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Corresponding author: Anne Isabelley Gondim ( anneisabelley@yahoo.com.br ) Academic editor: Yves Samyn
© 2014 Anne Isabelley Gondim, Martin Lindsey Christoffersen, Thelma Lucia Pereira Dias.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gondim A, Lindsey Christoffersen M, Dias T (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys 449: 1-56. https://doi.org/10.3897/zookeys.449.6813
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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.
Echinoderms, taxonomy, check list, sea-stars, northeastern Brazil
The Asteroidea represent the second most diverse group within the phylum Echinodermata, with an estimated number of 1900 living species (
The first paper to deal with the Echinodermata of the Brazilian littoral was published by Georg Marcgraf (1648) and entitled “Natural History of Brazil”. In this work, animals observed from an expedition to northeast Brazil during the stay of the Count of Nassau are described, sometimes in a rather fanciful way (
Thirty six years after the paper by
After these pioneer studies, several works focusing on taxonomy, ecology and geographical distribution were developed, mainly in the south and southeast (e.g.
For northeast Brazil, only three papers have focused on the Asteroidea:
Northeastern Brazil has a coastline with 3,400 km in extension (
Although the first record of the Asteroidea for the Brazilian littoral was made about 363 years ago, the knowledge of this fauna in the north and northeastern regions still remains unsatisfactory and punctual, from both a systematic and an ecological point of view. The aim of the present work is to expand our knowledge on the morphology of the species of Asteroidea from northeastern Brazil.
Species determinations were based mainly on
Illustrations are based on photos made with a Canon A640 10MP camera coupled with a Nikon stereomicroscope. Morphometrics were recorded with a digital EDC 6 caliper ruler.
The studied material is conserved in 70% alcohol or preserved dry and registered in the visited collections.
The littoral region of northeast Brazil extends for approximately 3,400 km (
The region has a great diversity of littoral environments, such as beaches, dunes, cliffs, estuaries, deltas, sandbanks, reefs and underwater biotopes, such as rhodolith beds, seagrass beds and coral habitats (Fig.
Due to the absence of large rivers and to the predominance of warm waters from the South Equatorial Current, offshore conditions in the northeast region are ideal for the formation of coastal coral reefs (
The northeastern continental shelf is narrow and shallow. Due to the reduced continental influence and to the tropical climate, an important sedimentation of biogenic carbonates dominates most of the middle and outer platform, particularly between Macau (RN) and Maceió (AL) (
The biogenic carbonates form sand and gravel, consisting mainly of incrusting and ramified calcareous algae, with local occurrences of Halimeda spp. These sediments may contain up to 5% of carbonatic mud derived from the desintegration of larger organisms. In certain areas, these sediments undergo a process of litification, also affecting the quartsoze sands of the inner platform, originating the extensive coastal reefs known as beach rocks. These then become overgrown by algae and corals (
Abbreviations: Brazilian states – Ceará (CE), Piauí (PI), Rio Grande do Norte (RN), Paraíba (PB), Pernambuco (PE), Bahia (BA), Paraná (PR), Rio de Janeiro (RJ), Rio Grande do Sul (RS), Santa Catarina (SC), São Paulo (SP).
Acronyms: UFPB.Ech: Echinodermata Collection from Universidade Federal da Paraíba. MZUFBA: Museum of the Federal University of Bahia. MZUSP: Museum of Zoology of the University of São Paulo. MNRJ: National Museum of Rio de Janeiro.
A total of 21 species, belonging to five orders, 10 families and 12 genera were identified. These are listed and described below, following the taxonomic organization of
Phylum Echinodermata Brugière, 1791
Class ASTEROIDEA de Blainville, 1830
Order Paxillosida Perrier, 1884
Family Luidiidae Verrill, 1900
Luidia alternata alternata (Say, 1825)
Luidia clathrata (Say, 1825)
Luidia ludwigi scotti Bell, 1917
Luidia senegalensis (Lamarck, 1816)
Family Astropectinidae Gray, 1840
Astropecten acutiradiatus Tortonese, 1956
Astropecten alligator Perrier, 1881
Astropecten brasiliensis Müller & Troschel, 1842
Astropecten cingulatus Sladen, 1833
Astropecten duplicatus Gray, 1840
Astropecten marginatus Gray, 1840
Order Valvatida Perrier, 1884
Family Asterinidae Gray, 1840
Asterinides folium (Lütken, 1860)
Family Mithrodiidae Viguier, 1878
Mithrodia clavigera (Lamarck, 1816)
Family Oreasteridae Fisher, 1911
Oreaster reticulatus (Linnaeus, 1758)
Family Goniasteridae Forbes, 184
Nymphaster arenatus (Perrier, 1881)
Plinthaster dentatus (Perrier, 1884)
Order Velatida Perrier, 1884
Family Pterasteridae Perrier, 1875
Calyptraster coa Sladen, 1882
Family Ophidiasteridae Verrill, 1870
Linckia guildingi Gray, 1840
Narcissia trigonaria Sladen, 1889
Order Spinulosida Perrier, 1884
Family Echinasteridae Verrill, 1867
Echinaster (Othilia) brasiliensis Müller & Troschel, 1842
Echinaster (Othilia) echinophorus (Lamarck, 1816)
Order Forcipulatida Perrier, 1884
Family Asteriidae Gray, 1840
Coscinasterias tenuispina (Lamarck, 1816)
Asterias alternata Say, 1825: 144–145.
Luidia alternata Lütken, 1859: 42–43.
Luidia granulosa Perrier, 1869: 109–110, pl. 2, fig. 18.
Luidia variegata Perrier, 1875: 337.
Luidia numidica Koehler, 1911: 3, pl. 1, figs 8–11.
Luidia quequenensis Bernasconi, 1942: 253.
Luidia bernasconiae A.H.Clark, 1945: 19–21.
Luidia alternata var. numidica Madsen, 1950: 206–209, fig. 9.
Luidia alternata numidica A.M.Clark, 1953: 388–389, pl. 41, fig. 1.
Luidia alternata alternata Clark & Downey, 1992: 8–9;
Paraíba: 6°46'S; 34°50'W, 1spec., UFPB/ECH.877, 13.II.1981, 14m; 7°01'S; 34°41'W, 1spec., UFPB/ECH.876, 13.II.1981, 24m; 7°04"S; 34°41'W, 1spec., UFPB/ECH.879, 16.II.1981, 22m.
Dry Tortugas, Florida Keys, Florida (
Body pentagonal (Fig.
Some species of the family Luidiidae recorded in northeastern Brazil. Luidia alternata alternata (A–D). A Abactinal view, in detail the paxilla B Actinal view, in detail the pedicellariae C Abactinal view of the arm D Actinal view of the arm; Luidia clathrata (E–H)E Abactinal view F Actinal view, in detail the mouth G Abactinal view of the arm, in detail the paxilla, and H Actinal view of the arm.
Colour. Abactinal surface white or cream-coloured, with dark transversal bands. The colour of these bands varies, and may be brown, black, green, or purple. Actinal surface whitish or beige.
North Carolina, the Bahamas, Florida, Gulf of Mexico, Cuba, Jamaica, Honduras, Puerto Rico, Panama, Colombia, Venezuela, Brazil, Uruguay, and Argentina (
The main characters distinguishing Luidia alternata alternata from the remaining species of the genus that occur in the Western Atlantic are their colour and the presence of long conical spines on the adradial paxillae. The abactinal colouration is distinctive. The nominal subspecies differs from its congeneric form Luidia alternata numidica Koehler, 1911, from West Africa, for attaining a larger size and for having longer paxillar spines.
This subspecies is primarily associated with sandy and muddy bottoms. It may also be found in mangroves or associated with fragments of shells and calcareous algae (
Asterias clathrata Say, 1825: 142.
Luidia clathratta Lütken, 1859: 37–39.
Luidia clathrata Lütken, 1859: 37.
Paraíba: 6°57'S; 34°41'W, 2 spec., UFPB/ECH.875, 12.II.1981, 26m.
Probably no longer existant (
Five long and narrow arms (Fig.
Colour. Abactinal surface bluish-gray, frequently with a darker line occupying the carinal region of the arm and disk.
Bermuda, Gulf of Mexico, Belize, Honduras, Nicaragua, Panama, Colombia, Venezuela, and Brazil (
Luidia clathrata differs from L. alternata alternata for not having the abactinal surface spinulose. It differs from L. senegalensis for having only 5 arms and from L. ludwigi scotti for not having pedicellariae. According to
This species lives in sandy or muddy areas with low hydrodynamism near the coast, such as bays and lagoons, and is also found in mangroves and regions with low salinity. Further away from the coast, it lives in substrates with sand, mud and gravel (
Luidia scotti Bell, 1917: 8–9.
Luidia doello-juradoi Bernasconi, 1941: 117; 1943: 8–11.
Luidia rosaurae Jonh & Clark, 1954: 142–145.
Luidia doello-juradol Brito, 1968: 12, pl. 3, fig. 5.
Luidia ludwigi Walenkamp, 1976: 32–37, fig. 9, pl. 2, figs 1–3, pl.4, fig. 3.
Luidia ludwigi scotti A.M.Clark, 1982: 171–173.
Luidia rosaurae John & Clark, 1954: 142–145, pl. 6, fig. 1.
Paraíba: 6°39'05"S; 34°49'W, 1 spec., UFPB/ECH.878, 28.V.1981, 20m.
Rio de Janeiro, Brazil (
Body flattened. Five arms that taper gradually towards their extremities (Fig.
Some species of the family Luidiidae recorded in northeastern Brazil. Luidia ludwigi scotti (A–D)A Abactinal view, in detail abactinal intermediate area B Actinal view, in detail the mouth C Abactinal view of the arm, in detail the paxilla D Actinal view. Luidia senegalensis (E–H)E Abactinal view F Actinal view, in detail the mouth G Abactinal view of the arm, in detail the paxilla, and H Actinal view of the arm.
Colour. The colour pattern on the aboral surface of the body is very variable. Some specimens may be whitish with black spots on arms, others are brown with white spots on arms. Most, however, have the arms more or less banded with alternating white and brown stripes (
Florida, Gulf of Mexico, Colombia, Venezuela, Guyana, French Guyana, Brazil, and Argentina (Mar del Plata) (
This subspecies may be distinguished from other taxa in the genus by the presence of pedicellariae with three or four well-developed valves on the actinal surface, the delicate structure of the paxillae, and the slightly triangular shape of the arms, which become narrow distally. Luidia clathrata is similar to L. ludwigi scotti, but may be distinguished from it by having a stronger actinal skeleton and by the absence of pedicellariae. The synonymy between Luidia rosaurae John & Clark, 1954 and L. scotti was proposed by
This subspecies occurs in non-consolidated sediments containing fine or coarse sand (
Asterias senegalensis Lamarck, 1816: 567.
Luidia senegalensis Müller & Troschel, 1842: 78.
Luidia marcgravii Steenstrup in
Rio Grande do Norte: Timbau Beach, 3 spec., UFPB/ECH.1582, 28.II.1980; Areia Branca, Ponta do Mel, 1spec., UFPB/ECH.1428, 23.VI.1982. Paraíba: Lucena, Costinha Beach, 5 spec., UFPB/ECH.1673, 08.XI.2003; Cabedelo, Miramar Beach, 4spec., UFPB/ECH.1256, 03.II.1983; Cabedelo, Santa Catarina Beach, 1spec., UFPB/ECH.1583, 18.V.2007; Paraíba do Norte River Estuary, 1spec., UFPB/ECH.89, 18.06.1980; 1spec., UFPB/ECH.1586. 13.VI.1983.
Supposedly Senegal (‘L’ océan d’Afrique, les côtes du Senegal’), but probably West Indies (
Body flattened. Disk rounded. Nine long and narrow arms (rarely 7) (Fig.
Colour. Dorsally bluish-grey or greenish-gray, with a strong dark line along the central region of the disk and of the arms. The actinal surface is white to cream-coloured.
Florida, Cuba, Jamaica, Honduras, Nicaragua, Costa Rica, Panama, Venezuela and Brazil (
Distinguished from the remaining species from Brazil by the presence of 7 to 9 arms. Luidia barbadensis Perrier, 1881, recorded for Bahamas, Florida, Gulf of Mexico and south Brazil is the closest species, but it may be distinguished from L. senegalensis for having 6 arms, inframarginal figs with two long and narrow spines, and the usual presence of pedicellariae.
The species lives in environments of low hydrodynamism, in sediment containing sand, mud, or a combination of both of these (
Astropecten alligator Perrier, 1881: 30.
Astropecten nuttingi Verrill, 1915: 181, pl. 12, fig. 2e, pl. 21, figs 1–2.
Paraíba: 7°01'S; 34°41'05"W, 1 spec., UFPB/ECH.881, 13.II.1981, 24m.
Alligator Reef, Florida Keys, Florida (
Body flattened dorso-ventrally. Disk small, with five long and narrow arms (~4.93 mm) (Fig.
Some species of the family Astropectinidae recorded in northeastern Brazil. Astropecten alligator (A–D)A Abactinal view, in detail of the paxilla B Actinal view, in detail of mouth C Abactinal view of the arm D Actinal view of the arm; Astropecten cingulatus (E–H)E Abactinal view, in detail the paxilla F Actinal view G Abactinal view of the arm, and H Actinal view of the arm.
Colour. Reddish-brown dorsally, with conspicuous dark bands along the center of each arm (
North Carolina, the Bahamas, Florida, Honduras, Nicaragua Colombia, and Brazil (
Astropecten alligator difers from the remaining species in the genus recorded for Brazil for presenting a single, elongate, erect conical spine on all the superomarginal figs. This character is present in both juvenile and adult individuals (dd = 41.0 mm). For a long time it was believed that
Usually inhabits muddy environments, but may also be found associated to communities of azooxanthelate corals (
Asterias articulata Say, 1825: 144.
Astropecten dubius Gray, 1840: 182.
Asterias aranciaca Gould, 1841: 349 (non Asterias aranciacaLinnaeus, 1758).
Astropecten articulates Müller & Troschel, 1842: 72.
Astropecten buschi Müller & Troschel, 1843 (a variety of Astropecten articulatus (Say, 1825) according to
Astropecten articulatus dubius Verrill, 1915: 165.
Astropecten articulatus var. valenciennesi A.H.Clark, 1939: 442.
Astropecten articulatus duplicatus Zoppi de Roa, 1967: 277, fig. 6. (non Astropecten duplicatus Gray, 1840).
Caraguatatuba, São Paulo, 1 spec., MZUSP484, 16.X.2001, 19m.
Florida (
Body pentagonal, flattened dorso-ventrally. Five long and narrow arms (length of arm corresponds to approximately four times its width), which become gradually narrow towards tips. Abactinal surface with paxillae. Paxillae disposed regularly, carenals smaller than adradials. 1–6 short, thick and blunt paxillar spines in center, 10–16 in margins. Supermarginal figs granulose, with a large spine (~1.54 mm) positioned internally on the interbrachial figs. Some distal supermarginal figs have a short spine positioned more externally. Inferomarginal figs with two elongate, flattened and pointed marginal spines, positioned horizontally. Small oral spine. Three adambulacral spines divergent, the median one being the largest.
Colour. Dorsally dark blue or purple in paxillar region. Superomarginal figs white or orange. Oral surface white of beige (
North Carolina, Florida, the Bahamas, Mexico, Cuba, Puerto Rico, Dominican Republic, Panama, Colombia, Venezuela, Brazil, and Uruguay (
Astropecten articulatus differs from the congeneric A. cingulatus for having a spine or tubercule on the distal supermarginal figs, two flat spines on each inferomarginal fig and subambulacral spines larger that the adambulacral spines. The original description of A. articulatus
Inhabits sandy environments. According to
Astropecten brasiliensis Müller & Troschel, 1842: 68.
Astropecten braziliensis Rathbun, 1879: 150.
Astropecten brasiliensis riensis Döderlein, 1917: 84.
Astropecten brasiliensis armatus Jonh, 1948: 503.
Astropecten armatus brasiliensis Tortonese, 1956: 329.
Astropecten brasiliensis brasiliensis Döderlein, 1917: 83.
Astropecten armatus riensis Tommasi, 1958: 13–14, pr. 2, fig. 4.
Astropecten riensis Tommasi, 1970: 7.
Ceará: Fortaleza, Mucuripe, 1 spec., MNRJ285, 1945. Rio Grande do Norte: Areia Branca, Ponta do Mel, 1 spec., UFPB/ECH.1919, 23.VI.182. Bahia: Salvador, 2 spec., UFBA00132, 01.III.2000.
São Sebastião Island, São Paulo, Brazil (
Disk small with long, slender, dorsally flattened arms (~12.44 mm). Abactinal surface densely covered by overlapping and irregularly arranged paxillae. Paxillae with 26–30 long, blunt spinelets, the central ones sometimes shorter and more rounded than the marginal ones. Carinal paxillae larger than the adradial ones. Superomarginal figs longer (~5.7 mm) than wide (~1.19 mm), covered by short, apically rounded spinelets (giving them a granulose aspect), and having two large spines (rarely one). Inferomarginal figs with two long and flattened spines positioned vertically and forming a marginal fringe, the most ventral one being larger than the dorsal one. Adambulacral figs with three ambulacral spines, the inner one being slightly longer than the other two. Without pedicellariae.
Colour. According to
Honduras, Panama, Suriname, Brazil, Uruguay, and Argentina (Mar del Plata) (
This species differs from the remaining species of the genus known from the Brazilian coast for presenting paxillary spinelets and spines of the marginal fringe long and thin and for having up to two spines on the supermarginal figs. The vast synonymy presented by Astropecten brasiliensis evidences the plasticity of some of its characters, such as the number of spines on the superomarginal figs. This character, together with the shape of the marginal spines and paxillae spinelets formed the basis for the establishment of the five known subspecies.
As a rule burrowed in sand substrates, where it is a generalist predator, feeding on a variety of organisms from the benthic endofauna, such as bivalves, gastropods, crustaceans, echinoderms, and polychaetes (
Astropecten cingulatus Sladen, 1883: 266.
Astropecten mesactus Studer, 1884: 46.
Astropecten jarli Madsen, 1950: 181.
Rio de Janeiro: Cabo Frio, 1 spec., MNRJ1853, 18.VI.1997.
Pernambuco, Brazil (
Body pentagonal, flattened dorso-ventrally. Five long (~37.70 mm) and narrow (~9.56 mm) arms (length of arm corresponds to approximately four times its width) (Fig.
Colour. According to
North Carolina, the Bahamas, Gulf of Mexico, Mexico, Nicaragua, Costa Rica, Panama, Colombia, Brazil, Uruguay, Argentina, and Africa (
Astropecten cingulatus differs from its closest species, A. articulatus, for having three rounded spines on each inferomarginal fig, and for having subambulacral spines that are smaller than the adambulacral spines and rounded.
This species lives on sandy or muddy bottoms of the littoral region up to depths of 50 m (
Astropecten marginatus Gray, 1840: 181.
Astropecten ciliatus Grube, 1857: 340.
Astropecten richardi Gary, 1840: 181.
Astropecten orans Sluiter, 1895: 54.
Rio Grande do Norte: Areia Branca, Ponta do Mel, 2 spec., UFPB/ECH.1842, 23.VI.1982. Paraíba: Cabedelo, Miramar Beach, 1 spec., UFPB/ECH.1840, 03.II.1983; João Pessoa, 7°7'23,3"S; 34°48'27,9"W, 1 spec., UFPB/ECH.1839, 14.IX.1980; João Pessoa, Tambaú Beach, 1spec., UFPB/ECH.864, 03.X.2007. Pernambuco: Goiana, Catuama, UFPB/ECH.1427, 1 spec., 31.X.1982.
Unknown (
Body flattened. Five broad, triangular arms (Fig.
Colour. Dorsally either blue with white margins (
Costa Rica, Puerto Rico, Colombia, Venezuela, Guyana, and Brazil (
Astropecten marginatus differs from the remaining species of the genus known from Brazil for having large, triangular arms and two long, thick and blunt spines on each inferomarginal fig. Unlike other species of Astropecten, A. marginatus shows little morphological variation in characters considered of taxonomic interest (
This species lives in substrates containing sand or sand with mud (
Asterina minuta Gray, 1840: 289.
Asteriscus folium Lütken, 1860: 60–61.
Asterina folium A. Agassiz, 1877: 106, pl. 14, figs 7–9.
Asterinides folium Verrill, 1913: 479;
Paraíba: 06°59'S; 34°47'W, 1spec., UFPB/ECH.572, 07.III.2006, 10m. Bahia: Camaçari, Guarajuba, Busca Vida Beach, 1spec., UFBA00685, 01.II.2006; 2spec., UFBA00983, I.2010, 23m; 1spec., UFBA01163, VII.2010, 25m; 2spec., UFBA01107, I.2010, 23m; Salvador, Todos os Santos Bay, 1spec., UFBA01107, 05.IV. 1997, 12m; Salvador: Itapuã, 12°57'28"S; 38°21'22"W, 1spec., UFBA00528, 19.XI.2007, 1m.
Saint Thomas, Virgin Islands (
Body inflated, pentagonal (Fig.
Colour. Juvenile specimens vary from white to cream-coloured, larger juveniles are yellow to reddish, while adults are blue or greenish-blue (
Bermudas, Florida, Bahamas, Belize, Panama, and Brazil (
Only two species of the genus Asterinides are known from the Atlantic Ocean, A. folium and A. hartmeyeri (Döderlein, 1910). The first is recorded for Bermudas and southern Brazil and the second only for the Caribbean region. According to
This species lives in association with coral reefs, being found particularly under rocks or corals of the reef flat (
Asterias clavigera Lamarck, 1816: 562.
Mithrodia clavigera Verrill, 1870: 289.
Mithrodia spinulosa Gray, 1840: 288.
Ophidiaster echinulatus Müller & Troschel, 1842: 32.
Echinaster echinulatus von Martens, 1866: 59.
Mithrodia clavigera Perrier, 1875: 378.
Mithrodia victoriae Bell, 1882: 123, pl. 6, fig. 2.
Paraíba: 7°04'S; 34°41'W, 1 spec., UFPB.ECH.880, 17.II.1981, 26m.
Unknown (
Disk small (Fig.
Colour. Arms banded with dark brown or red lines, unusually green (
Mexico, Cuba, Nicaragua, Brazil, Indo-Pacific (except Hawaii) (
Mithrodia clavigera is the only species of the family Mithrodiidae recorded for the Western Atlantic. For some time, two species were considered present: M. clavigera and M. victoriae. The later species was described by
This species lives on hard substrates covered by incrusting organisms and in reef gravel (
Asterias gigas Linnaeus, 1753: 114.
Asterias reticulata Linnaeus, 1758: 661.
Pentaceros reticulatus Gray, 1840: 276.
Oreaster reticulatus Linnaeus, 1758.
Oreaster aculeatus Müller & Troschel, 1842: 50.
Oreaster lapidarius Grube, 1857: 342.
Oreaster tuberosus Möbius, 1859: 6.
Oreaster gigas Lütken, 1860: 64–75.
Oreaster reticulatus var. bermudensis H.L. Clark, 1942: 372, figs 1–2.
Ceará: off Fortaleza, 1spec., UFPB/ECH.1255, Geomar XXIV, V.1985. Paraíba: 1spec., UFPB/ECH.1579, 26.X.1980; Cabedelo, Farol de Cabedelo Reef, UFPB/ECH.1254, 22.I.1981, 26m; 1spec., UFPB/ECH.1588, 26.X.1980; 6°39'S; 34°49'W, 1spec., UFPB/ECH.1575, 28.V.1981, 20m; 6°39'5"S; 34°46'W, 1spec., UFPB/ECH.1429, 1spec., 29.V.1981, 35m; 6°39´05"S; 34°49"W, 1spec., UFPB/ECH.1251, 28.V.1981, 20m; 6°39'05"S; 34°49'W, 1spec., UFPB/ECH.1578, 28.V.1981, 20m; 6°50'S; 34°47'W, 1spec., UFPB/ECH.1253, 11.V.1981, 18m; 6°52'S; 34°46'W, 1spec., UFPB/ECH.1872, 19.II.1981, 18m; 6°52'S; 34°48'W, 1spec., UFPB/ECH.1590, 04.II.1981, 10m; 6°52'S; 34°49'W, 1spec., UFPB/ECH.1593, 04.II.1981, 12m; 6°57'S; 34°41'W, 1spc., UFPB/ECH.1577, 12.II.1981, 26m; 7°01'S; 34°47'05"W, 2spec., UFPB/ECH.1252, 02.V.1981, 11m; 7°04'S; 34°41'W, 2spec., UFPB/ECH.1576, 17.II.1981, 22m; 07°04'24,4"S; 034°47'49"W, 1spec., UFPB/ECH.1871, 24.VI.2005, 6m; 7°07'S; 34°47'W, 1spec., UFPB/ECH.1430, 05.II.1981, 10m; 7°10'S; 34°38'W, 1spec., UFPB/ECH.1873, 26.03.1981, 25m; 7°13'S; 34°42'W, 1spec., UFPB/ECH.1574, 27.III.1981, 10m; 7°34'S; 34°39'W, 1spec., UFPB/ECH.1594, 1spec., 7°34'S; 34°39'W, Paraíba, 22.I.1981, 26m. Pernambuco: Itamaracá Island, 4spec., UFPB/ECH.1581, VIII.1980, 20m; Suape Beach, 1spec., UFPB/ECH.1580, I.1980.
‘Las Antillas Occidentales’ or ‘Spanish West Indies’ (
Body pentagonal. Disk high, inflated, concave on actinal side (Fig.
Some species of the order Valvatida recorded in northeastern Brazil. Asterinides folium (A–E)A Abactinal view, in detail the superomarginal figs B Actinal view, in detail actinal internidate area C Detail the abactinal intermediate are D Actinal view of the arm E Detail of the mouth; Mithrodia clavigera (F–I)F Abactinal view G Actinal view, in detail the mouth H Abactinal view of the arm, in detail a spine, and I Actinal view of the arm.
Colour. According to
North and South Carolina, the Bermudas, the Bahamas, Belize, Guatemala, Honduras, Cuba, Nicaragua, Costa Rica, Venezuela, Brazil, and Cabo Verde (
Only two species of the genus Oreaster are known for the Atlantic Ocean, Oreaster clavatus and O. reticulatus. The first is known only from the Island of Cape Verde, São Thomé and the Gulf of Guinea. The second, occurs widely throughout the West Atlantic, from North Carolina to the south of Brazil (
The species lives in shallow reef environments with calm water, coastal lagoons, seagrass beds (Thalassia, Halodule and Syringodium), and mangrove channels (
Calyptraster coa Sladen, 1882: 207.
Calyptraster personatus Madsen, 1947: 3–7, figs 1–2.
MZUSP (without registration number), 1spec., W Besnardi, dredging 5142. MZUSP (without registration number), 2spec., W Besnardi, dredging 5363.
Recife, Pernambuco, Brazil (
Body pentagonal (Fig.
Some species of the order Valvatida recorded in northeastern Brazil. Oreaster reticulatus (A–G)A Abactinal view B Actinal view C Detail view da abactinal surface D Detail of the mouth E Actinal view of the arm F Detail of the bivalve pedicellariae, and G Skeleton, in detail its arrangement into a reticulum.
Colour. From light brown to hey-coloured (
The Bahamas, Florida, and Brazil (
The genus Calyptraster presently contains five species (
This species is quite rare, from deep waters, with little known of its biology and ecology. The species was collected in bottoms containing red mud (
Linckia guildingii Gray, 1840: 285.
Linckia pacifica Gray, 1840: 285.
Ophidiaster ornithopus Müller & Troschel, 1842: 31
Ophidiaster ehrenbergi Müller & Troschel, 1842: 31
Linckia ornithopus Verrill, 1867: 344.
Linckia nicobarica Lütken, 1872: 265.
Linckia ehrenbergi Loriol, 1885: 31.
Linckia guildingi Verrill, 1907: 325.
Paraíba: 6°29'S; 34°48'W, 1spec., UFPB/ECH.1235, 04.VI.1981, 30m; 6°29'S; 34°51'W, 1spec., UFPB/ECH.1244, 05.VI.1981, 22m; 6°33'S; 34°51'W, 1spec., UFPB/ECH.1243, 04.VI.1981, 20m; 7°01'S; 34°30'W, 1spec., UFPB/ECH.1856, 13.II.1981, 26m; 7°01'S; 34°30'W, 3spec., UFPB/ECH.1238, 13.II.1981, 26m; 7°04'S; 34°44'W, 1spec., UFPB/ECH.1237, 16.II.1981, 16m; 7°12'5"S; 34°36'W, 1spec., UFPB/ECH.1241, 01.IV.1981, 26m; 7°28'S; 34°34'W, 1spec., UFPB/ECH.1242, 06.V.1981, 30m; João Pessoa, Cabo Branco Beach, 1spec., UFPB/ECH.1159, 19.II.2003. 1spec., UFPB/ECH.1160, 04.XI.2006, 1spec., UFPB/ECH.1161, 06.X.1979; 3spec., UFPB/ECH.1236, 25.X.2007, 1spec., UFPB/ECH.1245, 16.V.2007, 1spec., UFPB/ECH.1246, 19.IV.2005; 1spec., UFPB/ECH.1247, 08.IX.2006, 2spec., UFPB/ECH.1248, 25.X.2007, 1spec., UFPB/ECH.1250, 03.VII.2004, 1spec., UFPB/ECH.1479, 13.XI.2008, 1spec., UFPB/ECH.1502, 08.II.2009, 1spec., UFPB/ECH.1867, III.2007. Pernambuco: Paulista, Pau Amarelo reef, 1spec., UFPB/ECH.1845, 08.XI.1982. Alagoas: Paripueira, Paripueira Beach, 2spec., UFPB/ECH.1851, 01.II.1983; Maceió, Ponta Verde Beach, 1spec., UFSITAB-199, I.2007, 1spec., UFSITAB-200, I.2007; Marechal Deodoro, Francês Beach, 5spec., UFPB/ECH.1855, 29.I.1983, 2spec., UFPB/ECH.1847, 20.XII.1984, 1spec., UFPB/ECH.1849, 19.II.1985, 2spec., UFPB/ECH.1857, 19.II.2011. Bahia: Salvador, Itapoã Beach, 1spec., UFPB/ECH.1848, 21.XII.1984; Itaparica, Pedrão, 4spec., UFPB/ECH.1853, 18.IX.1982; Itaparica, Barra Grande, 1spec., UFPB/ECH.1854, 17.IX.1982; Santa Cruz da Cabrália, Ponta da Coroa Vermelha, 5spec., UFPB/ECH.1846, 15.X.1982; Santa Cruz da Cabrália, Ponta do Mutá reef, 5spec., UFPB/ECH.1850, 16.X.1982; Porto Seguro, Ponta Grande reef, 5spec., UFPB/ECH.1852, 15.X.1982.
Saint Vincent, West Indies (
Disk small. From four to six long, thin, cylindrical arms (Fig.
Some species of the order Velatida(A–H) and Forcipulatida(I–M) recorded in northeastern Brazil. Calyptraster coa (A–H)A Abactinal view, in detail the supradorsal membrane B Actinal view C Detail of the paxillae with long peduncles D Actinal view of the arm E Detail of the mouth F Detail of the actinal intermediate area G Abactinal view, in detail oscular valves H Actinal view; Coscinasterias tenuispina (I–M)I Abactinal view J Actinal view L Abactinal view of the arm, in detail the bivalve pedicallariae (optical microscopic image), and M Lateral view of the arms, in detail the bivalve pedicellariae (optical microscopic image).
Colour. Extremely variable, usually juveniles and adults having different colours (
Tropicopolitan (
Two species of the genus Linckia are known for the Brazilian coast, L. guildingi and L. nodosa Perrier, 1875, the latter recorded only for south and southeastern Brazil.
This species lives in environments with consolidated substrates or sand banks among reefs (
Narcissia trigonaria Sladen, 1889: 414, pl. 65, figs 5–8.
Narcissia trigonaria var. helenae Mortensen, 1933: 429.
Alagoas: Lagoa Azeda, Jequidá da Praia, 1spec., MNRJ (no registration number), 22.VI.2002. Bahia: Salvador, north coast, 1spec., UFBA00570, 2003; Salvador, Porto da Barra, 1spec., UFBA00929, II.2008, 23m; Salvador, Barra Beach, 1spec., UFBA00962, X.2008; Itaparica, Ponta de Areia, 12°52'S; 38°40'W, 1spec., UFBA00469; Camaçari, Guarajuba, 1spec., UFBA00190, VII.2005, 23m; Camaçari, Guarajuba, Busca Vida Beach, 2spec., UFBA00042, 04.VI.1994, 1spec., UFBA01043, VII.2008, 26m, 1spec., UFBA01089, VII.2008, 32m.
Bahia, Brazil (
Disk high and pyramidal (Fig.
Some species of the order Velatida recorded in northeastern Brazil. Linckia guildingi (A–E)A Abactinal view B Actinal view C Abactinal view of the arm D Actinal view of the arm E Detail of the mouth; Narcissia trigonaria (F–J)F Abactinal view G Actinal view, in detail the mouth H Lateral view I Abactinal view of the arm, and J Actinal view of the arm.
Colour. Live specimens are cream-coloured with red spots (
North Carolina, Florida, Panama, Colombia, and Brazil (
Narcissia trigonaria is a well established species, with a small list of synonyms and little morphological variation. It differs from N. canariensis (d’Orbigny, 1839) for having subambulacral spines arranged into three series and paired papulae.
The species lives in consolidated substrates, either rocks or coral (
Echinaster brasiliensis Müller & Troschel, 1842: 22.
Echinaster braziliensis Verrill, 1915: 41–42, pl. 26, fig. 1.
Echinaster antonioensis Bernasconi, 1955: 72–73, pl. 6, figs 1–2.
Echinaster sentus Bernasconi, 1956: 136–137, pl. 4, fig. 3.
Echinaster spinulosus Bernasconi, 1956: 138–139, pl. 4, fig.4.
Echinaster densispinulosus Tommasi, 1970: 18–19, figs 49–51.
Echinaster nudus Tommasi, 1970: 18–19, figs 52–54.
Echinaster (Othilia) brasiliensis Clark & Downey, 1992: 21–22, pl. 4a.
Rio Grande do Norte: Macau, Diogo Lopes, 1spec., UFPB/ECH.869, 09.XI.2007, 1spec., UFPB/ECH.872, 09.XI.2008, 1spec., UFPB/ECH.1426, 09.XI.2007. Paraíba: 7°01'02"S; 34°47'55"W, 1spec., UFPB/ECH.571, 03.VI.2003; 7°03'50"S; 34°47'19"W, 1spec., UFPB/ECH.569, 21.III.2006; Cabedelo, Farol de Cabedelo reef, 1spec., UFPB/ECH.729, 26.X.1980; Cabedelo, Areia Vermelha reef, 1spec., UFPB/ECH.1465, 22.II.2008; João Pessoa, Cabo Branco Beach, 2spec., UFPB/ECH.138, 04.XI.2006; João Pessoa, Seixas reef, 1spec., UFPB/ECH.1183, 22.XII.2007. Bahia: Santo Amaro, Cabuçu Beach, 3spec., UFPB/ECH.718, 19.IX.1987.
Ubatuta, São Paulo, Brazil (
Disk small (Fig.
Some species of the order Spinulosida recorded in northeastern Brazil. Echinaster (Othilia) brasiliensis(A–E)A Abactinal view B Actinal view C Abactinal view of the arm D Actinal view of the arm E Arranjo do endoqesquelto do braço; Echinaster (Othilia) echinophorus(F–J)F Abactinal view G Actinal view H Abactinal view of the arm I Actinal view of the arm; and H Arranjo do endoesqueleto do braço.
Colour. Quite variable, being light brown, reddish-brown, dark red or even yellowish red (
Florida, Cuba, Honduras, Panama, Colombia, Brazil, and Argentina (
Many characters of Echinaster (O.) brasiliensis and other species of the genus present great plasticity, with the consequence that the taxonomy of the genus cannot be considered well resolved. E. (O.) brasiliensis differs from E. (O.) echinophorus for presenting a larger number of rows of dorsal longitudinal spines (9–15) and actinal figs not mammiform. E. (O.) sentus differs for having more numerous and shorter dorsal spines and for having mammiform abactinal figs.
This species lives in sand, substrates of sand with mud, and consolidated substrates, often associated with the bivalves Mytillus sp. and Lithophaga sp. (
Asterias spinosa Retzius, 1805: 18.
Asterias echinophora Lamarck, 1816: 560.
Stellonia spinosa Nardo, 1834: 716.
Othilia spinosa Gray, 1840: 281.
Echinaster spinosus Müller & Troschel, 1842: 22.
Echinaster (Othilia) crassispina Verrill, 1868: 368.
Echinaster crassispinus Lütken, 1872: 285.
Echinaster echinophorus Perrier, 1875: 100–102.
Othilia echinophora Fisher, 1919: 432.
Echinaster (Othilia) echinophorus Clark & Downey, 1992: 367–371.
Rio Grande do Norte: Macau, Diogo Lopes, 4spec., UFPB/ECH.871, 09.XI.2007; Tubarão River, 1spec., UFPB/ECH.1905,14.XI.2009; Tubarão River Mangrove, 1spec., UFPB/ECH.1904, 31.I.2011; 1spec., UFPB/ECH.1913, 04.IX.2010; Mangrove on highway to Galinhos, 1spec., UFPB/ECH.1914, 22.VI.1982. Paraíba: 7°03'50"S; 34°47'19"W, 2spec., UFPB/ECH.568, 21.III.2006; Lucena, Fagundes Beach, 2spec., UFPB/ECH.717, 22.IX.1995, 1spec., UFPB/ECH.728, 22.IX.1985; Cabedelo, Areia Vermelha reef, 2spec., UFPB/ECH.1464, 06.IV.2008; Cabedelo, Poço Beach, reefs facing Ponta de Campina, 1spec., UFPB/ECH.1903, 28.II.2010; João Pessoa, 7°4'30” S; 34°46'56”, 1spec., UFPB/ECH.725, 26.IV.2005; João Pessoa, Cabo Branco Beach, 3spec., UFPB/ECH.704, 2002, 4spec., UFPB/ECH.705, 9spec., UFPB/ECH.706, 04.IV.1981, 4spec., UFPB/ECH.707, 03.VII.1985, 1spec., UFPB/ECH.708, 17.II.1980, 1spec., UFPB/ECH.710, 01.IV.2006, 1spec., UFPB/ECH.711, V.1980, 2spec., UFPB/ECH.713, 06.XI.1983, 10spec., UFPB/ECH.714, 29.IV.2002, 3spec., UFPB/ECH.715, 13.XII.1985, 1spec., UFPB/ECH.719, 22.IX.1980, 3spec., UFPB/ECH.722, 09.II.2001, 1spec., UFPB/ECH.727, 09.IX.2006, 3spec., UFPB/ECH.870, 25.XI.2007, 1spec., UFPB/ECH.1240, 22.IX.1980, 2spec., UFPB/ECH.1466, 13.XI.2008, 1spec., UFPB/ECH.1911, 21.III.2000; João Pessoa, Ponta Seixas, 6spec., UFPB/ECH.139, 03.XI.1982, 3spec., UFPB/ECH.703, 23.X.1984, 2spec., UFPB/ECH.712, 25.V.1998; Seixas reef, 1spec., UFPB/ECH.1172, 23.III.2008, 6spec., UFPB/ECH.1234, 03.XI.1982. 4spec., UFPB/ECH.1184, 22.XII.2007, 2spec., UFPB/ECH.1463, 12.I.2009. Pernambuco: Goiana, Catuama Beach, 1spec., UFPB/ECH.1912, 31.X.1982. UFPB/ECH.873; Tamandaré, Carneiros Beach, 4spec., 15.X.1981. Alagoas: Marechal Deodoro, Francês Beach, 1spec., UFPB/ECH.721, 29.I.1983. Bahia: Itaparica, Pedrão de Itaparica, 8spec., UFPB/ECH.716, 18.IX.1982; Itaparica, Ponta de Aratuba, 7spec., UFPB/ECH.723, 19.X.1982; Itaparica, Barra Grande, 2spec., UFPB/ECH.726, 19.IX.1982; Prado, coral reef between Camaruxatiba and Ponta de Imbaçuaba, 3spec., UFPB/ECH.709, 14.X.1982, 2 spec., UFPB/ECH.724, 28.XI.1982; Santa Cruz da Cabrália, Ponta da Coroa Vermelha, 5spec., UFPB/ECH.720, 15.X.1982.
‘Amérique du Nord’ (lectotype) (
Disk small (Fig.
Colour. Red, orange-red, becoming brownish-red when conserved in alcohol and dark brown when dry (
Florida, the Bahamas, Gulf of Mexico, Puerto Rico, Jamaica, Nicaragua, Colombia, Venezuela, and Brazil (
Some common starfishes in their natural habitat. A Luidia senegalensis in a sand beach B Astropecten marginatus in a hypersaline mangrove C Oreaster reticulatus in a sand beach D Linckia guildingi on coral reefs EEchinaster (O.) echinophorus in seagrass beds and FE. (O.) echinophorus over the muddy bottom of a hypersaline mangrove. Photos: Thelma LP Dias.
Echinaster (O.) echinophorus differs from E. (O.) sentus for having few large and conspicuous spines on arms and a uniform colour (
Lives usually in environments containing consolidated substrates (
Asterias tenuispina Lamarck, 1816: 561–562.
Asteracanthion tenuispinus Müller & Troschel, 1842: 16.
Asterias atlantica Verrill, 1868: 368.
Asterias (Stolasterias) tenuispina Sladen, 1889: 565, 583.
Polyasterias tenuispina Perrier, 1894: 108.
Lytaster inaequalis Perrier, 1894: 98–99.
Coscinasterias tenuispina Verrill, 1914: 45.
Coscinasterias tenuispina var. atlantica Verrill, 1915: 20–21.
Stolasterias tenuispina Verrill, 1907: 324.
Coscinasterias (Stolasterias) tenuispina Fisher, 1926: 197.
Rio de Janeiro: Cabo Frio, Formoso Beach, 1 spec., MZUSP (without registration number), VII.1956; Cabo Frio, Arraial do Cabo, Brava Beach, 1spec., MZUSP (without registration number), 29.I.2001.
‘I’ ocean eropéen’ (
Disk small, with 1–3 madreporites (in some cases up to 5 were observed). Six to nine (rarely 5 and unusually 7) thin, elongate (Fig.
Colour. Specimens from Brazil vary from brown to orange colour (
North Carolina, Gulf of Mexico, Antilles, Bermudas, Cuba, Brazil, Portugal, Spain, France and Montenegro (
Two species of the genus Coscinasterias Verrill, 1870 are known for the Atlantic, C. tenuispina and C. calamaria (Gray, 1840). The first is widely distributed through the Atlantic and Mediterranean, while the second is restricted to South Africa, Angola, and Madagascar (
Lives in consolidated substrates, including areas with strong hydrodynamism (
The fauna of Asteroidea recorded for northeastern Brazil is composed mainly by species with broad geographical and bathimetic distributions, and considered common species for the Brazilian littoral (
Among the four recorded orders, Paxillosida was the nost diverse (n = 10 spp), followed by Valvatida (n = 5 spp), Velatida (n = 3 spp), Spinulosida (n = 2 spp) and Forcipulatida (n = 1 spp). These results were expected, because Paxillosida represents the most diverse order and contains the most speciose genus (Astropecten with 150 spp) and abundant species in shallow waters (
The genus Astropecten represents one of the most complex taxa within the class Asteroidea, in which species exibit great morphological plasticity, making identification of species difficult. According to
Another taxonomicly complex genus is Echinaster that, similarly to Astropecten, presents large morphological variability, making species identification difficult. According to
In terms of diversity, the fauna of asteroids in northeastern Brazil represent only 27% of the species known for Brazil, the coast of Bahia (n = 14 spp) and Paraíba (n = 12 spp) being the most diverse, followed by Pernambuco (n = 9 spp), Ceará and Rio Grande do Norte (both with 6 spp), Alagoas (n = 4 spp), and Piauí (n = 2 spp). There are still no records of asteroids for the coasts of Maranhão and Sergipe. These numbers indicate the scarce knowledge available on the Asteroidea from northeast Brazil, which represents one of the least known areas regarding the benthic macrofauna (
Presently 374 species of sea-stars are known for the Atlantic Ocean (
Another three species recorded previously for northeast Brazil were not confirmed in the present study: Allostichaster hartti (as Leptasterias hartii Rathbun, 1879) was recorded as a rare species for the littoral of the State of Bahia (
Due to the negligible previous knowledge on the Asteroidea from the littoral of northeastern Brazil, the present study represents an important re-evaluation of the diversity of this group for this area. It should further be noted that the study of material deposited in scientific collections turned out to be of fundamental importance, permitting a historical panorama of the Asteroidea from northeast Brazil. Deep-water sampling in the studied region, restricted to sporadic expeditions, as accounted for in
We are grateful to Dr. Carlos R. R. Ventura (MNRJ), Aline Benetti (MZUSP), Carla M. Menengola (MZUFBA) and Cynthia L. C. Manso (UFS) for permitting us to examine material deposited under their charge. Special thanks to Rosana Cunha and Luciana Martins for sending photos of some of the studied species, and José Anderson Feijó for making the map. We also thank the logistical support provided by the colleagues Luis Carlos, Mariana Contins, and Carlo Magenta. We are also grateful to the Federal University of Paraíba and to the Post Graduate Program in Biological Sciences (Zoology), for providing the infrastructure enabling this research. Anne I. Gondim was supported by CAPES (the Brazilian Ministry of Education) through a Master’s scholarship and Martin L. Christoffersen is supported by a CNPq productivity research grant (Process number: 300198/2010-8). We wish to express our sincere gratitude to two anonymous reviewers for their critical reading of the manuscript and constructive comments. We also thank all the people who somehow contributed in this study.