The origin of baleen in mysticete whales represents a major transition in the phylogenetic history of Cetacea. This key specialization, a keratinous sieve that enables filter-feeding, permitted exploitation of a new ecological niche and heralded the evolution of modern baleen-bearing whales, the largest animals on Earth. To date, all formally described mysticete fossils conform to two types: toothed species from Oligocene-age rocks ( approximately 24 to 34 million years old) and toothless species that presumably utilized baleen to feed (Recent to approximately 30 million years old). Here, we show that several Oligocene toothed mysticetes have nutrient foramina and associated sulci on the lateral portions of their palates, homologous structures in extant mysticetes house vessels that nourish baleen. The simultaneous occurrence of teeth and nutrient foramina implies that both teeth and baleen were present in these early mysticetes. Phylogenetic analyses of a supermatrix that includes extinct taxa and new data for 11 nuclear genes consistently resolve relationships at the base of Mysticeti. The combined data set of 27,340 characters supports a stepwise transition from a toothed ancestor, to a mosaic intermediate with both teeth and baleen, to modern baleen whales that lack an adult dentition but retain developmental and genetic evidence of their ancestral toothed heritage. Comparative sequence data for ENAM (enamelin) and AMBN (ameloblastin) indicate that enamel-specific loci are present in Mysticeti but have degraded to pseudogenes in this group. The dramatic transformation in mysticete feeding anatomy documents an apparently rare, stepwise mode of evolution in which a composite phenotype bridged the gap between primitive and derived morphologies; a combination of fossil and molecular evidence provides a multifaceted record of this macroevolutionary pattern.

The extinct edentulous mysticete family Cetotheriidae historically has been viewed as a notoriously paraphyletic group, and only recently have rigorous studies been executed to rectify this issue. These problems do not necessarily just stem from lack of phylogenetic analyses, but are in part because of a general lack of complete specimens, poor descriptions of taxa, and long-lived taxonomic instability issues. The fossil mysticete genus Herpetocetus is a poster child of these problems as it is primarily only known from a few relatively incomplete and poorly described specimens. A new species of Herpetocetus from the upper Pliocene of California, Herpetocetus morrowi sp. nov., provides an archetypal model for the genus based on a multitude of well-preserved specimens. These specimens reveal a diminutive mysticete characterized by an elongate rostrum and roughly quadrate cranium. A mosaic of primitive and derived features preserved in this new species underscores its potential value in helping to resolve a number of taxonomic and phylogenetic problems. The occurrence of specimens assignable to juvenile through to mature adult individuals provides a basis for investigating ontogenetic changes. Functional analysis of the unusual craniomandibular anatomy of H. morrowi suggests a limited degree of mandibular gape and an enhanced capacity for longitudinal rotation of the dentary, features that support a hypothesis of suction feeding convergent with that of living grey whales. A phylogenetic analysis provides support for recognition of a redefined and monophyletic Cetotheriidae and Herpetocetinae, and also serves as a basis for evaluating the recent proposal that the pygmy right whale (Caperea marginata) is a living cetothere. Morphological features of Herpetocetus morrowi, including features of the cranium and petrosal, suggest that a number of the purported synapomorphies supporting a Caperea−cetothere grouping are either symplesiomorphies, nonhomologous features, or are highly variable.

BackgroundAnatomical comparisons of the ear region of baleen whales (Mysticeti) are provided through detailed osteological descriptions and high-resolution photographs of the petrotympanic complex (tympanic bulla and petrosal bone) of all extant species of mysticete cetaceans. Salient morphological features are illustrated and identified, including overall shape of the bulla, size of the conical process of the bulla, morphology of the promontorium, and the size and shape of the anterior process of the petrosal. We place our comparative osteological observations into a phylogenetic context in order to initiate an exploration into petrotympanic evolution within Mysticeti.Principal FindingsThe morphology of the petrotympanic complex is diagnostic for individual species of baleen whale (e.g., sigmoid and conical processes positioned at midline of bulla in Balaenoptera musculus; confluence of fenestra cochleae and perilymphatic foramen in Eschrichtius robustus), and several mysticete clades are united by derived characteristics. Balaenids and neobalaenids share derived features of the bulla, such as a rhomboid shape and a reduced anterior lobe (swelling) in ventral aspect, and eschrichtiids share derived morphologies of the petrosal with balaenopterids, including loss of a medial promontory groove and dorsomedial elongation of the promontorium. Monophyly of Balaenoidea (Balaenidae and Neobalaenidae) and Balaenopteroidea (Balaenopteridae and Eschrichtiidae) was recovered in phylogenetic analyses utilizing data exclusively from the petrotympanic complex.SignificanceThis study fills a major gap in our knowledge of the complex structures of the mysticete petrotympanic complex, which is an important anatomical region for the interpretation of the evolutionary history of mammals. In addition, we introduce a novel body of phylogenetically informative characters from the ear region of mysticetes. Our detailed anatomical descriptions, illustrations, and comparisons provide valuable data for current and future studies on the phylogenetic relationships, evolution, and auditory physiology of mysticetes and other cetaceans throughout Earth's history.

Toothed mysticetes of the family Aetiocetidae from Oligocene rocks of the North Pacific play a key role in interpretations of cetacean evolution because they are transitional in grade between dorudontine archaeocetes and edentulous mysticetes. The holotype skull of Aetiocetus weltoni from the late Oligocene (28–24 Ma) of Oregon, USA, has been further prepared, revealing additional morphological features of the basicranium, rostrum and dentary that have important implications for mysticete evolution and functional anatomy. The palate of Aetiocetus weltoni preserves diminutive lateral palatal foramina and associated delicate sulci which appear to be homologous with the prominent palatal foramina and sulci that occur along the lateral portion of the palate in extant mysticetes. In modern baleen whales these foramina allow passage of branches of the superior alveolar artery, which supplies blood to the epithelia of the developing baleen racks. As homologous structures, the lateral palatal foramina of A. weltoni suggest that baleen was present in this Oligocene toothed mysticete. Cladistic analysis of 46 cranial and dental characters supports monophyly of the Aetiocetidae, with toothed mysticetes Janjucetus and Mammalodon positioned as successive sister taxa. Morawanacetus is the earliest diverging aetiocetid with Chonecetus as sister taxon to Aetiocetus species. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 308–352.

The earliest dispersal of humans into North America is a contentious subject, and proposed early sites are required to meet the following criteria for acceptance: (1) archaeological evidence is found in a clearly defined and undisturbed geologic context; (2) age is determined by reliable radiometric dating; (3) multiple lines of evidence from interdisciplinary studies provide consistent results; and (4) unquestionable artefacts are found in primary context. Here we describe the Cerutti Mastodon (CM) site, an archaeological site from the early late Pleistocene epoch, where in situ hammerstones and stone anvils occur in spatio-temporal association with fragmentary remains of a single mastodon (Mammut americanum). The CM site contains spiral-fractured bone and molar fragments, indicating that breakage occured while fresh. Several of these fragments also preserve evidence of percussion. The occurrence and distribution of bone, molar and stone refits suggest that breakage occurred at the site of burial. Five large cobbles (hammerstones and anvils) in the CM bone bed display use-wear and impact marks, and are hydraulically anomalous relative to the low-energy context of the enclosing sandy silt stratum. Th/U radiometric analysis of multiple bone specimens using diffusion-adsorption-decay dating models indicates a burial date of 130.7 ± 9.4 thousand years ago. These findings confirm the presence of an unidentified species of Homo at the CM site during the last interglacial period (MIS 5e; early late Pleistocene), indicating that humans with manual dexterity and the experiential knowledge to use hammerstones and anvils processed mastodon limb bones for marrow extraction and/or raw material for tool production. Systematic proboscidean bone reduction, evident at the CM site, fits within a broader pattern of Palaeolithic bone percussion technology in Africa, Eurasia and North America. The CM site is, to our knowledge, the oldest in situ, well-documented archaeological site in North America and, as such, substantially revises the timing of arrival of Homo into the Americas.

Balaenidae (right whales) are large, critically endangered baleen whales represented by four living species. The evolutionary relationships of balaenids are poorly known, with the number of genera, relationships to fossil taxa, and position within Mysticeti in contention. This study employs a comprehensive set of morphological characters to address aspects of balaenid phylogeny. A sister‐group relationship between neobalaenids and balaenids is strongly supported, although this conflicts with molecular evidence, which may be an artifact of long‐branch attraction (LBA). Monophyly of Balaenidae is supported, and three major clades are recognized: (1) extinct genus Balaenula, (2) extant and extinct species of the genus Eubalaena, and (3) extant and extinct species of the genus Balaena plus the extinct taxon, Balaenella. The relationships of these clades to one another, as well as to the early Miocene stem balaenid, Morenocetus parvus, remain unresolved. Pliocene taxa, Balaenula astensis and Balaenula balaenopsis, form a clade that is the sister group to the Japanese Pliocene Balaenula sp. Eubalaena glacialis and Pliocene Eubalaena belgica, are in an unresolved polytomy with a clade including E. japonica and E. australis. Extant and fossil species of Balaena form a monophyletic group that is sister group to the Dutch Pliocene Balaenella, although phylogenetic relationships within Balaena remain unresolved.

The origin of baleen and filter feeding in mysticete cetaceans occurred sometime between approximately 34 and 24 million years ago and represents a major macroevolutionary shift in cetacean morphology (teeth to baleen) and ecology (raptorial to filter feeding). We explore this dramatic change in feeding strategy by employing a diversity of tools and approaches: morphology, molecules, development, and stable isotopes from the geological record. Adaptations for raptorial feeding in extinct toothed mysticetes provide the phylogenetic context for evaluating morphological apomorphies preserved in the skeletons of stem and crown edentulous mysticetes. In this light, the presence of novel vascular structures on the palates of certain Oligocene toothed mysticetes is interpreted as the earliest evidence of baleen and points to an intermediate condition between an ancestral condition with teeth only and a derived condition with baleen only. Supporting this step-wise evolutionary hypothesis, evidence from stable isotopes show how changes in dental chemistry in early toothed mysticetes tracked the changes in diet and environment. Recent discoveries also demonstrate how this transition was made possible by radical changes in cranial ontogeny. In addition, genetic mutations and the possession of dental pseudogenes in extant baleen whales support a toothed ancestry for mysticetes. Molecular and morphological data also document the dramatic developmental shifts that take place in extant fetal baleen whales, in skull development, resorption of a fetal dentition and growth of baleen. The mechanisms involved in this complex evolutionary transition that entails multiple, integrated aspects of anatomy and ecology are only beginning to be understood, and future work will further clarify the processes underlying this macroevolutionary pattern.

The origin of baleen in mysticetes heralded a major transition during cetacean evolution. Extant mysticetes are edentulous in adulthood, but rudimentary teeth develop in utero within open maxillary and mandibular alveolar grooves. The teeth are resorbed prenatally and the alveolar grooves close as baleen germ develops. Arteries supplying blood to highly vascularized epithelial tissue from which baleen develops pass through lateral nutrient foramina in the area of the embryonic alveolar grooves and rudimentary teeth. Those vessels are hypothesized to be branches of the superior alveolar artery, but branches of the greater palatine arteries may play a role in the baleen vascularization. Through a combination of latex injection, CT, and traditional dissection of the palate of a neonatal gray whale (Eschrichtius robustus), we confirm that the baleen receives blood from vessels within the superior alveolar canal via the lateral foramina. The greater palatine artery is restricted to its own passage with no connections to the baleen. This study has implications for the presence of baleen in extinct taxa by identifying the vessels and bony canals that supply blood to the epithelium from which baleen develops. The results indicate that the lateral foramina in edentulous mysticete fossils are bony correlates for the presence of baleen, and the results can be used to help identify bony canals and foramina that have been used to reconstruct baleen in extinct mysticetes that retained teeth in adulthood. Further comparisons are made with mammals that also possess oral keratin structures, including ruminants, ornithorhynchid monotremes, and sirenians. Anat Rec, 298:691-702, 2015. V C 2015 Wiley Periodicals, Inc.Key words: baleen; gray whale; Eschrichtius robustus; vascularization; palateIn this article, we investigate the blood supply to the baleen and surrounding epithelium covering the hard palate in a neonatal gray whale (Eschrichtius robustus). Baleen is a neomorphic keratinous structure that is unique to mysticete cetaceans (baleen whales) and functions as a natural sieve to trap prey during filter feeding (Pivorunas, 1979;Werth, 2001Werth, , 2004Werth, , 2013. Baleen develops as a series of transverse laminae (main and minor plates) that form a pair of bilateral racks along the ventrolateral borders of the maxillae. The racks of baleen laminae (or baleen racks) are oriented parasagittally, and they are deeply embedded within the highly Additional Supporting Information may be found in the online version of this article.