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.     

LATE DEVONIAN (FAMENNIAN) LUNGFISHES FROM THE CATSKILL FORMATION OF PENNSYLVANIA, USA

Abstract:  Occurrences of fossil lungfishes (Dipnoi: Sarcopterygii) in the Famennian Catskill Formation of Pennsylvania are reviewed. A nearly complete dermal skull roof is assigned to a new genus and species, Apatorhynchus opistheretmus . Other recently discovered lungfish specimens include an incomplete postcranium similar to that of the Frasnian genus Fleurantia , a small parasphenoid of uncertain affinities, and isolated toothplates. Previously described dipnoan remains from the Catskill Formation include a partial skull roof of Soederberghia groenlandica , toothplates assigned to several species of Dipterus , a putative rostral or symphysial region placed in the problematic form taxon Ganorhynchus , and sedimentary structures interpreted as burrows. The toothplates attributed to Dipterus are indeterminate and are placed in open nomenclature, while the specimen identified as Ganorhynchus is not convincingly dipnoan. The status of the burrows remains uncertain pending the discovery of lungfish remains within these or similar structures in Catskill deposits. The distinct ichthyofaunas within the Catskill Formation and their lungfish components are briefly reviewed. Lungfishes are found in the Holoptychius - and Bothriolepis -dominated faunas common in the Catskill succession, as well as in the compositionally distinctive Red Hill assemblage. Many of the Devonian continental faunas that contain tetrapods also include long-snouted, denticle-bearing lungfishes ('rhynchodipterids', fleurantiids, or both). The composition of Late Devonian ichthyofaunas may have predictive qualities that will allow researchers to identify localities likely to produce the remains of early tetrapods.     

On the structure of the lower jaw in dipnoans: with a description of an early Devonian dipnoan from Canada, Melanognathus canadenis gen. et sp.nov.

The lower jaw in Melanognathus gen. nov. and several other Devonian dipnoans is described and compared with that in Neoceratodus. It is a most conservative structure, which apart from the effect of the retrogressive development of the skeleton has hardly changed since Devonian times. A new interpretation of the sensory canal bones is given. With respect to these elements and the structure of the lower jaw as a whole, the dipnoans differ fundamentally from the rhipidistid crossopterygians, tetrapods and aetinopterygians. Several resemblances to the lower jaw in holocephalians are demonstrated. The results confirm the view that the Dipnoi comprise an early specialized and isolated group, perhaps more closely related to the elasmo-branchiomorphs than to the teleostomes and tetrapods.     

The relationships of Uronemus: a carboniferous dipnoan with highly modified tooth plates

Previous accounts of the dentition of the Carboniferous dipnoan Uronemus have stressed the significance of the scattered small denticles. These, together with the marginal teeth and ridges, have been interpreted as primitive characters of the dipnoan dentition shared with three other genera: the Devonian Uranlophus and Griphognathus and the Carboniferous to Permian Conchopoma. Genera with tooth plates have been considered to be a monophyletic group in which tooth plates are a derived character; Uronemus has been excluded from this group in all previous investigations dealing with the significance of the dentition for determining relationships among dipnoans. The macromorphology of the dentition of Uronemus has been re-examined and correlated with the histology of all the dental tissues. Optical study of thin sections and scanning electron microscope study of the adjacent cut surfaces has shown that the hard, wear-resistant dentine of the teeth and ridges is petrodentine. The arrangement, growth, wear and histology of the dental tissues have been compared with those of denticulated and tooth-plated genera. The arrangement of new teeth relative to the tooth ridge, the pattern of wear along the ridge, and the type of dentine and its growth indicate that the dentition of Uronemus is best interpreted as a tooth plate with one long lingual tooth ridge and reduced lateral tooth rows. Therefore the marginal tooth ridges are not considered to be homologous with those of denticulate dipnoans such as Uranolophus. The presence of petrodentine, a tissue type only found in forms with tooth plates, is consistent with the view that the dentition is derived by modification of a radiate tooth plate. The denticles covering restricted regions of the palate and lower jaw are considered to have been a secondary acquisition. The suggestion that Conchopoma is a close relative of Uronemus is not accepted, and possible new relationships have been proposed. New data on Scaumenacia and Phaneropleuron, two other genera previously compared with Uronemus, are presented. Rhinodipterus, a form with elongate lingual ridges, is also discussed. Phaneropleuron is shown to have radiate tooth plates and not a marginal row of conical teeth as previously described. It is proposed that the tooth plate of Uronemus is derived from a dipterid type of plate. A discussion of some of the other factors involved in determining the relationships of the genus is given.(ABSTRACT TRUNCATED AT 400 WORDS)     

广西下泥盆统南极鱼类(盾皮鱼纲:节甲鱼目)一新属(英文)

依据采自广西平果早泥盆世郁江组中部的化石材料,记述了盾皮鱼纲节甲鱼目南极鱼科的一新属、新种——小眼坡塘鱼(Potangaspis parvoculatusgen.et sp.nov.)。该新属的眶孔很小,且为颅顶甲骨片所完全包围;中点线沟发育,与眶上感觉沟、中心感觉沟和后点线沟一起共同向颈片的骨化中心辏合。系统发育分析表明新属和发现于南极洲中泥盆统的南极鱼属构成姐妹群关系,支持了南极鱼科位于节甲鱼目最基干位置的假说,同时为早埃姆斯期东冈瓦纳大陆与中国南方之间的古地理密切联系提供了新的证据。     

The identification of the dermal bones of the head

The discovery of the ichthyostegid Amphibia in Upper Devonian rocks by Säve-Söderbergh (1932) introduced further difficulties into the already complex problems of the dermal bones of the skull roof. For some years previously ideas about the origin of the tetrapods had been dominated by Watson's (1926) Croonian Lecture in which he had demonstrated beyond reasonable doubt that the crossopterygian fishes and not the Dipnoi were their ancestors, and had attempted to show that many of the features of the Carboniferous labyrinthodonts were a direct inheritance from these fishes. It was to be expected, therefore, that any Amphibia from the Upper Devonian would be intermediate in their structures between the Middle Devonian osteolepids and the Carboniferous labyrinthodonts, but when discovered the ichthyostegids did not conform at all well to this expectation. While their skulls showed some very primitive features which might have been expected, the pattern of the dermal bones did not conform to plan, for these new animals had lost, it seemed, the intertemporals, bones found in both the osteolepids and nearly all early labyrinthodonts, and had a single postparietal bone in place of the paired bones of all other early Amphibia. The osteolepid skull had many more bones than these earliest Amphibia.     

The structure and stratigraphy of<Emphasis Type="Italic">Speonesydrion</Emphasis> from New South Wales,Australia, and the dentition of primitive dipnoans

New material ofSpeonesydrion iani, an Early Devonian dipnoan from New South Wales, has provided additional Information on the dentition and jaws. Two new partial palates have been found, and X-rays of the parasphenoid shows that the structure is well preserved. The palatal teeth are well worn even in partly grown material, and they do not originate at a growth point, but at a thickening of the palate. More mandibles have been collected, and thin sections have been prepared to allow a discussion of their histology. On the mandible the teeth are clear, and they are much more defined than they are on the palate. The dental heel is variably developed, and grows in phases by thickening of the dentine at the contact with the bone. Dentine forms on the bone at the base of the heel, partly by Solution of the bone and the addition of dentine from the pulp canals, but also by direct growth from the pulp canals dorsal to the bone. In the latter case the dentine and bone are in contact, and the two tissues intermingle. The teeth are also formed on a thickened bone and consist of dentine capped with enamel making a crest. Dentine and bone are related as in the heel. We conclude that the teeth inSpeonesydrion are not homologous with the teeth in other dipnoans, and are formed by a different process involving the aggregation of denticles.     

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.     

Developmental plasticity and disparity in early dipnoan (lungfish) dentitions

Although the lungfish (Dipnoi) belong within the Osteichthyes, their dentitions are radically different from other osteichthyans. Lungfish dentitions also show a uniquely high structural disparity during the early evolution of the group, partly owing to the independent variation of odontogenic and odontoclastic processes that are tightly and stereotypically coordinated in other osteichthyans. We present a phylogenetic analysis of early lungfishes incorporating a novel approach to coding these process characters in preference to the resultant adult dental morphology. The results only partially resolve the interrelationships of Devonian dipnoans, but show that the widely discussed hypothesis of separate tooth-plated, dentine-plated, and denticulated lineages is unlikely to be true. The dipnoan status of Diabolepis is corroborated. Lungfish dentitions seem to have undergone extensive and nonparsimonious evolution during the early history of the group, but much of the resulting disparity can be explained by a modest number of evolutionary steps in the underlying developmental processes, those for dental formation (odontogenic) and those for the remodeling of dentine tissue (odontoclastic). Later in lungfish evolution, this disparity was lost as the group settled to a pattern of dental development that is just as stereotypic as, but completely different from, that of other osteichthyans.     

Air-breathing adaptation in a marine Devonian lungfish

Recent discoveries of tetrapod trackways in 395 Myr old tidal zone deposits of Poland (Niedźwiedzki et al. 2010 Nature 463, 43–48 (doi:10.1038/nature.08623)) indicate that vertebrates had already ventured out of the water and might already have developed some air-breathing capacity by the Middle Devonian. Air-breathing in lungfishes is not considered to be a shared specialization with tetrapods, but evolved independently. Air-breathing in lungfishes has been postulated as starting in Middle Devonian times (ca 385 Ma) in freshwater habitats, based on a set of skeletal characters involved in air-breathing in extant lungfishes. New discoveries described herein of the lungfish Rhinodipterus from marine limestones of Australia identifies the node in dipnoan phylogeny where air-breathing begins, and confirms that lungfishes living in marine habitats had also developed specializations to breathe air by the start of the Late Devonian (ca 375 Ma). While invasion of freshwater habitats from the marine realm was previously suggested to be the prime cause of aerial respiration developing in lungfishes, we believe that global decline in oxygen levels during the Middle Devonian combined with higher metabolic costs is a more likely driver of air-breathing ability, which developed in both marine and freshwater lungfishes and tetrapodomorph fishes such as Gogonasus.     

Sampling diverse characters improves phylogenies: Craniodental and postcranial characters of vertebrates often imply different trees

Morphological cladograms of vertebrates are often inferred from greater numbers of characters describing the skull and teeth than from postcranial characters. This is either because the skull is believed to yield characters with a stronger phylogenetic signal (i.e., contain less homoplasy), because morphological variation therein is more readily atomized, or because craniodental material is more widely available (particularly in the palaeontological case). An analysis of 85 vertebrate datasets published between 2000 and 2013 confirms that craniodental characters are significantly more numerous than postcranial characters, but finds no evidence that levels of homoplasy differ in the two partitions. However, a new partition test, based on tree‐to‐tree distances (as measured by the Robinson Foulds metric) rather than tree length, reveals that relationships inferred from the partitions are significantly different about one time in three, much more often than expected. Such differences may reflect divergent selective pressures in different body regions, resulting in different localized patterns of homoplasy. Most systematists attempt to sample characters broadly across body regions, but this is not always possible. We conclude that trees inferred largely from either craniodental or postcranial characters in isolation may differ significantly from those that would result from a more holistic approach. We urge the latter.     

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.     

云南武定节甲类的新材料

文中描述了采自云南武定中泥盆统一新的属种Yinostius maior gen.et sp.nov.属于短胸节甲类Heterosteidae科,这类化石在我国系初次发现。     

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.     

云南沾益早泥盆世肺鱼类一新种

记述了云南沾益地区下泥盆统穿洞组肺鱼化石一新种——多脊混磨鱼(Tarachomylax multicostatus sp.nov.),归入标本包括一件左上齿板、一块B骨、一块可能是3骨的骨片和6个零散鳞片。主要特征:齿板内侧及后侧边缘被小齿(denticles)覆盖;齿脊11-12列,齿板中缘与最后一列齿脊夹角约135°;内翼骨侧凹不明显,后侧部窄小呈尖状;B骨和J骨上的前凹线沟相连;鳞片出露区与被覆压区之间有小齿瘤带。