Mónica R. Buono

Juvenile morphology: A clue to the origins of the most mysterious of mysticetes?

The origin of the pygmy right whale (Caperea marginata) has long been one of the most vexing conundrums of marine mammal evolution. The extremely disparate skeletal structure of Caperea and a patchy fossil record have left morphology and molecules at odds: whereas most morphological analyses ally Caperea with right whales (Balaenidae), most molecular studies instead suggest a close relationship with rorquals (Balaenopteridae) and grey whales (Eschrichtiidae). The morphological evidence supporting a Caperea-balaenid clade consists of several shared features of the skull and mandible, as traditionally observed in adult individuals. Here, we show that at least two of these features, the ascending process of the maxilla and the coronoid process, arise from substantially different precursors early during ontogeny and therefore likely do not represent genuine synapomorphies. Both of these juvenile morphologies have adult counterparts in the fossil record, thus indicating that the ontogenetic variation in the living species may be a genuine reflection of differing ancestral states. This new evidence contradicts previous morphological hypotheses on the origins of Caperea and may help to reconcile morphological and molecular evidence.

A new beaked whale (Cetacea, Odontoceti) from the Late Miocene of Patagonia, Argentina

ABSTRACT A new genus and species of Ziphiidae, Notoziphius bruneti, gen. et sp. nov., from the late Miocene of Patagonia, is described on the basis of a well-preserved skull and partial left and right dentaries. It can be diagnosed by large, triangular, and markedly asymmetric nasals that strongly point anteroventrally, the ascending process of the maxilla not expanded posteriorly, supraoccipital strongly sloped posteroventrally; elliptical fossa on the nasal process of the premaxilla, small and anterolaterally directed premaxillary crest, and the presence of well-defined alveoli in the maxilla. A phylogenetic analysis including 25 ziphiid genera and 31 characters shows Notoziphius as nested within Ziphiidae in a basal clade with Aporotus, Beneziphius, Messapicetus, and Ziphirostrum. This clade is diagnosed by medial fusion of the premaxillae that closes the mesorostral groove, lateral margin of prenarial basin formed by a thick strip of maxilla, premaxillary crest anterolaterally directed, and reduced contact between nasal and premaxillary crest. The presence of Notoziphius in the Miocene of Patagonia increases our knowledge of ziphiid diversity in South America. Notoziphius bruneti and other Miocene records of ziphiids reinforce the idea that during the Miocene ziphiids were widely distributed and diverse.

Morphology of the Eye of the Southern Right Whales (Eubalaena australis)

Recently, there has been a growing interest in the anatomy and optics of the visual system of cetaceans. However, much of the new information has been focused on odontocetes, and relatively little is known about the visual anatomy of baleen whales. The aim of this study was describe the eye anatomy of the southern right whale (Eubalaena australis). Eye samples were collected from 26 calves, four adults with known body length, as well as two specimens of unknown body length that had stranded near their nursery ground at Península Valdés, Argentina, over 6 years. We provide anatomical descriptions of the eyeball and extraocular structures, as well as quantitative data in the form of eyeball, corneal, scleral, and lens measurements. To explore the sensitivity of the eye to light, the f‐number was estimated in one specimen. We found that the eyes of the calves differed from those of the adults in having less periorbital fat surrounding the eyeball. We also observed variations in the abundance of periorbital fat among the adult specimens. The regression analysis revealed a correlation between body length and eyeball size. By contrast, the dimensions of the cornea were only weakly correlated with body length. The estimated f‐number suggests that the optical sensitivity of the Eubalaena australis eye is relatively low. However, caution had to be taken in interpreting f‐number as a proxy of eye sensitivity because it depends on the lens size, which can be affected by the fixation methods used. Anat Rec, 2012.

Eocene basilosaurid whales from the La Meseta Formation, Marambio (Seymour) Island, Antarctica

Abstract. Basal fully aquatic whales, the basilosaurids are worldwide known from Bartonian—Priabonian localities, indicating that this group was widely distributed during the late middle Eocene. In the Northern Hemisphere, fossils of basilosaurids are abundant, while records in the Southern Hemisphere are scarce and, in some cases (i.e., Antarctica), doubtful. The presence of basilosaurids in Antarctica was, until now, uncertain because most of the records are based on fragmentary materials that preclude an accurate assignment to known archaeocete taxa. Here we report the findings of mandibles, teeth, and innominate bone remains of basilosaurids recovered from the La Meseta Formation (TELM 4 Lutetian—Bartonian and; TELM 7 Priabonian), in Marambio (Seymour) Island (James Ross Basin, Antarctic Peninsula). These findings confirm the presence of Basilosauridae in the marine realm of Antarctica, increasing our knowledge of the paleobiogeographic distribution of basilosaurids during the middle—late Eocene. In addition, one of these records is among the oldest occurrences of basilosaurids worldwide, indicating a rapid radiation and dispersal of this group since at least the early middle Eocene. Resumen. El registro fósil de los basilosáuridos está bien documentado durante el Bartoniano—Priaboniano en varias localidades del mundo, lo cual indica que este grupo estaba ampliamente distribuido durante el Eoceno medio tardío. En el Hemisferio Norte, el registro fósil de este grupo es abundante, a diferencia de lo que ocurre en el Hemisferio Sur donde es escaso y, en algunos casos (i.e., Antártida), dudoso. La presencia de basilosáuridos en Antártida es incierta ya que la mayoría de los registros están basados en materiales fragmentarios, lo cual imposibilita su asignación a algún grupo de arqueocetos. En la presente contribución se describen restos de basilosáuridos correspondientes a mandíbulas, dientes aislados y un hueso pélvico, recuperados de la Formación La Meseta (TELM 4 Lutetiano—Bartoniano; TELM 7 Priaboniano), Isla Marambio (Seymour), (Cuenca James Ross, Península Antártica). Este hallazgo confirma la presencia de Basilosauridae en la Antártida, contribuyendo al conocimiento de la distribución paleobiogeográfica de este grupo durante el Eoceno medio—tardío. Finalmente, uno de estos registros se encuentra entre los basilosáuridos más antiguos conocidos, indicando un rápida radiación y dispersión de este grupo al menos desde el Eoceno medio temprano.

A Late Miocene Potential Neobalaenine Mandible from Argentina Sheds Light on the Origins of the Living Pygmy Right Whale

The origins and evolutionary relationships of the pygmy right whale (Caperea marginata), the only living member of the Neobalaeninae, have been the subject of a long-standing debate. This phylogenetic uncertainty is compounded by a limited neobalaenine fossil record. Here, we report a Late Miocene mysticete mandible from Patagonia, Argentina, and provisionally refer it to Neobalaeninae, gen. et sp. indet. The new material represents only the third report of a fossil neobalaenine, and the first fossil occurrence of this lineage in the southwestern Atlantic. It is also the oldest specimen so far reported, thus corroborating the idea of an early divergence time for neobalaenines.

Miocene Marine Transgressions: Paleoenvironments and Paleobiodiversity

Two major marine transgressions covered part of Patagonia during the Miocene and both are recorded in the Peninsula Valdes region. The older (early Miocene) is represented by the volumetrically scarce outcrops of the Gaiman Formation, composed by shelf mudstones and fine sandstones. The late Miocene transgression is represented by the Puerto Madryn Formation, widely distributed in Peninsula Valdes and composed of mudstones, sandstones and shell beds, being the focus of this work. Sediments of this unit were deposited in inner shelf, nearshore, tidal channel and tidal flat environments. Fossil content is abundant and diverse, including palynomorphs, foraminifers, marine invertebrates (dominated by molluscs), cetaceans, pinnipeds, marine fishes and birds, as well as continental mammals, birds, and fishes. Isotopic and biostratigraphical data suggest a late Miocene age for the Puerto Madryn Formation, although some middle Miocene biostratigraphical indicators are present. Paleoenvironmental information suggests oceanic and continental temperatures warmer than present day, evidenced by the Caribbean molluscan association and the continental vertebrate and palinological associations, respectively. Instead, cetaceans, dinoflagellates, and some marine fishes, suggest colder oceanic temperatures. Precipitations were also higher than present, evidenced by the presence of freshwater mammals, birds, fishes, and plants. This work highlighted some gaps in the geological and paleontological knowledge including geochronology, stratigraphic control of paleontological studies and the knowledge of poorly known fossil groups, which should be the focus of future investigations.

Anatomy of nasal complex in the southern right whale, Eubalaena australis (Cetacea, Mysticeti).

The nasal region of the skull has undergone dramatic changes during the course of cetacean evolution. In particular, mysticetes (baleen whales) conserve the nasal mammalian pattern associated with the secondary function of olfaction, and lack the sound‐producing specializations present in odontocetes (toothed whales, dolphins and porpoises). To improve our understanding of the morphology of the nasal region of mysticetes, we investigate the nasal anatomy, osteology and myology of the southern right whale, Eubalaena australis, and make comparisons with other mysticetes. In E. australis external deflection surfaces around the blowholes appear to divert water off the head, and differ in appearance from those observed in balaenopterids, eschrichtiids and cetotherids. In E. australis the blowholes are placed above hypertrophied nasal soft tissues formed by fat and nasal muscles, a pattern also observed in balaenopterids (rorqual mysticetes) and a cetotherid (pygmy right whale, Caperea marginata). Blowhole movements are due to the action of five nasofacial muscles: dilator naris superficialis, dilator naris profundus, depressor alae nasi, constrictor naris, and retractor alae nasi. The dilator naris profundus found in E. australis has not been previously reported in balaenopterids. The other nasofacial muscles have a similar arrangement in balaenopterids, with minor differences. A novel structure, not reported previously in any mysticete, is the presence of a vascular tissue (rete mirabile) covering the lower nasal passage. This vascular tissue could play a role in warming inspired air, or may engorge to accommodate loss of respiratory space volume due to gas compression from increased pressure during diving.

The early Miocene balaenid Morenocetus parvus from Patagonia (Argentina) and the evolution of right whales

Balaenidae (right and bowhead whales) are a key group in understanding baleen whale evolution, because they are the oldest surviving lineage of crown Mysticeti, with a fossil record that dates back ∼20 million years. However, this record is mostly Pliocene and younger, with most of the Miocene history of the clade remaining practically unknown. The earliest recognized balaenid is the early Miocene Morenocetus parvus Cabrera, 1926 from Argentina. M. parvus was originally briefly described from two incomplete crania, a mandible and some cervical vertebrae collected from the lower Miocene Gaiman Formation of Patagonia. Since then it has not been revised, thus remaining a frequently cited yet enigmatic fossil cetacean with great potential for shedding light on the early history of crown Mysticeti. Here we provide a detailed morphological description of this taxon and revisit its phylogenetic position. The phylogenetic analysis recovered the middle Miocene Peripolocetus as the earliest diverging balaenid, and Morenocetus as the sister taxon of all other balaenids. The analysis of cranial and periotic morphology of Morenocetus suggest that some of the specialized morphological traits of modern balaenids were acquired by the early Miocene and have remained essentially unchanged up to the present. Throughout balaenid evolution, morphological changes in skull arching and ventral displacement of the orbits appear to be coupled and functionally linked to mitigating a reduction of the field of vision. The body length of Morenocetus and other extinct balaenids was estimated and the evolution of body size in Balaenidae was reconstructed. Optimization of body length on our phylogeny of Balaenidae suggests that the primitive condition was a relatively small body length represented by Morenocetus, and that gigantism has been acquired independently at least twice (in Balaena mysticetus and Eubalaena spp.), with the earliest occurrence of this trait in the late Miocene–early Pliocene as represented by Eubalaena shinshuensis.