Taxonomic reassessment of Tetrapygus niger (Arbacioida, Echinoidea): molecular and morphological evidence support its placement in Arbacia.

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.

Arbacia ballenensis sp. nov. (Echinoidea, Arbacioida): A new species reveals diversification of the genus in Central America.

A new species of Arbacia (Echinoidea, Arbacioida), A. ballenensis sp. nov., from the Late PlioceneEarly Pleistocene of Baja California Norte, Mxico, is described from a series of 62 specimens ranging in size from 6 to 20 mm test diameter. It differs from all extant species in the genus by its high number of interambulacral plates, numerous small interambulacral tubercles, and the size of the interambulacral tags at the peristomial edge. This new species yields new evidence concerning the diversification of the genus along the Pacific coast of North America where previously, only one extinct species was described, Arbacia abiquaensis from the Oligocene of Oregon. A. ballenensis sp. nov. is closely related to the recently described Arbacia quyllur from the late Miocene of Chile. Study of the new species in the context of events such as the mid-Pleistocene Transition helps shed light on patterns of extinction and speciation within the genus.

Phylogenomic analyses of echinoid diversification prompt a re-evaluation of their fossil record.

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.

A new species and comparative morphology of Philippine sea biscuits (Echinoidea: Clypeaster/).

A new species of clypeasterid sea biscuit, Clypeaster brigitteae n. sp., is described from material collected in the Philippines at depths between 100 and 200 m. The new taxon increases the number of Clypeaster species recorded from the Philippines to nine, representing nearly a quarter of the world's diversity of the genus. Other Philippine species include: C. annandalei Koehler, 1922; C. fervens Koehler, 1922; C. humilis (Leske, 1778); C. japonicus Döderlein, 1885; C. latissimus (Lamarck, 1816); C. pateriformis Mortensen, 1948; C. reticulatus (Linnaeus, 1758); and C. virescens Döderlein, 1885. Using type material where available, each of these species is compared and contrasted with C. brigitteae n. sp. in tables consisting of new data derived from general test shape and size, petal structure, food grooves, plate architecture, internal structure, and morphology of spines, pedicellariae, and tube feet.

An annotated species list of regular echinoids from Sri Lanka with notes on some rarely seen temnopleurids.

The first recorded regular echinoid species from Sri Lanka, "Salmacis virgulatus" (now known as S. virgulata L. Agassiz in L. Agassiz Desor, 1846), was listed by Agassiz Desor (1846). Knowledge of Sri Lankan regular echinoids continued to advance until the end of the 19th century. However, there is a gap in knowledge between the mid-20th and the beginning of the 21st centuries due to a lack of systematic studies, with the exception of two checklists published by the IUCN Red List in 2006 and 2012. In the present study, we compiled a species list combining published data and new data based on fieldwork between 2013 and 2015. Echinoids were sampled by snorkelling, diving, examination of fisheries bycatch, and collection of tests from beaches. The updated species list presented in this study includes 39 regular echinoids belonging to 28 genera, nine families, and five orders. Phyllacanthus imperialis (Lamarck, 1816) and Temnopleurus toreumaticus (Leske, 1778), which were not recorded during the last 90 years, were confirmed to still occur in Sri Lanka by the present study. We develop an updated species list of regular echinoids to form a basis for future systematic work. The study recommends further investigation to evaluate the status of deep-sea species and additional field work off the northern and eastern coasts of Sri Lanka.

First record of the heart urchin Metalia persica (Mortensen, 1940) (Spatangoida: Brissidae) from the Chennai coast, India.

The irregular urchin, Metalia persica (Mortensen, 1940), a heart urchin of the family Brissidae, is reported for the first time from the southeastern coast of India. A single specimen was collected from fishing by-catch at Chinna Neelankarai (12º 56' 29.7" N, 80º 15' 36.6" E),  Chennai on April 5, 2018. Metalia persica was previously recorded only from the Persian Gulf (Arabian Gulf). This study records Metalia persica for the first time from Indian waters, in the Bay of Bengal, and elucidates features of the anal fasciole.

Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka.

The earliest information on Sri Lankan echinoid species belonging to the Irregularia dates back to Alexander Agassiz (1872). However, the current knowledge of diversity and distribution of irregular echinoids from Sri Lanka (formerly Ceylon) remains sparse. In addition, there are no recent taxonomic studies or biodiversity surveys for irregular echinoids, and no illustrated field-guides or reference collections are available specifically for Sri Lanka. To address these gaps, left open for more than 100 years since the work of Clark (1915), this study was conducted as an island-wide systematic sampling survey. Over 200 echinoid specimens were collected from 24 localities in Sri Lankan coastal waters by snorkelling and SCUBA diving down to 33 m depth. The collected specimens were identified using existing keys and authenticated with specimens available at the Natural History Museum in Vienna, Austria. The present study records 22 irregular echinoid species belonging to 15 genera and nine families in four orders. Among the identified irregular echinoids, six species, Echinocyamus megapetalus H.L. Clark, 1914, Fibularia ovulum Lamarck, 1816, Fibulariella angulipora Mortensen, 1948, Echinodiscus cf. truncatus L. Agassiz, 1841, Peronella oblonga Mortensen, 1948 and Brissus cf. agassizii Döderlein, 1885, are new records for Sri Lanka. Four unidentified, possibly new species belonging to the genera Fibularia, Jacksonaster and Metalia are reported, but kept in open nomenclature until more material becomes available. At present, the diversity of irregular echinoids from Sri Lanka now stands at 37 species representing 11 families in four orders. A dichotomous key is presented for all Sri Lankan irregular echinoids.

A phylogenomic resolution of the sea urchin tree of life.

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.

Late Oligocene-Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species.

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.

Megafauna of the UKSRL exploration contract area and eastern Clarion-Clipperton Zone in the Pacific Ocean: Echinodermata.

BACKGROUND: There is growing interest in mining polymetallic nodules from the abyssal Clarion-Clipperton Zone (CCZ) in the tropical Pacific Ocean. Despite being the focus of environmental studies for decades, the benthic megafauna of the CCZ remain poorly known. In order to predict and manage the environmental impacts of mining in the CCZ, baseline knowledge of the megafauna is essential. The ABYSSLINE Project has conducted benthic biological baseline surveys in the UK Seabed Resources Ltd polymetallic-nodule exploration contract area (UK-1). Prior to these research cruises in 2013 and 2015, no biological studies had been done in this area of the eastern CCZ. NEW INFORMATION: Using a Remotely Operated Vehicle and Autonomous Underwater Vehicle, the megafauna within the UKSRL exploration contract area (UK-1) and at a site ~250 km east of the UK-1 area were surveyed, allowing us to make the first estimates of megafaunal morphospecies richness from the imagery collected. Here, we present an atlas of the abyssal echinoderm megafauna observed and collected during the ABYSSLINE cruises to the UK-1 polymetallic-nodule exploration contract area in the CCZ. There appear to be at least 62 distinct morphospecies (13 Asteroidea, 5 Crinoidea, 9 Echinoidea, 29 Holothuroidea and 6 Ophiuroidea) identified mostly by imagery but also using molecular barcoding for a limited number of animals that were collected. This atlas will aid the synthesis of megafaunal presence/absence data collected by contractors, scientists and other stakeholders undertaking work in the CCZ, ultimately helping to decipher the biogeography of the megafauna in this threatened habitat.

A review of previous studies on the Sri Lankan echinoid fauna, with an updated species list.

A comprehensive review and analysis of the literature on echinoids from Sri Lankan waters were conducted to compile an annotated list that integrates the existing published data with original data from recent research. According to the published literature, 115 echinoid species and one subspecies have been reported from Sri Lanka to date. However, the current study revealed that only 66 echinoid species and one subspecies belonging to 20 families can be verified to occur in Sri Lankan waters. According to the present analysis, 49 species were excluded from the list due to uncertain records (16) or synonymy (33) with other taxa known from the region. Of the 66 species and one subspecies occurring in Sri Lankan waters, 11 were first described from type material collected from this region. Six of the type specimens are "regular" echinoids and five are Irregularia. Out of these 11, Araeosoma coriaceum indicum has been recorded only from and appears to be endemic to Sri Lankan waters. However, 34 species of Sri Lankan echinoids have not been recorded in the last 90 years. Echinoid species recorded from Sri Lankan waters represent 6.7% of the currently accepted species of extant echinoids and include representatives of 28% of the extant echinoid families. Forty-five percent (45%) of echinoids recorded from the Indian coast (113 species and subspecies) are present in Sri Lankan waters. The current study highlights the need for systematic revision of echinoid records in Sri Lanka through field surveys and reconciliation of discrepancies in the existing literature. Offshore sampling is also needed due to lack of recent information on local deep-sea echinoids.

A new South American Miocene species of 'one-holed' sand dollar (Echinoidea: Clypeasteroida: Monophorasteridae).

A new species of monophorasterid sand dollar, Monophoraster telfordi n. sp., is described from the Early Miocene basal horizons of the Chenque Formation of Patagonia, Santa Cruz Province, in southern Argentina. The new taxon raises the number of known species in the family to six, and represents first unequivocal record of the genus for the Early Miocene of South America. It is therefore also the oldest member of the genus. M. telfordi is characterized by its test width to length ratio, which is much higher than for the other two described species in the genus, but less than that known for the extremely wide members of the sister taxon, Amplaster. M. telfordi is also unusual among monophorasterids in lacking broad continuity between basicoronal and post-basicoronal plates in the oral interambulacra. A key is provided to all the known species of Monophorasteridae.

Phylogenetic re-evaluation of fossil and extant micro-echinoids with revision of Tridium, Cyamidia, and Lenicyamidia (Echinoidea: Clypeasteroida).

Tridium kieri Tandon & Srivastava, 1980, a clypeasteroid micro-echinoid from the Middle Eocene of Kachchh, India, has an apical system with just 3 gonopores. This condition is otherwise almost unknown among clypeasteroids, yet the morphology of Tridium is very similar to that of extant Fibularia, including members of another relatively poorly known genus from the Indian subcontinent and Western Australia, Cyamidia Lambert & Thiéry, 1914. Re-examination of the type and additional material of T. kieri and Cyamidia paucipora Brunnschweiler, 1962, along with specimens identified as C. nummulitica nummulitica (Duncan & Sladen, 1884), allows for redescription of these forms. For the first time, maps of coronal plate architecture of Tridium and Cyamidia are developed, and SEM images of test surface details of the former are provided. Such new sources of data aid determination of their phylogenetic position among a subset of laganiform clypeasteroid taxa. During these analyses, new data were uncovered for two additional enigmatic, fibulariid taxa, Leniechinus herricki Kier, 1968 (Middle Eocene, North America), and Lenicyamidia compta Brunnschweiler, 1962 (Eocene, Western Australia). Both species were added to the overall analysis. These observations lead to the conclusion that Tridium falls within the genus Fibularia Lamarck, 1816, prompting reassignment of T. kieri to Fibularia along with refinement of the diagnosis of the Fibulariidae Gray, 1855. The Western Australian representative of Cyamidia, C. paucipora, was found to represent juveniles of Lenicyamidia compta; it is here synonymized with the latter. The genus Cyamidia thus appears to be restricted to the Indian Subcontinent. 

How Hox genes can shed light on the place of echinoderms among the deuterostomes.

BACKGROUND: The Hox gene cluster ranks among the greatest of biological discoveries of the past 30 years. Morphogenetic patterning genes are remarkable for the systems they regulate during major ontogenetic events, and for their expressions of molecular, temporal, and spatial colinearity. Recent descriptions of exceptions to these colinearities are suggesting deep phylogenetic signal that can be used to explore origins of entire deuterostome phyla. Among the most enigmatic of these deuterostomes in terms of unique body patterning are the echinoderms. However, there remains no overall synthesis of the correlation between this signal and the variations observable in the presence/absence and expression patterns of Hox genes. RESULTS: Recent data from Hox cluster analyses shed light on how the bizarre shift from bilateral larvae to radial adults during echinoderm ontogeny can be accomplished by equally radical modifications within the Hox cluster. In order to explore this more fully, a compilation of observations on the genetic patterns among deuterostomes is integrated with the body patterning trajectories seen across the deuterostome clade. CONCLUSIONS: Synthesis of available data helps to explain morphogenesis along the anterior/posterior axis of echinoderms, delineating the origins and fate of that axis during ontogeny. From this, it is easy to distinguish between 'seriality' along echinoderm rays and true A/P axis phenomena such as colinearity within the somatocoels, and the ontogenetic outcomes of the unique translocation and inversion of the anterior Hox class found within the Echinodermata. An up-to-date summary and integration of the disparate lines of research so far produced on the relationship between Hox genes and pattern formation for all deuterostomes allows for development of a phylogeny and scenario for the evolution of deuterostomes in general, and the Echinodermata in particular.

A new late Cenozoic species of Abertella (Echinoidea: Clypeasteroida) from Patagonia.

A new species of abertellid sand dollar, Abertella miskellyi n. sp., is described from the Miocene Camarones Formation of Patagonia, southern Argentina. The new taxon corroborates the existence of the genus in South America, given that Abertella is most common in the southeastern USA and the eastern coast of Central America. It is characterized by a unique basicoronal circle, in which the interambulacral basicoronal plates are very heterogeneous in size (small in interambulacrum 5, largest in interambulacra 2 and 3). Additionally, it features disjunct oral interambulacra involving two ambulacral plates in some of the interambulacra rather than one, thus being the most disjunct of all known species of Abertella. A key to the species of the genus is provided.

Origin and evolutionary plasticity of the gastric caecum in sea urchins (Echinodermata: Echinoidea).

BACKGROUND: The digestive tract of many metazoan invertebrates is characterized by the presence of caeca or diverticula that serve secretory and/or absorptive functions. With the development of various feeding habits, distinctive digestive organs may be present in certain taxa. This also holds true for sea urchins (Echinodermata: Echinoidea), in which a highly specialized gastric caecum can be found in members of a derived subgroup, the Irregularia (cake urchins, sea biscuits, sand dollars, heart urchins, and related forms). As such a specialized caecum has not been reported from "regular" sea urchin taxa, the aim of this study was to elucidate its evolutionary origin. RESULTS: Using morphological data derived from dissection, magnetic resonance imaging, and extensive literature studies, we compare the digestive tract of 168 echinoid species belonging to 51 extant families. Based on a number of characters such as topography, general morphology, mesenterial suspension, and integration into the haemal system, we homologize the gastric caecum with the more or less pronounced dilation of the anterior stomach that is observed in most "regular" sea urchin taxa. In the Irregularia, a gastric caecum can be found in all taxa except in the Laganina and Scutellina. It is also undeveloped in certain spatangoid species. CONCLUSIONS: According to our findings, the sea urchin gastric caecum most likely constitutes a synapomorphy of the Euechinoidea. Its occurrence in "regular" euechinoids is linked to the presence of an additional festoon of the anterior stomach in ambulacrum III. Both structures, the additional festoon and the gastric caecum, are absent in the sister taxon to the Euechinoidea, the Cidaroida. Since the degree of specialization of the gastric caecum is most pronounced in the predominantly sediment-burrowing irregular taxa, we hypothesize that its evolution is closely linked to the development of more elaborate infaunal lifestyles. We provide a comprehensive study of the origin and evolutionary plasticity of a conspicuous digestive tract structure, the gastric caecum, in a major taxon of the extant invertebrate macrozoobenthos.

Post-Paleozoic crinoid radiation in response to benthic predation preceded the Mesozoic marine revolution.

It has been argued that increases in predation over geological time should result in increases in defensive adaptations in prey taxa. Recent in situ and laboratory observations indicate that cidaroid sea urchins feed on live stalked crinoids, leaving distinct bite marks on their skeletal elements. Similar bite marks on fossil crinoids from Poland strongly suggest that these animals have been subject to echinoid predation since the Triassic. Following their near-demise during the end-Permian extinction, crinoids underwent a major evolutionary radiation during the Middle-Late Triassic that produced distinct morphological and behavioral novelties, particularly motile taxa that contrasted strongly with the predominantly sessile Paleozoic crinoid faunas. We suggest that the appearance and subsequent evolutionary success of motile crinoids were related to benthic predation by post-Paleozoic echinoids with their stronger and more active feeding apparatus and that, in the case of crinoids, the predation-driven Mesozoic marine revolution started earlier than in other groups, perhaps soon after the end-Permian extinction.

Arrays in rays: terminal addition in echinoderms and its correlation with gene expression.

The echinoderms are deuterostomes that superimpose radial symmetry upon bilateral larval morphology. Consequently, they are not the first animals that come to mind when the concepts of segmentation and terminal addition are being discussed. However, it has long been recognized that echinoderms have serial elements along their radii formed in accordance with the ocular plate rule (OPR). The OPR is a special case of terminal growth, forming elements of the ambulacra that define the rays in echinoderms. New elements are added at the terminus of the ray, which may or may not be marked by a calcified element called the terminal plate (the "ocular" of sea urchins). The OPR operates in every echinoderm, from the occasionally bizarre fossils of the Cambrian to the most familiar extant taxa. Using the OPR and other criteria of recognition, echinoderm body wall can be divided into two main regions: extraxial components are associated with the somatocoels, axial components (formed in accordance with the OPR) with the hydrocoel. We compare patterns of development in axial regions of echinoderms with those found in the anterior-posterior axes of the earliest echinoderms as well as other invertebrates. Although axial and extraxial skeletons appear to be composed of the same biomineral matrix, the genes involved in patterning these two skeletal components are likely distinct. During development of the axial skeleton, for instance, the genes engrailed and orthodenticle are expressed in spatial and temporal patterns consistent with the OPR. Other genes such as distal-less seem to demarcate early ontogenetic boundaries between the axial rudiment and the extraxial larval body. There is a complex and pervasive reorganization of gene expression domains to produce the highly divergent morphologies seen in the Echinodermata. We integrate morphological and genetic information, particularly with respect to the origins of radial symmetry in the rudiment, and the concomitant development of the rays.

Extinct and extant sand dollars (Clypeasteroida: Echinoidea) from Uruguay.

We summarize information concerning Recent and Pleistocene-Holocene Uruguayan sand dollars (Mellitidae), as well as Miocene taxa (Monophorasteridae). Recent and Pleistocene-Holocene species (Encope emarginata, Mellita quinquiesperforata, and Leodia sexisperforata) are at their southernmost limits of distribution, with only E. emarginata recorded further south than Uruguay. Lower temperatures to the south, and/or the Rio de la Plata salinity barrier are suggested as controlling factors of these distributions. During the Miocene, the sea temperatures were notably higher than at present, and it was at this time that the extinct genera Monophoraster and Amplaster reached their maximum diversity in Uruguay. The family Monophorasteridae is the basal sister group of the Mellitidae.