Morphological structure of the aortic wall in three Delphinid species with shallow or intermediate diving habits: Evidence for diving adaptation.

Some modifications in the vascular system of marine mammals provide adaptive advantages for diving. This study analyses the organisation of the aortic wall in dolphins, observing artery changes in volume and blood pressure for diving behaviour. Samples of three aortic segments (ascending, thoracic and abdominal) of three dolphin species were processed for histological and morphometric studies. The three dolphin species used, striped dolphin (Stenella coeruleoalba), Atlantic spotted dolphin (Stenella frontalis) and common dolphin (Delphinus delphis), have shallow or intermediate diving habits. Our results indicated that the components of the aortic wall of the dolphins had different dispositions in the three selected segments. The aortic wall decreased in thickness along its length due to a loss of the lamellar units in the tunica media and a thinning of the main elements of the lamellar units along the artery. The life stage had little influence on the thickness of the aortic wall except for the ascending aorta. The weight, body length, species or sex of the specimen did not significantly influence the thickness of the wall or the lamellar units. In summary, the histological and morphometric aortic structure in dolphins, in relation to the studied parameters, seems to be similar to that previously described of terrestrial mammals such as pigs, except for a larger difference in the proportion of lamellar units between the ascending and thoracic segments.

Diffusion tensor imaging of dolphin brains reveals direct auditory pathway to temporal lobe.

The brains of odontocetes (toothed whales) look grossly different from their terrestrial relatives. Because of their adaptation to the aquatic environment and their reliance on echolocation, the odontocetes' auditory system is both unique and crucial to their survival. Yet, scant data exist about the functional organization of the cetacean auditory system. A predominant hypothesis is that the primary auditory cortex lies in the suprasylvian gyrus along the vertex of the hemispheres, with this position induced by expansion of 'associative' regions in lateral and caudal directions. However, the precise location of the auditory cortex and its connections are still unknown. Here, we used a novel diffusion tensor imaging (DTI) sequence in archival post-mortem brains of a common dolphin (Delphinus delphis) and a pantropical dolphin (Stenella attenuata) to map their sensory and motor systems. Using thalamic parcellation based on traditionally defined regions for the primary visual (V1) and auditory cortex (A1), we found distinct regions of the thalamus connected to V1 and A1. But in addition to suprasylvian-A1, we report here, for the first time, the auditory cortex also exists in the temporal lobe, in a region near cetacean-A2 and possibly analogous to the primary auditory cortex in related terrestrial mammals (Artiodactyla). Using probabilistic tract tracing, we found a direct pathway from the inferior colliculus to the medial geniculate nucleus to the temporal lobe near the sylvian fissure. Our results demonstrate the feasibility of post-mortem DTI in archival specimens to answer basic questions in comparative neurobiology in a way that has not previously been possible and shows a link between the cetacean auditory system and those of terrestrial mammals. Given that fresh cetacean specimens are relatively rare, the ability to measure connectivity in archival specimens opens up a plethora of possibilities for investigating neuroanatomy in cetaceans and other species.

Cross-sectional anatomy, computed tomography and magnetic resonance imaging of the head of common dolphin (Delphinus delphis) and striped dolphin (Stenella coeruleoalba).

Computed tomography (CT) and low-field magnetic resonance imaging (MRI) were used to scan seven by-caught dolphin cadavers, belonging to two species: four common dolphins (Delphinus delphis) and three striped dolphins (Stenella coeruleoalba). CT and MRI were obtained with the animals in ventral recumbency. After the imaging procedures, six dolphins were frozen at -20°C and sliced in the same position they were examined. Not only CT and MRI scans, but also cross sections of the heads were obtained in three body planes: transverse (slices of 1 cm thickness) in three dolphins, sagittal (5 cm thickness) in two dolphins and dorsal (5 cm thickness) in two dolphins. Relevant anatomical structures were identified and labelled on each cross section, obtaining a comprehensive bi-dimensional topographical anatomy guide of the main features of the common and the striped dolphin head. Furthermore, the anatomical cross sections were compared with their corresponding CT and MRI images, allowing an imaging identification of most of the anatomical features. CT scans produced an excellent definition of the bony and air-filled structures, while MRI allowed us to successfully identify most of the soft tissue structures in the dolphin's head. This paper provides a detailed anatomical description of the head structures of common and striped dolphins and compares anatomical cross sections with CT and MRI scans, becoming a reference guide for the interpretation of imaging studies.

Hierarchical population structure and habitat differences in a highly mobile marine species: the Atlantic spotted dolphin.

Recent molecular studies have shown that highly mobile species with continuous distributions can exhibit fine-scale population structure. In this context, we assessed genetic structure within a marine species with high dispersal potential, the Atlantic spotted dolphin (Stenella frontalis). Using 19 microsatellite loci and mitochondrial control region sequences, population structure was investigated in the western North Atlantic, the Gulf of Mexico and the Azores Islands. Analyses of the microsatellite data identified four distinct genetic clusters, which were supported by the control region sequences. The highest level of divergence was seen between two clusters corresponding to previously described morphotypes that inhabit oceanic and shelf waters. The combined morphological and genetic evidence suggests these two lineages are on distinct evolutionary trajectories and could be considered distinct subspecies despite their parapatry. Further analysis of the continental shelf cluster resulted in three groups: animals inhabiting shelf waters in the western North Atlantic, the eastern Gulf of Mexico and the western Gulf of Mexico. Analyses of environmental data indicate the four genetic clusters inhabit distinct habitats in terms of depth and sea surface temperature. Contemporary dispersal rate estimates suggest all of these populations should be considered as distinct management units. Conversely, no significant genetic differentiation was observed between S. frontalis from offshore waters of the western North Atlantic and the Azores, which are separated by approximately 4500 km. Overall, the hierarchical structure observed within the Atlantic spotted dolphin shows that the biogeography of the species is complex because it is not shaped solely by geographic distance.

Cross-sectional anatomy, computed tomography and magnetic resonance imaging of the thoracic region of common dolphin (Delphinus delphis) and striped dolphin (Stenella coeruleoalba).

The aim of this study was to provide a detailed anatomical description of the thoracic region features in normal common (Delphinus delphis) and striped dolphins (Stenella coeruleoalba) and to compare anatomical cross-sections with computed tomography (CT) and magnetic resonance imaging (MRI) scans. CT and MRI were used to scan 7 very fresh by-caught dolphin cadavers: four common and three striped dolphins. Diagnostic images were obtained from dolphins in ventral recumbency, and after the examinations, six dolphins were frozen (-20°C) and sliced in the same position. As well as CT and MRI scans, cross-sections were obtained in the three body planes: transverse (slices of 1 cm thickness), sagittal (5 cm thickness) and dorsal (5 cm thickness). Relevant anatomical features of the thoracic region were identified and labelled on each section, obtaining a complete bi-dimensional atlas. Furthermore, we compared CT and MRI scans with anatomical cross-sections, and results provided a complete reference guide for the interpretation of imaging studies of common and striped dolphin's thoracic structures.

Small-scale estimation of relative abundance for the coastal spotted dolphins (Stenella attenuata) in Costa Rica: the effect of habitat and seasonality / Estimación a pequeña escala de la abundancia relativa del delfín manchado costero (Stenella attenuata) en Costa Rica: el efecto del hábitat y la estacionalidad

The coastal spotted dolphin (Stenella attenuata graffmani) is one of the most common species of dolphin in inshore Pacific waters of Costa Rica. We conducted surveys in protected waters of the Papagayo Gulf, Costa Rica, to determine relative abundance of dolphins in relation to environmental variables. We used Generalized Additive Models to investigate the influence of a particular set of environmental factors and determine inter-annual trends in relative abundance. School sizes ranged from 1 to 50 individuals ( mean 9.95, SD=10.28). The number of dolphins increased linearly with water depth and transparency, and non-linearly with the dissolved oxygen concentration. High variability in the relative abundance occurred during the dry season (January-April). A previous study on this population found that high number of groups are involved in foraging activities during the dry season. Seasonal changes in relative abundance probably are associated with food availability, a variable that we did not measure. Understanding local resident populations may have important implications for conservation and management strategies. Large-scale studies may overlook variables affecting the abundance of local resident populations that may be detected with studies on a smaller scale such as this one.

El delfín manchado costero (Stenella attenuata graffmani) es una de las especies de delfines mas comunes de las aguas costeras del Pacifico de Costa Rica. En este estudio realizamos muestreos dentro de las aguas protegidas del Golfo de Papagayo para determinar su abundancia relativa en relación a características físico-químicas de su hábitat. Usamos modelos aditivos generalizados para investigar la influencia de un juego de variables ambientales y determinar tendencias inter-anuales en la abundancia relativa. El tamaño de los grupos varió de 1 a 50 individuos (promedio 9.95, SD=10.28). La cantidad de delfines aumentó linealmente con la profundidad y claridad del agua, y de forma no lineal con el oxígeno disuelto. Durante la época seca (enero-abril) encontramos la mayor variabilidad en la abundancia relativa. Un estudio anterior sugiere que una cantidad importante de delfines se alimenta aquí en la época seca. Ello sugiere que los cambios estacionales en la abundancia relativa de delfines manchados costeros podría estar asociada con la disponibilidad de alimento, una variable que no medimos pero que podría explicar la variación observada debido a la naturaleza productiva estacional del área. Los estudios de poblaciones locales y residentes pueden tener un mayor impacto en estrategias de conservación y manejo. Por lo general, los estudios a mayor escala geográfica pueden pasar por alto variables importantes que afectan la abundancia local de poblaciones residentes, las cuales pueden ser detectadas en estudios de menor escala como el presente.

Development of the skull of the pantropical spotted dolphin (Stenella attenuata).

We describe the bony and cartilaginous structures of five fetal skulls of Stenella attenuata (pantropical spotted dolphin) specimens. The specimens represent early fetal life as suggested by the presence of rostral tactile hairs and the beginnings of skin pigmentation. These specimens exhibit the developmental order of ossification of the intramembranous and endochondral elements of the cranium as well as the functional and morphological development of specific cetacean anatomical adaptations. Detailed observations are presented on telescoping, nasal anatomy, and middle ear anatomy. The development of the middle ear ossicles, ectotympanic bone, and median nasal cartilage is of interest because in the adult these structures are morphologically different from those in land mammals. We follow specific cetacean morphological characteristics through fetal development to provide insight into the form and function of the cetacean body plan. Combining these data with fossil evidence, it is possible to overlie ontogenetic patterns and discern evolutionary patterns of the cetacean skull.

Bones as biofuel: a review of whale bone composition with implications for deep-sea biology and palaeoanthropology.

Whales are unique among vertebrates because of the enormous oil reserves held in their soft tissue and bone. These 'biofuel' stores have been used by humans from prehistoric times to more recent industrial-scale whaling. Deep-sea biologists have now discovered that the oily bones of dead whales on the seabed are also used by specialist and generalist scavenging communities, including many unique organisms recently described as new to science. In the context of both cetacean and deep-sea invertebrate biology, we review scientific knowledge on the oil content of bone from several of the great whale species: Balaenoptera musculus, Balaenoptera physalus, Balaenoptera borealis, Megaptera novaeangliae, Eschrichtius robustus, Physeter macrocephalus and the striped dolphin, Stenella coeruleoalba. We show that data collected by scientists over 50 years ago during the heyday of industrial whaling explain several interesting phenomena with regard to the decay of whale remains. Variations in the lipid content of bones from different parts of a whale correspond closely with recently observed differences in the taphonomy of deep-sea whale carcasses and observed biases in the frequency of whale bones at archaeological sites.

Histological structure of the adrenal gland of the bottlenose dolphin (Tursiops truncatus) and the striped dolphin (Stenella coeruleoalba) from the Adriatic Sea.

The structure of the adrenal gland was studied in 11 bottlenose dolphins (Tursiops truncatus), and five striped dolphins (Stenella coeruleoalba). These species are legally protected in Croatia. All examined animals died of natural causes and were found stranded along eastern Adriatic coast. In both species the adrenal gland consists of a cortex and a medulla; the cortex is divided into three zones. Whereas in the bottlenose dolphin, there is a zona arcuata which contains columnar cells arranged in the form of arches; in the striped dolphin this zone is replaced by zona glomerulosa containing rounded clusters of polygonal cells. In both species, the zona fasciculata consists of radially oriented cords of polygonal cells, whereas in zona reticularis cells are arranged in branching and anastomosing cords. The adrenal medulla in both species contains dark, epinephrine-secreting cells and light norepinephrine-secreting cells. Epinephrine-secreting cells are localized in the outer part of the medulla, whereas norepinephrine-secreting cells are found in the inner part, arranged in clusters and surrounded by septa of thin connective tissue. The gland is surrounded by a thick connective-tissue capsule, from where thick trabeculae extend towards the interior. In the bottlenose dolphin, group of cells resembling both medullar and cortical cells can be seen within the capsule; whereas only groups of cells resembling cortical cells are found within the capsule of the striped dolphin. In the bottlenose dolphin invagination of the adrenal cortex into the medulla is obvious as well as medullary protrusions extending through cortex to the connective tissue capsule.

Laminar and cytoarchitectonic features of the cerebral cortex in the Risso's dolphin (Grampus griseus), striped dolphin (Stenella coeruleoalba), and bottlenose dolphin (Tursiops truncatus).

The present investigation carried out Nissl, Klüver-Barrera, and Golgi studies of the cerebral cortex in three distinct genera of oceanic dolphins (Risso's dolphin, striped dolphin and bottlenose dolphin) to identify and classify cortical laminar and cytoarchitectonic structures in four distinct functional areas, including primary motor (M1), primary sensory (S1), primary visual (V1), and primary auditory (A1) cortices. The laminar and cytoarchitectonic organization of each of these cortical areas was similar among the three dolphin species. M1 was visualized as five-layer structure that included the molecular layer (layer I), external granular layer (layer II), external pyramidal layer (layer III), internal pyramidal layer (layer V), and fusiform layer (layer VI). The internal granular layer was absent. The cetacean sensory-related cortical areas S1, V1, and A1 were also found to have a five-layer organization comprising layers I, II, III, V and VI. In particular, A1 was characterized by the broadest layer I, layer II and developed band of pyramidal neurons in layers III (sublayers IIIa, IIIb and IIIc) and V. The patch organization consisting of the layer IIIb-pyramidal neurons was detected in the S1 and V1, but not in A1. The laminar patterns of V1 and S1 were similar, but the cytoarchitectonic structures of the two areas were different. V1 was characterized by a broader layer II than that of S1, and also contained the specialized pyramidal and multipolar stellate neurons in layers III and V.

A morphological and histological examination of the pan-tropical spotted dolphin (Stenella attenuata) and the spinner dolphin (Stenella longirostris) adrenal gland.

The morphology and histology of the cetacean adrenal gland are poorly understood. Therefore, this study examined 32 pairs of adrenal glands from 18 pan-tropical spotted dolphins (Stenella attenuata) and 14 spinner dolphins (Stenella longirostris). In both species, the cortex was pseudolobulated and contained a typical mammalian zonation. Medullary protrusions (0-3 per section) and a medullary band were identified in both species. For S. attenuata, no statistical differences were found in the cortex to medulla (CM) ratio or the percent cross-sectional area (PCA) of the adrenal glands compared with sex or sexual maturity. The mean CM ratio for S. attenuata was 2.34 and the PCA was 64.4% cortex, 29.4% medulla and 6.2%'other'. 'Other' indicates blood vessels, connective tissue and the gland capsule itself. For S. longirostris, there was no statistical difference in the CM ratio compared with sexual maturity. However, a statistical difference was found between the CM ratio and sex, suggesting sexual dimorphism (female CM ratio = 2.46 and males = 3.21). No statistical differences were found in the PCA of S. longirostris adrenal glands by sexual maturity. However, a statistical difference was found between the PCA by sex. Female S. longirostris adrenal glands consisted of 65.0% cortex, 27.3% medulla and 7.7% 'other', whereas male adrenal glands consisted of 71.7% cortex, 22.7% medulla and 5.6% 'other'.

Cranial variation in the pantropical spotted dolphin, Stenella attenuata , in the Pacific Ocean.

Cranial variation in 129 pantropical spotted dolphins (Stenella attenuata) from the western and eastern tropical Pacific was investigated morphometrically. This study revealed significant sexual dimorphism in the dolphin skulls. Differences between the genders were mostly recorded in the rostrum and braincase, which are wider and more robust in males. This sexual variation was interpreted in terms of differences in behavioral and life-history strategies between genders. Geographical variation among populations of the eastern tropical Pacific (ETP), Japan, and Taiwan was also significant, with different sexual patterns. While no clear trend in geographical variation was detected in males, there was a positive relationship in females between morphological differences and geographical distance. That is, Japanese and Taiwanese female specimens were morphologically more similar than revealed by other pairwise comparisons between them and the ETP specimens. Moreover, two canonical discriminant functions further discriminated the three populations for males and females, respectively. The two functions showed that the rostrum and braincase are the principal elements in discrimination of geographical variation in males, while only the rostrum is the major discriminant morph of geographical variation in females. In addition, tests based on Mahalanobis distance-squares were conducted to assign six specimens from the western tropical Pacific to the above three populations to clarify the similarities among them.

The external and middle ear of the striped dolphin Stenella coeruleoalba (Meyen 1833).

The ear of cetaceans, and especially the middle ear, is very different from that of terrestrial mammals and shows specific adaptations to diving. Our research, performed on six Striped dolphins (Stenella coeruleoalba) stranded along the Italian coasts of the Mediterranean Sea, concentrated on the morphology of the external and middle ear of this species. We report the findings using a proper Veterinary Anatomical Nomenclature and describe the characteristics of the auditory meatus of the external ear and the presence and morphology of the erectile tissue in the middle ear. Our anatomical and histological data highlight the structure and possible functions of the corpus cavernosum located in the middle ear of the Striped dolphin, and suggest a possible role for this structure in relation to pressure regulation during diving. Many of our observations indicate the existence of an internal regulatory system able to prevent barotraumas by regulating pressure and volume inside the middle ear cavity.

A morphological and 13C NMR study of the extramandibular fat bodies of the striped dolphin (Stenella coeruleoalba).

The molecular and histological structure of the fat bodies covering externally the posterolateral region of the jaw of the striped dolphin (Stenella coeruleoalba) was investigated by means of morphological and nuclear magnetic resonance techniques. The analyses of samples belonging to adult and juvenile individuals were performed with the aim of seeking the presence of age-related differences. In our study, the level of isovalerate (iso5:0) in the extramandibular fat of the juvenile individuals is comparable with those of the adult counterparts; conversely, longer isobranched fatty acids were detected in lower quantities in the juveniles together with a higher degree of unsaturation. The morphologic analyses revealed that, in both adults and juveniles, this fatty tissue is similar to univacuolar adipose tissue. However, in the juveniles, a muscular component was present, whereas only in adult subjects, enlarged and irregularly shaped cavities may be seen within the adipose tissue. These cavities, structurally organized as veins, may regulate blood flow in response to changing water temperature and stabilize thermal gradient within the jaw lipids. These data suggest that the molecular components and the histological organization can indicate a maturation of the organ with age that probably may reflect different sound reception properties.