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.

Novel muscle and connective tissue design enables high extensibility and controls engulfment volume in lunge-feeding rorqual whales.

Muscle serves a wide variety of mechanical functions during animal feeding and locomotion, but the performance of this tissue is limited by how far it can be extended. In rorqual whales, feeding and locomotion are integrated in a dynamic process called lunge feeding, where an enormous volume of prey-laden water is engulfed into a capacious ventral oropharyngeal cavity that is bounded superficially by skeletal muscle and ventral groove blubber (VGB). The great expansion of the cavity wall presents a mechanical challenge for the physiological limits of skeletal muscle, yet its role is considered fundamental in controlling the flux of water into the mouth. Our analyses of the functional properties and mechanical behaviour of VGB muscles revealed a crimped microstructure in an unstrained, non-feeding state that is arranged in parallel with dense and straight elastin fibres. This allows the muscles to accommodate large tissue deformations of the VGB yet still operate within the known strain limits of vertebrate skeletal muscle. VGB transverse strains in routine-feeding rorquals were substantially less than those observed in dead ones, where decomposition gas stretched the VGB to its elastic limit, evidence supporting the idea that eccentric muscle contraction modulates the rate of expansion and ultimate size of the ventral cavity during engulfment.

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 new balaenopterid whale from the late Miocene of the Southern North Sea Basin and the evolution of balaenopterid diversity (Cetacea, Mysticeti).

BACKGROUND: Balaenopterid mysticetes represent the most successful family-rank group of this clade. Their evolutionary history is characterized by a rich fossil record but the origin of the living genera is still largely not understood. Recent discoveries in the southern border of the North Sea revealed a number of well preserved fossil balaenopterid whales that may help resolving this problem. In particular, skull NMR 14035 shares morphological characters with the living humpback whale, Megaptera novaeangliae and, for this reason, its characteristics are investigated here. METHODS: The comparative anatomical analysis of the new specimen formed the basis of a new phylogenetic analysis of the Mysticeti based on a matrix including 350 morphological character states scored for 82 Operational Taxonomic Units. The stratigraphic age of the specimen was determined based on the analysis of the dinocyst assemblage recovered in the associated sediment. We assessed clade diversity in Balaenopteridae by counting the numbers of clades in given time intervals and then plotted the results. RESULTS: Nehalaennia devossi n. gen. et sp. is described for the first time from the late Tortonian (8.7-8.1 Ma) of the Westerschelde (The Netherlands). This new taxon belongs to Balaenopteridae and shows a surprisingly high number of advanced characters in the skull morphology. Nehalaennia devossi is compared to a large sample of balaenopterid mysticetes and a phylogenetic analysis placed it as the sister group of a clade including the genus Archaebalaenoptera. The inclusion of this fossil allowed to propose a phylogenetic hypothesis for Balaenopteridae in which (1) Eschrichtiidae (gray whales) represents a family of its own, (2) Balaenopteridae + Eschrichtiidae form a monophyletic group (superfamily Balaenopteroidea), (3) Cetotheriidae is the sister group of Balaenopteroidea, (4) living Balaenoptera species form a monophyletic group and (5) living M. novaeangliae is the sister group of Balaenoptera. Our work reveals a complex phylogenetic history of Balaenopteridae and N. devossi informs us about the early morphological transformations in this family. Over time, this family experienced a number of diversity pulses suggesting that true evolutionary radiations had taken place. The paleoecological drivers of these pulses are then investigated.

Using opportunistic sightings to infer differential spatio-temporal use of western Mediterranean waters by the fin whale.

The fin whale (Balaenoptera physalus) is a cosmopolitan species with a resident population in the Mediterranean Sea. Due to its habitat, open seas often far from ports and airfields, and its long-distance migratory behaviour, studying and monitoring its distribution is costly. Currently, many opportunistic sightings (OS) reports are available, which provide a source of potentially useful, low-cost information about the spatio-temporal distribution of this species. Since 1993, the Spanish Institute of Oceanography has compiled a dataset comprising 874 records of OS of nine species of cetaceans in the western Mediterranean Sea and adjacent waters. The aim of this study was to use this dataset to investigate the differential use of these waters by the fin whale when compared with other cetaceans. We compared the presence of fin whales with the presence of any other cetacean species in the dataset. Binary logistic regression was then used to model these occurrences according to several spatio-temporal variables expected to reflect their habitat use. Several significant models reveal that fin whales are more prone than other cetaceans to use the waters over the slope of the Gulf of Lion in summer. This finding confirms that the Gulf of Lion is an area of importance for this species and suggests that the slope of the continental shelf could be particularly important. Our study shows how OS can be a source of useful information when appropriately analyzed.

Allometric scaling of morphology and engulfment capacity in rorqual whales.

Body length is one of the most important factors that influence organismal function and ecological niche. Although larger animals tend to have a suite of physiological advantages, such as lower mass-specific metabolic rates and lower costs of transport, they may also experience significant limitations to unsteady locomotor performance or maneuverability because of the relative scaling of control surface areas and body mass. Rorqual whales are the largest of all animals and thus represent a unique study system for understanding how animals function at the extreme of body mass. Rorquals are characterized by an engulfment-filtration foraging strategy facilitated by a complex set of morphological adaptations. We studied the scaling of key morphological structures related to locomotion and feeding in six rorqual species in a comparative framework. Our analyses show that most rorqual species exhibit positive allometry of both the control surfaces and body length, but the large scaling differences between these parameters suggest that larger rorquals will predictably suffer from decreased maneuverability and unsteady locomotor performance. However, we found that the dimensions of the engulfment apparatus also exhibit positive allometry, and thus engulfment capacity was relatively greater in larger rorquals. We posit that the allometric growth in the engulfment apparatus may be an adaptation that ameliorates the detrimental effects of large size on maneuverability. Our analyses also reveal significant differences in the scaling of mass-specific engulfment capacity among rorqual species that may reflect the evolution of unique foraging behaviors and the exploitation of divergent ecological niches.

Stranding survey as a framework to investigate rare cetacean records of the north and north-eastern Brazilian coasts.

Marine mammal stranding events are used as an important tool for understanding cetacean biology worldwide. Nonetheless, there are vast gaps of knowledge to be filled in for a wide range of species. Reputable information is required regarding species from large baleen whales to sperm and beaked whales, as well as pelagic dolphins. This paper describes new cetacean records from north and north-eastern Brazil, which are both the least surveyed areas regarding aquatic mammals. Regular beach surveys were conducted to recover cetacean carcasses along the coast of Pará beginning November 2005. At the coasts of the Maranhão and Piauí states, the surveys were conducted between 2003 and 2013. From 2003 to 2014, 34 strandings of cetaceans were registered. The study provides four additional species records' in the area based on strandings (Balaenoptera borealis, Balaenoptera physalus, Peponocephala electra, and Pseudorca crassidens). A mass stranding of Guiana dolphins (Sotalia guianensis, N = 12), the most common species for the region, was reported for the first time. The records presented herein are of special concern, since they expand the knowledge on cetaceans from the Brazilian coast. In addition, this study conducted an analysis to verify the similarity between cetacean compositions described for north and north-eastern Brazil and the southern Caribbean region. The results showed a high similarity between these regions, proving the connection with the Caribbean cetacean fauna.

Largest baleen whale mass mortality during strong El Niño event is likely related to harmful toxic algal bloom.

While large mass mortality events (MMEs) are well known for toothed whales, they have been rare in baleen whales due to their less gregarious behavior. Although in most cases the cause of mortality has not been conclusively identified, some baleen whale mortality events have been linked to bio-oceanographic conditions, such as harmful algal blooms (HABs). In Southern Chile, HABs can be triggered by the ocean-atmosphere phenomenon El Niño. The frequency of the strongest El Niño events is increasing due to climate change. In March 2015, by far the largest reported mass mortality of baleen whales took place in a gulf in Southern Chile. Here, we show that the synchronous death of at least 343, primarily sei whales can be attributed to HABs during a building El Niño. Although considered an oceanic species, the sei whales died while feeding near to shore in previously unknown large aggregations. This provides evidence of new feeding grounds for the species. The combination of older and newer remains of whales in the same area indicate that MMEs have occurred more than once in recent years. Large HABs and reports of marine mammal MMEs along the Northeast Pacific coast may indicate similar processes in both hemispheres. Increasing MMEs through HABs may become a serious concern in the conservation of endangered whale species.

A new Miocene baleen whale from the Peruvian desert.

The Pisco-Ica and Sacaco basins of southern Peru are renowned for their abundance of exceptionally preserved fossil cetaceans, several of which retain traces of soft tissue and occasionally even stomach contents. Previous work has mostly focused on odontocetes, with baleen whales currently being restricted to just three described taxa. Here, we report a new Late Miocene rorqual (family Balaenopteridae), Incakujira anillodefuego gen. et sp. nov., based on two exceptionally preserved specimens from the Pisco Formation exposed at Aguada de Lomas, Sacaco Basin, southern Peru. Incakujira overall closely resembles modern balaenopterids, but stands out for its unusually gracile ascending process of the maxilla, as well as a markedly twisted postglenoid process of the squamosal. The latter likely impeded lateral (omega) rotation of the mandible, in stark contrast with the highly flexible craniomandibular joint of extant lunge-feeding rorquals. Overall, Incakujira expands the still meagre Miocene record of balaenopterids and reveals a previously underappreciated degree of complexity in the evolution of their iconic lunge-feeding strategy.

Diversidad de cetáceos en el paisaje marino costeros de Golfo Dulce, Península de Osa, Costa Rica / Diversity of cetaceans in coastal seascape Golfo Dulce, Osa Peninsula, Costa Rica

Resumen Golfo Dulce es un estuario en forma de fiordo, localizado en el Pacífico Sur de Costa Rica. Este ecosistema marino-costero contiene una importante biodiversidad marina que incluye cetáceos. Este estudio describe la estructura de la comunidad de cetáceos en Golfo Dulce, sustentado en la diversidad de hábitats. Datos sobre avistamientos colectados en muestreos periódicos sistemáticos, han resultado en 943 registros (2005-2014≈3 490h≈64 533km) de siete especies de la familia Delphinidae, una Balaenopteridae y una especie de la familia Kogiidae. Se observa un gradiente en la diversidad de cetáceos desde la cuenca interna de Golfo Dulce hasta la zona de transición- oceánica en la entrada al Golfo. La dominancia se establece por la forma costera del delfín nariz de botella y el delfín manchado en la cuenca interna, mientras que T. truncatus junto con ballenas jorobadas migrantes ejercen la dominancia en el área del Umbral. El área transicional oceánica sostiene la mayor diversidad, dominada por especies de delfínidos de naturaleza piscívora, incluyendo la forma oceánica del delfín nariz de botella y la subespecie "Centroamericana" del delfín tornillo. Nosotros proponemos la hipótesis de una forma potencial particular del delfín manchado pantropical para Golfo Dulce, en contraste con S. attenuatta graffmani en aguas abiertas costeras del Pacífico Costarricense. El patrón de la estructura comunitaria de cetáceos se establece por la dinámica en uso de área por las especies presentes, lo cual debería ser considerado para el diseño de estrategias de manejo y conservación.

Abstract Golfo Dulce is a fiord-lilke embayment located in the southern Pacific of Costa Rica, this coastal-marine ecosystem harbors an important wealth in marine biodiversity, including cetaceans. In this study we describe the cetacean community of Golfo Dulce, in relation to the diversity of habitats constituting the seascape. Sighting data collected in periodic systematic surveys has yielded 943 records (2005-2014≈3 490h≈64 533km), of 12 species of cetaceans. A gradient of cetacean diversity is observed from the inner basin to the transitional-oceanic area at the entrance of the Gulf, with coastal bottlenose dolphin and spotted dolphin dominating the inner basin, and bottlenose dolphins and migratory humpback whales the sill area. Transitional oceanic habitat holds the greatest species diversity dominated by oceanic species, particularly piscivorous delphinids, including the off-shore form of bottlenose dolphins and the "Central American" spinner dolphin. We hypothesize the potential occurrence of a morphotype of pantropical spotted dolphins within Golfo Dulce, in contrast with S. attenuatta graffmani outside in open Pacific coastal waters. The pattern of cetacean community structure is established by the dynamic of area use by the species, which should be considered for the establishment of management and conservation strategies. Rev. Biol. Trop. 63 (2): 395-406. Epub 2015 June 01.