THE BIOLOGY OF THE LOBE-FINNED FISHES

1. Interpretation of structural evolution in a group such as the Sarcopterygii requires consideration of a combination of all possible functions, rather than single functions. 2. The Dipnoi are probably more closely related to the Crossopterygii than to other groups of fishes. The Sarcopterygii are a ‘natural’ group. Certain characters in common between the elasmobranchs and the Dipnoi or Coelacanthini seem to be the result of convergent evolution. 3. Evolution of the skull, in connexion with both respiratory and feeding mechanisms, has resulted in extreme specialization in all Sarcopterygii. The crossopterygian intracranial kinesis has evolved from an earlier mobility between the skull and neck and is adapted for increasing the power of the bite and for enclosing the prey from both above and below, in addition to other factors. Adaptive radiation is seen in the feeding mechanisms of all forms. The evolution of the Amphibia proceeded through elongation of the anterior division of the skull (which is not correlated with any changes in brain morphology) and loss of the kinetic mechanism in this sequence is at least partially associated with improved buccal pumping mechanisms for lung ventilation. 4. Adaptive radiation of the respiratory system in Dipnoi shows a progressive increase in the use of aerial respiration. The aquatic condition seen in Neoceratodus is probably secondary. Comparison of the three living genera shows a striking correlation between respiratory physiology and habit. There is little indication of reduction of the branchial respiratory system in known Rhipidistia, in which respiration was probably primarily aquatic. In Dipnoi and Rhipidistia, evolution of the lung allowed a partial control of the hydrostatic properties of the body. In coelacanths, aerial respiration was abandoned, except in certain secondarily freshwater forms, and the single lung is modified as an organ of hydrostatic balance. These changes are reflected in the over-all body proportions. 5. Locomotion in Sarcopterygii (except the coelacanths Laugia and Piveteauia) is adapted for contact with the substrate in relatively shallow water in most cases. Adaptive radiation of the locomotor apparatus is seen with respect to the relative roles and functions of the paired and unpaired fins, over-all body shape, caudal fin shape, and absolute size. An important function of the pectoral fins in advanced Rhipidistia was in supporting the body in shallow water and thus aiding lung ventilation. 6. Aestivation is an early feature of dipnoan biology, but was not evolved in Rhipidistia. The common faculty of urea production via the ornithine cycle and urea retention in coelacanths and dipnoans are adaptations to conditions in which the body tissues may become dehydrated (salt water and desiccation, respectively). The common pattern of nitrogen metabolism seems to have evolved during a marine phase in sarcopterygian evolution. 7. There is evidence that the earliest members of all sarcopterygian lines included marine forms. However, the subsequent major radiations of Dipnoi and Rhipidistia occurred in fresh waters. The distribution of Sarcopterygii was entirely tropical. The late Palaeozoic distribution of the freshwater forms seems to offer evidence for the occurrence of Continental Drift. Coelacanths were primarily coastal fishes. 8. The evolution of a major group of organisms requires a different pattern of evolutionary change than that by which adaptive radiations are produced. It evolves the structural and temporal correlation of modification in a number of different functional systems rather than the separate modification of each system without reference to other systems. 9. The first tetrapods evolved in a highly seasonal swampy environment on the shores of inland lakes or rivers, in permanently moist conditions.     

The cranial morphology of Greererpeton burkemorani Romer (Amphibia: Temnospondyli)

The skull of Greererpeton burkemorani Romer, a temnospondyl amphibian from the Upper Mississippian at Greer, West Virginia is described. A detailed account of the stapes of a Mississippian amphibian is given for the first time and its function is discussed. It is suggested that the stapes formed the principal element of support for the back of the braincase and resisted potential dislocation of the otico-occipital region from the skull roof during contraction of the hypaxial musculature.
Greererpeton is included in the Colosteidae and an amended diagnosis of the family is given. Erpetosaurus differs from Colosteus, Greererpeton and Pholidogaster in the pattern of bones in the skull roof and palate, the dentition and the otic region and, consequently, it is removed from the Colosteidae. The Temnospondyli are considered to be a monophyletic group characterized by the development of a connection between the dorsal portion of the occipital arch, the exoccipital bones, and the skull roof. The loxommatids are removed from the Temnospondyli as they retain the plesiomorphic condition of braincase attachment which relies exclusively on derivatives of the auditory capsules.
On the basis of similarities in the structure of the braincase, palate and manus it is suggested that microsaurs are the collateral descendants (sister group) of temnospondyls. This relationship may account for the large number of similarities in the three living groups of Amphibia: Anura are generally believed to have descended from temnospondyls, while the Urodela and Apoda are often considered to have descended from microsaurs. These systematic conclusions endorse the recent suggestions that neither the Lepospondyli nor the Labyrinthodontia are natural groups, and both terms should be abandoned.     

Cranial Specialization and Locomotor Habit in the Lagomorpha

SYNOPSIS. The skull of modern leporid lagomorphs is structurallyspecialized to permit significant intracranial movement. Thisappears to be the first recorded instance of organized cranialkinesis in mammals. A well-defined intracranial joint encirclesthe braincase between the posterior occipito-otic complex andthe remainder of the cranium. It arises from the retention andelaboration of a zone of patency found in neonates. The jointallows the much heavier anterior region of the cranium ($ mandible)to move relative to the posterior region which is stabilizedby its muscular attachment to the neck. It is hypothesized thatthe kinesis functions as a shock-absorbing mechanism to minimizethe jarring effects (possibly on the visual apparatus) of thelarge impulsive loads associated with running and jumping. Thedissipation of kinetic energy through controlled skull deformationmay be augmented by the hydraulic displacement of intracranialblood through a specialized system of venous channels and sinuseslocated within the fenestrated posterior regions of the cranium.Both biomechanical considerations and behavioral observationsindicate that the relatively massive external ears of cursorialleporids may play a vital role in cranial kinesis by helpingto "reset’ the kinetic mechanism between loading cycles.The evolutionary origin of this specialized cranium, it is suggested,may be associated with the development of a cursorial, saltatorylocomotor habit in lagomorphs. Cranial kinesis is lacking insuch noncursorial forms as the living pikas (=Ochotonidae) andcertain primitive fossil leporids of North America (e.g., Paleolagus,Megalagus).     

The role of cranial kinesis in birds.

In birds, the ability to move the upper beak relative to the braincase has been the subject of many functional morphological investigations, but in many instances the adaptive significance of cranial kinesis remains unclear. Alternatively, cranial kinesis may be considered a consequence of the general design of the skull, rather than an adaptive trait as such. The present study reviews some results related to the mechanism and functional significance of cranial kinesis in birds. Quantitative three-dimensional X-ray has shown that in skulls morphologically as divers as paleognaths and neognaths the mechanism for elevation of the upper beak is very similar. One of the mechanisms proposed for avian jaw movement is a mechanical coupling of the upper and the lower jaw movement by the postorbital ligament. Such a mechanical coupling would necessitate upper beak elevation. However, independent control of upper and lower jaw has been shown to occur during beak movements in birds. Moreover, kinematic modeling and force measurements suggests that the maximum extensibility of collagen, in combination with the short distance of the insertion of the postorbital ligament to the quadrato-mandibular articulation do not constitute a block to lower jaw depression. The lower jaw ligaments serve to limit the maximal extension of the mandibula. It is suggested here that cranial kinesis in avian feeding may have evolved as a consequence of an increase in eye size. This increase in size led to a reduction of bony bars in the lateral aspect of the skull enabling the transfer of quadrate movement to the upper jaw. The selective forces favoring the development of a kinetic upper beak in birds may be subtle and act in different ecological contexts. Simultaneous movement of the upper and lower jaw not only increases the velocity of beak movements, but with elevated upper beak also less force is required to open the lower jaw. However, the penalty of increased mobility of elements in a lightweight skull and a large eye is potential instability of skull elements during biting, smaller bite forces and limitations on joint reaction forces. Such a lightly built, kinetic skull may have evolved in animals that feed on small plant material or insects. This type of food does not require the resistance of large external forces on the jaws as in carnivores eating large prey.     

Anatomical study of the skull of amphisbaenian Diplometopon zarudnyi (Squamata,Amphisbaenia)

This study investigates the amphisbaenian species skull which includes cranium, lower jaw and hyoid apparatus. The medial dorsal bones comprise the premaxilla, nasal, frontal and parietal. The premaxilla carries a large medial tooth and two lateral ones. The nasals are paired bones and separated by longitudinal suture. Bones of circumorbital series are frontal, orbitosphenoid and maxilla. The occipital ring consists of basioccipital, supraoccipital and exooccipital. Supraoccipital and basioccipital are single bones while the exo-occipitals are paired. The bones of the palate comprise premaxilla, maxilla, septomaxilla, palatine, pterygoid, ectopterygoid, basisphenoid, parasphenoid, orbitosphenoid and laterosphenoid. Prevomer and pterygoid teeth are absent. Palatine represent by two separate bones. The temporal bones are clearly visible. The lower jaw consists of the dentary, articular, coronoid, supra-angular, angular and splenial. The hyoid apparatus is represented by a Y-shaped structure. The mandible is long and is suspended from the braincase via relatively short quadrate. There is an extensive contact between the long angular and the large triangular coronoid. Thus inter-mandibular joint is bridged completely by the angular and consequently, the lower jaws are relatively rigid and kinetic. The maxillae are suspended from the braincase largely by ligaments and muscles rather than through bony articulation. In conclusion, the skull shape affects feeding strategy in Diplometopon zarudnyi. The prey is ingested and transported via a rapid maxillary raking mechanism.     

Why the short face? Developmental disintegration of the neurocranium drives convergent evolution in neotropical electric fishes

Convergent evolution is widely viewed as strong evidence for the influence of natural selection on the origin of phenotypic design. However, the emerging evo‐devo synthesis has highlighted other processes that may bias and direct phenotypic evolution in the presence of environmental and genetic variation. Developmental biases on the production of phenotypic variation may channel the evolution of convergent forms by limiting the range of phenotypes produced during ontogeny. Here, we study the evolution and convergence of brachycephalic and dolichocephalic skull shapes among 133 species of Neotropical electric fishes (Gymnotiformes: Teleostei) and identify potential developmental biases on phenotypic evolution. We plot the ontogenetic trajectories of neurocranial phenotypes in 17 species and document developmental modularity between the face and braincase regions of the skull. We recover a significant relationship between developmental covariation and relative skull length and a significant relationship between developmental covariation and ontogenetic disparity. We demonstrate that modularity and integration bias the production of phenotypes along the brachycephalic and dolichocephalic skull axis and contribute to multiple, independent evolutionary transformations to highly brachycephalic and dolichocephalic skull morphologies.     

Microsaurs as possible apodan ancestors

The specific ancestry and nature of the relationships of modern amphibians have not yet been established. Detailed comparisons of the anatomy of the skull roof, palate and braincase of living apodans and the Paleozoic microsaur Goniorhynchus demonstrate greater similarities than between apodans and any other group of amphibians, fossil or recent. Unlike any other amphibians, extensive pleurosphenoid ossifications are developed in the area of the Vth nerve, uniting the otic capsule with the sphenethmoid. Other important features that they share (although not uniquely) include the presence of all the primitive dermal elements of the palate, a solidly roofed temporal region, a row of palatal teeth parallel to the marginal dentition and a row of teeth on the medial surface of the lower jaw. The stapes has a similar configuration and position, totally different from that of frogs and salamanders. Such similarities do not necessarily prove close relationship, but indicate the necessity for considering that apodans may have an ancestry distinct from that of frogs and salamanders.     

The identification of the dermal bones of the head

The discovery of the ichthyostegid Amphibia in Upper Devonian rocks by Säve-Söderbergh (1932) introduced further difficulties into the already complex problems of the dermal bones of the skull roof. For some years previously ideas about the origin of the tetrapods had been dominated by Watson's (1926) Croonian Lecture in which he had demonstrated beyond reasonable doubt that the crossopterygian fishes and not the Dipnoi were their ancestors, and had attempted to show that many of the features of the Carboniferous labyrinthodonts were a direct inheritance from these fishes. It was to be expected, therefore, that any Amphibia from the Upper Devonian would be intermediate in their structures between the Middle Devonian osteolepids and the Carboniferous labyrinthodonts, but when discovered the ichthyostegids did not conform at all well to this expectation. While their skulls showed some very primitive features which might have been expected, the pattern of the dermal bones did not conform to plan, for these new animals had lost, it seemed, the intertemporals, bones found in both the osteolepids and nearly all early labyrinthodonts, and had a single postparietal bone in place of the paired bones of all other early Amphibia. The osteolepid skull had many more bones than these earliest Amphibia.     

Observations on two rhipidistian fishes from the Upper Devonian of Lode, Latvia

New finds of extremely well preserved skeletons of two rhipidistian crossopterygians, Panderichthys rhombolepis (Gross) and Laccognathus panderi Gross, in the Gauja beds (Upper Devonian of Latvia) make possible nearly complete reconstructions of these fishes, and amplify our knowledge of the morphology of the two main orders of Rhipidistia, the Osteolepidida and Holoprvchiida. Panderichthys rhambolepis is distinguished from other osteolepids by consolidation of the bones of the skull roof and cheek; a single attachment surface between the scapulocoracoid and cleithrum; unusual vertebrae; partial fusion of the distal elements in the endoskeleton of the pectoral fin; and the absence of scutes at the bases of the fins. P. rhombolepis may represent a new order of Rhipidistia. L. panderi (litters from known holoptychiids only in details. One unusual feature is the presence of a single large external narial opening, limited posteriorly by a previously unknown bone, the prelachrvinal.     

The braincase and palate of the tetrapodomorph sarcopterygian mandageria fairfaxi: morphological variability near the fish–tetrapod transition

The braincase of the Late Devonian tristichopterid sarcopterygian Mandageria fairfaxi , from Canowindra, NSW, Australia, differs radically from the conservative pattern present in other 'osteolepiforms' (stem–group tetrapodomorph fishes) and non–dipnoan sarcopterygian fishes in general. The basioccipital region is short, displaced anteriorly, and either unossified or loosely articulated to the exoccipital, leaving most or all of the notochordal tunnel open ventrally. The exoccipital complex, which is developed into a large saddle that would have rested on top of the notochord, carries large, triangular articular facets on its posterior face and appears to have formed part of a functional neck joint, a synovial articulation between the skull and vertebral column that allows the former to rotate against the latter. Such a joint is characteristic of post–Devonian tetrapods, but unknown in other sarcopterygians. We infer that the ventrally open notochordal tunnel allowed gentle flexion of the cranial notochord during (predominantly vertical) rotational movement at the occiput; this is a mechanically unique solution to the problem of creating a mobile neck. Other unusual features of Mandageria include a posteriorly located lateral commissure, and structures on the entopterygoid and lateral commissure that may have been associated with an elaborate spiracular tract.     

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.     

Disparity and geometry of the skull in Archosauria (Reptilia: Diapsida)

A metric comparison of 155 fossil and extant species in lateral view based on the proportions of three homologous units (braincase, orbit and rostrum) reveals the existence of an archosaurian skull geometry. An empirical morphospace depicting skull proportions shows that the most variable unit is the rostrum. Three skull types based on rostral proportion are proposed: meso-, longi- and brevirostral. These types depend, on one hand, on a direct numerical relationship between the braincase and the orbit, with a mean ratio of 1:1; never surpassing a 2:1 or 1:2 ratio limit. On the other hand, skull types show a significant negative correlation between braincase and rostrum proportions. Close relationships have been obtained between orbit and the rostrum, although with lower significance and a geometric meaning specific to each group. Skull types depend mainly on the proportional relationship between the rostrum and the braincase. Mesorostral types account for more natural occurrences within morphospace, implying a plesiomorphic condition in Archosauria. Skulls with highest longirostral values (flying forms) display a more restrictive braincase–orbit ratio relationship. Brevirostrals are limited to the smallest skull lengths, up to approximately 180 mm. 85% of brevirostral modern birds have altricial post-hatchling development. General allometric pattern is very similar for all sampled archosaurs, although giant taxa (i.e. non-avian theropods) display a different type of skull proportional growth, closer to isometry. Results reveal the existence of a constructional skull geometry, highlighting the importance of the deviance of the structural design from adaptive explanations on craniofacial morphology in macroevolution.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80, 67–88.     

Early Evolution of the Vertebrate Eye—Fossil Evidence

Evidence of detailed brain morphology is illustrated and described for 400-million-year-old fossil skulls and braincases of early vertebrates (placoderm fishes). Their significance is summarized in the context of the historical development of knowledge of vertebrate anatomy, both before and since the time of Charles Darwin. These ancient extinct fishes show a unique type of preservation of the cartilaginous braincase and demonstrate a combination of characters unknown in other vertebrate species, living or extinct. The structure of the oldest detailed fossil evidence for the vertebrate eye and brain indicates a legacy from an ancestral segmented animal, in which the braincase is still partly subdivided, and the arrangement of nerves and muscles controlling eye movement was intermediate between the living jawless and jawed vertebrate groups. With their unique structure, these placoderms fill a gap in vertebrate morphology and also in the vertebrate fossil record. Like many other vertebrate fossils elucidated since Darwin’s time, they are key examples of the transitional forms that he predicted, showing combinations of characters that have never been observed together in living species.     

节甲鱼的一内颅化石

本文记述了广西六景的一长胸节甲类的内颅化石,并和我国已记述的Szuaspis yunnanensis、Kueichowlepis sinensis的内颅作了形态比较,讨论了它们之间的系统关系。     

Studies on the skull of the Indian sisorid catfish, Erethistes pussilus

The skull of Erethistes pussilus is described in detail. The general disposition of the bones corresponds to the siluroid pattern. Among the siluroid fishes, E. pussilus approaches the advanced forms in certain features such as obliteration of myodomic space, edentulous palate, absence of entopterygoids and supratemporals, intimate sutural articulation of posttemporals and complex vertebra with the cranium, diminished cranial cavity and loss of sutural articulation among the palatopterygoquadrate elements. In certain characters like the hyomandibula exclusively supported from the sphenotic, solitary hypohyal on each hyoid cornu, absence of interhyals, reduced orbits, edentulous vomer, small gape of mouth, toothless ectopterygoid and in the small number of branchiostegals, E. pussilus stands specialized alone among the catfishes. A diagnosis of the salient cranial characters of the fish is given and its relationship discussed.     

Studies on the skull of the Indian sisorid catfish, Erethistes pussilus

The skull of Erethistes pussilus is described in detail. The general disposition of the bones corresponds to the siluroid pattern. Among the siluroid fishes, E. pussilus approaches the advanced forms in certain features such as obliteration of myodomic space, edentulous palate, absence of entopterygoids and supratemporals, intimate sutural articulation of posttemporals and complex vertebra with the cranium, diminished cranial cavity and loss of sutural articulation among the palatopterygoquadrate elements. In certain characters like the hyomandibula exclusively supported from the sphenotic, solitary hypohyal on each hyoid cornu, absence of interhyals, reduced orbits, edentulous vomer, small gape of mouth, toothless ectopterygoid and in the small number of branchiostegals, E. pussilus stands specialized alone among the catfishes. A diagnosis of the salient cranial characters of the fish is given and its relationship discussed.     

Cranial kinesis in lizards (Lacertilia): Origin,biomechanics, and evolution

Various methods of investigation of cranial kinesis are compared. The biomechanical model of the amphikinetic cranial mechanism of lizards developed by Frazzetta (1962) corresponds to the skulls of species with the dependent streptostyly. A new modification of this model is proposed for species with the independent streptostyly, which conforms to the results of experimental investigations of cranial kinesis in living lizards. The lacertilian amphikinesis has developed on the basis of the pleurokinesis inherited from fish ancestors of tetrapods. Movable connections of the maxillo-buccal segments with the axial skull persisted in the amphikinetic skull and were completed by the transversal flexible connections in the dermatocranial roof and loose connections of dermatocranium with the braincase. The development of new movable intracranial connections could have been preceded by transformations in the jaw musculature (formation of the pterygoideus muscle and inclined position of the external jaw adductor), which caused longitudinal jaw movements. Development of new movable connections within the skull was triggered by paedomorphosis processes. In various lacertilian groups, the cranial kinesis was improved by the development of various forms of streptostyly and flexipalatality.