Tail use improves performance on soft substrates in models of early vertebrate land locomotors

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Abstract

In the evolutionary transition from an aquatic to a terrestrial environment, early tetrapods faced the challenges of terrestrial locomotion on flowable substrates, such as sand and mud of variable stiffness and incline. The morphology and range of motion of appendages can be revealed in fossils; however, biological and robophysical studies of modern taxa have shown that movement on such substrates can be sensitive to small changes in appendage use. Using a biological model (the mudskipper), a physical robot model, granular drag measurements, and theoretical tools from geometric mechanics, we demonstrate how tail use can improve robustness to variable limb use and substrate conditions. We hypothesize that properly coordinated tail movements could have provided a substantial benefit for the earliest vertebrates to move on land.

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A Devonian tetrapod-like fish and the evolution of the tetrapod body plan.

Edward B Daeschler, Neil H Shubin, Farish Jenkins (2006)

The relationship of limbed vertebrates (tetrapods) to lobe-finned fish (sarcopterygians) is well established, but the origin of major tetrapod features has remained obscure for lack of fossils that document the sequence of evolutionary changes. Here we report the discovery of a well-preserved species of fossil sarcopterygian fish from the Late Devonian of Arctic Canada that represents an intermediate between fish with fins and tetrapods with limbs, and provides unique insights into how and in what order important tetrapod characters arose. Although the body scales, fin rays, lower jaw and palate are comparable to those in more primitive sarcopterygians, the new species also has a shortened skull roof, a modified ear region, a mobile neck, a functional wrist joint, and other features that presage tetrapod conditions. The morphological features and geological setting of this new animal are suggestive of life in shallow-water, marginal and subaerial habitats.

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Undulatory swimming in sand: subsurface locomotion of the sandfish lizard.

Yang Ding, Chen Li, Ryan D Maladen … (2009)

The desert-dwelling sandfish (Scincus scincus) moves within dry sand, a material that displays solid and fluidlike behavior. High-speed x-ray imaging shows that below the surface, the lizard no longer uses limbs for propulsion but generates thrust to overcome drag by propagating an undulatory traveling wave down the body. Although viscous hydrodynamics can predict swimming speed in fluids such as water, an equivalent theory for granular drag is not available. To predict sandfish swimming speed, we developed an empirical model by measuring granular drag force on a small cylinder oriented at different angles relative to the displacement direction and summing these forces over the animal movement profile. The agreement between model and experiment implies that the noninertial swimming occurs in a frictional fluid.

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The Devonian tetrapod Acanthostega gunnari Jarvik: postcranial anatomy, basal tetrapod interrelationships and patterns of skeletal evolution

M I Coates (1996)

The postcranial skeleton ofAcanthostega gunnarifrom the Famennian of East Greenland displays a unique, transitional, mixture of features conventionally associated with fishand tetrapod-like morphologies. The rhachitomous vertebral column has a primitive, barely differentiated atlas-axis complex, encloses an unconstricted notochordal canal, and the weakly ossified neural arches have poorly developed zygapophyses. More derived axial skeletal features include caudal vertebral proliferation and, transiently, neural radials supporting unbranched and unsegmented lepidotrichia. Sacral and post-sacral ribs reiterate uncinate cervical and anterior thoracic rib morphologies: a simple distal flange supplies a broad surface for iliac attachment. The octodactylous forelimb and hindlimb each articulate with an unsutured, foraminate endoskeletal girdle. A broad-bladed femoral shaft with extreme anterior torsion and associated flattened epipodials indicates a paddle-like hindlimb function. Phylogenetic analysis placesAcanthostegaas the sister-group ofIchthyostegaplus all more advanced tetrapods.Tulerpetonappears to be a basal stemamniote plesion, tying the amphibian-amniote split to the uppermost Devonian.Caerorhachismay represent a more derived stem-amniote plesion. Postcranial evolutionary trends spanning the taxa traditionally associated with the fish-tetrapod transition are discussed in detail. Comparison between axial skeletons of primitive tetrapods suggests that plesiomorphic fish-like morphologies were re-patterned in a cranio-caudal direction with the emergence of tetrapod vertebral regionalisation. The evolution of digited limbs lags behind the initial enlargement of endoskeletal girdles, whereas digit evolution precedes the elaboration of complex carpal and tarsal articulations. Pentadactylous limbs appear to have stabilised independently in amniote and amphibian lineages; the colosteidGreererpetonhas a pentadactylous manus, indicating that basal amphibian forelimbs may not be restricted to patterns of four digits or less.