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A review of the genus Narcissia Gray, 1840 (Echinodermata: Asteroidea: Ophidiasteridae)
Abstract
Since the taxonomic revision of the Ophidiasteridae by H.L. Clark (1921), in which 20 genera were recognized as valid, several new genera and species were included in this family, and the boundaries between some of them became largely arbitrary. This also applies to Narcissia Gray, 1840, whose morphology and taxonomy are still very poorly understood today. The main goals of this research were: review the taxonomy of the genus Narcissia, from morphological studies; redescribe N. trigonaria Sladen, 1889 s. str.; characterize N. canariensis (d'Orbigny, 1839) and N. gracilis A.H. Clark, 1916, the two valid species most similar to N. trigonaria; review the validity of N. ahearnae Pawson, 2007 and N. trigonaria var. helenae Mortensen, 1933; review the validity of N. gracilis malpeloensis Downey, 1975. Altogether 370 specimens were analysed. All morphological structures of taxonomic importance have been illustrated, and, for the first time, the internal characters of this group were studied, from electron microscopy and microtomography. A new diagnosis for the genus Narcissia was constructed. Narcissia trigonaria was redescribed from a large specimen, from the type locality (Bahia, Brazil). Narcissia ahearnae is considered a valid species, and N. canariensis and N. trigonaria distinct entities. Narcissia trigonaria var. helenae is placed in the synonymy of N. trigonaria, and N. gracilis malpeloensis into the synonymy of N. gracilis. The dataset obtained in this work is by far the most complete acquired for the genus Narcissia.
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Narcissia ahearnae is a starfish found on hard substrates with a thin surface of fine sediment. This species has been collected in Florida and The Bahamas at depths of 53–135 m. This paper presents a new record of this species for the Mexican Caribbean (23 • 17 04 N, 87 • 57 04 W), on the northern coast of Quintana Roo, which extends its distribution range to its southernmost limit. A bathymetric range extension is also presented. Resumen Narcissia ahearnae es una estrella de mar que habita sobre sustratos duros que presentan una capa fina de sedimento en su superficie. Esta especie ha sido recolectada en las Bahamas y Florida a profundidades de 53-135 m. Se presenta un nuevo registro de la especie para el Caribe mexicano (23 • 17 04 N, 87 • 57 04 O), costa norte del estado de Quintana Roo, la cual extiende su área de distribución al límite meridional. También se presenta una extensión del rango batimétrico.
The skeletal morphology and homologies of 24 extant asteroid taxa and one Palaeozoic outgroup (Calliasterella mira; Upper Carboniferous, Moscow area, Russia) are described in detail. This information is used to construct a character matrix to identify deep relationships within the Neoasteroidea. Particular attention is paid to the morphology and homologies of the character-rich ossicles of the mouth frame (orals, circumorals, odontophore) and of the ambulacral groove (adambulacrals, ambulacrals). In addition, new homologies between alveolar and forcipulate pedicellariae are identified. Soft-tissue characters are reviewed from the literature. Simultaneous unconstrained cladistic analysis of 128 characters yielded six equally parsimonious trees, and a consensus tree of these is presented. This places the Paxillosida as the most basal neoasteroid group, sister taxon to the Surculifera, which include all other neoasteroids. The Spinulosida (used here in its original sense) are a well-supported monophyletic group, and a newly identified clade here named the Tripedicellaria, unified by characters of the pedicellariae, includes both the Valvatida and Forcipulatida. 'Valvatids' are a paraphyletic group of taxa that are sister group to the Forcipulatida. The fossil record of the asteroid families to which the 24 investigated species belong is reviewed, and the stratigraphical information is used to constrain the positions of the nodes on the consensus tree. The enigmatic asteroid fossil record from the Triassic is reviewed, and it is concluded that the order Trichasteropsida is a poorly understood assemblage of diverse morphologies of which the relationships with other taxa are uncertain. Middle Triassic (Muschelkalk) asteroids are identified as neoasteroids incertae sedis and assigned to two families: Trichasteropsidae and Migmasteridae fam. nov. Well-preserved asteroid ossicles from the Carnian of the Italian Dolomites are described and demonstrate that much of the neoasteroid radiation had taken place by the early Carnian (Late Triassic) because ossicles of taxa close to ophidiasterids and asterinids are present. It is concluded that the sudden appearance of diverse taxa with close affinities to modern families in the Early and Mid-Jurassic is an artefact of the extensive record of marine sediments of these ages and not of contemporaneous radiation of the neoasteroids. Detailed ossicular morphology is extremely useful and underused, both in the identification of asteroid relationships and for the assignation of fossil neoasteroids to extant families. The adaptive evolution of the neoasteroid skeleton is reviewed, and the neoasteroids are considered to be primitively infaunal and only secondarily adapted to epifaunal modes of life, the opposite of the case of post-Palaeozoic echinoids. It is concluded that Xyloplax is a true neoasteroid and most closely related to the Caymanostellidae. A new genus of Xyloplacidae, Ankyloplax, is erected. Calliasterella mira, from the Upper Carboniferous of Moscow, is redescribed. The first Jurassic goniopectinid (Chrispaulia jurassica sp. nov.), a benthopectinid (Jurapecten hessi gen. et sp. nov.) and two pterasterids (Savignaster wardi gen. et sp. nov., Savignaster trimbachensis gen. et sp. nov.) are described from the upper Oxfordian (Upper Jurassic) of the French Jura. Terminaster cancriformis is redescribed on the basis of new material and assigned to a new family, the Terminasteridae.
Presently more than 1900 species of sea stars are recognized, of which 77 are recorded for the coast of Brazil. Although the first starfish record in Brazil was published 363 years ago, our knowledge of this fauna remains unsatisfactory from a systematic and ecological point of view, particularly in the north and northeastern regions of the country. This study provides the first annotated list of sea stars from northeastern Brazil. Material described herein is housed at the collections of the Federal University of Paraíba, Federal University of Sergipe, and the Federal University of Bahia, Museum of Zoology of the University of São Paulo and Museu Nacional do Rio de Janeiro. Twenty-one species were identified, belonging to 12 genera, 10 families, and 5 orders. Descriptions of species are provided. Three new occurrences were recorded for northeast Brazil: Astropecten
alligator, Luidia
ludwigi
scotti, and Mithrodia
clavigera. Highest diversities of Asteroidea were encountered for the states of Bahia (n = 14 spp), Paraíba (n = 12 spp) and Pernambuco (n = 9 spp). No species were recorded for the states of Maranhão and Sergipe. Sandy substrates and depths below 10 m were the least sampled areas over the continental shelf. Herein we provide a first panorama on the fauna of Asteroidea occurring in the northeast region of Brazil, hopefully to function as a basic reference for biodiversity studies in this poorly studied area.
Collections and inventories provide important and essential information for understanding the composition and distribution of biodiversity. This contribution presents an inventory of the species held in the Echinodermata collection in the Setor de Comunidades Bentônicas (LABMAR/ICBS), Universidade Federal de Alagoas. The collections were made over a period of more than 20 years, at several localities of coastal Alagoas. This collection now includes 16,201 catalogued individual specimens, with 50 species in the five extant classes Crinoidea, Asteroidea, Ophiuroidea, Echinoidea and Holothuroidea, based on 640 records; the class Ophiuroidea is best represented. The results of this inventory contribute significantly to knowledge of marine benthic diversity from the state of Alagoas, and the information presented here expands the data for Echinodermata from the northeastern coast of Brazil.
We investigated the current patterns of diversity by country and by class of echinoderms, and analyzed their biogeographical, depth, and habitat or substratum affinities, using the database of the appendix of this book. Traditionally, the area has been divided into five biogeographical Regions and nine Provinces that cover a wide climate range. Currently, the echinoderm fauna of Latin America and Canary islands is constituted by 1,539 species, with 82 species of Crinoidea, 392 species of Asteroidea, 521 species of Ophiuroidea, 242 species of Echinoidea and 302 species of Holothuroidea. Species richness is highly variable among the different countries. The number of species for the countries is highly dependent on its coast length. The echinoderm fauna of the Panamic, Galápagos and the Chilean regions are biogeographically related. Other regions that are closely related are the Caribbean, West Indian, Lusitania and Brazilian. Cosmopolitan species are an important component in all the regions. Affinities between faunas are a consequence of the combination of climatic and trophic factors, connectivity as a function of distance, currents patterns and historical processes. Moreover, different environmental factors would be responsible for the faunal composition and species distribution at different spatial scales. The bathymetrical distribution of the echinoderm classes and the species richness varies according to the depth range and the ocean. Most species occurred at depths between 20 and 200 m. The Caribbean-Atlantic regions are richest in shallow depths, while the Pacific coast has higher values in deeper waters. The domination of each class in each substrate and habitat categories also varies differentially along each coast.
A systematic list of echinoderms from Malpelo Fauna and Flora Sanctuary (MFFS) was prepared, based on localsampling, literature review and identification of specimens from Museo de Historia Natural Marina de Colombia(MHNMC) and from National Museum of Natural History, Smithsonian Institution, Washington D. C. (USNM).Standard methodology for monitoring the Eastern Tropical Pacific Seascape was used in December 2006 and March2007 to sample shallow water echinoderms (1-20 m deep). Malpelo is the richest locality of the Colombian Pacific with84 species reported (56 % of them deep water organisms), belonging to 22 orders, 42 families and 68 genera, distributedbetween the surface and a depth of 5000 m. Thirty-six new records for MFFS and Colombian Pacific are reported. InColombia the shallow water species are widely distributed in the Pacific, while deep forms only occur at Malpelo. Themajority of species are mostly related to the Panamanian and Galapagos region showing a possible connectivity. Thislist increases knowledge on the echinoderm biodiversity from the Malpelo Island in 68 %.
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.
Narcissia ahearnae, new species, is described from off the east coast of Florida and the Bahamas, in depths of 53–135 m. It is distinctive in having conspicuous undulating carinal ridges extending from the center of the disc along the upper surfaces of the arms.
A new phylogenetic analysis of the superorder Forcipulatacea is presented. Forcipulatacea is one of the three major groups of sea stars (Asteroidea: Echinodermata), composed of 400 extant species. The sampled taxa are thought to represent the morphological diversity of the group. Twenty-nine forcipulate taxa were sampled belonging to Asteriidae, Stichasteridae, Heliasteridae, Pedicellasteridae, Zoroasteridae and Brisingida. Specimens were dissected with bleach. Detailed description of the skeleton and the anatomy of the ossicles were investigated using scanning electron microscopy. Comparative anatomy allowed the scoring of 115 phylogenetically informative characters. The consensus tree resulting from the analysis recovers Asteriidae, Stichasteridae, Zoroasteridae and Brisingida as monophyletic. All types of morphological features contribute to tree resolution and may be appropriate for taxon diagnosis. The synapomorphies supporting different clades are described and discussed. Brisingida and Zoroasteridae are the best-supported clades. The potentially challenging position of Brisingida in the tree may be explained by homoplastic changes, but also by the presence of numerous non-applicable characters.
Narcissia ahearnae, new species, is described from off the east coast of Florida and the Bahamas, in depths of 53-135 m. It is distinctive in having conspicuous undulating carinal ridges extending from the center of the disc along the upper surfaces of the arms.
Introduction ... 3 List of the stations, where Asteroidea were collected ... 4 Taxonomic report ... 11 Survey of the species ... 18 Literature cited ... 82 Explanation of the plates ... 88 Index ... 90 INTRODUCTION The present paper deals with the Asteroidea (Echinodermata), collected during two surveys of the continental shelf of Surinam (O. C. P. S. I & II) carried out in the spring of 1966 and 1969 by H . N L . M . S. Snellius and H. N L . M . S. Luymes, respectively. During my studies I have identified and classified the sea-stars of the bottom-samples. I have also tried to solve nomenclatural problems and to correct some nomenclatural conceptions, with which I did not agree. For each species I have composed the distributional pattern from records in the literature. During the surveys "it proved to be possible to correlate the zoological work with the geomorphological and sedimentological survey of the shelf bottom" (Vervoort, 1967, 1971, these papers can be considered as an introduction to this report).
CONTENTS Introduction.......... 3 Acknowledgements........ 4 List of the stations where Asteroidea were collected........ 4 Taxonomic report......... 7 Identification.......... 9 Survey of the species........ 11 Literature cited......... 88 Index............ 95 INTRODUCTION In a previous paper (Walenkamp, 1976) a revision was given of the Asteroidea of the coastal waters of Surinam. Since the preparation of that paper new material, collected in 1970 by the "Luymes" Guyana Shelf Expedition, became available. The present report is based on this new material. The 1970 expedition not only explored the waters off Surinam, but also the continental shelf off Guyana and French Guyana, and, moreover, investigated deeper waters than the O.C.P.S. expeditions dealt with in my 1976 paper. As a result fourteen species were taken, which were not represented in the collections of the previous expeditions. The present report, which can be considered a supplement to the 1976 report, has the same set-up. Of the newly found species synonymy and full descriptions are given. For the other species I may refer to the 1976 descriptions. In addition to the material mentioned above, a small collection of Asteroidea, property of the Surinaams Museum, Paramaribo, is included. These
The morphology of the following eight major ossicle types is described and illustrated for the goniopectinid asteroid Ctenodiscus crispatus: terminal plates, superomarginal and inferomarginal ossicles, adambulacral and ambulacral ossicles, odontophores, oral intermediate plates, and superambulacral ossicles. Development, variation, and relationships with soft body-parts and with other ossicles are embphasized. Each ossicle type is distinguished by numerous structures related to its function and to articulation with adjoining skeletal elements. Because major structures (such as pustules, alveoli, and articulation surfaces) distinguishing ossicle types develop early during ontogeny, immature ossicles are readily identifiable. However, changes in form and orientation of these structures occur during ossicle growth. Ontogenetic changes are influenced by development of associated skeletal and soft parts. Ambulacral and adambulacral ossicles near the peristome are highly modified but retain the basic characteristics of structure and orientation which define these major types of skeletal elements.
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