Comparative embryology of Delphinapterus leucas (beluga whale), Balaena mysticetus (bowhead whale), and Stenella attenuata (pan-tropical spotted dolphin) (Cetacea: Mammalia).

Embryogenesis of cetaceans (whales, dolphins, porpoises) is best known in Stenella attenuata, the pan-tropical spotted dolphin, based on a remarkably complete and well-studied prenatal ontogenetic series. Our study expands understanding of cetacean embryology by adding two additional cetacean taxa: the beluga whale (Delphinapterus leucas, Odontoceti), and the bowhead whale (Balaena mysticetus, Mysticeti). We identify key features that characterize these taxa at specific stages and highlight heterochrony between the odontocetes and mysticetes. The toothed whales are more similar in developmental timing to each other than either is to Balaena. The two odontocete taxa, Stenella and Delphinapterus, share similar developmental trajectories while early Balaena specimens differ from the odontocetes. This developmental variation proves challenging to ascribe to the existing Carnegie staging system. Most notably, flippers, hindlimbs, and flukes all provide morphological traits for characterization within the Carnegie staging system. A presomitic Delphinapterus embryo is also described. This study applies the Carnegie staging system to two more cetacean taxa and forms a framework for future research on cetacean developmental genetics and the modeling of fetal growth.

Serological Detection of Marine Origin Brucella Exposure in Two Alaska Beluga Stocks.

Among emerging threats to the Arctic is the introduction, spread, or resurgence of disease. Marine brucellosis is an emerging disease concern among free-ranging cetaceans and is less well-studied than terrestrial forms. To investigate marine-origin Brucella sp. exposure in two beluga stocks in Alaska, USA, this study used serological status as well as real-time polymerase chain reaction (rtPCR) and bacterial culture. In total, 55 live-captured-released belugas were tested for Brucella exposure in Bristol Bay (2008-2016) and 112 (8 live-captured; 104 subsistence-harvested) whales were tested in the eastern Chukchi Sea (2007-2017). In total, 73% percent of Bristol Bay live captures, 50% of Chukchi Sea live captures, and 66% of Chukchi Sea harvested belugas were positive on serology. Only 10 of 69 seropositive belugas were rtPCR positive in at least one tissue. Only one seropositive animal was PCR positive in both the spleen and mesenteric lymph node. All animals tested were culture negative. The high prevalence of seropositivity detected suggests widespread exposure in both stocks, however, the low level of rtPCR and culture positive results suggests clinical brucellosis was not prevalent in the belugas surveyed. Continued detection of Brucella exposure supports the need for long-term monitoring of these and other beluga populations.

Olfactory epithelium and ontogeny of the nasal chambers in the bowhead whale (Balaena mysticetus).

In a species of baleen whale, we identify olfactory epithelium that suggests a functional sense of smell and document the ontogeny of the surrounding olfactory anatomy. Whales must surface to breathe, thereby providing an opportunity to detect airborne odorants. Although many toothed whales (odontocetes) lack olfactory anatomy, baleen whales (mysticetes) have retained theirs. Here, we investigate fetal and postnatal specimens of bowhead whales (Balaena mysticetus). Computed tomography (CT) reveals the presence of nasal passages and nasal chambers with simple ethmoturbinates through ontogeny. Additionally, we describe the dorsal nasal meatuses and olfactory bulb chambers. The cribriform plate has foramina that communicate with the nasal chambers. We show this anatomy within the context of the whole prenatal and postnatal skull. We document the tunnel for the ethmoidal nerve (ethmoid foramen) and the rostrolateral recess of the nasal chamber, which appears postnatally. Bilateral symmetry was apparent in the postnatal nasal chambers. No such symmetry was found prenatally, possibly due to tissue deformation. No nasal air sacs were found in fetal development. Olfactory epithelium, identified histologically, covers at least part of the ethmoturbinates. We identify olfactory epithelium using six explicit criteria of mammalian olfactory epithelium. Immunohistochemistry revealed the presence of olfactory marker protein (OMP), which is only found in mature olfactory sensory neurons. Although it seems that these neurons are scarce in bowhead whales compared to typical terrestrial mammals, our results suggest that bowhead whales have a functional sense of smell, which they may use to find prey.

A comparative analysis of cone photoreceptor morphology in bowhead and beluga whales.

The cetacean visual system is a product of selection pressures favoring underwater vision, yet relatively little is known about it across taxa. Previous studies report several mutations in the opsin genetic sequence in cetaceans, suggesting the evolutionary complete or partial loss of retinal cone photoreceptor function in mysticete and odontocete lineages, respectively. Despite this, limited anatomical evidence suggests cone structures are partially maintained but with absent outer and inner segments in the bowhead retina. The functional consequence and anatomical distributions associated with these unique cone morphologies remain unclear. The current study further investigates the morphology and distribution of cone photoreceptors in the bowhead whale and beluga retina and evaluates the potential functional capacity of these cells' alternative to photoreception. Refined histological and advanced microscopic techniques revealed two additional cone morphologies in the bowhead and beluga retina that have not been previously described. Two proteins involved in magnetosensation were present in these cone structures suggesting the possibility for an alternative functional role in responding to changes in geomagnetic fields. These findings highlight a revised understanding of the unique evolution of cone and gross retinal anatomy in cetaceans, and provide prefatory evidence of potential functional reassignment of these cells.

Group structure and kinship in beluga whale societies.

Evolutionary explanations for mammalian sociality typically center on inclusive-fitness benefits of associating and cooperating with close kin, or close maternal kin as in some whale societies, including killer and sperm whales. Their matrilineal structure has strongly influenced the thinking about social structure in less well-studied cetaceans, including beluga whales. In a cross-sectional study of group structure and kinship we found that belugas formed a limited number of distinct group types, consistently observed across populations and habitats. Certain behaviours were associated with group type, but group membership was often dynamic. MtDNA-microsatellite profiling combined with relatedness and network analysis revealed, contrary to predictions, that most social groupings were not predominantly organized around close maternal relatives. They comprised both kin and non-kin, many group members were paternal rather than maternal relatives, and unrelated adult males often traveled together. The evolutionary mechanisms that shape beluga societies are likely complex; fitness benefits may be achieved through reciprocity, mutualism and kin selection. At the largest scales these societies are communities comprising all ages and both sexes where multiple social learning pathways involving kin and non-kin can foster the emergence of cultures. We explore the implications of these findings for species management and the evolution of menopause.

Multi-year assessment (2006-2015) of persistent organic pollutant concentrations in blubber and muscle from Western Arctic bowhead whales (Balaena mysticetus), North Slope, Alaska.

Blubber and muscle were collected from male bowhead whales (n = 71) landed near Utqiagvik (Barrow), Alaska, between 2006 and 2015 and analyzed for lipid content and concentrations of persistent organic pollutants (POPs) in order to determine levels and trends over the collection period. Collection year was a significant predictor of blubber concentrations for most classes of POPs, while for a few classes, animal length (proxy for age) was also a significant predictor. This is the first report on levels of PBDEs in bowhead whales; concentrations of these compounds are low (≤55 ng/g wet weight). Blubber concentrations were lower than those reported in samples collected between 1992 and 2000, and many POP classes in blubber declined significantly between 2006 and 2015. Concentrations of POPs in bowhead whale tissues, which are subsistence foods for Native Alaskan communities, appear to be declining at rates comparable with previously reported temporal trends in Arctic biota.

Whale tear glands in the bowhead and the beluga whales: Source and function.

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.

The role of desmosomes in the ear plug formation in the bowhead whale (Balaena mysticetus).

The external acoustic meatus (EAM) of most baleen whales accumulates cellular debris annually in the lumen as whales age, forming a lamellated ear plug. The bowhead whale ear plug is formed from annually molting lining of the EAM as the entire epithelium releases at the level of the stratum basale during the spring migration. Epithelial regeneration is mostly completed by the fall migration, remaining intact for 6-7 months before being torn off the following spring. Desmosomes are integral to cell-cell adhesion with connecting desmosomal cadherins desmoglein (dsg) and desmocollin (dsc). Paraffin sections of the oral cavity and EAM lining of spring and fall adult bowhead whales, as well as the EAM of spring-caught juvenile, were immunohistochemically examined for the presence of these cadherins. In all fall specimens, both cadherins occurred in all layers except the superficial keratinous layer of the oral cavity. In spring, three different conditions existed: (a) oral cavity of spring-caught adults had reduced cadherins, with superficial fissuring in its keratinized layer and vacuolation in the upper stratum spinosum; (b) EAM of juvenile spring-caught whales displayed fissuring with accompanying reduction of both cadherins in its superficial lining; and (c) EAM lining of spring-caught adults displayed deep fissures, reduced cadherins, and absence of dsc1 in the fissuring zone. These results suggest that shedding of skin layers in mammals, whether normal molting, pathological, or the result of injury and wound repair all revolve around desmosome function. The specific role, structure, and location of these two cadherins need to be further addressed.

Intestinal polycyclic aromatic hydrocarbon-DNA adducts in a population of beluga whales with high levels of gastrointestinal cancers.

Carcinogenic polycyclic aromatic hydrocarbons (PAHs) were disposed directly into the Saguenay River of the St. Lawrence Estuary (SLE) by local aluminum smelters (Quebec, Canada) for 50 years (1926-1976). PAHs in the river sediments are likely etiologically related to gastrointestinal epithelial cancers observed in 7% of 156 mature (>19-year old) adult beluga found dead along the shorelines. Because DNA adduct formation provides a critical link between exposure and cancer induction, and because PAH-DNA adducts are chemically stable, we hypothesized that SLE beluga intestine would contain PAH-DNA adducts. Using an antiserum specific for DNA modified with several carcinogenic PAHs, we stained sections of paraffin-embedded intestine from 51 SLE beluga (0-63 years), 4 Cook Inlet (CI) Alaska beluga (0-26 years), and 20 beluga (0-46 years) living in Arctic areas (Eastern Beaufort Sea, Eastern Chukchi Sea, Point Lay Alaska) and aquaria, all with low PAH contamination. Stained sections showed nuclear light-to-dark pink color indicating the presence of PAH-DNA adducts concentrated in intestinal crypt epithelial lining cells. Scoring of whole tissue sections revealed higher values for the 51 SLE beluga, compared with the 20 Arctic and aquarium beluga (P = 0.003). The H-scoring system, applied to coded individual photomicrographs, confirmed that SLE beluga and CI beluga had levels of intestinal PAH-DNA adducts significantly higher than Arctic and aquarium beluga (P = 0.003 and 0.02, respectively). Furthermore, high levels of intestinal PAH-DNA adducts in four SLE beluga with gastrointestinal cancers, considered as a group, support a link of causality between PAH exposure and intestinal cancer in SLE beluga. Environ. Mol. Mutagen. 60:29-41, 2019. Published 2018. This article is a U.S. Government work and is in the public domain in the USA.

Structure of the external auditory meatus of the Bowhead whale (Balaena mysticetus) and its relation to their seasonal migration.

The external auditory meatus (EAM) in many species of mysticete whales is filled with a waxy ear plug. Though this lamellated structure is often used to age a whale, its formation and development remain undescribed. It is thought that growth layer groups (GLGs) are laid down annually, thereby increasing the size of this structure. Since some mysticete whales are migratory and many undergo molting, we hypothesized that the cyclical production of these GLGs may be related to these processes. The epithelia of both EAM and glove finger (a part of the tympanic membrane protruding into the EAM) of one juvenile and multiple adult bowhead whales from both fall (October: non-molting) and spring (May: molting) seasons were dissected and examined anatomically and histologically. These tissue samples were compared with the adult oral epithelia at the same time periods. These epithelia shared a similar basic broad structure, though there were differences in thickness and presence of intraepithelial structures. All epithelia in the October specimens were rich in both glycogen and lipid. The parakeratinized epithelium of the oral cavity in the juvenile and some May specimens shed via the production of several superficial epithelial fissures. Other adult May specimens exhibited deep epithelial fissures, reminiscent of pressure ulcers, which would cause the detachment of the entire epithelium from the dermis. We propose that sloughed epithelial lining is the source of the GLGs in the ear plug. Correlating a potential molting sequence with these observations explained the presence of epidermal glycogen, deep epidermal fissures and dermal glycolipid, and to some extent calls into question the origin and structure of the ear plug itself. Further morphological characterization of ear plugs in bowheads is needed to better understand cell origin and ear plug formation.

Migratory culture, population structure and stock identity in North Pacific beluga whales (Delphinapterus leucas).

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.

Validation of Growth Layer Group (GLG) depositional rate using daily incremental growth lines in the dentin of beluga (Delphinapterus leucas (Pallas, 1776)) teeth.

Counts of Growth Layer Groups (GLGs) in the dentin of marine mammal teeth are widely used as indicators of age. In most marine mammals, observations document that GLGs are deposited yearly, but in beluga whales, some studies have supported the view that two GLGs are deposited each year. Our understanding of beluga life-history differs substantially depending on assumptions regarding the timing of GLG deposition; therefore, resolving this issue has important considerations for population assessments. In this study, we used incremental lines that represent daily pulses of dentin mineralization to test the hypothesis that GLGs in beluga dentin are deposited on a yearly basis. Our estimate of the number of daily growth lines within one GLG is remarkably close to 365 days within error, supporting the hypothesis that GLGs are deposited annually in beluga. We show that measurement of daily growth increments can be used to validate the time represented by GLGs in beluga. Furthermore, we believe this methodology may have broader applications to age estimation in other taxa.

Habitat selection by two beluga whale populations in the Chukchi and Beaufort seas.

There has been extensive sea ice loss in the Chukchi and Beaufort seas where two beluga whale (Delphinapterus leucas) populations occur between July-November. Our goal was to develop population-specific beluga habitat selection models that quantify relative use of sea ice and bathymetric features related to oceanographic processes, which can provide context to the importance of changing sea ice conditions. We established habitat selection models that incorporated daily sea ice measures (sea ice concentration, proximity to ice edge and dense ice) and bathymetric features (slope, depth, proximity to the continental slope, Barrow Canyon, and shore) to establish quantitative estimates of habitat use for the Eastern Chukchi Sea ('Chukchi') and Eastern Beaufort Sea ('Beaufort') populations. We applied 'used v. available' resource selection functions to locations of 65 whales tagged from 1993-2012, revealing large variations in seasonal habitat selection that were distinct between sex and population groups. Chukchi whales of both sexes were predicted to use areas in close proximity to Barrow Canyon (typically <200 km) as well as the continental slope in summer, although deeper water and denser ice were stronger predictors for males than females. Habitat selection differed more between sexes for Beaufort belugas. Beaufort males selected higher ice concentrations (≥40%) than females (0-40%) in July-August. Proximity to shore (<200 km) strongly predicted summer habitat of Beaufort females, while distance to the ice edge was important for male habitat selection, especially during westward migration in September. Overall, our results indicate that sea ice variables were rarely the primary drivers of beluga summer-fall habitat selection. While diminished sea ice may indirectly affect belugas through changes in the ecosystem, associations with bathymetric features that affect prey availability seemed key to habitat selection during summer and fall. These results provide a benchmark by which to assess future changes in beluga habitat use of the Pacific Arctic.

Evolutionary aspects of the development of teeth and baleen in the bowhead whale.

In utero, baleen whales initiate the development of several dozens of teeth in upper and lower jaws. These tooth germs reach the bell stage and are sometimes mineralized, but toward the end of prenatal life they are resorbed and no trace remains after birth. Around the time that the germs disappear, the keratinous baleen plates start to form in the upper jaw, and these form the food-collecting mechanism. Baleen whale ancestors had two generations of teeth and never developed baleen, and the prenatal teeth of modern fetuses are usually interpreted as an evolutionary leftover. We investigated the development of teeth and baleen in bowhead whale fetuses using histological and immunohistochemical evidence. We found that upper and lower dentition initially follow similar developmental pathways. As development proceeds, upper and lower tooth germs diverge developmentally. Lower tooth germs differ along the length of the jaw, reminiscent of a heterodont dentition of cetacean ancestors, and lingual processes of the dental lamina represent initiation of tooth bud formation of replacement teeth. Upper tooth germs remain homodont and there is no evidence of a secondary dentition. After these germs disappear, the oral epithelium thickens to form the baleen plates, and the protein FGF-4 displays a signaling pattern reminiscent of baleen plates. In laboratory mammals, FGF-4 is not involved in the formation of hair or palatal rugae, but it is involved in tooth development. This leads us to propose that the signaling cascade that forms teeth in most mammals has been exapted to be involved in baleen plate ontogeny in mysticetes.

Decadal shifts in autumn migration timing by Pacific Arctic beluga whales are related to delayed annual sea ice formation.

Migrations are often influenced by seasonal environmental gradients that are increasingly being altered by climate change. The consequences of rapid changes in Arctic sea ice have the potential to affect migrations of a number of marine species whose timing is temporally matched to seasonal sea ice cover. This topic has not been investigated for Pacific Arctic beluga whales (Delphinapterus leucas) that follow matrilineally maintained autumn migrations in the waters around Alaska and Russia. For the sympatric Eastern Chukchi Sea ('Chukchi') and Eastern Beaufort Sea ('Beaufort') beluga populations, we examined changes in autumn migration timing as related to delayed regional sea ice freeze-up since the 1990s, using two independent data sources (satellite telemetry data and passive acoustics) for both populations. We compared dates of migration between 'early' (1993-2002) and 'late' (2004-2012) tagging periods. During the late tagging period, Chukchi belugas had significantly delayed migrations (by 2 to >4 weeks, depending on location) from the Beaufort and Chukchi seas. Spatial analyses also revealed that departure from Beaufort Sea foraging regions by Chukchi whales was postponed in the late period. Chukchi beluga autumn migration timing occurred significantly later as regional sea ice freeze-up timing became later in the Beaufort, Chukchi, and Bering seas. In contrast, Beaufort belugas did not shift migration timing between periods, nor was migration timing related to freeze-up timing, other than for southward migration at the Bering Strait. Passive acoustic data from 2008 to 2014 provided independent and supplementary support for delayed migration from the Beaufort Sea (4 day yr-1 ) by Chukchi belugas. Here, we report the first phenological study examining beluga whale migrations within the context of their rapidly transforming Pacific Arctic ecosystem, suggesting flexible responses that may enable their persistence yet also complicate predictions of how belugas may fare in the future.

Navigating under sea ice promotes rapid maturation of diving physiology and performance in beluga whales.

Little is known about the postnatal development of the physiological characteristics that support breath-hold in cetaceans, despite their need to swim and dive at birth. Arctic species have the additional demand of avoiding entrapment while navigating under sea ice, where breathing holes are patchily distributed and ephemeral. This is the first investigation of the ontogeny of the biochemistry of the locomotor muscle in a year-round Arctic-dwelling cetacean (beluga whale, Delphinapterus leucas). Compared with what we know about other cetaceans, belugas are born with high myoglobin content (1.56±0.02 g 100 g-1 wet muscle mass, N=2) that matures rapidly. Myoglobin increased by 452% during the first year after birth and achieved adult levels (6.91±0.35 g 100 g-1 wet muscle mass, N=9) by 14 months postpartum. Buffering capacity was 48.88±0.69 slykes (N=2) at birth; adult levels (84.31±1.38 slykes, N=9) were also achieved by 14 months postpartum. As the oxygen stores matured, calculated aerobic dive limit more than doubled over the first year of life, undoubtedly facilitating the movements of calves under sea ice. Nonetheless, small body size theoretically continues to constrain the diving ability of newly weaned 2 year olds, as they only had 74% and 69% of the aerobic breath-hold capacity of larger adult female and male counterparts. These assessments enhance our knowledge of the biology of cetaceans and provide insight into age-specific flexibility to alter underwater behaviors, as may be required with the ongoing alterations in the Arctic marine ecosystem associated with climate change and increased anthropogenic activities.

Severe Bone Loss as Part of the Life History Strategy of Bowhead Whales.

The evolution of baleen constituted a major evolutionary change that made it possible for baleen whales to reach enormous body sizes while filter feeding on tiny organisms and migrating over tremendous distances. Bowhead whales (Balaena mysticetus) live in the Arctic where the annual cycle of increasing and decreasing ice cover affects their habitat, prey, and migration. During the nursing period, bowheads grow rapidly; but between weaning and approximately year 5, bowhead whales display sustained baleen and head growth while limiting growth in the rest of their bodies. During this period, they withdraw resources from the skeleton, in particular the ribs, which may lose 40% of bone mass. Such dramatic changes in bones of immature mammals are rare, although fossil cetaceans between 40 and 50 million years ago show an array of rib specializations that include bone loss and are usually interpreted as related to buoyancy control.

Morphometric Correlates of the Ovary and Ovulatory Corpora in the Bowhead Whale, Balaena mysticetus.

Gross morphology and morphometry of the bowhead whale ovary, including ovulatory corpora, were investigated in 50 whales from the Chukchi and Beaufort seas off the coast of Alaska. Using the presence of ovarian corpora to define sexual maturity, 23 sexually immature whales (7.6-14.2 m total body length) and 27 sexually mature whales (14.2-17.7 m total body length) were identified. Ovary pair weights ranged from 0.38 to 2.45 kg and 2.92 to 12.02 kg for sexually immature and sexually mature whales, respectively. In sexually mature whales, corpora lutea (CLs) and/or large corpora albicantia (CAs) projected beyond ovary surfaces. CAs became increasingly less interruptive of the surface contour as they regressed, while remaining identifiable within transverse sections of the ovarian cortex. CLs formed large globular bodies, often with a central lumen, featuring golden parenchymas enfolded within radiating fibrous cords. CAs, sometimes vesicular, featured a dense fibrous core with outward fibrous projections through the former luteal tissue. CLs (never more than one per ovary pair) ranged from 6.7 to 15.0 cm in diameter in 13 whales. Fetuses were confirmed in nine of the 13 whales, with the associated CLs ranging from 8.3 to 15.0 cm in diameter. CLs from four whales where a fetus was not detected ranged from 6.7 to 10.6 cm in diameter. CA totals ranged from 0 to 22 for any single ovary, and from 1 to 41 for an ovary pair. CAs measured from 0.3 to 6.3 cm in diameter, and smaller corpora were more numerous, suggesting an accumulating record of ovulation. Neither the left nor the right ovary dominated in the production of corpora. Anat Rec, 299:769-797, 2016. © 2016 Wiley Periodicals, Inc.

Current Status of Development of Methods to Assess Effects of Cumulative or Aggregated Underwater Sounds on Marine Mammals.

There are no standards for assessment of the cumulative effects of underwater sound. Quantitative assessments typically consider a single source, whereas qualitative assessments may include multiple sources but rarely identify response variables. As a step toward understanding the cumulative effects of underwater sound, we assessed the aggregated sounds of multiple sources received by migrating bowhead whales (Balaena mysticetus). The quantitative method models the sound field from multiple sources and simulates movement of a population through it. The qualitative method uses experts to assess the responses of individuals and populations to sound sources and identify the potential mechanisms. These methods increase the transparency of assessments.