when corticosterone concentrations were examined by year, age class and distribution between locations. When examined by year, corticosterone concentrations for male humpback whales were higher in Hawaii in 2004 than in 2005 and 2006 (p < 0.05). Corticosterone concentration also varied by age class with initially high concentrations at birth which subsequently tapered off and remained relatively low until sexual maturity was reached around age 8-10 years. Corticosterone concentrations appeared to peak in male humpback whales around 15-25 years of age. Blubber biopsies from Alaska and Hawaii had similar mean corticosterone concentrations, yet the variability in these samples was much greater for whales located in Hawaii. It is clear that much work remains to be done in order to accurately define or monitor a stress response in male humpback whales and that specific attention is required when looking at age, sex, and yearly trends. Our results suggest that a stress response may be most impacted by age and yearly oceanographic conditions and needs to be initially examined at the individual level.

Understanding reproductive profiles and timing of reproductive events is essential in the management and conservation of humpback whales (Megaptera novaeangliae). Yet compared to other parameters and life history traits, such as abundance, migratory trends, reproductive rates, behavior and communication, relatively little is known about variations in reproductive physiology, especially in males. Here, an enzyme immunoassay (EIA) for testosterone was validated for use in biopsy samples from male humpback whales. Analyses were conducted on 277 North Pacific male humpback whale blubber samples, including 268 non-calves and 9 calves that were collected in the Hawaiian breeding grounds and the Southeast Alaskan feeding grounds from 2004 to 2006. Testosterone concentrations (ng/g) were significantly different between non-calves sampled in Hawaii (n = 182) and Alaska (n = 86, p < 0.05) with peak testosterone concentrations occurring in the winter (January-March) and the lowest concentrations occurring in the summer (June-September). Fall and spring showed increasing and decreasing trends in testosterone concentrations, respectively. Blubber testosterone concentrations in non-calves and calves sampled in Alaska were not significantly different. Blubber and skin from the same individual biopsies (n = 37) were also compared, with blubber having significantly higher testosterone concentrations (p < 0.05) than skin samples. We found variability in testosterone concentration with age, suggesting that male humpbacks reach peak lifetime testosterone concentrations in the breeding grounds around age 8-25 years. The testosterone profile of male humpback whales follows a predictable pattern for capital breeders, where testosterone begins to increase prior to the breeding season, stimulating the onset of spermatogenesis. Incorporation of reproductive hormonal profiles into our overall understanding of humpback whale physiology will shed additional light on the timing of reproduction and overall health of the recently delisted Hawaii distinct population segment (DPS).

The rise of inexpensive, user-friendly cameras and editing software promises to revolutionize data collection with minimal disturbance to marine mammals. Video sequences recorded by aerial drones and GoPro cameras provided close-up views and unique perspectives of humpback whales engulfing juvenile salmon at or just below the water surface in Southeast Alaska and Prince William Sound. Although humpback feeding is famous for its flexibility, several stereotyped events were noted in the 47 lunges we analyzed. Engulfment was rapid (mean 2.07 s), and the entrance through which the tongue inverts into the ventral pouch was seen as water rushes in. Cranial elevation was a major contributor to gape, and pouch contraction sometimes began before full gape closure, with reverberating waves indicating rebounding flow of water within the expanded pouch. Expulsion of filtered water began with a small splash at the anterior of the mouth, followed by sustained excurrent flow in the mouth's central or posterior regions. Apart from a splash of rebounding water, water within the mouth was surprisingly turbulence-free during engulfment, but submersion of the whale's head created visible surface whirlpools and vortices which may aggregate prey for subsequent engulfment.