Increasing control over biomineralization in conodont evolution.

Vertebrates use the phosphate mineral apatite in their skeletons, which allowed them to develop tissues such as enamel, characterized by an outstanding combination of hardness and elasticity. It has been hypothesized that the evolution of the earliest vertebrate skeletal tissues, found in the teeth of the extinct group of conodonts, was driven by adaptation to dental function. We test this hypothesis quantitatively and demonstrate that the crystallographic order increased throughout the early evolution of conodont teeth in parallel with morphological adaptation to food processing. With the c-axes of apatite crystals oriented perpendicular to the functional feeding surfaces, the strongest resistance to uniaxial compressional stress is conferred along the long axes of denticles. Our results support increasing control over biomineralization in the first skeletonized vertebrates and allow us to test models of functional morphology and material properties across conodont dental diversity.

Taxonomy and stratigraphic distribution of Lotagnostus (Agnostida: Agnostidae) and associated trilobites and conodonts in the Upper Cambrian (Furongian) of Laurentia.

Two Lotagnostus-dominated faunas from the Windfall Formation at Ninemile Canyon in the Antelope Range of Nevada, USA, are described: an older Lotagnostus nolani Fauna and younger L. rushtoni Fauna. The former is dominated by two morphs of Lotagnostus, one strongly scrobiculate and the other smooth to weakly scrobiculate. Both morphs fall within the broad concept advocated for L. americanus by Peng et al. (2015). The numerous (>1400 sclerites) specimens of Lotaganostus in collections of the L. nolani Fauna confirm that the two morphs do not intergrade and remain distinct throughout ontogeny. Both display multiple traits that distinguish them from the type material of L. americanus, justifying treatment as separate species. Similarly unique, diagnostic features were identified to restore the Asian species L. punctatus and L. asiaticus to full species status, whereas deficiencies in the type material for L. americanus warrant restriction of the name to the holotype. New species described from the Windfall include five agnostoids (Lotagnostus nolani, L. clarki, L. morrisoni, L. rushtoni, and Neoagnostus parki) and one trilobite (Bienvillia eurekensis). Plicatolina nyensis Taylor is reassigned to Mendoparabolina on the form of its pygidium. Conodonts from the Catlin Member of the Windfall Formation and overlying informal Caryocaris shale member of the Goodwin Formation at Ninemile Canyon provide a late Sunwaptan (Eoconodontus Zone) age for the Lotagnostus rushtoni Fauna and assign the entire Caryocaris shale to the early Ordovician Rossodus manitouensis Zone. Combined with published data on trilobite faunas, the conodont faunas confirm strong diachroneity for the top of the Catlin, and a lack of overlap in age between the Caryocaris shale and Bullwhacker Member of the Windfall in ranges to the north and east. Co-occurrence of Lotagnostus nolani and Mendoparabolina nyensis establishes age equivalence of the L. nolani Fauna with the Hedinaspis-Charchaqia (HC) Fauna at the base of the Hales Limestone in the Hot Creek Range, and earlier correlations of the latter with the L. punctatus Zone in Asia are supported. However, isolation of the HC Fauna in starved-basin deposits above a major sequence boundary at the base of the Hales, and ecologic restriction of Lotagnostus to lower slope and basinal environments that prevented association with endemic shallow marine taxa, renders correlation into the biostratigraphy of Laurentian upper slope and platform imprecise on the order of 10s, if not 100s of meters.

Growth and feeding ecology of coniform conodonts.

Conodonts were the first vertebrates to develop mineralized dental tools, known as elements. Recent research suggests that conodonts were macrophagous predators and/or scavengers but we do not know how this feeding habit emerged in the earliest coniform conodonts, since most studies focus on the derived, 'complex' conodonts. Previous modelling of element position and mechanical properties indicate they were capable of food processing. A direct test would be provided through evidence of in vivo element crown tissue damage or through in vivo incorporated chemical proxies for a shift in their trophic position during ontogeny. Here we focus on coniform elements from two conodont taxa, the phylogenetically primitive Proconodontus muelleri Miller, 1969 from the late Cambrian and the more derived Panderodus equicostatus Rhodes, 1954 from the Silurian. Proposing that this extremely small sample is, however, representative for these taxa, we aim to describe in detail the growth of an element from each of these taxa in order to the test the following hypotheses: (1) Panderodus and Proconodontus processed hard food, which led to damage of their elements consistent with prey capture function; and (2) both genera shifted towards higher trophic levels during ontogeny. We employed backscatter electron (BSE) imaging, energy-dispersive X-ray spectroscopy (EDX) and synchrotron radiation X-ray tomographic microscopy (SRXTM) to identify growth increments, wear and damage surfaces, and the Sr/Ca ratio in bioapatite as a proxy for the trophic position. Using these data, we can identify whether they exhibit determinate or indeterminate growth and whether both species followed linear or allometric growth dynamics. Growth increments (27 in Pa. equicostatus and 58 in Pr. muelleri) were formed in bundles of 4-7 increments in Pa. equicostatus and 7-9 in Pr. muelleri. We interpret the bundles as analogous to Retzius periodicity in vertebrate teeth. Based on applied optimal resource allocation models, internal periodicity might explain indeterminate growth in both species. They also allow us to interpret the almost linear growth of both individuals as an indicator that there was no size-dependent increase in mortality in the ecosystems where they lived e.g., as would be the case in the presence of larger predators. Our findings show that periodic growth was present in early conodonts and preceded tissue repair in response to wear and damage. We found no microwear and the Sr/Ca ratio, and therefore the trophic position, did not change substantially during the lifetimes of either individual. Trophic ecology of coniform conodonts differed from the predatory and/or scavenger lifestyle documented for "complex" conodonts. We propose that conodonts adapted their life histories to top-down controlled ecosystems during the Nekton Revolution.

The origin of conodonts and of vertebrate mineralized skeletons.

Conodonts are an extinct group of jawless vertebrates whose tooth-like elements are the earliest instance of a mineralized skeleton in the vertebrate lineage, inspiring the 'inside-out' hypothesis that teeth evolved independently of the vertebrate dermal skeleton and before the origin of jaws. However, these propositions have been based on evidence from derived euconodonts. Here we test hypotheses of a paraconodont ancestry of euconodonts using synchrotron radiation X-ray tomographic microscopy to characterize and compare the microstructure of morphologically similar euconodont and paraconodont elements. Paraconodonts exhibit a range of grades of structural differentiation, including tissues and a pattern of growth common to euconodont basal bodies. The different grades of structural differentiation exhibited by paraconodonts demonstrate the stepwise acquisition of euconodont characters, resolving debate over the relationship between these two groups. By implication, the putative homology of euconodont crown tissue and vertebrate enamel must be rejected as these tissues have evolved independently and convergently. Thus, the precise ontogenetic, structural and topological similarities between conodont elements and vertebrate odontodes appear to be a remarkable instance of convergence. The last common ancestor of conodonts and jawed vertebrates probably lacked mineralized skeletal tissues. The hypothesis that teeth evolved before jaws and the inside-out hypothesis of dental evolution must be rejected; teeth seem to have evolved through the extension of odontogenic competence from the external dermis to internal epithelium soon after the origin of jaws.

Fossilized embryos are widespread but the record is temporally and taxonomically biased.

We report new discoveries of embryos and egg capsules from the Lower Cambrian of Siberia, Middle Cambrian of Australia and Lower Ordovician of North America. Together with existing records, embryos have now been recorded from four of the seven continents. However, the new discoveries highlight secular and systematic biases in the fossil record of embryonic stages. The temporal window within which the embryos and egg capsules are found is of relatively short duration; it ends in the Early Ordovician and is roughly coincident with that of typical "Orsten"-type faunas. The reduced occurrence of such fossils has been attributed to reducing levels of phosphate in marine waters during the early Paleozoic, but may also be owing to the increasing depth of sediment mixing by infaunal metazoans. Furthermore, most records younger than the earliest Cambrian are of a single kind-large eggs and embryos of the priapulid-like scalidophoran Markuelia. We explore alternative explanations for the low taxonomic diversity of embryos recovered thus far, including sampling, size, anatomy, ecology, and environment, concluding that the preponderance of Markuelia embryos is due to its precocious development of cuticle at an embryonic stage, predisposing it to preservation through action as a substrate on which microbially mediated precipitation of authigenic calcium phosphate may occur. The fossil record of embryos may be limited to a late Neoproterozoic to early Ordovician snapshot that is subject to dramatic systematic bias. Together, these biases must be considered seriously in attempts to use the fossil record to arbitrate between hypotheses of developmental and life history evolution implicated in the origin of metazoan clades.