Crossed Tracks: Mesolimulus, Archaeopteryx, and the Nature of Fossils

Organisms leave a variety of traces in the fossil record. Among these traces, vertebrate and invertebrate paleontologists conventionally recognize a distinction between the remains of an organism’s phenotype (body fossils) and the remains of an organism’s life activities (trace fossils). The same convention recognizes body fossils as biological structures and trace fossils as geological objects. This convention explains some curious practices in the classification, as with the distinction between taxa for trace fossils and for tracemakers. I consider the distinction between “parallel taxonomies,” or parataxonomies, which privileges some kinds of fossil taxa as “natural” and others as “artificial.” The motivations for and consequences of this practice are inconsistent. By comparison, I examine an alternative system of classification used by paleobotanists that regards all fossil taxa as “artificially” split. While this system has the potential to inflate the number of taxa with which paleontologists work, the system offers greater consistency than conventional practices. Weighing the strengths and weaknesses of each system, I recommend that paleontologists should adopt the paleobotanical system more broadly

Biomechanical evolution of solid bones in large animals: a microanatomical investigation

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Using Models to Correct Data: Paleodiversity and the Fossil Record

Despite an enormous philosophical literature on models in science, surprisingly little has been written about data models and how they are constructed. In this paper, I examine the case of how paleodiversity data models are constructed from the fossil data. In particular, I show how paleontologists are using various model-based techniques to correct the data. Drawing on this research, I argue for the following related theses: First, the 'purity' of a data model is not a measure of its epistemic reliability. Instead it is the fidelity of the data that matters. Second, the fidelity of a data model in capturing the signal of interest is a matter of degree. Third, the fidelity of a data model can be improved 'vicariously', such as through the use of post hoc model-based correction techniques. And, fourth, data models, like theoretical models, should be assessed as adequate (or inadequate) for particular purposes

Declining Volatility, a General Property of Disparate Systems: From Fossils, to Stocks, to the Stars

There may be structural principles pertaining to the general behavior of systems that lead to similarities in a variety of different contexts. Classic examples include the descriptive power of fractals, the importance of surface area to volume constraints, the universality of entropy in systems, and mathematical rules of growth and form. Documenting such overarching principles may represent a rejoinder to the Neodarwinian synthesis that emphasizes adaptation and competition. Instead, these principles could indicate the importance of constraint and structure on form and evolution. Here we document a potential example of a phenomenon suggesting congruent behavior of very different systems. We focus on the notion that universally there has been a tendency for more volatile entities to disappear from systems such that the net volatility in these systems tends to decline. We specifically focus on origination and extinction rates in the marine animal fossil record, the performance of stocks in the stock market, and the characters of stars and stellar systems. We consider the evidence that each is experiencing declining volatility, and also consider the broader significance of this.Comment: Accepted for publication in Palaeontology. 13 pages, 3 figure

Using models to correct data: paleodiversity and the fossil record

Despite an enormous philosophical literature on models in science, surprisingly little has been written about data models and how they are constructed. In this paper, I examine the case of how paleodiversity data models are constructed from the fossil data. In particular, I show how paleontologists are using various model-based techniques to correct the data. Drawing on this research, I argue for the following related theses: first, the ‘purity’ of a data model is not a measure of its epistemic reliability. Instead it is the fidelity of the data that matters. Second, the fidelity of a data model in capturing the signal of interest is a matter of degree. Third, the fidelity of a data model can be improved ‘vicariously’, such as through the use of post hoc model-based correction techniques. And, fourth, data models, like theoretical models, should be assessed as adequate for particular purposes

Integrating genomics with the fossil record to explore the evolutionary history of Echinoidea

Echinoidea constitutes one of five major clades of living echinoderms, marine animals uniquely characterized by a pentaradial symmetry. Approximately 1,000 living and 10,000 extinct species have been described, including many commonly known as sea urchins, heart urchins and sand dollars. Today, echinoids are ubiquitous in benthic marine environments, where they strongly affect the functioning of biodiverse communities such as coral reefs and kelp forests. Given the quality of their fossil record, their remarkable morphological complexity and our thorough understanding of their development, echinoids provide unparalleled opportunities to explore evolutionary questions in deep-time, providing access to the developmental and morphological underpinnings of evolutionary innovation. These questions cannot be addressed without first resolving the phylogenetic relationships among living and extinct lineages. The goal of this dissertation is to advance our understanding of echinoid relationships and evolutionary history, as well as to explore more broadly the integration of phylogenomic, morphological and paleontological data in phylogenetic reconstruction and macroevolutionary inference.In Chapter 1, I report the results of the first phylogenomic analysis of echinoids based on the sequencing of 17 novel echinoid transcriptomes. Phylogenetic analyses of this data resolve the position of several clades—including the sand dollars—in disagreement with traditional morphological hypotheses. I demonstrate the presence of a strong phylogenetic signal for these novel resolutions, and explore scenarios to reconcile these findings with morphological evidence. In Chapter 7, I extend this approach with a more thorough taxon sampling, resulting in a robust topology with a near-complete sampling of major echinoid lineages. This effort reveals that apatopygids, a clade of three species with previously unclear affinities, represent the only living descendants of a once diverse Mesozoic clade. I also perform a thorough time calibration analysis, quantifying the relative effects of choosing among alternative models of molecular evolution, gene samples and clock priors. I introduce the concept of a chronospace and use it to reveal that only the last among the aforementioned choices affects significantly our understanding of echinoid diversification. Molecular clocks unambiguously support late Permian and late Cretaceous origins for crown group echinoids and sand dollars, respectively, implying long ghost ranges for both. Fossils have been shown to improve the accuracy of phylogenetic comparative methods, warranting their inclusion alongside extant terminals when exploring evolutionary processes across deep timescales. However, their impact on topological inference remains controversial. I explore this topic in Chapter 3 with the use of simulations, which show that morphological phylogenies are more accurate when fossil taxa are incorporated. I also show that tip-dated Bayesian inference, which takes stratigraphic information from fossils into account, outperforms uncalibrated methods. This approach is complemented in Chapter 2 with the analysis of empirical datasets, confirming that incorporating fossils reshapes phylogenies in a manner that is entirely distinct from increased sampling of extant taxa, a result largely attributable to the occurrence of distinctive character combinations among fossils. Even though phylogenomic and paleontological data are complementary resources for unraveling the relationships and divergence times of lineages, few studies have attempted to fully integrate them. Chapter 4 revisits the phylogeny of crown group Echinoidea using a total-evidence dating approach combining phylogenomic, morphological and stratigraphic information. To this end, I develop a method (genesortR) for subsampling molecular datasets that selects loci with high phylogenetic signal and low systematic biases. The results demonstrate that combining different data sources increases topological accuracy and helps resolve phylogenetic conflicts. Notably, I present a new hypothesis for the origin and early morphological evolution of the sand dollars and close allies. In Chapter 6, I compare the behavior of genesortR against alternative subsampling strategies across a sample of phylogenomic matrices. I find this method to systematically outperform random loci selection, unlike commonly-used approaches that target specific evolutionary rates or minimize sources of systematic error. I conclude that these methods should not be used indiscriminately, and that multivariate methods of phylogenomic subsampling should be favored. Finally, in Chapter 5, I explore the macroevolutionary dynamics of echinoid body size across 270 million years using data for more than 5,000 specimens in a phylogenetically explicit context. I also develop a method (extendedSurface) for parameterizing adaptive landscapes that overcomes issues with existing approaches and finds better fitting models. While echinoid body size has been largely constrained to evolve within a single adaptive peak, the disparity of the clade was generated by regime shifts driving the repeated evolution of miniaturized and gigantic forms. Most innovations occurred during the latter half of the Mesozoic, and were followed by a drastic slowdown in the aftermath of the Cretaceous-Paleogene mass extinction

Macroevolutionary Patterns In The Evolutionary Radiation Of Archosaurs (Tetrapoda: Diapsida)

The rise of archosaurs during the Triassic and Early Jurassic has been treated as a classic example of an evolutionary radiation in the fossil record. This paper reviews published studies and provides new data on archosaur lineage origination, diversity and lineage evolution, morphological disparity, rates of morphological character change, and faunal abundance during the Triassic–Early Jurassic. The fundamental archosaur lineages originated early in the Triassic, in concert with the highest rates of character change. Disparity and diversity peaked later, during the Norian, but the most significant increase in disparity occurred before maximum diversity. Archosaurs were rare components of Early–Middle Triassic faunas, but were more abundant in the Late Triassic and pre-eminent globally by the Early Jurassic. The archosaur radiation was a drawn-out event and major components such as diversity and abundance were discordant from each other. Crurotarsans (crocodile-line archosaurs) were more disparate, diverse, and abundant than avemetatarsalians (bird-line archosaurs, including dinosaurs) during the Late Triassic, but these roles were reversed in the Early Jurassic. There is no strong evidence that dinosaurs outcompeted or gradually eclipsed crurotarsans during the Late Triassic. Instead, crurotarsan diversity decreased precipitously by the end-Triassic extinction, which helped usher in the age of dinosaurian dominance

Molecular evidence that phoronids are a subtaxon of brachiopods (Brachiopoda: Phoronata) and that genetic divergence of metazoan phyla began long before the early Cambrian

Concatenated SSU (18S) and partial LSU (28S) sequences (~2 kb) from 12 ingroup taxa, comprising 2 phoronids, 2 members of each of the craniid, discinid, and lingulid inarticulate brachiopod lineages, and 4 rhynchonellate, articulate brachiopods (2 rhynchonellides, 1 terebratulide and 1 terebratellide) were aligned with homologous sequences from 6 protostome, deuterostome and sponge outgroups (3964 sites). Regions of potentially ambiguous alignment were removed, and the resulting data (3275 sites, of which 377 were parsimony-informative and 635 variable) were analysed by parsimony, and by maximum and Bayesian likelihood using objectively selected models. There was no base composition heterogeneity. Relative rate tests led to the exclusion (from most analyses) of the more distant outgroups, with retention of the closer pectinid and polyplacophoran (chiton). Parsimony and likelihood bootstrap and Bayesian clade support values were generally high, but only likelihood analyses recovered all brachiopod indicator clades designated a priori. All analyses confirmed the monophyly of (brachiopods+phoronids) and identified phoronids as the sister-group of the three inarticulate brachiopod lineages. Consequently, a revised Linnean classification is proposed in which the subphylum Linguliformea comprises three classes: Lingulata, ‘Phoronata’ (the phoronids), and ‘Craniata’ (the current subphylum Craniiformea). Divergence times of all nodes were estimated by regression from node depths in non-parametrically rate-smoothed and other chronograms, calibrated against palaeontological data, with probable errors not less than 50 My. Only three predicted brachiopod divergence times disagree with palaeontological ages by more than the probable error, and a reasonable explanation exists for at least two. Pruning long-branched ingroups made scant difference to predicted divergence time estimates. The palaeontological age calibration and the existence of Lower Cambrian fossils of both main brachiopod clades together indicate that initial genetic divergence between brachiopod and molluscan (chiton) lineages occurred well before the Lower Cambrian, suggesting that much divergence between metazoan phyla took place in the Proterozoic

The evolution of large-bodied theropod dinosaurs during the Mesozoic in Asia

The fossil record of large-bodied, apex carnivorous theropod dinosaurs in Eastern Asia is now among the best understood in the world, thanks to new discoveries and reinterpretations of long-neglected fossils. Asia boasts the most complete record of Middle Jurassic theropods globally, as well as one of the best-studied Late Cretaceous theropod faunas, and new research is helping to fill what was previously a 60-million-year gap in the Early-mid Cretaceous fossil record of large Asian predators. In general, the bio-geographic affinities of large-bodied Asian theropods over time were intimately related to physical geography, and progressively more derived theropod clades evolved large body size and occupied the apex predator niche throughout the Jurassic and Cretaceous. During the Middle Jurassic, largely endemic clades of basal tetanurans were prevalent in Asia, whereas during the Late Jurassic mid Cretaceous more derived “intermediate” tetanuran theropods with cosmopolitan affinities occupied the large predator role, including sinraptorids, spinosauris, and carcharodontosaurians. Finalli, during the final 20 million years of the Cretaceous, more derived, bird-like coelurosaurs attained large body size. Foremost among these were the tyrannosaurids, a radiation of northern (Asian and North American) megapredators whose ascent into the apex predator niche was a delayed event restricted to the Campanian-Masastrichian. As Asia is the focus of intense ongoing dinosaur fieldwork, our undestarnding of large-bodied theropod evolution will continue to be refined with future discoveries.El registro fósil de los dinosaurios carnívoros terópodos de gran talla en el este de Asia es uno de los mejor conocidos del mundo, gracias a nuevos descubrimientos y reinterpretaciones de fósiles que han permanecido pobremente estudiados durante mucho tiempo. Globalmente, Asia comprende el registro fósil mas completo de terópodos del Jurasico Medio, así como una de las faunas finicretácicas mejor estudiadas. Asimismo, las nuevas investigaciones están contribuyendo a completar un hiato de 60 millones de años en el registro fósil de grandes depredadores asiáticos correspondientes al Cretácico inferior-medio. En general las afinidades biogeográficas de los grandes terópodos asiáticos a través del tiempo se hallan íntimamente ligadas a la geografía física. Progresivamente, varios clados derivados de terópodos evolucionaron grandes tallas corporales, ocupando la cima del nicho de depredador durante todo el Jurasico y el Cretácico. Durante el Jurasico Medio prevalecieron clados de tetanuros basales mayormente endémicos, mientras que durante el Jurásico Superior-Cretácico Medio clados más derivados de terópodos tetanuros “intermedios” de afinidades cosmopolitas ocuparon el papel de gran depredador, incluyendo sinraptoridos, espinosauridos y carcharodontosauridos. Finalmente, durante los ultimos 20 millones de anos del Cretacico, coelurosaurios mas derivados con aspecto reminiscente a las aves alcanzaron grandes tallas corporales. Pirmordialmente entre estas formas se hallaban los tiranosauridos, una radiación septentrional (asiáticos y norteamericanos) de megadepredadores cuyo ascenso a la cumbre del nicho de gran depredador se retraso hasta el Campaniense y Maastrichtiense. Mientras Asia continua constituyendo el foco de una intensa actividad paleontológica, nuestros conocientos sobre la evolución de los grandes terópodos continuará refinándose con el estudio de futuros hallazgos