Nanomechanical variability in the early evolution of vertebrate dentition.

Conodonts are an extinct group of primitive jawless vertebrates whose elements represent the earliest examples of a mineralized feeding apparatus in vertebrates. Their relative relationship within vertebrates remains unresolved. As teeth, conodont elements are not homologous with the dentition of vertebrates, but they exhibit similarities in mineralization, growth patterns, and function. They clearly represent an early evolutionary experiment in mineralized dentition and offer insight into analogous dentition in other groups. Unfortunately, analysis of functional performance has been limited to a handful of derived morphologies and material properties that may inform ecology and functional analysis are virtually unknown. Here we applied a nanoscale approach to evaluate material properties of conodont bioapatite by utilizing Atomic Force Microscopy (AFM) nanoindentation to determine Young's modulus (E) along multiple elements representing different ontogenetic stages of development in the coniform-bearing apparatus of Dapsilodus obliquicostatus. We observed extreme and systematic variation in E along the length (oral to aboral) of each element that largely mirrors the spatial and ontogenetic variability in the crystalline structure of these specimens. Extreme spatial variability of E likely contributed to breakage of elements that were regularly repaired/regrown in conodonts but later vertebrate dentition strategies that lacked the ability to repair/regrow likely required the development of different material properties to avoid structural failure.

Atlanticalymene, a new genus of Middle Ordovician (Darriwilian) calymenine trilobites, and revision of the calymenoidean genus Protocalymene Ross.

Middle Ordovician (Darriwilian) species representing early Laurentian occurrences of the Subfamily Calymeninae Milne Edwards, 1840 (=Flexicalymeninae Siveter, 1977) are assigned to Atlanticalymene n. gen. (type species: A. bardensis n. sp. from the Table Cove Formation, western Newfoundland, Canada). They have routinely been confused with the older (Dapingian) calymenoidean taxon Protocalymene Ross, 1967. Revision of the type species of Protocalymene, P. mcallisteri Ross, 1967, from the Antelope Valley Formation, Funeral Mountains, California, indicates that it is not a calymenine, and that while it is clearly a calymenoidean its close affinity is otherwise difficult to determine. A single genuine calymenine species is known from the Laurentian Dapingian, and revised here as "Calymeninae n. gen.? n. sp. A" from the Antelope Valley Formation, Nevada, USA. A species from the Dapingian of Tarim, known from a single partial cranidium, appears to represent an older, extra-Laurentian species of Atlanticalymene.

Ontogenetic variability in crystallography and mosaicity of conodont apatite: implications for microstructure, palaeothermometry and geochemistry.

X-ray diffraction data from Silurian conodonts belonging to various developmental stages of the species Dapsilodus obliquicostatus demonstrate changes in crystallography and degree of nanocrystallite ordering (mosaicity) in both lamellar crown tissue and white matter. The exclusive use of a single species in this study, combined with systematic testing of each element type at multiple locations, provided insight into microstructural and crystallographic differentiation between element type (Sa , Sb -c , M) as well as between juveniles and adults. A relative increase in the unit cell dimensions a/c ratio of nanocrystallites during growth was apparent in areas demonstrating single-crystal behaviour, but no such relationship was seen in dominantly polycrystalline areas. Systematic variations in mosaicity were identified, with mosaicity (as a proxy for disorder) increasing during growth, as well as along elements from tip to base. These results provide potential insight into the integrity of conodont apatite as a recorder of palaeoseawater chemistry, as well as demonstrate the need to consider the influence of ontogeny and element type on the use of conodonts in palaeothermometry and geochemical investigations.

The pliomerid trilobite Ibexaspis and related new genera, with species from the Early Ordovician (Floian; Tulean, Blackhillsian) of the Great Basin, western USA.

Field-based revision and phylogenetic analysis demonstrate that the pliomerid trilobite taxon Ibexaspis Pribyl and Vanek (in Pribyl et al., 1985), previously known from a single formally named species (I. brevis [Young, 1973]), belongs to a complex of 14 mostly newly discovered, related species from the Early Ordovician (Floian; Tulean and Blackhillsian) of the northern Laurentian margin. The species are known from silicified samples recovered from sections in eastern Nevada, western Utah, and southeastern Idaho. The stratigraphically early Tuleaspis n. gen. (type species: T. jeneki n. sp.; Tulean; low Protopliomerella contracta Zone) includes its type and two species described in open nomenclature. Tuleaspis is sister to the remainder of the clade. Ibexaspis now includes three additional species: I. coadyi n. sp. (Blackhillsian; Carolinites nevadensis Zone), I. leuppi n. sp. (Blackhillsian; Presbynileus ibexensis Zone), and I. rupauli n. sp. (Blackhillsian; "Pseudocybele nasuta Zone"). Ibexapsis is sister to a clade of Millardaspis n. gen. + Deltapliomera n. gen. Millardaspis (type species M. milsteadi n. sp.; Tulean; Heckethornia hyndeae Zone) also includes M. knoxi n. sp. (Tulean; Panisaspis sevierensis Zone). Deltapliomera (type species D. humphriesi n. sp.; Blackhillsian, Carolinites nevadensis Zone) also includes D. inglei n. sp. (Tulean; Heckethornia bowiei Zone), D. heimbergi (Tulean; Panisaspis sevierensis Zone), D. eppersoni n. sp. (Blackhillsian; Bathyurina plicolabeona Zone), and a species described in open nomenclature.