Morphology and development of the cranium of Felis silvestris f. catus Linné 1758--a contribution to the comparative anatomy of the Carnivora. III: Ear region and occipital region

In Felis, the otic region of younger embryonic stages up to Felis 1 is characterized by extremely medial extended cochleae, compressing the basal plate to a slender trabeculum. As a result of a quite strong rostrad convergence of the long axis of the ear capsules, the Commissura praefacialis fuses with the Commissura orbitoparietalis laterally. Until now, this has been found in whales only. Continuing embryogenesis towards Felis 2, the cochlea moves laterally and slightly ventrally, so the angle of convergence between the whole Capsula otica and the skull base decreases. The problem of interpreting these positional changes of the Capsula otica during phylogenesis and ontogenesis is discussed in detail. Up to recent literature, there is a discussion about homology of the Foramen perilymphaticum and allied structures in reptiles referring to the openings in the Capsula otica in mammals. Configuration of these structures in fissiped carnivores and the appearance of a "limitating membrane" in Felis 2, gives reason for a new discussion of these problems. Composition of the Bulla tympani is a very important feature for investigation of phylogeny and systematics in fissiped carnivores. In Felis 2, there appears a caudal entotympanic, consisting of young cartilaginous tissue. The development of the caudal entotympanic has impact on 2 structures in the occipital region: The Lamina alaris and the Processus paracondyloideus. Felis 1 shows a distinct Lamina alaris and a short Processus paracondyloideus. With Felis 2, either element is reduced largely, probably to the extent as the caudal entotympanic develops.     

Measurements of the tympanic cavity

The angle between the annulus fibrocartilaginous and the superior crest of the petrous bone was measured with 14 (10 to 19) degrees. The height of the annulus was in our material 10.04 mm, the width 9.45 mm. Measured were also the length of the plicae malleares and the height of the recessus membrane tympani anterior, posterior et superior with (2.5, 3.0 and 1.5 mm). In about 74%, the chorda tympani was imbedded in different zones of the plicae malleares. Its entrance and exit areas, the length of its intratympanal course and its width were measured. Included are also values of the ligaments of the otic ossicles, measurements of the ostium tympanicum tubae auditivae, of the epitympanic recess, the aditus ad antrum, and the antrum mastoideum. The eminentia pyramidalis, the sinus posterior and the sinus tympani, ponticuli and subicula, the fossula fenestrae cochleae and the fenestra cochleae, the fenestra vestibuli and prominences of the facial canal in the lateral semicircular canal have been researched and measured. Also the tympanic nerve and its course on the promontorium have been estimated.     

The braincase of Typhlops and Leptotyphlops (Reptilia: Serpentes)

The structure of the braincases and associated nerves and blood-vessels is described. Typhlops combines primitive lacertilian features, notably the retention of a palatine artery, with specialisations such as the structure of the recessus scalae tympani. Leptotyphlops is more completely snake-like, but the two genera show in common a specialised intracranial course of the hyomandibular branch of the facial nerve, due to lateral closure of the juxtastapedial recess by the overgrowing crista circumfenestralis. The lateral closure of the juxtastapedial recess is considered as an adaptation to burrowing habits. The Vidian canal in scolecophidians is poorly defined, due to the lack of lateral ascending wings of the parasphenoid. This might constitute an archaic character contradicting the descent of snakes from any Recent lacertilian group.     

Ontogeny and cranial morphology of the tympanic region of the Tupaiidae, with special reference to Ptilocercus

The structure of the tympanic region of the skull of Ptilocercus lowii was studied in an embryo of 30 mm crown-rump length and in 5 osteocrania. As in Tupaia, the anterior wall of the bulla of Ptilocercus is not completed by a tympanic process of the alisphenoid, contrary to earlier reports. Ptilocercus resembles Tupaia in the following derived characters. The ventral wall of the tympanic cavity is formed by a rostral entotympanic and by a caudal tympanic process of the petrosal. The entotympanic develops in primary connection with the tubal cartilage. The tympanic aperture of the auditory tube is bordered by the entotympanic. The ring-shaped tympanicum is covered by the entotympanicum and is aphaneric. The musculus tensor tympani is lacking. Among mammals, these characters can be regarded as synapomorphic for the Tupaiidae, that is, to have been present in the common ancestor of the two subfamilies. From the evidence of the tympanic region, the Tupaiidae, therefore, form a monophyletic group. Besides these synapomorphies, there are remarkable differences between Ptilocercus and Tupaia in the structure of the bulla. In Ptilocercus the bulla is smaller and less pneumatized than in Tupaia. An anterior intrabullar septum, present in Tupaia, is lacking in Ptilocercus. The epitympanic wing of the alisphenoid is smaller in Ptilocercus than in Tupaia. A lateral prefacial commissure of the tegmen tympani is present in Ptilocercus, but absent in Tupaia. The caudal tympanic process of the petrosal is larger in Ptilocercus than in Tupaia. These characters are autapomorphic for the Ptilocercinae and for the Tupaiinae, respectively. They demonstrate that the auditory bulla of Ptilocercus and that of Tupaia have evolved independently to a considerable extent. An early phylogenetic separation of their respective ancestors seems likely. The tympanic region of the skull provides no evidence for close relationships of the tree shrews to the primates or to any other eutherians. The classification of the Tupaiidae in a separate order, Scandentia, is supported.     

Morphogenesis of the cranium of Cavia porcellus L. I: Introduction, classification and descriptive part

The craniogenesis of Cavia porcellus has been investigated in 7 embryos of different ages. From the developmental stage of 36 mm CRL, a reconstruction of the chondrocranium is described. As there is still a gap in the tectum posterius, the foramen magnum is not closed posteriorly. The course of the notochord is retrobasilar at the beginning, more rostrally it runs intrabasilar. After ossification, the place where the notochord enters the basioccipital is marked by a cavity. This cavity is not identical with the posterior basicranial fenestra. The laminae alares build up strong paracondylar processes. The auditory capsule is connected with the occipital region only by the exoccipitocapsular commissure, but there is no supraoccipitocapsular commissure. The fossa subarcuata is pierced by a subarcuate foramen. The very short lamina parietalis is not in contact with the orbital wing as there is no orbitoparietal commissure. The processus recessus divides the perilymphatic foramen into fenestra rotunda and aquaeductus cochleae. The suprafacial commissure arises from the upper margin of the canalicular part of the auditory capsule, but it does not reach the superior surface of the cochlear part at this stage. The tegmen tympani is well developed. The foramen singulare is not yet separated from the inferior acoustic foramen. At the stage of 36 mm CRL there is no septum spirale cartilagineum. Mm. tensor tympani et stapedius are developed in the typical way. The auditory capsule is connected with the basal plate by the anterior basicochlear and the alicochlear commissures. The floor of the orbitotemporal region is incomplete, the temporal wings are small. The alicochlear commissure, the alar process, the anterior basicochlear commissure, and the basal plate surround the carotid foramen, but there is no blood vessel passing through it. The temporal wing is at the beginning of ossification and shows a fissura ovalis for the mandibular nerve. Although the hypophysial canal is still present, there are no structures passing through it. There is no foramen rotundum. The orbital wing is still cartilagineous. The basal plate and the pre- and postoptic roots border the optical foramen. The straight muscles of the eyeball are attached to the ala hypochiasmatica which projects from the postoptic root. The interorbital septum is absent. The ophthalmic artery, which instead of the obliterated internal carotid artery, contributes to the circulus arteriosus cerebri is well developed. There is no orbitonasal commissure . The epiphanial foramina are present.(ABSTRACT TRUNCATED AT 400 WORDS)     

The evolution of the basicranium in the Henophidia (Reptilia: Serpentes)

In lizards, a short Vidian canal pierces the base of the basipterygoid process. In snakes, the anterior opening of the primitively short Vidian canal lies on the dorsal surface of the basisphenoid, trapped in an intracranial position by downgrowths of the frontal and parietal which meet the lateral edges of the basisphenoid-parasphenoid complex. This condition is observed in anilioid snakes which retain other primitive features in the braincase: the paroccipital process and the spheno-occipital tubercle (Aniliidae only) and participation of the basioccipital in the apertura lateralis of the recessus scalae tympani.
Subsequent evolution within booid snakes shows a shift of the anterior opening of the Vidian canal around the anterior edge of the basisphenoid, thus acquiring a secondary extracranial position. This occurs in parallel within boine and pythonine snakes. Dinilysia shows a parallel development of die condition observed in advanced booid snakes. Pseudoboa , with its short Vidian canal opening in-tracranially, demonstrates that caenophidians originated from a basal henophidian or pre-henophidian stock. The Acrochordidae show a basicranium that can be interpreted as either primitive henophidian or primitive caenophidian.     

The skull of the Upper Cretaceous snake Dinilysia patagonica Smith‐Woodward, 1901, and its phylogenetic position revisited

The cranial anatomy of Dinilysia patagonica, a terrestrial snake from the Upper Cretaceous of Argentina, is redescribed and illustrated, based on high‐resolution X‐ray computed tomography and better preparations made on previously known specimens, including the holotype. Previously unreported characters reinforce the intriguing mosaic nature of the skull of Dinilysia, with a suite of plesiomorphic and apomorphic characters with respect to extant snakes. Newly recognized plesiomorphies are the absence of the medial vertical flange of the nasal, lateral position of the prefrontal, lizard‐like contact between vomer and palatine, floor of the recessus scalae tympani formed by the basioccipital, posterolateral corners of the basisphenoid strongly ventrolaterally projected, and absence of a medial parietal pillar separating the telencephalon and mesencephalon, amongst others. We also reinterpreted the structures forming the otic region of Dinilysia, confirming the presence of a crista circumfenestralis, which represents an important derived ophidian synapomorphy. Both plesiomorphic and apomorphic traits of Dinilysia are treated in detail and illustrated accordingly. Results of a phylogenetic analysis support a basal position of Dinilysia, as the sister‐taxon to all extant snakes. The fossil taxa Yurlunggur, Haasiophis, Eupodophis, Pachyrhachis, and Wonambi appear as derived snakes nested within the extant clade Alethinophidia, as stem‐taxa to the crown‐clade Macrostomata. The hypothesis of a sister‐group relationship between Dinilysia and Najash rionegrina, as suggested by some authors, is rejected by the results of our analysis. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 194–238.     

Cloning of full-length coding sequence of tree shrew CD3E and prediction of its molecular characteristics

The use of tree shrews (Tupaia belangeri) in human disease studies demands essential research tools, in particular cellular markers and their monoclonal antibodies for immunological studies. Here we cloned the full-length cDNAs encoding CD3E from total RNA of the spleen, liver and peripheral blood of tree shrews and analyzed their structural characteristics in comparison with other mammals by Discovery Studio software. The results showed that the open reading frame sequence of tree shrew CD3E was 582 bp, encoding 194 amino acids. The overall structure of tree shrew CD3E protein was similar to its counterparts of other mammals, intracellular and transmembrane domain highly conserved. However, detailed analysis revealed two potential glycosylation sites and different surface charges in the extracellular domain. Availability of the entire open-reading-frame and related sequence information would therefore facilitate the preparation of monoclonal antibodies against tree shrew CD3 and further studies for its function.     

Evolution of the tetrapod ear: an analysis and reinterpretation

The dominant view of tetrapod otic evolution–the “standard view”–holds that the tympanum developed very early in tetrapod history and is homologous in all tetrapods and that the opercular process of the rhipidistian hyomandibula is homologous to the tympanic process of the stapes in lower tetrapods. Under that view, the labyrinthodont amphibians of the Paleozoic are usually considered ancestral to reptiles, and thus the “otic notch” of labyrinthodonts and the tympanum it presumably contained form the starting-point for middle ear evolution in reptiles. Four problems have classically been identified with the standard view: the differing relationships of the internal mandibular branch of N. VII (chorda tympani) to the processes of the stapes in amniotes and anurans; the differing orientations of the stapes in key fossil and living groups; the location of the tympanum in early fossil reptiles; and the transferral of the tympanum, during the origin of mammals, from the stapes to the articular bone of the lower jaw. An examination of these problems and of the solutions proposed under the standard view reveals the ad hoc, and therefore unsatisfactory, nature of the proposed solutions. To organize and review alternative hypotheses of otic evolution an analytical table is constructed, using three characters (tympanic process, Nerve VII, tympanum), each with two possible states. A total of eight hypotheses about middle ear evolution are possible under this system, one of which is the standard view. The seven “non-standard” hypotheses, only five of which have been argued in the literature, are briefly examined. Six of the “non-standard” hypotheses appear unattractive for various reasons, including reliance on ad hoc arguments. The seventh was first proposed by Gaupp in 1898. It is today almost universally ignored but apparently largely for historical rather than scientific reasons. This hypothesis, her called the “alternative view”, appears to rest on assumptions equally as plausible as those of the standard view. Moreover, it offers a solution of the problems associated with the standard view without, apparently, raising any similarly serious problems. This paper compares the standard and alternative views of middle ear evolution in detail. Comparison proceeds on two levels. On one level, they are compared in terms of the hypotheses of phyletic tetrapod relationships each promotes and how strongly each supports its hypothesis. Both views promote the same hypothesis of tetrapod relationships. The alternative view is the more parsimonious, but the difference is not considered sufficient to provide a choice. On another level, the two views are compared in terms of their implications for: (1) the evolution of relative and absolute auditory perceptive ability; (2) the origin of reptiles; (3) the evolution of the suspensorium and cranial kinesis; and (4) the origin and evolution of recent amphibians. The nature of the data required for a test of the implications of the two views is specified in each case. Where data are available. the alternative view is consistent and the standard view is inconsistent with these data. We conclude that the alternative view is the preferable hypothesis of middle-ear evolution. This conclusion implies the following: the tympanic membranes and the tympanic processes of the stapes in recent mammals, reptiles + birds. and frogs. are not homologous; the evolution of “special periotic systems” in the ancestors of amphibians and amniotes were independent events and preceded the evolution of tympanic membranes; the amphibian tympanic membrane. probably including that of labyrinthodonts. is not ancestral to that of amniotes. and that labyiinthodonts with an otic notch are not suitable as amniote ancestors; the stapes of early reptiles functioned primarily as part of the jaw suspension rather than in hearing; the mechanisms and abilities of sound perception in recent tetrapods are likely to be diverse rather than forming parts of a cline; and the lack of a tympanum in Gymnophiona and Caudata may be a retention of a primitive condition.     

Morphogenesis of the nasal apparatus of the red deer (Cervus elaphus L.)

Three stages of morphogenesis of the nasal apparatus of the red deer (Cervus elaphus L.) were studied. Many ancestral traits reminiscent of relationships in other mammals and even in reptiles were found, including a cart, ectochoanalis, paraseptal cartilages, the septum nasi and its ventral trabecular enlargement, a lamina transversalis ant., clear separation of the cart, parietotectalis and cart, paranasalis from each other and a crista semicircularis. A maxilloturbinale, was present, but not a nasoturbinale. The main specific features were a completely rostrally localized, peculiar cartilaginous structure in the preseptal space, for which there is as yet no morphological explanation, and pronounced bulging of the cartilaginous wall of the nasal capsule in a ventrolateral direction, level with the rostral region of the olfactory labyrinth (caudally to the aboral end of the maxilloturbinale). In the early stages of morphogenesis, it was found that the ethmoturbinalia might be formed by fusion of the edges of the anlagen of the paranasal cartilage and the lamina orbitonasalis. The structure of the olfactory labyrinth was reminiscent of its organization in the sheep embryo; the recessus frontalis was completed by a series of frontoturbinal recesses and frontoturbinalia, which are poorly developed in the red deer, however. The floor of the caudal part of the nasal capsule was very little developed and there was no cart, paraseptalis post.     

Structure and postnatal development of photoreceptors and their synapses in the retina of the tree shrew (Tupaia belangen)

Summary The all cone retina of the tree shrew (Tupaia belangeri) was examined in the adult and early postnatal stages by light and electron microscopy. Rods are not as rare as previously thought, but make up about 4% of the photoreceptors. They are relatively short and narrow cells, which stain (toluidine blue) more intensively and lie more proximal than cones. Among the cones three morphological varieties could be distinguished. Most cones stain lightly but have a light or a dark giant mitochondrion in their inner segment; a third type stains darker but occurs only rarely. All cones possess extensive radial processes (lateral fins) around the basal part of their inner segments. Such fins are well known from reptiles and birds, but have only once been described in a mammal (gray squirrel).The maturation of the retina in Tupaia belangeri proceeds centrifugally, i.e., from the vitreal to the scleral side, as in most mammals. A few synapses are already present at birth in the outer and inner plexiform layers, but seem to be more advanced in the latter. Such early synapses are small and have only few synaptic vesicles; they appear almost mature by day 14. The light-sensitive outer segments develop last. The first disks are seen by day 10, but regular membrane stacks are only present by day 18. Thus, it seems that the retina is functional when the young first open their eyes, which occurs around day 18.     

Homology between the recessus lateralis and cephalic sensory canals, with the proposition of additional synapomorphies for the Clupeiformes and the Clupeoidei

The recessus lateralis , a complex structure in the otic region of the skull that is probably associated with detection and analysis of small vibrational pressures and displacements, is widely recognized as a synapomorphy of the Clupeiformes. The Clupeiformes includes the Denticipitoidei, with one living species, Denticeps clupeoides , and the Clupeoidei, with about 360 living species commonly known as herrings and anchovies. Comparisons between details of the recessus lateralis of the Clupeoidei and Denticipitoidei, and the sensory cephalic canals of other teleosts, lead to hypotheses of a series of transformations of the cephalic sensory canals . Treating that complex as a single binary 'presence vs. absence' character as was traditional practice obscures important phylogenetically informative variation. Specific synapomorphies in that system exist for the Clupeiformes and the Clupeoidei. Hypothesized synapomorphies in the recessus lateralis for the Clupeiformes are the presence of a dilated internal temporal sensory canal in the pterotic, a postorbital branch of the supraorbital sensory canal located in a bony groove in the lateral wing of the frontal, and the terminal portions of preopercular and infraorbital sensory canals closely positioned. Hypothesized synapomorphies for the Clupeoidei are the presence of a postorbital branch of the supraorbital sensory canal located deep within the body of the lateral wing of the frontal, with the distal portion of that branch totally internal on the cranium, and the expanded distal portion of the postorbital branch of the supraorbital sensory canal. The homology of the sinus temporalis of Clupeoidei, and of the dermosphenotic of both Denticeps and the Clupeoidei, with those of other teleosts is also considered.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 257–270.     

Evolution of electrosensory ampullary organs: conservation of Eya4 expression during lateral line development in jawed vertebrates

The lateral line system of fishes and amphibians comprises two ancient sensory systems: mechanoreception and electroreception. Electroreception is found in all major vertebrate groups (i.e. jawless fishes, cartilaginous fishes, and bony fishes); however, it was lost in several groups including anuran amphibians (frogs) and amniotes (reptiles, birds, and mammals), as well as in the lineage leading to the neopterygian clade of bony fishes (bowfins, gars, and teleosts). Electroreception is mediated by modified “hair cells,” which are collected in ampullary organs that flank lines of mechanosensory hair cell containing neuromasts. In the axolotl (a urodele amphibian), grafting and ablation studies have shown a lateral line placode origin for both mechanosensory neuromasts and electrosensory ampullary organs (and the neurons that innervate them). However, little is known at the molecular level about the development of the amphibian lateral line system in general and electrosensory ampullary organs in particular. Previously, we identified Eya4 as a marker for lateral line (and otic) placodes, neuromasts, and ampullary organs in a shark (a cartilaginous fish) and a paddlefish (a basal ray‐finned fish). Here, we show that Eya4 is similarly expressed during otic and lateral line placode development in the axolotl (a representative of the lobe‐finned fish clade). Furthermore, Eya4 expression is specifically restricted to hair cells in both neuromasts and ampullary organs, as identified by coexpression with the calcium‐buffering protein Parvalbumin3. As well as identifying new molecular markers for amphibian mechanosensory and electrosensory hair cells, these data demonstrate that Eya4 is a conserved marker for lateral line placodes and their derivatives in all jawed vertebrates.     

Small Mammal Investigation in Spotted Fever Focus with DNA-Barcoding and Taxonomic Implications on Rodents Species from Hainan of China

Although mammals are a well-studied group of animals, making accurate field identification of small mammals is still complex because of morphological variation across developmental stages, color variation of pelages, and often damaged osteological and dental characteristics. In 2008, small mammals were collected for an epidemiological study of a spotted fever outbreak in Hainan, China. Ten species of small mammals were identified by morphological characters in the field, most using pelage color characters only. The study is extended here, in order to assess whether DNA barcoding would be suitable as an identification tool in these small mammals. Barcode clusters showed some incongruence with morphospecies, especially for some species of Rattus and Niviventer, so molecular delineation was carried out with an expanded dataset of combined cytochrome b (Cyt-b) and cytochrome c oxidase subunit I (COI) sequences. COI sequences were successfully amplified from 83% of collected mammals, but failed in all specimens of Suncus murinus, which were thus excluded in DNA barcoding analysis. Of note, ten molecular taxonomic units were found from samples of nine morphologically identified species. Accordingly, 11 species of small mammals were present in the investigated areas, including four Rattus species, three Niviventer species, Callosciurus erythraeus, Neohylomys hainanensis, Tupaia belangeri, and Suncus murinus. Based on the results of the phylogenetic and molecular delineation analyses, the systematic status of some rodent species should be redefined. R. rattus hainanicus and R. rattus sladeni are synonyms of R. andamanensis. R. losea from China and Southeast Asia comprises two independent species: R. losea and R. sakeratensis. Finally, the taxonomic status of three putative species of Niviventer should be further confirmed according to morphological, molecular and ecological characters.     

Co-localization of xenopsin and gastrin immunoreactivity in gastric antral G-cells

Studies indicating evidence for the presence of the amphibian octapeptide xenopsin in gastric mucosa of mammals prompted us to investigate the cellular localization of this peptide. Using the peroxidase-antiperoxidase method and a specific antiserum to xenopsin (Xen-7) on paraffin and adjacent semithin sections of gastric antral mucosa from man, dog, and Tupaia belangeri, we found numerous epithelial cells showing a specific positive immunoreaction. These cells were of typical pyramidal shape and could be classified as of the "open" type. Cell quantification in serial sections processed for xenopsin and gastrin immunoreactivity, respectively, revealed an identical number of cells per section and an identical distribution of these cells in the middle zone of the antral mucosa. Furthermore, adjacent semithin sections demonstrated the colocalization of xenopsin and gastrin immunoreactivity within the same G-cell. The xenopsin antiserum could be completely absorbed with synthetic xenopsin but not with gastrin. Preabsorption tests with neurotensin, a xenopsin related peptide, or with somatostatin, glucagon, and enkephalins gave no evidence for crossreactivity of the xenopsin antiserum with these peptides. It is concluded that gastric antral G-cells in addition to gastrin also contain the amphibian peptide xenopsin.