Bellwethers of change: population modelling of North Pacific humpback whales from 2002 through 2021 reveals shift from recovery to climate response.

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.

Temporal dynamics of mother-offspring relationships in Bigg's killer whales: opportunities for kin-directed help by post-reproductive females.

Age-related changes in the patterns of local relatedness (kinship dynamics) can be a significant selective force shaping the evolution of life history and social behaviour. In humans and some species of toothed whales, average female relatedness increases with age, which can select for a prolonged post-reproductive lifespan in older females due to both costs of reproductive conflict and benefits of late-life helping of kin. Killer whales (Orcinus orca) provide a valuable system for exploring social dynamics related to such costs and benefits in a mammal with an extended post-reproductive female lifespan. We use more than 40 years of demographic and association data on the mammal-eating Bigg's killer whale to quantify how mother-offspring social relationships change with offspring age and identify opportunities for late-life helping and the potential for an intergenerational reproductive conflict. Our results suggest a high degree of male philopatry and female-biased budding dispersal in Bigg's killer whales, with some variability in the dispersal rate for both sexes. These patterns of dispersal provide opportunities for late-life helping particularly between mothers and their adult sons, while partly mitigating the costs of mother-daughter reproductive conflict. Our results provide an important step towards understanding why and how menopause has evolved in Bigg's killer whales.

A collaborative and near-comprehensive North Pacific humpback whale photo-ID dataset.

We present an ocean-basin-scale dataset that includes tail fluke photographic identification (photo-ID) and encounter data for most living individual humpback whales (Megaptera novaeangliae) in the North Pacific Ocean. The dataset was built through a broad collaboration combining 39 separate curated photo-ID catalogs, supplemented with community science data. Data from throughout the North Pacific were aggregated into 13 regions, including six breeding regions, six feeding regions, and one migratory corridor. All images were compared with minimal pre-processing using a recently developed image recognition algorithm based on machine learning through artificial intelligence; this system is capable of rapidly detecting matches between individuals with an estimated 97-99% accuracy. For the 2001-2021 study period, a total of 27,956 unique individuals were documented in 157,350 encounters. Each individual was encountered, on average, in 5.6 sampling periods (i.e., breeding and feeding seasons), with an annual average of 87% of whales encountered in more than one season. The combined dataset and image recognition tool represents a living and accessible resource for collaborative, basin-wide studies of a keystone marine mammal in a time of rapid ecological change.

Integrating technologies provides insight into the subsurface foraging behaviour of a humpback whale (Megaptera novaeangliae) feeding on walleye pollock (Gadus chalcogrammus) in Juan de Fuca Strait, Canada.

Subsurface foraging is an important proportion of the activity budget of rorqual whales, yet information on their behaviour underwater remains challenging to obtain. Rorquals are assumed to feed throughout the water column and to select prey as a function of depth, availability and density, but there remain limitations in the precise identification of targeted prey. Current data on rorqual foraging in western Canadian waters have thus been limited to observations of prey species amenable to surface feeding, such as euphausiids and Pacific herring (Clupea pallasii), with no information on deeper alternative prey sources. We measured the foraging behaviour of a humpback whale (Megaptera novaeangliae) in Juan de Fuca Strait, British Columbia, using three complimentary methods: whale-borne tag data, acoustic prey mapping, and fecal sub-sampling. Acoustically detected prey layers were near the seafloor and consistent with dense schools of walleye pollock (Gadus chalcogrammus) distributed above more diffuse aggregations of pollock. Analysis of a fecal sample from the tagged whale confirmed that it had been feeding on pollock. Integrating the dive profile with the prey data revealed that the whale's foraging effort followed the general pattern of areal prey density, wherein the whale had a higher lunge-feeding rate at the highest prey abundance and stopped feeding when prey became limited. Our findings of a humpback whale feeding on seasonally energy-dense fish like walleye pollock, which are potentially abundant in British Columbia, suggests that pollock may be an important prey source for this rapidly growing whale population. This result is informative when assessing regional fishing activities for semi-pelagic species as well as the whales' vulnerability to fishing gear entanglements and feeding disturbances during a narrow window of prey acquisition.

Acoustic and visual cetacean surveys reveal year-round spatial and temporal distributions for multiple species in northern British Columbia, Canada.

Cetaceans spend most of their time below the surface of the sea, highlighting the importance of passive acoustic monitoring as a tool to facilitate understanding and mapping their year-round spatial and temporal distributions. To increase our limited knowledge of cetacean acoustic detection patterns for the east and west coasts of Gwaii Haanas, a remote protected area on Haida Gwaii, BC, Canada, acoustic datasets recorded off SG̱ang Gwaay (Sep 2009-May 2011), Gowgaia Slope (Jul 2017-Jul 2019), and Ramsay Island (Aug 2018-Aug 2019) were analyzed. Comparing overlapping periods of visual surveys and acoustic monitoring confirmed presence of 12 cetacean species/species groups within the study region. Seasonal patterns were identified for blue, fin, humpback, grey and sperm whale acoustic signals. Killer whale and delphinid acoustic signals occurred year-round on both coasts of Haida Gwaii and showed strong diel variation. Cuvier's, Baird's, beaked whale and porpoise clicks, were identified in high-frequency recordings on the west coast. Correlations between environmental factors, chlorophyll-a and sea surface temperature, and cetacean acoustic occurrence off Gwaii Haanas were also examined. This study is the first to acoustically monitor Gwaii Haanas waters for an extended continuous period and therefore serves as a baseline from which to monitor future changes.

Evidence from sperm whale clans of symbolic marking in non-human cultures.

Culture, a pillar of the remarkable ecological success of humans, is increasingly recognized as a powerful force structuring nonhuman animal populations. A key gap between these two types of culture is quantitative evidence of symbolic markers-seemingly arbitrary traits that function as reliable indicators of cultural group membership to conspecifics. Using acoustic data collected from 23 Pacific Ocean locations, we provide quantitative evidence that certain sperm whale acoustic signals exhibit spatial patterns consistent with a symbolic marker function. Culture segments sperm whale populations into behaviorally distinct clans, which are defined based on dialects of stereotyped click patterns (codas). We classified 23,429 codas into types using contaminated mixture models and hierarchically clustered coda repertoires into seven clans based on similarities in coda usage; then we evaluated whether coda usage varied with geographic distance within clans or with spatial overlap between clans. Similarities in within-clan usage of both "identity codas" (coda types diagnostic of clan identity) and "nonidentity codas" (coda types used by multiple clans) decrease as space between repertoire recording locations increases. However, between-clan similarity in identity, but not nonidentity, coda usage decreases as clan spatial overlap increases. This matches expectations if sympatry is related to a measurable pressure to diversify to make cultural divisions sharper, thereby providing evidence that identity codas function as symbolic markers of clan identity. Our study provides quantitative evidence of arbitrary traits, resembling human ethnic markers, conveying cultural identity outside of humans, and highlights remarkable similarities in the distributions of human ethnolinguistic groups and sperm whale clans.

Assessing social structure: a data-driven approach to define associations between individuals.

Our interpretation of animal social structures is inherently dependent on our ability to define association criteria that are biologically meaningful. However, association thresholds are often based upon generalized preconceptions of a species' social behaviour, and the impact of using these arbitrary definitions has been largely overlooked. In this study we suggest a probability-based method for defining association thresholds using lagged identification rates on photographic records of identifiable individuals. This technique uses a simple model of emigration/immigration from photographable clusters to identify the time-dependent lag value between identifications of two individuals that corresponds to approximately 75% probability of being in close spatial proximity and likely associating. This lag value is then used as the threshold to define associations for social analyses. We applied the technique to a dataset of northern resident killer whales (Orcinus orca) in the Northeast Pacific and tested its performance against two arbitrary thresholds. The probabilistic association maximized the variation in association strengths at different levels of the social structure, in line with known social patterns in this population. Furthermore, variability in inferred social structure metrics generated by different association criteria highlighted the consequential effect of choosing arbitrary thresholds. Data-driven association thresholds are a promising approach to study populations without the need to subjectively define associations in the field, especially in societies with prominent fission-fusion dynamics. This method is applicable to any dataset of sequential identifications where it can be assumed that associated individuals will tend to be identified in close proximity. Supplementary Information: The online version contains supplementary material available at 10.1007/s42991-022-00231-9.

A long postreproductive life span is a shared trait among genetically distinct killer whale populations.

The extended female postreproductive life span found in humans and some toothed whales remains an evolutionary puzzle. Theory predicts demographic patterns resulting in increased female relatedness with age (kinship dynamics) can select for a prolonged postreproductive life span due to the combined costs of intergenerational reproductive conflict and benefits of late-life helping. Here, we test this prediction using >40 years of longitudinal demographic data from the sympatric yet genetically distinct killer whale ecotypes: resident and Bigg's killer whales. The female relatedness with age is predicted to increase in both ecotypes, but with a less steep increase in Bigg's due to their different social structure. Here, we show that there is a significant postreproductive life span in both ecotypes with >30% of adult female years being lived as postreproductive, supporting the general prediction that an increase in local relatedness with age predisposes the evolution of a postreproductive life span. Differences in the magnitude of kinship dynamics however did not influence the timing or duration of the postreproductive life span with females in both ecotypes terminating reproduction before their mid-40s followed by an expected postreproductive period of about 20 years. Our results highlight the important role of kinship dynamics in the evolution of a long postreproductive life span in long-lived mammals, while further implying that the timing of menopause may be a robust trait that is persistent despite substantial variation in demographic patterns among populations.

Postreproductive killer whale grandmothers improve the survival of their grandoffspring.

Understanding why females of some mammalian species cease ovulation prior to the end of life is a long-standing interdisciplinary and evolutionary challenge. In humans and some species of toothed whales, females can live for decades after stopping reproduction. This unusual life history trait is thought to have evolved, in part, due to the inclusive fitness benefits that postreproductive females gain by helping kin. In humans, grandmothers gain inclusive fitness benefits by increasing their number of surviving grandoffspring, referred to as the grandmother effect. Among toothed whales, the grandmother effect has not been rigorously tested. Here, we test for the grandmother effect in killer whales, by quantifying grandoffspring survival with living or recently deceased reproductive and postreproductive grandmothers, and show that postreproductive grandmothers provide significant survival benefits to their grandoffspring above that provided by reproductive grandmothers. This provides evidence of the grandmother effect in a nonhuman menopausal species. By stopping reproduction, grandmothers avoid reproductive conflict with their daughters, and offer increased benefits to their grandoffspring. The benefits postreproductive grandmothers provide to their grandoffspring are shown to be most important in difficult times where the salmon abundance is low to moderate. The postreproductive grandmother effect we report, together with the known costs of late-life reproduction in killer whales, can help explain the long postreproductive life spans of resident killer whales.