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Echinoids are key components of modern marine ecosystems. Despite a remarkable fossil record, the emergence of their crown group is documented by few specimens of unclear affinities, rendering their early history uncertain. The origin of sand dollars, one of its most distinctive clades, is also unclear due to an unstable phylogenetic context. We employ 18 novel genomes and transcriptomes to build a phylogenomic dataset with a near-complete sampling of major lineages. With it, we revise the phylogeny and divergence times of echinoids, and place their history within the broader context of echinoderm evolution. We also introduce the concept of a chronospace - a multidimensional representation of node ages - and use it to explore methodological decisions involved in time calibrating phylogenies. We find the choice of clock model to have the strongest impact on divergence times, while the use of site-heterogeneous models and alternative node prior distributions show minimal effects. The choice of loci has an intermediate impact, affecting mostly deep Paleozoic nodes, for which clock-like genes recover dates more congruent with fossil evidence. Our results reveal that crown group echinoids originated in the Permian and diversified rapidly in the Triassic, despite the relative lack of fossil evidence for this early diversification. We also clarify the relationships between sand dollars and their close relatives and confidently date their origins to the Cretaceous, implying ghost ranges spanning approximately 50 million years, a remarkable discrepancy with their rich fossil record.

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Introduction: Scutelliforms were diverse and widespread in shallow marine environments during Neogene times in South America. Nevertheless, they have almost never been used as biostratigraphic tools. Objective: To provide a refined stratigraphic frame useful for calibrating temporal dimensions of scutelliform diversity from Argentina and Uruguay and its correlation with the molluscan assemblages previously proposed. Methods: A detailed survey of their geographic and stratigraphic provenance was carried out. We revised both the bibliography and collections (institutional and from our own field work). Results: The group is represented by 14 species belonging to six genera, and four assemblages were identified. Numerical dates of the Neogene marine rocks obtained recently allowed their placement in a chronological scheme: "Iheringiella" sp. A is restricted to the late Oligocene, the genera Camachoaster and "Eoscutella" and the species Monophoraster telfordi to the early Miocene, Abertella gualichensis and Abertella miskellyi to the middle Miocene, and Monophoraster duboisi, Amplaster coloniensis and Amplaster ellipticus to the late Miocene. Non-lunulate scutelliforms are not restricted to the late Oligocene as previously supposed. The oldest occurrence of the genus Monophoraster corresponds to the early Miocene, and along with Iheringiella are long-living taxa that embrace the 25.3 Ma-18.1 Ma (Iheringiella patagonensis) and approximately 15 Ma-6.48 Ma (Monophoraster darwini) intervals. The presence of Iheringiella in the early Miocene of northeastern Patagonia is corroborated, reaching there its northernmost distribution. Monophoraster darwini has a temporal range from the late Miocene (where it was previously thought to be restricted) back to the middle Miocene, since this is the species yielded in the well-known and discussed "Monophoraster and Venericor Beds". Conclusions: The Paleogene-Neogene scutelliforms of Argentina and Uruguay range from the late Oligocene to the late Miocene. There is a good correspondence among the numerical ages, molluscan biozones and scutelliform assemblages.

Introducción: Realizamos una síntesis actualizada de la composición taxonómica de las asociaciones de escutélidos del Oligoceno tardío-Mioceno tardío de Argentina y Uruguay. Objetivo: Ubicar los escutélidos en un marco estratigráfico refinado, a los efectos de observar su diversidad a lo largo del tiempo y la correlación de sus ensambles con los ya conocidos de moluscos. Métodos: Revisamos en detalle la procedencia geográfica y estratigráfica de los ejemplares y su asignación taxonómica, basándonos tanto en la bibliografía como en colecciones institucionales y producto de nuestras campañas de colecta. Resultados: El grupo está representado por 14 especies agrupadas en seis géneros, siendo identificadas en cuatro ensambles. Las edades numéricas recientemente obtenidas permitieron su ubicación en un esquema cronológico: "Iheringiella" sp. A se restringe al Oligoceno tardío, Camachoaster y "Eoscutella", y Monophoraster telfordi al Mioceno temprano, Abertella gualichensis y Abertella miskellyi al Mioceno medio, y Monophoraster duboisi, Amplaster coloniensis y Amplaster ellipticus al Mioceno tardío. Los escutélidos sin lúnula no están restringidos al Oligoceno tardío como se suponía. Monophoraster aparece en el Mioceno temprano y junto con Iheringiella abarcan largos períodos de tiempo, encontrándoselos entre los 25.3 Ma-18.1 Ma (Iheringiella patagonensis) y aproximadamente los 15 Ma-6.48 Ma (Monophoraster darwini). Se corrobora la presencia de Iheringiella en el Mioceno temprano del noreste de Patagonia, donde alcanza su punto más septentrional. Monophoraster darwini se encuentra desde el Mioceno medio hasta el Mioceno tardío (lapso para el cual previamente se consideraba restringido), ya que se trata de la especie presente en los bien conocidos "estratos con Monophoraster y Venericor". Conclusiones: Los escutélidos del Paleógeno-Neógeno de Argentina y Uruguay se encuentran desde el Oligoceno tardío hasta el Mioceno tardío. Hay una buena correspondencia entre las edades numéricas, las biozonas de moluscos y los ensambles de escutélidos.

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BACKGROUND: Echinoidea is a clade of marine animals including sea urchins, heart urchins, sand dollars and sea biscuits. Found in benthic habitats across all latitudes, echinoids are key components of marine communities such as coral reefs and kelp forests. A little over 1000 species inhabit the oceans today, a diversity that traces its roots back at least to the Permian. Although much effort has been devoted to elucidating the echinoid tree of life using a variety of morphological data, molecular attempts have relied on only a handful of genes. Both of these approaches have had limited success at resolving the deepest nodes of the tree, and their disagreement over the positions of a number of clades remains unresolved. RESULTS: We performed de novo sequencing and assembly of 17 transcriptomes to complement available genomic resources of sea urchins and produce the first phylogenomic analysis of the clade. Multiple methods of probabilistic inference recovered identical topologies, with virtually all nodes showing maximum support. In contrast, the coalescent-based method ASTRAL-II resolved one node differently, a result apparently driven by gene tree error induced by evolutionary rate heterogeneity. Regardless of the method employed, our phylogenetic structure deviates from the currently accepted classification of echinoids, with neither Acroechinoidea (all euechinoids except echinothurioids), nor Clypeasteroida (sand dollars and sea biscuits) being monophyletic as currently defined. We show that phylogenetic signal for novel resolutions of these lineages is strong and distributed throughout the genome, and fail to recover systematic biases as drivers of our results. CONCLUSIONS: Our investigation substantially augments the molecular resources available for sea urchins, providing the first transcriptomes for many of its main lineages. Using this expanded genomic dataset, we resolve the position of several clades in agreement with early molecular analyses but in disagreement with morphological data. Our efforts settle multiple phylogenetic uncertainties, including the position of the enigmatic deep-sea echinothurioids and the identity of the sister clade to sand dollars. We offer a detailed assessment of evolutionary scenarios that could reconcile our findings with morphological evidence, opening up new lines of research into the development and evolutionary history of this ancient clade.

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Material for all seven South American species of late Oligocene and Miocene scutelliform, non-lunulate sand dollars (Echinoidea: Clypeasteroida) was examined, resulting in revision of several taxa, and description of a new family, Placatenellidae n. fam., to house Placatenella n. gen. and Camachoaster n. gen. Known only from the state of Pará, Brazil, and previously described as a member of Abertella Durham, 1953, A. complanata Brito, 1981 is redescribed on the basis of analysis of the holotype and subsequently described material from the same region. A. complanata is removed from synonymy with A. pirabensis (Marchesini Santos, 1958), and transferred as the type species of Placatenella n. gen. A new taxon, Camachoaster n. gen., is described to hold C. maquedensis n. sp. The Abertellidae Durham, 1955 is revised and the concept of the genus Abertella restricted to include only those forms with a posterior notch and all interambulacra discontinuous, including A. pirabensis, which is redescribed from the only known specimen, confirming that this species lacks any trace of an anal lunule but possesses features fully congruent with its placement in Abertella. Iheringiellidae n. fam. is established to house the common but taxonomically challenging taxon, Iheringiella Berg, 1898. New data on the occurrence and location of the types of Iheringiella are discussed, and comments on the status of Eoscutella mirandae Parma, 1985 provided. A tabular key to all seven South American non-lunulates is given, along with discussion of the relationship between the posterior notch and the anal lunule.

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