


Vol 46, No 7 (2019)
- Year: 2019
- Articles: 10
- URL: https://ogarev-online.ru/1062-3590/issue/view/11443
Article
Vertebrate Head Metamerism: The Current State of an Old Problem
Abstract
The problem of vertebrate head metamerism was first posed more than 200 years ago. This issue gave rise to two approaches: segmentalism, which relies on the model of the typical head segment, and antisegmentalism, which rejects the existence of cephalic segments altogether. The truth, probably, lies somewhere in the middle. Data on the development of the head mesoderm, cranial neural crest, and cranial nerves lead to the conclusion that there are five metameres in the vertebrate head: premandibular, mandibular, hyoid, 1st metahyoid, and 2nd metahyoid. According to the hypothesis of F. Edgeworth, this number of head metameres is primary; the head spreading back occurred as a result of the polymerization of visceral arches inside the nervus vagus metamere which had originally included only the 2nd metahyoid arch. According to the hypothesis of C. Gegenbaur, initially, the visceral arches were more numerous in this area and the branchial nerves merged into a single nervus vagus system only secondarily. The Cambrian paleontology may assist in resolving this controversy. In any case, vertebrate head metamerism is primarily heteronomous (as interpreted by P.P. Ivanov): the premandibular and mandibular metameres are morphologically atypical and lie beyond the Hox gene expression zone. Developmental biology shows that the bodies of other metameric animals are arranged on the basis of similar principles.



Functional Morphology and Adaptive Significance of the Jaw Apparatus of the Parakeet Auklet (Aethia psittacula, Alcidae)
Abstract
The jaw apparatus of the parakeet auklet (Aethia psittacula) has been investigated in comparison to that of the crested auklet (Aethia cristatella). The mechanical and biological effects of the distinctive bill structure in the parakeet auklet have been analyzed. The oral slit is concave; therefore, equilibrium is achieved without the need for disabling the posterior or anterior parts of the dorsal adductor of the lower jaw and the adductor can act with the maximal force regardless of the position of the object clamped in the bill. Shearing of large food objects, such as polychaetes and jellyfish, by the parakeet auklet bill is a jaw apparatus specialization unique among birds. Strengthening of the jaw apparatus, which was initially adapted for capturing small planktonic crustaceans, involved the development of additional aponeuroses in the internal adductor, consolidation of the latter, shortening of the bill, and changes of the shape of the oral slit opening.



Functional Morphology of the Hyoid Apparatus in Old World Suboscines (Eurylaimides): 1. Anatomical Description
Abstract
The hyoid apparatus of the Old World suboscines (Eurylaimides) was studied by a layer-by-layer preparation of the hypoglossal muscles with subsequent comparative analysis of the morphological features found. The characteristic features of the tongue structure of representatives of all three families (Pittidae, Eurylaimidae, and Philepittidae) were described in detail. It was shown that, in general, the structure of the hyoid apparatus of these birds is similar to songbirds, which are characterized by considerable mobility of the tongue and which have a pronounced ability to manipulate objects with their beak. In addition, details specific to each of the studied groups were identified. The tongue of pittas (Pittidae) has a structure that is the most typical of passerines, while differing in by a significant relative length of some parts of its skeleton. Broadbills (Eurylaimidae), especially carnivorous species, are distinguished by a specifically broad and thick tongue. The tongue of asities (Philepittidae) is characterized by differentiation: its distal part is curled up into a tube, whereas the proximal part forms a platform.



Possible Directions of Further Development of F.Ya. Dzerzhinsky’s Program in Research on the Jaw Apparatus of Birds
Abstract
The works by F.Ya. Dzerzhinsky and his disciples have rendered the jaw apparatus of birds a conventional and thoroughly characterized research object for functional morphology in Russia. The author, who communicated with Dzerzhinsky for many years, attempts to reconsider his scientific heritage in a critical manner and to use it as a foundation for outlining the most promising directions of further research. The analysis shows that underestimation of the problem of muscle mass economy in the jaw apparatus, despite the critical importance of any excess ballast for birds, was a serious drawback of Dzerzhinsky’s method. A new model based on the separation of the kinematics of bill opening and food object compression provides a solution for the problem of economical use of muscle mass in the avian jaw apparatus. The length of the major jaw adductor muscles remains virtually unchanged when the bill is opened, so that these muscles act as inextensible links of the closed multilink kinematic chain of the kinetic skull. Length change upon mouth opening being unnecessary, these muscles can be short-fibered, that is, lightweight but strong. The large force of these muscles is only used by the bird when the object is fixed in the bill and the jaw joint locking mechanism is switched on; a different kinematic chain used for object compression only is formed in this case. If the nonparallel arrangement of the pterygoid muscle and the jugal bar is also taken into account, the avian capacity for using the full force potential of the jaw adductor muscles without careful adjustment to each food object and the specific conditions of food object capturing becomes easy to understand. The main elements of the proposed model, which were found out by Dzerzhinsky himself, acquire a new meaning in light of the concept of economical use of the muscle mass. This enables us to make a major generalization concerning the adaptive significance of the cranial kinesis in birds as a means of reducing the jaw adductors’ mass without decreasing their force. In addition to the new model of jaw mechanics, other promising directions are proposed for the development of Dzerzhinsky’s work based on the expansion of the set of methods. The analysis of the aponeurotic framework of jaw muscles from the standpoint of optimizing the spatial packaging of muscle fibers may be a promising topic for research based on a combination of computer tomography and computational modeling.



Pathways and Level of Morphological Adaptations in Modern Diatomyidae and Ctenodactylidae (Rodentia)
Abstract
Based on published and original data, several morphofunctional systems were compared in the extant species of Diatomyidae and Ctenodactylidae, both families being considered as sister taxa. The subcutaneous and auricular muscles, otic capsule, jaw apparatus, and distal limbs were examined. These groups were shown to differ significantly in both the level and the direction of the morphofunctional transformations of the above systems. Ctenodactylids are a much more specialized group than Laonastes. They have an otic capsule, a jaw apparatus, and distal limb sections that reach the maximum level of morphological and functional specialization in the rodents, whereas in Laonastes, they correspond approximately to the average level of their development. Both groups are characterized by different pathways of morphological transformations of all systems considered, even those of them (jaw apparatus and limbs) that are associated with adaptations to similar ecological conditions, i.e., life on stones and herbivory. In Laonastes, the structure of the above systems retains archaic characteristics that are combined with features specific only to this group. In gundis, the direction of morphological transformations is completely specific with regard to some of the parameters, while in other respects it corresponds to the trends typical of hystricognathous rodents, this having led to a large number of structural parallelisms. In the structure of each morphological system examined, synapomorphies that support the monophyly of Ctenohystrica were revealed. There is no single-valued morphological evidence for the close relationship between Ctenodactylidae and Laonastes relative to Hystricognathi, although in the structure of almost all of the systems examined there are common features distinguishing these taxa from other rodents, in particular from Hystricognathi. Characters reflecting the pattern of differentiation of the subcutaneous muscle and the features of mastoid pneumatization can be regarded as the most significant for assessing the phylogenetic relationships of Diatomyidae, Ctenodactylidae, and Hystricognathi. However, for a more reliable assessment of the relationships based on morphological data, more extensive material is needed to cover the diversity of the structures considered in hystricognathous rodents.



Morphological Specificity of the Auditory Capsule of Sciurid (Sciuridae, Rodentia)
Abstract
Specific features of the sciurid auditory capsule have been analyzed based on 63 species from 20 genera. Its morphological specificity is characterized by a peculiar combination of primitive and advanced features stable within the group. They are (1) a rounded and inflated bulla with internal septa and a primitive attachment of the tympanic ring to the inner tympanic wall; (2) a transverse position of the bulla relative to the axis of the promontorium, resulting in the presence of deep petrosal fossae in front of and above the promontorium which are not covered by the tympanic bone; (3) a fully formed osseus facial canal and a developed bone tube of the stapedial artery; (4) the presence of a meato-cochlear bridge connecting the promontorium to the posterior wall of the acoustic duct which, in a fully developed form, does not occur in any other recent rodents; (5) pneumatization of the mastoid due to the expanding of the epitympanic chamber both anteriorly and posteriorly, with the formation of a premeatal pocket and a large epitympano-mastoid chamber divided into parts. The sciurid auditory capsule can generally be considered as morphologically advanced. Its diversity within the family concerns functionally significant features and is manifested in the structure of the external acoustic meatus, the size of the tympanic membrane, the length of the processes of auditory ossicles, and, most importantly, the degree of pneumatization of the auditory capsule. This is associated with quantitative variations of the characters and does not affect its structural plan, as a rule.



Modern Data on the Origin and Early Radiation of Mammals
Abstract
Modern fossil evidence shows extensive ecomorphological radiation of Mesozoic mammals. By the mid-Cretaceous, mammals had evolved all the main adaptations of modern small-sized Mammalia except for true powered flight. In addition, new paleontological and embryological data on the morphogenesis of the auditory ossicles demonstrate parallel developments of the definitive mammal middle ear in the placentals, marsupials, and monotremes, as well as the independent origins of a number of early groups: Allotheria (Haramiyida and Multituberculata) within Mammaliamorpha (irrespective of the lineage including eucynodonts Brasilodontidae and the mammaliaform Apotheria, Prototheria, and Acrotheria) and Prototheria (including Monotremata) within the Mammaliaformes (parallel to the Acrotheria lineage).



Peculiarities of the Structure and Locomotor Function of the Tail in Sauropterygia
Abstract
Among ancient and modern marine reptiles, several structural types of the locomotor apparatus were or are present, supporting different styles of swimming. Ichthyosaurs, mosasaurs, saltwater crocodiles, and representatives of many other groups swam or swim with horizontal undulations of the body primarily using the tail with a vertical caudal fin. Sea turtles with a reduced tail and their body completely immobilized by the shell use only limbs transformed into flippers for swimming. Sauropterygians had a substantially immobilized trunk and a horizontal caudal fin. They used a unique type of subaquatic locomotion with the leading role of two pairs of powerful flippers and vertical undulations of the tail, partial analogs of which are found in sea turtles, sirens, cetaceans, and some semi-aquatic mammals.



Biomechanical Analysis of Fur as a Tool for Study of Thermal Insulation in Semi-Aquatic Mammals
Abstract
One of the means for body insulation in mammals during their transition from terrestrial to semi-aquatic and then aquatic life is based on the use of fur in water as an effective “dry” insulator. Fur functional conditions are changed drastically and instantaneously when fur makes contact with water. The physical mechanisms underlying fur waterproofing were analyzed, and their relations to both fur structure and animal body size were established. The results of the analysis allow us to define to what extent the structural features of fur of a mammal in relation to its body size are capable of trapping an insulating air layer when the animal swims in water. On the basis of our analysis, both the body size of the ancestors of fur seals and the type of thermal insulation changing in the postnatal ontogeny of fur seals are hypothesized.



Microstructure of the Primary Remex of Owls (Strigiformes)
Abstract
An electron microscopic study of the microstructure of the primary remiges in owls (Strigiformes) was conducted using 13 species: Nyctea scandiaca, Bubo bubo, Asio otus, Otus scops, O. sunia, Aegolius funereus, Athene noctua, Glaucidium passerinum, Surnia ulula, Strux aluco, S. uralensis, S. nebulosa, and Tyto alba. Owls have a number of species-specific microstructural primary remex characteristics that are taxonomically important. First of all, these are the structural features of the pennaceous barb, which differ markedly not only at the ordinal, but also at the species level: cross-section configuration, pith architectonics in cross- and longitudinal sections, and cuticular barb structures. This work emphasizes that the identified elements are of taxonomic significance only when comparing the specific, strictly analogous sections of the barbs in different bird species. Based on the present study, we conclude that along with the typical elements of feather architectonics characteristic of representatives of other bird orders, the owls show a number of peculiar microstructural features. Thus, as the result of an analysis of the microstructure of the owls primary remex pennaceous barb vanule, the unique features of the distal barbules and the structure of the apical section of the barb with tightly adjacent and elongate proximal and distal barbules could be distinguished. These characteristics are revealed to cause a dense fleecy structure of the vane dorsal surface and the presence of a complex of peculiar “bunches” that form the cleft edge, i.e., the purely specific traits of the owls feather that are not found in representatives of any other bird orders we have investigated.


