The pelvic fin and girdle of Panderichthys and the origin of tetrapod locomotion

One of the most marked transformations in the vertebrate transition to land was that of fins to limbs. This transformation involved not only the generation of morphological novelties (digits, sacrum) but also a shift in locomotory dominance from the pectoral to the pelvic appendage. Despite its importance, the transformation from pelvic fin to hindlimb is the least studied and least well-documented part of this transformation, which is bracketed by the osteolepiform Eusthenopteron and the early tetrapods Ichthyostega and Acanthostega, but is not directly illuminated by any intermediate form. Panderichthys is the closest tetrapod relative currently represented by complete fossils, but its pelvic fin skeleton has not been described. Here, I present the only known articulated pelvic fin endoskeleton and associated partial pelvis of Panderichthys. The pelvic girdle is even less tetrapod-like than that of the osteolepiform Eusthenopteron, but the pelvic fin endoskeleton shares derived characteristics with basal tetrapods despite being more primitive than the pectoral fin of Panderichthys. The evolution of tetrapod locomotion appears to have passed through a stage of body-flexion propulsion, in which the pelvic fins played a relatively minor anchoring part, before the emergence of hindlimb-powered propulsion in the interval between Panderichthys and Acanthostega.     

The pectoral fin of Tiktaalik roseae and the origin of the tetrapod limb

Wrists, ankles and digits distinguish tetrapod limbs from fins, but direct evidence on the origin of these features has been unavailable. Here we describe the pectoral appendage of a member of the sister group of tetrapods, Tiktaalik roseae, which is morphologically and functionally transitional between a fin and a limb. The expanded array of distal endochondral bones and synovial joints in the fin of Tiktaalik is similar to the distal limb pattern of basal tetrapods. The fin of Tiktaalik was capable of a range of postures, including a limb-like substrate-supported stance in which the shoulder and elbow were flexed and the distal skeleton extended. The origin of limbs probably involved the elaboration and proliferation of features already present in the fins of fish such as Tiktaalik.     

An exceptional Devonian fish from Australia sheds light on tetrapod origins

The transition from fishes to tetrapods was one of the most dramatic events in the evolution of vertebrates, but many pivotal fossils are incomplete, resulting in gaps in the data that are used for phylogenetic reconstruction. Here we present new observations from the most complete, acid-prepared Devonian tetrapodomorph fish yet discovered, Gogonasus, which was previously placed just crownward of Kenichthys and rhizodontids, the most primitive taxa on the tetrapod lineage. Unexpectedly, Gogonasus shows a mosaic of plesiomorphic and derived tetrapod-like features. Whereas the braincase and dermal cranial skeleton exhibit generalized morphologies with respect to Eusthenopteron or Panderichthys, taxa that are traditionally considered to be phyletically close to tetrapods, the presence of a deeply invaginated, wide spiracle, advanced internal spiracular architecture and near-horizontal hyomandibula are specialized features that are absent from Eusthenopteron. Furthermore, the pectoral fin skeleton of Gogonasus shares several features with that of Tiktaalik, the most tetrapod-like fish. A new phylogenetic analysis places Gogonasus crownward of Eusthenopteron as the sister taxon to the Elpistostegalia. Aspects of the basic tetrapod limb skeleton and middle ear architecture can now be traced further back within the tetrapodomorph radiation.     

An autopodial-like pattern of Hox expression in the fins of a basal actinopterygian fish

Comparative analyses of Hox gene expression and regulation in teleost fish and tetrapods support the long-entrenched notion that the distal region of tetrapod limbs, containing the wrist, ankle and digits, is an evolutionary novelty. Data from fossils support the notion that the unique features of tetrapod limbs were assembled over evolutionary time in the paired fins of fish. The challenge in linking developmental and palaeontological approaches has been that developmental data for fins and limbs compare only highly derived teleosts and tetrapods; what is lacking are data from extant taxa that retain greater portions of the fin skeletal morphology considered primitive to all bony fish. Here, we report on the expression and function of genes implicated in the origin of the autopod in a basal actinopterygian, Polyodon spathula. Polyodon exhibits a late-phase, inverted collinear expression of 5' HoxD genes, a pattern of expression long considered a developmental hallmark of the autopod and shown in tetrapods to be controlled by a 'digit enhancer' region. These data show that aspects of the development of the autopod are primitive to tetrapods and that the origin of digits entailed the redeployment of ancient patterns of gene activity.     

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

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.     

Tetrapod-like middle ear architecture in a Devonian fish

Few fossils show the incipient stages of complex morphological transformations. For example, the earliest stages in the remodelling of the spiracular tract and suspensorium (jaw suspension) of osteolepiform fishes into the middle ear of tetrapods have remained elusive. The most primitive known tetrapods show a middle ear architecture that is very different from osteolepiforms such as Eusthenopteron, with little indication of how this transformation took place. Here we present an analysis of tetrapod middle ear origins that is based on a detailed study of Panderichthys, the immediate sister taxon of tetrapods. We show that the spiracular region is radically transformed from osteolepiforms and represents the earliest stages in the origin of the tetrapod middle ear architecture. The posterior palatoquadrate of Panderichthys is completely tetrapod-like and defines a similarly tetrapod-like spiracular tract. The hyomandibula has lost its distal portion, representing a previously unrecognized advance towards a stapes-like morphology. This spiracular specialization suggests that the middle ear of early tetrapods evolved initially as part of a spiracular breathing apparatus.     

Evolutionary origins of vertebrate appendicular muscle

The evolution of terrestrial tetrapod species heralded a transition in locomotor strategies. While most fish species use the undulating contractions of the axial musculature to generate propulsive force, tetrapods also rely on the appendicular muscles of the limbs to generate movement. Despite the fossil record generating an understanding of the way in which the appendicular skeleton has evolved to provide the scaffold for tetrapod limb musculature, there is, by contrast, almost no information as to how this musculature arose. Here we examine fin muscle formation within two extant classes of fish. We find that in the teleost, zebrafish, fin muscles arise from migratory mesenchymal precursor cells that possess molecular and morphogenetic identity with the limb muscle precursors of tetrapod species. Chondrichthyan dogfish embryos, however, use the primitive mechanism of direct epithelial somitic extensions to derive the muscles of the fin. We conclude that the genetic mechanism controlling formation of tetrapod limb muscles evolved before the Sarcopterygian radiation.     

Ventastega curonica and the origin of tetrapod morphology

The gap in our understanding of the evolutionary transition from fish to tetrapod is beginning to close thanks to the discovery of new intermediate forms such as Tiktaalik roseae. Here we narrow it further by presenting the skull, exceptionally preserved braincase, shoulder girdle and partial pelvis of Ventastega curonica from the Late Devonian of Latvia, a transitional intermediate form between the 'elpistostegids' Panderichthys and Tiktaalik and the Devonian tetrapods (limbed vertebrates) Acanthostega and Ichthyostega. Ventastega is the most primitive Devonian tetrapod represented by extensive remains, and casts light on a part of the phylogeny otherwise only represented by fragmentary taxa: it illuminates the origin of principal tetrapod structures and the extent of morphological diversity among the transitional forms.     

An early tetrapod from 'Romer's Gap'

The fossil record of early tetrapods has been increased recently by new finds from the Devonian period and mid-late Early Carboniferous period. Despite this, understanding of tetrapod evolution has been hampered by a 20-million-year gap ('Romer's Gap') that covers the crucial, early period when many key features of terrestrial tetrapods were acquired. Here I describe the only articulated skeleton of a tetrapod, Pederpes, yet found from the Tournaisian epoch (354-344 million years ago (Myr)). The new taxon includes a pes with five robust digits, but a very small, possibly supernumerary digit preserved on the manus suggests the presence of polydactyly. Polydactylous early tetrapods may have survived beyond the end of the Devonian and pentadactyly cannot be assumed for the pes. However, the pes has characteristics that distinguish it from the paddle-like feet of the Devonian forms and resembles the feet of later, more terrestrially adapted Carboniferous forms. Pederpes is the earliest-known tetrapod to show the beginnings of terrestrial locomotion and was at least functionally pentadactyl. With its later American sister-genus, Whatcheeria, it represents the next most primitive tetrapod clade after those of the Late Devonian, bridging the temporal, morphological and phylogenetic gaps that have hitherto separated Late Devonian and mid-Carboniferous tetrapod faunas.     

A primitive fish close to the common ancestor of tetrapods and lungfish

The relationship of the three living groups of sarcopterygians or lobe-finned fish (tetrapods, lungfish and coelacanths) has been a matter of debate. Although opinions still differ, most recent phylogenies suggest that tetrapods are more closely related to lungfish than to coelacanths. However, no previously known fossil taxon exhibits a concrete character combination approximating the condition expected in the last common ancestor of tetrapods and lungfish -- and it is still poorly understood how early sarcopterygians diverged into the tetrapod lineage (Tetrapodomorpha) and the lungfish lineage (Dipnomorpha). Here we describe a fossil sarcopterygian fish, Styloichthys changae gen. et sp. nov., that possesses an eyestalk and which exhibits the character combination expected in a stem group close to the last common ancestor of tetrapods and lungfish. Styloichthys from the Lower Devonian of China bridges the morphological gap between stem-group sarcopterygians (Psarolepis and Achoania) and basal tetrapodomorphs/basal dipnomorphs. It provides information that will help in the study of the relationship of early sarcopterygians, and which will also help to resolve the tetrapod-lungfish divergence into a documented sequence of character acquisition.     

Sonic hedgehog function in chondrichthyan fins and the evolution of appendage patterning

The genetic mechanisms regulating tetrapod limb development are well characterized, but how they were assembled during evolution and their function in basal vertebrates is poorly understood. Initial studies report that chondrichthyans, the most primitive extant vertebrates with paired appendages, differ from ray-finned fish and tetrapods in having Sonic hedgehog (Shh)-independent patterning of the appendage skeleton. Here we demonstrate that chondrichthyans share patterns of appendage Shh expression, Shh appendage-specific regulatory DNA, and Shh function with ray-finned fish and tetrapods. These studies demonstrate that some aspects of Shh function are deeply conserved in vertebrate phylogeny, but also highlight how the evolution of Shh regulation may underlie major morphological changes during appendage evolution.     

The cranial endoskeleton of Tiktaalik roseae

Among the morphological changes that occurred during the 'fish-to-tetrapod' transition was a marked reorganization of the cranial endoskeleton. Details of this transition, including the sequence of character acquisition, have not been evident from the fossil record. Here we describe the braincase, palatoquadrate and branchial skeleton of Tiktaalik roseae, the Late Devonian sarcopterygian fish most closely related to tetrapods. Although retaining a primitive configuration in many respects, the cranial endoskeleton of T. roseae shares derived features with tetrapods such as a large basal articulation and a flat, horizontally oriented entopterygoid. Other features in T. roseae, like the short, straight hyomandibula, show morphology intermediate between the condition observed in more primitive fish and that observed in tetrapods. The combination of characters in T. roseae helps to resolve the relative timing of modifications in the cranial endoskeleton. The sequence of modifications suggests changes in head mobility and intracranial kinesis that have ramifications for the origin of vertebrate terrestriality.     

A primitive sarcopterygian fish with an eyestalk

The discovery of two Early Devonian osteichthyan (bony fish) fossils has challenged established ideas about the origin of osteichthyans and their divergence into actinopterygians (teleosts and their relatives) and sarcopterygians (tetrapods, coelacanths, lungfishes and related groups). Psarolepis from China and an unnamed braincase from Australia combine derived sarcopterygian and actinopterygian characters with primitive features previously restricted to non-osteichthyans, suggesting that early osteichthyan evolution may have involved substantial parallellism between sarcopterygians and actinopterygians. But interpretation of these fossils has been hampered by poor phylogenetic resolution. Here we describe a basal sarcopterygian fish, Achoania gen, et sp. nov., that fills the morphological gap between Psarolepis and higher sarcoptergyians. We also report the presence of eyestalk attachments in both Achoania and Psarolepis, showing that this supposedly non-osteichthyan feature occurs in basal sarcopterygians as well as the actinoptergyian-like Australian braincase.     

Palaeogeography: Devonian tetrapod from western Europe

Several discoveries of Late Devonian tetrapods (limbed vertebrates) have been made during the past two decades, but each has been confined to one locality. Here we describe a tetrapod jaw of about 365 million years (Myr) old from the Famennian of Belgium, which is the first from western continental Europe. The jaw closely resembles that of Ichthyostega, a Famennian tetrapod hitherto known only from Greenland. The environment of this fossil provides information about the conditions that prevailed just before the virtual disappearance of tetrapods from the fossil record for 20 Myr.     

Mechanical design and implementation of a new biomimetic robot fish

A mechanical design method of robot fish is introduced in this paper.Based on this method an antonomous 3-Dimension(3D)locomotion robot fish with two pectoral fins and a caudal fin is developed.The pectoral fin mechanism has 3 degrees of freedom(3-DOFs),which enables the robot fish to realize yawing and pitching motions by controlling two pectoral fins.And the caudal fin mechanism is designed based on fish body wave curve fitting.The forward velocity can be adjusted by changing the caudal mechanism's oscillating frequency.Finally a physical implementation of the robot fish and experimental results are given.     

Three-dimensional limb joint mobility in the early tetrapod Ichthyostega

The origin of tetrapods and the transition from swimming to walking was a pivotal step in the evolution and diversification of terrestrial vertebrates. During this time, modifications of the limbs—particularly the specialization of joints and the structures that guide their motions—fundamentally changed the ways in which early tetrapods could move. Nonetheless, little is known about the functional consequences of limb anatomy in early tetrapods and how that anatomy influenced locomotion capabilities at this very critical stage in vertebrate evolution. Here we present a three-dimensional reconstruction of the iconic Devonian tetrapod Ichthyostega and a quantitative and comparative analysis of limb mobility in this early tetrapod. We show that Ichthyostega could not have employed typical tetrapod locomotory behaviours, such as lateral sequence walking. In particular, it lacked the necessary rotary motions in its limbs to push the body off the ground and move the limbs in an alternating sequence. Given that long-axis rotation was present in the fins of tetrapodomorph fishes, it seems that either early tetrapods evolved through an initial stage of restricted shoulder and hip joint mobility or that Ichthyostega was unique in this respect. We conclude that early tetrapods with the skeletal morphology and limb mobility of Ichthyostega were unlikely to have made some of the recently described Middle Devonian trackways.     

仿生机器鱼胸鳍的三维设计及数控加工

胸鳍是机器鱼实现沉浮、变向、倒退和推进的关键部件,胸鳍的三维设计及型面质量直接影响机器鱼实现稳定的各种运动.本文运用Soildworks2007对仿生机器鱼的胸鳍进行三维设计,通过CAM功能生成了胸鳍数控加工的代码,并用数控机床实现了机器鱼胸鳍的数控加工.     

Discovery of the earliest-known tetrapod stapes

The evolution of the middle ear is central to the discussion of how the first tetrapods adapted to life on land as well as their phylogeny. Here I report the discovery of the stapes of Acanthostega gunnari, from the Upper Devonian of east Greenland. This is the earliest tetrapod stapes so far described, and it throws new light on both these aspects of early tetrapod biology. It has been assumed that the common inheritance of all early tetrapods was a light, rod-like stapes associated with a temporal notch in the otic region that was thought to have supported a tympanum, or eardrum. The stapes would have conducted vibrations from the tympanum to the otic capsule. By contrast, the stapes of Acanthostega was stout with a broad distal ramus associated with the temporal notch. I suggest that the temporal notch of Acanthostega and other early tetrapods supported a spiracular opening rather than a tympanum, and that the stapes controlled palatal and spiracular movements in ventilation.     

贵州省洞穴高原鳅属一新种

描记采于贵州省盘县保田镇一地下洞穴高原鳅属一新种,命名保田高原鳅Triplophysa baotianensis sp. nov..新种是贵州省南盘江水系的首个高原鳅发现.与贵州已记载的5种高原鳅比较,新种与北盘江水系的贞丰高原鳅性状相近,但有明显特征可区别.（1）新种胸腹鳍平展与躯体呈“十”字形,贞丰高原鳅胸腹鳍走向正常;（2）新种背鳍起点在腹鳍起点之前上方,贞丰高原鳅在腹鳍起点之后上方;（3）新种体表裸露无鳞,贞丰高原鳅体被明显细鳞;（4）新种鼻瓣发育成短须状,贞丰高原鳅鼻瓣不呈须状;（5）新种属于南盘江水系,贞丰高原鳅属于北盘江水系.