A primitive fish provides key characters bearing on deep osteichthyan phylogeny

Osteichthyans, or bony vertebrates, include actinopterygians (teleosts and their relatives) and sarcopterygians (coelacanths, lungfishes and tetrapods). Despite features found in basal actinopterygians (for example, Dialipina and Ligulalepis) and basal sarcopterygians (for example, Psarolepis and Achoania), the morphological gap between the two lineages remains wide and how sarcopterygians developed a dermal surface covering known as cosmine (composed of a pore-canal network and a single layer of odontodes and enamel) is still poorly known. Here we describe a primitive fossil fish, Meemannia eos gen. et sp. nov., that possesses an actinopterygian-like skull roof and a cosmine-like dermal surface combining a pore-canal network (found in various fossil sarcopterygians) with superimposed layers of odontodes and enamel (previously known in actinopterygians and some acanthodians). This 405-million-year-old fish from the Lower Devonian of Yunnan (China) demonstrates that cosmine in many fossil sarcopterygians arose step by step through the acquisition of a pore-canal network followed by the subsequently developed ability to resorb previous generations of odontodes and enamel. Meemannia provides key characters for studying deep osteichthyan phylogeny and indicates a possible morphotype for the common ancestor of actinopterygians and sarcopterygians.     

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.     

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.     

The pectoral fin of Panderichthys and the origin of digits

One of the identifying characteristics of tetrapods (limbed vertebrates) is the presence of fingers and toes. Whereas the proximal part of the tetrapod limb skeleton can easily be homologized with the paired fin skeletons of sarcopterygian (lobe-finned) fish, there has been much debate about the origin of digits. Early hypotheses interpreted digits as derivatives of fin radials, but during the 1990s the idea gained acceptance that digits are evolutionary novelties without direct equivalents in fish fin skeletons. This was partly based on developmental genetic data, but also substantially on the pectoral fin skeleton of the elpistostegid (transitional fish/tetrapod) Panderichthys, which appeared to lack distal digit-like radials. Here we present a CT scan study of an undisturbed pectoral fin of Panderichthys demonstrating that the plate-like 'ulnare' of previous reconstructions is an artefact and that distal radials are in fact present. This distal portion is more tetrapod-like than that found in Tiktaalik and, in combination with new data about fin development in basal actinopterygians, sharks and lungfish, makes a strong case for fingers not being a novelty of tetrapods but derived from pre-existing distal radials present in all sarcopterygian fish.     

The most primitive osteichthyan braincase?

Most living vertebrates, from teleosts to tetrapods, are osteichthyans (bony fishes), but the origin of this major group is poorly understood. The actinopterygians (ray-finned bony fishes) are the most successful living vertebrates in terms of diversity. They appear in the fossil record in the Late Silurian but are poorly known before the Late Devonian. Here we report the discovery of the oldest and most primitive actinopterygian-like osteichthyan braincase known, from 400-million-year-old limestone in southeastern Australia. This specimen displays previously unknown primitive conditions, in particular, an opening for a cartilaginous eyestalk. It provides an important and unique counterpart to the similarly aged and recently described Psarolepis from China and Vietnam. The contrasting features of these specimens, and the unusual anatomy of the new specimen in particular, provide new insights into anatomical conditions close to the evolutionary radiation of all modern osteichthyan groups.     

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.     

Cloning the interferon regulatory factor I gene in lungfish （Protopterus annectens） and its molecular evolution among sarcopterygians

Sarcopterygians is an important vertebrate clade that includes crossopterygians and tetrapods. Crossopterygians are lobe-finned fish that include lungfish and coelacanths. Tetrapods include amphibians, reptiles, avians and mammals. To compare the interferon regulatory factor 1 (irf-1) gene structure and to explore phylogenetic relationships among sarcopterygians, we cloned the cDNA sequence of irf-1 from lungfish and compared it with irf-1 orthologs in other sarcopterygian species. The lungfish is a primitive sarcopterygian that occupies a very important position in vertebrate phylogeny. Interferon regulatory factors (IRFs) are a family of proteins involved in innate immunity. To date, 11 IRF family members have been reported. All IRFs share homology in the first 115 amino acids, which encompasses a DNA binding domain containing a characteristic repeat of 5 tryptophan residues separated by 10–18 amino acids. IRF-1 and IRF-2 were the first members of this family to be reported and they have a very important role in innate immunity. However, studies of the irf-1 and irf-2 genes are mostly confined to mammals; very few non-mammalian irf-1 genes have been reported. Consistent with the irf-1 gene sequences already published, the first 345 nucleotides of lungfish irf-1 are highly conserved. At the carboxyl terminal a C-terminal transactivating region motif and an interferon associated domain (IAD2) were identified. 417 million years separate the present from the closest common ancestor of lungfish and tetrapods; however, the irf-1 genes among sarcopterygians are highly conserved and have very obvious phylogenetic relationships. Also the interrelationship tree of sarcopterygians, based on IRF-1 amino acid sequences, is identical with trees produced using other data, such as morphological characteristics or mitochondrial gene sequences.     

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.     

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 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.     

The avian nature of the brain and inner ear of Archaeopteryx

Archaeopteryx, the earliest known flying bird (avialan) from the Late Jurassic period, exhibits many shared primitive characters with more basal coelurosaurian dinosaurs (the clade including all theropods more bird-like than Allosaurus), such as teeth, a long bony tail and pinnate feathers. However, Archaeopteryx possessed asymmetrical flight feathers on its wings and tail, together with a wing feather arrangement shared with modern birds. This suggests some degree of powered flight capability but, until now, little was understood about the extent to which its brain and special senses were adapted for flight. We investigated this problem by computed tomography scanning and three-dimensional reconstruction of the braincase of the London specimen of Archaeopteryx. Here we show the reconstruction of the braincase from which we derived endocasts of the brain and inner ear. These suggest that Archaeopteryx closely resembled modern birds in the dominance of the sense of vision and in the possession of expanded auditory and spatial sensory perception in the ear. We conclude that Archaeopteryx had acquired the derived neurological and structural adaptations necessary for flight. An enlarged forebrain suggests that it had also developed enhanced somatosensory integration with these special senses demanded by a lifestyle involving flying ability.     

An Early Cambrian tunicate from China

Like the Burgess Shales of Canada, the Chengjiang Lagerst?tte from the Lower Cambrian of China is renowned for the detailed preservation as fossils of delicate, soft-bodied creatures, providing an insight into the Cambrian explosion. The fossils of possible hemichordate chordates and vertebrates have attracted particular attention. Tunicates, or urochordates, comprise the most basal chordate clade, and details of their evolution could be important in understanding the sequence of character acquisition that led to the emergence of chordates and vertebrates. However, definitive fossils of tunicates from the Cambrian are scarce or debatable. Here we report a probable tunicate Cheungkongella ancestralis from the Chengjiang fauna. It resembles the extant ascidian tunicate genus Styela whose morphology could be useful in understanding the origin of the vertebrates.     

Microtubular metazoan fossils with multi-branches in Weng’an Biota

Microtubular metazoan fossils with multibranches are found at least 4 times from the Sinian （Ediacarian） Doushantuo Formation at Weng＇an, Guizhou, China. Their living mode should be the benthonic colony with higher ability of asexual reproduction. The inner of the tube with cross-walls and chamber shows that these fossils are basal or stemgroup cnidarians, similar to extinct tabulates. The multi-branching microtubular fossils are real metazoan found from Weng＇an biota, which is very important for studying the origin and evolvement of metazoan.     

A deep-bodied ginglymodian fish from the Middle Triassic of eastern Yunnan Province, China, and the phylogeny of lower neopterygians

The Ginglymodi are a group of ray-finned fishes that make up one of three major subdivisions of the infraclass Neopterygii.Ex- tant ginglymodians are represented by gars,which inhabit freshwater environments of North and Central America and Cuba.Here, we report the discovery of well-preserved fossils of a new ginglymodian,Kyphosichthys grandei gen.et sp.nov.,from the Middle Triassic(Anisian)marine deposits(Guanling Formation)in Luoping,eastern Yunnan Province,China.The discovery documents the first known fossil record of highly deep-bodied ginglymodians,adding new information on the early morphological diversity of this group.The studies of functional morphology of extant deep-bodied fishes indicate that Kyphosichthys is not a fast swim- mer but has a good performance in precise maneuvering,representing a morphological adaptation to structurally complex habitats (e.g.thick macrophyte beds,rocky areas,or coral reefs),which differs from the other members of this group.A cladistic analysis with the new fish taxon included supports the hypothesis that the Ginglymodi are more closely related to the Halecomorphi than to the Teleostei.Represented by Felberia,Kyphosichthys,and Dapedium,a highly deep and short fish body type has inde- pendently evolved at least three times in the stem-group neopterygians,ginglymodians,and basal teleosts within the lower neop- terygians of the Triassic.     

Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary

Estimates of the time of origin for placental mammals from DNA studies span nearly the duration of the Cretaceous period (145 to 65 million years ago), with a maximum of 129 million years ago and a minimum of 78 million years ago. Palaeontologists too are divided on the timing. Some support a deep Cretaceous origin by allying certain middle Cretaceous fossils (97-90 million years old) from Uzbekistan with modern placental lineages, whereas others support the origin of crown group Placentalia near the close of the Cretaceous. This controversy has yet to be addressed by a comprehensive phylogenetic analysis that includes all well-known Cretaceous fossils and a wide sample of morphology among Tertiary and recent placentals. Here we report the discovery of a new well-preserved mammal from the Late Cretaceous of Mongolia and a broad-scale phylogenetic analysis. Our results exclude Cretaceous fossils from Placentalia, place the origin of Placentalia near the Cretaceous/Tertiary (K/T) boundary in Laurasia rather than much earlier within the Cretaceous in the Southern Hemisphere, and place afrotherians and xenarthrans in a nested rather than a basal position within Placentalia.     

First record of Lower Triassic <Emphasis Type="Italic">Undichna</Emphasis> spp. fish swimming traces from Emei,Sichuan Province,China

We discuss traces from the ichnospecies Undichna unisulca recorded from the Triassic that are reported for the first time in this paper. These fossils are also some of the oldest vertebrate ichnites discovered in China to date. The fish swimming traces (Undichna unisulca) described originate from the Lower Triassic. These traces were found in the Jialingjiang Formation and are located at a gorge ~7.5 km from Emei, Sichuan Province, China. The ichnospecies Undichna unisulca consists of a regular sinusoidal wave. The wavelengths and amplitudes are quite constant between each specimen ranging between 28-32 mm and 9-12 mm, respectively. The fish swimming traces and distribution within the same bed are preserved as hypichnial ridges at the undersurface.     

Transitional mammalian middle ear from a new Cretaceous Jehol eutriconodont

The transference of post-dentary jaw elements to the cranium of mammals as auditory ossicles is one of the central topics in evolutionary biology of vertebrates. Homologies of these bones among jawed vertebrates have long been demonstrated by developmental studies; but fossils illuminating this critical transference are sparse and often ambiguous. Here we report the first unambiguous ectotympanic (angular), malleus (articular and prearticular) and incus (quadrate) of an Early Cretaceous eutriconodont mammal from the Jehol Biota, Liaoning, China. The ectotympanic and malleus have lost their direct contact with the dentary bone but still connect the ossified Meckel's cartilage (OMC); we hypothesize that the OMC serves as a stabilizing mechanism bridging the dentary and the detached ossicles during mammalian evolution. This transitional mammalian middle ear narrows the morphological gap between the mandibular middle ear in basal mammaliaforms and the definitive mammalian middle ear (DMME) of extant mammals; it reveals complex changes contributing to the detachment of ear ossicles during mammalian evolution.     

A uniquely specialized ear in a very early tetrapod

The Late Devonian genus Ichthyostega was for many decades the earliest known tetrapod, and the sole representative of a transitional form between a fish and a land vertebrate. However, despite being known since 1932 (ref. 1) from a large collection of specimens, its morphology remained enigmatic and not what was expected of a very primitive tetrapod. Its apparent specializations led it to be considered as a "blind offshoot" or "sidebranch" off the tetrapod family tree, and recent cladistic analyses have disagreed about its exact phylogenetic position within the tetrapod stem group. In particular, its braincase and ear region defied interpretation, such that conventional anatomical terms seemed inapplicable. Using new material collected in 1998 (ref. 9), preparation of earlier-collected material, and high-resolution computed tomography scanning, here we identify and interpret these problematic anatomical structures. They can now be seen to form part of a highly specialized ear, probably a hearing device for use in water. This represents a structurally and functionally unique modification of the tetrapod otic region, unlike anything seen in subsequent tetrapod evolution. The presence of deeply grooved gill bars as in its contemporary Acanthostega suggest that Ichthyostega may have been more aquatically adapted than previously believed.     

Eocene lizard from Germany reveals amphisbaenian origins

Amphisbaenia is a speciose clade of fossorial lizards characterized by a snake-like body and a strongly reinforced skull adapted for head-first burrowing. The evolutionary origins of amphisbaenians are controversial, with molecular data uniting them with lacertids, a clade of Old World terrestrial lizards, whereas morphology supports a grouping with snakes and other limbless squamates. Reports of fossil stem amphisbaenians have been falsified, and no fossils have previously tested these competing phylogenetic hypotheses or shed light on ancestral amphisbaenian ecology. Here we report the discovery of a new lacertid-like lizard from the Eocene Messel locality of Germany that provides the first morphological evidence for lacertid-amphisbaenian monophyly on the basis of a reinforced, akinetic skull roof and braincase, supporting the view that body elongation and limblessness in amphisbaenians and snakes evolved independently. Morphometric analysis of body shape and ecology in squamates indicates that the postcranial anatomy of the new taxon is most consistent with opportunistically burrowing habits, which in combination with cranial reinforcement indicates that head-first burrowing evolved before body elongation and may have been a crucial first step in the evolution of amphisbaenian fossoriality.