Mechanical properties of the California sea lion (Zalophus californianus) and northern elephant seal (Mirounga angustirostris) lower jaws explain trophic plasticity.

The fossil record of pinnipeds documents a suite of morphological changes that facilitate their ecological transition from a terrestrial to an aquatic lifestyle. Among these is the loss of the tribosphenic molar and the behavior typically associated with it in mammals: mastication. Instead, modern pinnipeds exhibit a broad range of feeding strategies that facilitate their distinct aquatic ecologies. Here, we examine the feeding morphology of two species of pinnipeds with disparate feeding ecologies: Zalophus californianus, a specialized raptorial biter, and Mirounga angustirostris, a suction specialist. Specifically, we test whether the morphology of the lower jaws facilitates trophic plasticity in feeding for either of these species. We used finite element analysis (FEA) to simulate the stresses during the opening and closing of the lower jaws in these species to explore the mechanical limits of their feeding ecology. Our simulations demonstrate that both jaws are highly resistant to the tensile stresses experienced during feeding. The lower jaws of Z. californianus experienced the maximum stress at the articular condyle and the base of the coronoid process. The lower jaws of M. angustirostris experienced the maximum stress at the angular process and were more evenly distributed throughout the body of the mandible. Surprisingly, the lower jaws of M. angustirostris were even more resistant to the stresses experienced during feeding than those of Z. californianus. Thus, we conclude that the superlative trophic plasticity of Z. californianus is driven by other factors unrelated to the mandible's tensile resistance to stress during feeding.

Lateral palatal foramina do not indicate baleen in fossil whales.

Today's mysticetes filter-feed using baleen, a novel integumentary structure with no apparent homolog in any living mammal. The origins of filter-feeding and baleen can be informed by the fossil record, including rare instances of soft tissue preservation of baleen and also by potential osteological correlates of baleen. Lateral palatal foramina on the roof of the mouth have been proposed as potential osteological correlates of baleen and their presence in some tooth-bearing stem mysticetes has led to the hypothesis that these early mysticetes possessed both teeth and incipient baleen. Here, we test this hypothesis by examining lateral palatal foramina in both filter-feeding and non-filter-feeding cetaceans, including crown and stem odontocetes and in stem cetaceans (or archaeocetes). We also confirm the presence of lateral palatal foramina in 61 species of terrestrial artiodactyls. CT scanning demonstrates consistent internal morphology across all observed taxa, suggesting that the lateral palatal foramina observed in extant mysticetes are homologous to those of terrestrial artiodactyls. The presence of lateral palatal foramina in terrestrial artiodactyls and non-filter-feeding whales (odontocetes and archaeocetes) suggests that these structures are not unique predictors for the presence of baleen in fossil whales; instead, these structures are more probably associated with gingiva or other oral tissue.

Puncture performance tests reveal distinct feeding modes in pinniped teeth.

Marine mammals have undergone a dramatic series of morphological transformations throughout their evolutionary history that facilitated their ecological transition to life in the water. Pinnipeds are a diverse clade of marine mammals that evolved from terrestrial carnivorans in the Oligocene (∼27 million years ago). However, pinnipeds have secondarily lost the dental innovations emblematic of mammalian and carnivoran feeding, such as a talonid basin or shearing carnassials. Modern pinnipeds do not masticate their prey, but can reduce prey size through chopping behavior. Typically, small prey are swallowed whole. Nevertheless, pinnipeds display a wide breadth of morphology of the post-canine teeth. We investigated the relationship between dental morphology and pinniped feeding by measuring the puncture performance of the cheek-teeth of seven extant pinniped genera. Puncture performance was measured as the maximum force and the maximum energy required to puncture a standardized prey item (Loligo sp.). We report significant differences in the puncture performance values across the seven genera, and identify three distinct categories based on cheek-teeth morphology and puncture performance: effective, ineffective and moderate puncturers. In addition, we measured the overall complexity of the tooth row using two different metrics, orientation patch count rotated (OPCR) and relief index (RFI). Neither metric of complexity predicted puncture performance. Finally, we discuss these results in the broader context of known pinniped feeding strategies and lay the groundwork for subsequent efforts to explore the ecological variation of specific dental morphologies.

Morphological variation of the relictual alveolar structures in the mandibles of baleen whales.

Living baleen whales (mysticetes) are bulk filter feeders that use keratinous baleen plates to filter food from prey laden water. Extant mysticetes are born entirely edentulous, though they possess tooth buds early in ontogeny, a trait inherited from toothed ancestors. The mandibles of extant baleen whales have neither teeth nor baleen; teeth are resorbed in utero and baleen grows only on the palate. The mandibles of extant baleen whales also preserve a series of foramina and associated sulci that collectively form an elongated trough, called the alveolar groove. Despite this name, it remains unclear if the alveolar groove of edentulous mysticetes and the dental structures of toothed mammals are homologous. Here, we describe and quantify the anatomical diversity of these structures across extant mysticetes and compare their variable morphologies across living taxonomic groups (i.e., Balaenidae, Neobalaenidae, Eschrichtiidae, and Balaenopteridae). Although we found broad variability across taxonomic groups for the alveolar groove length, occupying approximately 60-80 percent of the mandible's total curvilinear length (CLL) across all taxa, the relictual alveolar foramen showed distinct patterns, ranging between 15-25% CLL in balaenids, while ranging between 3-12% CLL in balaenopterids. This variability and the morphological patterning along the body of the mandible is consistent with the hypothesis that the foramina underlying the alveolar groove reflect relictual alveoli. These findings also lay the groundwork for future histological studies to examine the contents of these foramina and clarify their potential role in the feeding process.

What are the limits on whale ear bone size? Non-isometric scaling of the cetacean bulla.

The history of cetaceans demonstrates dramatic macroevolutionary changes that have aided their transformation from terrestrial to obligate aquatic mammals. Their fossil record shows extensive anatomical modifications that facilitate life in a marine environment. To better understand the constraints on this transition, we examined the physical dimensions of the bony auditory complex, in relation to body size, for both living and extinct cetaceans. We compared the dimensions of the tympanic bulla, a conch-shaped ear bone unique to cetaceans, with bizygomatic width-a proxy for cetacean body size. Our results demonstrate that cetacean ears scale non-isometrically with body size, with about 70% of variation explained by increases in bizygomatic width. Our results, which encompass the breadth of the whale fossil record, size diversity, and taxonomic distribution, suggest that functional auditory capacity is constrained by congruent factors related to cranial morphology, as opposed to allometrically scaling with body size.

Norrisanima miocaena, a new generic name and redescription of a stem balaenopteroid mysticete (Mammalia, Cetacea) from the Miocene of California.

Rorqual whales are among the most species rich group of baleen whales (or mysticetes) alive today, yet the monophyly of the traditional grouping (i.e., Balaenopteridae) remains unclear. Additionally, many fossil mysticetes putatively assigned to either Balaenopteridae or Balaenopteroidea may actually belong to stem lineages, although many of these fossil taxa suffer from inadequate descriptions of fragmentary skeletal material. Here we provide a redescription of the holotype of Megaptera miocaena, a fossil balaenopteroid from the Monterey Formation of California, which consists of a partial cranium, a fragment of the rostrum, a single vertebra, and both tympanoperiotics. Kellogg (1922) assigned the type specimen to the genus Megaptera Gray (1846), on the basis of its broad similarities to distinctive traits in the cranium of extant humpback whales (Megaptera novaeangliae (Borowski, 1781)). Subsequent phylogenetic analyses have found these two species as sister taxa in morphological datasets alone; the most recent systematic analyses using both molecular and morphological data sets place Megaptera miocaena as a stem balaenopteroid unrelated to humpback whales. Here, we redescribe the type specimen of Megaptera miocaena in the context of other fossil balaenopteroids discovered nearly a century since Kellogg's original description and provide a morphological basis for discriminating it from Megaptera novaeangliae. We also provide a new generic name and recombine the taxon as Norrisanima miocaena, gen. nov., to reflect its phylogenetic position outside of crown Balaenopteroidea, unrelated to extant Megaptera. Lastly, we refine the stratigraphic age of Norrisanima miocaena, based on associated microfossils to a Tortonian age (7.6-7.3 Ma), which carries implications for understanding the origin of key features associated with feeding and body size evolution in this group of whales.

Borealodon osedax, a new stem mysticete (Mammalia, Cetacea) from the Oligocene of Washington State and its implications for fossil whale-fall communities.

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.

Tooth Loss Precedes the Origin of Baleen in Whales.

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.

Salishicetus meadi, a new aetiocetid from the late Oligocene of Washington State and implications for feeding transitions in early mysticete evolution.

Living baleen whales, or Mysticeti, lack teeth and instead feed using keratinous baleen plates to sieve prey-laden water. This feeding strategy is profoundly different from that of their toothed ancestors, which processed prey using the differentiated dentition characteristic of mammals. The fossil record of mysticetes reveals stem members that include extinct taxa with dentition, illuminating the morphological states that preceded the loss of teeth and the subsequent origin of baleen. The relationships among stem mysticetes, including putative clades such as Mammalodontidae and Aetiocetidae, remain debatable. Aetiocetids are among the more species-rich clade of stem mysticetes, and known only from fossil localities along the North Pacific coastline. Here, we report a new aetiocetid, Salishicetus meadi gen. et sp. nov, from the late Oligocene of Washington State, USA. Salishicetus preserves a near-complete lower dentition with extensive occlusal wear, indicating that it processed prey using shearing cheek teeth in the same way as its stem cetacean ancestors. Using a matrix with all known species of aetiocetids, we recover a monophyletic Aetiocetidae, crownward of a basal clade of Mammalodontidae. The description of Salishicetus resolves phylogenetic relationships among aetiocetids, which provides a basis for reconstructing ancestral feeding morphology along the stem leading to crown Mysticeti.

Diversity of late Neogene Monachinae (Carnivora, Phocidae) from the North Atlantic, with the description of two new species.

While the diversity of 'southern seals', or Monachinae, in the North Atlantic realm is currently limited to the Mediterranean monk seal, Monachus monachus, their diversity was much higher during the late Miocene and Pliocene. Although the fossil record of Monachinae from the North Atlantic is mainly composed of isolated specimens, many taxa have been erected on the basis of fragmentary and incomparable specimens. The humerus is commonly considered the most diagnostic postcranial bone. The research presented in this study limits the selection of type specimens for different fossil Monachinae to humeri and questions fossil taxa that have other types of bones as type specimens, such as for Terranectes parvus. In addition, it is essential that the humeri selected as type specimens are (almost) complete. This questions the validity of partial humeri selected as type specimens, such as for Terranectes magnus. This study revises Callophoca obscura, Homiphoca capensis and Pliophoca etrusca, all purportedly known from the Lee Creek Mine, Aurora, North Carolina, in addition to their respective type localities in Belgium, South Africa and Italy, respectively. C. obscura is retained as a monachine seal taxon that lived both on the east coast of North America and in the North Sea Basin. However, H. capensis from North America cannot be identified beyond the genus level, and specimens previously assigned to Pl. etrusca from North America clearly belong to different taxa. Indeed, we also present new material and describe two new genera of late Miocene and Pliocene Monachinae from the east coast of North America: Auroraphoca atlantica nov. gen. et nov. sp., and Virginiaphoca magurai nov. gen. et nov. sp. This suggests less faunal interchange of late Neogene Monachinae between the east and west coasts of the North Atlantic than previously expected.

Alveoli, teeth, and tooth loss: Understanding the homology of internal mandibular structures in mysticete cetaceans.

The evolution of filter feeding in baleen whales (Mysticeti) facilitated a wide range of ecological diversity and extreme gigantism. The innovation of filter feeding evolved in a shift from a mineralized upper and lower dentition in stem mysticetes to keratinous baleen plates that hang only from the roof of the mouth in extant species, which are all edentulous as adults. While all extant mysticetes are born with a mandible lacking a specialized feeding structure (i.e., baleen), the bony surface retains small foramina with elongated sulci that often merge together in what has been termed the alveolar gutter. Because mysticete embryos develop tooth buds that resorb in utero, these foramina have been interpreted as homologous to tooth alveoli in other mammals. Here, we test this homology by creating 3D models of the internal mandibular morphology from terrestrial artiodactyls and fossil and extant cetaceans, including stem cetaceans, odontocetes and mysticetes. We demonstrate that dorsal foramina on the mandible communicate with the mandibular canal via smaller canals, which we explain within the context of known mechanical models of bone resorption. We suggest that these dorsal foramina represent distinct branches of the inferior alveolar nerve (or artery), rather than alveoli homologous with those of other mammals. As a functional explanation, we propose that these branches provide sensation to the dorsal margin of the mandible to facilitate placement and occlusion of the baleen plates during filer feeding.