Correction: Redescription of three fossil baleen whale skulls from the Miocene of Portugal reveals new cetotheriid phylogenetic insights.

[This corrects the article DOI: 10.1371/journal.pone.0298658.].

A vertebra of a small species of Pachycetus from the North Sea and its inner structure and vascularity compared with other basilosaurid vertebrae from the same site.

In the Western Scheldt Estuary near the Belgian-Dutch border, middle to late Eocene strata crop out at the current seafloor. Most vertebrae of large Eocene basilosaurid taxa from this area were previously described in several papers. They represent three morphotypes: elongated vertebrae of a large species of Pachycetus (Morphotype 1b), a not-elongated vertebra of a large 'dorudontid' basilosaurid (Morphotype 2) and 'shortened' vertebrae of a new, unnamed taxon (Morphotype 3). This article deals with a still undescribed, smaller vertebra, NMR-16642, from this site. Our first aim was to date it by dinoflagellate cysts in adhering sediments. Yielding an age of about 38 Ma, it is one of the very few remains of basilosaurids from Europe, of which the age could be assessed with reasonable certainty. The vertebra, Morphotype 1a, is assigned to a small species of Pachycetus. High-quality CT scans are used to differentiate between NMR-16642, Morphotype 1a, and the large species of Pachycetus, Morphotype 1b. Another aim of this paper is to investigate the inner structure and vascularity of the study vertebra and that of the other morphotypes (1b, 2, 3) from this area by using high-quality CT scans. Notwithstanding differences in size, shape and compactness, the vertebral inner structure with a multi-layered cortex of periosteal bone, surrounding two cones of endosteal bone appears to be basically similar in all morphotypes. Apparently, this inner structure reflects the ontogenetic vertebral growth. An attempt to reconstruct the vascularity of the vertebrae reveals a remarkable pattern of interconnected vascular systems. From the dorsal and, if present, ventral foramina, vascular canals are running to a central vascular node. From this node a system of vascular canals goes to the epiphyseal ends, giving rise to separate systems for cortex and cones. It is the first time that the vascularity of vertebrae of archaeocetes is investigated.

Redescription of three fossil baleen whale skulls from the Miocene of Portugal reveals new cetotheriid phylogenetic insights.

Cetotheriidae is a family of baleen whales that went nearly extinct during the Pleistocene (excluding Caperea marginata). For a long time, the Cetotheriidae family has been seen as a problematic clade, but in the past two decades there have been various studies trying to resolve the phylogeny of this group. In 1831, Alexandre Vandelli described three cetotheriid skulls, found during a gold exploration at Adiça beach (Portugal). These specimens constituted the first Portuguese vertebrate fossils ever published in the literature. Another skull was added to the "Vandelli skulls" by Jacinto Pedro Gomes, in 1914, during a survey of the Museu Nacional de História Natural collections without giving information on the origin of this skull. In 1941, Remington Kellogg states that one of the original "Vandelli skulls" is no longer present in the Museu Nacional de História Natural collections. Until today, there is no information on how, or exactly when, the fourth skull and one of the original three "Vandelli skulls" appeared and disappeared, respectively. Since their discovery, all the attempts to describe these specimens were not based on direct observations and no comprehensive phylogenetic analysis have included the three skulls. Here we provide a detailed anatomic description, a new phylogenetic analysis and a palaeoecological reconstruction of these specimens, clarifying their relationships within the Cetotheriidae family and fostering the importance of these historical specimens to the modern comprehension of fossil whale evolution. In addition, our results support that Cephalotropis nectus is a valid species with an emended diagnosis. We also concluded that two specimens belong to a new genus, forming two new fossil species (new combinations).

The tympanoperiotic complex of the blue whale, Balaenoptera musculus.

The tympanoperiotic complex of a blue whale Balaenoptera musculus is described and compared to the homologous structures in the other extant and fossil baleen whale species. The periotic and the tympanic bulla represent informative anatomical regions in both functional and phylogenetic studies and for this reason a micro-CT scan of the bones was performed in order to better characterize their external aspect and to reconstruct the inner structures. In particular, the cochlea, the semicircular canals and associated portions of the periotic are reconstructed so that these structures may be used in phylogenetic analyses. We observed that the blue whale periotic is characterized by the presence of a strong dorsal protrusion which is posteriorly bordered by a previously undescribed morphological character that we name the posterotransverse fossa. The peculiar shape of the anterior process and the en echelon organization of the posterior foramina of the pars cochlearis are also described and compared. From a phylogenetic perspective, the blue whale is confirmed to be closely related to the fin whale, Balaenoptera physalus, but it is suggested, based on ear bone characters only, that it diverged before the other balaenopterid species in the phylogeny of Balaenopteridae. This placement supports a series of morphological observations suggesting that the extant blue whale was an early-diverging member of Balaenoptera. Our results help to decipher the evolutionary origin of the blue whale, the largest living animal, by allowing new and more detailed morphological analyses of the balaenopterid fossil record.

A new species of rorqual whale (Cetacea, Mysticeti, Balaenopteridae) from the Late Miocene of the Southern North Sea Basin and the role of the North Atlantic in the paleobiogeography of Archaebalaenoptera.

BACKGROUND: The rich fossil record of rorqual and humpback whales (Cetacea, Mysticeti, Balaenopteridae) is mainly characterized by monotypic genera since genera including more than one species are extremely rare. The discovery of new species belonging to known genera would be of great importance in order to better understand ancestor-descendant relationships and paleobiogeographic patterns in this diverse group. Recent discoveries in the southern North Sea Basin yielded a number of reasonably well preserved fossil balaenopterids from the Late Miocene; this sample includes a balaenopterid skull from Liessel, The Netherlands, which shares key characters with Archaebalaenoptera castriarquati from the Pliocene of Mediterranean. This skull is permanently held by Oertijdmuseum, Boxtel, The Netherlands, with the number MAB002286 and is investigated here. METHODS: A detailed comparative anatomical analysis of the skull MAB002286 is performed in order to understand its relationships. The age of the skull is determined by dinocyst analysis of the associated sediment. A paleobiogeographic analysis is performed to understand paleobiogeographic patterns within the balaenopterid clade the new skull belongs to. RESULTS: Our work resulted in the description of Archaebalaenoptera liesselensis new species. The geological age of the holotype skull is between 8.1 and 7.5 Ma. The phylogenetic relationships of this species reveals that it is monophyletic with Archaebalaenoptera castriarquati from the Italian Pliocene. Moreover, in combination with a more basal species of Archaebalaenoptera from the late Miocene of Peru, our paleobiogeographic analysis suggests that the North Atlantic ocean played a major role as a center of origin of a number of balaenopterid clades including Protororqualus, Archaebalaenoptera and more advanced balaenopterid taxa. From a North Atlantic center of origin, two dispersal events are inferred that led to the origins of Archaebalaenoptera species in the South Pacific and Mediterranean. The distribution of Archaebalaenoptera was antitropical in the late Miocene. The role played by the Mediterranean salinity crisis is also investigated and discussed.

A large Late Miocene cetotheriid (Cetacea, Mysticeti) from the Netherlands clarifies the status of Tranatocetidae.

Cetotheriidae are a group of small baleen whales (Mysticeti) that evolved alongside modern rorquals. They once enjoyed a nearly global distribution, but then largely went extinct during the Plio-Pleistocene. After languishing as a wastebasket taxon for more than a century, the concept of Cetotheriidae is now well established. Nevertheless, the clade remains notable for its variability, and its scope remains in flux. In particular, the recent referral of several traditional cetotheriids to a new and seemingly unrelated family, Tranatocetidae, has created major phylogenetic uncertainty. Here, we describe a new species of Tranatocetus, the type of Tranatocetidae, from the Late Miocene of the Netherlands. Tranatocetus maregermanicum sp. nov. clarifies several of the traits previously ascribed to this genus, and reveals distinctive auditory and mandibular morphologies suggesting cetotheriid affinities. This interpretation is supported by a large phylogenetic analysis, which mingles cetotheriids and tranatocetids within a unified clade. As a result, we suggest that both groups should be reintegrated into the single family Cetotheriidae.

Northern pygmy right whales highlight Quaternary marine mammal interchange.

The pygmy right whale, Caperea marginata, is the most enigmatic living whale. Little is known about its ecology and behaviour, but unusual specialisations of visual pigments [1], mitochondrial tRNAs [2], and postcranial anatomy [3] suggest a lifestyle different from that of other extant whales. Geographically, Caperea represents the only major baleen whale lineage entirely restricted to the Southern Ocean. Caperea-like fossils, the oldest of which date to the Late Miocene, are exceedingly rare and likewise limited to the Southern Hemisphere [4], despite a more substantial history of fossil sampling north of the equator. Two new Pleistocene fossils now provide unexpected evidence of a brief and relatively recent period in geological history when Caperea occurred in the Northern Hemisphere (Figure 1A,B).

Revision of "Balaena" belgica reveals a new right whale species, the possible ancestry of the northern right whale, Eubalaena glacialis, and the ages of divergence for the living right whale species.

In 1941, Abel established Balaena belgica based on a series of fused cervical vertebrae and citing other cranial fragments from the late Neogene of the Antwerp harbor (northern Belgium). Later, Plisnier-Ladame & Quinet (1969) added a neurocranium and other skeletal remains from the same area to this species. Recently, the neurocranium was re-assigned to the genus Eubalaena thanks to newer phylogenetic analyses. Here, a new description is provided of materials previously assigned to "Balaena" belgica together with taxonomic revisions. Our work suggests that the cervical complex originally designated as the type of "Balaena" belgica is too poorly preserved to be used as such and is assigned to Balaenidae gen. et sp. indet., thus making "Balaena" belgica a nomen dubium. In addition to the neurocranium, the other remains consist in a fragment of maxilla assigned to Balaenidae gen. et sp. indet. and in a humerus assigned to Eubalaena sp. Discovered in the Kruisschans Sands Member of the Lillo Formation (3.2-2.8 Ma, Piacenzian, Late Pliocene), the neurocranium is designated as the holotype of the new species Eubalaena ianitrix. Our phylogenetic analysis supports a sister-group relationship of Eubalaena ianitrix and Eubalaena glacialis, and helps constraining the ages of origin for balaenid clades. Ecological and phylogenetic data suggest that Eubalaena ianitrix may represent the direct ancestor of Eubalaena glacialis, the latter having evolved through phyletic transformation including body size increase during the temperature decline of the Late Pliocene.

Cetopirus complanatus (Cirripedia: Coronulidae) from the late Middle Pleistocene human settlement of Pinnacle Point 13B (Mossel Bay, South Africa).

The late Middle Pleistocene cave site of Pinnacle Point 13B (PP13B, South Africa) has provided the archaeologically oldest evidences yet known of human consumption of marine resources. Among the marine invertebrates recognised at PP13B, an isolated whale barnacle compartment was tentatively determined as Coronula diadema and regarded as indirect evidence of human consumption of a baleen whale (likely Megaptera novaeangliae). In this paper we redetermine this coronulid specimen as Cetopirus complanatus. This record significantly extends the fossil history of C. complanatus back by about 150 ky, thus partially bridging the occurrence of Cetopirus fragilis in the early Pleistocene to the latest Quaternary record of C. complanatus. Since C. complanatus is currently known as a highly specific phoront of right whales (Eubalaena spp.), we propose that the late Middle Pleistocene human groups that inhabited PP13B fed on a stranded southern right whale. Therefore, the whale barnacle from PP13B suggests the persistence of a southern right whale population off South Africa during the predominantly glacial MIS 6, thus evoking the continuity of cetacean migrations and antitropical distribution during that global cold phase. Interestingly, the most ancient evidence of humans feeding on a whale involves Eubalaena, historically the most exploited cetacean genus, and currently still seriously threatened with extinction due to human impact.

The whale barnacle Cryptolepas rhachianecti (Cirripedia: Coronulidae), a phoront of the grey whale Eschrichtius robustus (Cetacea: Eschrichtiidae), from a sandy beach in The Netherlands.

An isolated compartment of a whale barnacle is herein described from Recent beach deposits in Zoutelande (Walcheren, The Netherlands). This specimen is identified as belonging to the extant coronulid species Cryptolepas rhachianecti, currently known as an epizoic symbiont of the grey whale Eschrichtius robustus. This find represents the first occurrence of C. rhachianecti outside the North Pacific, and the first one as a (sub)fossil. In view of the fact that E. robustus, which is currently confined to the North Pacific, is known as a subfossil from the northeastern Atlantic between late Late Pleistocene (c. 45,000 years ago) and historical (c. 1700 AD) times, we propose a similar (late Quaternary) age for the isolated compartment. The find indicates that the extinct late Quaternary northeastern Atlantic population of E. robustus was infected by Cryptolepas rhachianecti. Our find is, therefore, compatible with the hypothesis of an ancient grey whale migration route running between the subtropical/temperate waters of the northeast Atlantic (or Mediterranean Basin), and the cold waters of the Baltic Sea (or southern Arctic Ocean), through the southern North Sea. Finally, we discuss the systematic placement of the fossil barnacle species Cryptolepas murata and propose the possibility of its removal from the genus Cryptolepas pending further investigations.

A new species of Metopocetus (Cetacea, Mysticeti, Cetotheriidae) from the Late Miocene of the Netherlands.

The family Cetotheriidae has played a major role in recent discussions of baleen whale phylogenetics. Within this group, the enigmatic, monotypic Metopocetus durinasus has been interpreted as transitional between herpetocetines and other members of the family, but so far has been restricted to a single, fragmentary cranium of uncertain provenance and age. Here, we expand the genus and shed new light on its phylogenetic affinities and functional morphology by describing Metopocetus hunteri sp. nov. from the Late Miocene of the Netherlands. Unlike the holotype of M. durinasus, the material described here is confidently dated and preserves both the tympanic bulla and additional details of the basicranium. M. hunteri closely resembles M. durinasus, differing primarily in its somewhat less distally expanded compound posterior process of the tympanoperiotic. Both species are characterised by the development of an unusually large fossa on the ventral surface of the paroccipital process, which extends anteriorly on to the compound posterior process and completely floors the facial sulcus. In life, this enlarged fossa may have housed the posterior sinus and/or the articulation of the stylohyal. Like other cetotheriids, Metopocetus also bears a well-developed, posteriorly-pointing dorsal infraorbital foramen near the base of the ascending process of the maxilla, the precise function of which remains unclear.