Description and interpretation of interphalangeal lines in tetrapods

A previously unnoticed geometric pattern is present in the extremities of all tetrapods. Sets of straight and typically uninterrupted hinge lines pass through neighboring interphalangeal joints and across ungual tips. Four sets of these lines appear in basal polydactyl tetrapods, two medial sets, a transverse set and a lateral set. The two medial sets merge in primitive pentadactyl tetrapods. The resulting three line sets persist in later taxa, even when digits shrink and disappear. Primitively and typically the lines in each set are more or less parallel, but lines may converge, merge and shift as phalanges disappear or phalangeal proportions change. Confirming this geometric pattern, complex interphalangeal joint surfaces typically align with hinge lines and pad divisions parallel them. In addition, unguals rarely cross extensions of hinge lines and longer unguals may divert medially or laterally rather than cross them. Exceptions occur most commonly on ungual II. Line sets may exist because phalanges appear to flex and extend most efficiently in unison. Hinge line patterns appear to identify clades so they may, to a limited extent, be used taxonomically. Hinge lines also have predictive value in that missing phalanges, including unguals, can be reconstructed with confidence using hinge lines as size guides. Correct digit spread and metapodial configuration can also be determined in extinct taxa by seeking the appearance of continuous interphalangeal hinge lines in tested reconstructions.     

Interpreting the autopodia of tetrapods: interphalangeal lines hinge on too many assumptions

Recently Peters proposed the concept of ‘interphalangeal lines’, defined as sub-parallel lines that could supposedly be drawn across the joints of the digits of all tetrapods. The lines were viewed as potential axes of rotation, and it was suggested that they could be used to determine the resting position of the digits, reconstruct missing digital elements of fossil tetrapods, and provide information on systematic relationships. Evidence was adduced from the skeletons of recent and fossil vertebrates and from footprints. However, detailed analysis shows that these claims are largely unfounded. Linear alignments of joints on neighbouring digits are not consistently present in tetrapods, especially across locomotor cycles. Even if present, interphalangeal (IP) lines would rarely be in an appropriate orientation to facilitate joint movements during locomotion. There is no reason to believe that IP lines would be homologous across different taxa, so they cannot be used to infer systematic relationships. Finally, the alleged support from the ichnological record is undermined by the uncertain relationship between the joint structure of the skeleton and the form of the print. We conclude that IP lines cannot be consistently constructed on tetrapod extremities, and would have minimal functional relevance or predictive power in any case.     

Bone Cells in Birds Show Exceptional Surface Area,a Characteristic Tracing Back to Saurischian Dinosaurs of the Late Triassic

Background

Dinosaurs are unique among terrestrial tetrapods in their body sizes, which range from less than 3 gm in hummingbirds to 70,000 kg or more in sauropods. Studies of the microstructure of bone tissue have indicated that large dinosaurs, once believed to be slow growing, attained maturity at rates comparable to or greater than those of large mammals. A number of structural criteria in bone tissue have been used to assess differences in rates of osteogenesis in extinct taxa, including counts of lines of arrested growth and the density of vascular canals.

Methodology/Principal Findings

Here, we examine the density of the cytoplasmic surface of bone-producing cells, a feature which may set an upper limit to the rate of osteogenesis. Osteocyte lacunae and canaliculi, the cavities in bone containing osteocytes and their extensions, were measured in thin-sections of primary (woven and parallel fibered) bone in a diversity of tetrapods. The results indicate that bone cell surfaces are more densely organized in the Saurischia (extant birds, extinct Mesozoic Theropoda and Sauropodomorpha) than in other tetrapods, a result of denser branching of the cell extensions. The highest postnatal growth rates among extant tetrapods occur in modern birds, the only surviving saurischians, and the finding of exceptional cytoplasmic surface area of the cells that produce bone in this group suggests a relationship with bone growth rate. In support of this relationship is finding the lowest cell surface density among the saurischians examined in Dinornis, a member of a group of ratites that evolved in New Zealand in isolation from mammalian predators and show other evidence of lowered maturation rates.     

Are lungfishes the sister group of tetrapods?

It is generally accepted that rhipaidistian crossopterygians are the closest relatives of tetrapods. Rosen, Forey, Gardiner & Patterson (1981) challenge this view and contend that lungfishes are the sister group of tetrapods. They present a detailed cladistic analysis and claim to identify a large number of synapomorphies shared by lungfishes and tetrapods but not by rhipidistians. Their analysis is faulty. Although Rosen et al. (1981) correctly emphasize that cladistic relationships must be based on shared derived characters, they often fail to take intragroup variation into account in postulating synapomorphies. They also use evidence inconsistently by attributing greater significance to similarities between lungfishes and tetrapods than to even more detailed similarities between rhipidistians and tetrapods. They misinterpret the skeletal pattern of the paired appendages. The many synapomorphies that they claim to have identified are either invalid, irrelevant, or are characters involving reduction or loss (which have a high probability of convergence). Consequently, they make an unconvincing case for a sister-group relationship between lungfishes and tetrapods. On the other hand, Rosen et al. (1981) do show that evidence for the orthodox view of rhipidistian-tetrapod relationships is not as strong as generally believed. The uncertain interrelationships among rhipidistians is a major problem. Tetrapod-fish relationships need to be re-examined by means of a properly conducted cladistic analysis.     

Evolution of tetrapod locomotion

Fish-like ancestors of tetrapods did not need strong limb musculature because they inhabited waters and were practically imponderable. In the primitive tetrapods, principal function of the limbs was initially restricted to passive anchoring in the course of animal movements on the substrate by means of lateral bending of the body (undulation). However, progressive development of carrying function of tetrapod limbs lead to clearing the body off the substrate which reduced friction costs and made the tetrapods less dependent on the substrate properties. Along with this, the limbs became more important as the active locomotory organs. But at the beginning, this diminished locomotory speed as the momentum caused by undulation could no longer provide additional forward sliding. Locomotory function of the tetrapod limb could be carried out due to both retraction and pronation at the shoulder joint. Relatively short humerus of the primitive tetrapods made it indifferent which of these two particular actions lead to elongation of the steps. In most of the recent tetrapods with sprawling limbs (Urodela, Lacertilia Sphenodontia, Crocodilia), step elongation was carried out mainly by retraction at the shoulder joint. Contrary to this, in Tachyglossidae (Mammalia: Monotremata) retraction is absent while pronation at the shoulder joint becomes the most important component of step elongation. This made it possible to recognize two principal types, pronatory and retractory, of locomotion on the basis of the main movement in the phase of support. A mathematical model describing changes in step length during the phase of support in both of these types is elaborated. It takes into account relative sizes of stylopodium and zeugopodium, the angles of pronation and retraction at the shoulder joint, the angle of adduction at the elbow joint, and the angle of body undulation arc. It is shown on the basis of this model, varying of which of the above parameters is advantageous and which is disadvantageous in each of the locomotory types. In the pronatory locomotory type, adduction (lateral mobility) at the elbow joint is employed. It leads to special changes in morphology of the elbow joint due to which humeral condyle becomes spherical and promotes both adduction and rotation of the entire antebrachium. In the retractory locomotory type, amplification of pronation is to be limited in order to provide step elongation, so certain morphological adaptations occur in the elbow joint which prevent adduction at this joint. For step elongation, retraction at the shoulder joint is usually more advantageous than pronation, therefore historical emergence of the pronatory type could be considered as inadaptive. However, transversal horizontal axis of rotation at the shoulder joint appeared to be a prerequisite of the subsequent appearance of the most perfect locomotion in the therian mammals with their parasagittal limbs. Transition to the parasagittal limb construction was associated with adaptation to jumping asymmetric locomotion. It caused elongation of the shoulder bone downward which lead to widening of rotation cone of the humerus and, at the same time, to reduction of the coracoid portion of the glenoid fossa, the latter became horizontal rather than lateral. As a part of this process, the longitudinal axis of the scapula was displacing caudally with destruction of the suture-like articulation of the acromion process with the clavicle. The latter became articulated with the sternum directly or via much reduced interclavicle (or via procoracoid rudiment). This increases amortisatory function of the shoulder girdle during landing at the final stage of jump.     

Dermal bone in early tetrapods: a palaeophysiological hypothesis of adaptation for terrestrial acidosis

The dermal bone sculpture of early, basal tetrapods of the Permo-Carboniferous is unlike the bone surface of any living vertebrate, and its function has long been obscure. Drawing from physiological studies of extant tetrapods, where dermal bone or other calcified tissues aid in regulating acid-base balance relating to hypercapnia (excess blood carbon dioxide) and/or lactate acidosis, we propose a similar function for these sculptured dermal bones in early tetrapods. Unlike the condition in modern reptiles, which experience hypercapnia when submerged in water, these animals would have experienced hypercapnia on land, owing to likely inefficient means of eliminating carbon dioxide. The different patterns of dermal bone sculpture in these tetrapods largely correlates with levels of terrestriality: sculpture is reduced or lost in stem amniotes that likely had the more efficient lung ventilation mode of costal aspiration, and in small-sized stem amphibians that would have been able to use the skin for gas exchange.     

Better than fish on land? Hearing across metamorphosis in salamanders

Early tetrapods faced an auditory challenge from the impedance mismatch between air and tissue in the transition from aquatic to terrestrial lifestyles during the Early Carboniferous (350 Ma). Consequently, tetrapods may have been deaf to airborne sounds for up to 100 Myr until tympanic middle ears evolved during the Triassic. The middle ear morphology of recent urodeles is similar to that of early ‘lepospondyl’ microsaur tetrapods, and experimental studies on their hearing capabilities are therefore useful to understand the evolutionary and functional drivers behind the shift from aquatic to aerial hearing in early tetrapods. Here, we combine imaging techniques with neurophysiological measurements to resolve how the change from aquatic larvae to terrestrial adult affects the ear morphology and sensory capabilities of salamanders. We show that air-induced pressure detection enhances underwater hearing sensitivity of salamanders at frequencies above 120 Hz, and that both terrestrial adults and fully aquatic juvenile salamanders can detect airborne sound. Collectively, these findings suggest that early atympanic tetrapods may have been pre-equipped to aerial hearing and are able to hear airborne sound better than fish on land. When selected for, this rudimentary hearing could have led to the evolution of tympanic middle ears.     

THE ORIGIN OF TETRAPODS-PAST ANDPRESENT HYPOTHESES

HistoricalreviewoneshouldexpectthatanewtheorychangesorimprovestheunderstandingofPhylogeneticquestions.ThatdoesnotseemtobetrueoftheoriginoftetrapodsasRosenetal.(l98l)havealreadyshowninthecaseoftheapPearanceofDarwin's'ontheoriginofsPecies"inl859.Incontrast,thehistoryofthedevelopmentofhypothesesontheoriginoftetrapodsdemonstratesthatdiscoveryofnewextantorfos-silforms(Tab.l)shapesourunderstandingoftherelationshipoftetrapodstofishes.Thefrstextantlungfishwasdiscoveredinl836inSouthAmerica(Fitzinge…     

Recent Progress in Understanding Early Tetrapods

This paper reviews significant discoveries and interpretationsmade for Paleozoic tetrapods over the past twenty-five years.In that span twelve significant, new localities have been found,including the oldest ever at about 370 million years in age.About 60 new genera have been described; five providing importantinsight into the early evolution of land vertebrates. The numberof exceptionally well known taxa with multiple, excellentlypreserved specimens has doubled to eight from four. The veryearliest tetrapods have been discovered to have been polydactylous,the ear region to have had a complicated early evolution andthe specialized tooth type found in Recent amphibians has beenfound in a group of Lower Permian temnospondyl amphibians, indicatingan evolutionary relationship. Perhaps the most significant advancein understanding the evolution of early tetrapods is that thebasal amniote groups have become better characterized and astart has been made in providing a defensible hypothesis fortheir relationships. The ascendancy of cladistics, functionalmorphology and plate tectonics has changed the way paleontologistsview fossils resulting in more defensible phylogenies and behavioraland biogeographical scenarios. These approaches to understandinghave, perhaps, had a more profound impact than any of the newfossil discoveries.     

Sequences,stratigraphy and scenarios: what can we say about the fossil record of the earliest tetrapods?

Past research on the emergence of digit-bearing tetrapods has led to the widely accepted premise that this important evolutionary event occurred during the Late Devonian. The discovery of convincing digit-bearing tetrapod trackways of early Middle Devonian age in Poland has upset this orthodoxy, indicating that current scenarios which link the timing of the origin of digited tetrapods to specific events in Earth history are likely to be in error. Inspired by this find, we examine the fossil record of early digit-bearing tetrapods and their closest fish-like relatives from a statistical standpoint. We find that the Polish trackways force a substantial reconsideration of the nature of the early tetrapod record when only body fossils are considered. However, the effect is less drastic (and often not statistically significant) when other reliably dated trackways that were previously considered anachronistic are taken into account. Using two approaches, we find that 95 per cent credible and confidence intervals for the origin of digit-bearing tetrapods extend into the Early Devonian and beyond, spanning late Emsian to mid Ludlow. For biologically realistic diversity models, estimated genus-level preservation rates for Devonian digited tetrapods and their relatives range from 0.025 to 0.073 per lineage-million years, an order of magnitude lower than species-level rates for groups typically considered to have dense records. Available fossils of early digited tetrapods and their immediate relatives are adequate for documenting large-scale patterns of character acquisition associated with the origin of terrestriality, but low preservation rates coupled with clear geographical and stratigraphic sampling biases caution against building scenarios for the origin of digits and terrestrialization tied to the provenance of particular specimens or faunas.     

Proposed habitats of early tetrapods: gills, kidneys, and the water-land transition

Recent finds of early tetrapods have established that the most primitive form, Acanthostega, retained internal gills and other fish-like features; this has led to the conclusion that it was a primarily aquatic animal. Other Late Devonian tetrapods, such as lchthyostega and Tulerpeton, provide no evidence of internal gills, but have also been interpreted as inhabiting an aquatic environment. The probable aquatic habits of a diversity of Devonian tetrapods has led to the suggestion that the entire early tetrapod radiation may have been an aquatic one, with terrestriality having evolved in later forms. However, consideration of the physiology of living amphibious vertebrates suggests that this scenario is unlikely. The use of the gills for the excretion of carbon dioxide and ammonia appears to be a fundamental feature of all primarily aquatic vertebrates. No living fish loses its internal gills, even if it excretes a significant portion of its nitrogenous waste as urea via the kidney in the water. Gills are simply too valuable to be lost by an aquatic animal, even in those air-breathing fishes that no longer use the gills for oxygen uptake. We suggest that the apparent loss of the gills in tetrapods more derived than Acanthostega signals their descent from a more terrestrial phase in tetrapod evolution, following the primary assumption by the kidney of the excretion of nitrogenous wastes. Without this new role of the kidney, loss of the gills would have been impossible. With this new kidney role, loss of the gills may have been advantageous in reducing desiccation on land.     

Which came first, the lung or the breath?

Lungs are the characteristic air-filled organs (AO) of the Polypteriformes, lungfish and tetrapods, whereas the swimbladder is ancestral in all other bony fish. Lungs are paired ventral derivatives of the pharynx posterior to the gills. Their respiratory blood supply is the sixth branchial artery and the venous outflow enters the heart separately from systemic and portal blood at the sinus venosus (Polypteriformes) or the atrium (lungfish), or is delivered to a separate left atrium (tetrapods). The swimbladder, on the other hand, is unpaired, and arises dorsally from the posterior pharynx. It is employed in breathing in Ginglymodi (gars), Halecomorphi (bowfin) and in basal teleosts. In most cases, its respiratory blood supply is homologous to that of the lung, but the vein drains to the cardinal veins. Separate intercardiac channels for oxygenated and deoxygenated blood are lacking. The question of the homology of lungs and swimbladders and of breathing mechanisms remains open. On the whole, air ventilatory mechanisms in the actinopterygian lineage are similar among different groups, including Polypteriformes, but are distinct from those of lungfish and tetrapods. However, there is extreme variation within this apparent dichotomy. Furthermore, the possible separate origin of air breathing in actinopterygian and 'sarcopterygian' lines is in conflict with the postulated much more ancient origin of vertebrate air-breathing organs. New studies on the isolated brainstem preparation of the gar (Lepisosteus osseus) show a pattern of efferent activity associated with a glottal opening that is remarkably similar to that seen in the in-vitro brainstem preparation of frogs and tadpoles. Given the complete lack of evidence for AO in chondrichthyans, and the isolated position of placoderms for which buoyancy organs of uncertain homology have been demonstrated, it is likely that homologous pharyngeal AO arose in the ancestors of early bony fish, and was pre-dated by behavioral mechanisms for surface (water) breathing. The primitive AO may have been the posterior gill pouches or even the modified gills themselves, served by the sixth branchial artery. Further development of the dorsal part may have led to the respiratory swimbladder, whereas the paired ventral parts evolved into lungs.     

Molecules,fossils, and the origin of tetrapods

Summary Since the discovery of the coelacanth, Latimeria chalumnae, more than 50 years ago, paleontologists and comparative morphologists have debated whether coelacanths or lungfishes, two groups of lobe-finned fishes, are the closest living relatives of land vertebrates (Tetrapoda). Previously, Meyer and Wilson (1990) determined partial DNA sequences from two conservative mitochondrial genes and found support for a close relationship of lungfishes to tetrapods. We present additional DNA sequences from the 12S rRNA mitochondria gene for three species of the two lineages of lungfishes that were not represented in the first study: Protopterus annectens and Protopterus aethiopicus from Africa and Neoceratodus forsteri (kindly provided by B. Hedges and L. Maxson) from Australia. This extended data set tends to group the two lepidosirenid lungfish lineages (Lepidosiren and Protopterus) with Neoceratodus as their sister group. All lungfishes seem to be more closely related to tetrapods than the coelacanth is. This result appears to rule out the possibility that the coelacanth lineage gave rise to land vertebrates. The common ancestor of lungfishes and tetrapods might have possessed multiple morphological traits that are shared by lungfishes and tetrapods [Meyer and Wilson (1990) listed 14 such traits]. Those traits that seem to link Latimeria and tetrapods are arguably due to convergent evolution or reversals and not to common descent. In this way, the molecular tree facilitates an evolutionary interpretation of the morphological differences among the living forms. We recommended that the extinct groups of lobe-finned fishes be placed onto the molecular tree that has lungfishes and not the coelacanth more closely related to tetrapods. The placement of fossils would help to further interpret the sequence of morphological events and innovations associated with the origin of tetrapods but appears to be problematic because the quality of fossils is not always high enough, and differences among paleontologists in the interpretation of the fossils have stood in the way of a consensus opinion for the branching order among lobefinned fishes. Marshall and Schultze (1992) criticized the morphological analysis presented by Meyer and Wilson (1990) and suggest that 13 of the 14 morphological traits that support the sister group relationship of lungfishes and tetrapods are not shared derived characters. Here we present further alternative viewpoints to the ones of Marshall and Schultze (1992) from the paleontological literature. We argue that all available information (paleontological, neontological, and molecular data) and rigorous cladistic methodology should be used when relating fossils and extant taxa in a phylogenetic framework. Offprint requests to: Axel Meyer     

Lessons from parasitic flatworms about evolution and historical biogeography of their vertebrate hosts

Cophylogenetic studies investigate the evolutionary trends within host-parasite associations. Examination of the different levels of fidelity between host and parasite phylogenies provides a powerful tool to inspect patterns and processes of parasite diversification over host evolution and geological times. Within the phylum Platyhelminthes, the monogeneans are mainly fish parasites. The Polystomatidae, however, are known from the sarcopterygian Australian lungfish and tetrapods such as amphibians, freshwater turtles, and the African hippopotamus. Cophylogenetic and biogeographic vicariance analyses, supplemented by molecular calibrations, showed that the Polystomatidae may track the evolutionary history of the first aquatic tetrapods in the Palaeozoic age. Evolutionary lines of the major polystome lineages would also be intimately related to the evolution of their hosts over hundreds of millions years. Since the Mesozoic, evolution of polystomes would have been shaped mainly by plate tectonics during the break-up of Gondwanaland and subsequent dispersal of ancestral neobatrachian host lineages. Therefore the Polystomatidae could serve as a novel model to improve cophylogenetic tools and to inspect a suite of questions about the evolution of vertebrate hosts. To cite this article: O. Verneau et al., C. R. Biologies 332 (2009).     

The complete mitochondrial DNA sequence of the shark Mustelus manazo: evaluating rooting contradictions to living bony vertebrates

A remarkable example of a misleading mitochondrial protein tree is presented, involving ray-finned fishes, coelacanths, lungfishes, and tetrapods, with sea lampreys as an outgroup. In previous molecular phylogenetic studies on the origin of tetrapods, ray-finned fishes have been assumed as an outgroup to the tetrapod/lungfish/coelacanth clade, an assumption supported by morphological evidence. Standard methods of molecular phylogenetics applied to the protein-encoding genes of mitochondria, however, give a bizarre tree in which lamprey groups with lungfish and, therefore, ray-finned fishes are not the outgroup to a tetrapod/lungfish/coelacanth clade. All of the dozens of published phylogenetic methods, including every possible modification to maximum likelihood known to us (such as inclusion of site heterogeneity and exclusion of potentially misleading hydrophobic amino acids), fail to place the ray-finned fishes in a biologically acceptable position. A likely cause of this failure may be the use of an inappropriate outgroup. Accordingly, we have determined the complete mitochondrial DNA sequence from the shark, Mustelus manazo, which we have used as an alternative and more proximal outgroup than the lamprey. Using sharks as the outgroup, lungfish appear to be the closest living relative of tetrapods, although the possibility of a lungfish/coelacanth clade being the sister group of tetrapods cannot be excluded.      

The stapes of the Goal Measures embolomere Pholiderpeton scutigerum Huxley (Amphibia: Anthracosauria) and otic evolution in early tetrapods

Middle ear structure has been of interest for a long time in studies of the origins and relationships of early tetrapod groups. The model of a dorsally-directed, rod-like stapes with a tympanum, thought common to labyrinthodont amphibians, was taken to be primitive for tetrapods. The stapes of embolomeres and other early anthracosaurs were assumed to be of this form, but difficulties resulted if the middle ear structure of fossil and living reptiles was considered ultimately derived from this source.
The embolomere stapes has been identified and does not conform to the predicted model. It most closely resembles that of Greererpeton , an early notchless temnospondyl. The stapes is compared with those of other tetrapods in terms of the theoretical five processes. An interpretation is put forward in which all but the opercular are seen as potentially present. The embolomere stapes is compared with that of Greererpeton in terms of recent theories of mechanical function and is seen to weaken them. They are then compared as part of a possible acoustic mechanism. The embolomere middle ear structure is reinterpreted as a receiver for low-frequency sound and the 'otic notch' is not considered to have housed a tympanum.
The resemblance between the stapes of these two animals seems best explained by their closeness to the plesiomorphic condition for tetrapods, a conclusion which forces the abandonment of the concept of a 'labyrinthodont middle ear'. The middle ear structure of later groups can be interpreted as having evolved from one similar to that seen in these two animals. The conclusion supports those reached in other recent papers that tympana were not primitive for tetrapods but have been independently derived in several groups.     

Getting a grip on tetrapod grasping: form,function, and evolution

Human beings have been credited with unparalleled capabilities for digital prehension grasping. However, grasping behaviour is widespread among tetrapods. The propensity to grasp, and the anatomical characteristics that underlie it, appear in all of the major groups of tetrapods with the possible exception of terrestrial turtles. Although some features are synapomorphic to the tetrapod clade, such as well‐defined digits and digital musculature, other features, such as opposable digits and tendon configurations, appear to have evolved independently in many lineages. Here we examine the incidence, functional morphology, and evolution of grasping across four major tetrapod clades. Our review suggests that the ability to grasp with the manus and pes is considerably more widespread, and ecologically and evolutionarily important, than previously thought. The morphological bases and ecological factors that govern grasping abilities may differ among tetrapods, yet the selective forces shaping them are likely similar. We suggest that further investigation into grasping form and function within and among these clades may expose a greater role for grasping ability in the evolutionary success of many tetrapod lineages.     

Overexpression of homologous phytochrome genes in tomato: exploring the limits in photoperception

Transgenic tomato [Lycopersicon esculentum (=Solanum lycopersicum)] lines overexpressing tomato PHYA, PHYB1, or PHYB2, under control of the constitutive double-35S promoter from cauliflower mosaic virus (CaMV) have been generated to test the level of saturation in individual phytochrome-signalling pathways in tomato. Western blot analysis confirmed the elevated phytochrome protein levels in dark-grown seedlings of the respective PHY overexpressing (PHYOE) lines. Exposure to 4 h of red light resulted in a decrease in phytochrome A protein level in the PHYAOE lines, indicating that the chromophore availability is not limiting for assembly into holoprotein and that the excess of phytochrome A protein is also targeted for light-regulated destruction. The elongation and anthocyanin accumulation responses of plants grown under white light, red light, far-red light, and end-of-day far-red light were used for characterization of selected PHYOE lines. In addition, the anthocyanin accumulation response to different fluence rates of red light of 4-d-old dark-grown seedlings was studied. The elevated levels of phyA in the PHYAOE lines had little effect on seedling and adult plant phenotype. Both PHYAOE in the phyA mutant background and PHYB2OE in the double-mutant background rescued the mutant phenotype, proving that expression of the transgene results in biologically active phytochrome. The PHYB1OE lines showed mild effects on the inhibition of stem elongation and anthocyanin accumulation and little or no effect on the red light high irradiance response. By contrast, the PHYB2OE lines showed a strong inhibition of elongation, enhancement of anthocyanin accumulation, and a strong amplification of the red light high irradiance response.     

Devonian tetrapod trackways and trackmakers; a review of the fossils and footprints

The earliest tetrapods are known from the Upper Devonian. Their remains are becoming better known from increasing numbers of specimens, localities, environments and ichnofossils. Each of the eight (or possibly nine) genera now represented by skeletal fossils is reviewed in its sedimentological, faunal and stratigraphic context, with an assessment of what might be inferred about the habitus and locomotory capabilities of each. Fossil trackways and their interpretations are then re-examined in the context of the known body forms, and consideration given to the degree of fit between the skeletal fossils, the trackways and their interpretations. The currently known Devonian tetrapods are unlikely to have made any of the known tracks, unless they were produced under water. Neither the skeletal fossils nor the trackways show good evidence of terrestrial locomotion among Devonian tetrapods. When the fossil material and recent phylogenetic analyses are taken in combination, it appears that neither tetrapods nor limbs with digits are likely to have arisen before the Frasnian. This should be borne in mind in palaeoecological studies of these animals.     

Skeletal Morphogenesis of Microbrachis and Hyloplesion (Tetrapoda: Lepospondyli), and Implications for the Developmental Patterns of Extinct,Early Tetrapods

The ontogeny of extant amphibians often is used as a model for that of extinct early tetrapods, despite evidence for a spectrum of developmental modes in temnospondyls and a paucity of ontogenetic data for lepospondyls. I describe the skeletal morphogenesis of the extinct lepospondyls Microbrachis pelikani and Hyloplesion longicostatum using the largest samples examined for either taxon. Nearly all known specimens were re-examined, allowing for substantial anatomical revisions that affect the scoring of characters commonly used in phylogenetic analyses of early tetrapods. The palate of H. longicostatum is re-interpreted and suggested to be more similar to that of M. pelikani, especially in the nature of the contact between the pterygoids. Both taxa possess lateral lines, and M. pelikani additionally exhibits branchial plates. However, early and rapid ossification of the postcranial skeleton, including a well-developed pubis and ossified epipodials, suggests that neither taxon metamorphosed nor were they neotenic in the sense of branchiosaurids and salamanders. Morphogenetic patterns in the foot suggest that digit 5 was developmentally delayed and the final digit to ossify in M. pelikani and H. longicostatum. Overall patterns of postcranial ossification may indicate postaxial dominance in limb and digit formation, but also more developmental variation in early tetrapods than has been appreciated. The phylogenetic position and developmental patterns of M. pelikani and H. longicostatum are congruent with the hypothesis that early tetrapods lacked metamorphosis ancestrally and that stem-amniotes exhibited derived features of development, such as rapid and complete ossification of the skeleton, potentially prior to the evolution of the amniotic egg.