Individual-level assessments of wild animal health, vital rates, and foraging ecology are critical for understanding population-wide impacts of exposure to stressors. Large whales face multiple stressors, including, but not limited to, ocean noise, pollution, and ship strikes. Because baleen is a continuously growing keratinized structure, serial extraction, and quantification of hormones and stable isotopes along the length of baleen provide a historical record of whale physiology and foraging ecology. Furthermore, baleen analysis enables the investigation of dead specimens, even decades later, allowing comparisons between historic and modern populations. Here, we examined baleen of five sub-adult gray whales and observed distinct patterns of oscillations in δ15N values along the length of their baleen plates which enabled estimation of baleen growth rates and differentiation of isotopic niche widths of the whales during wintering and summer foraging. In contrast, no regular patterns were apparent in δ13C values. Prolonged elevation of cortisol in four individuals before death indicates that chronic stress may have impacted their health and survival. Triiodothyronine (T3) increased over months in the whales with unknown causes of death, simultaneous with elevations in cortisol, but both hormones remained stable in the one case of acute death attributed to killer whale predation. This parallel elevation of cortisol and T3 challenges the classic understanding of their interaction and might relate to increased energetic demands during exposure to stressors. Reproductive hormone profiles in subadults did not show cyclical trends, suggesting they had not yet reached sexual maturity. This study highlights the potential of baleen analysis to retrospectively assess gray whales\' physiological status, exposure to stressors, reproductive status, and foraging ecology in the months or years leading up to their death, which can be a useful tool for conservation diagnostics to mitigate unusual mortality events.

Filter-feeding has been present for hundreds of millions of years, independently evolving in aquatic vertebrates\' numerous times. Mysticete whales are a group of gigantic, marine filter-feeders that are defined by their fringed baleen and are divided into two groups: balaenids and rorquals. Recent studies have shown that balaenids likely feed using a self-cleaning, cross-flow filtration mechanism where food particles are collected and then swept to the esophagus for swallowing. However, it is unclear how filtering is achieved in the rorquals (Balaenopteridae). Lunging rorqual whales engulf enormous masses of both prey and water; the prey is then separated from the water through baleen plates lining the length of their upper jaw and positioned perpendicular to flow. Rorqual baleen is composed of both major (larger) and minor (smaller) keratin plates containing embedded fringe that extends into the whale\'s mouth, forming a filtering fringe. We used a multimodal approach, including microcomputed tomography (µCT) and scanning electron microscopy (SEM), to visualize and describe the variability in baleen anatomy across five species of rorqual whales, spanning two orders of magnitude in body length. For most morphological measurements, larger whales exhibited hypoallometry relative to body length. µCT and SEM revealed that the major and minor plates break away from the mineralized fringes at variable distances from the gums. We proposed a model for estimating the effective pore size to determine whether flow scales with body length or prey size across species. We found that pore size is likely not a proxy for prey size but instead, may reflect changes in resistance through the filter that affect fluid flow.

Quantification of contaminant concentrations in baleen whales is important for individual and population level health assessments but is difficult due to large migrations and infrequent resighings. The use of baleen allows for a multiyear retrospective analysis of contaminant concentrations without having to collect repeated samples from the same individual. Here we provide case studies of mercury analysis using cold vapor atomic absorption spectroscopy in three individual humpback whales (Megaptera novaeangliae), a 44.5-year-old female and two males aged ≥35 and 66 years, over approximately three years of baleen growth. Mercury concentrations in the female\'s baleen were consistently 2-3 times higher than in either male. Age did not affect mercury concentrations in baleen; the younger male had comparable levels to the older male. In the female, mercury concentrations in the baleen did not change markedly during pregnancy but mercury did spike during the first half of lactation. Stable isotope profiles suggest that diet likely drove the female\'s high mercury concentrations. In conclusion, variations in baleen mercury content can be highly individualistic. Future studies should compare sexes as well as different populations and species to determine how the concentrations of mercury and other contaminants vary by life history parameters and geography.

Isotopic analyses of the incrementally growing baleen in Mysticeti have been used to learn about their feeding and movement patterns. Using methods previously applied to Pacific minke whales, stable δ15N and δ13C isotope values were measured along the baleen plates of male and female minke whales from two locations in the Northeast Atlantic. The sample sizes used in this study are comparable to those previously used in the literature, and, although limited in size, the evidence suggests differences in isotopic signatures between whales caught at different locations. Both the δ15N and δ13C data suggest whales at the higher latitude site of Svalbard have a narrower diet than the whales from Lofoten/Vesterålen in Norway. Across all whales, the δ15N data indicate the whales primarily prey on fish for much of the year, only switching to zooplankton during the spring bloom. The δ13C data fail to confirm whether the whales migrate over long distances.

This study established the first baseline of changing elemental concentrations in bowhead whale baleen over time (1958-1999). From previously published stable isotope data, year, season (summer or winter), and location (Beaufort or Bering/Chukchi seas) were attributed to each sample. Thirteen elements (Al, Cd, Co, Cr, Cu, Fe, Hg, Mn, Ni, Pb, Se, V, Zn) in baleen from nine subsistence-harvested bowhead whales (n = 138) were detected. Al, Cu, and Fe were the highest concentrations while Cd and V were among the lowest. Our data supports absorption as the main route of exposure to environmental elements rather than biomagnification due to bowhead whales\' low trophic position. A linear mixed-effects model confirmed most elements\' concentrations increased with time, while location and sex were insignificant explanatory factors. These temporal fluctuations were most likely a product of environmental changes due to a warming climate and human activities.

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

Baleen whales (mysticetes) lack teeth as adults and instead filter feed using keratinous baleen plates. They do not echolocate with ultrasonic frequencies like toothed whales but are instead known for infrasonic acoustics. Both baleen and infrasonic hearing are separately considered key innovations linked to their gigantism, evolutionary success and ecological diversity. The earliest mysticetes had teeth, and the phylogenetic position of many so-called toothed mysticetes remains debated, including those belonging to the nominal taxonomic groups Llanocetidae, Mammalodontidae and Aetiocetidae. Here, we report a new stem mysticete, Borealodon osedax gen. et sp. nov., from the Oligocene of Washington State, USA. Borealodon preserves multi-cusped teeth with apical wear; microCT scans of the inner ear indicate that the minimum frequency hearing limit of Borealodon was similar to mammalodontids. Borealodon is not recovered within a monophyletic Mammalodontidae nor a monophyletic Aetiocetidae; instead, it represents an unnamed lineage of stem Mysticeti, adding to the diversity of stem mysticetes, especially across the Rupelian-Chattian boundary. Furthermore, the presence of a putative chemosynthetic bivalve along with Osedax, a bone-boring annelid, found in association with the type specimen of Borealodon, offer more insights into the evolution of deep-sea whale-fall communities.

Whales use baleen, a novel integumentary structure, to filter feed; filter feeding itself evolved at least five times in tetrapod history but demonstrably only once in mammals [1]. Living baleen whales (mysticetes) are born without teeth, but paleontological and embryological evidence demonstrate that they evolved from toothed ancestors that lacked baleen entirely [2]. The mechanisms driving the origin of filter feeding in tetrapods remain obscure. Here we report Maiabalaena nesbittae gen. et sp. nov., a new fossil whale from early Oligocene rocks of Washington State, USA, lacking evidence of both teeth and baleen. The holotype possesses a nearly complete skull with ear bones, both mandibles, and associated postcrania. Phylogenetic analysis shows Maiabalaena as crownward of all toothed mysticetes, demonstrating that tooth loss preceded the evolution of baleen. The functional transition from teeth to baleen in mysticetes has remained enigmatic because baleen decays rapidly and leaves osteological correlates with unclear homology; the oldest direct evidence for fossil baleen is ∼25 million years younger [3] than the oldest stem mysticetes (∼36 Ma). Previous hypotheses for the origin of baleen [4, 5] are inconsistent with the morphology and phylogenetic position of Maiabalaena. The absence of both teeth and baleen in Maiabalaena is consistent with recent evidence that the evolutionary loss of teeth and origin of baleen are decoupled evolutionary transformations, each with a separate morphological and genetic basis [2, 6]. Understanding these macroevolutionary patterns in baleen whales is akin to other macroevolutionary transformations in tetrapods such as scales to feathers in birds.

Whale baleen is a keratin-based biological material; it provides life-long (40-100 years) filter-feeding for baleen whales in place of teeth. This study reveals new aspects of the contribution of the baleen\'s hierarchical structure to its fracture toughness and connects it to the unique performance requirements, which require anisotropy of fracture resistance. Baleen plates are subjected to competing external effects of hydration and varying loading rates and demonstrate a high fracture toughness in transverse loading, which is the most important direction in the filtering function; in the longitudinal direction, the toughness is much lower since delamination and controlled flexure are expected and desirable. The compressive strength is also established and results support the fracture toughness measurements: it is also highly anisotropic, and exhibits a ductile-to-brittle transition with increasing strain rate in the dry condition, which is absent in the hydrated condition, conferring impact resistance to the baleen. Using 3D-printing prototypes that replicate the three principal structural features of the baleen plate (hollow medulla, mineralized tubules, and sandwich-tubular structure) are created, and the role of its structure in determining its mechanical behavior is demonstrated. These findings suggest new bioinspired engineering materials.

Male baleen whales have long been suspected to have annual cycles in testosterone, but due to difficulty in collecting endocrine samples, little direct evidence exists to confirm this hypothesis. Potential influences of stress or adrenal stress hormones (cortisol, corticosterone) on male reproduction have also been difficult to study. Baleen has recently been shown to accumulate steroid hormones during growth, such that a single baleen plate contains a continuous, multi-year retrospective record of the whale\'s endocrine history. As a preliminary investigation into potential testosterone cyclicity in male whales and influences of stress, we determined patterns in immunoreactive testosterone, two glucocorticoids (cortisol and corticosterone), and stable-isotope (SI) ratios, across the full length of baleen plates from a bowhead whale (Balaena mysticetus), a North Atlantic right whale (Eubalaena glacialis) and a blue whale (Balaenoptera musculus), all adult males. Baleen was subsampled at 2 cm (bowhead, right) or 1 cm (blue) intervals and hormones were extracted from baleen powder with methanol, followed by quantification of all three hormones using enzyme immunoassays validated for baleen extract of these species. Baleen of all three males contained regularly spaced peaks in testosterone content, with number and spacing of testosterone peaks corresponding well to SI data and to species-specific estimates of annual baleen growth rate. Cortisol and corticosterone exhibited some peaks that co-occurred with testosterone peaks, while other glucocorticoid peaks occurred independent of testosterone peaks. The right whale had unusually high glucocorticoids during a period with a known entanglement in fishing gear and a possible disease episode; in the subsequent year, testosterone was unusually low. Further study of baleen testosterone patterns in male whales could help clarify conservation- and management-related questions such as age of sexual maturity, location and season of breeding, and the potential effect of anthropogenic and natural stressors on male testosterone cycles.