Fossil evidence for the ancient link between clonal fragmentation, six-fold symmetry and an epizoic lifestyle in asterozoan echinoderms.

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.

Regeneration of the digestive system in the crinoid Lamprometra palmata (Mariametridae, Comatulida).

The morphology and regeneration of the digestive system and tegmen after autotomy of the visceral mass in the crinoid Lamprometra palmata (Clark 1921) was studied. The gut has a five-lobed shape and is covered by a tegmen. The tegmen consists of epidermis and underlying connective tissue. The digestive tube can be divided into three parts: esophagus, intestine, and rectum. At 6 h post-autotomy, the calyx surface is covered by a layer of amoebocytes and juxtaligamental cells (JLCs). At 14-18 h, post-autotomy transdifferentiation of JLCs begins and give rise to the epidermis and cells of digestive system. On days 1-2 post-autotomy, JLCs undergo the mesenchymal-epithelial transition. Some JLCs turn into typical epidermal cells, while other JLCs form small closed epithelial structures that represent the gut anlage. On day 4 post-autotomy, the animals have a mouth opening and a small anal cone. On day 7 post-autotomy, the visceral mass and the digestive system become fully formed but are smaller than normal. A 24-h exposure of L. palmata individuals to a 10-7 M colchicine solution did not slow down regeneration, and the timing of gut formation was similar to that in the control animals. We conclude that JLCs are the major cell source for gut and epidermis regeneration in L. palmata. The main mechanisms of morphogenesis are cell migration, mesenchymal-epithelial transition, and transdifferentiation.

Cambrian edrioasteroid reveals new mechanism for secondary reduction of the skeleton in echinoderms.

Echinoderms are characterized by a distinctive high-magnesium calcite endoskeleton as adults, but elements of this have been drastically reduced in some groups. Herein, we describe a new pentaradial echinoderm, Yorkicystis haefneri n. gen. n. sp., which provides, to our knowledge, the oldest evidence of secondary non-mineralization of the echinoderm skeleton. This material was collected from the Cambrian Kinzers Formation in York (Pennsylvania, USA) and is dated as ca 510 Ma. Detailed morphological observations demonstrate that the ambulacra (i.e. axial region) are composed of flooring and cover plates, but the rest of the body (i.e. extraxial region) is preserved as a dark film and lacks any evidence of skeletal plating. Moreover, X-ray fluorescence analysis reveals that the axial region is elevated in iron. Based on our morphological and chemical data and on taphonomic comparisons with other fossils from the Kinzers Formation, we infer that the axial region was originally calcified, while the extraxial region was non-mineralized. Phylogenetic analyses recover Yorkicystis as an edrioasteroid, indicating that this partial absence of skeleton resulted from a secondary reduction. We hypothesize that skeletal reduction resulted from lack of expression of the skeletogenic gene regulatory network in the extraxial body wall during development. Secondary reduction of the skeleton in Yorkicystis might have allowed for greater flexibility of the body wall.

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.

The evolutionary relationship between arm vertebrae shape and ecological lifestyle in brittle stars (Echinodermata: Ophiuroidea).

Ophiuroidea are one of the most diverse classes among extant echinoderms, characterized by their flexible arms composed of a series of ossicles called vertebrae, articulating with each other proximally and distally. Their arms show a wide range of motion, important for feeding and locomotion, associated with their epizoic and non-epizoic lifestyles. It remains to be explored to what degree the phenotypic variation in these ossicles also reflects adaptations to these lifestyles, rather than only their phylogenetic affinity. In this study, we analyzed the 3D shape variation of six arm vertebrae from the middle and distal parts of an arm in 12 species, belonging to the intertidal, subtidal and bathyal zones and showing epizoic and non-epizoic behaviors. A PERMANOVA indicated a significant difference in ossicle morphology between species and between lifestyles. A principal component analysis showed that the morphology of epizoic ophiuroids is distinct from non-epizoic ones; which may reflect variation in arm function related to these different lifestyles. The Phylogenetic MANOVA and phylogenetic signal analysis showed that shape variation in the vertebral articulation seems to reflect ecological and functional adaptations, whereas phylogeny controls more the lateral morphology of the vertebrae. This suggests a convergent evolution through ecological adaptation to some degree, indicating that some of these characters may have limited taxonomic value.

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.

Problems and puzzles in echinoderm demography / Problemas y acertijos en la demografía de los equinodermos

Introduction: There are problems and puzzles in understanding reproduction, growth and mortality in echinoderm life cycles. Objective: Explore problems and puzzles in life cycles that are important and challenging. Methods: The literature is used to elucidate problems associated with all life stages. Results: Sources of larvae that settle at a site are explored using oceanographic modelling and genetic methods. There are few studies that have estimated larval mortality in the plankton under field conditions and results differ from experimental results or patterns of settlement. In a small number of studies, mortality rate of newly settled larvae appears to change rapidly as individuals grow. There are problems measuring size, and measurement bias that interferes with many tagging methods used to estimate growth. There also are problems with the use of natural growth lines and commonly used software to estimate both growth and mortality from size-frequency data. An interesting puzzle is that echinoderms may show negative senescence with mortality rate decreasing with size. There is a problem in fertilization success based on density so there should not be rare species where sexes are separate with free spawning of gametes yet there seem to be rare echinoderms. Conclusions: All parts of echinoderm life cycles provide problems and puzzles that are important and challenging.

Introducción: Existen problemas y acertijos en la comprensión de la reproducción, el crecimiento y la mortalidad en los ciclos de vida de los equinodermos. Objetivo: Explorar los problemas y acertijos en los ciclos de vida que son importantes y desafiantes. Métodos: La literatura es usada para dilucidar los problemas asociados con todas las etapas de vida. Resultados: Las fuentes de larvas que se asientan en un sitio se exploran usando modelos oceanográficos y métodos genéticos. Existen pocos estudios que han estimado la mortalidad larval del plancton bajo condiciones de campo y los resultados difieren de los resultados experimentales o los patrones de asentamiento. En un número pequeño de estudios, la tasa de mortalidad de las larvas recién asentadas parece cambiar rápidamente a medida que los organismos crecen. Existen problemas para medir tamaños y el sesgo de medición interfiere con el uso de muchos métodos de marcado para estimar el crecimiento. También hay problemas con el uso de las líneas de crecimiento natural y con los programas comúnmente usados para estimar tanto el crecimiento como la mortalidad a partir de datos de frecuencia de tamaño. Un acertijo interesante es que los equinodermos pueden mostrar senescencia negativa con una tasa de mortalidad que disminuye con el tamaño. Existe un problema con el éxito de la fertilización basado en la densidad, por lo que no debería haber especies raras cuando los sexos están separados y existe un desove libre de gametos, sin embargo, parece haber equinodermos raros. Conclusiones: Todas las partes de los ciclos de vida de los equinodermos proveen problemas y acertijos que son importantes y desafiantes.

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.

Heart urchins from the depths: Corparva lyrida gen. et sp. nov. (Palaeotropidae), and new records for the southwestern Atlantic Ocean / Erizos corazón de las profundidades: Corparva lyrida gen. et sp. nov. (Palaeotropidae) y nuevos registros para el Océano Atlántico sudoccidental

Introduction: Sea urchins in the order Spatangoida are the most diverse group of extant echinoids. Objective: Describe a new genus and species of Spatangoida from abyssal depths, and add new records for known species. Methods: Specimens were collected during several cruises at different areas of the southwestern Atlantic Ocean (SWAO), among 37-55° S latitude at depths ranging from 55 to 3 000 m. We present morphological and ultrastructure analyses. Results: Corparva lyrida gen. et sp. nov. (Palaeotropidae) is described from the Mar del Plata Canyon on the Argentine continental slope (2 950 m depth), the first record of this family from Argentina. Corparva gen. nov. differs in having an apical system semi-ethmolytic, and labrum reaching to rear part of second adjacent ambulacral plate. We also report the northernmost distribution and deepest record for Brisaster moseleyi (38° S latitude, 2 212 m depth), the northward extension of the distribution range of Tripylus excavatus (39° S latitude, 74 m depth), and the first record of Abatus philippii and Abatus agassizii at the Burdwood Bank/MPA Namuncurá. Conclusions: The present work brings novel and updated data about the diversity and distribution of spatangoids from the SWAO, including the description of C. lyrida gen. et sp. nov., and new records of species. This shows how much remains to be known about the diversity and distribution of heart urchins in the SWAO, especially from the deep-sea.

Introducción: Los erizos de mar del orden Spatangoida son el grupo más diverso de equinoideos recientes. Objetivo: Describir un nuevo género y una nueva especie de Spatangoida de profundidades abisales, y reportar nuevos registros para especies conocidas. Métodos: Los ejemplares fueron recolectados durante varias expediciones a diferentes áreas del Océano Atlántico sudoccidental (OAS), entre las latitudes 37-55° S y abarcando profundidades desde 55 a 3 000 metros. Presentamos análisis morfológicos y de ultraestructura. Resultados: Corparva lyrida gen. et sp. nov. (Palaeotropidae) fue descripta para el cañón submarino Mar del Plata en el talud continental de Argentina (2 950 m de profundidad), el primer registro de esta familia para Argentina. Corparva gen. nov. difiere en tener un sistema apical semi-etmolítico y labrum que llega a la parte posterior de la segunda placa ambulacral adyacente. También informamos la distribución más septentrional y el registro más profundo para Brisaster moseleyi (latitud 38° S, 2 212 m de profundidad), la extensión hacia el norte del rango de distribución de Tripylus excavatus (latitud 39° S, 74 m de profundidad) y el primer registro de Abatus philippii y Abatus agassizii en el Banco Burdwood/AMP Namuncurá. Conclusiones: El presente trabajo aporta datos novedosos y actualizados sobre la diversidad y distribución de erizos de mar espatangoideos del OAS, incluyendo la descripción de C. lyrida gen. et sp. nov., y nuevos registros para especies conocidas. Esto muestra cuánto queda por conocer sobre la diversidad y distribución de los erizos corazón en el OAS, especialmente de las profundidades marinas.

Área de conservación Guanacaste Echinoderms, North Pacific of Costa Rica / Equinodermos del Área de Conservación Guanacaste, Pacífico Norte de Costa Rica

Introduction: The study of the marine diversity of the North Pacific of Costa Rica began with isolated foreign expeditions in the 1930s and was systematically developed in the mid-1990s by the Center for Research in Marine Sciences and Limnology, Universidad de Costa Rica, as consequence there are now a total of 1 479 reported species in this region. Objective: Present an update to the echinoderm richness of the Guanacaste Conservation Area. Methods: We sampled 25 localities exhaustively and estimated similarity between sites based on the family richness and environmental heterogeneity. Results: We found 61 taxa, which represent 26 % of the echinoderm reported species for the country's Pacific coast. Of these, 43 species are new records for the Guanacaste Conservation Area, and seven for Costa Rica and Central American Pacific coasts. We found three morpho-species that do not match to available descriptions of the Eastern Tropical Pacific echinoderm species. We also found the holothuroid Epitomapta tabogae, and the ophiuroid Ophioplocus hancocki, previously thought endemic to Panama and the Galapagos Islands, respectively. The proximity of the sampled sites and the redundancy of certain families may explain why we did not find important differences among localities. Conclusions: The echinoderm richness of this conservation area is at least 20 % higher than previously reported, reaching similar levels to those in other high diversity sites of the Eastern Tropical Pacific.

Introducción: El estudio de la diversidad marina del Pacífico Norte de Costa Rica inició con expediciones extranjeras aisladas en la década de 1930, y fue desarrollado sistemáticamente a mediados de la década de 1990 por el Centro de Investigaciones en Ciencias del Mar y Limnología de la Universidad de Costa Rica, como consecuencia ahora se reporta un total de 1 479 especies en esta región. Objetivo: Presentar una actualización de la riqueza de equinodermos del Área de Conservación Guanacaste. Métodos: Realizamos muestreos exhaustivos en 25 localidades y estimamos la similitud entre sitios con base en la riqueaza de familias y la heterogeneidad ambiental. Resultados: Encontramos 61 taxa, que representan el 26% de las especies reportadas para la costa pacífica del país. De estas, 43 especies son nuevos registros para el Área de Conservación Guanacaste y siete para las costas de Costa Rica y el Pacífico centroamericano. Tres morfoespecies no coinciden con las descripciones disponibles para las especies del Pacífico Tropical Oriental. Por último, hallamos un ejemplar del holoturoideo Epitomapta tabogae y otro del ofiuroideo Ophioplocus hancocki, considerados endémicos para Panamá y las Islas Galápagos respectivamente. La proximidad entre los sitios muestreados y la redundancia de ciertas familias pueden explicar por qué no se encontraron diferencias entre las localidades. Conclusiones: La riqueza de equinodermos de esta área de conservación es al menos 20% mayor que la reportada anteriormente, alcanzando niveles similares a los de otros sitios de alta diversidad del Pacífico Tropical Oriental.

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.

First record of Ubaghsicystis (Eocrinoidea - Echinodermata) from the Cambrian (Miaolingian, Wuliuan) of Sonora, Mexico: Biostratigraphical and paleoecological considerations / Primer registro de Ubaghsicystis (Eocrinoidea - Echinodermata) del Cámbrico (Miaolingio, Wuliuan) de Sonora, México: Consideraciones bioestratigráficas y paleoecológicas

Introduction: The present work constitutes the first record of Ubaghsicystis Gil-Cid & Domínguez-Alonso (Echinodermata-Eocrinoidea) in the El Gavilán Formation, exposed in the El Sahuaral Hill, a new Cambrian locality of central Sonora, Mexico. The El Gavilán Formation is a sedimentary sequence mainly made up of shale interbedded with limestone, with abundant invertebrate fossils, deposited in an open shelf marine environment. Objective: The main objective of this research work is to describe the species Ubaghsicystis cf. U. segurae for the first time in Mexico, in addition to biostratigraphic and paleoecological considerations. Methods: A composite stratigraphic section of the El Gavilán Fm. was measured in central Sonora, where samples corresponding to eocrinods were collected, the material was examined in the Microscope Leica MZ10. Results: The biotic association is composed of reticulosan sponges, chancelloriids (Chancelloria eros, Allonnia tintinopsis, Archiasterella sp.), hyolithids, brachiopods, trilobites (Quadragnostus depressa, Peronopsis sp., Tonkinella valida, and Elrathina sp.), and echinoderms (Ubaghsicystis cf. U. segurae). Regarding the age, the trilobite association establishes a chronostratigraphic position from the middle Cambrian (Miaolingian, Wuliuan), Altiocculus Subzone, Ehmaniella Zone. The material studied is constituted by isolated echinoderm plates, which probably represent various cycles of theca plates assigned to Ubaghsicystis cf. U. segurae. The genus Ubaghsicystis has been scarcely recorded in a few Cambrian regions (e.g., Burgess Shale in Canada and Los Barrios de Luna in Spain), so that the information about this taxon is limited. Conclusions: The Cambrian succession of Sonora, where Ubaghsicystis occurred, was part of the border from the Laurentia craton during this period, located near the Paleoequator during this period, where ideal conditions for increasing diversity and abundance of species were developed, including echinoderms.

Introducción: El presente trabajo constituye el primer registro en México de Ubaghsicystis Gil-Cid & Domínguez-Alonso (Echinodermata-Eocrinoidea) procedente de la Formación El Gavilán, la cual aflora en el Cerro El Sahuaral, una nueva localidad del Cámbrico ubicada en la parte central del estado de Sonora, México. La Formación El Gavilán es una secuencia sedimentaria depositada en un ambiente marino de plataforma exterior, la cual consiste principalmente de lutita intercalada con caliza, conteniendo abundantes fósiles de invertebrados. Objetivo: El principal objetivo del presente trabajo de investigación es describir la especie Ubaghsicystis cf. U. segurae por primera vez para México, así como indicar las consideraciones bioestratigráficas y paleoecológicas. Métodos: Una sección estratigráfica compuesta fue medida para la Formación El Gavilán en Sonora central, donde muestras correspondientes a eocrinoides fueron colectadas, el material fue examinado en el Microscopio Leica MZ10. Resultados: La asociación biótica está conformada por esponjas reticulosas, chancellóridos (Chancelloria eros, Allonnia tintinopsis, Archiasterella sp.), hiolítidos, braquiópodos, trilobites (Quadragnostus depressa, Tonkinella valida, Elrathina sp.), y equinodermos (Ubaghsicystis cf. U. segurae). La asociación de trilobites confiere a estas rocas una edad del Cámbrico medio (Miaolingiano, Wuliuano), Subzona de Altiocculus, Zona de Ehmaniella. El material estudiado está constituido por placas aisladas y desarticuladas las cuales probablemente representen varios ciclos de placas de la teca, las cuales son asignadas a Ubaghsicystis cf. U. segurae. El género Ubaghsicystis ha sido documentado previamente en otras localidades del Cámbrico como Burgess Shale en Canadá y Los Barrios de Luna en España, para los cuales existen escasos registros. Conclusiones: La sucesión cámbrica de Sonora, donde ocurre Ubaghsicystis, formó parte del borde del cratón de Laurentia durante este período, el cual se localizaba cerca del Paleo Ecuador, donde se generaron las condiciones necesarias para una gran diversidad y abundancia de especies, incluyendo equinodermos.

Embryonic and larval development is conditioned by water temperature and maternal origin of eggs in the sea urchin Arbacia dufresnii (Echinodermata: Echinoidea) / El desarrollo embrionario y larvario está condicionado por la temperatura del agua y el origen materno de los huevos en el erizo de mar Arbacia dufresnii (Echinodermata: Echinoidea)

Introduction: Embryonic and larval development in sea urchins is highly dependent on maternal nutritional status and on the environmental conditions of the seawater. Objective: To compare the development of Arbacia dufresnii in two different water temperatures and in progeny with varying maternal origins. Methods: We induced A. dufresnii females and males from Nuevo Gulf to spawn, collected the eggs of each female individually (progeny), separated them into two seawater temperatures (12 and 17 °C), and fertilized them. We recorded the percentage of fertilized eggs and embryos per developmental stage according to time, temperature and progeny. We measured larval growth by total length (TL) and midline body length (M) according to time post fecundation (DPF), temperature, and progeny. Results: Temperature did not affect fertilization, but embryo development was faster and more synchronized in the high temperature treatment. The generalized linear models indicate that embryo development depends on a quadruple interaction between the embryonic stage, time (h), seawater temperature and progeny. Larval growth was faster, producing larger larvae at the highest temperature. Larval growth depends on a triple interaction between time (DPF), seawater temperature and progeny. Conclusions: We found a temperature and progeny impact during embryonic and larval development and, in both cases, these factors generate a synergistic effect on developmental timing and larval size. This probably provides a survival advantage as a more rapid speed of development implies a decrease in the time spent in the water column, where the sea urchins are vulnerable to biotic and abiotic stressors.

Introducción: El desarrollo embrionario y larvario de los erizos de mar depende en gran medida del estado nutricional materno y de las condiciones ambientales del agua de mar. Objetivo: Comparar el desarrollo de Arbacia dufresnii en dos temperaturas de agua diferentes y en progenies con diferentes orígenes maternos. Métodos: Indujimos a las hembras y machos de A. dufresnii del Golfo Nuevo a desovar, recolectamos los huevos de cada hembra individualmente (progenie), los separamos en dos temperaturas de agua de mar (12 y 17 ° C) y los fertilizamos. Registramos el porcentaje de óvulos fecundados y el porcentaje de embriones por etapa de desarrollo según tiempo, temperatura y descendencia. Medimos el crecimiento larvario según la longitud total (TL) y la longitud corporal de la línea media (M) de acuerdo con el tiempo en días post fecundación, la temperatura y la progenie. Resultados: La temperatura no afectó la fertilización, pero el desarrollo del embrión fue más rápido y más sincronizado en el tratamiento de alta temperatura. Los modelos lineales generalizados indican que el desarrollo del embrión depende una interacción cuádruple entre el estadio embrionario, el tiempo (h), la temperatura del agua de mar y la progenie. El crecimiento larvario fue más rápido, produciendo larvas más grandes a la temperatura más alta. El crecimiento de las larvas depende de una triple interacción entre el tiempo (DPF), la temperatura del agua de mar y la progenie. Conclusiones: Encontramos un impacto en la temperatura y en la progenie durante el desarrollo embrionario y larvario y, en ambos casos, estos factores generaron un efecto sinérgico sobre el tiempo de desarrollo y el tamaño de las larvas. Esto probablemente proporciona una ventaja de supervivencia, ya que una velocidad de desarrollo más rápida implica una disminución en el tiempo que pasan en la columna de agua, donde los erizos de mar son vulnerables a los factores estresantes bióticos y abióticos.

Morfología y crecimiento relativo en Clypeaster europacificus (Clypeasteroida: Clypeasteridae) del Golfo de California / Morphology and relative growth in Clypeaster europacificus (Clypeasteroida: Clypeasteridae) from the Gulf of California

Introducción: Clypeaster europacificus es una especie de galleta de mar que se distribuye de 0 a 200 m en el Golfo de California. Objetivo: Describir la morfología y el crecimiento relativo en la testa de C. europacificus. Métodos: Medimos 1 846 especímenes de museo de C. europacificus (N = 1 846) y analizamos el patrón de crecimiento en 32 especímenes de Baja California Sur, México (64 m de profundidad). Estudiamos cinco variables: radio mayor (R) (variable independiente), radio menor (r), longitud del pétalo anterior (LP), área total de prueba (A) y peso seco (P). Resultados: El área de cobertura del petaloide y el contorno de la testa son los caracteres taxonómicos importantes para C. europacificus: los especímenes van desde casi redondos en especímenes pequeños (≤ 3 cm de radio) a ondulados, enfatizando un contorno pentagonal, en organismos más grandes (≥ 3,1 cm de radio). Conclusiones: Existe un crecimiento alométrico negativo en las relaciones R vs. r, R vs. A y R vs. P; mientras que la relación R vs. LP tiene un crecimiento alométrico positivo.

Introduction: Clypeaster europacificus is a species of sand dollar found from 0 to 200 m in the Gulf of California. Objective: Describe the morphology and relative growth in the test of C. europacificus. Methods: We measured 1 846 museum specimens of C. europacificus (N = 1 846) and analyzed growth pattern in 32 specimens from Baja California Sur, Mexico (64 m depth). We studied five variables: major radius (R) (independent variable), minor radius (r), length of anterior petal (LP), total test area (A) and dry weight (P). Results: The petaloid cover area and the outline of the test are the important taxonomic characters for C. europacificus: they range from almost round in small specimens (≤ 3 cm in radius) to wavy, emphasizing a pentagonal outline, in larger organisms (≥ 3.1 cm in radius). Conclusions: There is negative allometric growth in the relationships R vs. r, R vs. A, and R vs. P; while the relationship R vs. LP has positive allometric growth.

Morphogenesis and histogenesis during the arm regeneration in a basket star Astrocladus dofleini (Euryalida, Ophiuroidea, Echinodermata).

Basket stars, that is, Ophiuroidea in Echinodermata, exhibit distinctive morphological characteristics with their complicatedly branched arms that can regenerate immediately after mutilation. Although, in brittle stars, that is, ophiuroids with nonbranched arms, the arm regeneration process following accidental trauma or autotomy have been morphologically and histologically observed in several species, few studies have so far been carried out on the regeneration of branched arms in basket stars. In this study, the developmental and morphological features of arm regeneration in Astrocladus dofleini (Gorgonocephalidae, Euryalida, Euryophiurida), one of the most common basket star species in Japanese waters, was anatomically and histologically investigated. Results clearly showed the following phases during the arm regeneration: (a) repair phase, (b) early regenerative phase, (c) intermediate regenerative phase, (d) advanced regenerative Phase I, and (e) advanced regenerative Phase II. The morphogenetic process during the arm regeneration in the basket star showed similar patterns to those of nonbranched arms observed in other ophiuroids. However, differences were also seen between the two ophiuroid types, that is, there were some developmental features specific to the basket star. In the early regenerative phase, branching of coelomic cavities was observed prior to the formation of other tissues, probably inducing the later morphogenesis of branched arms. In addition, hard skeletal ossicles form rapidly at the advanced regenerative Phase II. These developmental features may have led the evolution of bizarre morphologies seen in basket stars, probably contributing to the adaptation to shallow waters from deep-sea habitats.

Population size structure and abnormalities in the number of rays of the Sea Star Pentaceraster cumingi (Valvatida: Oreasteridae) in Bahía Chamela, Mexican Pacific / Estructura de tallas poblacional y anormalidades en el número de radios de la estrella de mar Pentaceraster cumingi (Valvatida: Oreasteridae) en Bahía Chamela, Pacífico mexicano

Introduction: The Panamic Cushion Star Pentaceraster cumingi is widely distributed along the Tropical Eastern Pacific. This species strictly produces only five arms, but sometimes, this number varies or show another kind of abnormality. Objective: We aimed to evaluate the population size structure and abnormalities occurrence in the radial pattern of P. cumingi in Bahía Chamela, Jalisco, Mexico. Methods: The population was monitored along four years (2016-2019), in two seasonal periods (warm and cold). During fieldwork, a random sample of individuals was collected. Every starfish was measured, weighted, and evaluated to identify any abnormality on its radial pattern. Results: The highest density of P. cumingi was found in October 2019 (2.03 ± 0.05 ind/m-2), the lower in March 2017 (0.66 ± 0.13 ind/m-2). A total of 849 individuals were collected. For 5-armed starfishes, the average length was 123.8 ± 15.2 mm and the average weight of 326.0 ± 62.4 g. The most frequent length classes ranged from 110 to 120 mm. Of the total of individuals sampled 0.82 % had four arms, 1.06 % six, and 1.41 % had one bifurcated arm. Conclusions: There were differences in the population density and size structure of P. cumingi between seasons. The main causes of abnormalities in the starfish could be due to the changes that occur during larval metamorphosis or by an abnormal regeneration of the arms after a predation attempt.

Introducción: La Estrella Cojín Pentaceraster cumingi está ampliamente distribuida a lo largo del Pacífico Oriental Tropical. Esta especie produce estrictamente cinco brazos, pero en algunas ocasiones el número puede ser menor o mayor de cinco, o mostrar otro tipo de anormalidad. Objetivo: Evaluar la estructura de tallas poblacional y la presencia de anormalidades en el patrón radial de P. cumingi en la Bahía Chamela, Jalisco, México. Métodos: La población fue monitoreada a lo largo de cuatro años (2016-2019), durante dos periodos estacionales (cálido y templado). Durante el trabajo de campo se recolectó una muestra aleatoria de individuos. Cada estrella de mar fue medida, pesada y revisada para identificar alguna anormalidad en el patrón radial. Resultados: La mayor densidad de individuos se encontró en octubre de 2019 (2.03 ± 0.05 ind/m-2), la menor en marzo de 2017 (0.66 ± 0.13 ind/m-2). Un total de 849 individuos fueron recolectados. Para las estrellas de mar con 5 brazos, la longitud promedio fue de 123.8 ± 15.2 mm y el peso promedio de 326.0 ± 62.4 g. La clase de talla más frecuente varió entre 110 y 120 mm. Del total de individuos muestreados el 0.82 % tuvieron cuatro brazos, el 1.06 % con seis, y el 1.41 % tuvieron un brazo bifurcado. Conclusiones: Se encontraron diferencias en la densidad poblacional y en la estructura de tallas de P. cumingi entre estaciones. Las principales causas de anormalidades en la estrella de mar pueden ser debidas a los cambios que ocurren durante la metamorfosis de las larvas o por una regeneración anormal de los brazos a causa de la depredación.

Axial complex of Crinoidea: Comparison with other Ambulacraria.

The anatomy of Crinoidea differs from that of the other modern echinoderms. In order to see, whether such differences extend to the axial complex as well, we studied the axial complex of Himerometra robustipinna (Himerometridae, Comatulida) and compared it with modern Eleutherozoa. The axial coelom is represented by narrow spaces lined with squamous coelothelium, and surrounds the extracellular haemocoelic lacunae of the axial organ. The latter is located, for the most part, along the central oral-aboral axis of the body. The axial organ can be divided into the lacunar and tubular region. The tubular coelomic canals penetrating the thickness of the axial organ have cuboidal epithelial lining, and end blindly both on the oral and aboral sides. The axial coelom, perihaemal coelom, and genital coelom are clearly visible, but they connect with the general perivisceral coelom and with each other via numerous openings. The haemocoelic spaces of the oral haemal ring pass between the clefts of the perihaemal coelom, and connect with the axial organ. In addition, the axial organ connects with intestinal haemal vessels and with the genital haemal lacuna. Numerous thin stone canaliculi pierce the spongy tissue of the oral haemal ring. They do not connect with the environment. On the oral side, each stone canaliculus opens into the water ring. The numerous slender tegmenal pores penetrate the oral epidermis of the calyx and open to the environment. Tegmenal canaliculi lead into bubbles of the perivisceral coelom. Some structures of the crinoid axial complex (stone canaliculi, communication between different coeloms) are numerous whereas in other echinoderms these structures are fewer or only one. The arrangement of the circumoral complex of Crinoidea is most similar to Holothuroidea. The anatomical structure and histology of the axial complex of Crinoidea resembles the "heart-kidney" of Hemichordata in some aspects.

Genomic insights of body plan transitions from bilateral to pentameral symmetry in Echinoderms.

Echinoderms are an exceptional group of bilaterians that develop pentameral adult symmetry from a bilaterally symmetric larva. However, the genetic basis in evolution and development of this unique transformation remains to be clarified. Here we report newly sequenced genomes, developmental transcriptomes, and proteomes of diverse echinoderms including the green sea urchin (L. variegatus), a sea cucumber (A. japonicus), and with particular emphasis on a sister group of the earliest-diverged echinoderms, the feather star (A. japonica). We learned that the last common ancestor of echinoderms retained a well-organized Hox cluster reminiscent of the hemichordate, and had gene sets involved in endoskeleton development. Further, unlike in other animal groups, the most conserved developmental stages were not at the body plan establishing phase, and genes normally involved in bilaterality appear to function in pentameric axis development. These results enhance our understanding of the divergence of protostomes and deuterostomes almost 500 Mya.

Evolution and Development at the Origin of a Phylum.

Quantifying morphological evolution is key to determining the patterns and processes underlying the origin of phyla. We constructed a hierarchical morphological character matrix to characterize the radiation and establishment of echinoderm body plans during the early Paleozoic. This showed that subphylum-level clades diverged gradually through the Cambrian, and the distinctiveness of the resulting body plans was amplified by the extinction of transitional forms and obscured by convergent evolution during the Ordovician. Higher-order characters that define these body plans were not fixed at the origin of the phylum, countering hypotheses regarding developmental processes governing the early evolution of animals. Instead, these burdened characters were flexible, enabling continued evolutionary innovation throughout the clades' history.