ABSTRACT
We assessed long-term changes in the feeding ecology and mercury (Hg) accumulation in Eastern High Arctic-Baffin Bay beluga (Delphinapterus leucas) using total Hg and stable isotope (δ13C, δ15N) assays in teeth samples from historical (1854-1905) and modern (1985-2000) populations. Mean δ13C values in teeth declined significantly over time, from -13.01 ± 0.55 historically to -14.41 ± 0.28 in 2000, while no consistent pattern was evident for δ15N due to high individual variability within each period. The temporal shift in isotopic niche is consistent with beluga feeding ecology changing in recent decades to a more pelagic and less isotopically diverse diet or an ecosystem wide change in isotope profiles. Mercury concentrations in modern beluga teeth were 3-5 times higher on average than those in historical beluga. These results are similar to the long-term trends of Hg and feeding ecology reported in other beluga populations and in other Arctic marine predators. Similar feeding ecology shifts across regions and species indicate a consistent increased pelagic diet response to climate change as the Arctic Ocean progressively warmed and lost sea ice. Previously, significant temporal Hg increase in beluga and other Arctic animals was attributed solely to direct inputs of anthropogenic Hg from long-range sources. Recent advances in understanding the Arctic marine Hg cycle suggest an additional, complementary possibilityâincreased inputs of terrestrial Hg of mixed anthropogenic-natural origin, mobilized from permafrost and other Arctic soils by climate warming. At present, it is not possible to assign relative importance to the two processes in explaining the rise of Hg concentrations in modern Arctic marine predators.
Subject(s)
Beluga Whale , Mercury , Water Pollutants, Chemical , Animals , Arctic Regions , Beluga Whale/physiology , Ecosystem , Environmental Monitoring , Mercury/analysis , Water Pollutants, Chemical/analysisABSTRACT
Cook Inlet, Alaska, is home to an endangered and declining population of 279 belugas (Delphinapterus leucas). Recovery efforts highlight a paucity of basic ecological knowledge, impeding the correct assessment of threats and the development of recovery actions. In particular, information on diet and foraging habitat is very limited for this population. Passive acoustic monitoring has proven to be an efficient approach to monitor beluga distribution and seasonal occurrence. Identifying acoustic foraging behavior could help address the current gap in information on diet and foraging habitat. To address this conservation challenge, eight belugas from a comparative, healthy population in Bristol Bay, Alaska, were instrumented with a multi-sensor tag (DTAG), a satellite tag, and a stomach temperature transmitter in August 2014 and May 2016. DTAG deployments provided 129.6 hours of data including foraging and social behavioral states. A total of 68 echolocation click trains ending in terminal buzzes were identified during successful prey chasing and capture, as well as during social interactions. Of these, 37 click trains were successfully processed to measure inter-click intervals (ICI) and ICI trend in their buzzing section. Terminal buzzes with short ICI (minimum ICI <8.98 ms) and consistently decreasing ICI trend (ICI increment range <1.49 ms) were exclusively associated with feeding behavior. This dual metric was applied to acoustic data from one acoustic mooring within the Cook Inlet beluga critical habitat as an example of the application of detecting feeding in long-term passive acoustic monitoring data. This approach allowed description of the relationship between beluga presence, feeding occurrence, and the timing of spawning runs by different species of anadromous fish. Results reflected a clear preference for the Susitna River delta during eulachon (Thaleichthys pacificus), Chinook (Oncorhynchus tshawytscha), pink (Oncorhynchus gorbuscha), and coho (Oncorhynchus kisutch) salmon spawning run periods, with increased feeding occurrence at the peak of the Chinook and pink salmon runs.
Subject(s)
Beluga Whale/physiology , Echolocation/physiology , Ecosystem , Feeding Behavior/physiology , Alaska , Animals , Female , MaleABSTRACT
Belugas (Delphinapterus leucas) and narwhals (Monodon monoceros) are highly social Arctic toothed whales with large vocal repertoires and similar acoustic profiles. Passive Acoustic Monitoring (PAM) that uses multiple hydrophones over large spatiotemporal scales has been a primary method to study their populations, particularly in response to rapid climate change and increasing underwater noise. This study marks the first acoustic comparison between wild belugas and narwhals from the same location and reveals that they can be acoustically differentiated and classified solely by echolocation clicks. Acoustic recordings were made in the pack ice of Baffin Bay, West Greenland, during 2013. Multivariate analyses and Random Forests classification models were applied to eighty-one single-species acoustic events comprised of numerous echolocation clicks. Results demonstrate a significant difference between species' acoustic parameters where beluga echolocation was distinguished by higher frequency content, evidenced by higher peak frequencies, center frequencies, and frequency minimums and maximums. Spectral peaks, troughs, and center frequencies for beluga clicks were generally > 60 kHz and narwhal clicks < 60 kHz with overlap between 40-60 kHz. Classification model predictive performance was strong with an overall correct classification rate of 97.5% for the best model. The most important predictors for species assignment were defined by peaks and notches in frequency spectra. Our results provide strong support for the use of echolocation in PAM efforts to differentiate belugas and narwhals acoustically.
Subject(s)
Beluga Whale/physiology , Echolocation/physiology , Vocalization, Animal/physiology , Whales/physiology , Acoustics , Animals , Greenland , Species SpecificityABSTRACT
Echolocation signals of wild beluga whales (Delphinapterus leucas) were recorded in 2013 using a vertical, linear 16-hydrophone array at two locations in the pack ice of Baffin Bay, West Greenland. Individual whales were localized for 4:42 minutes of 1:04 hours of recordings. Clicks centered on the recording equipment (i.e. on-axis clicks) were isolated to calculate sonar parameters. We report the first sonar beam estimate of in situ recordings of wild belugas with an average -3 dB asymmetrical vertical beam width of 5.4°, showing a wider ventral beam. This narrow beam width is consistent with estimates from captive belugas; however, our results indicate that beluga sonar beams may not be symmetrical and may differ in wild and captive contexts. The mean apparent source level for on-axis clicks was 212 dB pp re 1 µPa and whales were shown to vertically scan the array from 120 meters distance. Our findings support the hypothesis that highly directional sonar beams and high source levels are an evolutionary adaptation for Arctic odontocetes to reduce unwanted surface echoes from sea ice (i.e., acoustic clutter) and effectively navigate through leads in the pack ice (e.g., find breathing holes). These results provide the first baseline beluga sonar metrics from free-ranging animals using a hydrophone array and are important for acoustic programs throughout the Arctic, particularly for acoustic classification between belugas and narwhals (Monodon monoceros).
Subject(s)
Beluga Whale/physiology , Sound , Animals , Echolocation , Geography , GreenlandABSTRACT
Emergence of new technologies in remote sensing give scientists a new way to detect and monitor wildlife populations. In this study we assess the ability to detect and classify two emblematic Arctic cetaceans, the narwhal (Monodon monoceros) and beluga whale (Delphinapterus leucas), using very high-resolution (VHR) satellite imagery. We analyzed 12 VHR images acquired in August 2017 and 2019, collected by the WorldView-3 satellite, which has a maximum resolution of 0.31 m per pixel. The images covered Clearwater Fiord (138.8 km2), an area on eastern Baffin Island, Canada where belugas spend a large part of the summer, and Tremblay Sound (127.0 km2), a narrow water body located on the north shore of Baffin Island that is used by narwhals during the open water season. A total of 292 beluga whales and 109 narwhals were detected in the images. This study contributes to our understanding of Arctic cetacean distribution and highlights the capabilities of using satellite imagery to detect marine mammals.
Subject(s)
Animal Migration/physiology , Beluga Whale/physiology , Whales/physiology , Animals , Arctic Regions , Canada , SpacecraftABSTRACT
The Distributed Biological Observatory (DBO) was established to detect environmental changes in the Pacific Arctic by regular monitoring of biophysical responses in each of 8 DBO regions. Here we examine the occurrence of bowhead and beluga whale vocalizations in the western Beaufort Sea acquired by acoustic instruments deployed from September 2008-July 2014 and September 2016-October 2018 to examine inter-annual variability of these Arctic endemic species in DBO Region 6. Acoustic data were collected on an oceanographic mooring deployed in the Beaufort shelfbreak jet at ~71.4°N, 152.0°W. Spectrograms of acoustic data files were visually examined for the presence or absence of known signals of bowhead and beluga whales. Weekly averages of whale occurrence were compared with outputs of zooplankton, temperature and sea ice from the BIOMAS model to determine if any of these variables influenced whale occurrence. In addition, the dates of acoustic whale passage in the spring and fall were compared to annual sea ice melt-out and freeze-up dates to examine changes in phenology. Neither bowhead nor beluga whale migration times changed significantly in spring, but bowhead whales migrated significantly later in fall from 2008-2018. There were no clear relationships between bowhead whales and the environmental variables, suggesting that the DBO 6 region is a migratory corridor, but not a feeding hotspot, for this species. Surprisingly, beluga whale acoustic presence was related to zooplankton biomass near the mooring, but this is unlikely to be a direct relationship: there are likely interactions of environmental drivers that result in higher occurrence of both modeled zooplankton and belugas in the DBO 6 region. The environmental triggers that drive the migratory phenology of the two Arctic endemic cetacean species likely extend from Bering Sea transport of heat, nutrients and plankton through the Chukchi and into the Beaufort Sea.
Subject(s)
Acoustics , Beluga Whale/physiology , Bowhead Whale/physiology , Ecosystem , Animals , Arctic Regions , Humans , Ice CoverABSTRACT
Orbital glands are found in many tetrapod vertebrates, and are usually separate structures, consisting of individual glands lying in the eyelids and both canthi of the orbit. In cetaceans, however, the orbital glandular units are less distinct and have been described by numerous authors as a single, periorbital mass. There are few histochemical and immunhistochemical studies to date of these structures. In this study, we examined the orbital glandular region of both the bowhead whale (Balaena mysticetus: Mysticeti) and the beluga whale (Delphinapterus leucas: Odontoceti) using histological, histochemical, and immunohistochemical techniques. Histologically, in the bowhead, three glandular areas were noted (circumorbital, including Harderian and lacrimal poles), palpebral (midway in the lower eyelid), and rim (near the edge of the eyelid). In the beluga, there was only a large, continuous mass within the eyelid itself. Histochemical investigation suggests neither sexual dimorphism nor age-related differences, but both whales had two cell types freely intermingling with each other in all glandular masses. Large cells (cell type 1) were distended by four histochemically distinct intracellular secretory granules. Smaller cells (cell type 2) were not distended (fewer granules) and had two to three histochemically distinct intracellular secretory granules. The beluga orbital glands had additional lipid granules in cell type 1. Counterintuitively, both lipocalin and transferrin were localized to cell type 2 only. This intermingling of cell types is unusual for vertebrates in whom individual orbital glands usually have one cell type with one to two different secretory granules present. The heterogeneity of the orbital fluid produced by cetacean orbital glands implies a complex function, or series of functions, for these orbital glands and their role in producing the tear fluid.
Subject(s)
Beluga Whale/anatomy & histology , Beluga Whale/physiology , Bowhead Whale/anatomy & histology , Bowhead Whale/physiology , Animal Structures/anatomy & histology , Animals , Female , MaleABSTRACT
The effects of adapting sounds (pip trains or pure tones) on auditory evoked potentials (the rate following response, RFR) were investigated in a beluga whale. During RFR acquisition, adapting signals lasting 128 ms each were alternated with test signals lasting 16 ms each; the test signal levels varied randomly. Adapting signals were trains of cosine-enveloped tone pips or pure tones. Pip rate varied with the envelope cosine cycle maintained at 0.125 of pip intervals and the cosine rise-fall time maintained at 0.0625 of pip intervals. Adapting signals shifted the amplitude-level function upward compared to the baseline (no adapting signal) function. The higher the adapting signal level was, the bigger the shift in the amplitude-level function was. The slower the pips were in the adapting signal, the smaller the adaptation effect was. A train of pips with a 0.0625-ms rise-fall time and 125 dB SPL shifted the function by 35-40 dB, whereas a train of pips with a 1-ms rise-fall time or a pure tone with the same SPL shifted the function by approximately 15 dB. The difference between the "fast" and "slow" adapting signals is supposed to be associated with their abilities to stimulate the auditory system in odontocetes.
Subject(s)
Adaptation, Physiological/physiology , Beluga Whale/physiology , Evoked Potentials, Auditory/physiology , Hearing/physiology , Animals , Female , SoundABSTRACT
In a beluga whale, the positions of sound receiving areas on the head surface were determined by comparing the acoustic delays from different sound source positions. For this investigation, auditory evoked potentials (AEPs) in response to short tone pips were recorded. Latencies of the first AEP wave that presumably reflected the activity of the auditory nerve were measured at different sound source azimuths. For AEPs of equal amplitudes, the difference in AEP latencies was attributed to the difference in the acoustic delays. These delay differences were used to compute the azimuths of sound receiving points. Measurements were conducted at frequencies from 22.5 to 90 kHz in half-octave steps. At all stimulus frequencies, the receiving points were located 24-38 cm caudal of the melon tip, which is near a proximal part of the lower jaw. Thus, the results indicated the latero-mandibular acoustic window. Possible causes for not finding a lateral or ventro-mandibular window are discussed.
Subject(s)
Auditory Perception/physiology , Beluga Whale/physiology , Evoked Potentials, Auditory/physiology , Hearing/physiology , Acoustic Stimulation/methods , Acoustics , Animals , Auditory Threshold/physiology , Cochlear Nerve/physiology , Humans , SoundABSTRACT
The topic of individual differences in animal behavior has garnered a great deal of recent attention across many species, but questions remain concerning the degree to which behavioral differences vary over time or by age and sex. The present study focused on white whales (Delphinapterus leucas), a species in which a high degree of behavioral variability may be expected due to the fact that belugas are large-brained, long-lived, and highly social in nature. A suite of 23 behavioral measures related to boldness, playfulness, sociability, and other traits were assessed in 41 seaquarium-based belugas that were housed in mixed age/sex grouping. The goals were to assess consistency within individuals over time as well as variations by age and sex. Nineteen of the 23 measures showed significant within-subject consistency over time, suggesting that stable behavioral differences appear to exist in this species. However, very few measures showed significant correlations with each other, indicating that they could not be grouped into identifiable factors that comprised recognizable clusters. It is suggested, therefore, that individual differences are best viewed as a complex array of characteristics that depend on an individual's age and contextual variables that influence the degree to which specific sets of behaviors are expressed and co-vary.
Subject(s)
Behavior, Animal/physiology , Beluga Whale/psychology , Individuality , Personality/physiology , Animals , Beluga Whale/physiology , Female , MaleABSTRACT
Arctic marine ecosystems are currently undergoing rapid environmental changes. Over the past 20â years, individual growth rates of beluga whales (Delphinapterus leucas) have declined, which may be a response to climate change; however, the scarcity of physiological data makes it difficult to gauge the adaptive capacity and resilience of the species. We explored relationships between body condition and physiological parameters pertaining to oxygen (O2) storage capacity in 77 beluga whales in the eastern Beaufort Sea. Muscle myoglobin concentrations averaged 77.9â mgâ g-1, one of the highest values reported among mammals. Importantly, blood haematocrit, haemoglobin and muscle myoglobin concentrations correlated positively to indices of body condition, including maximum half-girth to length ratios. Thus, a whale with the lowest body condition index would have â¼27% lower blood (26.0 versus 35.7â mlâ kg-1) and 12% lower muscle (15.6 versus 17.7â mlâ kg-1) O2 stores than a whale of equivalent mass with the highest body condition index; with the conservative assumption that underwater O2 consumption rates are unaffected by body condition, this equates to a >3â min difference in maximal aerobic dive time between the two extremes (14.3 versus 17.4â min). Consequently, environmental changes that negatively impact body condition may hinder the ability of whales to reach preferred prey sources, evade predators and escape ice entrapments. The relationship between body condition and O2 storage capacity may represent a vicious cycle, in which environmental changes resulting in decreased body condition impair foraging, leading to further reductions in condition through diminished prey acquisition and/or increased foraging efforts.
Subject(s)
Beluga Whale/physiology , Muscles/chemistry , Oxygen/analysis , Oxygen/blood , Animals , Body Composition , Female , Hematocrit , Hemoglobins/analysis , Male , Myoglobin/analysis , Northwest TerritoriesABSTRACT
Although many animals, including odontocetes, exhibit interactions involving mouths (e.g., mouthing, nuzzling, biting), a limited number of animals display mouth-to-mouth social interactions that involve mutual coordination and collaboration. The purpose of this paper is to describe briefly a spontaneous and unexpected mouth-to-mouth social interaction between beluga calves in human care during their first year of life. Forty-seven independent events were identified after event sampling from more than 345 hr of observations of four mother-calf pairs and their companions. Unique aspects of this behavior included early emergence presumably without the benefit of a model and a preference for similar aged partners. Adult belugas did not display this social interaction. Based on its early emergence, the presence of similar-aged partner preferences, and affiliative cooperative contexts, this unusual mouth-to-mouth social interaction may play a significant role in beluga social and physical development. To evaluate this possibility, additional research is needed in which the complete topography, possible functions, and potential outcomes associated with this rare but conspicuous behavior exhibited by beluga calves are assessed.
Subject(s)
Aging , Behavior, Animal/physiology , Beluga Whale/physiology , Social Behavior , Animals , Animals, Zoo , Female , Male , MouthABSTRACT
The determination of body condition is a fundamental component of an evaluation during postmortem examinations of cetaceans. Three veterinarians experienced in beluga postmortem techniques subjectively evaluated 62 carcasses of beluga whales ( Delphinapterus leucas) in duplicate, using a visual analog scale. The result of evaluation and scoring was repeatable in cachectic animals and animals in very good condition but did not capture the continuum of body condition determinations between the obvious visual conditions.
Subject(s)
Beluga Whale/physiology , Body Composition/physiology , Animals , Conservation of Natural Resources , Endangered Species , Visual Analog ScaleABSTRACT
We measured respiratory flow, breath duration, and calculated tidal volume (VT) in nine belugas (Delphinapterus leucas, mean measured body mass: 628 ± 151 kg, n = 5) housed in managed care facilities. Both spontaneous (resting at station) and trained maximal respirations (chuffs) were measured. The mean (±s.d.) inspiratory VT for spontaneous breaths (16.7 ± 4.7 l, range: 7.5-18.7 l) was larger than those predicted based on respiratory scaling equations from terrestrial mammals and was 32 ± 10% of estimated total lung capacity (TLCest) based on an equation from static measurements made on a range of cetaceans and pinniped lungs, and 52 ± 18% of estimated vital capacities (VC, mean: 27.7 ± 8.9 l, range: 16.7-40.3 l) based on respiratory measurements obtained during trained maximal respirations. Expiratory flow (VËexp, spontaneous: 26.1 ± 5.5 l s-1, chuff: 66.8 ± 22.5 l s-1) was significantly higher as compared with inspiratory flow (VËinsp, spontaneous: 22.3 ± 4.6 l s-1, chuff: 30.1 ± 8.4 l s-1), and the maximal expiratory flow recorded was 212 l s-1. The breath duration was shorter for forced breaths (Expiration: 518 ± 101 ms; Inspiration: 905 ± 170 ms; Total: 1423 ± 227 ms) as compared with spontaneous breaths (Expiration: 995 ± 176 ms; Inspiration: 1098 ± 219 ms; Total: 2093 ± 302 ms). These data provide baseline estimates of the respiratory capacity of belugas.
Subject(s)
Beluga Whale/physiology , Respiration , Vital Capacity/physiology , Animals , Female , Male , Respiratory Function Tests/instrumentation , Respiratory Function Tests/methods , Tidal Volume/physiologyABSTRACT
The frequency resolving power of hearing (FRP) of the beluga whale Delphinapterus leucas was studied as dependent on influence of lasting low-intensity sounds (of the ultrasonic range from -20 to +10 dB). Testing of the spectrum ripple-phase reversal was used in conjunction with a noninvasive recording of auditory evoked potentials. FRP parameters were found to depend nonmonotonically on the intensity of the background noise. The resultant adaptation effects can be explained by the fact that, in response to the high-intensity signals, the auditory system sensitivity is reduced to the level optimal for analysis of these signals.
Subject(s)
Beluga Whale/physiology , Noise , Acoustic Stimulation , Animals , Evoked Potentials, Auditory/physiologyABSTRACT
The effects of prolonged sound stimuli (tone pip trains) on evoked potentials (the rate following response, RFR) were investigated in a beluga whale. The stimuli (rhythmic tone pips) were of 64 kHz frequency at levels from 80 to 140 dB re 1 µPa. During stimulation, every 1000 ms stimulus level either was kept constant (the steady-state stimulation) or changed up/down by 20 or 40 dB. With such stimulus presentation manner, RFR amplitude varied as follows. (i) After a stimulus level increase, the response amplitude increased quickly and then decayed slowly. The more the level increased, the higher the response amplitude increased. (ii) After a stimulus level decrease, the response amplitude was suppressed and then recovered slowly. The more the level decreased, the stronger was the response suppression. (iii) At the end of the 1000 ms window, the response amplitude approached, but did not reach, the amplitude characteristic of the steady-state stimulation. As a result, both after a sound level increase and decrease, the responses were almost stabilized during an analysis time as short as 1 s. This stabilization is attributed to an adaptation process. RFR decay after initial increase could be approximated by an exponent with a time constant of 59.4 ±1.8 (standard error) ms; RFR recovery after initial decrease could be approximated by an exponent with a time constant of 139.2 ±9.9 ms.
Subject(s)
Adaptation, Physiological , Auditory Perception/physiology , Beluga Whale/physiology , Brain/physiology , Acoustic Stimulation , Animals , Auditory Pathways/physiology , Evoked Potentials, Auditory , FemaleABSTRACT
The resolution of spectral patterns in adaptation background noise was investigated in a beluga whale, Delphinapterus leucas, using the evoked-potential technique. The resolution of spectral patterns was investigated using rippled-spectrum test stimuli of various levels and ripple densities and recording the rhythmic evoked responses (the rate following response, RFR) to ripple phase reversals. In baseline (no adaptation background noise) experiments, the highest RFR magnitude was observed at signal sound pressure levels (SPLs) of 100-110 dB re 1 µPa; at SPLs both below the optimum (down to 80 dB re 1 µPa) and above the optimum (up to 140 dB re 1 µPa), the RFR magnitude decreased. For high signal levels (above 110 dB re 1 µPa), low-level adaptation background noise (from -10 to -20 dB re signal level) increased RFR magnitude compared to baseline. This effect is considered to be a result of the optimization of the sensation level of the high-SPL signals due to decreasing hearing sensitivity caused by the adaptation background noise.
Subject(s)
Acoustic Stimulation/methods , Adaptation, Physiological/physiology , Auditory Threshold/physiology , Beluga Whale/physiology , Evoked Potentials, Auditory/physiology , Animals , FemaleABSTRACT
Documenting hearing abilities is vital to understanding a species' acoustic ecology and for predicting the impacts of increasing anthropogenic noise. Cetaceans use sound for essential biological functions such as foraging, navigation and communication; hearing is considered to be their primary sensory modality. Yet, we know little regarding the hearing of most, if not all, cetacean populations, which limits our understanding of their sensory ecology, population level variability and the potential impacts of increasing anthropogenic noise. We obtained audiograms (5.6-150â kHz) of 26 wild beluga whales to measure hearing thresholds during capture-release events in Bristol Bay, AK, USA, using auditory evoked potential methods. The goal was to establish the baseline population audiogram, incidences of hearing loss and general variability in wild beluga whales. In general, belugas showed sensitive hearing with low thresholds (<80â dB) from 16 to 100â kHz, and most individuals (76%) responded to at least 120â kHz. Despite belugas often showing sensitive hearing, thresholds were usually above or approached the low ambient noise levels measured in the area, suggesting that a quiet environment may be associated with hearing sensitivity and that hearing thresholds in the most sensitive animals may have been masked. Although this is just one wild population, the success of the method suggests that it should be applied to other populations and species to better assess potential differences. Bristol Bay beluga audiograms showed substantial (30-70â dB) variation among individuals; this variation increased at higher frequencies. Differences among individual belugas reflect that testing multiple individuals of a population is necessary to best describe maximum sensitivity and population variance. The results of this study quadruple the number of individual beluga whales for which audiograms have been conducted and provide the first auditory data for a population of healthy wild odontocetes.
Subject(s)
Auditory Threshold , Beluga Whale/physiology , Hearing , Alaska , Animals , Female , Hearing Tests/veterinary , Male , Reference ValuesABSTRACT
The annual return of beluga whales, Delphinapterus leucas, to traditional seasonal locations across the Arctic may involve migratory culture, while the convergence of discrete summering aggregations on common wintering grounds may facilitate outbreeding. Natal philopatry and cultural inheritance, however, has been difficult to assess as earlier studies were of too short a duration, while genetic analyses of breeding patterns, especially across the beluga's Pacific range, have been hampered by inadequate sampling and sparse information on wintering areas. Using a much expanded sample and genetic marker set comprising 1,647 whales, spanning more than two decades and encompassing all major coastal summering aggregations in the Pacific Ocean, we found evolutionary-level divergence among three geographic regions: the Gulf of Alaska, the Bering-Chukchi-Beaufort Seas, and the Sea of Okhotsk (Φst = 0.11-0.32, Rst = 0.09-0.13), and likely demographic independence of (Fst-mtDNA = 0.02-0.66), and in many cases limited gene flow (Fst-nDNA = 0.0-0.02; K = 5-6) among, summering groups within regions. Assignment tests identified few immigrants within summering aggregations, linked migrating groups to specific summering areas, and found that some migratory corridors comprise whales from multiple subpopulations (PBAYES = 0.31:0.69). Further, dispersal is male-biased and substantial numbers of closely related whales congregate together at coastal summering areas. Stable patterns of heterogeneity between areas and consistently high proportions (~20%) of close kin (including parent-offspring) sampled up to 20 years apart within areas (G = 0.2-2.9, p>0.5) is the first direct evidence of natal philopatry to migration destinations in belugas. Using recent satellite telemetry findings on belugas we found that the spatial proximity of winter ranges has a greater influence on the degree of both individual and genetic exchange than summer ranges (rwinter-Fst-mtDNA = 0.9, rsummer-Fst-nDNA = 0.1). These findings indicate widespread natal philopatry to summering aggregation and entire migratory circuits, and provide compelling evidence that migratory culture and kinship helps maintain demographically discrete beluga stocks that can overlap in time and space.
Subject(s)
Animal Migration/physiology , Beluga Whale/physiology , Animals , Female , Male , Pacific OceanABSTRACT
Asymmetrical intraguild predation (AIGP), which combines both predation and competition between predator species, is pervasive in nature with relative strengths varying by prey availability. But with species redistributions associated with climate change, the response by endemic predators within an AIGP context to changing biotic-abiotic conditions over time (i.e. seasonal and decadal) has yet to be quantified. Furthermore, little is known on AIGP dynamics in ecosystems undergoing rapid directional change such as the Arctic. Here, we investigate the flexibility of AIGP among two predators in the same trophic guild: beluga (Delphinapterus leucas) and Greenland halibut (Reinhardtius hippoglossoides), by season and over 30 years in Cumberland Sound-a system where forage fish capelin (Mallotus villosus) have recently become more available. Using stable isotopes, we illustrate different predator responses to temporal shifts in forage fish availability. On a seasonal cycle, beluga consumed less Greenland halibut and increased consumption of forage fish during summer, contrasting a constant consumption rate of forage fish by Greenland halibut year-round leading to decreased AIGP pressure between predators. Over a decadal scale (1982-2012), annual consumption of forage fish by beluga increased with a concomitant decline in the consumption of Greenland halibut, thereby indicating decreased AIGP pressure between predators in concordance with increased forage fish availability. The long-term changes of AIGP pressure between endemic predators illustrated here highlights climate-driven environmental alterations to interspecific intraguild interactions in the Arctic.