Corresponding author: Anne Isabelley Gondim (
Academic editor: Y. Samyn
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:
Gondim AI, Christoffersen ML, Dias TLP (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys 449: 1–56. doi:
The
The first paper to deal with the
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
Northeastern Brazil has a coastline with 3,400 km in extension (
Although the first record of the
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 (
Map of the Americas showing the northeast region of Brazil divided in the nine states.
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.
Some littoral environmnets and shallow-water biotopes used as habitats by sea stars in northeastern Brazil.
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
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
Paraíba:
Dry Tortugas, Florida Keys, Florida (
Body pentagonal (Fig.
Some species of the family
North Carolina, the Bahamas, Florida, Gulf of Mexico, Cuba, Jamaica, Honduras, Puerto Rico, Panama, Colombia, Venezuela, Brazil, Uruguay, and Argentina (
The main characters distinguishing
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 (
Paraíba:
Probably no longer existant (
Five long and narrow arms (Fig.
Bermuda, Gulf of Mexico, Belize, Honduras, Nicaragua, Panama, Colombia, Venezuela, and Brazil (
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 (
Paraíba:
Rio de Janeiro, Brazil (
Body flattened. Five arms that taper gradually towards their extremities (Fig.
Some species of the family
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.
This subspecies occurs in non-consolidated sediments containing fine or coarse sand (
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.
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.
The species lives in environments of low hydrodynamism, in sediment containing sand, mud, or a combination of both of these (
Paraíba:
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
North Carolina, the Bahamas, Florida, Honduras, Nicaragua Colombia, and Brazil (
Usually inhabits muddy environments, but may also be found associated to communities of azooxanthelate corals (
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.
North Carolina, Florida, the Bahamas, Mexico, Cuba, Puerto Rico, Dominican Republic, Panama, Colombia, Venezuela, Brazil, and Uruguay (
Inhabits sandy environments. According to
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.
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
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 (
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.
North Carolina, the Bahamas, Gulf of Mexico, Mexico, Nicaragua, Costa Rica, Panama, Colombia, Brazil, Uruguay, Argentina, and Africa (
This species lives on sandy or muddy bottoms of the littoral region up to depths of 50 m (
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,
Unknown (
Body flattened. Five broad, triangular arms (Fig.
Some species of the family
Costa Rica, Puerto Rico, Colombia, Venezuela, Guyana, and Brazil (
This species lives in substrates containing sand or sand with mud (
Paraíba:
Saint Thomas, Virgin Islands (
Body inflated, pentagonal (Fig.
Bermudas, Florida, Bahamas, Belize, Panama, and Brazil (
Only two species of the genus
This species lives in association with coral reefs, being found particularly under rocks or corals of the reef flat (
Paraíba:
Unknown (
Disk small (Fig.
Mexico, Cuba, Nicaragua, Brazil, Indo-Pacific (except Hawaii) (
This species lives on hard substrates covered by incrusting organisms and in reef gravel (
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;
‘Las Antillas Occidentales’ or ‘Spanish West Indies’ (
Body pentagonal. Disk high, inflated, concave on actinal side (Fig.
Some species of the order
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
The species lives in shallow reef environments with calm water, coastal lagoons, seagrass beds (
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
The Bahamas, Florida, and Brazil (
The genus
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 (
Paraíba:
Saint Vincent, West Indies (
Disk small. From four to six long, thin, cylindrical arms (Fig.
Some species of the order
Tropicopolitan (
Two species of the genus
This species lives in environments with consolidated substrates or sand banks among reefs (
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,
Bahia, Brazil (
Disk high and pyramidal (Fig.
Some species of the order
North Carolina, Florida, Panama, Colombia, and Brazil (
The species lives in consolidated substrates, either rocks or coral (
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:
Ubatuta, São Paulo, Brazil (
Disk small (Fig.
Some species of the order
Florida, Cuba, Honduras, Panama, Colombia, Brazil, and Argentina (
Many characters of
This species lives in sand, substrates of sand with mud, and consolidated substrates, often associated with the bivalves
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:
‘Amérique du Nord’ (lectotype) (
Disk small (Fig.
Florida, the Bahamas, Gulf of Mexico, Puerto Rico, Jamaica, Nicaragua, Colombia, Venezuela, and Brazil (
Some common starfishes in their natural habitat.
Lives usually in environments containing consolidated substrates (
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.
North Carolina, Gulf of Mexico, Antilles, Bermudas, Cuba, Brazil, Portugal, Spain, France and Montenegro (
Two species of the genus
Lives in consolidated substrates, including areas with strong hydrodynamism (
The fauna of
Among the four recorded orders,
The genus
Another taxonomicly complex genus is
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
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:
Due to the negligible previous knowledge on the
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.