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1.
A parsimony analysis of 'higher' temnospondyls (all temnospondyls descended from the common ancestor of Eryops and Parotosuchus ) was performed using 37 terminal taxa and 121 osteological characters. Bremer support values for each internal node were calculated as a measure of clade strength. Additionally, the shortest trees that conformed to some alternative hypotheses were searched for. The following new taxa are established on the basis of the results: Euskelia (the clade containing the Eryopoidea and Dissorophoidea), Limnarchia (the clade containing Trimerorhachidae, Dvinosauroidea, Archegosauroidea and Stereospondyli), Dvinosauria (the clade containing Trimerorhachidae and Dvinosauroidea), Stereo-spondylomorpha (the clade containing Archegosauroidea and Stereospondyli), Capitosauria (the clade containing Lydekkerina and 'capitosauroids'), and Trematosauria (the clade containing Trematosauroidea, Rhytidosteidae, Plagiosauroidea, Metoposauroidea and Brachyopoidea). The monophyly of the assemblage of Mesozoic families called the Stereospondyli by Romer is supported. The dominance of the Stereospondyli in the Mesozoic and its rarity in the Palaeozoic is discussed. It is suggested that the radiation of the diverse stereospondyl clades, the Capitosauria and Trematosauria, began in the Late Permian of Gondwana, in a 'safe haven' that was less severely affected by the Late Permian extinction event. It is further speculated that the 'safe haven' was located in Antarctica, or possibly Australia.  相似文献   

2.
The fossil Tiktaalik roseae from the Late Devonian induces clear definition of the biomechanical and functional preconditions for a terrestrial lifestyle including quadrupedal standing and locomotion on limbs. Therefore, we determined the internal stresses in this model organism under the influence of gravity using the finite element method. Stress patterns during symmetrical two-forelimb support result from bending of trunk and neck. During asymmetrical one-forelimb support, as occurs during terrestrial locomotion, torsional stresses are higher than those caused by bending. The observed patterns of compressive stresses correspond well with the arrangement of compression-resistant materials: vertebral column, shoulder girdle and ribs. The tensile stresses are in accordance with the arrangement of longitudinal and oblique muscles forming the body wall. Torsional stresses concentrate along the periphery of the trunk, leaving its cavity free from mechanical stresses. Theoretical mechanics indicate that the flat skull and the mobility of the neck were advantageous for lateral snapping, similar to crocodiles. The same movement on land requires sprawling and flexed forelimbs. Our results can be interpreted as explanations for the tetrapod bauplan as well as confirmation and refinement of existing hypotheses about the lifestyle at the border between water and land of this early predecessor of terrestrial tetrapods.  相似文献   

3.
Witzmann F. (2011). Morphological and histological changes of dermal scales during the fish‐to‐tetrapod transition. —Acta Zoologica (Stockholm) 92 : 281–302. The gastral scales of limbed tetrapodomorphs evolved from the ‘elpistostegid’‐type of scale by an enlargement and differentiation of the articulation facets and a shortening and broadening of the keel. These changes caused a tighter connection between gastral scales within a scale row and a greater overlap between the rows. Dorsal round scales of limbed tetrapodomorphs developed from a gastral scale‐type by an alteration of the ontogenetic pathway. The posterolateral direction of scale rows in ‘elpistostegids’ was retained in the gastral scalation of most limbed tetrapodomorphs, whereas the arrangement of round dorsal scales is modified to a transverse orientation. Both gastral and dorsal scales of limbed tetrapodomorphs consist solely of parallel‐fibred bone with circumferential growth marks. The proportionally larger overlap surfaces of gastral scales and their mode of articulation in the ventral midline indicate that the body of limbed tetrapodomorphs might have been more flexible than that of their finned relatives. The alteration of dermal scales was one of the most rapid morphological changes during the fish‐to‐tetrapod transition. Once established, gastral and dorsal scales were retained as a conservative character in different lineages of basal tetrapods, in both the amphibian and the amniote lineages.  相似文献   

4.
Extinction risk in the modern world and extinction in the geological past are often linked to aspects of life history or other facets of biology that are phylogenetically conserved within clades. These links can result in phylogenetic clustering of extinction, a measurement comparable across different clades and time periods that can be made in the absence of detailed trait data. This phylogenetic approach is particularly suitable for vertebrate taxa, which often have fragmentary fossil records, but robust, cladistically‐inferred trees. Here we use simulations to investigate the adequacy of measures of phylogenetic clustering of extinction when applied to phylogenies of fossil taxa while assuming a Brownian motion model of trait evolution. We characterize expected biases under a variety of evolutionary and analytical scenarios. Recovery of accurate estimates of extinction clustering depends heavily on the sampling rate, and results can be highly variable across topologies. Clustering is often underestimated at low sampling rates, whereas at high sampling rates it is always overestimated. Sampling rate dictates which cladogram timescaling method will produce the most accurate results, as well as how much of a bias ancestor–descendant pairs introduce. We illustrate this approach by applying two phylogenetic metrics of extinction clustering (Fritz and Purvis's D and Moran's I) to three tetrapod clades across an interval including the Permo‐Triassic mass extinction event. These groups consistently show phylogenetic clustering of extinction, unrelated to change in other quantitative metrics such as taxonomic diversity or extinction intensity.  相似文献   

5.
Marine reptiles and mammals are phylogenetically so distant from each other that their marine adaptations are rarely compared directly. We reviewed ecophysiological features in extant non-avian marine tetrapods representing 31 marine colonizations to test whether there is a common pattern across higher taxonomic groups, such as mammals and reptiles. Marine adaptations in tetrapods can be roughly divided into aquatic and haline adaptations, each of which seems to follow a sequence of three steps. In combination, these six categories exhibit five steps of marine adaptation that apply across all clades except snakes: Step M1, incipient use of marine resources; Step M2, direct feeding in the saline sea; Step M3, water balance maintenance without terrestrial fresh water; Step M4, minimized terrestrial travel and loss of terrestrial feeding; and Step M5, loss of terrestrial thermoregulation and fur/plumage. Acquisition of viviparity is not included because there is no known case where viviparity evolved after a tetrapod lineage colonized the sea. A similar sequence is found in snakes but with the haline adaptation step (Step M3) lagging behind aquatic adaptation (haline adaptation is Step S5 in snakes), most likely because their unique method of water balance maintenance requires a supply of fresh water. The same constraint may limit the maximum body size of fully marine snakes. Steps M4 and M5 in all taxa except snakes are associated with skeletal adaptations that are mechanistically linked to relevant ecophysiological features, allowing assessment of marine adaptation steps in some fossil marine tetrapods. We identified four fossil clades containing members that reached Step M5 outside of stem whales, pinnipeds, sea cows and sea turtles, namely Eosauropterygia, Ichthyosauromorpha, Mosasauroidea, and Thalattosuchia, while five other clades reached Step M4: Saurosphargidae, Placodontia, Dinocephalosaurus, Desmostylia, and Odontochelys. Clades reaching Steps M4 and M5, both extant and extinct, appear to have higher species diversity than those only reaching Steps M1 to M3, while the total number of clades is higher for the earlier steps. This suggests that marine colonizers only diversified greatly after they minimized their use of terrestrial resources, with many lineages not reaching these advanced steps. Historical patterns suggest that a clade does not advance to Steps M4 and M5 unless these steps are reached early in the evolution of the clade. Intermediate forms before a clade reached Steps M4 and M5 tend to become extinct without leaving extant descendants or fossil evidence. This makes it difficult to reconstruct the evolutionary history of marine adaptation in many clades. Clades that reached Steps M4 and M5 tend to last longer than other marine tetrapod clades, sometimes for more than 100 million years.  相似文献   

6.
One of the most important physiological changes during the conquest of land by vertebrates was the increasing reliance on lung breathing, with the concomitant decrease in importance of gill breathing. The main problem involved here was to cope with the excessive accumulation of CO2 in the body and to avoid respiratory acidosis. In the past, several often mutually contradicting hypotheses of CO2‐elimination via skin, lungs and gills in early tetrapods have been proposed, based on theoretical physiological considerations and comparison with extant air‐breathing fishes and amphibians. This study proposes a revised scenario of CO2‐elimination in early tetrapods based on fossil evidence, that is recently identified osteological correlates of gills, skin structure and mode of lung ventilation. In stem tetrapods of the Devonian and Carboniferous, O2‐uptake via the lungs by buccal pumping was decoupled from CO2‐release via internal gills, and the rather gas‐impermeable skin played a minor role in gaseous exchange. The two main lineages of crown‐group tetrapods, the amphibian and amniote lineage, used different strategies of CO2‐elimination. As in stem tetrapods, O2‐uptake and CO2‐release remained always largely decoupled in temnospondyls, which ventilated their lungs via buccal pumping and relied mainly on their internal gills for CO2‐release. Temnospondyls were not able to reduce their internal gills before their skin became more gas permeable and their body size was reduced, to shift from internal gills to the skin as the major site of CO2‐elimination, a pattern that is retained in most lissamphibians. In contrast, internal gills were lost very early in stem amniote evolution. This was associated with the evolution of the more effective aspiration pump that allowed the elimination of the bulk of CO2 via the lungs, leading to a coupled O2‐uptake and CO2‐loss in stem amniotes and later in amniotes.  相似文献   

7.
A partial skull from the Lower Keuper (Middle Triassic) of Germany is recognized as belonging to a new genus and species of plagiosaurid temnospondyl. It is readily identified by the following autapomorphies: (1) extremely large orbits medially extended to give very thin interorbital region and cheek; (2) posterior skull table abbreviated, with splint‐like supratemporals, postparietals and parietals; (3) supraorbital lateral line sulcus absent on frontal, with blind ending on parietal, and continued at the anterior margin of the postorbital; (4) occiput sloping posteriorly, with subtympanic fossa exposed in ventral view; (5) cultriform process and basicranial suture extremely narrow; (6) mandibular and maxillary teeth very long with crowns markedly curved inwards. The new taxon shares the following derived character states with Plagiosternum: (1) the pentagonal shape of orbits; (2) the slender interorbital region and cultriform process; (3) the small‐scale, polygonal pit‐and‐ridge ornament on posterior skull table, with single prominent tubercles rising from ridges. Phylogenetic analysis finds plagiosaurids to be monophyletic within a broad range of temnospondyls, nesting within a clade of short‐skulled stereospondyls. Plagio‐suchus is the most basal plagiosaurid, and Plagiosternum forms the sister group to the new taxon. The biological significance of the large orbits of plagiosaurids and their relationships to the eyeballs is discussed. The only type of eyeball that would be feasible for all known plagiosaurids would be a small spherical structure possibly situated near the anterior edge of the orbit. For those plagiosaurs with extremely large shallow orbits like Plagiosternum and Megalophthalma, a possible adaptation would be a lens‐less eye comprising a flat retinal plate across the entire orbit.  相似文献   

8.
Abstract: The complete neurocranium plus palatoquadrate of the plagiosaurid temnospondyl Gerrothorax pulcherrimus from the Middle Triassic of Germany is described for the first time, based on outer morphological observations and micro‐CT scanning. The exoccipitals are strong elements with paroccipital processes and well‐separated occipital condyles. Anterolaterally, the exoccipitals contact the otics, which are mediolaterally elongated and have massive lateral walls. The otics contact the basisphenoid, which shows well‐developed sellar processes. Anteriorly, the basisphenoid is continuous with the sphenethmoid region. In its posterior portion, the sphenethmoid gives rise to robust, laterally directed laterosphenoid walls, a unique morphology among basal tetrapods. The palatoquadrate is extensively ossified. The quadrate portion overlaps the descending lamina of squamosal and ascending lamina of pterygoid anteriorly, almost contacting the epipterygoid laterally. The epipterygoid is a complex element and may be co‐ossified with otics and laterosphenoid walls. It has a broad, sheet‐like footplate and a horizontally aligned ascending process that contacts the laterosphenoid walls. The degree of ossification of the epipterygoid, however, is subject to individual variation obviously independent from ontogenetic changes. The stapes of Gerrothorax is a large, blade‐like element that differs conspicuously from the plesiomorphic temnospondyl condition. It has a prominent anterolateral projection which has not been observed in other basal tetrapods. Morphology of neurocranium and palatoquadratum of Gerrothorax most closely resembles that of the Russian plagiosaurid Plagiosternum danilovi, although the elements are less ossified in the latter. The extensive endocranial ossification of Gerrothorax is consistent with the general high degree of ossification in the exo‐ and endoskeleton of this temnospondyl and supports the view that a strong endocranial ossification cannot be evaluated as a plesiomorphic character in basal tetrapods.  相似文献   

9.
In the present study, effects of the trematode eye fluke Diplostomum spathaceum infection and parasite‐induced cataract on energetic costs and feeding of hatchery‐bred one‐summer‐old Lake Saimaa Arctic charr Salvelinus alpinus were examined. The energetic effects (measured as oxygen uptake) of acute eye fluke infection on S. alpinus were negligible, but in the chronic infections mean oxygen consumption of infected fish was significantly higher than that of controls. Feeding capability of S. alpinus was studied in aquarium experiments, and uninfected healthy‐eyed fish reacted to zooplankton more rapidly and caught zooplankton with better success compared to individuals with cataracts in the same school. The results suggested that to compensate energetic costs and impaired feeding capability caused by eye fluke, cataract‐bearing fish spend more time foraging, which may result in increased predation risk.  相似文献   

10.
Summary. We used two approaches to identify sequence variants in ionotropic glutamate receptor (IGR) genes: high-throughput screening and resequencing techniques, and information mining of public (e.g. dbSNP, ENSEMBL) and private (i.e. Celera Discovery System) sequence databases. Each of the 16 known IGRs is represented in these databases, their positions on a canonical physical map are established. Comparisons of mouse, rat, and human sequences revealed substantial conservation among these genes, which are located on different chromosomes but found within syntenic groups of genes. The IGRs are members of a phylogenetically ancient gene family, sharing similarities with glutamate-like receptors in plants. Parsimony analysis of amino acid sequences groups the IGRs into three distinct clades based on ligand-binding specificity and structural features, such as the channel pore and membrane spanning domains. A collection of 38 variants with amino acid changes was obtained by combining screening, resequencing, and informatics approaches for several of the IGR genes. This represents only a fraction of the sequence variation across these genes, but in fact these may constitute a large fraction of the common polymorphisms at these genes and these polymorphisms are a starting point for understanding the role of these variants in function.Genetically influenced human neurobehavioral phenotypes are likely to be linked to IGR genetic variants. Because ionotropic glutamate receptor activation leads to calcium entry, which is fundamental in brain development and in forms of synaptic plasticity essential for learning and memory and is essential for neuronal survival, it is likely that sequence variants in IGR genes may have profound functional roles in neuronal activation and survival mechanisms.  相似文献   

11.
12.
Butterflies of the genus Maculinea are highly endangered in Europe. The cuckoo species M. rebeli has been thoroughly investigated through both empirical and modelling studies, but less is known about the population ecology of predatory Maculinea. We present the findings of a 2-year research study on sympatric populations of two endangered butterflies: Maculinea teleius and M. nausithous in the Kraków region, southern Poland. The study comprised mark–release–recapture sampling and laboratory rearing of butterflies from larvae collected in the field. For both species the sex ratio was slightly, but consistently, female-biased and there was little year-to-year change in the seasonal population sizes. Daily numbers showed greater variation between the 2 years of the study due to the differences in daily survival rate. The average life span of laboratory-raised butterflies kept in ideal conditions was more than 6 days, compared to only 2–3 days in the field. The recruitment of both males and females consistently followed a bimodal pattern. A small proportion of individuals (maximum 25%) changed sites, in spite of relatively short distances of ca. 100 m separating them. The results indicate that populations of both species are typically stable within their sites, possibly due to larval polymorphism, but there is little inter-site mobility and thus landscape corridors seem necessary to enhance metapopulation viability. A further problem to be considered in the conservation of Maculinea butterflies is the fact that their very short life span in relation to flight-period length reduces the effective population size.  相似文献   

13.
The origin and evolution of the vertebrate skull have been topics of intense study for more than two centuries. Whereas early theories of skull origin, such as the influential vertebral theory, have been largely refuted with respect to the anterior (pre‐otic) region of the skull, the posterior (post‐otic) region is known to be derived from the anteriormost paraxial segments, i.e. the somites. Here we review the morphology and development of the occiput in both living and extinct tetrapods, taking into account revised knowledge of skull development by augmenting historical accounts with recent data. When occipital composition is evaluated relative to its position along the neural axis, and specifically to the hypoglossal nerve complex, much of the apparent interspecific variation in the location of the skull–neck boundary stabilizes in a phylogenetically informative way. Based on this criterion, three distinct conditions are identified in (i) frogs, (ii) salamanders and caecilians, and (iii) amniotes. The position of the posteriormost occipital segment relative to the hypoglossal nerve is key to understanding the evolution of the posterior limit of the skull. By using cranial foramina as osteological proxies of the hypoglossal nerve, a survey of fossil taxa reveals the amniote condition to be present at the base of Tetrapoda. This result challenges traditional theories of cranial evolution, which posit translocation of the occiput to a more posterior location in amniotes relative to lissamphibians (frogs, salamanders, caecilians), and instead supports the largely overlooked hypothesis that the reduced occiput in lissamphibians is secondarily derived. Recent advances in our understanding of the genetic basis of axial patterning and its regulation in amniotes support the hypothesis that the lissamphibian occipital form may have arisen as the product of a homeotic shift in segment fate from an amniote‐like condition.  相似文献   

14.
15.
Buchwitz, M., Witzmann, F., Voigt, S. and Golubev, V. 2012. Osteoderm microstructure indicates the presence of a crocodylian‐like trunk bracing system in a group of armoured basal tetrapods. —Acta Zoologica (Stockholm) 93 : 260–280. The microstructure of dorsal osteoderms referred to the chroniosuchid taxa Chroniosuchus, Chroniosaurus, Madygenerpeton and cf. Uralerpeton is compared to existing data on the bystrowianid chroniosuchian Bystrowiella and further tetrapods. Chroniosuchid osteoderms are marked by thin internal and relatively thick external cortices that consist of lowly vascularised parallel‐fibred bone. They are structured by growth marks and, in case of Madygenerpeton, by lines of arrested growth. The cancellous middle region is marked by a high degree of remodelling and a primary bone matrix of parallel‐fibred bone that may include domains of interwoven structural fibres. Whereas the convergence of Bystrowiella and chroniosuchid osteoderms is not confirmed by our observations, the internal cortex of the latter displays a significant peculiarity: It contains distinct bundles of shallowly dipping Sharpey’s fibres with a cranio‐ or caudoventral orientation. We interpret this feature as indicative for the attachment of epaxial muscles which spanned several vertebral segments between the medioventral surface of the osteoderms and the transversal processes of the thoracic vertebrae. This finding endorses the hypothesis that the chroniosuchid osteoderm series was part of a crocodylian‐like trunk bracing system that supported terrestrial locomotion. According to the measured range of osteoderm bone compactness, some chroniosuchian species may have had a more aquatic lifestyle than others.  相似文献   

16.
Twenty modified‐Whittaker plots were stratified at different sampling locations from February to May of 2008 in the central zone of Korup National Park, Cameroon. Our interest was to assess floristic diversity and investigate their relationship with environmental variables. Diversity profiles and rank abundance–curves were used for diversity analysis while canonical correspondence analysis and species–response curves were used to investigate the relationships between the response and explanatory variables. Of the 66 families identified, the Rubiaceae (999 species) were the most abundant. The Sterculiaceae (basal area = 10.482 m2 ha?1) were the dominant family, while the co‐dominant families included the Ebenaceae (basal area = 9.092 m2 ha?1) and the Euphorbiaceae (basal area = 8.168 m2 ha?1). Soil variables explained 54.3% of total variation in family distribution. Canonical axes were related to different environmental gradients: axis1 was related to increasing canopy cover (r = 0.6951); axis 2, increasing Magnesium (r = 0.8465) and effective cation exchange capacity (r = 0.5899); axis 3, increasing effective cation exchange capacity (r = 0.5536); while axis 4, increasing Phosphorus concentration (r = 0.5232). Our results demonstrate the advantage which diversity profiles have over single or combination of indices, and the importance of using a combination of methodologies in diversity analysis.  相似文献   

17.
Morphological and molecular phylogenetic studies show that there is a close relationship between Coffea and Psilanthus. In this study we reassess species relationships based on improved species sampling for Psilanthus, including P. melanocarpus, a species that shares morpho‐taxonomic characters of both genera. Analyses are performed using parsimony and Bayesian inference, on sequence data from four plastid regions [trnL–F intron, trnL–F IGS, rpl16 intron and accD–psa1 intergenic spacer (IGS)] and the internal transcribed spacer (ITS) region of nuclear ribosomal DNA (ITS 1/5.8S/ITS 2). Several major lineages with geographical coherence, as identified in previous studies based on smaller and larger data sets, are supported. Our results also confirm previous studies showing that the level of sequence divergence between Coffea and Psilanthus species is negligible, particularly given the much longer branch lengths separating other genera of tribe Coffeeae. There are strong indications that neither Psilanthus nor Coffea is monophyletic. Psilanthus melanocarpus is nested with the CoffeaPsilanthus clade, which means that there is only one critical difference between Coffea and Psilanthus; the former has a long‐emergent style and the latter a short, included style. Based on these new data, in addition to other systematically informative evidence from a broad range of studies, and especially morphology, Psilanthus is subsumed into Coffea. This decision increases the number of species in Coffea from 104 to 124, extends the distribution to tropical Asia and Australasia and broadens the morphological characterization of the genus. The implications for understanding the evolutionary history of Coffea are discussed. A group of closely related species is informally named the ‘Coffea liberica alliance’. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 357–377.  相似文献   

18.
Mature males of nurseryfish have a hook on their head to which the eggs become attached and are carried like a bunch of grapes. This paper examines the anatomy and histology of the hook. The osteological basis of the hook is shown to be a modification of the supraoccipital crest of the skull covered by typical teleost skin. The integument in the cleft of the hook, where the eggs are attached, is considerably different from ordinary fish skin. The stratified epidermis is devoid of secretory mucus and neurosensory cells and is folded into crypts that extend deeply into the dermis. This may be a specialization that facilitates adhesion of the sticky egg mass. Field observations show that this cleft area of the hook is edematous, and histology confirms that the area is highly vascularized. We speculate that this may facilitate gas exchange and/or nutrition between the male and the egg mass, but this can only be confirmed by physiological experiments with pregnant males in captivity. Engorgement with blood in the highly vascularised dermis of the hook may help hold the egg mass in place.  相似文献   

19.
The Upper Permian seymouriamorph tetrapod Karpinskiosaurus from European Russia includes two species: Karpinskiosaurus secundus and Karpinskiosaurus ultimus. Karpinskiosaurus secundus is represented by two specimens with skull lengths of about 75 mm. All specimens of K. ultimus are smaller than those of K. secundus. Revision of the cranial anatomy of all previously known and several new specimens of Karpinskiosaurus shows that the specimens of K. secundus and most of the specimens of K. ultimus represent the ontogenetic series of one species: K. secundus. The holotype specimen of K. ultimus requires revision, with the aim to find out whether it represents a second species of Karpinskiosaurus or not. The available material permits new reconstructions of the largest, holotype skull, and one smaller skull with a length of about 36 mm. Karpinskiosaurus secundus is included in a cladistic analysis for the first time here. The analysis shows it to form a sister taxon to Discosauriscidae. The clade comprising Karpinskiosaurus secundus plus Discosauriscidae forms a sister group to Seymouriidae. Karpinskiosaurus secundus has a large postorbital and a short preorbital region, and the orbits are placed in the posterior portion of the anterior half of the skull length. Among all seymouriamorphs, such cranial proportions are exhibited only by the largest known specimens of Discosauriscus austriacus. None of the specimens of K. secundus described here exhibits the presence of sensory grooves; thus, all specimens composing the ontogenetic sequence of K. secundus are considered to be terrestrial. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.  相似文献   

20.
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