A pan-cetacean MHC amplicon sequencing panel developed and evaluated in combination with genome assemblies.

The major histocompatibility complex (MHC) is a highly polymorphic gene family that is crucial in immunity, and its diversity can be effectively used as a fitness marker for populations. Despite this, MHC remains poorly characterised in non-model species (e.g., cetaceans: whales, dolphins and porpoises) as high gene copy number variation, especially in the fast-evolving class I region, makes analyses of genomic sequences difficult. To date, only small sections of class I and IIa genes have been used to assess functional diversity in cetacean populations. Here, we undertook a systematic characterisation of the MHC class I and IIa regions in available cetacean genomes. We extracted full-length gene sequences to design pan-cetacean primers that amplified the complete exon 2 from MHC class I and IIa genes in one combined sequencing panel. We validated this panel in 19 cetacean species and described 354 alleles for both classes. Furthermore, we identified likely assembly artefacts for many MHC class I assemblies based on the presence of class I genes in the amplicon data compared to missing genes from genomes. Finally, we investigated MHC diversity using the panel in 25 humpback and 30 southern right whales, including four paternity trios for humpback whales. This revealed copy-number variable class I haplotypes in humpback whales, which is likely a common phenomenon across cetaceans. These MHC alleles will form the basis for a cetacean branch of the Immuno-Polymorphism Database (IPD-MHC), a curated resource intended to aid in the systematic compilation of MHC alleles across several species, to support conservation initiatives.

Diel spatio-temporal patterns of humpback whale singing on a high-density breeding ground.

Humpback whale song chorusing dominates the marine soundscape in Hawai'i during winter months, yet little is known about spatio-temporal habitat use patterns of singers. We analysed passive acoustic monitoring data from five sites off Maui and found that ambient noise levels associated with song chorusing decreased during daytime hours nearshore but increased offshore. To resolve whether these changes reflect a diel offshore-onshore movement or a temporal difference in singing activity, data from 71 concurrently conducted land-based theodolite surveys were analysed. Non-calf pods (n = 3082), presumably including the majority of singers, were found further offshore with increasing time of the day. Separately, we acoustically localized 217 nearshore singers using vector-sensors. During the day, distances to shore and minimum distances among singers increased, and singers switched more between being stationary and singing while travelling. Together, these findings suggest that the observed diel trends in humpback whale chorusing off Maui represent a pattern of active onshore-offshore movement of singers. We hypothesize that this may result from singers attempting to reduce intraspecific acoustic masking when densities are high nearshore and avoidance of a loud, non-humpback, biological evening chorus offshore, creating a dynamic of movement of singers aimed at increasing the efficiency of their acoustic display.

A novel RT-qPCR health assay reveals differential expression of stress and immunoregulatory genes between the seasonal migrations of humpback whales (Megaptera novaeangliae).

Health information is essential for the conservation management of whale species. However, assessing the health of free-ranging whales is challenging as samples are primarily limited to skin and blubber tissue. Reverse transcription quantitative polymerase chain reaction (RT-qPCR) offers a method to measure health from blubber RNA, providing insights into energetic status, stress and immune activity. To identify changes in health, natural differences in baseline gene expression linked to an individual's sex, reproductive status and life-history stage must first be quantified. This study aimed to establish baseline gene expression indices of health in migrating humpback whales (Megaptera novaeangliae). To do this, we developed an assay to quantify seven health-related gene transcripts (Leptin, Leptin Receptor, Adiponectin, Aryl Hydrocarbon Receptor, Tumour Necrosis Factor-α, Interleukin-6, Heat Shock Protein-70) and used Bayesian mixed effect models to assess differential baseline expression based on sex, lactation status and migration stage (northbound to and southbound from the annual breeding grounds). Results showed no significant contribution of sex to differential baseline expression. However, lactating individuals exhibited downregulated AhR and HSP-70 compared to non-lactating conspecifics. Additionally, southbound individuals demonstrated significantly upregulated HSP-70 and downregulated TNF-alpha, suggesting a relationship between these inflammation-linked transcripts and migratory fasting. Our results suggest that baseline differences due to migratory stage and lactation status should be considered in health applications of this assay. Future monitoring efforts can use our baseline measurements to better understand how gene expression is tied to population-level impacts, such as reduced prey availability or migratory stressors.

Interactive bioacoustic playback as a tool for detecting and exploring nonhuman intelligence: "conversing" with an Alaskan humpback whale.

Here we report on a rare and opportunistic acoustic turn-taking with an adult female humpback whale, known as Twain, in Southeast Alaska. Post hoc acoustic and statistical analyses of a 20-min acoustic exchange between the broadcast of a recorded contact call, known as a 'whup/throp', with call responses by Twain revealed an intentional human-whale acoustic (and behavioral) interaction. Our results show that Twain participated both physically and acoustically in three phases of interaction (Phase 1: Engagement, Phase 2: Agitation, Phase 3: Disengagement), independently determined by blind observers reporting on surface behavior and respiratory activity of the interacting whale. A close examination of both changes to the latency between Twain's calls and the temporal matching to the latency of the exemplar across phases indicated that Twain was actively engaged in the exchange during Phase 1 (Engagement), less so during Phase 2 (Agitation), and disengaged during Phase 3 (Disengagement). These results, while preliminary, point to several key considerations for effective playback design, namely the importance of salient, dynamic and adaptive playbacks, that should be utilized in experimentation with whales and other interactive nonhuman species.

New Marginal Spectrum Feature Information Views of Humpback Whale Vocalization Signals Using the EMD Analysis Methods.

Marginal spectrum (MS) feature information of humpback whale vocalization (HWV) signals is an interesting and significant research topic. Empirical mode decomposition (EMD) is a powerful time-frequency analysis tool for marine mammal vocalizations. In this paper, new MS feature innovation information of HWV signals was extracted using the EMD analysis method. Thirty-six HWV samples with a time duration of 17.2 ms were classified into Classes I, II, and III, which consisted of 15, 5, and 16 samples, respectively. The following ratios were evaluated: the average energy ratios of the 1 first intrinsic mode function (IMF1) and residual function (RF) to the referred total energy for the Class I samples; the average energy ratios of the IMF1, 2nd IMF (IMF2), and RF to the referred total energy for the Class II samples; the average energy ratios of the IMF1, 6th IMF (IMF6), and RF to the referred total energy for the Class III samples. These average energy ratios were all more than 10%. The average energy ratios of IMF1 to the referred total energy were 9.825%, 13.790%, 4.938%, 3.977%, and 3.32% in the 2980-3725, 3725-4470, 4470-5215, 10,430-11,175, and 11,175-11,920 Hz bands, respectively, in the Class I samples; 14.675% and 4.910% in the 745-1490 and 1490-2235 Hz bands, respectively, in the Class II samples; 12.0640%, 6.8850%, and 4.1040% in the 2980-3725, 3725-4470, and 11,175-11,920 Hz bands, respectively, in the Class III samples. The results of this study provide a better understanding, high resolution, and new innovative views on the information obtained from the MS features of the HWV signals.

Diversity of mitochondrial DNA in 3 species of great whales before and after modern whaling.

The 20th century commercial whaling industry severely reduced populations of great whales throughout the Southern Hemisphere. The effect of this exploitation on genetic diversity and population structure remains largely undescribed. Here, we compare pre- and post-whaling diversity of mitochondrial DNA (mtDNA) control region sequences for 3 great whales in the South Atlantic, such as the blue, humpback, and fin whale. Pre-whaling diversity is described from mtDNA extracted from bones collected near abandoned whaling stations, primarily from the South Atlantic island of South Georgia. These bones are known to represent the first stage of 20th century whaling and thus pre-whaling diversity of these populations. Post-whaling diversity is described from previously published studies reporting large-scale sampling of living whales in the Southern Hemisphere. Despite relatively high levels of surviving genetic diversity in the post-whaling populations, we found evidence of a probable loss of mtDNA lineages in all 3 species. This is evidenced by the detection of a large number of haplotypes found in the pre-whaling samples that are not present in the post-whaling samples. A rarefaction analysis further supports a loss of haplotypes in the South Atlantic humpback and Antarctic blue whale populations. The bones from former whaling stations in the South Atlantic represent a remarkable molecular archive for further investigation of the decline and ongoing recovery in the great whales of the Southern Hemisphere.

A surplus no more? Variation in krill availability impacts reproductive rates of Antarctic baleen whales.

The krill surplus hypothesis of unlimited prey resources available for Antarctic predators due to commercial whaling in the 20th century has remained largely untested since the 1970s. Rapid warming of the Western Antarctic Peninsula (WAP) over the past 50 years has resulted in decreased seasonal ice cover and a reduction of krill. The latter is being exacerbated by a commercial krill fishery in the region. Despite this, humpback whale populations have increased but may be at a threshold for growth based on these human-induced changes. Understanding how climate-mediated variation in prey availability influences humpback whale population dynamics is critical for focused management and conservation actions. Using an 8-year dataset (2013-2020), we show that inter-annual humpback whale pregnancy rates, as determined from skin-blubber biopsy samples (n = 616), are positively correlated with krill availability and fluctuations in ice cover in the previous year. Pregnancy rates showed significant inter-annual variability, between 29% and 86%. Our results indicate that krill availability is in fact limiting and affecting reproductive rates, in contrast to the krill surplus hypothesis. This suggests that this population of humpback whales may be at a threshold for population growth due to prey limitations. As a result, continued warming and increased fishing along the WAP, which continue to reduce krill stocks, will likely impact this humpback whale population and other krill predators in the region. Humpback whales are sentinel species of ecosystem health, and changes in pregnancy rates can provide quantifiable signals of the impact of environmental change at the population level. Our findings must be considered paramount in developing new and more restrictive conservation and management plans for the Antarctic marine ecosystem and minimizing the negative impacts of human activities in the region.

Multiscale relationships between humpback whales and forage species hotspots within a large marine ecosystem.

Fluctuations in prey abundance, composition, and distribution can impact predators, and when predators and fisheries target the same species, predators become essential to ecosystem-based management. Because of the difficulty in collecting concomitant predator-prey data at appropriate scales in patchy environments, few studies have identified strong linkages between cetaceans and prey, especially across large geographic areas. During summer 2018, a line-transect survey for cetaceans and coastal pelagic species was conducted over the continental shelf and slope of British Columbia, Canada, and the US West Coast, allowing for a large-scale investigation of predator-prey spatial relationships. We report on a case study of humpback whales (Megaptera novaeangliae) and their primary prey-Pacific herring (Clupea pallasii), northern anchovy (Engraulis mordax), and krill-using generalized additive models to explore the relationships between whale abundance on 10-km transect segments and prey metrics. Prey metrics included direct measures of biomass densities on segments and an original hotspot metric. For each prey species, segments in the upper fifth percentile for biomass density (across all segments) were designated hotspots, and whale counts on a segment were evaluated for their relationship to number of hotspot segments (species-specific and multispecies) within 25, 50, or 100 km. Whale abundance was not strongly related to direct measures of biomass densities, whereas models using hotspot metrics were more effective at describing variation in whale abundance, underscoring that evaluating prey at relevant and measurable scales is critical in patchy, dynamic marine environments. Our analysis highlighted differences in the distribution and prey availability for three humpback whale distinct population segments (DPSs) as defined under the US Endangered Species Act, including threatened and endangered DPSs that forage within the California Current Large Marine Ecosystem. These linkages provide insights into which prey species whales may be targeting in different regions and across multiple scales and, consequently, how climatic variability and anthropogenic risks may differentially impact these distinct predator-prey assemblages. By identifying scale-appropriate prey hotspots that co-occur with humpback whale aggregations, and with targeted, consistent prey sampling and estimations of potential consumption rates by whales, these findings can help inform the conservation and management of humpback whales within an ecosystem-based management framework.

Expert assessment of the impact of ship-strikes on cetacean welfare using the Welfare Assessment Tool for Wild Cetaceans.

Human activities are increasingly impacting our oceans and the focus tends to be on their environmental impacts, rather than consequences for animal welfare. Global shipping density has quadrupled since 1992. Unsurprisingly, increased levels of vessel collisions with cetaceans have followed this global expansion of shipping. This paper is the first to attempt to consider the severity of ship-strike on individual whale welfare. The methodology of the 'Welfare Assessment Tool for Wild Cetaceans' (WATWC) was used, which is itself based upon the Five Domains model. Expert opinion was sought on six hypothetical but realistic case studies involving humpback whales (Megaptera novaeangliae) struck by ships. Twenty-nine experts in the cetacean and welfare sector took part. They were split into two groups; Group 1 first assessed a case we judged to be the least severe and Group 2 first assessed the most severe. Both groups then additionally assessed the same four further cases. This was to investigate whether the severity of the first case influenced judgements regarding subsequent cases (i.e. expert judgements were relative) or not (i.e. judgements were absolute). No significant difference between the two groups of assessors was found; therefore, the hypothesis of relative scoring was rejected. Experts judged whales may suffer some level (>1) of overall (Domain 5) harm for the rest of their lives following a ship-strike incident. Health, closely followed by Behaviour were found to be the welfare aspects most affected by ship-strikes. Overall, the WATWC shows a robust potential to aid decision-making on wild cetacean welfare.

Humpback whale song revolutions continue to spread from the central into the eastern South Pacific.

Cultural transmission of behaviour is an important aspect of many animal communities ranging from humans to birds. Male humpback whales (Megaptera novaeangliae) sing a repetitive, stereotyped, socially learnt and culturally transmitted song display that slowly evolves each year. Most males within a population sing the same, slow-evolving song type; but in the South Pacific, song 'revolutions' have led to rapid and complete replacement of one song type by another introduced from a neighbouring population. Songs spread eastwards, from eastern Australia to French Polynesia, but the easterly extent of this transmission was unknown. Here, we investigated whether song revolutions continue to spread from the central (French Polynesia) into the eastern (Ecuador) South Pacific region. Similarity analyses using three consecutive years of song data (2016-2018) revealed that song themes recorded in 2016-2018 French Polynesian song matched song themes sung in 2018 Ecuadorian song, suggesting continued easterly transmission of song to Ecuador, and vocal connectivity across the entire South Pacific Ocean basin. This study demonstrates songs first identified in western populations can be transmitted across the entire South Pacific, supporting the potential for a circumpolar Southern Hemisphere cultural transmission of song and a vocal culture rivalled in its extent only by our own.

Intra-seasonal variation in feeding rates and diel foraging behaviour in a seasonally fasting mammal, the humpback whale.

Antarctic humpback whales forage in summer, coincident with the seasonal abundance of their primary prey, the Antarctic krill. During the feeding season, humpback whales accumulate energy stores sufficient to fuel their fasting period lasting over six months. Previous animal movement modelling work (using area-restricted search as a proxy) suggests a hyperphagic period late in the feeding season, similar in timing to some terrestrial fasting mammals. However, no direct measures of seasonal foraging behaviour existed to corroborate this hypothesis. We attached high-resolution, motion-sensing biologging tags to 69 humpback whales along the Western Antarctic Peninsula throughout the feeding season from January to June to determine how foraging effort changes throughout the season. Our results did not support existing hypotheses: we found a significant reduction in foraging presence and feeding rates from the beginning to the end of the feeding season. During the early summer period, feeding occurred during all hours at high rates. As the season progressed, foraging occurred mostly at night and at lower rates. We provide novel information on seasonal changes in foraging of humpback whales and suggest that these animals, contrary to nearly all other animals that seasonally fast, exhibit high feeding rates soon after exiting the fasting period.

Decadal-scale phenology and seasonal climate drivers of migratory baleen whales in a rapidly warming marine ecosystem.

Species' response to rapid climate change can be measured through shifts in timing of recurring biological events, known as phenology. The Gulf of Maine is one of the most rapidly warming regions of the ocean, and thus an ideal system to study phenological and biological responses to climate change. A better understanding of climate-induced changes in phenology is needed to effectively and adaptively manage human-wildlife conflicts. Using data from a 20+ year marine mammal observation program, we tested the hypothesis that the phenology of large whale habitat use in Cape Cod Bay has changed and is related to regional-scale shifts in the thermal onset of spring. We used a multi-season occupancy model to measure phenological shifts and evaluate trends in the date of peak habitat use for North Atlantic right (Eubalaena glacialis), humpback (Megaptera novaeangliae), and fin (Balaenoptera physalus) whales. The date of peak habitat use shifted by +18.1 days (0.90 days/year) for right whales and +19.1 days (0.96 days/year) for humpback whales. We then evaluated interannual variability in peak habitat use relative to thermal spring transition dates (STD), and hypothesized that right whales, as planktivorous specialist feeders, would exhibit a stronger response to thermal phenology than fin and humpback whales, which are more generalist piscivorous feeders. There was a significant negative effect of western region STD on right whale habitat use, and a significant positive effect of eastern region STD on fin whale habitat use indicating differential responses to spatial seasonal conditions. Protections for threatened and endangered whales have been designed to align with expected phenology of habitat use. Our results show that whales are becoming mismatched with static seasonal management measures through shifts in their timing of habitat use, and they suggest that effective management strategies may need to alter protections as species adapt to climate change.

Intra-season variations in distribution and abundance of humpback whales in the West Antarctic Peninsula using cruise vessels as opportunistic platforms.

Fine-scale knowledge of spatiotemporal dynamics in cetacean distribution and abundance throughout the Western Antarctic Peninsula (WAP) is sparse yet essential for effective ecosystem-based management (EBM). Cruise vessels were used as platforms of opportunity to collect data on the distribution and abundance of humpback whales (Megaptera novaeangliae) during the austral summer of 2019/2020 in a region that is also important for the Antarctic krill (Euphausia superba) fishery, to assess potential spatiotemporal interactions for future use in EBM. Data were analyzed using traditional design-based line transect methodology and spatial density surface hurdle models fitted using a set of physical environmental covariates to estimate the abundance and distribution of whales in the area, and to describe their temporal dynamics. Our results indicate a rapid increase in humpback whale abundance in the Bransfield and Gerlache Straits through December, reaching a stable abundance by mid-January. The distribution of humpback whales appeared to change from a patchier distribution in the northern Gerlache Strait to a significantly concentrated presence in the central Gerlache and southern Bransfield Straits, followed by a subsequent dispersion throughout the area. Abundance estimates agreed well with previous literature, increasing from approximately 7000 individuals in 2000 to a peak of 19,107 in 2020. Based on these estimates, we project a total krill consumption of between 1.4 and 3.7 million tons based on traditional and contemporary literature on per capita krill consumption of whales, respectively. When taken in the context of krill fishery catch data in the study area, we conclude that there is minimal spatiotemporal overlap between humpback whales and fishery activity during our study period of November-January. However, there is potential for significant interaction between the two later in the feeding season, but cetacean survey efforts need to be extended into late season in order to fully characterize this potential overlap.

Fine-scale diversity of prey detected in humpback whale feces.

Predator diets are largely influenced by prey availability and abundance. Yet, in heterogenous marine environments, identifying the prey species consumed by diving mammals remains a fundamental challenge. For rorqual whales, the energetic constraints of prey engulfment require that the whales seek areas of high prey abundance and execute discrete lunge feeding events on patches of high-density prey. Prey occurrences in feces should therefore provide meaningful insight into the dominant taxa in food patches selected by the animal. We investigated the prey consumed by humpback whales in three regions in southern British Columbia (BC), Canada, using opportunistic fecal sampling, microscopy, and DNA metabarcoding of 14 fecal samples. Fish including Pacific herring (Clupea pallasii), hake (Merluccius productus), and eulachon (Thaleichthys pacificus) were the most common fish species potentially targeted by humpback whales in two regions. The krill Euphausia pacifica was the most prevalent invertebrate DNA detected in all three regions, while sergestid and mysid shrimp may also be important. High DNA read abundances from walleye pollock (Gadus chalcogrammus) and sablefish (Anoplopoma fimbria) were also recovered in one sample each, suggesting that juveniles of these semi-pelagic species may occasionally be targeted. In general, we observed heavily digested fecal material that drove substantial dissimilarities in taxonomic resolution between polymerase chain reaction-based and morphological analyses of the feces. Pacific herring and walleye pollock were the only prey species confirmed by both methods. Our results highlight that molecular and visual analyses of fecal samples provide a complementary approach to diet analysis, with each method providing unique insight into prey diversity.

Bidirectional Interactions With Humpback Whale Singer Using Concrete Sound Elements.

We describe an art-science project called "Feral Interactions-The Answer of the Humpback Whale" inspired by humpback whale songs and interactions between individuals based on mutual influences, learning process, or ranking in the dominance hierarchy. The aim was to build new sounds that can be used to initiate acoustic interactions with these whales, not in a one-way direction, as playbacks do, but in real interspecies exchanges. Thus, we investigated how the humpback whales generate sounds in order to better understand their abilities and limits. By carefully listening to their emitted vocalizations, we also describe their acoustic features and temporal structure, in a scientific way and also with a musical approach as it is done with musique concrète, in order to specify the types and the morphologies of whale sounds. The idea is to highlight the most precise information to generate our own sounds that will be suggested to the whales. Based on the approach developed in musique concrète, similarities with the sounds produced by bassoon were identified and then were processed to become "concrete sound elements." This analysis also brought us to design a new music interface that allows us to create adapted musical phrases in real-time. With this approach, interactions will be possible in both directions, from and to whales.

Using an omnidirectional video logger to observe the underwater life of marine animals: Humpback whale resting behaviour.

Animal-borne video loggers are powerful tools for investigating animal behaviour because they directly record immediate and extended peripheral animal activities; however, typical video loggers capture only a limited area on one side of an animal being monitored owing to their narrow field of view. Here, we investigated the resting behaviour of humpback whales using an animal-borne omnidirectional video camera combined with a behavioural data logger. In the video logger footage, two non-tagged resting individuals, which did not spread their flippers or move their flukes, were observed above a tagged animal, representing an apparent bout of group resting. During the video logger recording, the swim speed was relatively slow (0.75 m s-1), and the tagged animal made only a few strokes of very low amplitude during drift diving. We report the drift dives as resting behaviour specific to baleen whales as same as seals, sperm whales and loggerhead turtles. Overall, our study shows that an omnidirectional video logger is a valuable tool for interpreting animal ecology with improved accuracy owing to its ability to record a wide field of view.

Validation of presence-only models for conservation planning and the application to whales in a multiple-use marine park.

Identification of species' Biologically Important Areas (BIAs) is fundamental to conservation planning and species distribution models (SDMs) are a powerful tool commonly used to do this. Presence-only data are increasingly being used to develop SDMs to aid the conservation decision-making process. The application of presence-only SDMs for marine species' is particularly attractive due to often logistical and economic costs of obtaining systematic species' distribution data. However, robust model validation is important for conservation management applications that require accurate and reliable species' occurrence data (e.g., spatially explicit risk assessments). This is commonly done using a random subset of the data and less commonly with fully independent test data. Here, we apply a spatial block cross-validation (CV) approach to validate a MaxEnt presence-only model using independent presence/absence survey data for a highly mobile, marine species (humpback whale, Megaptera novaengliae) in the Great Barrier Reef (GBR). A MaxEnt model was developed using opportunistic whale sightings (2003-2007) and then used to identify areas differing in habitat suitability (low, medium, high) to conduct a systematic, line-transect, aerial survey (2012) and derive a density surface model. A spatial block CV buffering strategy was used to validate the MaxEnt model, using the opportunistic sightings as training data and independent aerial survey sightings data as test data. Moderate performance measures indicate MaxEnt was reliable in identifying the distribution patterns of a mobile whale species on their breeding ground, indicated by areas of high density aligned to areas of high habitat suitability. Furthermore, we demonstrate that MaxEnt models can be useful and cost-effective for designing a sampling scheme to undertake systematic surveys that significantly reduces sampling effort. In this study, higher quality information on whale reproductive class (calf vs. non-calf groups) was obtained that the presence-only data lacked, while sampling only 18% of the GBR World Heritage Area. The validation approach using fully independent data provides greater confidence in the MaxEnt model, which indicates significant overlap with the main breeding ground of humpback whales and the inner shipping route. This is important when evaluating presence-only models within certain conservation management applications, such as spatial risk assessments.

Ontogenetic niche shifts as a driver of seasonal migration.

Ontogenetic niche shifts have helped to understand population dynamics. Here we show that ontogenetic niche shifts also offer an explanation, complementary to traditional concepts, as to why certain species show seasonal migration. We describe how demographic processes (survival, reproduction and migration) and associated ecological requirements of species may change with ontogenetic stage (juvenile, adult) and across the migratory range (breeding, non-breeding). We apply this concept to widely different species (dark-bellied brent geese (Branta b. bernicla), humpback whales (Megaptera novaeangliae) and migratory Pacific salmon (Oncorhynchus gorbuscha) to check the generality of this hypothesis. Consistent with the idea that ontogenetic niche shifts are an important driver of seasonal migration, we find that growth and survival of juvenile life stages profit most from ecological conditions that are specific to breeding areas. We suggest that matrix population modelling techniques are promising to detect the importance of the ontogenetic niche shifts in maintaining migratory strategies. As a proof of concept, we applied a first analysis to resident, partial migratory and fully migratory populations of barnacle geese (Branta leucopsis). We argue that recognition of the costs and benefits of migration, and how these vary with life stages, is important to understand and conserve migration under global environmental change.

Corticosterone in central North Pacific male humpback whales (Megaptera novaeangliae): Pairing sighting histories with endocrine markers to assess stress.

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.

Vessel noise levels drive behavioural responses of humpback whales with implications for whale-watching.

Disturbance from whale-watching can cause significant behavioural changes with fitness consequences for targeted whale populations. However, the sensory stimuli triggering these responses are unknown, preventing effective mitigation. Here, we test the hypothesis that vessel noise level is a driver of disturbance, using humpback whales (Megaptera novaeangliae) as a model species. We conducted controlled exposure experiments (n = 42) on resting mother-calf pairs on a resting ground off Australia, by simulating whale-watch scenarios with a research vessel (range 100 m, speed 1.5 knts) playing back vessel noise at control/low (124/148 dB), medium (160 dB) or high (172 dB) low frequency-weighted source levels (re 1 µPa RMS@1 m). Compared to control/low treatments, during high noise playbacks the mother's proportion of time resting decreased by 30%, respiration rate doubled and swim speed increased by 37%. We therefore conclude that vessel noise is an adequate driver of behavioural disturbance in whales and that regulations to mitigate the impact of whale-watching should include noise emission standards.

Whale-watching is a multi-billion-dollar industry that is growing around the world. Typically, tour operators use boats to transport tourists into coastal waters to see groups of whales, dolphins or porpoises. There is, however, accumulating evidence that boat-based whale-watching negatively affects the way these animals behave and so many countries have put guidelines in place to mitigate activities that may disturb the animals. These guidelines generally stipulate the boat's angle of approach, how close the boat can get and the speed at which it can pass by the animals. In general, these guidelines are based on the assumption that the animals are disturbed by the closeness of the whale-watching boats. However, whales, dolphins and porpoises have very sensitive hearing, and only have a short range of vision underwater. Therefore, it seems plausible that the animals hear whale-watching boats long before they see them and so the loudness of underwater noise from the boats may be enough to disturb these animals' behaviour. To test this hypothesis, Sprogis et al. performed experiments where they simulated a whale-watching vessel approaching humpback whale mothers and calves who were resting off the northwest coast of Australia. A small motorised research boat travelling at a low speed passed different mother-calf pairs at a target distance of 100 meters, which is a common whale-watching distance guideline in many countries. The boat had an underwater speaker that played recordings of the boat noise at different volumes, while a drone with a video camera flew overhead to record the whales' behaviours in detail and to identify individual animals. These "controlled exposure experiments" showed that the quiet boat noise did not appear to disturb the mothers and calves. However, compared to when the quiet boat passed the animals the louder boat noise decreased how long the mother whale rested on the surface by 30%, made her swim 37% faster, and doubled the number of breaths she took per minute. If there are many disturbances from humans, then it can negatively impact the energy the mother and calf have available for nursing, fending off males and predators, and migrating back to their feeding ground nearer the Earth's poles. Based on these findings, it is shown that the loudness of the underwater noise from boats can explain why whales may be disturbed during whale-watching activities. To help reduce this disturbance, Sprogis et al. recommend that noise emission standards should be added to the current whale-watching regulations such that boats should be as quiet as possible and ideally around the volume of the ambient background noise. This would allow operators to approach the animals in a responsible, sustainable manner and offer tourists a view of undisturbed wildlife.