Mechanical factors in the evolution of the mammalian secondary palate: a theoretical analysis

The secondary palate of mammals is a bony shelf that closes the ventral aspect of the rostrum. The rostrum, therefore, approximates to a tapered semicylindrical tube that is theoretically a mechanically efficient structure for resisting the forces of biting, including the more prolonged bouts of mastication typical of mammals. Certain mammal-like reptiles illustrate stages in the development of the palate in which the shelves projecting medially from each premaxilla and maxilla do not meet in the midline. We evaluate several geometric properties of sections through the rostrum of the American opossum (Didelphis virginiana). For loading at the incisors and canines, these properties indicate the structural strength and stiffness in both bending and torsion of the rostrum and of single maxillae. We then repeat the analysis but progressively omit segments of the palatal shelf, a procedure which simulates, in reverse, the evolutionary development of the structure. The results demonstrate that the secondary palate contributes significantly to the torsional strength and stiffness of the rostrum of Didelphis and to the strength of each maxilla in lateromedial bending. The major evolutionary implications of the results are that the rapid increase in rostral strength with small increments of the palatal shelves may have been a significant factor in the development of the complete structure. The results indicate that there was a marked jump in torsional strength and stiffness when the shelves met in the midline, which is likely to have been important in the subsequent development of the diverse masticatory mechanisms of cynodonts and mammals. On the basis of this analysis the mammalian secondary palate may be interpreted as one of a number of methods, seen in the mammal-like reptiles, for strengthening the rostrum.     

The skull of Morganucodon

Morganucodon is a triconodont (atherian) mammal from the Lower Jurassic. Two species are described: M. oehleri from China and M. watsoni from Wales. The skull in M. walsoni is 26 mm long; M. oehleri is slightly larger. The dentition is differentiated functionally into incisors, canines, premolars and molars. The pineal foramen is closed. The prefrontals, postfrontals and postorbitals are lost. Septomaxilla, quadratojugal, tabular and pterygoid flanges are retained. The bony external nares are unpaired. The nasal cavity had the mammalian complement of turbinals. The posterior palate has ridges and troughs similar to those in tritylodonts, triconodonts and multituberculates. The alisphenoid ascending process is narrow and is not in contact with the anterior lamina of the petrosal, lying lateral to it. There is a cavum epiptericum, as in late therapsids. The anterior lamina forms the lateral braincase wall, perforated by the foramina pseudovale and pseudorotundum. There is a squamosal-dentary articulation, but the reptilian jaw joint is retained. The ear resembles that in later therapsids, with the tympanum in the lower jaw. The small quadrate was moveable, buttressed medially be a large stapes. Sound conduction from the tympanum was via articular, quadrate and stapes. The systematic position of Morganucodon is discussed.     

Development of the supralaryngeal vocal tract in Japanese macaques: implications for the evolution of the descent of the larynx

The configuration of the supralaryngeal vocal tract depends on the nonuniform growth of the oral and pharyngeal portion. The human pharynx develops to form a unique configuration, with the epiglottis losing contact with the velum. This configuration develops from the great descent of the larynx relative to the palate, which is accomplished through both the descent of the laryngeal skeleton relative to the hyoid and the descent of the hyoid relative to the palate. Chimpanzees show both processes of laryngeal descent, as in humans, but the evolutionary path before the divergence of the human and chimpanzee lineages is unclear. The development of laryngeal descent in six living Japanese macaque monkeys, Macaca fuscata, was examined monthly during the first three years of life using magnetic resonance imaging, to delineate the present or absence of these two processes and their contributions to the development of the pharyngeal topology. The macaque shows descent of the hyoid relative to the palate, but lacks the descent of the laryngeal skeleton relative to the hyoid and that of the EG from the VL. We argue that the former descent is simply a morphological consequence of mandibular growth and that the latter pair of descents arose in a common ancestor of extant hominoids. Thus, the evolutionary path of the great descent of the larynx is likely to be explained by a model comprising multiple and mosaic evolutionary pathways, wherein these developmental phenomena may have contributed secondarily to the faculty of speech in the human lineage.     

Form and function of the masticatory musculature in the tree sloths, Bradypus and Choloepus

The tree sloths, Bradypus and Choloepus, show unusual masticatory specializations, compared to each other and to other mammals. Both have an incomplete zygomatic arch with descending jugal process, a complex superficial masseter, a large temporalis and medial pterygoid musculature, and a lateral pterygoid with two heads. In Choloepus the deep masseter and zygomaticomandibularis are typical when compared to other mammals. However, in Bradypus there is an ascending jugal process from which enlarged and vertically oriented deep masseter and zygomaticomandibularis muscles originate. Although both sloths are folivores, the anterior teeth in Choloepus are caniniform, while those of Bradypus have lost such elongation. In both sloths the glenoid cavity is similarly located; however, in Bradypus the craniomandibular joint is raised above the occlusal plane, and the pterygoid flanges are elongated. Prediction of the evolutionary sequence of cranial changes from Choloepus-like (primitive) to Bradypus-like (derived) morphology is based upon the most parsimonious model of masseter-medial pterygoid complex changes for masticatory efficiency improvement. The model proposes that the condylar neck in Bradypus was elongated and that this single change predicated a series of other structural changes. Mandibular movement patterns in both sloths showed anteromedially directed unilateral power strokes as in other mammals. Puncture-crushing, tooth-sharpening, and chewing cycles are distinct in Choloepus, less so in Bradypus. The masticatory rate is slow in sloths compared to other mammals of similar body size, averaging 590 ms per cycle for Choloepus and 510 ms for Bradypus.     

Mammals with a long diastema typically also have dominant masseter and pterygoid muscles

A few orders of mammals contain many individuals with dominant masseter and pterygoid muscles that pull up and forward as they close the jaw. A dominant temporalis muscle that pulls the jaw up and to the rear is the more common condition in mammals. A long toothless region (diastema) is present in almost all mammals with a large masseter/pterygoid complex. The presence of a diastema, when few teeth have been lost and their size has not changed significantly over evolutionary time, implies that the jaws have lengthened, as in horses and selenodont artiodactyls. (A long jaw with a shorter diastema will also form if very long incisors develop as in rodents.) The sum of the forces of all the jaw muscles (represented by an arrow) typically divides the jaw into a posterior, toothless region and an anterior region where the teeth are located. In most mammals, the sum of all the bite forces at the teeth is maximized when the lengths of the projections of these two regions, onto a line perpendicular to the arrow, are in the ratio of 3 : 7. If the tooth-bearing region of the jaws becomes longer over evolutionary time this ratio will obviously be disturbed. A change in the location of some basic bony features of the jaw mechanism could maintain this ratio, but this requires major disruption of the skull and jaws. Alternatively, simply changing the masses of the muscles that close the jaw (smaller temporalis, larger masseter and/or pterygoid, or some combination), so that the lower jaw is pulled up and forward, rather than backward, also maintains the ratio. According to this view, if the jaw lengthens over evolutionary time, the relative sizes of the jaw muscles will change so that the masseter/pterygoid complex will become dominant.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 625–629.     

内蒙古大青山地区二齿兽类化石的发现

本文记述内蒙古大青山地区脑包沟组中首次发现的脊椎动物化石——边缘大青山兽(Da-qingshanodon limbus)。它具有翼骨腭上枝、基蝶骨小侧架、卷曲的鳞骨等独特构造。依其颅基中轴长、翼间窝长、翼间孔窄,眶前部分短,泪骨与隔颌骨分离等特征,以及在较高层位中发现的大鼻龙类、兽头类等化石,确认大青山兽属典型的晚二叠世二齿兽类型。文中还讨论了二齿兽类鼻区、蝶筛区、翼骨区等构造的演变,“萨拉齐系”与“石千峰群”的岩性和生物的可比性。     

The origin and early radiation of the therapsid mammal-like reptiles: a palaeobiological hypothesis

The replacement of the basal synapsid pelycosaurs by the more 'mammal-like' therapsids in the Permian was an important event in the history of tetrapods because it initiated the eventual transition to the mammals. It is also an example of taxon replacement in the fossil record that is unusually amenable to explanation, based on a combination of analysis of the biological significance of the inferred character changes, with the stratigraphic, palaeogeographic and palaeoecological circumstances of the time. An hypothesis is presented in which the origin of the therapsids resulted from a correlated progression of character evolution leading to higher levels of metabolic activity and homeostatic regulation of the body. It was a response to the availability of a seasonally arid, savanna-like biome. The subsequent explosive radiation of therapsids was associated with habitat expansion made possible by the Mid-Permian development of geographical continuity between that biome and the temperate biomes. The final extinction of the pelycosaurs was a case of incumbent replacement by the new therapsid lineages.     

Histomorphological structure of the palate and histochemical profiles of the salivary palatine glands in the Chukar partridge (Alectoris chukar,Gray 1830)

Morphology of the palatine mucosa and its secretion was described in Chukar partridges, by gross morphology and histochemistry techniques. For this purpose, 10 healthy adults (five male and five female) were sacrificed. After sacrifice, the palatine tissues were extirpated and fixed in 10% formol‐alcohol for 18 h and were embedded longitudinally and transversally in paraffin. The 5‐μm sections were employed histological and histochemical staining techniques. The lateral rims of the caudal part of the choanal cleft were bordered by large conical papillae. In the periphery of the choanal and the infundibular cleft, small papillae were scattered across the palatine mucosa. The palate was lined by keratinized stratified squamous epithelium, which contained conical papillae of varying height. However, the folds of the keratinized stratified squamous epithelial layer covering the choanal and infundibular cleft were nonkeratinized. The rostral aspect of the choanal cleft contained simple branched tubulo‐alveolar glands of both mucous and sero‐mucous characteristic, whilst the caudal aspect included mucous simple branched tubular glands. Furthermore, it was ascertained that the secretion of the palatine glands contained glycoproteins, carboxylated proteoglycans, weakly and strongly sulphated mucins, sialic acid and hyaluronic acid, but lacked glycogen. In conclusion, it was demonstrated that the histological structure of the mucosal epithelium and the supporting elements displayed similarity to those of other domestic avian species.     

Anatomy of the auditory tube and related structures in the rhesus monkey (Macaca mulatta).

The primate nasopharynx-auditory tube-middle ear complex is being used by several researchers to model both normal and pathologic functions of the human auditory tube. An extensive search of the literature has indicated little detailed information on the primate tube/middle ear system. This study was undertaken to define the anatomical characteristic of the system in the rhesus monkey (Macaca mulatta) and to determine the limits on the use of the monkey as a model of human tubal function. Although the direct application of morphologic data to account for the observed function of a system is a tenous one, the data on the rhesus monkey auditory tube appear to be consistent with those published for other mammals. The tensor veli palatini muscle appears to be the only muscle to act directly on the tube and effect tubal dilation. The muscle is attached to the lateral membranous tubal wall along its extrabullar extension. The muscle has an inferior attachment to the posterior hard palate and thus possesses a vector directed inferolaterally; contraction would appear to pull the membranous wall inferiorly and laterally, resulting in tubal dilation. The auditory tube relationships of the salpingopharyngeus, levator veli palatini, and internal pterygoid muscles are described. Their possible role in primate tubal function is minimal at best.     

The radiation of cynodonts and the ground plan of mammalian morphological diversity

Cynodont therapsids diversified extensively after the Permo-Triassic mass extinction event, and gave rise to mammals in the Jurassic. We use an enlarged and revised dataset of discrete skeletal characters to build a new phylogeny for all main cynodont clades from the Late Permian to the Early Jurassic, and we analyse models of morphological diversification in the group. Basal taxa and epicynodonts are paraphyletic relative to eucynodonts, and the latter are divided into cynognathians and probainognathians, with tritylodonts and mammals forming sister groups. Disparity analyses reveal a heterogeneous distribution of cynodonts in a morphospace derived from cladistic characters. Pairwise morphological distances are weakly correlated with phylogenetic distances. Comparisons of disparity by groups and through time are non-significant, especially after the data are rarefied. A disparity peak occurs in the Early/Middle Triassic, after which period the mean disparity fluctuates little. Cynognathians were characterized by high evolutionary rates and high diversity early in their history, whereas probainognathian rates were low. Community structure may have been instrumental in imposing different rates on the two clades.     

Secondary Male Sex Characteristics of Hoplolaimus galeatus

The secondary male sex characteristics of Hoplolaimus galeatus consisted of caudal alae, two independently retractable spicules and a gubernaculum with two bilobed titillae. The spicules were dimorphic, with the outer one possessing a velum. When both spicules were completely extruded, the only open orifice on the ventral surface of the posterior region was formed by the close association of these two appendages. In specimens where the inner spicule was slightly retracted, the velum almost completely surrounded the inner spicule. When the inner spicule was retracted further, the velum appeared to convolute, closing the orifice described above.     

Surgical anatomy of the levator veli palatini: a previously undescribed tendinous insertion of the anterolateral fibers

The purpose of this study was to describe the previously unreported tendinous insertion of the anterolateral fibers of the levator veli palatini (levator) and discuss possible implications for levator function and cleft palate repair. The velopharyngeal anatomy in normal adult cadavers was studied, with histologic confirmation of anatomical findings. These findings were compared with a more limited study of levator anatomy in cleft palates at the time of intraoperative muscle dissection. Just before entering the velum, the levator divides into two parts. The smaller bundle of muscle fibers (anterolateral part) runs anteriorly, close to the lateral pharyngeal wall, and inserts into the palatine aponeurosis through a number of fine tendons. The main part of the muscle runs medially into the velum, where it fans out and forms the levator sling with the contralateral levator. The possible function of the anterolateral part of the levator is discussed. Inadequate release of the tendinous insertions at the time of palate repair may tether the levator anteriorly and compromise muscle retropositioning or may result in splitting of the levator, so that only part of the levator is retropositioned.     

Cranial structure and relationships of the Liassic mammal Sinoconodon*

The skull of the 'Rhaeto-Liassic' mammal Sinoconodon changchiawaensis (Young) from the lower Lufeng Series of China is described. It is characterized by a relatively larger and more robust dentary condyle and a greater reduction of the post-dentary bones than are present in the Morganucodontidae, Kuehneotheriidae, and Dinnetherium. In other aspects Sinoconodon is more primitive; precise post-canine occlusion is lacking, the mandibular symphysis is deep, the jaw articulation lies below a line projected through the apices of the teeth, the pterygoparoccipital foramen is large and the post-canine teeth cannot be divided into molars and premolars. The jaw articulation and braincase of Sinoconodon are compared with those of the two cynodont therapsids Probainognathus and Thrinaxodon. It is concluded that in the transition from therapsid to mammal the medial surface of the groove in the squamosal housing the quadrate was lost and, as a result, in Sinoconodon, Morganucodon and Dinnetherium the hollow medial surface of the quadrate abutted directly against the paroccipital process. A definition for the Class Mammalia is given. It is suggested that the three-boned middle ear was present in Cretaceous triconodonts, and that it probably arose independently in the lines leading to multituberculates and Cretaceous therians. The structure of recently discovered 'Rhaeto-Liassic' mammals and that of Sinoconodon indicates that there was greater diversity among the earliest known mammals than was previously thought.     

Selective Factors Associated with the Origin of Fur and Feathers

Conventional wisdom notwithstanding, fur and feathers are unlikelyto have arisen in direct association with elevated metabolicrates in early mammals, birds, or their ancestors. A completeinsulative fur coat probably appeared first in the earliestmammals long after mammalian ancestors (therapsids) had attainedmammalian, or near-mammalian, metabolic rates. The evolutionof feathers was unlinked to the evolution of modern avian metabolicrates since early, fully flighted birds (i.e., Archaeopteryx)retained an ectothermic metabolic status. Recent claims of "feathereddinosaurs" should be regarded with caution.     

Diversity of functions of proteins with internal symmetry in spatial arrangement of secondary structural elements.

We carry out a systematic analysis of the correlation between similarity of protein three-dimensional structures and their evolutionary relationships. The structural similarity is quantitatively identified by an all-against-all comparison of the spatial arrangement of secondary structural elements in nonredundant 967 representative proteins, and the evolutionary relationship is judged according to the definition of superfamily in the SCOP database. We find the following symmetry rule: a protein pair that has similar folds but belong to different superfamilies has (with a very rare exception) certain internal symmetry in its common similar folds. Possible reasons behind the symmetry rule are discussed.     

A new fossil from the Jurassic of Patagonia reveals the early basicranial evolution and the origins of Crocodyliformes

Extant crocodylians have a limited taxonomic and ecological diversity but they belong to a lineage (Crocodylomorpha) that includes basal and rather generalized species and a highly diverse clade, Crocodyliformes. The latter was among the most successful groups of Mesozoic tetrapods, both in terms of taxonomic and ecological diversity. Crocodyliforms thrived in terrestrial, semiaquatic, and marine environments, and their fossil diversity includes carnivorous, piscivorous, insectivorous, and herbivorous species. This remarkable ecological and trophic diversity is thought only to occur in forms with a completely akinetic skull, characterized by a functionally integrated and tightly sutured braincase‐quadrate‐palate complex. However, the patterns of evolutionary change that led to the highly modified skull of crocodyliforms and that likely enabled their diversification remain poorly understood. Herein, a new basal crocodylomorph from the Late Jurassic of Patagonia is described, Almadasuchus figarii gen. et sp. nov. The new taxon is known from a well‐preserved posterior region of the skull as well as other craniomandibular and postcranial remains. Almadasuchus figarii differs from all other crocodylomorphs in the presence of six autapomorphic features, including the presence of a large lateral notch on the upper temporal bar, an otic shelf of the squamosal that is wider than long, a deep subtriangular concavity on the posterolateral surface of the squamosal, and an elongated pneumatopore on the ventral surface of the quadrate. Phylogenetic analysis focused on the origin of Crocodyliformes places Almadasuchus as the sister group of Crocodyliformes, supported by synapomorphic features of the skull (e.g. subtriangular basisphenoid, absence of basipterygoid process, absence of a sagittal ridge on the frontal, and a flat anterior skull roof with an ornamented dorsal surface). New braincase information provided by Almadasuchus and other crocodylomorphs indicates that most of the modifications on the posterior region of the skull of crocodyliforms, including the strongly sutured braincase, quadrate, and the extensive secondary palate appeared in a stepwise manner, and pre‐dated the evolutionary changes in the snout, jaws, and dentition. This indicates that the progressively increased rigidity of the skull provided the structural framework that allowed the great ecological diversification of crocodyliforms during the course of the Mesozoic. The phylogenetic pattern of character acquisition inferred for the strongly sutured (akinetic) skull and the appearance of more diverse feeding behaviours that create high mechanical loads on the skull provides another interesting parallel between the evolution of Mesozoic crocodyliforms and the evolutionary origins of mammals.     

Morphogenetic features in the tail region of the rat embryo.

The secondary (direct) body formation is a mechanism of development in which morphogenesis of various organs occurs directly from a mass of undifferentiated mesenchymal cells (blastema) without previous formation of germ layers. It is characteristic of the posterior end of the embryonic body, i.e. of the tail bud of tailless and the tail of tailed mammals. Development of the neural tube occurring by this mechanism (secondary neurulation) has been previously explained. We investigated the morphogenetic mechanism by which two other axial structures in the rat tail develop: the tail gut and the notochord. Both structures develop from an axial condensation of undifferentiated mesenchymal cells (tail cord) of tail bud origin. The tail gut forms in a similar way to the secondary neural tube: cells in the ventral part of the tail cord elongate, acquire an apicobasal polarity and form a rosette-like structure around a lumen in the centre. The notochord forms by detachment of a group of cells of the tail cord dorsally to the developing tail gut. The peculiarities of this morphogenetic mechanism in comparison with those in other parts of the embryo are discussed. Causal (including evolutionary) explanations of this mechanism are ruled out.     

Epithelium of lips and associated structures of the Indian major carp,Catla catla

Structural organization of the epithelium of the lips and associated structures of the Indian major carp,Catla catla, is described. The upper lip is thin and is associated on its dorsal side with a membranous fold of skin and the rostral cap. In contrast, the lower lip is thick and very conspicuous. It is associated on its ventral side with a fold of skin between it and the ventral head skin. The lower lip is divided into a non-projectile portion, a projectile portion and an intermediate groove region. The projectile portion remains folded covering a part of the ventral head skin when the mouth is closed. Their role in relation to the formation of the characteristic feeding tube is discussed. The epithelium of the lips and associated structures is stratified in nature and is composed of the epithelial cells, mucous cells, club cells, lymphocytes and the taste buds. The mucous cells are small, few or even absent and do not appear to secrete profusely at the surfaces of the upper and the lower lips. This suggests that the lips inCatla catla, which feeds on micro-organisms, do not need extra lubrication for protection against abrasion during feeding. In the epithelium at the folds of skin, the voluminous mucous cells secrete profusely and provide extra lubrication to their surface. This reduces the resistance to surface drag during stretching and enables the jaws to protrude with increasing efficiency and swiftness. The club cells are developed additionally to complement the mucous cells in the rostral cap and the upper lip epithelium. Their primary function appears protective in some way, which needs further confirmation. The taste buds, though few in the lower lip, are located in a good number in the upper lip on the characteristic epithelial papillae-like projections, and are projected at the surface. These have been associated with the acute gustatory sense of the fish. The taste buds are absent on the folds of skin where they may not be of much significance.