Devonian Vertebrates From Colombia

Vertebrate remains are reported from the Emsian–Eifelian Floresta Formation and the Late Devonian (?Frasnian) Cuche Formation of north‐eastern Colombia. The material from the Floresta Formation is associated with a marine invertebrate fauna and includes an arthrodire and probably a rhenanid. Several vertebrate‐bearing localities are recorded from the Cuche Formation; vertebrates occur with plant remains and lingulid fragments. They include an acanthodian (Cheiracanthoides? sp.), a chondrichthyan (Antarctilamna? sp.), placoderms (Bothriolepis sp., Asterolepis? sp. and an undetermined groenlandaspidid or primitive brachythoracid arthrodire), a stegotrachelid actinopterygianand three sarcopterygians (a cosmine‐covered form tentatively referred to an osteolepidid, the porolepiform Holoptychiussp., and the rhizodontid Strepsodus? sp.). This assemblage suggests a Late Frasnian age and is surprisingly similar to Late Devonian vertebrate assemblages found in similar facies of Europe and North America, notwithstanding the presence of the Gondwanan chondrichthyan Antarctilamna?. key words: Vertebrata, Devonian, Colombia, South America, biostratigraphy, palaeobiogeography.     

First selenosteid placoderms from the eastern Anti‐Atlas of Morocco; osteology,phylogeny and palaeogeographical implications

Abstract: A new placoderm assemblage is reported from the Kellwasser facies of the eastern Anti‐Atlas, Morocco. This is the first record of an abundant Frasnian placoderm assemblage from Africa. The following new selenosteid taxa are described: Draconichthys elegans gen. et sp. nov., Enseosteus marocanensis sp. nov. and Walterosteus lelievrei sp. nov. In addition, material of Rhinosteus cf. parvulus is figured and described for the first time from Gondwana. The genera Enseosteus, Rhinosteus and Walterosteus are reviewed. ‘Wildungenichthys’ is regarded as a junior subjective synonym of Walterosteus. The new material and a revision of the taxa from Bad Wildungen, Germany, are the basis for a new phylogenetic analysis of the selenosteid arthrodires. The monophyly of the selenosteids is confirmed. The North American selenosteids are a sister group of Moroccan and European selenosteids, excluding Pachyosteus. The Moroccan vertebrate assemblage resembles that of Bad Wildungen. This supports the close palaeogeographical positions of Gondwana and Laurussia during the Late Devonian and indicates that biogeographical barriers for placoderms that could have separated the two assemblages were absent.     

Further evidence for the presence of holoptychiid porolepiforms (Sarcopterygii,Dipnomorpha) from the Frasnian of Colombia

The Devonian (Frasnian) of Colombia has provided important information on the taxonomical diversity and palaeobiogeographic relationships of the Late Devonian marginal marine vertebrate faunas of South America (northwestern margin of Gondwana). This fauna is mostly composed of Gondwanan endemics, but includes two taxa also known in Euramerica: the antiarchan placoderm Asterolepis and the porolepiform sarcopterygian Holoptychius. The occurrence of holoptychiid porolepiforms in Colombia was previously suggested based solely on the presence of scales attributed to Holoptychius, although with caution. Here, we describe further holoptychiid remains that include a large isolated tooth with dendrodont microstructure, typical of porolepiforms, and additional Holoptychius scales. These new findings increase the record of holoptychiid porolepiform occurrences in Gondwana and suggest that dispersion of fish faunas between Euramerica and Gondwana by the beginning of the Late Devonian was possible through South America.     

The fish swimming trace Undichna unisulca from the Silurian of Sweden: probably the oldest vertebrate locomotion trace fossil

The fish swimming trace Undichna unisulca is reported from a sandstone block collected from the Upper Silurian (Ludlow) Burgsvik Formation of Gotland, Sweden. It represents the oldest record of a vertebrate trace (apart from coprolites), predating previous finds from the Lower Devonian by at least 10 Ma. A thelodont or acanthodian fish may have produced the sinusoidal trails with the aid of their caudal or anal fins while browsing slowly over the sediment whilst searching for food.     

A three‐dimensional placoderm (stem‐group gnathostome) pharyngeal skeleton and its implications for primitive gnathostome pharyngeal architecture

The pharyngeal skeleton is a key vertebrate anatomical system in debates on the origin of jaws and gnathostome (jawed vertebrate) feeding. Furthermore, it offers considerable potential as a source of phylogenetic data. Well‐preserved examples of pharyngeal skeletons from stem‐group gnathostomes remain poorly known. Here, we describe an articulated, nearly complete pharyngeal skeleton in an Early Devonian placoderm fish, Paraplesiobatis heinrichsi Broili, from Hunsrück Slate of Germany. Using synchrotron light tomography, we resolve and reconstruct the three‐dimensional gill arch architecture of Paraplesiobatis and compare it with other gnathostomes. The preserved pharyngeal skeleton comprises elements of the hyoid arch (probable ceratohyal) and a series of branchial arches. Limited resolution in the tomography scan causes some uncertainty in interpreting the exact number of arches preserved. However, at least four branchial arches are present. The final and penultimate arches are connected as in osteichthyans. A single median basihyal is present as in chondrichthyans. No dorsal (epibranchial or pharyngobranchial) elements are observed. The structure of the pharyngeal skeleton of Paraplesiobatis agrees well with Pseudopetalichthys from the same deposit, allowing an alternative interpretation of the latter taxon. The phylogenetic significance of Paraplesiobatis is considered. A median basihyal is likely an ancestral gnathostome character, probably with some connection to both the hyoid and the first branchial arch pair. Unpaired basibranchial bones may be independently derived in chondrichthyans and osteichthyans.     

A new find of the placoderm genus <Emphasis Type="Italic">Bothriolepis</Emphasis> Eichwald in the Upper Devonian of Uzbekistan

A new placoderm species, Bothriolepis sanzarensis sp. nov., from the Upper Frasnian (Upper Devonian) of the western marginal area of the Turkestan Mountain Ridge (Samarkand Region, Uzbekistan) is described. The new species is distinguished by the isometric anterior medio-dorsal plate, with a broad posterior region, and the undulating sutures between this and other trunk plates. This is the second find of Bothriolepididae in Uzbekistan.     

The placoderm <Emphasis Type="Italic">Plourdosteus livonicus</Emphasis> (Eastman) in the early Frasnian of the Central Devonian Field and the trophic structure of the Mikhailovskii Fish Assemblage

The placoderm Plourdosteus livonicus (Eastman) (Euarthrodira, Coccosteina) from the Upper Timanian Mikhailovskii Fish Assemblage of the Central Devonian Field is described. The composition and structure of the assemblage are discussed and compared with later Frasnian fish assemblages of the Central Devonian Field.     

云南丽江早泥盆世棘鱼类微体化石的首次报道

记述了云南丽江县鸣音区阿冷初早泥盆世山江组 (Lochkovian EarlyPragian)和阿冷初组(Middle LatePragian)棘鱼亚纲微体化石。这些化石被归属于 4属 5种 ,其中包含 1新属 2新种。这是西藏—滇西区棘鱼类化石的首次报道。不同于华南区棘鱼类的组成 ,丽江早泥盆世棘鱼类以栅棘鱼目属种为主 ,缺少锉棘鱼目属种。它与国外对比 ,则与澳大利亚新南威尔士中部同时代的棘鱼类表现出较多的相似性     

The Upper Devonian tetrapodomorph Gogonasus andrewsae from Western Australia: Reconstruction of the shoulder girdle and opercular series using X-ray Micro-Computed Tomography

The tetrapodomorph fish, Gogonasus andrewsae is a three dimensionally well-preserved sarcopterygian from the Gogo Formation (Frasnian, early Upper Devonian, ∼380 million years ago) in Western Australia. High-resolution X-ray Micro-Computed Tomography and 3D printouts were used to obtain a digital reconstruction of its shoulder girdle and opercular series. Our new findings show the opercular series in a close fit against the upper bones of the shoulder girdle only if the anocleithrum, supracleithrum and post-temporal are aligned more horizontally than in previous reconstructions. The lowermost subopercular bone also differs, in partly covering the clavicle of the shoulder girdle. The ascending process of the clavicle, and the ventral process of the anocleithrum, do not fit closely inside the cleithrum, and perhaps functioned for ligamentous attachment. A rugose area on the anocleithral process is in a similar relative position to the attachment of a muscle ligament on the shoulder girdle of various living actinopterygians. Our manipulation of 3D printouts permits testing of the morphological fit of extremely fragile acid-etched bones, and indicates a new way to investigate the constructional morphology of one or more mechanical units of the vertebrate skeleton. It is suggested that Micro-CT imaging, reconstruction, visualisation and 3D printing techniques will provide a rigorous new test leading to modification of previous reconstructions of extinct vertebrates that were based on graphical methods and 2D imaging.     

A microanatomical and histological study of the fin long bones of the Devonian sarcopterygian Eusthenopteron foordi

Meunier F.J. and Laurin M. 2012. A microanatomical and histological study of the fin long bones of the Devonian sarcopterygian Eusthenopteron foordi. —Acta Zoologica (Stockholm) 93 : 88–97. A paleohistological study of the endoskeletal bones of the dorsal and pelvic fins shows that Eusthenopteron foordi had true long bones that grew in length and thickness through endochondral and periosteal ossification, respectively. Endochondral ossification required cartilaginous epiphyses with a growth plate system whose presence is confirmed by both calcified cartilage and thin endochondral bony trabeculae that overlaid the erosive bays located in hypertrophic calcified cartilage. Articulations between axial mesomeres in paired fins were diarthroses. This microanatomical organization, i.e. longitudinal growth of diaphysis and articulations between epiphyses, can be considered an exaptation for terrestrial locomotion as it can produce skeletal elements able to support strong mechanical stress.     

Late Devonian - Early Carboniferous vertebrate microremains from the Carnic Alps, northern Italy

Vertebrate microremains from the Late Devonian-Early Carboniferous of the Carnic Alps are predominantly chondrichthyan, with minor placoderm and actinopterygian remains. The faunas are sparse and, with very few exceptions, occur only in conodont-rich pelagic limestones (Pramosio Limestone) representative of the palmatolepid-bispathodid conodont biofacies. Phoebodont and jalodont chondrichthyans, also reflecting open-ocean environments, predominated during the Famennian, and eventually symmoriids seem to predominate during the Early Carboniferous. The presence of Siamodus in this assemblage gives a new locality for this genus known from few regions in the world and allows confirming its stratigraphical range (limpidus Zone) and its relation to deep-water environments. The Late Devonian vertebrate faunas are tropical and cosmopolitan, having much in common with coeval taxa from the North-Gondwanan margins and Asian terranes. Composition of the vertebrate faunas is consistent with the Carnic Alps terrane having occupied a position intermediate between Gondwana and Laurussia, as hypothesized by various authors, but because of sparsity of the taxa represented and the pronounced cosmopolitan nature of both the conodont and vertebrate faunas, the data are not compelling.     

Diet selectivity in a terrestrial forest invertebrate,the Auckland tree wētā, across three habitat zones

Insects are important but overlooked components of forest ecosystems in New Zealand. For many insect species, information on foraging patterns and trophic relationships is lacking. We examined diet composition and selectivity in a large‐bodied insect, the Auckland tree wētā Hemideina thoracica, in three habitat zones in a lowland New Zealand forest. We asked whether H. thoracica selectively forage from available plant food sources, and whether these choices were lipid‐rich compared to nonpreferred available plants. We also identified the proportion of invertebrates in their frass as a proxy for omnivory. From reconnaissance plot sampling, together with fecal fragment analysis, we report that more than 93% of individual tree wētā had eaten invertebrates before capture. Additionally, wētā in the highest elevation hillslope habitat zone consumed significantly fewer species of plants on average than wētā on the low‐elevation terrace habitat. Upper hillslope wētā also had the highest average number of invertebrate fragments in their frass, significantly more than wētā in the low‐elevation terrace habitat zone. Wētā showed high variability in the consumption of fruit and seeds across all habitat zones. Generally, we did not observe diet differences between the sexes (although it appears that male wētā in the mid‐hillslope habitat ate fruits and seeds more voraciously than females), suggesting that the sexes have similar niche breadths and display similar degrees of omnivorous behavior. Extraction of leaf lipids demonstrated a range of lipid content values in available plants, and Ivlev's Electivity Index indicated that plant species which demonstrated high electivity tended to have higher concentrations of lipids in their leaves. Our findings indicate that H. thoracica forage omnivorously and selectively, and hence play multiple roles in native ecosystems and food webs.     

Primitive bony fishes, with especial reference to Cheirolepis and palaeonisciform actinopterygians

The relationships of the Devonian palaeonisciform fish Cheirolepis are examined and the early evolutionary trends within the Actinopterygii and the Osteichthyes are considered.
Cheirolepis is the most primitive known actinopterygian. The contemporary stegotrachelid palaeonisciforms are more advanced in their cranial and locomotor anatomy. The general directions of these advances are similar to those subsequently displayed by later palaeonisciforms over the stegotrachelids themselves. Cheirolepis , furthermore, possesses many characters which can be logically interpreted as primitive for the Osteichthyes by extrapolation of trends in actinopterygian and sarcopterygian lineages. 11 is the most primitive known osteichthyan.
The Osteichthyes are considered to have arisen from a micromerically-scaled acanthodian or acanthodian-like ancestor at the end of the Silurian period.     

Fossilized ontogenies: the contribution of placoderm ontogeny to our understanding of the evolution of early gnathostomes

Placoderms, representing phylogenetically more inclusive jawed vertebrates and successive sister taxa to crown‐group gnathostomes, are critical to our understanding of character evolution within the crown‐group (chondrichthyans + osteichthyans), including developmental characters. Early ontogenetic stages of placoderms are generally poorly known, although some exceptional faunas preserve both embryonic (e.g. from the Gogo Formation, Western Australia) and post‐embryonic individuals (the Miguasha Formation, Canada; Lode Formation, Latvia; Merriganowry Formation, Gogo Formation, Australia). Information provided by these ontogenies is relevant to questions of placoderm taxonomy and phylogeny, but also to broader questions pertinent to vertebrate evolution as a whole, for example, evolution of bone development, evolution of the axial skeleton and evolution of reproduction.     

DISTRIBUTION OF VERTEBRATES IN THE SILURIAN OF ESTONIA

A rich collection of Silurian vertebrate remains, including abundant thelodont and acanthodian scales, has been made from recent borings on the Estonian island of Saaremaa (Ösel). Thelodonts have been reported for the first time at the top of the Jaagarahu Stage, as have anaspids from the Rootsiküla (formerly Kaarma) Stage, and acanthodians from the Paadla Stage. Arthrodires as well as actinopterygians (?), represented by the genus Andreolepis, have been found for the first time on Saaremaa. Arthrodires appear to be the oldest known members of this group. The recent stratigraphical classification of the Silurian of Estonia has been taken into account, and the stratigraphical distribution of the vertebrate genera and species is given. The correlation between some of the vertebrate-bearing beds of Saaremaa, Gotland, and the southern Baltic is also discussed.     

Trace fossil evidence of predation upon bone‐eating worms on a baleen whale skeleton from the Oligocene of New Zealand

The osteophagous worm Osedax (Annelida: Siboglinidae) colonizes vertebrate bones in deep‐sea environments globally. Osedax bioerosion of modern bones suggests a potentially destructive agent in the marine vertebrate fossil record, but the dearth of published reports of abundant Osedax traces suggests an uncertain taphonomic influence of this organism. This study reports Osedax traces (Osspecus boreholes, pockmarks and collapsed galleries) in an Oligocene baleen whale (Cetacea: Eomysticetidae) from New Zealand, which constitute the first record of fossil Osedax traces from the southern hemisphere. Some Osedax traces are cross‐cut by linear biogenic scrape marks, implying that sharks or bony fish fed upon Osedax worms, a process which compounds or potentially accelerates worm‐inflicted damage to vertebrate bones in marine environments.     

“Lungs” in Placoderms,a persistent palaeobiological myth related to environmental preconceived interpretations

The presence of an aerial breathing organ in Placoderms is noticed in many textbooks on the history of breathing in Vertebrates. The origin of this interpretation is from a paper published in 1941 dealing with the interpretation of the differential sedimentary infilling of the armour of the antiarch Bothriolepis canadensis from the Late Devonian Escuminac formation of Canada. A revision of this material shows that if some sedimentary structures could be interpreted as traces of some digestive organs, none could be interpreted as putative lungs. The original proposal was based mainly on a presupposed mode of life of Bothriolepis in a freshwater environment of an alluvial plain. Recent studies of the Escuminac Formation environment conclude that it represents a marginal marine environment. Moreover, Bothriolepis has a worldwide distribution, notably in strictly marine environment. Thus, the presence of lungs in Bothriolepis remains highly questionable: it cannot be supported by anatomical, phylogenetic, nor biological arguments.