CT scan reconstruction of the palate region of Latimeria chalumnae

Synopsis The palate of Latimeria chalumnae is described based mainly on three-dimensional CT scan reconstruction. It is compared with that of other osteichthyans. The palate of L. chalumnae compares best with that of rhipidistians; it is more advanced than that of actinopterygians in having fewer bones. This tendency toward bone reduction in the palate is even more pronounced in dipnoans. The interpretation of features of the Early Devonian genus Diabolepis determines if authors consider dipnoans or actinistians more closely related to tetrapods. Both groups are only distant relatives of tetrapods.     

A near-tetrapod from the Baltic Middle Devonian

The tetrapodomorph sarcopterygian Livoniana multidentata gen. et sp. nov. is described on the basis of lower jaw fragments from the Middle Devonian(late Givetian) of Latvia and Estonia. It possesses a suite of derived characters previously only known from tetrapods, which first appear in the late Devonian (late Frasnian), and a phylogenetic analysis places it on the internode between Panderichthys and the base of the Tetrapoda. The analysis also reveals that the 'Elpistostegalia' are paraphyletic to Tetrapoda, with Elpistostege closer to tetrapods than is Panderichthys. Owing to incompleteness of the material, there is almost no overlap between the data sets for Elpistostege Livoniana ; the analysis places the two genera in an unresolved trichotomy. In addition to the tetrapod features, Livoniana has a strikingly autapomorphic dentary dentition comprising multiple tooth rows. It thus provides evidence both for the unexpectedly early evolution of tetrapod characteristics and for morphological radiation around the fish-tetrapod transition.     

The First Virtual Cranial Endocast of a Lungfish (Sarcopterygii: Dipnoi)

Lungfish, or dipnoans, have a history spanning over 400 million years and are the closest living sister taxon to the tetrapods. Most Devonian lungfish had heavily ossified endoskeletons, whereas most Mesozoic and Cenozoic lungfish had largely cartilaginous endoskeletons and are usually known only from isolated tooth plates or disarticulated bone fragments. There is thus a substantial temporal and evolutionary gap in our understanding of lungfish endoskeletal morphology, between the diverse and highly variable Devonian forms on the one hand and the three extant genera on the other. Here we present a virtual cranial endocast of Rhinodipterus kimberleyensis, from the Late Devonian Gogo Formation of Australia, one of the most derived fossil dipnoans with a well-ossified braincase. This endocast, generated from a Computed Microtomography (µCT) scan of the skull, is the first virtual endocast of any lungfish published, and only the third fossil dipnoan endocast to be illustrated in its entirety. Key features include long olfactory canals, a telencephalic cavity with a moderate degree of ventral expansion, large suparaotic cavities, and moderately enlarged utricular recesses. It has numerous similarities to the endocasts of Chirodipterus wildungensis and Griphognathus whitei, and to a lesser degree to ''Chirodipterus'' australis and Dipnorhynchus sussmilchi. Among extant lungfish, it consistently resembles Neoceratodus more closely than Lepidosiren and Protopterus. Several trends in the evolution of the brains and labyrinth regions in dipnoans, such as the expansions of the utricular recess and telencephalic regions over time, are identified and discussed.     

A NEW DIPNOAN FISH FROM THE MIDDLE DEVONIAN (EIFELIAN) OF SCOTLAND

Abstract:  A new dipnoan fish, Pinnalongus saxoni gen. et sp. nov., is described from a suite of recently discovered specimens from Caithness and East Sutherland, northern Scotland. The stratum where Pinnalongus occurs is within the middle Eifelian of the Middle Devonian. Pinnalongus has a skull roof consisting of a mosaic of small bones at the anterior end, a feature more typically found in Early Devonian dipnoans. The posterior part of the skull roof is more typical of later advanced Middle Devonian dipnoans with the B-bone separating the I-bones. The postcranial body is now considered not to be so important with respect to the classification of dipnoans as originally proposed by Dollo in 1895, who thought there was an evolutionary transformation series. Nevertheless, the postcranial body of Pinnalongus has more in common with later Middle and Late Devonian dipnoans such as Scaumenacia , with a very long second dorsal fin. Based on the skull morphology, Pinnalongus is most closely related to Tarachomylax from the Lower Devonian of Severnaya Zemlya, Russia. Pinnalongus represents the earliest known complete articulated dipnoan with tooth plates.     

Cytogenetic and molecular studies in a lungfish,Protopterus annectens (Osteichthyes,Dipnoi)

A neontological approach to the problem of the origin of tetrapods consists in the examination of the available cytological and molecular data on the genome of these vertebrates. Dipnoans are a group of osteichthyian fishes, the evolutionary relationships of which with tetrapods have been disputed since their discovery. In the past, they were variously considered as being related to actinistians, tetrapods, and lower actinopterygians, though nowadays they are considered a monophyletic group, the sister group of crossopterygians. Dipnoans first appeared in the geologic record in the Early Devonian with 50 extinct genera, surviving up to date, with only three genera: Lepidosiren, Neoceratodus and Protopterus, including only six recognized species. Nothing is known of the genome of the early tetrapods, except that they and the Choanoichthyes exhibited a remarkable interspecific variability of the karyotype and of DNA content. These characteristics are often found in dipnoans and in the extant lissamphibians. Very little is known about the evolutionary karyology in the four Protopterus species and in the dipnoan clade in general. In this paper, we karyotyped ten male and female specimens of P. annectens (2n=34) from Nigeria. Moreover, we localized heterochromatin and nucleolar organizer regions by using base-specific fluorochromes and detected the human telomeric (TTAGGG)(n) sequences on all the telomeric sites of P. annectens chromosomes. DNA was also extracted and digested with seven restriction enzymes, which revealed the probable presence of almost three different families of satellite DNA. Nuclear DNA content was identified from blood samples by flow cytometry. New genomic and karyological data were compared and discussed with those on closer genera and taxa available in literature.     

Oldest coelacanth, from the Early Devonian of Australia

Coelacanths are well-known sarcopterygian (lobe-finned) fishes, which together with lungfishes are the closest extant relatives of land vertebrates (tetrapods). Coelacanths have both living representatives and a rich fossil record, but lack fossils older than the late Middle Devonian (385-390 Myr ago), conflicting with current phylogenies implying coelacanths diverged from other sarcopterygians in the earliest Devonian (410-415 Myr ago). Here, we report the discovery of a new coelacanth from the Early Devonian of Australia (407-409 Myr ago), which fills in the approximately 20 Myr 'ghost range' between previous coelacanth records and the predicted origin of the group. This taxon is based on a single lower jaw bone, the dentary, which is deep and short in form and possesses a dentary sensory pore, otherwise seen in Carboniferous and younger taxa.     

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.     

New finds of vertebrates in the Middle Devonian Brandenberg Group (Sauerland, Northwest Germany)

Placoderms (Ptyctodontida), acanthodians (incl.Atopacanthus?ambrockensis n. sp.), actinopterygians, dipnoans and crossopterygians — including the first Middle Devonian osteolepids from the Rheinisches Schiefergebirge — are described from the lower Brandenberg Group of Hagen-Ambrock and Lasbeck (northern Sauerland, Northwest Germany). The composition of the fauna is similar to that of contemporaneous Scottish and Baltic Old Red localities, even though typical marine vertebrates are present too (e.g. selachians). This indicates — like the invertebrates — rather a marginal marine deposition.     

New discoveries of tetrapods (ichthyostegid‐like and whatcheeriid‐like) in the Famennian (Late Devonian) localities of Strud and Becco (Belgium)

The origin of tetrapods is one of the key events in vertebrate history. The oldest tetrapod body fossils are Late Devonian (Frasnian–Famennian) in age, most of them consisting of rare isolated bone elements. Here we describe tetrapod remains from two Famennian localities from Belgium: Strud, in the Province of Namur, and Becco, in the Province of Liège. The newly collected material consists of an isolated complete postorbital, fragments of two maxillae, and one putative partial cleithrum, all from Strud, and an almost complete maxilla from Becco. The two incomplete maxillae and cleithrum from Strud, together with the lower jaw previously recorded from this site, closely resemble the genus Ichthyostega, initially described from East Greenland. The postorbital from Strud and the maxilla from Becco do not resemble the genus Ichthyostega. They show several derived anatomical characters allowing their tentative assignment to a whatcheeriid‐grade group. The new tetrapod records show that there are at least two tetrapod taxa in Belgium and almost certainly two different tetrapod taxa at Strud. This locality joins the group of Devonian tetrapod‐bearing localities yielding more than one tetrapod taxon, confirming that environments favourable to early tetrapod life were often colonized by several tetrapod taxa.     

The inferognathal in the Middle Devonian arthrodire Homostius

Mark-Kurik, E. 1992 04 15: The inferognathal in the Middle Devonian arthrodire Homostius. Lethaia , Vol. 25, pp. 173–178. Oslo. ISSN 0024–1164.
The inferognathal or lower jaw bone of the gigantic Middle Devonian arthrodire Homostius (family Homostiidae) is described in detail. The bone is distinctive in shape, being convex posteriorly but concave anteriorly. Inferognathals of the same sinuous shape are rare in arthrodires. They are known also in Angarichrhys and Brachyosteus . An explanation is given of the nature of the supposed shearing surface (described first by Heintz in 1934) at the anterior end of the bone. It is concluded that Homostius had neither teeth nor shearing surfaces. * Middle Devonian, Estonia, Arthrodira, Homostiidae, lower jaw bone, restoration, structure .     

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.     

The postcranial skeleton of the Devonian tetrapod Tulerpeton curtum Lebedev

Postcranial remains of the Russian Late Devonian tetrapod Tulerpeton include the hexadactylous fore limb, hind limb, anocleithral pectoral girdle, squamation, and associated disarticulated postcranial bones. A cladistic analysis indicates that Tulerpeton is a reptiliomorph stem-group amniote and the earliest known crown-group tetrapod: Acanthostega and Ichthyostega are successively more derived plesion stem-group tetrapods and do not consititute a monophyletic ichthyostegalian radiation. Previous analyses suggesting a profound split in tetrapod phylogeny are thereby corroborated, and likewise the interpretation of Westlothiana as a stem-group amniote. The divergence of reptiliomorphs from batrachomorphs occurred before the Devonian-Carboniferous boundary. Tulerpeton originates from an entirely aquatic environment with a diverse fish fauna. The morphologies of its limbs and those of Devonian stem-tetrapods suggest that dactyly predates the elaboration of the carpus and tarsus, and that Polydactyly persisted after the evolutionary divergence of the principal lineages of living tetrapods. The apparent absence of a branchial lamina and gill skeleton suggests that Tulerpeton was primarily air-breathing, whereas contemporary stem-group tetrapods and more recent batrachomorphs retained greater emphasis on gill-breathing.     

Postcranial remains of Baphetes and their bearing on the relationships of the Baphetidae (= Loxommatidae)

The first unequivocal postcranial remains of baphetids (= loxommatids) associated with skull roof and lower jaw material are reported from a specimen of Baphetes from the English Upper Carboniferous, Duckmantian. Characters of the mandible, including the incorporation in the symphysis of paired parasymphysial plates, permit the identification of a previously indeterminate jaw from the Langsettian (Westphalian A) of Nova Scotia, as baphetid. The postcranial remains include vertebrae, pectoral and pelvic limb and limb girdle elements that present a combination of unique characters extending the diagnosis of the family Baphetidae, together with plesiomophic and derived characters which suggest that baphetids are primitive stem group tetrapods.     

Estuarine fish and tetrapod evolution: insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent

The Waterloo Farm lagerstätte in South Africa provides a uniquely well‐preserved record of a Latest Devonian estuarine ecosystem. Ecological evidence from it is reviewed, contextualised, and compared with that available from the analogous Swartvlei estuarine lake, with a particular emphasis on their piscean inhabitants. Although the taxonomic affinities of the estuarine species are temporally very different, the overall patterns of utilisation prove to be remarkably congruent, with similar trophic structures. Significantly, both systems show evidence of widespread use of estuaries as fish nurseries by both resident and marine migrant taxa. Holocene estuaries are almost exclusively utilised by actinopterygians which are overwhelmingly dominated by oviparous species. Complex strategies are utilised by estuarine resident species to avoid exposure of eggs to environmental stresses that characterize these systems. By contrast, many of the groups utilising Devonian estuaries were likely live bearers, potentially allowing them to avoid the challenges faced by oviparous taxa. This may have contributed to dominance of these systems by non‐actinoptergians prior to the End Devonian Mass Extinction. The association of early aquatic tetrapods at Waterloo Farm with a fish nursery environment is consistent with findings from North America, Belgium and Russia, and may be implied by the estuarine settings of a number of other Devonian tetrapods. Tetrapods apparently replace their sister group, the elpistostegids, in estuaries with both groups having been postulated to be adaptated to shallow water habitats where they could access small piscean prey. Correlation of tetrapods (and elpistostegids) with fish nursery areas in the Late Devonian lends strong support to this hypothesis, suggesting that adaptations permitting improved access to the abundant juvenile fish within the littoral zone of estuarine lakes and continental water bodies may have been pivotal in the evolution of tetrapods.     

COSMINE AND COSMINE GROWTH

From the point of view of phylogeny, the development of cosmine superficially on the dermal elements of certain early lower vertebrates is considered to be a specialized rather than a primitive condition. An analysis of cosmine formation in the squamation of Devonian dipnoans, based on material from the Plattenkalk of Bergisch Gladbach in the Rhineland, shows that (a) separate cosmine sheets on the individual scales, bounded-off from each other by Westoll lines, conform to the pattern of areal growth as displayed in odontode distribution on the dermal elements of various other lower vertebrates, and (b) each consecutive period of cosmine formation began in certain specific scale-areas and subsequently spread to the remainder of the squamation.     

Relative importance of molecular,neontological, and paleontological data in understanding the biology of the vertebrate invasion of land

Summary Meyer and Wilson's (1990) 12S rRNA phylogeny unites lungfish and tetrapods to the exclusion of the coelacanth. These workers also provide a list of morphological features shared in common between modern lungfish and tetrapods, and they conclude that these traits were probably present in their last common ancestor. However, the exquisite fossil records of the abundant extinct lungfishes and rhipidistians show that at least 13 out of Meyer and Wilson's 14 supposed ancestral traits were not present in the last common ancestor of lungfishes and tetrapods. Using extant taxa to infer ancestral morphologies is fraught with difficulties; just like molecular sequences, ancestral character states of morphological traits may be severely overprinted by subsequent modifications. Modern lungfish are air-breathing nonmarine forms, yet their Devonian forebears were marine fish that did not breathe air. Fossils dating from the time of origin of tetrapods in the Devonian offer the only hope of understanding the morphological innovations that led to tetrapods; morphological analysis of the living fossils, the coelacanth and lungfish, only lends confusion.     

Origin of dental occlusion in tetrapods: signal for terrestrial vertebrate evolution?

Evolutionary changes of the dentition in tetrapods can be associated with major events in the history of terrestrial vertebrates. Dental occlusion, the process by which teeth from the upper jaw come in contact with those in the lower jaw, appears first in the fossil record in amniotes and their close relatives near the Permo-Carboniferous boundary approximately 300 million years ago. This evolutionary innovation permitted a dramatic increase in the level of oral processing of food in these early tetrapods, and has been generally associated with herbivory. Whereas herbivory in extinct vertebrates is based on circumstantial evidence, dental occlusion provides direct evidence about feeding strategies because jaw movements can be reconstructed from the wear patterns of the teeth. Examination of the evolution of dental occlusion in Paleozoic tetrapods within a phylogenetic framework reveals that this innovation developed independently in several lineages of amniotes, and is represented by a wide range of dental and mandibular morphologies. Dental occlusion also developed within diadectomorphs, the sister taxon of amniotes. The independent, multiple acquisition of this feeding strategy represents an important signal in the evolution of complex terrestrial vertebrate communities, and the first steps in the profound changes in the pattern of trophic interactions in terrestrial ecosystems.     

The Evolutionary Origin and Homologues of the Supracochlear Lamina: a Contribution to our Knowledge of Mammalian Ancestry

In cetaceans, each otic capsule is bound anteriorly to the remainder of the endocranium by an aliform structure called the supracochlear lamina. This lamina has been established here as the homologue of the sphenocochlear commissure of other mammals. Most researchers believe that these two structures arose within mammals. However, a comparative study involving data from both extant and extinct craniates suggests that the supracochlear lamina and the sphenocochlear commissure have a long premammalian history. The conclusion is that the lower of the two pharyngeal segments at the upper end of the mandibular branchial arch first became incorporated in the endocranium, constituting the cartilago trabecularis in the embryos of jawed craniates. The distal, or posterior, extremity of the trabecular cartilage projected outwardly to form the basitrabecular process. This process, which when ossified is called the processus basipterygoideus, functioned as a jaw support. Later, one more jaw support, termed the basivincular, evolved immediately in front of the basipterygoid process. Both these supports for the upper jaw existed in the piscine forerunners of the tetrapods. Subsequently, in the lineages leading to mammals, the basivincular jaw support became the processus alaris to which is attached an upper jaw endoskeletal remnant, called the ala temporalis. The basipterygoid jaw support, however, vanished as a consequence of the evolution of an exoskeletal joint system for the jaws of mammals. Yet the basipterygoid process persisted and as the sphenocochlear commissure or the supracochlear lamina assumed the new function of supporting the cochlear part of the auditory capsule.     

Myoseptal architecture of sarcopterygian fishes and salamanders with special reference to Ambystoma mexicanum

During axial undulatory swimming in fishes and salamanders muscular forces are transmitted to the vertebral axis and to the tail. One of the major components of force transmission is the myoseptal system. The structure of this system is well known in actinopterygian fishes, but has never been addressed in sarcopterygian fishes or salamanders. In this study we describe the spatial arrangement and collagen fiber architecture of myosepta in Latimeria, two dipnoans, and three salamanders in order to gain insight into function and evolution of the myoseptal system in these groups. Salamander myosepta lack prominent cones, and consist of homogenously distributed collagen fibers of various orientations that never form distinct tendons. Fiber orientations are difficult to homologize with those of fish myosepta. The myosepta of Latimeria and dipnoans (Protopterus and Neoceratodus) illustrate that major changes in architecture occurred in the sarcopterygian clade (loss of horizontal septum), in the rhipidistian (dipnoans + tetrapods) clade (loss of epineural and epipleural tendon), and in tetrapods (loss of lateral tendons and myoseptal folding). When compared to fishes, the myosepta of wholly aquatic salamanders (Ambystoma mexicanum, Amphiuma tridactylum, Necturus maculosus) do not have the lateral tendons we suppose serve to transfer muscular forces posteriorly. We propose that alternative structures (most conspicuously present in Ambystoma) perform this function: posteriorly the relative amount of connective tissue increases considerably, and myosepta are disintegrated to horizontal lamellae of connective tissue. The structures thought to be involved in modulation of body stiffness in fishes during swimming are also absent in salamanders. Our data also have implications for the hypothesis that salamander hypaxial myosepta are designed to increase shortening amplification of the hypaxial muscle fibers. The posterior hypaxial myosepta of all three salamander species possess only mediolaterally directed collagen fibers, which would indeed amplify the shortening of the associated muscle.     

The First Record of a Lungfish (Dipnoi,Sarcopterygii) in the Famennian Deposits (Upper Devonian) of the Tver Oblast

Paleontological Journal - A new dipnoan genus and species Anchidipterus dariae Krupina, gen. et sp. nov. (Dipteridae) is described based on a lower jaw specimen from the Famennian (Upper Devonian)...