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Assembly of publically available next-generation sequence data facilitated the generation of three camarodont echinoid mitogenomes: two for the Green Urchin (Lytechinus variegatus) and one for the Red Urchin (Mesocentrotus franciscanus). The data generated are exploited in a phylogenomic analysis of the superfamily Odontophora, originally proposed for echinoids with tooth supports on the epiphyses of the jaw. The analysis highly supports this taxon and its current subdivision into three families: the Echinometridae, Toxopneustidae, and Strongylocentrotidae. The analysis furthermore implies that historical taxonomic issues between two members of the genus Strongylocentrotus (S. pallidus and S. droebachiensis) may have a genetic basis. The novel mitogenomes for the model species L. variegatus complements the draft genome available for this taxon, one of only three genome-enabled echinoid species. The assembly method applied herein, follows a divide-and-conquer approach that provides for reduced computational requirements and facilitates resolving assembly problems when processing ultra-high coverage next-generation sequence data., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

The evolutionary origin and history of metazoan nervous systems has been at the heart of numerous scientific debates for well over a century. This has been a particularly difficult issue to resolve within the deuterostomes, chiefly due to the distinct neural architectures observed within this group of animals. Indeed, deuterosomes feature central nervous systems, apical organs, nerve cords, and basiepidermal nerve nets. Comparative analyses investigating the anatomy and molecular composition of deuterostome nervous systems have nonetheless succeeded in identifying a number of shared and derived features. These analyses have led to the elaboration of diverse theories about the origin and evolutionary history of deuterostome nervous systems. Here, we provide an overview of these distinct theories. Further, we argue that deciphering the adult nervous systems of representatives of all deuterostome phyla, including echinoderms, which have long been neglected in this type of surveys, will ultimately provide answers to the questions concerning the ancestry and evolution of deuterostome nervous systems.

The phylogenetic relationships among deuterostome animals have been debated for many years, and a diversity of hypotheses have been proposed based on both morphological and molecular data. Here we have assembled sequences of 217 nuclear-encoded proteins to address specific questions concerning their relationships and times of origin. We recovered significant support for urochordates as the closest relative of vertebrates with an analysis of 59 proteins (17,400 amino acids) and suggest that the basal position of urochordates found in previous molecular studies may have been the result of long-branch attraction biases. Our results also support Ambulacraria, the pairing of hemichordates with echinoderms (nine proteins; 2,382 amino acids), and Cyclostomata, the pairing of lampreys with hagfish (25 proteins; 6,895 amino acids). In addition, 325 shared proteins (102,110 amino acids) were obtained from the complete genomes of six vertebrates and a urochordate for phylogenetic analysis and divergence time estimation. An evolutionary timescale was estimated using a local (Bayesian) molecular clock method. We found that most major lineages of deuterostomes arose prior to the Cambrian Explosion of fossils (approximately 520 MYA) and that several lineages had originated before periods of global glaciation in the Precambrian.

Endo-alpha-D-mannosidase is a processing enzyme which in contrast to other glycosidases involved in the trimming of N-linked oligosaccharides of glycoproteins acts at an internal position by cleaving the linkage between the glucose-substituted mannose and the internal portion of the polymannose unit and thereby provides an alternate deglucosylating pathway. In order to evaluate at what stage in evolution this unusual enzyme first emerged, we have carried out a phylogenetic survey of its distribution among a broad group of eukaryotes ranging from unicellular organisms to highly developed animals and plants, all of which are known to have the capacity to N-glycosylate proteins and subsequently trim the nascent glucosylated polymannose oligosaccharides. It became evident from enzyme assays and in vivo studies that endomannosidase is limited in its distribution to members of the chordate phylum, including placental and marsupial mammals, birds, reptiles, amphibians, and fish, with the single except of the Mollusca in which it was detected in three distinct classes. The enzyme's absence in all other invertebrates examined as well as in yeast, various protozoa and higher plants, stands in contrast to glucosidase II and alpha 1,2-mannosidase which were found to be present in all eukaryotes studied. The observation that endomannosidase activity was not present in insects was confirmed by radiolabeling experiments with Sf9 cells in culture. These cells, which are widely employed for the expression of mammalian genes, were in distinction to mouse cells unable to circumvent a castanospermine (CST)-induced glucosidase blockade. Moreover we observed that Tetrahymenae, which synthesize glycoproteins with truncated N-linked oligosaccharides, could not process these beyond the Glc3Man5GlcNAc2 stage in the presence of CST. The late appearance of endomannosidase during evolution suggests a need for an alternate deglucosylation route in higher animals which parallels the development of elaborate complex N-linked oligosaccharides. Such carbohydrate units are believed to carry out vital biological functions and deglucosylation is a prerequisite to the further processing steps required for their formation.

The Sox gene family, a collection of transcription factors widely distributed throughout the animal kingdom, plays a crucial role in numerous developmental processes. Echinoderms occupy a pivotal position in many research fields, such as neuroscience, sex determination and differentiation, and embryonic development. However, to date, no comprehensive study has been conducted to characterize and analyze Sox genes in echinoderms. In the present study, the evolution and expression of Sox family genes across 11 echinoderms were analyzed using bioinformatics methods. The results revealed a total of 70 Sox genes, with counts ranging from 5 to 8 across different echinoderms. Phylogenetic analysis revealed that the identified Sox genes could be categorized into seven distinct classes: the SoxB1 class, SoxB2 class, SoxC class, SoxD class, SoxE class, SoxF class and SoxH class. Notably, the SoxB1, SoxB2, and SoxF genes were ubiquitously present in all the echinoderms studied, which suggests that these genes may be conserved in echinoderms. The spatiotemporal expression patterns observed for Sox genes in the three echinoderms indicated that various Sox members perform distinct functional roles. Notably, SoxB1 is likely involved in echinoderm ovary development, while SoxH may play a crucial role in testis development in starfish and sea cucumber. In general, the present investigation provides a molecular foundation for exploring the Sox gene in echinoderms, providing a valuable resource for future phylogenetic and genomic studies. [ABSTRACT FROM AUTHOR]

A large and taxonomically diverse collection of Ordovician Asterozoa from France and Morocco enables new insights into the early evolution of the subphylum. Available specimens of both Asteroidea and Stenuroidea are comparatively few, the collection dominated by Ophiuroidea and to a lesser extent, Somasteroidea. Nearly all the asteroid specimens are assigned to Petraster Billings, 1858; P. caidramiensis n. sp. is differentiated based on the complexity of the extraxial skeleton. The genus Petraster is reviewed. Another asteroid fragment, Euaxosida sp. A, is too incomplete to assign below the ordinal level. Among species of Petraster, the extraxial skeleton suggests homoplastic emergence of ossicular series definition whereas evolution of ambulacral axial-adaxial series expressions was conservative. The stemward asteroid axial/adaxial configuration is considered largely a plesiomorphic derivative of the somasteroid condition. Ossicular details of all series, however, differ among Petraster species. Euaxosida sp. A also offers oral-surface expressions suggestive of a transitional status. Differences among ossicular series expressions exemplify concerns surrounding interpretation of early asterozoan phylogeny and taxonomy. [ABSTRACT FROM AUTHOR]

Asexual reproduction by means of splitting, also called fissiparity, is a common feature in some asterozoan groups, especially in ophiactid brittle stars. Most fissiparous brittle stars show six instead of the usual five rays, live as epibionts on host organisms, and use clonal fragmentation to rapidly colonize secluded habitats and effectively expand the margins of their distribution area. While the biology and ecology of clonal fragmentation are comparatively well understood, virtually nothing is known about the evolution and geological history of that phenomenon. Here, we describe an exceptional fossil of an articulated six-armed brittle star from the Late Jurassic of Germany, showing one body half in the process of regeneration, and assign it to the new species Ophiactis hex sp. nov. Phylogenetic inference shows that the fossil represents the oldest member of the extant family Ophiactidae. Because the Ophiactis hex specimen shows an original six-fold symmetry combined with a morphology typically found in epizoic ophiuroids, in line with recent fissiparous ophiactid relatives, we assume that the regenerating body half is an indication for fissiparity. Ophiactis hex thus shows that fissiparity was established as a means of asexual reproduction in asterozoan echinoderms by the Late Jurassic. [ABSTRACT FROM AUTHOR]

Great strides have been made in understanding the phylogeny of the five extant echinoderm classes, however, many Palaeozoic groups have yet to be examined in a rigorous, quantitative framework. The aberrant morphologies of Paracrinoidea, an unusual group of Palaeozoic echinoderms, have hindered their inclusion in large‐scale phylogenetic and morphologic studies. This study uses a combined approach of phylogenetic analysis and morphological disparity to elucidate species relationships within the clade. Findings from this study suggest that Paracrinoidea is a monophyletic group and that respiratory structures, oral plate arrangement, and ambulacral morphologies are important for defining subclades within Paracrinoidea. Examination of paracrinoids in a quantitative framework, facilitates their inclusion in larger projects examining Palaeozoic echinoderm evolution, ecology and biogeography. [ABSTRACT FROM AUTHOR]

Elongation of very long-chain fatty acid (Elovl) proteins is the critical rate-limiting enzyme that catalyzes the biosynthesis of long-chain polyunsaturated fatty acids (LC-PUFAs). Echinoderms are extremely important research objects in various fields, such as regeneration studies and developmental biology. However, to date, the genome-wide characterization and analysis of Elovl genes in echinoderms have not been investigated. In this study, the identification and analysis of Elovl genes in 10 representative echinoderms were performed using bioinformatics methods. A total of 117 Elovl genes have been found in the studied echinoderms. The phylogenetic tree showed that all Elovl genes from echinoderms can be subdivided into 6 classes, Elovl3-like class, Elovl4-like class, Elovl5-like class, Elovl6-like class, Elovl7-like class, and Elovl8-like class. Furthermore, the selection pressure analysis suggested that Elovl genes in echinoderms were mainly constrained by strong purifying selection events. In general, this study provides a molecular basis for echinoderm Elovl genes and may serve as a reference for in-depth phylogenomics. [ABSTRACT FROM AUTHOR]

Previous molecular phylogenetic reconstructions have highlighted frequent convergences and iterations of external morphological characters during the evolution of crinoids. Classifications of extant and fossil crinoids based on these characters need to be thoroughly revised. This is particularly the case for the Isocrinida. We propose a new classification of this order based on molecular data and ontogenetic analysis of the derived characters displayed on ossicles and their articulations. The argument is strengthened by the discovery and description of a species belonging to a new genus (Panglaocrinus isseliformis gen. nov. sp. nov.) closely related to the genus Isselicrinus , which was abundant and widespread during the Late Cretaceous and Eocene. Extant Isocrinida appear to be divided into two major lineages that we interpret as two distinct families: Isocrinidae and Balanocrinidae. Isocrinidae are represented by the subfamily Metacrininae (genera Metacrinus and Saracrinus), while Balanocrinidae comprise four subfamilies: Balanocrininae (Neocrinus), Diplocrininae (Endoxocrinus and Teliocrinus), Isselicrininae (Panglaocrinus), and Proisocrininae (Proisocrinus). In the absence of molecular data, the genus Cenocrinus has not been attributed to any subfamily, but its close relationship to Balanocrinidae is suggested. Possible relationships between Mesozoic and modern taxa are discussed, although descriptions of arms in the palaeontological record are often important details. [ABSTRACT FROM AUTHOR]

Deep-sea holothurian specimens were collected during five scientific expeditions (2018–2023) using the submersible vehicle ‘Shenhaiyongshi’. Our examination of specimens of Deimatidae from the South China Sea and the Mariana fore-arc area revealed three new species, which were described as Oneirophanta idsseica sp. nov., Oneirophanta brunneannulata sp. nov., and Oneirophanta lucerna sp. nov. These species were distinguished from each other and from congeners by the arrangement, and number of ventrolateral tube feet and ossicle types. We also reported Oneirophanta mutabilis mutabilis Théel, 1879 for the first time from the Mariana fore-arc area, and we recorded Deima validum validum for the second time from the South China Sea. The taxonomy of these new species and new records is discussed, and a phylogenetic analysis based on a concatenated dataset of 16S and COI genes was conducted. Additionally, the inter- and intraspecific genetic divergences we calculated among deimatid species. The results support the assignment of these new species to the genus Oneirophanta and their separation from congeners. A description of the main morphological characters of Oneirophanta species is also provided. The data were collected from geographically diverse areas and suggest that species of Deimatidae were abundant in the Pacific Ocean and occupied a wide range of depths. [ABSTRACT FROM AUTHOR]

Ophiuroids are a diversified benthic taxon in the deep sea. Given their various dispersal strategies, they are considered an adequate group to assess genetic connectivity, especially in the seamounts that function as islands. Ophioleila elegans A.H. Clark, 1949, in the family Ophiothamnidae, was previously reported from the Caiwei Guyot, a seamount in the northwest Pacific Ocean. Here, we described the mitochondrial genome of O. elegans collected from another seamount in the northwest Pacific. The whole mitogenome is 16,376 bp in length and encodes 13 protein-coding genes, two ribosomal RNA genes, and 22 transfer RNA genes. Phylogenetic analysis based on the mitogenome sequences showed that O. elegans was clustered with Histampica sp., the only species for which mitogenome sequence has been reported within the family Ophiothamnidae. The complete mitogenome of O. elegans first reported in the present study provides useful information for population genetics and evolutionary relationship of this taxon, especially in the northwest Pacific seamounts. [ABSTRACT FROM AUTHOR]

An increasing number of Antarctic invertebrate taxa have been revealed as cryptic species complexes following DNA-based assessments. This ultimately necessitates a morphological reassessment to find traits that will help identify these cryptic or pseudocryptic species without the need for sequencing every individual. This work concerns comatulid crinoid echinoderms long considered to represent a single, circum-Antarctic species, Promachocrinus kerguelensis. The first molecular studies sought to distinguish the diversity in the complex and understand the constituent species distributions but stopped short of formal taxonomic assessment. Here, we continued to increase sample representation around the Southern Ocean and sequenced the mitochondrial COI gene for all new specimens, and additional genes for a few representatives. We also elucidated previously unappreciated features, particularly body pigmentation and morphology of the centrodorsal ossicle in an attempt to diagnose some species morphologically and based on DNA data. The species complex within Promachocrinus is here resolved into P. kerguelensis Carpenter, 1879, P. vanhoeffenianus Minckert, 1905, P. joubini Vaney, 1910, P. mawsoni (Clark, 1937) comb. nov. (transferred from Florometra) and four previously unnamed species, P. fragarius sp. nov., P. unruhi sp. nov., P. uskglassi sp. nov. and P. wattsorum sp. nov. Although most species can be distinguished morphologically, several cannot be reliably separated without DNA data. All sequenced species are essentially circum-Antarctic, with the notable exception of P. wattsorum sp. nov. that is restricted to the Prince Edward Islands in the sub-Antarctic Indian Ocean and P. vanhoeffenianus that is only known from the type locality in the Davis Sea. The vast nature of the Antarctic and Southern Ocean ecosystem dictates large scale sampling to understand the full extent of the biodiversity. ZooBank: urn:lsid:zoobank.org:pub:F871CDC8-973B-48CE-8A61-33658D4EB4B1 The Antarctic feather star Promachocrinus kerguelensis arguably comprises several species but taxonomic revision has not been attempted. Here we use DNA data and morphological characters to differentiate the species and formally described those that are new. The species complex was resolved into eight species, four of which are newly described, though not all can be reliably distinguished based on morphology. One of the new species has a restricted distribution, but the remainder appear to be circum-Antarctic. [ABSTRACT FROM AUTHOR]

The family Zoroasteridae is a clade of extant sea stars found exclusively in deep marine environments (i.e. bathyal environments deeper than 200 m). In the fossil record, seven species have been compared and/or assigned to the Zoroasteridae, on a time span ranging from the Middle Jurassic to the Miocene. This study describes a new taxon, Viridisaster guerangeri gen. et sp. nov., from the Cenomanian of Le Mans (Sarthe, France) and reappraises two Mesozoic species, the Jurassic Terminaster cancriformis and the Early Cretaceous Protothyraster priscus. To test the relationships of the putative fossil zoroasterids with modern taxa, a phylogenetic analysis was performed combining six of the best-known fossil species and 29 forcipulatacean species, including four Zoroasteridae. Many of the characters that define the extant Zoroasteridae do not appear in the fossil record before the Cenozoic. The family name is maintained for the crown group only. The family Terminasteridae, that accounted for the genera Terminaster and Alkaidia, is found paraphyletic and is rejected here. The order Zorocallida, initially erected for the family Zoroasteridae only, is redefined as the sister clade to the Forcipulatida that includes the crown-group Zoroasteridae and its stem relatives. Mesozoic Zorocallida are small in size compared to the Cenozoic Zorocallida, and they express morphological characters that are typical of juvenile Zoroasteridae; thus, the emergence of modern forms is likely associated with peramorphic evolution. The palaeoecology of the Zoroasteridae and their restriction to deep-sea environments after the Eocene is discussed. [ABSTRACT FROM AUTHOR]

Evaluating the diversity of crinoids (Echinodermata: Crinoidea) in Singapore accurately has been challenging due in part to inadequacies in crinoid taxonomy. Most species descriptions have neither incorporated morphological variation of juveniles into diagnostic characters nor analysed species' molecular phylogenetic affinities. In this study, we performed detailed morphological examination and phylogenetic analyses of the mitochondrial cytochrome c oxidase subunit I marker on Himerometroidea crinoids collected from the Singapore Strait, and showed that specimens could not be reliably identified to species based on existing taxonomic descriptions. Specifically, juveniles identified as Heterometra spp. and Zygometra sp., and adults as H. cf. producta, solely based on morphological features were revealed to comprise three distinct species: Zygometra cf. comata, Heterometra schlegelii, and Dichrometra sp. The juveniles were found to have sizes smaller than the ranges given in existing descriptions, with considerable variabilities in traits that limited the utility of morphology for species taxonomy. Features contributing to misidentification of specimens and their application in past species diagnoses are discussed. Revised descriptions of these species, as well as the recently revised Homalometra crenulata, are also presented. More broadly, this study emphasises the need to revisit diagnoses of crinoid species according to various ontogenetic stages and to uncover more congruent morphological characters based on a robust molecular phylogeny. [ABSTRACT FROM AUTHOR]

Ophiuroidea is the most speciose of all classes of Echinoderma. It is an important component in benthic ecosystems, occurring in almost all ecological niches of modern seas. To date, the phylogeny and complete evolutionary history of the ophiuroids have not yet been fully resolved. In this study, we sequenced the complete mitochondrial genomes (mitogenomes) of Ophiothrix (Ophiothrix) exigua and two deep‐sea species Histampica sp. CS049 and Ophioplinthaca sp. M5261. These two deep‐sea ophiuroids displayed reversed strand‐compositional bias and rearranged gene orders. Thirteen distinct patterns of mitochondrial gene order among ophiuroid mitogenomes were detected, with two gene order newly found in Ophiuroidea. Our data supported the gene order found in all sampled Ophiuridae as the most likely ancestral order of all Ophiuroidea. To improve phylogenetic accuracy based on nucleotide differences, two different criteria were used for the analyses: (i) nucleotide sequence from all codon positions (PCG123); (ii) the NTE method ("Neutral Transitions Excluded") for ameliorating the misleading effects of a reverse strand bias in the data. The two methods confirmed the polyphyly of the orders Ophiacanthida and Amphilepidia. At family and genus level, Ophiuridae, Ophionotus and Ophioplinthus were not monophyletic. The most notable exception was that the NTE phylogeny showed low variation of branch length. NTE dataset generated younger age for most lower‐level nodes than that from PCG123 dataset. All analyses suggested that the ophiuroids radiation occurred around the Permian–Triassic mass extinction event, and the divergence time of the deep‐sea lineages was during the Cretaceous. [ABSTRACT FROM AUTHOR]

'Asturicystis' havliceki Fatka & Kordule from the middle Cambrian of Bohemia (Czech Republic) is re‐described based on its type material. Several features, including the extension of the food grooves and presence of ventral swellings suggest that 'A.' havliceki does not belong to Asturicystis, and is placed in the new genus Bohemiacinctus. To ascertain the phylogenetic position of Bohemiacinctus havliceki, we conducted Bayesian fossil tip‐dating and parsimony‐based phylogenetic analyses of 24 species spanning all major groups of cinctan higher taxa. Results show a high degree of congruence between tree topologies recovered by both tip‐dated and parsimony‐based analyses. Both methods indicate B. havliceki is phylogenetically distant from Asturicystis and is most likely to be an early representative of the family Sucocystidae. Overall, our phylogeny is broadly similar to previous estimates of cinctan relationships, including a more conventional phylogenetic position of controversial taxa such as Protocinctus. These results point to the sensitivity of small clades such as cinctans to taxon sampling effects, and highlights the importance of taxonomy and accurate morphological character descriptions in phylogenetic analyses of fossil taxa. [ABSTRACT FROM AUTHOR]

The common ancestor of Ambulacraria was a mobile bilaterally symmetrical organism. Its body was subdivided into a preoral (proboscis) region, a perioral (collar) region with ciliated tentacles, and a trunk region with a metameric coelom and metameric gill slits. Like in other Bilateria, the preoral and tentacular regions of Ambulacraria are free of Hox gene expression, which begins in the area of the first pair of gill slits in compliance with the colinearity. The axial complex is a synapomorphy of the Ambulacraria clade. A bilaterally symmetrical ancestor of Echinodermata lay on its dorsal side, so its anus was shifted to the ventral side as in present-day echinoderm larvae. This stage of evolution corresponds to the Early Paleozoic bilaterally symmetrical forms, such as Protocinctus, Ctenocystis, and Ctenoimbricata. The common ancestor of echinoderms had a symmetrical tentacular apparatus consisting of five ciliated tentacles on each side of a collar. At the next stage, the ancestors of echinoderms lay on the right side that resulted in the reduction of the tentacles on the right side and the right hydrocoel. This evolution stage includes various Early Paleozoic forms (Cothurnocystis, Dendrocystoides, Syringocrinus, Castericystis, Coleicarpus, Rhenocystis, etc.). The next stage is related to the sedentary lifestyle. During this stage, the mouth and tentacles occupied an apical position, the anus has shifted up, and thus an intestinal loop was formed. The five primary tentacles of the left side of the ancestor of Ambulacraria predestined the formation of pentaradial symmetry of echinoderms. The primary tentacles remained only in Holothuroidea. The secondary ciliary grooves were formed between the primary tentacles. These grooves were accompanied by hydrocoelic canals, which gave rise to the water-vascular (ambulacral) system. The present-day echinoderms are characterized by several metameric rings formed by the derivatives of the left somatocoel. They derive from the coelomic segments of the left side of the trunk of the ambulacrarian ancestor and, thus, the echinoderms retain coelomic metamerism. The attachment of the crinoid larvae by the preoral lobe reflects the ancient method of locomotion of deuterostomes using the proboscis, but, in reality, the Pelmatozoa stalk is homologous to the Pterobranchia stalk, i.e., the posterior end of the body, because the right somatocoel grows into it. During metamorphosis, the internal coelomic complex of the larva is inverted by 180°. The significance of this inversion for the translocation of the anterior genes of the Hox cluster is discussed. The ancestors of Eleutherozoa began to crawl on the oral surface, which led to a shift of the anus to the aboral side. Thus, the ontogeny and phylogeny of echinoderms exhibits a change from bilateral symmetry to dissymmetry, and then the development of pentameric symmetry. [ABSTRACT FROM AUTHOR]

The complete mitochondrial genome of the feather star Cenometra bella was sequenced in this study. The mitogenome is 15,872 bp in length, with 13 PCGs, 22 tRNA, and two rRNA, and nucleotide composition was as follows: 24.38% A, 47.79% T, 11.16% C, and 16.68% G. Phylogenetic analyses place C. bella as closely related to Stephanometra indica, consistent with previous inferences.

A proposed new Ordovician asterozoan genus and species, Cantabrigiaster fezouataensis , has been recently described and assigned to the class Somasteroidea. An accompanying phylogenetic analysis treated twenty-two asterozoans and sixteen early non-asterozoans, one of the latter providing the outgroup. The phylogenetic hypothesis focused on derivation of the subphylum Asterozoa and emphasized the Crinoidea; Cantabrigiaster fezouataensis was interpreted as documenting a critical transition stage in the origin of the Asterozoa. The diagnosis for the proposed new generic name posits absence of ambital framework ossicles as the single character expression that distinguishes Cantabrigiaster among somasteroid genera. Ambital framework ossicular presence is documented here in the holotype and other members of type suite; Cantabrigiaster is synonymized with Villebrunaster Spencer and V. fezouataensis nov. comb. is redescribed. The data matrix for the former Cantabrigiaster phylogenetic analysis relies on an earlier study treating overall echinoderm construction. Expressions outlined in the earlier study are not amenable to transfer to a data matrix without evaluation of level of universality of each cited expression; such evaluations were not provided thereby leading to flawed phylogenetic conclusions that are rejected. An alternative hypothesis for the early history of the Asterozoa supported by aspects of the fossil record, that subphylum diversification preceded the origin of readily preserved skeletons, and therefore potentially no tenable pre-asterozoan outgroup candidate survives in the fossil record, has not been falsified. [ABSTRACT FROM AUTHOR]

Acorn worms, also known as enteropneust (literally, 'gut-breathing') hemichordates, are marine invertebrates that share features with echinoderms and chordates. Together, these three phyla comprise the deuterostomes. Here we report the draft genome sequences of two acorn worms, Saccoglossus kowalevskii and Ptychodera flava. By comparing them with diverse bilaterian genomes, we identify shared traits that were probably inherited from the last common deuterostome ancestor, and then explore evolutionary trajectories leading from this ancestor to hemichordates, echinoderms and chordates. The hemichordate genomes exhibit extensive conserved synteny with amphioxus and other bilaterians, and deeply conserved non-coding sequences that are candidates for conserved gene-regulatory elements. Notably, hemichordates possess a deuterostome-specific genomic cluster of four ordered transcription factor genes, the expression of which is associated with the development of pharyngeal 'gill' slits, the foremost morphological innovation of early deuterostomes, and is probably central to their filter-feeding lifestyle. Comparative analysis reveals numerous deuterostome-specific gene novelties, including genes found in deuterostomes and marine microbes, but not other animals. The putative functions of these genes can be linked to physiological, metabolic and developmental specializations of the filter-feeding ancestor.

Within all major taxa of Bilateria, there are forms with coelomic metamerism. This suggests that coelomic metamerism was characteristic of the common ancestor of Bilateria. Among deuterostomes, metamerism is clearly expressed in chordates, and elements of metamerism are present in hemichordates. Do echinoderms have remnants of coelomic metamerism that was inherited from the common ancestor of Bilateria? The coelomic system of echinoderms includes several metameric coelomic rings located along the oral-aboral axis, namely: the axocoelomic ring, the hydrocoelomic ring, 2 to 6 coelomic rings originating from the left somatocoel, and one epigastric ring originating from the right somatocoel. Thus, in echinoderms, there is a dissymmetrical metamerism, derived from the original metamerism of the common ancestors of Deuterostomia and, possibly, the common ancestors of Bilateria. The problem of dexiothetism as the cause for the formation of coelomic dissymmetry in echinoderms is discussed. [ABSTRACT FROM AUTHOR]

Chemosensation is a critical signalling process in animals and especially important in sea cucumbers, a group of ecologically and economically important marine echinoderms (class Holothuroidea), which lack audio and visual organs and rely on chemical sensing for survival, feeding and reproduction. The ionotropic receptors are a recently identified family of chemosensory receptors in insects and other protostomes, related to the ionotropic glutamate receptor family (iGluR), a large family of membrane receptors in metazoan. Here we characterize the echinoderm iGluR subunits and consider their possible role in chemical communication in sea cucumbers. Sequence similarity searches revealed that sea cucumbers have in general a higher number of iGluR subunits when compared to other echinoderms. Phylogenetic analysis and sequence comparisons revealed GluH as a specific iGluR subfamily present in all echinoderms. Homologues of the vertebrate GluA (aka α‐amino‐3‐hydroxy‐5‐methyl‐4‐isoxazolepropionic acid, AMPA), GluK (aka kainate) and GluD (aka delta) were also identified. The GluN (aka N‐methyl‐d‐aspartate, NMDA) as well as the invertebrate deuterostome subfamily GluF (aka phi) are absent in echinoderms. The echinoderm GluH subfamily shares conserved structural protein organization with vertebrate iGluRs and the ligand binding domain (LBD) is the most conserved region; genome analysis indicates evolution via lineage and species‐specific tandem gene duplications. GluH genes (named Grih) are the most highly expressed iGluRs subunit genes in tissues in the sea cucumber Holothuria arguinesis, with Griha1, Griha2 and Griha5 exclusively expressed in tentacles, making them candidates to have a chemosensory role in this species. The multiple GluH subunits may provide alternative receptor assembly combinations, thus expanding the functional possibilities and widening the range of compounds detected during aggregation and spawning in echinoderms. [ABSTRACT FROM AUTHOR]

The echinoids of the order Bothriocidaroida represent the initial burst of sea urchin diversification. They were the first echinoids to achieve widespread biogeographical dispersal and achieved high levels of species richness compared to other clades of stem group echinoids. Following long-standing controversy regarding their phylogenetic affinities within the phylum Echinodermata, bothriocidaroids are now regarded as echinoids. The species- and genus-level phylogenetic relationships of the bothriocidaroids, are, however, less clearly known. We herein compile a database of bothriocidaroid occurrences, undertake detailed phylogenetic analyses of all named species within the Bothriocidaroida, and provide updated diagnoses for all species. Our phylogenetic analysis includes species belonging to three genera: Bothriocidaris, Unibothriocidaris and Neobothriocidaris. Furthermore, we describe a new species, Neobothriocidaris pentlandensis sp. nov., from the Silurian of Scotland and Sweden, using micro-computed tomography and scanning electron microscopy. This species has implications for understanding the diversity dynamics of bothriocidaroids before and after the Late Ordovician mass extinction and the palaeobiogeographical distribution of bothriocidaroids in the Silurian. Using Bayesian and parsimony-based phylogenetics, we demonstrate that Neobothriocidaris is a distinct clade, and depending upon the analytical technique, Unibothrioidaris and Bothriocidaris are also resolved as monophyletic. Unibothriocidaris appears to have become extinct in the Sandbian, while Neobothriocidaris and Bothriocidaris survived the Late Ordovician extinction events and lasted at least until the Ludlow. Furthermore, we performed statistical time-calibration of our phylogenetic trees and show that bothriocidaroids originated in the Dapingian or Darriwilian, diversifying in the Darriwilian to Sandbian and reaching peak diversity in the Sandbian and Katian. This Sandbian and Katian peak is similar to that observed in other echinoderm clades, as well as other animal groups during the Ordovician radiation, and may be linked to heightened sea level and expansive tropical shelves present in the Upper Ordovician, and to new inhabitable environments linked with Ordovician oxygenation. [ABSTRACT FROM AUTHOR]

BackgroundRecent genomic information has revealed that neuroglobin and cytoglobin are the two principal lineages of vertebrate hemoglobins, with the latter encompassing the familiar myoglobin and α-globin/β-globin tetramer hemoglobin, and several minor groups. In contrast, very little is known about hemoglobins in echinoderms, a phylum of exclusively marine organisms closely related to vertebrates, beyond the presence of coelomic hemoglobins in sea cucumbers and brittle stars. We identified about 50 hemoglobins in sea urchin, starfish and sea cucumber genomes and transcriptomes, and used Bayesian inference to carry out a molecular phylogenetic analysis of their relationship to vertebrate sequences, specifically, to assess the hypothesis that the neuroglobin and cytoglobin lineages are also present in echinoderms.ResultsThe genome of the sea urchin Strongylocentrotus purpuratus encodes several hemoglobins, including a unique chimeric 14-domain globin, 2 androglobin isoforms and a unique single androglobin domain protein. Other strongylocentrotid genomes appear to have similar repertoires of globin genes. We carried out molecular phylogenetic analyses of 52 hemoglobins identified in sea urchin, brittle star and sea cucumber genomes and transcriptomes, using different multiple sequence alignment methods coupled with Bayesian and maximum likelihood approaches. The results demonstrate that there are two major globin lineages in echinoderms, which are related to the vertebrate neuroglobin and cytoglobin lineages. Furthermore, the brittle star and sea cucumber coelomic hemoglobins appear to have evolved independently from the cytoglobin lineage, similar to the evolution of erythroid oxygen binding globins in cyclostomes and vertebrates.ConclusionThe presence of echinoderm globins related to the vertebrate neuroglobin and cytoglobin lineages suggests that the split between neuroglobins and cytoglobins occurred in the deuterostome ancestor shared by echinoderms and vertebrates.

Understanding of the evolutionary history of the ophiuroids, or brittle stars, is hampered by a patchy knowledge of the fossil record. Especially, the stem members of the living clades are poorly known, resulting in blurry concepts of the early clade evolution and imprecise estimates of divergence ages. Here, we describe new ophiuroid fossil from the Lower Jurassic of France, Luxembourg and Austria and introduce the new taxa Ophiogojira labadiei gen. et sp. nov. from lower Pliensbachian shallow sublittoral deposits, Ophiogojira andreui gen. et sp. nov. from lower Toarcian shallow sublittoral deposits and Ophioduplantiera noctiluca gen. et sp. nov. from late Sinemurian to lower Pliensbachian bathyal deposits. A Bayesian morphological phylogenetic analysis shows that Ophiogojira holds a basal position within the order Ophiurida, whereas Ophioduplantiera has a more crownward position within the ophiurid family Ophiuridae. The position of Ophioduplantiera in the evolutionary tree suggests that family-level divergences within the Ophiurida must have occurred before the late Sinemurian, and that ancient slope environments played an important role in fostering early clade evolution.

Despite descending from heavily calcified ancestors, the holothuroid skeleton is fully internal and composed of microscopic ossicles and a ring of plates bound by connective tissue, the calcareous ring. The calcareous ring exhibits a complex and poorly understood morphology; as a result, establishing unambiguous homology statements about its macrostructure has been challenging and phylogenetic studies have had to simplify this important structure. Here, we provide the first broad comparative study of Dendrochirotida calcareous rings using micro‐computed tomography (μCT). A detailed description of the three‐dimensional macrostructure of the calcareous ring of 10 sea cucumber species, including rare and type specimens, is presented. The structures observed were highly variable at the subfamily level, especially at the point of tissue attachment. The relationship between the calcareous ring and its associated organs, and their functional morphology are discussed. To aid future phylogenetic studies, we listed 22 characters and performed a preliminary cladistic analysis. The topology obtained supports the idea that the simple, cucumariid ring is ancestral to the mosaic‐like phyllophorid ring; however, it did not support the monophyly of the cucumariids. It also did not support the family Sclerodactylidae, which was described based on the ring morphology. Differently from the dermal ossicles, which are highly homoplastic, the general homoplasy index of the calcareous ring characters was relatively low. This result highlights the importance of this structure for phylogenetic inference. Unfortunately, time since collection, rough collection methods and fixation can damage the skeleton, and the calcareous ring is often overlooked in taxonomic descriptions. The data presented here will improve our understanding of holothuroid relationships and facilitate studies on holothuroid functional morphology and biomechanics. [ABSTRACT FROM AUTHOR]

The phylogenetic relationships of spherical and sub-spherical asteroids, belonging to the valvatidan family Sphaerasteridae (Early Jurassic–Recent), are revised in the light of abundant new fossil material from Europe and North Africa. The family had radiated by the Early Jurassic and the family Stauranderasteridae is its sister group. Morphological changes involved the formation of a domed body, the absence of differentiated marginal ossicles and the transformation of dorsal abactinal ossicles into a tessellation of large, thin plates. The family Podosphaerasteridae is placed in synonomy with the Sphaerasteridae. New taxa include Eosphaeraster amellagensis gen. et sp. nov., Bulbosphaeraster valettei gen. et sp. nov., Pouzaster pocknotata gen. et sp. nov., Rugosphaeraster rugenensis gen. et sp. nov., Echinosphaeraster gen. nov. (type species: Asterias scutata Goldfuss, 1833) and Valettaster thuyi sp. nov. It is suggested that the morphology of Sphaerasteridae is related to their close association with sponges. [ABSTRACT FROM AUTHOR]

[Display omitted] • A new phylogeny of Brisingida with broad coverage is reported. • Both Brisingida families and four genera are recovered as non-monopheletic. • Brisingida is revised to encompass 5 families and 17 genera. • New genus, species and combinations are described and a diagnostic key is provided. • Convergence caused by paedomorphosis is found in phylogenetically distant genera. Brisingida Fisher 1928 is one of the seven currently recognised starfish orders, and one of the least known taxa as being exclusive deep-sea inhabitants. Modern deep-sea expeditions revealed their common occurrences in various deep-sea settings including seamounts, basins and hydrothermal vent peripheral, underlining the necessity of clarifying their global diversity and phylogeny. In this study, we present a comprehensive molecular phylogeny of Brisingida which encompasses the highest taxonomic diversity to date. DNA sequences (COI, 16S, 12S and 28S) were obtained from 225 specimens collected in the global ocean, identified as 58 species spanning 15 of the 17 extant genera. Phylogenetic relationship was inferred using both maximum likelihood and Bayesian inference methods, revealing polyphyletic families and genera and indicating nonnegligible bias in prior morphology-based systematics. Based on the new phylogeny, a novel classification of the order, consisting of 5 families and 17 genera, is proposed. Families Odinellidae, Brisingasteridae and Novodiniidae (sensu Clark and Mah, 2001) were resurrected to encompass the genera Odinella , Brisingaster and Novodinia. Brisingidae and Freyellidae were revised to include 11 and 3 genera, respectively. A new genus and species, two new subgenera and seven new combinations are described and a key to each genus and family is provided. Transformations of morphological traits were evaluated under the present phylogenetic hypothesis. A series of paedomorphic characters were found in many genera and species, which led to a high degree of homoplasy across phylogenetically distant groups. Our results provide new insights in the phylogeny and ontogeny of the order, and highlight the necessity to evaluate character convergence under sound phylogenetic hypothesis. [ABSTRACT FROM AUTHOR]

The complete mitochondrial genome of the feather star Cenometra bella was sequenced in this study. The mitogenome is 15,872 bp in length, with 13 PCGs, 22 tRNA, and two rRNA, and nucleotide composition was as follows: 24.38% A, 47.79% T, 11.16% C, and 16.68% G. Phylogenetic analyses place C. bella as closely related to Stephanometra indica, consistent with previous inferences. [ABSTRACT FROM AUTHOR]

Ontogeny of non-muscular brachial articulations in extant species of Balanocrininae, i.e., Neocrinus decorus, Neocrinus blakei and Hypalocrinus naresianus (Crinoidea, Isocrinida), is described using SEM observations. All three species share embayed synarthries and symplexies (previously only known in crinoid stalks) showing a radiating crenularium pattern in their proximal arms but differ in several important ways. Neocrinus decorus has a shallow simple symmorphy affecting symplexies, and embayed synarthries. During the latest ontogeny of embayed synarthries, irregular syzygial ridges appear on the aboral segment of the fulcral ridge. Neocrinus blakei and H. naresianus share a peculiar sharp deep symmorphy superimposed on symplexies, and synarthries with a more complete single fulcral ridge that only appears late in ontogeny. Comparison with other crinoid taxa that have more advanced arm axial synarthries shows that this ontogenetic trajectory is restricted to paedomorphic stages in extant balanocrinins. An embayed synarthry seems to be derived from the earliest developmental stage of the radiating symplexial crenularium via hypermorphosis of a single crenula. An embayed synarthry is, therefore, a symplesiomorphy based on paedomorphic stage of development; it thus lacks phylogenetic significance, and should be abandoned as a major character in the classification of Isocrinida. The most advanced brachial synarthries shared by distant crinoid taxa mainly represent a homoplasy under morphofunctional constraints. However, they could result from different ontogenetic trajectories, which have only rarely been investigated. Another distinctive articulation feature, the peculiar sharp deep symmorphy observed in extant balanocrinins is a derived character known in a few fossil isocrinids beginning in the Middle Jurassic. We question its phylogenetic significance and suggest that it has developed repeatedly via iterative evolution in Isocrinida. Therefore, because these three extant balanocrinin species share the same ontogenetic trajectories of arm and stalk ligamentary articulations, and differ only in various states of heterochronic development of a few characters, we treat them as belonging to the same genus. We, therefore, consider Hypalocrinus as a junior synonym of the genus Neocrinus. [ABSTRACT FROM AUTHOR]

"Pan-dichoporites" (new informal term) is proposed to unite Cambrian blastozoans, such as Cambrocrinus, Ridersia, and Sanducystis, glyptocystitoid and hemicosmitoid rhombiferans, coronates, blastoids, and Lysocystites. Pan-dichoporite ambulacra have double biserial main axes with brachiole facets shared by pairs of floor (glyptocystitoids), side (blastoid) or trunk (hemicosmitoids, coronates) plates. These axial plates are the first two brachiolar plates modified to form the ambulacral axes. In glyptocystitoids the first brachiole facet in each ambulacrum is shared by an oral and another plate. Hence, these are also two modified brachiolar plates and part of the axial skeleton under the Extraxial Axial Theory (EAT). Pan-dichoporites are also characterized by thecae composed of homologous plate circlets. The unique glyptocystitoid genus Rhombifera bears ambulacral facets on five radial plates, which alternate with five orals. The oral area of Lysocystites (blastoid sensu lato) is very similar, which suggests that rhombiferan radials are homologous with "ambulacrals" of Lysocystites and hence with blastoid lancet plates. This implies derivation of blastoids from glyptocystitoids and suggests that blastoid and coronate radials and deltoids are homologous with rhombiferan infralaterals and laterals. Thus, homologous plate circlets occur in all pan-dichoporites, which strengthens the validity of a pan-dichoporite clade. Under Universal Elemental Homology (UEH), deltoids were homologized with rhombiferan orals, but this is inconsistent with the EAT. Deltoids bear respiratory pore structures and so are perforate extraxial skeletal plates, whereas rhombiferan orals are axial skeleton. The new plate homologies also explain why only five plates form the oral frames of coronates, blastoids and Lysocystites, whereas glyptocystitoids (except Rhombifera) have six oral frame plates; all glyptocystitoids have only five laterals. Hemicosmitoids arose by paedomorphic ambulacral reduction, but the paedomorphosis also affected the thecal plates and stem. Paedomorphosis poses special problems for cladistic character analysis, since the new characters often appear to be plesiomorphic. [ABSTRACT FROM AUTHOR]

Anneissia intermedia (A.H. Clark, 1916) is a common crinoid found in waters along the coastlines of China, Japan, and Korea. In this study, we determined the complete mitogenome of A. intermedia. The genome was found to be 15,874 bp in length and consists of 13 protein-coding genes, 22 tRNAs, and 2 rRNAs. With the exception of Antedon mediterranea, the gene order and genetic characteristics of the A. intermedia mitogenome are identical with those of the mitogenomes of other crinoids. The complete mitogenome of A. intermedia will contribute to enhance our understanding of the phylogeny of crinoids.

The complete mitochondrial genome of the widespread and common Indo-Pacific sea star Archaster typicus has been sequenced in this study. The mitogenome is 16,230 base pairs (bp) in length, with 13 protein coding genes (PCGs), 22 tRNAs and 2 rRNAs. Gene order of its PCGs and rRNAs matches those of nine other asteroid taxa included for comparison in this study, and it has a similar nucleotide composition of 33.08% A, 26.38% T, 25.53% C and 15.01% G nucleotides. Phylogenetic analyses place A. typicus as the sister group to Acanthaster spp., consistent with previous inferences.

A new phylogenetic analysis of the superorder Forcipulatacea is presented. Forcipulatacea is one of the three major groups of sea stars (Asteroidea: Echinodermata), composed of 400 extant species. The sampled taxa are thought to represent the morphological diversity of the group. Twenty-nine forcipulate taxa were sampled belonging to Asteriidae, Stichasteridae, Heliasteridae, Pedicellasteridae, Zoroasteridae and Brisingida. Specimens were dissected with bleach. Detailed description of the skeleton and the anatomy of the ossicles were investigated using scanning electron microscopy. Comparative anatomy allowed the scoring of 115 phylogenetically informative characters. The consensus tree resulting from the analysis recovers Asteriidae, Stichasteridae, Zoroasteridae and Brisingida as monophyletic. All types of morphological features contribute to tree resolution and may be appropriate for taxon diagnosis. The synapomorphies supporting different clades are described and discussed. Brisingida and Zoroasteridae are the best-supported clades. The potentially challenging position of Brisingida in the tree may be explained by homoplastic changes, but also by the presence of numerous non-applicable characters. [ABSTRACT FROM AUTHOR]

Morphologic and molecular data often lead to different hypotheses of phylogenetic relationships. Such incongruence has been found in the echinoderm class Echinoidea. In particular, the phylogenetic status of the order Clypeasteroida is not well resolved. Complete mitochondrial genomes are currently available for 29 echinoid species, but no clypeasteroid had been sequenced to date. DNA extracted from a single live individual of Sinaechinocyamus mai was sequenced with 10× Genomics technology. This first complete mitochondrial genome (mitogenome) for the order Clypeasteroida is 15,756 base pairs in length. Phylogenomic analysis based on 34 ingroup taxa belonging to nine orders of the class Echinoidea show congruence between our new genetic inference and published trees based on morphologic characters, but also includes some intriguing differences that imply the need for additional investigation. • The first mitochondrial genome (15,756 bp) of the enigmatic sand dollar Sinaechinocyamus mai. • Phylogenomic analyses of the class Echinoidea based on complete mitogenomes. • Filtering protocol for extracting valid echinoid data from contaminated 10× Genomics raw data prior to genome assembly. [ABSTRACT FROM AUTHOR]

Eublastoids are a large clade of blastoids; stemmed blastozoan echinoderms diagnosed by their conservative body plan (three basals, four deltoid plates and five radial plates), lancet plate supporting the ambulacra, and hydrospire respiratory structures. Although Eublastoidea was a highly successful clade in the middle and late Palaeozoic it is absent from early echinoderm radiations seen in the Cambrian and Ordovician record. Here we provide a re‐evaluation of Macurdablastus uniplicatus Broadhead from the Ordovician, using detailed morphological assessment based on advanced synchrotron tomography and phylogenetic analysis. Macurdablastus uniplicatus falls outside Eublastoidea because of the morphological differences in lancet plate and respiratory structures. The oldest recorded eublastoid is thus middle Silurian in age. The re‐evaluation of the morphology of Macurdablastus provides a basis for revising blastoid phylogeny and classification. [ABSTRACT FROM AUTHOR]

Inclusion of fossils can be crucial to address evolutionary questions, because their unique morphology, often drastically modified in recent species, can improve phylogenetic resolution. We performed a cladistic analysis of 45 cassidulids with 98 characters, which resulted in 24 most parsimonious trees. The strict consensus recovers three major cassiduloid clades, and the monophyly of the family Cassidulidae is not supported. Ancillary analyses to determine the sensitivity of the phylogeny to missing data do not result in significantly different topologies. The taxonomic implications of these results, including the description of a new cassiduloid family and the evolution of some morphological features, are discussed. Cassiduloids (as defined here) most probably originated in the Early Cretaceous, and their evolutionary history has been dominated by high levels of homoplasy and a dearth of unique, novel traits. Despite their high diversity during the Palaeogene, there are only seven extant cassiduloid species, and three of these are relicts of lineages dating back to the Eocene. Future studies of the biology of these poorly known species, some of which brood their young, will yield further insights into the evolutionary history of this group. [ABSTRACT FROM AUTHOR]

Next-generation sequencing was used to obtain the complete mitogenome of Henricia pachyderma (Hayashi, ). The mitogenome form was found to be a circular molecule 16,192 bp long with a 57.1% AT bias. Gene arrangement and composition of H. pachyderma were exactly the same as those of previously reported mitogenomes from other species of the family Echinasteridae, including H. leviuscula and Echinaster (Othilia) brasiliensis. Phylogenetic analysis with maximum likelihood and Bayesian inference revealed that H. pachyderma formed a monophyletic clade with H. leviuscula and E. (O.) brasiliensis belonging the same taxon, family Echinasteridae.

Arachnoides placenta is the most common sand dollar species in western Taiwan. Due to its lack of fossil record, the arrival of this species in Taiwan could not be estimated until now. Fragments of A. placenta was recovered sorting through a 1981- museum collection of sand dollars collected off Mailiao county, western Taiwan. Additional echinoderm specimens were recovered from Holocene sediments nearby, but no fragments of A. placenta were found, indicating a decline in the living population. A total of 11 echinoderm samples, including five specimens that were assigned to A. placenta with certainty, were sent for radiocarbon dating. Results showed that studied echinoderm fragments range from modern (after 1950) to as old as 2680 ± 55 a BP and the oldest date is from A. placenta. It is evident that Taiwan is an active migration front for A. placenta due to the relatively young arrival age for this pandemic species which is widespread in the Indo-West Pacific region. Additionally, the DNA sequence, including the cytochrome c oxidase subunit I (cox1) gene, of the species was obtained to conduct the first molecular phylogenetic study of sand dollars in Taiwan. [ABSTRACT FROM AUTHOR]

The phylogenetic relationships of Paleozoic blastozoan echinoderms are poorly understood and many of the traditionally ascribed groups are likely polyphyletic. Diploporitans, those blastozoans with double pore (diplopore) respiratory structures, have never been placed within a rigorous phylogenetic framework, and their highly variable morphologies suggest that they do not represent a natural clade. A maximum parsimony phylogenetic analysis, spanning a wide range of diploporitan and related taxa, indicates that diplopore-bearing blastozoans are a polyphyletic grouping and, consequently, that diplopore respiratory structures have evolved more than once within the echinoderms. Constraint analyses indicate that a single diplopore-bearing clade bearing the traditionally defined Glyptosphaeritida, Sphaeronitida, Asteroblastida is less parsimonious than multiple diplopore-bearing clades inferred by the unconstrained analysis. [ABSTRACT FROM AUTHOR]

The subgenus Setihenricia Chichvarkhin and Chichvarkhina, 2017 currently consists of 32 out of 94 species of sea stars in the genus Henricia Gray, 1840. However, only 10 have been molecularly ascertained as to the phylogenetic position. Based on seven mitochondrial gene markers (16S rRNA, COI, tRNA-Ala, tRNA-Leu, tRNA-Asn, tRNA-Gln, and tRNA-Pro), we performed molecular analyses to assess the monophyly of Setihenricia using 16 species of Henricia from Japanese waters, as well as other congeners for which sequences were available in public databases. The monophyletic Setihenricia was only weakly recovered because the support values were low, but we assume this taxon may be valid in consideration of a morphological key. Henricia kinkasana Hayashi, 1940 is herein identified as a new member of Setihenricia, based on molecular and morphological evidence. Sequences in the public databases previously identified as H. compacta (Sladen, 1889) and H. obesa (Sladen, 1889) were nested in the Setihenricia clade; these two species are also likely members of Setihenricia . Our morphological assessment yielded three findings: (1) the number of the rows of spines on each inferomarginal plate, being three or more, appears to be a useful diagnostic character for Setihenricia, as has been suggested in previous studies, (2) the degree of spine tapering turned out to be less relevant in diagnosing the subgenus than previously supposed, (3) two newly introduced characters--( i ) the width of adambulacral plates and ( ii ) the ratio of two measurements in ambulacral plates to the length of interradius--may be effective to distinguish members of this apparent Henricia subclade from other members of the genus. [ABSTRACT FROM AUTHOR]

Order Diplobathrida is a major clade of camerate crinoids spanning the Ordovician–Mississippian, yet phylogenetic relationships have only been inferred for Ordovician taxa. This has hampered efforts to construct a comprehensive tree of life for crinoids and develop a classification scheme that adequately reflects diplobathrid evolutionary history. Here, I apply maximum parsimony and Bayesian phylogenetic approaches to the fossil record of diplobathrids to infer the largest tree of fossil crinoids to date, with over 100 genera included. Recovered trees provide a framework for evaluating the current classification of diplobathrids. Notably, previous suborder divisions are not supported, and superfamily divisions will require significant modification. Although numerous revisions are required for families, most can be retained through reassignment of genera. In addition, recovered trees were used to produce phylogeny‐based estimates of diplobathrid lineage diversity. By accounting for ghost lineages, phylogeny‐based richness estimates offer greater insight into diversification and extinction dynamics than traditional taxonomy‐based approaches alone and provide a detailed summary of the ~150 million‐year evolutionary history of Diplobathrida. This study constitutes a major step toward producing a phylogeny of the Crinoidea and documenting crinoid diversity dynamics. In addition, it will serve as a framework for subsequent phylogeny‐based investigations of macroevolutionary questions. [ABSTRACT FROM AUTHOR]

Recent debates over the evolutionary relationships of early echinoderms have relied heavily on morphological evidence from the feeding ambulacral system. Eumorphocystis, a Late Ordovician diploporitan, has been a focus in these debates because it bears ambulacral features that show strong morphological similarity to early crinoid arms. Undescribed and well‐preserved specimens of Eumorphocystis from the Bromide Formation (Oklahoma, USA) provide new data illustrating that composite arms supported by a radial plate that bear a triserial arrangement of axial and extraxial components encasing a coelomic extension can also be found in blastozoans. Previous reports have considered these arm structures to be restricted to crinoids; these combined features have not been previously observed in blastozoan echinoderms. Phylogenetic analyses suggest that Eumorphocystis and crinoids are sister taxa and that shared derived features of these taxa are homologous. The evidence from the arms of Eumorphocystis suggests that crinoid arms were derived from a specialized blastozoan ambulacral system that lost feeding brachioles and strongly suggests that crinoids are nested within blastozoans. [ABSTRACT FROM AUTHOR]

Mitochondrial genomes provide an additional data source for nuclear phylogenomics. Recently, the taxonomic classification of Ophiuroidea was changed dramatically based on a molecular phylogenetic analysis that utilized a huge nuclear dataset of transcriptome and target-capture approaches. However, the mitochondrial genome analysis of Ophiuroidea was not conducted sufficiently in depth to allow comparison with current phylogenetic relationships, especially for the Ophintegrida. In this study, eight mitogenomes were newly reported in Amphilepidida and Ophiacanthida. Phylogenetic analyses were undertaken based on the nucleotide sequences of 13 protein coding genes (PCGs), 22 tRNA, and two rRNA, and the amino acid translated sequences of 13 PCGs. The results of the phylogenetic analysis suggested that the amino acid translated sequences were more useful than the nucleotide sequences for the analysis of higher phylogenetic relationships. The mitochondrial gene order of 13 PCGs and two rRNA in Amphilepidida and Ophiacanthida was relatively conserved. However, more complex gene orders occurred in the phylogeny of Amphilepidida and Ophiacanthida, including 22 tRNA. The results of our study suggest that Ophiuroidea has undergone more complex gene rearrangements than other classes of echinoderms. [ABSTRACT FROM AUTHOR]

The common sunstar, Crossaster papposus, belongs to the family Solasteridae whose ordinal classification has been unstable. Here, for the first time, we assembled and annotated the complete mitochondrial genome of the common sunstar, C. papposus Linnaeus, 1767. The circular genome of C. papposus is 16,335 bp in length and contains 13 protein-coding genes (PCGs), 22 transfer RNA (tRNA) genes, a control region, and large and small ribosomal subunits. The overall genomic structure and gene arrangement were identical to the reported mitochondrial genomes of sea star species, and a phylogenetic analysis of 13 PCGs recovers a closest relationship with the derived cluster of the paraphyletic order Valvatida. [ABSTRACT FROM AUTHOR]