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Comparative morphology and evolution of the otic region in toothed whales (Cetacea, Mammalia).

The otic region in the skull of archeocetes and odontocetes is compared and interpreted with special emphasis on the morphology and suspension of the ear bones. In archeocetes, the periotic was obviously separate from the mastoid but still integrated within the skull via a long anterior and posterior process. The rotation of the cochlear part of the periotic was already obvious. The tympanic bone was attached to a decreasing number of neighboring elements, with the periotic becoming more and more important in the later archeocetes. The accessory air sacs of the tympanic cavity had invaded some of the adjacent skeletal elements and attained a moderate-to-remarkable extension. In the evolution of the odontocetes, the periotic and tympanic were successively uncoupled from the skull and combined to a new morphological and functional unit (tympanoperiotic complex). This uncoupling was mainly achieved by shortening the periotical processes and simultaneously extending the tympanic air sacs. For functional reasons, however, the periotic (posterior process) stayed in immediate contact with the mastoid, the latter remaining in the lateral wall of skull. In advanced marine dolphins, the bony sheaths of the accessory air sacs are largely reduced, presumably because of volume fluctuations in the tympanic cavity during diving. The perfect uncoupling of the ear bones from the skull obviously was an essential prerequisite for directional hearing, for effective ultrasound orientation and communication, and finally, for the striking development of the dolphin brain.

Animals↗

Studies on the fetal development of the gubernaculum in cetacea.

BACKGROUND: Adult cetacean males, like non-mammalian vertebrates and other testicond mammals, have intra-abdominal testes. There is no evidence of a processus vaginalis in them. Testicondia in cetaceans is considered secondary as they are judged, evolutionarily, the descendants of terrestrial mammals (ungulates) with testis descent. A possible argument in support of the latter contention would be that cetacean fetuses develop gubernacula which are the primordia of the processus vaginalis and other structures associated with testis descent in other placental mammals. The present study intended to analyse cetacean fetuses in this respect. METHODS: Serial sections of 25 fetuses (total body length between 39.5 and 160 mm) of 4 cetacean species (Delphinus delphis, Phocoena phocoena, Eschrichtius robustus, Physeter catodon) were examined with special attention to the presence or absence of structures homologous to the gubernaculum of other placental mammals (rats and humans). RESULTS: Gubernacular primordia were observed in fetuses from about the time of onset of sexual differentiation. Their shape and anatomical relationship with the surrounding structures were similar as those in mammals with testis descent. The gubernaculum in males developed into a large mass of dense connective tissue in the ventral-caudal abdominal region at the site of the insertion of the mesonephric inguinal ligament and associated to the tip of the internal abdominal oblique muscle. No (or only very little) development of a processus vaginalis was noticed. CONCLUSIONS: The results demonstrate initial emergence of mammalian-like gubernacular primordia in cetacean fetuses without their further development to elaborate structures required for testis descent. The findings support the view that cetaceans are secondarily testicond. It is suggested that (1) absence of the pelvic girdle together with (2) the development of structures in and beyond the caudal abdominal region, particularly the caudal hypaxial musculature, precludes the outgrowth, into caudal direction, of hollow organs (such as the processus vaginalis) from the abdominal cavity.

Animals↗

Arterial circulation of the spinal cord and brain in the Monodontidae (order Cetacea).

In this paper we document retial supply of the spinal cord and describe the arterial vascular pattern of the brain in the whale family Monodontidae. Observations are based on gross dissections of four brains, two each of Monodon monoceros and Delphinapterus leucas, and one spinal cord from M. monoceros. Vessels of the spinal cord arise from extradural retia in the neural canal. Arteries originating from the retia penetrate the dura between successive spinal roots (mainly ventral) and not in association with them, unlike radicular arteries of other mammals. Also, these vessels are uniformly distributed and contribute equally to a plexus surrounding the cord. An A. radicularis magna is not present, and neither are distinct anterior or posterior spinal arteries. Circulation to the brain is effected by two pairs of arteries originating from intracranial retia. The rostral pair supplies most of the forebrain (prosencephalon), whereas the more caudal pair supplies mainly the midbrain (mesencephalon) and hindbrain (rhombencephalon). The circulatory pattern is characterized by 1) complete independence of anterior cerebral arteries (no anastomoses); 2) extensive cortical supply by the anterior choroidal arteries; 3) absence of subdural communicating vessels between rostral and caudal trunks; 4) union of caudal trunks to form a small basilar artery; and 5) absence of vertebral arteries and hence of a vertebral basilar system. There are some obvious differences between subdural arteries in the Monodontidae and those in other mammals; however, their general patterns of distribution are similar, and we suggest that most of the vessels, at least in the cranium, are homologous.

Animals↗

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.

Age Determination by Skeleton↗

The dinosaurian origin of feathers: perspectives from dolphin (Cetacea) collagen fibers.

The early origin of birds is a hotly disputed debate and may be broadly framed as a conflict between paleontologists and ornithologists. The paleontological emphasis has shifted from Archaeopteryx and its origins to recent finds of Cretaceous birds and "feathered" dinosaurs from China. The identification of alleged feathers has, however, relied principally on the visual image. Some workers have interpreted these integumentary structures as collagen fibers. To test the latter hypothesis, using light microscopy, collagen from the hypodermis (blubber) and subdermal connective tissue sheath was examined from a dolphin that had been buried for a year as part of an experiment. Within the blubber, toward the central thicker parts of the material, the collagen fibers had compacted and the three-dimensional latticework of normal blubber had more or less collapsed. Chromatographic analysis of the blubber revealed pronounced oxidation of the unsaturated lipids, probably accounting for the collapse of the latticework. Fibers normally bound together in bundles became separated into individual fibers or smaller bundles by degradation of the glue-like substance binding them together. These degraded collagen fibers show, in many instances, feather-like patterns, strikingly reminiscent of many of those identified as either "protofeathers" or "modern" feathers in dromaeosaurid dinosaurs. The findings throw serious doubt on the virtually complete reliance on visual image by supporters of the feathered dinosaur thesis and emphasize the need for more rigorous methods of identification using modern feathers as a frame of reference. Since collagen is the main fiber type found in most supporting tissues, the results have wide implications regarding the degradation and fossilization of vertebrate integument, such as that of the ichthyosaurs, dinosaurs and birds.

Adipose Tissue↗

Feeding kinematics of Kogia and Tursiops (Odontoceti: Cetacea): characterization of suction and ram feeding.

The feeding kinematics of dwarf and pygmy sperm whales (Kogia sima and K. breviceps) and bottlenose dolphins Tursiops truncatus were characterized and compared incorporating the Ram-Suction Index (RSI). Mean RSI data support a suction feeding strategy for Kogia (-0.67+/-0.29; mean +/- s.d.) and a ram feeding strategy for Tursiops (0.94+/-0.11; mean +/- s.d.). Tursiops displayed two ram-based feeding behaviours: open gape approach, where gape was at least 50% of maximum in the first video field, and closed gape approach, where gape increased near food items. Four feeding phases were identified in both odontocetes: preparatory, jaw opening, gular depression and jaw closing. The mean Kogia feeding cycle duration (470+/-139 ms) was significantly shorter (P<0.003) than all Tursiops groups (pooled: 863+/-337 ms; open gape approach: 1211+/-207 ms; closed gape approach: 662+/-207 ms). Kogia mean maximum gape angle (39.8+/-18.9 degrees), mean maximum opening and closing gape angle velocities (293+/-261 deg. s(-1) and 223+/-121 deg. s(-1), respectively) were significantly greater (P<0.005) than pooled Tursiops mean maximum gape angle (24.8+/-6.6 degrees), mean maximum opening and closing gape angle velocities (84+/-56 deg. s(-1) and 120+/-54 deg. s(-1), respectively). Negative Kogia RSI values were correlated with increasing maximum gular depression and retraction, wide gape angle, and rapid opening gape angle velocity. Kinematic data support functional hypotheses that odontocetes generate suction by rapid depression of the hyoid and tongue.

Analysis of Variance↗