Unveiling ophiuroid biodiversity across North Atlantic habitats via an integrative perspective.

The depths of the North Atlantic Ocean host a species-rich fauna providing heterogeneous habitats from thermal vent fields to cold-water coral reefs. With the increasing threat of destruction of deep-sea habitats due to human impacts, such as demersal fishing and the beginning of deep-sea mining, an analysis of the diversity and distribution of species is crucial for conservation efforts. Brittle stars occur in high biomasses, contributing to the biodiversity of the seafloor. Specimens were collected during several scientific expeditions to gain a more detailed insight into the brittle star diversity in the North Atlantic Ocean. An integrative approach to identify the species with DNA barcoding (mtCOI) in combination with morphological studies revealed 24 species. Most species have been previously identified in the North Atlantic, but sequences for 13 species are newly added to public repositories. Additionally, the MALDI-TOF-MS proteomic analysis was successfully applied for 197 specimens with known COI barcodes. Results are congruent with other molecular species delimitations demonstrating the functionality of proteomics for the identification of brittle stars. This dataset significantly expands our understanding of the taxonomic and genetic diversity of brittle stars and contributes to publicly available data. It emphasizes the importance of considering habitat heterogeneity for large scale patterns of biodiversity.

Riqueza y densidad de equinodermos en los arrecifes rocosos de Punta Amapala, El Salvador / Richness and density of echinoderms in the rocky reefs of Punta Amapala, El Salvador

Resumen Introducción: Los equinodermos son animales estrictamente acuáticos y de los grupos de invertebrados marinos cuya importancia ecológica es crucial para los ambientes en donde habitan. La característica más destacada es su simetría pentarradial. Estos organismos pueden distribuirse en una amplia variedad de ambientes oceánicos, y no ser exclusivamente habitantes de las formaciones arrecifales. Se han registrado el total de 1 539 especies, de las cuales 717 pertenecen a Pacífico y 60 de ellas se han registrado en El Salvador. Objetivo: El objetivo del trabajo fue registrar la riqueza y densidad de especies de equinodermos en arrecifes rocosos de Punta Amapala (La Unión). Métodos: El muestreo fue llevado a cabo en cinco sitios, dos a 3 m de profundidad y tres a 6 m. Para esto, se utilizó snorkeling y buceo autónomo, recorriendo dos transectos de banda de 30 x 1 m. Dentro de los transectos, se realizó la identificación (riqueza) y contabilización (abundancia) de los organismos. Resultados: La especie que presentó mayor densidad en los arrecifes de 3 m fue Echinometra vanbrunti (0.83 indv/m2), seguido por el ofiuroideo Ophiocomella alexandri (0.16 indv/m2) y Ophiocoma aethiops (0.16 indv/m2). Y en los arrecifes a 6 m fueron Ophiothela mirabilis (0.83 indv/m2), Ophionereis annulata (0.21 indv/m2), Ophiothrix rudis (0.08 indv/m2) y Ophiocomella alexandri (0.01 indv/m2). Conclusiones: Estos organismos, cumplen con diversos roles ecológicos, como ser bioindicadores de calidad de agua al tolerar ciertos cambios en las condiciones físico-químicas, así como cumplir el papel de hospederos de macrofauna por las cavidades que forman algunos erizos. Es por ello que los esfuerzos de muestreo deben incrementarse en estas zonas para monitorear la calidad y salud de los arrecifes rocosos, cuyo papel es clave, por ser junto con Los Cóbanos, los de mayor extensión en el país.

Abstract Introduction: Echinoderms are strictly aquatic animals and one of the groups of marine invertebrates whose ecological importance is crucial for the environments where they inhabit. The most outstanding characteristic is their pentaradial symmetry. These organisms can be distributed in a wide variety of oceanic environments, and are not exclusive inhabitants of reef formations. A total of 1 539 species have been registered, of which 717 belong to the Pacific and 60 of them have been registered for El Salvador. Objective: The objective of the work was to record the richness and density of echinoderm species in rocky reefs of Punta Amapala (La Unión). Methods: The sampling was conducted in five sites, two of them at 3 m of depth and three at 6 m. For this, snorkeling and autonomous diving were used, covering two band transects of 30 x 1 m. Within the transects, the identification (richness) and counting (abundance) of the organisms was carried out. Results: The species that presented the highest density at 3 m reefs was Echinometra vanbrunti (0.83 indv/m2), followed by the ophiuroid Ophiocomella alexandri (0.16 indv/m2) and Ophiocoma aethiops (0.16 indv/m2). And at the 6 m reefs were Ophiothela mirabilis (0.83 indv/m2), Ophionereis annulata (0.21 indv/m2), Ophiothrix rudis (0.08 indv/m2) and Ophiocomella alexandri (0.01 indv/m2). Conclusions: These organisms fulfill various ecological roles, such as being bioindicators of water quality by tolerating certain changes in physicochemical conditions and fulfilling the role of macrofauna hosts due to the cavities formed by some sea urchins. That is why sampling efforts must increase in these areas to monitor the quality and health of the rocky reefs, whose role is key, as they are, together with Los Cóbanos, the largest in the country.

Equinodermos de la colección biológica del Laboratorio de Ecología Molecular, Microbiología y Taxonomía (LEMITAX) de la Universidad de Guadalajara, México / Echinoderms of the biological collection of the Laboratorio de Ecología Molecular, Microbiología y Taxonomía (LEMITAX) of the Universidad de Guadalajara, Mexico

Resumen Introducción: Las colecciones biológicas representan los cimientos para el conocimiento y manejo de la biodiversidad de una región. No obstante, en México, y en particular en el Pacífico central mexicano (PCM), las colecciones regionales enfocadas en equinodermos, son escasas. La colección biológica del Laboratorio de Ecología Molecular, Microbiología y Taxonomía (LEMITAX) pertenece al Centro Universitario de Ciencias Biológicas y Agropecuarias de la Universidad de Guadalajara, Jalisco, México y sirve como referencia de la biodiversidad marina de la región. Objetivo: Dar a conocer la riqueza de equinodermos resguardada en la colección del LEMITAX. Métodos: Los organismos depositados en LEMITAX se han recolectado mediante buceo libre, SCUBA, y arrastres de fondo con dragas biológicas en diversas áreas del Pacífico mexicano. Los especímenes están preservados en húmedo (alcohol al 70 %; Asteroidea, Echinoidea, Holothuroidea y Ophiuroidea) o en seco (Asteroidea y Echinoidea). Resultados: Los especímenes provienen de los estados de Sinaloa, Nayarit, Jalisco y Colima, incluyendo dos áreas naturales protegidas, el Parque Nacional Isla Isabel y el Santuario de las islas e islotes de Bahía de Chamela. La colección dispone de 20 761 ejemplares de equinodermos distribuidos en 75 especies (10 asteroideos, 17 ofiuroideos, 17 equinoideos y 31 holoturoideos). El estado mejor representado es Jalisco (64 especies) seguido de Nayarit (31), Colima (20) y Sinaloa (11). Bahía de Chamela es la mejor representada (60), seguido de Isla Isabel (22). Se aportan 34 registros nuevos, la mayor contribución es para Bahía de Chamela con 14 registros nuevos, seguido del estado de Jalisco (siete), Nayarit (cuatro), Colima (tres) e Isla Isabel (tres). Para el PCM, se reporta por primera vez la presencia de Astropecten ornatissimus, Luidia phragma, Cucumaria crax y Holothuria (Cystipus) casoae, lo que actualiza su riqueza de equinodermos a 197 especies. Se amplía el intervalo de distribución batimétrica de Ophiactis simplex, Ophiocomella alexandri y Holothuria (Cystipus) casoae, así como el intervalo de distribución geográfica de Cucumaria crax. Conclusiones: Las colecciones biológicas de las universidades contribuyen de manera sustancial al conocimiento de la biodiversidad, como se refleja en la colección LEMITAX, cuya revisión resultó en la actualización de la riqueza de equinodermos de la región.

Abstract Introduction: The biological collections represent the foundation for the knowledge and management of the biodiversity of a region. However, regional collections focused on echinoderms are scarce in Mexico, particularly in the Central Mexican Pacific (CMP). The biological collection of the Laboratorio de Ecología Molecular, Microbiología y Taxonomía (LEMITAX) belongs to the Centro Universitario de Ciencias Biológicas y Agropecuarias of the Universidad de Guadalajara, Jalisco, Mexico and it aims to serve as a reference for the region's marine biodiversity. Objective: To state the richness of echinoderms in the LEMITAX collection. Methods: The organisms deposited at the LEMITAX have been collected by SCUBA, free-diving, and bottom trawls with biological dredges in different areas of the Mexican Pacific. The specimens are wet-preserved (70 % ethanol; Asteroidea, Echinoidea, Holothuroidea, and Ophiuroidea) or dried (Asteroidea and Echinoidea). Results: The specimens are from the states of Sinaloa, Nayarit, Jalisco, and Colima, including two natural protected areas (NPAs), the Isabel Island National Park and the sanctuary of the Islands and Islets of Bahía de Chamela. The collection has 20 761 specimens of echinoderms distributed in 75 species (10 asteroids, 17 ophiuroids, 17 echinoids, and 31 holothuroids). The best-represented state is Jalisco (64 species), followed by Nayarit (31), Colima (20), and Sinaloa (11). Concerning the NPAs, Chamela is the best represented (60), followed by Isabel Island (22). Thirty-four new records are added; the largest contribution is for Chamela, with 14 new records, followed by the state of Jalisco (seven), Nayarit (four), Colima (three), and Isabel Island (three). For the CMP, the presence of Astropecten ornatissimus, Luidia phragma, Cucumaria crax, and Holothuria (Cystipus) casoae, is reported for the first time, updating the echinoderm richness to 197 species. The bathymetric distribution range of Ophiactis simplex, Ophiocomella alexandri, and Holothuria (Cystipus) casoae is extended, as well as the geographic range of Cucumaria crax. Conclusions: The biological collections deposited in the universities contribute substantially to the knowledge of biodiversity, as reflected in the LEMITAX collection, whose revision resulted in the updating of the echinoderm richness in the region.

Crinoidea y Asteroidea (Echinodermata) del talud de la península de Yucatán, de la Colección Nacional de Equinodermos de México / Crinoidea and Asteroidea (Echinodermata) from the slope of the Yucatan Peninsula, Colección Nacional de Equinodermos, Mexico

Resumen Introducción: Las especies de las clases Crinoidea y Asteroidea se distribuyen en gran variedad de hábitats en todos los océanos desde la zona intermareal hasta grandes profundidades, existen pocos registros en aguas mexicanas, a profundidades superiores a los 200 m, los escasos registros existentes datan de documentos históricos de la literatura especializada y los ejemplares se resguardan en colecciones científicas extranjeras. Objetivo: Contribuir a la recopilación de información de los ejemplares resguardados en la Colección Nacional de Equinodermos (CNE) del Instituto de Ciencias del Mar y Limnología (ICML), UNAM y proveer un inventario de las especies de crinoideos y asteroideos que habitan en zonas profundas del talud de la Península de Yucatán, México. Métodos: Del año 2005 al 2014 se recolectaron ejemplares de crinoideos y asteriodeos en cuatro cruceros en el B/O "Justo Sierra" BIOREPES 1 y 2 y COBERPES 2 y 6. Se realizaron arrastres con red camaronera en 80 estaciones de muestreo. Resultados: El listado taxonómico comprende 1146 ejemplares, de los cuales 204 son crinoideos del orden Comatulida, distribuidos en tres familias tres géneros y tres especies y 942 asteroideos distribuidos en seis órdenes, 11 familias, 21 géneros y 28 especies. Se obtuvieron los registros nuevos para la CNE: Democrinus rawsonii y Atelecrinus balanoides en crinoideos. Astropecten alligator, Hymenaster modestus, Calyptraster personatus, Pteraster militarioides militarioides, Sclerasterias contorta en asteroideos. Con esta información se incrementan los registros de crinoideos y asteroideos (de profundidad mayor a 200 m) para los estados de Yucatán (16) y Quintana Roo (16).

Abstract Introduction: The species of the Crinoidea and Asteroidea classes are distributed in a wide variety of habitats in all oceans from the intertidal zone to great depths, there are few Mexican records in depths greater than 200 m, the few existing records date from historical documents in the specialized literature and the specimens are kept in foreign scientific collections. Objective: Contribute to the collection of information on the specimens kept in the Colección Nacional de Equinodermos (CNE) of Instituto de Ciencias del Mar y Limnología (ICML), UNAM and provide an inventory of the species of crinoids and asteroids that inhabit deep areas of the slope of the Yucatan Peninsula, Mexico. Methods: From 2005 to 2014, specimens of crinoids and asterioids were collected through four cruises in the B/O "Justo Sierra" BIOREPES 1 and 2 and COBERPES 2 and 6, shrimp net trawls were carried out at 80 sampling stations. Results: The taxonomic list includes 1146 specimens, of which 204 crinoids are of the order Comatulida, distributed in three families, three genera and three species and 942 asteroids distributed in six orders, 11 families, 21 genera and 28 species. New records were obtained for CNE: Democrinus rawsonii and Atelecrinus balanoides in crinoids. Astropecten alligator, Hymenaster modestus, Calyptraster personatus, Pteraster militarioides militarioides, Sclerasterias contorta in asteroids. With this information, the records of crinoids and asteroids (deeper than 200 m) increase for the states of Yucatan (16) and Quintana Roo (16).

Evaluating the reliability of DNA Barcoding for Central American Pacific shallow water echinoderms identification: a molecular taxonomy and database accuracy analysis / Evaluando la fiabilidad del Código de Barras de ADN para la identificación de equinodermos en aguas poco profundas del Pacífico de América Central: un análisis de taxonomía molecular y precisión de bases de datos

Abstract Introduction: Molecular divergence thresholds have been proposed to distinguish recently separated evolutive units, often displaying more accurate putative species assignments in taxonomic research compared to traditional morphological approaches. This makes DNA barcoding an attractive identification tool for a variety of marine invertebrates, especially for cryptic species complexes. Although GenBank and the Barcode of Life Data System (BOLD) are the major sequence repositories worldwide, very few have tested their performance in the identification of echinoderm sequences. Objective: We use COI echinoderm sequences from local samples and the molecular identification platforms from GenBank and BOLD, in order to test their accuracy and reliability in the DNA barcoding identification for Central American shallow water echinoderms, at genus and species level. Methods: We conducted sampling, tissue extraction, COI amplification, sequencing, and taxonomic identification for 475 specimens. The 348 obtained sequences were individually enquired with BLAST in GenBank as well as using the Identification System (IDS) in BOLD. Query sequences were classified depending on the best match result. McNemar's chi-squared, Kruskal-Wallis's and Mann-Whitney's U tests were performed to prove differences between the results from both databases. Additionally, we recorded an updated list of species reported for the shallow waters of the Central American Pacific. Results: We found 324 echinoderm species reported for Central American Pacific shallow waters. Only 118 and 110 were present in GenBank and BOLD databases respectively. We proposed 325 solved morphology-based identities and 21 provisional identifications in 50 putative taxa. GenBank retrieved 348 molecular-based identifications in 58 species, including twelve provisional identifications in tree taxa. BOLD recovered 170 COI identifications in 23 species with one provisional identification. Nevertheless, 178 sequences retrieved unmatched terms (in 34 morphology-based taxa). Only 86 sequences (25 %) were retrieved as correct identifications and 128 (37 %) as identification errors in both platforms. We include 84 sequences for eleven species not represented in GenBank and 65 sequences for ten species in BOLD Echinoderm COI databases. The identification accuracy using BLAST (175 correct and 152 incorrect identifications) was greater than with IDS engine (110 correct and 218 identification errors), therefore GenBank outperforms BOLD (Kruskal-Wallis = 41.625, df = 1, p < 0.001). Conclusions: Additional echinoderm sample references are needed to improve the utility of the evaluated DNA barcoding identification tools. Identification discordances in both databases may obey specific parameters used in each search algorithm engine and the available sequences. We recommend the use of barcoding as a complementary identification source for Central American Pacific shallow water echinoderm species.

Resumen Introducción: Se han propuesto los umbrales de divergencia molecular para distinguir unidades evolutivas recientemente separadas, que a menudo muestran asignaciones de especies putativas más precisas en la investigación taxonómica en comparación con los enfoques morfológicos tradicionales. Esto hace que los Códigos de Barras de ADN sean una herramienta de identificación atractiva para una variedad de invertebrados marinos, especialmente para complejos de especies crípticas. Aunque GenBank y Barcode of Life Data System (BOLD) son los principales repositorios de secuencias en todo el mundo, muy pocos han probado su desempeño en la identificación de secuencias de equinodermos. Objetivo: Utilizamos secuencias de equinodermos COI de muestras locales y las plataformas de identificación molecular de GenBank y BOLD, para probar su precisión y confiabilidad en la implementación de códigos de barras de ADN para equinodermos de aguas someras de Centroamérica, a nivel de género y especie. Métodos: Realizamos muestreo, extracción de tejido, amplificación de COI, secuenciación e identificación taxonómica de 475 especímenes. Las 348 secuencias obtenidas fueron consultadas individualmente con BLAST en GenBank así como utilizando el Sistema de Identificación (IDS) en BOLD. Las secuencias consultadas se clasificaron según el mejor resultado de coincidencia. Se realizaron las pruebas chi-cuadrado de McNemar, Kruskal-Wallis y U de Mann-Whitney para comprobar diferencias entre los resultados de ambas bases de datos. Además, registramos una lista actualizada de especies reportadas para las aguas someras del Pacífico Centroamericano. Resultados: Encontramos 324 especies de equinodermos reportadas para aguas someras (< 200 m) del Pacífico centroamericano. Sólo 118 y 110 estaban presentes en las bases de datos GenBank y BOLD respectivamente. Propusimos 325 identidades resueltas basadas en morfología y 21 identificaciones provisionales en 50 taxones putativos. GenBank recuperó 348 identificaciones de base molecular en 58 especies, incluidas doce identificaciones provisionales en tres taxones. BOLD recuperó 170 identificaciones de COI en 23 especies con una identificación provisional. Sin embargo, 178 secuencias recuperaron términos no coincidentes (en 34 taxones basados en morfología). Sólo 86 secuencias (25 %) se recuperaron como identificaciones correctas y 128 (37 %) como errores de identificación en ambas plataformas. Incluimos 84 secuencias para once especies no representadas en GenBank y 65 secuencias para diez especies ausentes en las bases de datos BOLD Echinoderm COI. La precisión de la identificación usando BLAST (175 identificaciones correctas y 152 incorrectas) fue mayor que con el motor IDS (110 correctas y 218 errores de identificación), por lo tanto, GenBank supera a BOLD (Kruskal-Wallis = 41.625, df = 1, p < 0.001). Conclusiones: Se necesitan muestras adicionales de equinodermos de referencia para mejorar la utilidad de las herramientas de identificación de códigos de barras de ADN evaluadas. Las discordancias de identificación en ambas bases de datos pueden obedecer a parámetros específicos utilizados en cada algoritmo de búsqueda y a las secuencias disponibles. Recomendamos el uso de códigos de barras como fuente de identificación complementaria para las especies de equinodermos de aguas someras del Pacífico centroamericano.

Earliest known ophiuroids from high palaeolatitude, southern Gondwana, recovered from the Pragian to earliest Emsian Baviaanskloof Formation (Table Mountain Group, Cape Supergroup) South Africa.

For the first time, ophiuroids have been found in South African strata predating the lowermost Bokkeveld Group. These comprise natural moulds and casts from two localities in the 'upper unit' of the Baviaanskloof Formation (Table Mountain Group). As a Pragian to earliest Emsian age has been inferred for this member, the new taxa comprise the earliest high-palaeolatitude ophiuroid records from southern Gondwana. Morphological analysis of the specimens revealed the presence of two distinct taxa. One is here described as Krommaster spinosus gen. et sp. nov., a new encrinasterid characterised by very large spines on the dorsal side of the disc, the ventral interradial marginal plates and the arm midlines. The second taxon is a poorly preserved specimen of Hexuraster weitzi, a cheiropterasterid previously described from the slightly younger Bokkeveld Group.

Continent-wide declines in shallow reef life over a decade of ocean warming.

Human society is dependent on nature1,2, but whether our ecological foundations are at risk remains unknown in the absence of systematic monitoring of species' populations3. Knowledge of species fluctuations is particularly inadequate in the marine realm4. Here we assess the population trends of 1,057 common shallow reef species from multiple phyla at 1,636 sites around Australia over the past decade. Most populations decreased over this period, including many tropical fishes, temperate invertebrates (particularly echinoderms) and southwestern Australian macroalgae, whereas coral populations remained relatively stable. Population declines typically followed heatwave years, when local water temperatures were more than 0.5 °C above temperatures in 2008. Following heatwaves5,6, species abundances generally tended to decline near warm range edges, and increase near cool range edges. More than 30% of shallow invertebrate species in cool latitudes exhibited high extinction risk, with rapidly declining populations trapped by deep ocean barriers, preventing poleward retreat as temperatures rise. Greater conservation effort is needed to safeguard temperate marine ecosystems, which are disproportionately threatened and include species with deep evolutionary roots. Fundamental among such efforts, and broader societal needs to efficiently adapt to interacting anthropogenic and natural pressures, is greatly expanded monitoring of species' population trends7,8.

Miniaturization during a Silurian environmental crisis generated the modern brittle star body plan.

Pivotal anatomical innovations often seem to appear by chance when viewed through the lens of the fossil record. As a consequence, specific driving forces behind the origination of major organismal clades generally remain speculative. Here, we present a rare exception to this axiom by constraining the appearance of a diverse animal group (the living Ophiuroidea) to a single speciation event rather than hypothetical ancestors. Fossils belonging to a new pair of temporally consecutive species of brittle stars (Ophiopetagno paicei gen. et sp. nov. and Muldaster haakei gen. et sp. nov.) from the Silurian (444-419 Mya) of Sweden reveal a process of miniaturization that temporally coincides with a global extinction and environmental perturbation known as the Mulde Event. The reduction in size from O. paicei to M. haakei forced a structural simplification of the ophiuroid skeleton through ontogenetic retention of juvenile traits, thereby generating the modern brittle star bauplan.

Echinobase: leveraging an extant model organism database to build a knowledgebase supporting research on the genomics and biology of echinoderms.

Echinobase (www.echinobase.org) is a third generation web resource supporting genomic research on echinoderms. The new version was built by cloning the mature Xenopus model organism knowledgebase, Xenbase, refactoring data ingestion pipelines and modifying the user interface to adapt to multispecies echinoderm content. This approach leveraged over 15 years of previous database and web application development to generate a new fully featured informatics resource in a single year. In addition to the software stack, Echinobase uses the private cloud and physical hosts that support Xenbase. Echinobase currently supports six echinoderm species, focused on those used for genomics, developmental biology and gene regulatory network analyses. Over 38 000 gene pages, 18 000 publications, new improved genome assemblies, JBrowse genome browser and BLAST + services are available and supported by the development of a new echinoderm anatomical ontology, uniformly applied formal gene nomenclature, and consistent orthology predictions. A novel feature of Echinobase is integrating support for multiple, disparate species. New genomes from the diverse echinoderm phylum will be added and supported as data becomes available. The common code development design of the integrated knowledgebases ensures parallel improvements as each resource evolves. This approach is widely applicable for developing new model organism informatics resources.

Catalogue of living crinoids (Echinodermata: Crinoidea) from Brazil.

Knowledge of living crinoids from Brazil is chiefly restricted to the contributions of Dr Luis Roberto Tommasi between the decades of 1950 and 1970. Herein we present an updated catalogue of the crinoids occurring along the Brazilian coast, including data on synonyms, type localities, type material, geographical distribution, bathymetric ranges, and occurrences of species along the coast. The data is based on an extensive revision of the literature and on a survey of several databases. A total of 20 species were catalogued, of which two are endemic (Phrixometra longipinna var. brasiliensis and Thaumatometra minutissima). According to information from the literature, P. longipinna var. brasiliensis represents an undescribed taxon. Although Brazil supports the greatest number of crinoid species in the Southwestern Atlantic, its crinoid fauna remains understudied and likely underestimated. Research on crinoids remains a subject of great potential, not only for taxonomy but also for other areas of knowledge.

Deep-sea Ophiuroidea (Echinodermata) from the Danish Galathea II Expedition, 1950-52, with taxonomic revisions.

The brittle star samples collected by the Danish cruise 'Galathea II' (1950-52) had not been studied completely. We examined the remaining deep-sea samples (400 m) and present the species inventory, discussing taxonomic issues in relation to recent phylogenetic data. About 235 samples were examined, over 9,300 individuals, from 67 species and 74 sampling localities, at depths of 425-5340 m. The species complex Amphiophiura bullata (Thomson, 1877) is morphologically not well separated, but molecular data suggest at least two clades. We propose to apply A. bullata for Atlantic and Australian populations and A. convexa (Lyman, 1878) for the North Pacific clade. We consider A. bullata pacifica Litvinova, 1971 conspecific with A. convexa. Ophiuroglypha irrorata (Lyman, 1878) and its subspecies are a polyphyletic group with unclear morphological boundaries. We propose to transfer Ophiura ossiculata (Koehler, 1908), Ophiura plana (Lütken Mortensen, 1899) and Ophiura scomba Paterson, 1985 to Ophiuroglypha. Silax Fell, 1962, until now synonymised with Amphioplus Verrill, 1899, is proposed as a valid genus with the species S. verrilli (Lyman, 1879), S. consors (Koehler, 1908), S. daleus (Lyman, 1879), S. patulus (Lyman, 1879) and S. magnificus (Koehler, 1907). Triplodia Turner Hallen, 2011 (a replacement name for Triodia A. M. Clark, 1970, due to homonymy) is synonymised with Silax, and possible specimens of its type species Triodia abdita A. M. Clark, 1970 are analysed. The species limits of Ophiacantha cosmica Lyman, 1879 and Ophiacantha pacifica Lütken Mortensen, 1899 could not be confirmed morphologically, but published molecular data suggest two clades. We propose to apply O. pacifica to the Northern/Central Pacific population and O. cosmica to the Southern Pacific/Antarctic population.

A diverse crinoid fauna (Echinodermata, Crinoidea) from the Lower Eocene of the Gulf of Languedoc (Corbières, Aude, southern France).

Detailed studies of the middle Ilerdian (lower Ypresian) blue marls of the Gulf of Languedoc (Corbières, Aude, France), belonging to the north Pyrenean foreland basin, have revealed a more abundant and diverse crinoid fauna than previously documented from the Lower Eocene. Here we describe five species of stalked crinoids in the family Rhizocrinidae (Cherbonniericrinus requiensis n. sp., ?Democrinus elongatus, Globulocrinus amphoraformis n. gen., n. sp., Pseudoconocrinus doncieuxi and P. lavadensis n. sp.), one barnacle-like species in the stalkless family Holopodidae (Holopus plaziati n. sp.) and a single feather star in the family Conometridae (Amphorometra atacica). Several sites have yielded brachials and rhizoids in addition to abundant aboral cups and columnals indicating in situ fossilisation of the dissociated skeletal elements. P. lavadensis n. sp. and ?D. elongatus have been collected only from outcrops located in the upper part of the middle blue marls, while P. doncieuxi predominates, with a wide range of morphological variation, in the lower blue marls. The fossil assemblage at the locality of Réqui near Montlaur differs from the others in the smaller size of most individuals and the presence of H. plaziati n. sp., C. requiensis n. sp., G. amphoraformis n. gen., n. sp., and P. doncieuxi suboblongus n. subsp. This particular association with high juvenile mortality corresponds to an unstable environment with mixed substrates (muddy and rocky). The crinoid fauna of the Corbières appears to be the most diverse of Early Eocene age known to date. With the fauna of the London Clay, a boreal formation of the same age, it shares the presence of the genera Democrinus and Amphorometra in an open-sea environment. A comparison with extant faunas allows the depth of deposition at the Ypresian sites in the Gulf of Languedoc to be estimated between from 100 and 140 meters.

Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species.

Describing ontogenetic morphological change is an important part of integrative taxonomy; still, most taxonomic studies are based only on adult characters. Here, we provide illustrations and a morphological description of the sea stars from Bahia, including ontogenetic and intraspecific variation, and identify taxonomic issues. A total of 293 specimens from different localities along the Bahia State coastline and comparative material from other localities were examined. Eighteen species (11 genera, eight families) of Asteroidea were identified; Astropectinidae was the most representative family. All species identified also occur in subtropical Brazilian waters and most species are from shallow water habitats with soft bottoms. Most observed ontogenetic variation was quantitative in nature, such as the increase in the number of spines in the furrow and of spinelets in the paxillae with specimen growth. Genera that require further taxonomic studies are Astropecten and Othilia, whose specimens are commonly misidentified in local studies. One third of the species from Bahia are currently classified as "Vulnerable" in the Brazilian Red List, but baseline data on the population biology of these species are scarce. An illustrated identification key to the 65 Brazilian sea star species is also provided. This taxonomic study will facilitate the identification of specimens occurring along the Brazilian coast and help scientists and policy makers to establish the conservation status of the Brazilian species.

New hosts and morphological data for the star pearlfish Carapus mourlani (Ophidiiformes: Carapidae) from collections made in the North Pacific coast of Costa Rica / Nuevos hospederos y datos morfológicos para el Pez perla estrella Carapus mourlani (Ophidiiformes: Carapidae) a partir de recolectas realizadas al norte de la costa Pacífico de Costa Rica

Introduction: The family Carapidae includes about 40 species of marine fishes distributed in coastal habitats worldwide. The family includes some free-living species, however, most of them are found as commensal inquilines or parasites of marine invertebrates, including several echinoderm species. In the Eastern Tropical Pacific, the biology and host use of the representatives of the Carapidae is relatively poorly known. Objective: The present study reports the occurrence of the Star pearlfish Carapus mourlani within three previously unknown hosts in the region: the sea stars Nidorellia armata, Phataria unifascialis, and the sea cucumber Stichopus horrens. Some ecological implications and considerations regarding such symbiotic relationships are raised and discussed. Additional morphometric and meristic data for the fish and the echinoderms are also provided and discussed. Methods: Echinoderms were collected, from 25 localities along the North Pacific coast of Costa Rica, and were carefully examined searching for commensal/parasitic fishes. Echinoderms and fishes were identified and characterized in accordance with the specialized literature. Results: A total of 497 echinoderms, including about 60 species, were collected and examined. Commensal/parasitic fish (a single species represented by 13 specimens) were found in three echinoderm specimens/species. Conclusions: The list of echinoderm hosts for this carapid fish, through its whole distribution range, rises to 12 species (six sea stars and six sea cucumbers) and that could be a consequence of its wide geographic distribution, its generalist feeding habits and opportunistic commensal behavior.

Introducción: La familia Carapidae está compuesta por alrededor de 40 especies de peces marinos, presentes en hábitats costeros alrededor de todo el mundo. Se incluyen dentro de esta familia algunas especies de vida libre, no obstante, la mayoría de carápidos son inquilinos oportunistas o parásitos de algunos grupos de invertebrados marinos, incluyendo varias especies de equinodermos. En el Pacífico Tropical Oriental (PTO), se sabe relativamente poco sobre la biología de estos peces, así como de las diversas asociaciones existentes y de los hospederos utilizados. Objetivo: En este trabajo reportamos la ocurrencia del Pez perla estrella Carapus mourlani en tres nuevos hospederos: las estrellas de mar Nidorellia armata y Phataria unifascialis, y el pepino de mar Stichopus horrens. También se discuten algunas implicaciones y consideraciones ecológicas relacionadas a estas asociaciones simbióticas. Además, se proveen y discuten datos morfométricos y merísticos de los peces y sus hospederos. Métodos: Se realizaron recolectas de equinodermos, entre 2018 y 2019, en un total de 25 localidades distribuidas al norte de la costa Pacífico de Costa Rica, los cuales fueron cuidadosamente revisados en búsqueda de peces comensales/parásitos. Los equinodermos y los peces fueron identificados y caracterizados de acuerdo con la literatura especializada. Resultados: Se recolectaron y examinaron un total de 497 equinodermos, incluyendo alrededor de 60 especies, de los cuales solo tres individuos-especies estuvieron ocupados por peces comensales/parásitos. Conclusiones: La lista de hospederos equinodermos de C. mourlani a lo largo de su ámbito de distribución geográfico llega a 12 especies (seis estrellas de mar y seis pepinos de mar), lo cual podría ser un reflejo de su amplia distribución geográfica, de sus hábitos de alimentación generalistas y de su comportamiento oportunista en lo relativo al uso de hospederos.

Expansion of the genus Massinium (Holothuroidea: Thyonidae) to the American continent and description of a new species / Expansión del género Massinium (Holothuroidea: Thyonidae) al continente americano y descripción de una nueva especie

Introduction: The genus Massinium includes 11 species, most from the Indo Pacific Ocean, and had not previously being reported from the American continent. Objective: To present the new record of the genus Massinium and describe a new species of this genus. Methods: Material collection was done by SCUBA-diving to depths of a maximum of 9 m. Results: Massinium ocumichoensis sp. nov. is described from seven specimens that extends the range of the genus to the American continent (Guerrero, Michoacan and Jalisco, Mexico). The new species lives in sandy-rocky substrata, from 2 to 16 m deep. It is distinguished from its congeneric species by a combination of morphological characters: mid-dorsal slightly bigger pseudobuttons (40-60 μm), and tables in the body wall. A taxonomic key for distinguishing the species of Massinium is provided. Conclusions: The geographic range of the genus Massinium is extended to the Mexican Pacific with M. ocumichoensis sp. nov.

Introducción : El género Massinium incluye 11 especies, la mayoría del Océano Indo-Pacífico, y no se había reportado previamente en el continente americano. Objetivo: Presentar el nuevo registro del género Massinium y describir una nueva especie de este. Métodos: La recolección de material se realizó mediante buceo a una profundidad máxima de 9 m. Resultados: Massinium ocumichoensis sp. nov. se describe a partir de siete ejemplares que extiende el rango de distribución del género al Continente Americano (Guerrero, Michoacán y Jalisco, México). La nueva especie vive en sustratos arenosos-rocosos, de 2 a 16 m de profundidad. Se distingue de sus especies congenéricas por una combinación de caracteres morfológicos: pseudobotones de la zona dorsal media ligeramente más grandes (40-60 μm) y tablas en la pared del cuerpo. Se proporciona una clave taxonómica para distinguir las especies de Massinium. Conclusiones: El rango geográfico del género Massinium se extiende al Pacífico mexicano con M. ocumichoensis sp. nov.

Los ofiuroideos (Echinodermata: Ophiuroidea) de la bahía de Chamela, Jalisco, México / Ophiuroids (Echinodermata: Ophiuroidea) from Chamela bay, Jalisco, Mexico

Introducción: La bahía de Chamela está ubicada en la parte central de la costa del estado de Jalisco, México y es un importante área natural protegida. Pese a ello, el estudio de los equinodermos en la zona es incipiente. Objetivo: Proporcionar una lista de verificación actualizada, descripciones de las especies y una clave taxonómica de los ofiuroideos de la bahía de Chamela. Métodos: Recolectamos en la bahía de 1998 a 2018, y también analizamos información de la literatura y especímenes en colecciones científicas internacionales. Resultados: se reportan 19 especies de ofiuroides para el área de estudio, distribuidas en dos órdenes, siete familias y 11 géneros. Ocho de estos corresponden a nuevos registros para el área y tres posiblemente representan nuevas especies. La biodiversidad de ofiuroideos en la bahía de Chamela representa el 15.2% de las especies de ofiuroides del Pacífico mexicano y el 73% de Jalisco. El número de equinodermos conocidos en la bahía de Chamela aumenta a 48 especies. Conclusiones: Se presenta el listado más completo de la clase Ophiuroidea en la bahía de Chamela, Jalisco. La información proporcionada se puede utilizar para otras áreas del Pacífico oriental. La bahía de Chamela es importante en términos de riqueza de especies debido a su heterogeneidad ambiental y numerosas islas.

Introduction: Chamela bay is located in the central portion of the coast from the Jalisco state, Mexico and is an important protected natural area. Despite this, the study of echinoderms in the area is incipient. Objective: To provide an updated checklist, descriptions of the species and a taxonomic key of the ophiuroids from Chamela bay. Methods: We collected specimens from 1998 to 2018, and also analyzed information from the literature and specimens deposited at international scientific collections from Chamela bay. Results: 19 species of ophiuroids are reported for the study area, distributed in two orders, seven families, and 11 genera. Seven of these correspond to new records for the area, one to a new record for Jalisco state, and two possibly represent new species. The biodiversity of ophiuroids in Chamela bay represents 15.2 % of ophiuroid species from the Mexican Pacific and 73 % from Jalisco. The number of known echinoderms in Chamela bay is increased to 48 species. Conclusions: The most complete checklist of the class Ophiuroidea in Chamela bay, Jalisco, is presented. The information provided can be used for other areas of the eastern Pacific. Chamela bay is important in terms of species richness due to its environmental heterogeneity and numerous islands.

Echinoderms (Echinodermata) from the Central Mexican Pacific / Equinodermos (Echinodermata) del Pacífico Central mexicano

Introduction: The echinoderms from the Central Mexican Pacific are of high scientific interest and, prior to this present work, there was a lack of basic information that included incomplete checklists with inconsistencies in systematics and spatial distribution. Objective: To provide a historical review, and an updated checklist with a more complete richness of echinoderms for each state and island of the region. Methods: A checklist was elaborated based on an exhaustive literature search of the Echinodermata, and was complemented with taxonomical revisions of Ophiuroidea scientific collections. All the geographical coordinates of the records were validated. Results: The region harbors 187 species of Echinodermata: three Crinoidea, 35 Asteroidea, 67 Ophiuroidea, 32 Echinoidea, and 50 Holothuroidea. We detected 52 records in the literature that must be considered as invalid and five as doubtful. We provide 16 new records of Ophiuroidea from different states and islands; of them, four are new records for the region. Jalisco presented the highest number of species (84), followed by the coast of Nayarit (74), Michoacán (63), and Colima (55); among the islands, Revillagigedo showed the major number of species (85) followed by Marías (81), Marietas (48), and Isabel (44). Conclusions: The numbers of species known in the region are mostly related to both sampling effort and environmental characteristics that promote high biodiversity. The Central Mexican Pacific is an oceanographic region with mixed conditions from the North and South of the Mexican Pacific, and therefore, with a biogeographical importance reflected in its species richness.

Introducción: Los equinodermos del Pacífico Central mexicano poseen un alto interés científico, pero anterior al presente trabajo existía un vacío en la información básica como listados de especies incompletos con inconsistencias sistemáticas y de distribución espacial. Objetivo: Proporcionar una revisión histórica y un listado de especies actualizado con la riqueza completa de equinodermos para cada estado e islas de la región. Métodos: Se elaboró un listado de especies basado en una búsqueda exhaustiva de Echinodermata, y fue complementada con la revisión taxonómica de Ophiuroidea de colecciones científicas. Todas las coordenadas geográficas de los registros fueron validadas. Resultados: La región posee 187 especies de Echinodermata: tres Crinoidea, 35 Asteroidea, 67 Ophiuroidea, 32 Echinoidea, y 50 Holothuroidea. Detectamos 52 registros en la literatura considerados como inválidos y cinco como dudosos. Proveemos 16 nuevos registros de Ophiuroidea para diversos estados e islas; de estos, cuatro son nuevos registros para la región. Jalisco presentó el mayor número de especies (84), seguido por las costas de Nayarit (74), Michoacán (63), y Colima (55); entre las islas, Revillagigedo presentó el mayor número de especies (85) seguido por Marías (81), Marietas (48), e Isabel (44). Conclusiones: Los números de especies de la región están mayormente relacionados tanto con el esfuerzo de muestreo como con características ambientales que promueven una alta biodiversidad. El Pacífico Central mexicano es una región oceanográfica con condiciones mixtas del norte y sur del Pacífico mexicano, y por lo tanto con una importancia biogeográfica reflejada en la riqueza de la región.

Morphological variability of recent species of the order Cassiduloida (Echinodermata: Echinoidea) of Mexico / Variabilidad morfológica de especies recientes del orden Cassiduloida (Echinodermata: Echinoidea) de México

Introduction: In Mexico, there are two recorded living species of Cassiduloida: Cassidulus caribaearum and Rhyncholampas pacifica. Most of the taxonomic studies on cassiduloids have used external morphology, pedicellariae and morphometric characters; however, the intraspecific variation of quantitative and qualitative characters has been poorly evaluated. Objective: To compare the basic morphology of R. pacifica and C. caribaearum. Methods: We examined a total of 2 158 specimens of R. pacifica and C. caribaearum, selecting 50 to evaluate shape and size with linear regression and Principal Component analysis. We selected an additional 62 specimens per species to identify significant character correlations and morphological groups within species. Results: There is a direct relationship between Test length and Test width. Test height/Test width, and Total length (oral view)/Distance from the ambitus to the peristome apex, are the two main ratios to distinguish both species. C. caribaearum is more dorsoventrally compressed and has a round peristome base; versus R. pacifica has a tall and triangular one. There are four morphological groups of C. caribaearum and two groups for R. pacifica. Conclusions: These two species can be distinguished with reliable morphological characters, in which peristome shape suggests that R. pacifica is more adapted to burrowing deeper into certain types of substratum.

Introducción: En México, hay dos especies vivientes registradas de Cassiduloida: Cassidulus caribaearum y Rhyncholampas pacifica. La mayoría de los estudios taxonómicos sobre casiduloides han utilizado morfología externa, pedicelarios y caracteres morfométricos; sin embargo, la variación intraespecífica de caracteres cuantitativos y cualitativos ha sido poco evaluada. Objetivo: Comparar la morfología básica de R. pacifica y C. caribaearum. Métodos: Examinamos un total de 2 158 especímenes de R. pacifica y C. caribaearum, seleccionando 50 para evaluar la forma y el tamaño con regresión lineal y análisis de componentes principales. Seleccionamos 62 especímenes adicionales por especie para identificar correlaciones significativas de caracteres y grupos morfológicos dentro de las especies. Resultados: Existe una relación directa entre la longitud de la testa y el ancho de la testa. La Altura de la testa / Anchura de la testa y la Longitud total (vista oral) / Distancia desde el ambitus hasta el ápice del peristoma, son las dos proporciones principales para distinguir ambas especies. C. caribaearum está más comprimido dorsoventralmente y tiene una base del peristoma redonda; versus R. pacifica que tenía una alta y triangular. Hay cuatro grupos morfológicos de C. caribaearum y dos grupos de R. pacifica. Conclusiones: Estas dos especies se pueden distinguir con caracteres morfológicos confiables, en los que la forma del peristoma sugiere que R. pacifica está más adaptada para excavar más profundamente en ciertos tipos de sustratos.

Biodiversidad de ofiuroideos (Echinodermata: Ophiuroidea) del talud de la Península de Yucatán, México / Biodiversity of ophiuroids (Echinodermata: Ophiuroidea) from the slope of the Yucatan Peninsula, Mexico

Introduction: The Ophiuroidea class is the most diverse of the Phylum Echinodermata, with 2 064 described species, which can be found from intertidal zones to abyssal depths, but the fauna of Yucatan peninsula needs more study. Objective: To describe the diversity of ophiuroids in the Yucatan continental slope. Methods: Information was compiled from four oceanographic surveys carried out in August 2005, June 2007 (BIOREPES 1 and 2: BRP1 and BRP2), April 2011 and August 2014 (COBERPES 2 and 6: CBP2 and CBP6). The specimens were obtained by trawling with a shrimp net in soft substrates and with a skimmer-type trawl dredge at a depth range of 200 to 1 071 m off the states of Yucatán and Quintana Roo, Mexico. Results: We found members of two superorders, five orders, six suborders, five superfamilies, 18 families, 28 genera and 43 species. Conclusions: The northeast zone Yucatan Peninsula has at least 43 species, of which 19 are new records for the area.

Introducción: La clase Ophiuroidea es la más diversa del Phylum Echinodermata, con 2 064 especies descritas, que se pueden encontrar desde zonas intermareales hasta profundidades abisales, pero la fauna de la península de Yucatán necesita más estudio. Objetivo: Describir la diversidad de ofiuroideos en el talud continental de Yucatán. Métodos: Se recopiló información de cuatro estudios oceanográficos llevados a cabo en agosto de 2005, junio de 2007 (BIOREPES 1 y 2: BRP1 y BRP2), abril de 2011 y agosto de 2014 (COBERPES 2 y 6: CBP2 y CBP6). Los especímenes se obtuvieron mediante la pesca de arrastre con una red camaronera en sustratos blandos y con una draga de arrastre tipo skimmer en un rango de profundidad de 200 a 1 071 m frente a los estados de Yucatán y Quintana Roo, México. Resultados: Encontramos miembros de dos superórdenes, cinco órdenes, seis subórdenes, cinco superfamilias, 18 familias, 28 géneros y 43 especies. Conclusiones: La zona noreste de la Península de Yucatán tiene al menos 43 especies, de las cuales 19 son nuevos registros para el área.

Diversity and biostratigraphy of the late Oligocene-late Miocene sand dollars (Echinoidea: Scutelliformes) of Argentina and Uruguay / Diversidad y bioestratigrafía de las galletas de mar (Echinoidea: Scutelloida: Scutellifomes) del Oligoceno tardío-Mioceno tardío de Argentina y Uruguay

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

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