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ABSTRACT AimThe aim of the current study is to conduct a comprehensive phylogenetic analysis of the genusArbaciato elucidate the evolution and phylogenetic relationships among all extant species and reevaluate the presence of geographic structure within species that have wide, fragmented distributions. LocationSpecimens ofArbaciawere collected from 34 localities spanning the Atlantic and Pacific Oceans, and the Mediterranean Sea. MethodsWe obtained sequences from three mitochondrial markers (COI, 16S and the control region and adjacent tRNAs) and two nuclear markers (28S and 18S; the latter ultimately excluded from the final analyses). Phylogenetic trees were constructed using maximum likelihood and Bayesian inference approaches. A time‐calibrated phylogenetic tree was inferred using a relaxed Bayesian molecular clock and three fossil calibration points. ResultsOur analysis supports the monophyly of the genusArbacia, including the speciesArbacia nigra(previously assigned to the monotypic genusTetrapygus). The new phylogenetic topology suggests an alternative biogeographic scenario of initial divergence between Atlantic and Pacific subclades occurring approximately 9 million years ago. The dispersal and subsequent diversification of the Pacific subclade to the southeast Pacific coincides with the onset of glacial and interglacial cycles in Patagonia. In the Atlantic subclade, the split betweenA. punctulataandA. lixulaoccurred 3.01–6.30 (median 3.74 million years ago), possibly associated with the strengthening of the Gulf Stream current connecting the western and eastern Atlantic. Our study also reveals significant genetic and phylogeographic structures within both Atlantic species, indicating ongoing differentiation processes between populations. Main ConclusionOur study provides valuable insights into the evolutionary history and biogeography of the genusArbaciaand highlights the complex interplay between historical climate changes and oceanic currents in shaping the distribution and diversification of echinoids in the Atlantic and Pacific Oceans.more » « lessFree, publicly-accessible full text available March 1, 2026
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Sixty-five species and nine indeterminate taxa of Florida Paleogene echinoids are discussed, and their geographic and stratigraphic distributions provided. These include 49 species documented from the Eocene and 16 from the Oligocene. Ten new species are described: Prionocidaris robertsi n. sp., Rhyncholampas mariannaensis n. sp., Rhyncholampas bao n. sp., Weisbordella inglisensis n. sp., Weisbordella libum n. sp., Durhamella tetrapora n. sp., and Brissus jonesi n. sp. from the Eocene; and Plagiobrissus cassadyi n. sp., Eupatagus dumonti n. sp., and Schizaster carlsoni n. sp. from the Oligocene. We reconsidered subjective junior synonyms of all species and resurrect Neolaganum archerensis, Echinocyamus macneili, and Eupatagus mooreanus. Furthermore, we updated the taxonomy for all included species and their known distributions and provide emended diagnoses for the genera and species of Florida Neolaganidae. In addition, we herein report the occurrence of Porpitella micra in Cretaceous strata of the subsurface of Florida. This remarkable finding makes P. micra the earliest known of all the scutelloids. Echinoids within the Ocala Limestone are placed in five echinoid biozones, which are defined within, these include the Oligopygus phelani, Oligopygus haldemani, Oligopygus wetherbyi, Wythella eldridgei, and Haimea brooksi Zones. This document complements the Neogene (including the Quaternary) fossil echinoid fauna of Florida we published in 2020 and represents a compilation of the known Florida Paleogene echinoid record. The region is currently known to have the most speciose and diverse assemblage of Paleogene echinoids in the United States.more » « lessFree, publicly-accessible full text available February 17, 2026
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The methane seeps on the Pacific margin of Costa Rica support extensive animal diversity and offer insights into deep-sea biogeography. During five expeditions between 2009 and 2019, we conducted intensive faunal sampling via 63 submersible dives to 11 localities at depths of 300–3600 m. Based on these expeditions and published literature, we compiled voucher specimens, images, and 274 newly published DNA sequences to present a taxonomic inventory of macrofaunal and megafaunal diversity with a focus on invertebrates. In total 488 morphospecies were identified, representing the highest number of distinct morphospecies published from a single seep or vent region to date. Of these, 131 are described species, at least 58 are undescribed species, and the remainder include some degree of taxonomic uncertainty, likely representing additional undescribed species. Of the described species, 38 are known only from the Costa Rica seeps and their vicinity. Fifteen range extensions are also reported for species known from Mexico, the Galápagos seamounts, Chile, and the western Pacific; as well as 16 new depth records and three new seep records for species known to occur at vents or organic falls. No single evolutionary narrative explains the patterns of biodiversity at these seeps, as even morphologically indistinguishable species can show different biogeographic affinities, biogeographic ranges, or depth ranges. The value of careful molecular taxonomy and comprehensive specimen-based regional inventories is emphasized for biodiversity research and monitoring.more » « less
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Davoult, D (Ed.)Abstract: The two main approaches in interpreting the significance of non-radial fossil echinoderms (carpoids) have consistently produced the same two distinct clusters of results for over a century of investigation. Studies based on conceptual models imbued with Haeckelian precepts consider bilaterally symmetric or asymmetric morphologies of carpoids plesiomorphic for the phylum. These approaches do not find a place for carpoids within an existing phylogenetic framework for the phylum because it is assumed that they primitively lack pentaradiality. Emphasis on differences between these taxa and pentaradial echinoderms forces them outside of and downward from more crownward clades. It is crucial to examine the data supporting these supposed differences. Is it worth considering non-pentaradial echinoderms as members of a well-known group of echinoderms, the blastozoans, which already includes many secondarily-derived, non-pentaradial members? Followers of an empirical model think so, presenting an integration of paleontology, embryology, body wall homology, and image analysis that derives echinoderms from a bilaterian, archimeric larva, not bilateral adults. Unprecedented modification of a single mesocoel (hydrocoel) initiated the pentaradial adult echinoderm, most parsimoniously with five primary lobes in stem forms of each major clade within the phylum. The unique water vascular system led to rearrangement of adult axes that literally have no parallel with those of any other invertebrate, representing an iconic synapomorphy for the Echinodermata. There are few, if any, developmental or stratigraphic data defending carpoids as 'bilateral precursors'. Their free appendage is now shown to be an ambulacrum, undermining any supposition of a 'head', 'tail', or 'gill slits'. Pentaradiality is plesiomorphic for the phylum, obviating the requirement for a triradial intermediate (helicoplacoids) between carpoids and pentaradial forms. Carpoids, a subset of blastozoans, exploited motility as a feeding mode, leading to extraordinary adaptations that belie their interpretation as ancestral echinoderms.more » « less
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The Brissidae is a family of heart urchins that includes shallow‑water, infaunal species, some of which are rarely observed. One species,Rhynobrissus cuneusCooke, 1957, has only been recorded from the western Atlantic off the coast of North Carolina, USA and in the Gulf of Mexico off the coast of Veracruz, Mexico. This work identifies new records from the coast of Florida, USA, increasing our knowledge of this species’ geographic distribution.more » « less
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Davoult, D (Ed.)Abstract: Echinoderms are so highly derived compared with other deuterostomes, including their sister group, hemichordates, that comparisons of body plans are sometimes accompanied by points of view enjoying varying levels of morphological, paleontological, and especially, embryological support. No echinoderm taxon has been the subject of more contentious debate than the carpoids, a disparate assemblage of non-pentaradial, flattened echinoderms that includes the Cincta, Ctenocystoidea, Soluta, and Stylophora. Because of their unusual morphologies, the phylogenetic position and significance of carpoids concerning the origins of the Echinodermata are still being evaluated. A detailed review of carpoid research over the past century and a half reveals that the debate largely results from methodological issues employing two basic, but very different models. Conceptual models, usually imbued with Haeckelian principles, consider the absence of a single character (pentaradial symmetry) as a recapitulation of the pre-metamorphic larval stage of echinoderms, forcing unusual taxa that also lack pentaradiality down the phylum's phylogenetic tree. Such scenarios assume that first echinoderms had a bilaterian-type anterior-posterior axis. Empirical models rely on comparison of non-pentaradial early forms with a wide array of data obtained from extant and fossil echinoderms. These data support a view in which larval morphologies of echinoderms are not represented in the fossil record of echinoderms, and that pentaradial symmetry was secondarily lost in carpoids, just as it was in many other coeval types of echinoderms.more » « less
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Davoult, D (Ed.)Abstract: Several levels of the Lorraine Group (Upper Ordovician) in upstate New York (USA) have yielded low-diversity, exceptionally preserved, pyritized invertebrate assemblages dominated by the trilobite Triarthrus eatoni. Sedimentological and taphonomic features suggest dysoxic bottom-water conditions, with limited transport and rapid burial by distal turbidites. Echinoderms are extremely rare in these strata. Here we report, for the first time, the occurrence of the anomalocystitid mitrate Enoploura popei in the Konservat-Lagerstätte of Beecher's Trilobite Bed. A pyritized specimen of this stylophoran was CT-scanned and three-dimensionally reconstructed. The mitrate is laterally compressed, but its 3D-rendering provided several insights into its internal anatomy and taphonomy. The recurved position of the single feeding appendage (aulacophore) is consistent with ligament-induced, post mortem contraction. This posture and the collapse of one lateral series of cover plates indicate that the individual was probably not buried alive. Nevertheless, a portion of the distal aulacophore shows clear evidence of exceptionally preserved soft parts (ambulacral system) in between two sets of slightly open cover plates and the underlying ossicles. One of the most intriguing features of this specimen is its close association with a sinuous, elongated, pyritized trace fossil, which enters the stylophoran through the mouth and disappears into the proximal aulacophore. In marked contrast with other skeletal parts of the specimen (theca and distal part of the aulacophore), the proximal rings of the aulacophore are heavily disrupted and disarticulated. Proximal rings are usually decay-resistant skeletal regions in stylophorans. Therefore, close association of this disrupted region with a trace fossil penetrating it suggests the action of an unknown infaunal scavenger. Location of this trace suggests targeting during early decay of the large muscular proximal aulacophore.more » « less
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The echinoid genus Tetrapygus was initially described by L. Agassiz (1841) based on a single species, Tetrapygus niger Molina, 1782. Since the extensive work conducted by Mortensen (1935), Tetrapygus has received limited taxonomic attention over the past century. Recent discoveries of new fossil species of Arbacia Gray, 1835 from the upper Pliocene of northern Chile revealed striking morphological similarities between the two distinct Arbaciidae genera Arbacia and Tetrapygus. These findings compelled new investigations to evaluate the taxonomic status of these genera. Based on molecular mitochondrial (COI), nuclear (28S), and morphological evidence, Tetrapygus niger is here recovered as the sister species to Arbacia dufresnii, both species forming a clade within the phylogeny of South American species of Arbacia. Consequently, the diagnosis and description of Tetrapygus niger are here revised, and the species is reattributed to Arbacia, as previously proposed by A. Agassiz in Agassiz & Desor (1846) under the species name Arbacia nigra. An emended diagnosis of Arbacia is also proposed in light of these new findings.more » « less
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Abstract Evidence from the earliest-known crinoids (Tremadocian, Early Ordovician), called protocrinoids, is used to hypothesize initial steps by which elements of the calyx evolved. Protocrinoid calyces are composed of extraxial primary and surrounding secondary plates (both of which have epispires along their sutures) that are unlike those of more crownward fossil and extant crinoids in which equivalent calycinal plating is strongly organized. These reductions inspired several schemes by which to name the plates in these calyces. However, the primary-secondary systems seen in protocrinoids first appeared among Cambrian stem radial echinoderms, with primaries representing centers around which secondaries were sequentially added during ontogeny. Therefore, the protocrinoid calyx represents an intermediate condition between earliest echinoderms and crownward crinoids. Position and ontogeny indicate certain primaries remained as loss of secondaries occurred, resulting in abutting of primaries into the conjoined alternating circlets characteristic of crinoids. This transformative event included suppression of secondary plating and modification or, more commonly, elimination of respiratory structures. These data indicate subradial calyx plate terminology does not correspond with most common usage, but rather, supports an alternative redefinition of these traditional expressions. Extension and adoral growth of fixed rays during calyx ontogeny preceded conjoined primaries in earliest crinoids. Restriction with modification or elimination of calyx respiratory structures also accompanied this modification. Phylogenetic analyses strongly support crinoid origination from early pentaradiate echinoderms, separate from blastozoans. Accordingly, all Tremadocian crinoids express a distinctive aggregate of plesiomorphic and apomorphic commonalities; all branch early within the crinoid clade, separate from traditional subclass-level clades. Nevertheless, each taxon within this assemblage expresses at least one diagnostic apomorphy of camerate, cladid, or disparid clades.more » « less
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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.more » « less
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