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Gross and microscopic visceral anatomy of the male Cape fur seal, Arctocephalus pusillus pusillus (Pinnipedia: Otariidae), with reference to organ size and growth.

The gross and microscopic anatomy of the Cape fur seal heart, lung, liver, spleen, stomach, intestine and kidneys (n = 31 seals) is described. Absolute and relative size of organs from 30 male seals are presented, with histological examination conducted on 7 animals. The relationship between log body weight, log organ weight and age was investigated using linear regression. Twenty five animals were of known age, while 6 were aged from counts of incremental lines observed in the dentine of tooth sections. For the range of ages represented in this study, body weight changes were accurately described by the exponential growth equation, weight = w(o)r(t), with body weight increasing by 23 % per annum until at least 9-10 y of age. Organ weight increased at a rate of between 25 % and 33 % per annum until at least 9-10 y of age, with the exception of the intestines, where exponential increase appeared to have ceased by about 7 y. The relationship between body weight and organ weight was investigated using logarithmic transformations of the allometric equation, y = ax(b), where the exponent b is 1 if organ weight is proportional to body weight. Most organs increased in proportion to the body. However, the heart, liver and spleen had exponents b > 1, suggesting that these organs increased at a faster rate than the body. The basic anatomical features of the viscera were similar to those of other pinnipeds, with some exceptions, including the arrangement of the multilobed lung and liver. Apart from the large liver and kidneys, relative size of the organs did not differ greatly from similar sized terrestrial carnivores. The histological features of the organs were generally consistent with those previously described for this species and other otariids. The heart, as in other pinnipeds, was unlike that of cetacea in not having unusually thick endocardium or prominent Purkinje cells. Notable histological features of the lungs included prominent fibrous septa, prominent smooth muscle bundles, cartilage extending to the level of the alveolar sacs and ample lymphoid tissue. The spleen had a thick capsule, well developed trabeculae and plentiful plasma cells. Abundant parietal cells were present in the fundic glands and lymphoid follicles were present in the gastric lamina propria, particularly in the pyloric region. Small intestinal villi were very long but this could have resulted from underlying chronic inflammation. Lymphoid follicles were prominent in the colon. The kidney reniculi each had a complete cortex, medulla and calyx, but a sportaperi medullaris musculosa was not identified.

Animals↗

On the development of Cetacean extremities: I. Hind limb rudimentation in the Spotted dolphin (Stenella attenuata).

The Cetacea are group of animals which have completely lost their hind limbs during the course of evolution as a result of their entirely aquatic mode of life. It is known, however, that during their embryonal period, the hind limb buds are temporarily present. The control mechanisms of this regression are not yet understood, and vestigial limbs can sometimes be found in adults. The aim of the present study is to describe the course of hind limb rudimentation during prenatal development of Stenella attenuata (Spotted dolphin) at tissue and cell levels and compare the results with other natural or experimentally induced amelias. Hind limb buds of dolphin embryos, CRL 10-30 mm, were examined histologically. Before total disappearance, they show histodifferentiation comparable with other mammals. Initially, they form the apical ectodermal ridge, which soon regresses. The mesenchyme undergoes the process of condensation to form anlagens of prospective skeletal elements. These condensations are surrounded by vascular plexuses. During the course of rudimentation, some mesenchymal cells die, while the others are incorporated into the body wall. Nerve ingrowth into rudimentary limb buds was also detected. The temporary presence of hind limb rudiments in cetacean embryos can be regarded as a good example of recapitulation of phylogenesis in ontogenesis.

Animals↗

On the development of Cetacean extremities: II. Morphogenesis and histogenesis of the flippers in the spotted dolphin (Stenella attenuata).

Externally, the flippers of Cetacea resemble fish fins, but their internal structure is entirely mammalian. They show, however, some adaptative deviations from the typical pattern of the mammalian extremities, the most striking of which is an increased number of phalanges. The aim of this study is to describe the course of the development of flippers in the spotted dolphin (Stenella attenuata) and compare its features with other similar species from an evolutionary perspective. Early stages of flipper development were studied histologically. Differentiation of cartilaginous anlagens of the skeleton progresses proximodistally, condensation in digital rays being evident sooner than chondrogenesis in the carpal region. In one specimen, the temporary presence of cartilaginous rudiments of two carpal elements, which are not found in adults, was observed. At all examined stages, phalangeal number progressively increases up to (radial to ulnar) 3, 7, 7, 5, 3 in the most advanced stage. The reason for this condition is the specialised function of these limb-like structures. It is a classical example of convergence, in which mammalian extremities change their form to emulate the fin function. A similar condition is found in another group of originally terrestrial animals secondarily fully adapted to the aquatic mode of life-Ichyosauria (Reptilia).

Adaptation, Physiological↗

Mitochondrial phylogenetics and evolution of mysticete whales.

The phylogenetic relationships among baleen whales (Order: Cetacea) remain uncertain despite extensive research in cetacean molecular phylogenetics and a potential morphological sample size of over 2 million animals harvested. Questions remain regarding the number of species and the monophyly of genera, as well as higher order relationships. Here, we approach mysticete phylogeny with complete mitochondrial genome sequence analysis. We determined complete mtDNA sequences of 10 extant Mysticeti species, inferred their phylogenetic relationships, and estimated node divergence times. The mtDNA sequence analysis concurs with previous molecular studies in the ordering of the principal branches, with Balaenidae (right whales) as sister to all other mysticetes base, followed by Neobalaenidae (pygmy right whale), Eschrichtiidae (gray whale), and finally Balaenopteridae (rorquals + humpback whale). The mtDNA analysis further suggests that four lineages exist within the clade of Eschrichtiidae + Balaenopteridae, including a sister relationship between the humpback and fin whales, and a monophyletic group formed by the blue, sei, and Bryde's whales, each of which represents a newly recognized phylogenetic relationship in Mysticeti. We also estimated the divergence times of all extant mysticete species, accounting for evolutionary rate heterogeneity among lineages. When the mtDNA divergence estimates are compared with the mysticete fossil record, several lineages have molecular divergence estimates strikingly older than indicated by paleontological data. We suggest this discrepancy reflects both a large amount of ancestral polymorphism and long generation times of ancestral baleen whale populations.

Animals↗

A comprehensive and validated molecular taxonomy of beaked whales, family Ziphiidae.

DNA sequences from orthologous loci can provide universal characters for taxonomic identification. Molecular taxonomy is of particular value for groups in which distinctive morphological features are difficult to observe or compare. To assist in species identification for the little known family Ziphiidae (beaked whales), we compiled a reference database of mitochondrial DNA (mtDNA) control region (437 bp) and cytochrome b (384 bp) sequences for all 21 described species in this group. This mtDNA database is complemented by a nuclear database of actin intron sequences (925 bp) for 17 of the 21 species. All reference sequences were derived from specimens validated by diagnostic skeletal material or other documentation, and included four holotypes. Phylogenetic analyses of mtDNA sequences confirmed the genetic distinctiveness of all beaked whale species currently recognized. Both mitochondrial loci were well suited for species identification, with reference sequences for all known ziphiids forming robust species-specific clades in phylogenetic reconstructions. The majority of species were also distinguished by nuclear alleles. Phylogenetic comparison of sequence data from "test" specimens to these reference databases resulted in three major taxonomic discoveries involving animals previously misclassified from morphology. Based on our experience with this family and the order Cetacea as a whole, we suggest that a molecular taxonomy should consider the following components: comprehensiveness, validation, locus sensitivity, genetic distinctiveness and exclusivity, concordance, and universal accessibility and curation.

Actins↗

Acidic glycolipids from dolphin kidney.

The composition and contents of acidic glycolipids in the kidney of a striped dolphin (Stenella coeruleoalba, the order Cetacea, whales) were determined. The following eight acidic glycolipids were isolated and characterized: SM4s (124.2 nmol/g tissue), SM3 (8.7), GM3 (NeuAc) (12.3), GM3 (NeuGc) (31.6), GD3 (NeuAc-NeuAc) (14.7), GD3 (NeuAc-NeuGc) (II3 alpha(NeuAc alpha 2-8NeuGc)-LacCer) (9.8), GD3 (NeuGc-NeuAc) (II3 alpha(NeuGc alpha 2-8NeuAc)-LacCer) (5.3), and GD3 (NeuGc-NeuGc) (15.8). The assignment of the four types of GD3 was further confirmed as described below. Evidence indicating 2-8 linkages of the disialosyl residues of GD3 was obtained on methylation analysis of sialic acid. GD3 (NeuAc-NeuAc) and GD3 (NeuGc-NeuAc) were degraded to GM3 (NeuAc), and GD3 (NeuAc-NeuGc) and GD3 (NeuGc-NeuGc) yielded GM3 (NeuGc) on mild acid hydrolysis. Fragment ions characteristic of the carbohydrate and lipophilic moieties of the permethylated GD3 were observed in direct inlet-electron impact-mass spectra. The presence of these four types of GD3 in a tissue has not been reported previously. GD3 contained non-hydroxy (69-84%) and hydroxy fatty acids (16-31%) with 16-24 carbons. The long chain base of all GD3, except GD3 (NeuGc-NeuAc) (not analyzed), consisted of 4-sphingenine (d18:1) and 4-hydroxysphinganine (t18:0) in almost equal amounts. The total amount of renal lipid-bound acidic groups (sulfate and sialic acid) of the dolphin (190 mumol/animal) is considerably higher than that of a terrestrial mammal (88 mumol/animal) with a body weight comparable to that of the dolphin. This deviation suggests that the amount of renal acidic amphiphiles required to maintain the osmotic balance of body fluids in marine mammals might be higher than that in terrestrial ones.

Animals↗

The phylogenetic position of the Talpidae within eutheria based on analysis of complete mitochondrial sequences.

The complete mitochondrial (mt) genome of the mole Talpa europaea was sequenced and included in phylogenetic analyses together with another lipotyphlan (insectivore) species, the hedgehog Erinaceus europaeus, and 22 other eutherian species plus three outgroup taxa (two marsupials and a monotreme). The phylogenetic analyses reconstructed a sister group relationship between the mole and fruit bat Artibeus jamaicensis (order Chiroptera). The Talpa/Artibeus clade constitutes a sister clade of the cetferungulates, a clade including Cetacea, Artiodactyla, Perissodactyla, and Carnivora. A monophyletic relationship between the hedgehog and the mole was significantly rejected by maximum parsimony and maximum likelihood. Consistent with current systematic schemes, analyses of complete cytochrome b genes including the shrew Sorex araneus (family Soricidae) revealed a close relationship between Talpidae and Soricidae. The analyses of complete mtDNAs, along with the findings of other insectivore studies, challenge the maintenance of the order Lipotyphla as a taxonomic unit and support the elevation of the Soricomorpha (with the families Talpidae and Soricidae and possibly also the Solenodontidae and Tenrecidae) to the level of an order, as previously proposed in some morphological studies.

Animals↗

Beaked whales echolocate on prey.

Beaked whales (Cetacea: Ziphiidea) of the genera Ziphius and Mesoplodon are so difficult to study that they are mostly known from strandings. How these elusive toothed whales use and react to sound is of concern because they mass strand during naval sonar exercises. A new non-invasive acoustic ording tag was attached to four beaked whales(two Mesoplodon densirostris and two Ziphius cavirostris) and recorded high-frequency clicks during deep dives. The tagged whales only clicked at depths below 200 m, down to a maximum depth of 1267 m. Both species produced a large number of short, directional, ultrasonic clicks with significant energy below 20 kHz. The tags recorded echoes from prey items; to our knowledge, a first for any animal echolocating in the wild. As far as we are aware, these echoes provide the first direct evidence on how free-ranging toothed whales use echolocation in foraging. The strength of these echoes suggests that the source level of Mesoplodon clicks is in the range of 200-220 dB re 1 microPa at 1 m. This paper presents conclusive data on the normal vocalizations of these beaked whale species, which may enable acoustic monitoring to mitigate exposure to sounds intense enough to harm them.

Animals↗

Comparative morphology of the tapetum lucidum (among selected species).

OBJECTIVES: The phenomenon of 'eye-shine' is seen in a variety of animal species, and is generally thought to be related to the presence of an intraocular reflecting structure, the tapetum lucidum. The tapetum lucidum is a biologic reflector system that is a common feature in the eyes of vertebrates. It normally functions to provide the light-sensitive retinal cells with a second opportunity for photon-photoreceptor stimulation, thereby enhancing visual sensitivity at low light levels. The tapetum lucidum is presented here according to a classification based on the location, as well as the composition, of this reflective layer. Finally, the physical and chemical properties, as well as the origins of the different tapeta lucida, are discussed and compared. METHODS: The anatomic and biochemical aspects of the tapetum lucidum in various vertebrates are examined. Morphologic observations were made from paraffin and plastic embedded specimens. Specimens were treated with traditional stains and observed by light and transmission electron microscopy. RESULTS: Some species (primates, squirrels, birds, red kangaroo and pig) do not have this structure and they usually are diurnal animals. In vertebrates, the tapetum lucidum exhibits diverse structure, organization and composition. Therefore, the retinal tapetum (teleosts, crocodilians, marsupials, fruit bat), the choroidal guanine tapetum (elasmobranchs), the choroidal tapetum cellulosum (carnivores, rodents, cetacea), and the choroidal tapetum fibrosum (cow, sheep, goat, horse) are described. CONCLUSIONS: The tapetum lucidum represents a remarkable example of neural cell and tissue specialization as an adaptation to a dim light environment and, despite these differences, all tapetal variants act to increase retinal sensitivity by reflecting light back through the photoreceptor layer. These variations regarding both its location and structure, as well as the choice of reflective material, may represent selective visual adaptations associated with their feeding behavior, in response to the use of specific wavelengths and amount of reflectance required.

Animals↗

Simulating the effect of high-intensity sound on cetaceans: modeling approach and a case study for Cuvier's beaked whale (Ziphius cavirostris).

A finite element model is formulated to study the steady-state vibration response of the anatomy of a whale (Cetacea) submerged in seawater. The anatomy was reconstructed from a combination of two-dimensional (2D) computed tomography (CT) scan images, identification of Hounsfield units with tissue types, and mapping of mechanical properties. A partial differential equation model describes the motion of the tissues within a Lagrangean framework. The computational model was applied to the study of the response of the tissues within the head of a neonate Cuvier's beaked whale Ziphius cavirostris. The characteristics of the sound stimulus was a continuous wave excitation at 3500 Hz and 180 dB re: 1 microPa received level, incident as a plane wave. We model the beaked whale tissues embedded within a volume of seawater. To account for the finite dimensions of the computational volume, we increased the damping for viscous shear stresses within the water volume, in an attempt to reduce the contribution of waves reflected from the boundaries of the computational box. The mechanical response of the tissues was simulated including: strain amplitude; dissipated power; and pressure. The tissues are not likely to suffer direct mechanical or thermal damage, within the range of parameters tested.

Acoustics↗

Origin of whales from early artiodactyls: hands and feet of Eocene Protocetidae from Pakistan.

Partial skeletons of two new fossil whales, Artiocetus clavis and Rodhocetus balochistanensis, are among the oldest known protocetid archaeocetes. These came from early Lutetian age (47 million years ago) strata in eastern Balochistan Province, Pakistan. Both have an astragalus and cuboid in the ankle with characteristics diagnostic of artiodactyls; R. balochistanensis has virtually complete fore- and hind limbs. The new skeletons are important in augmenting the diversity of early Protocetidae, clarifying that Cetacea evolved from early Artiodactyla rather than Mesonychia and showing how early protocetids swam.

Animals↗

Chromosome homologies between man and mountain zebra (Equus zebra hartmannae) and description of a new ancestral synteny involving sequences homologous to human chromosomes 4 and 8.

Using human chromosome painting probes, we looked for homologies between human and mountain zebra (Equus zebra hartmannae, Equidae, Perissodactyla) karyotypes. Except for two very short segments, all euchromatic regions were found to have a human homologous chromosome segment. Conserved syntenies previously described in various mammalian orders were detected. Each synteny corresponded to a chromosomal region homologous to two parts of human chromosomes: HSA3 and HSA21, HSA7 and HSA16, HSA12 and HSA22, and HSA16 and HSA19. Chromosomal segments homologous to a part of HSA11 and HSA19p are found syntenic in zebra, horse and donkey, suggesting that this group of synteny has been inherited from an Equidae or Perissodactyla common ancestor. A synteny of segments homologous to parts of HSA4 and HSA8 was observed in zebra and horse. It also exists in the rabbit (Lagomorpha) and several Carnivora species. A second group of taxa which does not have this region of synteny is composed of primates, Chiroptera and Insectivora, and possibly also Cetacea and Scandantia. Thus, the presence or absence of this region of synteny may separate two groups of eutherian mammals.

Animals↗

Mitogenomic analyses of eutherian relationships.

Reasonably correct phylogenies are fundamental to the testing of evolutionary hypotheses. Here, we present phylogenetic findings based on analyses of 67 complete mammalian mitochondrial (mt) genomes. The analyses, irrespective of whether they were performed at the amino acid (aa) level or on nucleotides (nt) of first and second codon positions, placed Erinaceomorpha (hedgehogs and their kin) as the sister group of remaining eutherians. Thus, the analyses separated Erinaceomorpha from other traditional lipotyphlans (e.g., tenrecs, moles, and shrews), making traditional Lipotyphla polyphyletic. Both the aa and nt data sets identified the two order-rich eutherian clades, the Cetferungulata (comprising Pholidota, Carnivora, Perissodactyla, Artiodactyla, and Cetacea) and the African clade (Tenrecomorpha, Macroscelidea, Tubulidentata, Hyracoidea, Proboscidea, and Sirenia). The study corroborated recent findings that have identified a sister-group relationship between Anthropoidea and Dermoptera (flying lemurs), thereby making our own order, Primates, a paraphyletic assembly. Molecular estimates using paleontologically well-established calibration points, placed the origin of most eutherian orders in Cretaceous times, 70-100 million years before present (MYBP). The same estimates place all primate divergences much earlier than traditionally believed. For example, the divergence between Homo and Pan is estimated to have taken place approximately 10 MYBP, a dating consistent with recent findings in primate paleontology.

Animals↗

Phylogeny through brain traits: more characters for the analysis of mammalian evolution.

We have assembled data on nine brain traits, in addition to the fifteen we have previously described, which provide new evidence for assessing mammalian relationships. States of these characters are tabulated as they occur in each of 152 mammalian species, providing data in numerically ordered form, useful for multiple analyses of phylogenetic relationships in programs which take into account variations in several different characters simultaneously. Derived states of each of the nine traits are characteristic of certain restricted groups of mammals; (1) mirroring of the complete SI body representation in isocortex (anthropoid primates); (2) loss of the accessory olfactory bulbs (sirenians, cetaceans, most bats, catarrhine primates); (3) Rindenkerne, clumps of cell bodies in layer 6 of cerebral cortex (sirenians); (4) posteriorly-pointing digits in the SI body representation (bats, both mega- and micro-); (5) equivalent tectopetal connections to the anterior colliculus of one side from both retinas, rather than predominantly from the contralateral retina (primates and megabats); (6) loss of lamination in dorsal cochlear nuclei (anthropoid primates, bats, seals, sirenians, cetaceans); (7) separation of claustrum from cerebral cortex (diprotodont marsupials, carnivores, artiodactyls, perissodactyls, hyracoids, cetaceans and primates), (8) presence of a complete secondary (SII) somatic sensory region of cerebral cortex (therians-all extant mammals other than monotremes), and (9) presence of a distinct external cuneate nucleus among the nuclei of the dorsal columns (all mammalian groups except monotremes and sirenians). Two examples of phylogenetic trees derived from these data are presented. These sample trees maintain the segregation of the monotremes and the marsupials, and the basic dichotomy of placentals seen in our earlier trees based entirely on brain data. They also show: an orderly sequence of bifurcations (rather than the commonly seen multifurcation near the base of the radiation) in the reconstruction of placental relationships; extremes of derivation for the Cetacea, the Chiroptera, and the Sirenia (in concordance with trees based on other data); a ferungulate association of Carnivora, Perissodactyla, Artiodactyla, Hyracoidea and Sirenia; and an assemblage of related Dermoptera, Primates, Scandentia, and Chiroptera which in this model also includes Insectivora and Macroscelidea. Analyses based on brain characters can reinforce conclusions based on other data, while at the same time introducing new ideas about relationships. Neural traits provide a source of data independent of those commonly used in phylogenetic analysis, and are extremely valuable for testing old hypotheses and for introducing new ones.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Size and shape of the cerebral cortex in mammals. II. The cortical volume.

The geometry of the brain and cerebral cortex in mammals has been studied from an evolutionary perspective and is described in mathematical terms. The volume of the cerebral cortex, in contrast to the cortical surface area, scales to brain volume in a similar way, irrespective of the degree of cortical folding. Among mammals, Cetacea form a subgroup, in that their volumetric data fit an isometric model better than an allometric model. An index of corticalization is presented which contains information about both the mass of interconnective nerve fibers and the degree of intracortical processing. It is shown, furthermore, that a semilogarithmic equation appropriately describes the relationship between mean cortical thickness and brain volume. Finally, allometric equations between brain volume and cortical parameters, which can be used for predictive purposes, are presented.

Animals↗

Size and shape of the cerebral cortex in mammals. I. The cortical surface.

The evolution of the brain in mammals has been accompanied by a progressive enlargement of the cerebral cortex. Allometric analysis of the volume, surface and convolutedness of this cortex shows that among mammals two major groups can be distinguished: (1) species with lissencephalic brains, where surface-volume relationships are determined by linear geometric laws, and (2) species with gyrencephalic brains, where strict geometric similarity no longer applies. Different mathematical models are required to describe the brain morphology in these groups. It has been shown, moreover, that among species with convoluted brains, marine mammals (Cetacea) form a subgroup in that the cerebrocortical surface in these animals is more folded than in terrestrial mammals of similar brain size. Thus it appears that the use of a single allometric relation in studying the effect of size increase on the geometry of the brain - irrespective of convolutedness or ecological strategy - as has been done in previous studies, is unjustified and obscures the differences in the structural organization of mammalian brains. Finally, a dimensionless index of cortical folding is proposed, which indicates that the degree of cortical folding depends not only on the volume of the brain or cortex but on cortical thickness as well.

Animals↗

The West Indian manatee (Trichechus manatus) lacks a vomeronasal organ.

Completely aquatic marine mammals of the order Cetacea such as whales and dolphins have a reduced or absent olfactory system and neither a vomeronasal organ nor an accessory olfactory bulb. In comparison, seals, which are only partially aquatic, have olfactory and accessory olfactory systems including the vomeronasal organ. Thus, there seems to be a strong relation between the degree of adaptation to an aquatic environment and the degree of reduction in olfactory structures. Sirenians, such as manatees and dugongs, are another family of marine mammals which have secondarily adapted to a fully aquatic existence, yet there is dispute about the status of their olfactory structures. In the present study there was no evidence for a vomeronasal organ in the adult West Indian Manatee, Trichechus manatus. Additionally, the main olfactory system appeared quite rudimentary. These observations support the hypothesis that, in mammals, secondary adaptation to an aquatic environment leads to the reduction or loss of the olfactory senses.

Adaptation, Biological↗

Histopathologic and immunocytochemical studies of distemper in harbor porpoises.

During 1988 thousands of harbor seals (Phoca vitulina) died in European seas as a result of morbillivirus infection. Six harbor porpoises (Phocoena phocoena) found stranded on the coast of Northern Ireland in late 1988 were submitted to our laboratory for necropsy. Pneumonia was the main necropsy finding in three of these animals. Microscopic lung lesions characterized by necrosis of bronchial and bronchiolar epithelium and infiltration of alveoli with leukocytes, lymphoid cells, macrophages, and multinucleate syncytia were seen in all six porpoises. Cytoplasmic and nuclear acidophilic inclusions characteristic of morbillivirus infection were common in bronchial and bronchiolar epithelial cells and in alveolar macrophages and syncytia. Brain alterations included degeneration and necrosis of neurons, microglial infiltration, and perivascular cuffing. There were cytoplasmic and nuclear acidophilic inclusions in many neurons. Immunoperoxidase staining of morbillivirus antigen was seen in many tissues including lung, brain, spleen, and urinary bladder. Alterations in our porpoises were similar to those seen in distemper in seals and many species of terrestrial mammals. Systemic viral disease has not previously been documented in Cetacea.

Animals↗