Gene duplications and gene loss in the epidermal differentiation complex during the evolutionary land-to-water transition of cetaceans.

Major protein components of the mammalian skin barrier are encoded by genes clustered in the Epidermal Differentiation Complex (EDC). The skin of cetaceans, i.e. whales, porpoises and dolphins, differs histologically from that of terrestrial mammals. However, the genetic regulation of their epidermal barrier is only incompletely known. Here, we investigated the EDC of cetaceans by comparative genomics. We found that important epidermal cornification proteins, such as loricrin and involucrin are conserved and subtypes of small proline-rich proteins (SPRRs) are even expanded in numbers in cetaceans. By contrast, keratinocyte proline rich protein (KPRP), skin-specific protein 32 (XP32) and late-cornified envelope (LCE) genes with the notable exception of LCE7A have been lost in cetaceans. Genes encoding proline rich 9 (PRR9) and late cornified envelope like proline rich 1 (LELP1) have degenerated in subgroups of cetaceans. These data suggest that the evolution of an aquatic lifestyle was accompanied by amplification of SPRR genes and loss of specific other epidermal differentiation genes in the phylogenetic lineage leading to cetaceans.

Learning to play: A review and theoretical investigation of the developmental mechanisms and functions of cetacean play.

Play is a behavioral phenomenon most commonly observed in the young of both solitary and social species. Documentation of play in cetaceans varies across species and settings. Cetacean play behavioral repertoires include a broad range of actions, such as the manipulation of diverse objects, blowing bubbles, chasing conspecifics, and swimming in spirals through the water. As is common in research on animal play, cetacean play has been grouped into categories by its form, including locomotor play, object play, and different variations of social play, such as affiliative games, play fighting, and socio-sexual play. Research has primarily focused on recording the topography of cetacean play and the demographics of the individuals engaging in play. However, these classifications are insufficient to address the possible developmental and societal functions of cetacean play behaviors, or the mechanisms with which play behaviors are spread between conspecifics and acquired by young members of cetacean populations. This article applies several developmental and social learning theories in order to organize current knowledge and guide future research.

Finding sacral: Developmental evolution of the axial skeleton of odontocetes (Cetacea).

Axial morphology was dramatically transformed during the transition from terrestrial to aquatic environments by archaeocete cetaceans, and again during the subsequent odontocete radiation. Here, we reconstruct the sequence of developmental events that underlie these phenotypic transitions. Archaeocete innovations include the loss of primaxial/abaxial interaction at the sacral/pelvic articulation and the modular dissociation of the fluke from the remainder of the tail. Odontocetes subsequently integrated lumbar, sacral, and anterior caudal vertebrae into a single torso module, and underwent multiple series-specific changes in vertebral count. The conservation of regional proportions despite regional fluctuations in count strongly argues that rates of somitogenesis can vary along the column and that segmentation was dissociated from regionalization during odontocete evolution. Conserved regional proportions also allow the prediction of the location and count of sacral homologs within the torso module. These predictions are tested with the analysis of comparative pudendal nerve root location and geometric morphometrics. We conclude that the proportion of the column represented by the sacral series has been conserved, and that its vertebrae have changed in count and relative centrum length in parallel with other torso vertebrae. Although the sacral series of odontocetes is de-differentiated, it is not de-regionalized.

Inner ear development in cetaceans.

Cetaceans face the challenge of maintaining equilibrium underwater and obtaining sensory input within a dense, low-visibility medium. The cetacean ear represents a key innovation that marked their evolution from terrestrial artiodactyls to among the most fully aquatic mammals in existence. Using micro-CT and histological data, we document shape and size changes in the cetacean inner ear during ontogeny, and demonstrate that, as a proportion of gestation time, the cetacean inner ear is precocial in its growth compared with that of suid artiodactyls. Cetacean inner ears begin ossifying and reach near-adult shape as early as at 32% of the gestation period, and near-adult dimensions as early as at 27% newborn total length. Our earliest embryos with measurable inner ears (13% newborn length) exhibit a flattened cochlea (i.e. smaller distance from cochlear apex to round window) compared with later and adult stages. Inner ears of Sus scrofa have neither begun ossifying nor reached near-adult dimensions at 55% of the gestation period, but have an adult-like ratio of cochlear diameters to each other, suggesting an adult-like shape. The precocial development of the cetacean inner ear complements previous work demonstrating precocial development of other cetacean anatomical features such as the locomotor muscles to facilitate swimming at the moment of birth.

Phylogeny and adaptive evolution of the brain-development gene microcephalin (MCPH1) in cetaceans.

BACKGROUND: Representatives of Cetacea have the greatest absolute brain size among animals, and the largest relative brain size aside from humans. Despite this, genes implicated in the evolution of large brain size in primates have yet to be surveyed in cetaceans. RESULTS: We sequenced ~1240 basepairs of the brain development gene microcephalin (MCPH1) in 38 cetacean species. Alignments of these data and a published complete sequence from Tursiops truncatus with primate MCPH1 were utilized in phylogenetic analyses and to estimate ω (rate of nonsynonymous substitution/rate of synonymous substitution) using site and branch models of molecular evolution. We also tested the hypothesis that selection on MCPH1 was correlated with brain size in cetaceans using a continuous regression analysis that accounted for phylogenetic history. Our analyses revealed widespread signals of adaptive evolution in the MCPH1 of Cetacea and in other subclades of Mammalia, however, there was not a significant positive association between ω and brain size within Cetacea. CONCLUSION: In conjunction with a recent study of Primates, we find no evidence to support an association between MCPH1 evolution and the evolution of brain size in highly encephalized mammalian species. Our finding of significant positive selection in MCPH1 may be linked to other functions of the gene.

Occipital ossification of balaenopteroid mysticetes.

The bones of the posterior portion of the mammalian skull often exhibit incomplete ossification of the joints between the bones at the time of birth, with complete ossification at some point after birth. The sequence of ossification of these joints in mysticetes can be used to characterize the relative age in the calf and early juvenile ontogenetic stages. This study examined occipital joints ossification of 38 dry prepared neonate specimens in four mysticete species from two families (Eschrichtiidae: Eschrichtius robustus; Balaenopteridae: Balaenoptera acutorostrata, Balaenoptera physalus, and Megaptera novaeangliae). Each of the joints responsible for the fusion of the occiput were examined and rated for degree of ossification. The cranial ossification analysis indicates that E. robustus calves have open occipital joints until ~6 months of age and are born at a less mature stage than closely related balaenopterids. All of the species followed the same sequence of ossification: basioccipital/exoccipital joint, followed by the basioccipital/basisphenoid joint, and completed by the supraoccipital/exoccipital joint.

Movements of the bottlenose Dolphin (Tursiops truncatus) in the Rio de Janeiro State, Southeastern Brazil / Deslocamentos do golfinho-nariz-de-garrafa (Tursiops truncatus) no estado do Rio de Janeiro, sudeste do Brasil

Aiming to verify the movements of the bottlenose dolphin (Tursiops truncatus) at Rio de Janeiro State coast, southeastern Brazil, we performed a photoidentification comparison between the catalogued individuals of the Cagarras Archipelago (23º 02' S and 43º 12' W) in 2004 and 2006 (n = 26) and the images obtained (n = 179) during the Southeastern Cetaceans Expedition, conducted during months of June and November of 2005. Eight individuals (three females and five dolphins of unknown gender) identified in the Cagarras Archipelago were resighted in the Grande Island (23º 21'S and 44º 15' W), about 100 km southwestwards from Cagarras Archipelago. The observed movements include distances commonly recorded for the species elsewhere and are probably related to search for prey.

Com o objetivo de verificar os deslocamentos do golfinho-nariz-de-garrafa (Tursiops truncatus) no estado do Rio de Janeiro, sudeste do Brasil, foi feita uma análise entre os indivíduos catalogados no arquipélago das Cagarras em 2004 e 2006 (n = 26) e as fotografias (n = 179) obtidas durante a Expedição Cetáceos do Sudeste, realizada em junho e novembro de 2005. Oito indivíduos (três fêmeas e cinco golfinhos de sexo indeterminado) identificados no arquipélago das Cagarras (23º 02' S e 43º 12' W) foram reavistados na Ilha Grande (23º 21'S e 44º 15' W), aproximadamente 100 km a sudoeste do arquipélago. Os deslocamentos observados estão dentro das distâncias comumente registradas para a espécie e, provavelmente, são relacionados com a busca de recursos alimentares.

Diversity of cetaceans as tool in monitoring environmental impacts of seismic surveys / Diversidade de cetáceos como ferramenta no monitoramento dos impactos ambientais das sísmicas marítimas

New information about the effects of seismic surveys on cetaceans is causing increasing concern about the impact of this type of activity on marine life. The effects described include behavioral responses and changes in vocalization patterns, diversion of migratory routes, damage to the auditory system, and an increase in strandings. Although such effects could affect the diversity of species in areas where seismic research has been carried out, there is no scientific information on this subject. This study aims to evaluate the relationship between seismic surveys, oceanographic data and diversity of cetaceans recorded in Brazil following the stepping up of seismic survey activities between 1999 and 2004. The study is based on oceanographic data from the Pilot Research Moored Array in the Tropical Atlantic (PIRATA Project), sightings made during seismic surveys, progress reports from Brazilian research projects to the International Whaling Commission, Brazilian seismic survey reports available at the Escritório de Licenciamento de Petróleo e Nuclear of the Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis (ELPN-Ibama) and complementary data from the webpage of the Agencia Nacional de Petróleo e Gás Natural (ANP). The results suggest a decrease in the diversity of species in the face of an increase in the number of seismic surveys during the years 2000 and 2001, even though there was no significant change in oceanographic patterns in this period, and that a relationship exists between diversity of cetaceans and intensity of seismic surveys between 1999 and 2004. It is recommended that data collection be improved in order to evaluate this hypothesis properly. The results suggest that species diversity might be used as a long-term indicator of the impact of seismic surveys on cetaceans.

Tem sido constante o surgimento de novas informações sobre o efeito das sísmicas marítimas em cetáceos, demonstrando uma crescente preocupação com essa atividade. Os efeitos da atividade incluem respostas comportamentais e mudanças nos padrões de vocalização, alterações nas rotas migratórias, danos no sistema auditivo e aumento dos encalhes. Embora todos esses efeitos possam afetar a diversidade de espécies nas áreas de realização de sísmicas marítimas, não existem informações acadêmicas nesse sentido. Este estudo buscou verificar as relações entre as sísmicas marítimas, dados oceanográficos e a diversidade de cetáceos registrados no Brasil após a intensificação das operações a partir de 1999 até o ano 2004. Foram utilizados dados de observações de cetáceos realizadas durante as sísmicas marítimas e complementadas por relatórios de progresso do Brasil enviados para a Comissão Internacional Baleeira, dados oceanográficos disponibilizados pelo projeto Pilot Research Moored Array in the Tropical Atlantic (PIRATA) e informações de sísmicas marítimas arquivadas no Escritório de Licenciamento de Petróleo e Nuclear do Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Sustentáveis (ELPN-Ibama) e outras disponibilizadas na página da Internet da Agencia Nacional de Petróleo e Gás Natural (ANP). Os resultados indicam decréscimo na diversidade de espécies durante os anos 2000 e 2001, enquanto uma intensificação das sísmicas marítimas foi registrada para os mesmos anos. Os dados oceanográficos não apresentaram variações significativas entre os anos estudados. Os resultados sugerem alguma relação de efeito e causa entre a diversidade de cetáceos e a intensidade de sísmicas marítimas ocorridas no Brasil entre 1999 e 2004. O aprimoramento na coleta de dados com foco mais específico para avaliar os efeitos imediatos na diversidade é recomendado para melhor avaliação da hipótese e utilização da diversidade de espécies como indicador de longo...

Fishes associated with spinner dolphins at Fernando de Noronha Archipelago, tropical Western Atlantic: an update and overview

An update is presented for fish species associated with spinner dolphins at Fernando de Noronha Archipelago, tropical Western Atlantic, providing a general view of their diversity. The associates are mostly reef-dwelling fishes that feed on the dolphin wastes. Twelve species are habitual or occasional plankton-eaters and two species are herbivores that occasionally forage on floating pieces of algae. One species is a strict carnivore, one species is a hitchhiker that forages on a variety of foods including parasites and dead tissue from the dolphins, and one species is a carnivore that joins the dolphin groups to forage on schools of small fishes or squids. We predict that the list of fish associated with spinner dolphins will expand mostly with addition of habitual or occasional plankton-eaters.

Uma atualização é apresentada sobre as espécies de peixes associados a golfinhos-rotadores no Arquipélago de Fernando de Noronha, Atlântico Ocidental, fornecendo uma visão geral sobre sua diversidade. Os peixes associados são principalmente habitantes recifais que se alimentam de dejetos dos golfinhos. Doze espécies são planctófagas habituais ou ocasionais e duas são herbívoras que se alimentam ocasionalmente de algas à deriva. Uma espécie é carnívora estrita, uma vive fixada aos golfinhos e seu alimento é variado, incluindo parasitos e tecido morto dos golfinhos, e uma é carnívora que se junta aos grupos de golfinhos durante a caça a peixes e lulas. Prevemos que a lista de peixes associados a golfinhos-rotadores seja expandida principalmente com adição de planctófagos habituais ou ocasionais.

Bycatch of marine mammals in U.S. and global fisheries.

Fisheries bycatch poses a significant threat to many populations of marine mammals, but there are few published estimates of the magnitude of these catches. We estimated marine mammal bycatch in U.S. fisheries from 1990 to 1999 with data taken from the stock assessment reports required by the U.S. Marine Mammal Protection Act. The mean annual bycatch of marine mammals during this period was 6215 +/- 448 (SE). Bycatch of cetaceans and pinnipeds occurred in similar numbers. Most cetacean (84%) and pinniped (98%) bycatch occurred in gill-net fisheries. Marine mammal bycatch declined significantly over the decade, primarily because of a reduction in the bycatch of cetaceans. Total marine mammal bycatch was significantly lower after the implementation of take reduction measures in the latter half of the decade. We derived a crude first estimate of marine mammal bycatch in the world's fisheries by expanding U.S. bycatch with data on fleet composition from the Food and Agriculture Organization. The global bycatch of marine mammals is in the hundreds of thousands. Bycatch is likely to have significant demographic effects on many populations of marine mammals. Better data are urgently needed to fully understand the impact of these interactions.

Development of the cetacean nasal skull.

The adaptation of cetaceans to aquatic life habits is reflected, in their nasal region, in three marked changes from the original relations found in land mammals. These changes include (1) the loss of the sense of smell, (2) translocation of the nostrils from the tip of the rostrum to the vertex of the head, and (3) elongation of the anterior head to form a rostrum protruding far towards anterior. The morphogenetic processes taking place during embryogenesis of the nasal skull play a decisive part in the development of all these changes. The lateral parts of the embryonic nasal capsule, encompassing the nasal passages, change their position from horizontal to vertical. At the same time, the structures of the original nasal floor (the solum nasi) are shifted in front of the nasal passages towards the rostrum. The structures of the original nasal roof (the tectum nasi) and of the nasal side wall (the paries nasi) are translocated behind the nasal passages towards the neurocranium. The medial nasal septum (the septum nasi) mostly loses its connection to the nasal passages and is produced into a point protruding far towards anterior. The transformed embryonic nasal skull of the Cetacea can be divided into three sections: 1. The median structures. These include the cartilaginous structures, viz., the rostrum nasi, the septum interorbitale and the spina mesethmoidalis, which are accompanied by the dermal bones, the vomer and the praemaxillare. In adult cetaceans the rostrum nasi is mostly preserved as a robust cartilage of the skull, which may possibly serve as a sound transmitting structure of the sonar system, or it may be responsible for the sensing of water streams and vibrations. 2. The posterior side wall structures. These include the following cartilaginous structures that are mostly heavily reduced or mutually fused: the cupula nasi anterior, the tectum nasi, the lamina cribrosa, the paries nasi, the commissura orbitonasalis, the cupula nasi posterior, the processus paraseptalis posterior, the crista semicircularis, the frontoturbinale, the ethmoturbinale I and the maxilloturbinale. The cartilaginous structures are largely accompanied by the dermal bone, the maxillare. Of these embryonic elements, very little is preserved in adult cetaceans. The cartilages of the cupula nasi anterior form the variable skeleton around the nostrils. In Physeter the tectum nasi forms a very long cartilaginous bar that passes through the whole giant anterior head of the sperm whale as a structure accompanying the left nasal passage. 3. The anterior side wall structures. These include the cartilaginous structures, viz., the cartilago ductus nasopalatini, the cartilago paraseptalis, the processus lateralis ventralis and the lamina transversalis anterior, accompanied by the dermal bones, the praemaxillare and the vomer. These structures participate in the formation of the robust rostrum of the cetacean skull, and they are partly preserved even in adults in the form of the isolated ossa pararostralia (the Meckelian ossicles). The comparison of morphogeny of the nasal skull has also made it possible to draw certain conclusions on the phylogeny and systematics of Cetacea. Already the earliest embryonic stages permit us to discern weighty transformations of the original nasal skull of land mammals. These transformations are common to all embryos examined. This fact indicates a common origin of all Cetacea, which thus form a single monophyletic order. However, later embryonic stages show some different modifications of the nasal capsule according to which at least three major groups can be distinguished within the order Cetacea, probably ranking as superfamilies: Balaenopteroidea, Physeteroidea and Delphinoidea. Our observations, being in full accordance with other morphological, and embryological, as well as molecular biological results, suggest that the division of the order Cetacea into two suborders, Mysticeti and Odontoceti, is no longer tenable.

[Comparative analysis of the neocortex during the ontogenesis of cetaceae and primates]. / Sravnitel'nyi analiz novoi kory mozga v ontogeneze kitoobraznykh i primatov.

Comparative ontogenetic investigation of cytoarchitectonics of the cerebral neocortex has been performed in Cetacea and Primates using paraffin frontal and sagittal cerebral sections stained after Nissl. Cerebral hemispheres of dolphins, whales, monkeys and human being have been studied at various periods of prenatal development and in mature individuals. The comparison has been made at similar stages of cytoarchitectonical differentiation of the cortical plate. At two first stages of the prenatal ontogenesis (formation of the cortical plate and its differentiation into layers) there is not any principle differences between the Cetacea and Primates. Peculiarities of the cerebral cortical plate differentiation in the Cetacea (absence of the internal granular layer IV) is determined at the stage of stratification. Similar agranular character of the cerebral cortex differentiation is maintained during the whole subsequent ontogenesis in the Cetacea (heterogenetic type of the neocortex after Brodman). Absence of the layer IV in the cerebral neocortex determines some other principles in the spatial organization of the cortical-subcortical and in the intracortical connections in the Cetacea brain. This is confirmed by modern data of morphological and electrophysiological investigations. Perhaps, a comparatively more simple initial architectonics of the Cetacea brain limited the level of their functional possibilities, the latter is comparable only with anthropoid apes.

[Comparison of growth functions as exemplified by the southern smooth whale Eubalaena australis]. / Vergleich von Wachstumsfunktionen am Beispiel des Südlichen Glattwals (Eubalaena australis).

6 year's observations of the southern right whale in the San José Golf off Argentina by an American research team have yielded graphical growth curves. As the values are in good agreement with older results from 1908 to 1969 mathematical approximations are performed giving interesting results and allowing for a better insight into the properties of the growth functions applied. The different course of 3 functions as gained by nonlinear regressions is demonstrated by a graph. The results for 9 growth functions are combined in tables together with the values of the parameters and characteristic data.

Roentgenographic indicators of skeletal maturity in marine mammals (Cetacea).

A new roentgenographic classification (grading) scheme is presented for utilization in studies of skeletal development and maturation in marine mammals, particularly cetaceans. This is based on adequate description of the extent of development and maturation of the various secondary ossification centers, their eventual patterns of fusion, and subsequent remodeling with the metaphysis. The six stages are illustrated schematically and roentgenographically. This scheme may be applied to any cetacean longitudinal bone developing proximal and distal epiphyseal ossification centers.

Heart weight as a function of body weight in mammals.

Linear regression analysis was carried out on logarithmically transformed heart weight and body weight data in 104 mammalian species. It was shown that heart weight varies as the 0.98 power of body weight over essentially the whole mammalian weight range. The coefficient of correlation between heart weight and body weight is 0.99. Student's test was employed to compare the slopes of the several regression lines as between male and female animals, and as between terrestrial and aquatic mammals. In neither case were the differences in slope found to be statistically significant (df greater than 177, p less than 0.5).