Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea)

Abstract

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.

Full text

Taxonomic guide and historical review of starshes in northeastern Brazil... 1
Taxonomic guide and historical review of starfishes in
northeastern Brazil (Echinodermata, Asteroidea)
Anne Isabelley Gondim1, Martin Lindsey Christoersen1,
elma Lúcia Pereira Dias2
1 Universidade Federal da Paraíba, Programa de Pós-Graduação em Ciências Biológicas (Zoologia), Depar-
tamento de Sistemática e Ecologia, Laboratório de Invertebrados Marinhos Paulo Young, Bairro Cidade Uni-
versitária s/n, CEP. 58059-900, João Pessoa, PB, Brasil 2 Universidade Estadual da Paraíba, CCBS, Depar-
tamento de Biologia, Laboratório de Biologia Marinha, Campus I, Rua Baraúnas, 351, Bairro Universitário,
CEP 58429-500, Campina Grande, PB, Brasil
Corresponding author: Anne Isabelley Gondim (anneisabelley@yahoo.com.br)
Academic editor: Y. Samyn|Received 15 December 2013|Accepted 8 October 2014|Published 22 October 2014
http://zoobank.org/75DDC584-63EB-4BF1-BBF9-08C1D2954CAC
Citation: Gondim AI, Christoersen ML, Dias TLP (2014) Taxonomic guide and historical review of starshes in
northeastern Brazil (Echinodermata, Asteroidea). ZooKeys 449: 1–56. doi: 10.3897/zookeys.449.6813
Abstract
Presently more than 1900 species of sea stars are recognized, of which 77 are recorded for the coast of
Brazil. Although the rst starsh 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 north-
eastern regions of the country. is study provides the rst 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 identied, belonging to 12 genera,
10 families, and 5 orders. Descriptions of species are provided. ree 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 rst 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.
Keywords
Echinoderms, taxonomy, check list, sea-stars, northeastern Brazil
ZooKeys 449: 1–56 (2014)
doi: 10.3897/zookeys.449.6813
http://zookeys.pensoft.net
Copyright Anne Isabelley Gondim et al. 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.
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Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
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Introduction
e Asteroidea represent the second most diverse group within the phylum Echino-
dermata, with an estimated number of 1900 living species (Mah and Blake 2012). Of
these, 77 species are recorded for the Brazilian coastline (Ventura et al. 2012).
e rst 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 (Hadel et al. 1999). Among the diverse
groups described and illustrated by Marcgraf are two species of starsh (Luidia senegalen-
sis as Stella senegalensis, and Oreaster reticulatus). Later C. F. Hartt collected 16 species of
echinoderms from Abrolhos (Bahia), of which four were asteroids (Oreaster reticulatus,
Linckia guildingi, Echinaster (Othilia) echinophorus, and Coscinasterias tenuispina) (Verrill
1868). Between 1875 and 1877 this same author collected another 62 species of echi-
noderms, which were described by Rathbun (1879). at paper furnished the rst list of
echinoderms from Brazil and in it 12 species of asteroids were recorded.
irty six years after the paper by Rathbun (1879), Verrill (1915) published a new
list with descriptions and illustrations of 125 species of sea-stars from the West Indies,
Florida, and Brazil. Later H. P. Oliveira (1940) furnished a new list of the Brazilian as-
teroids. Since then, several contributions on the marine benthic fauna were published
by diverse authors, which increased our knowledge of the Brazilian asteroids. Concom-
itantly, several papers were published on the Echinodermata from Brazil by Dr. Luiz
Roberto Tommasi. e paper published by Tommasi (1970) is his main contribution
to the Class Asteroidea. In this work he produced descriptions and illustrations of the
42 species then known for the coast of Brazil.
After these pioneer studies, several works focusing on taxonomy, ecology and ge-
ographical distribution were developed, mainly in the south and southeast (e.g. Brito
1960, Carrera-Rodríguez and Tommasi 1977, Monteiro and Pardo 1991, Ventura and
Fernandes 1995, Ventura et al. 1997, Nobre and Campos-Creasey 2000, Alves et al.
2002, Carvalho and Ventura 2002, Gibran 2002, Calil et al. 2009, Mariante et al. 2010).
For northeast Brazil, only three papers have focused on the Asteroidea: Lima-Verde
and Matthews (1969), who studies the feeding habits of Luidia senegalensis in the State
of Ceará; Manso (2006), who recorded the rst Goniasteridae for the Potiguar Basin in
the Cretaceous of Brazil, and Matthews and Lima-Verde (1969), who furnished eco-
logical information on Oreaster reticulatus from the northeastern region. Knowledge on
diversity along the northeastern littoral stems from species inventories. Among these:
Lima-Verde (1969) recorded seven species along Ceará, Rio Grande do Norte, Per-
nambuco and Alagoas; Tommasi (1970) recorded six species for several northeastern
states; Tommasi and Aron (1988) expanded to ten the known species from Bahia; Fer-
nandes et al. (2002) record six species from Pernambuco; Magalhães et al. (2005) cited
eight species for Bahia; Gondim et al. (2008) recorded ve species for Paraíba; Gondim
and Giacometti (2010) and Gondim et al. (2013) signaled two species for the coast of
Piauí; and Miranda etal. (2012) recorded eight species from Alagoas.

Taxonomic guide and historical review of starshes in northeastern Brazil... 3
Northeastern Brazil has a coastline with 3,400 km in extension (Pinheiro et al.
2008), which represents 42.5% of the entire Brazilian coastline. is region contains a
great variety of ecosystems and has one of the largest reef environments of the South-
western Atlantic Ocean. Notwithstanding, many areas remain unexplored (Marques
and Lamas 2006) and our knowledge of the macrozoobenthos below 20 m is still limit-
ed, mainly regarding the outer platform and the continental slope (Migotto and Tiago
1999). Brazil has organized few marine expeditions (an example being the Programa
de Avaliação do Potencial Sustentável de Recursos Vivos na Zona Econômica Exclu-
siva - REVIZEE), but most results continue concentrated in the south and southeast-
ern regions, the addition of information for the north and northeastern regions being
minimal below 20 m depth (Marques and Lamas 2006).
Although the rst 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. e aim of the present work is to expand our knowledge on the morphology
of the species of Asteroidea from northeastern Brazil.
Materials and methods
Species determinations were based mainly on Tommasi (1970), Clark and Downey
(1992), and Hendler et al. (1995). Synonyms were compiled from Tommasi (1970),
Clark and Downey (1992), and Mah (2013). We provide a reference to the rst work
mentioning the synonym and to the main papers on the Brazilian fauna that subse-
quently cite the species.
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.
e studied material is conserved in 70% alcohol or preserved dry and registered
in the visited collections.
Study area
e littoral region of northeast Brazil extends for approximately 3,400 km (Pinheiro
et al. 2008), beginning in the Parnaíba River Delta and extending rst to the east and
then to the south up to the border of between the States of Bahia and Espírito Santo.
e region encompasses nine coastal States: Maranhão, Piauí, Ceará, Rio Grande do
Norte, Paraíba, Pernambuco, Alagoas, Sergipe, and Bahia (Fig. 1).
e region has a great diversity of littoral environments, such as beaches, dunes,
clis, estuaries, deltas, sandbanks, reefs and underwater biotopes, such as rhodolith beds,
seagrass beds and coral habitats (Fig. 2). ey share Tertiary sediments from the Barreiras
Formation, beach rocks, and coral reefs (Suguio 2003). is coastal diversity, allied to

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
4
the favorable climatic conditions and year-round warm waters, makes this region one of
the most important tourist destinations in the country (Pinheiro et al. 2008).
Due to the absence of large rivers and to the predominance of warm waters from
the South Equatorial Current, oshore conditions in the northeast region are ideal
for the formation of coastal coral reefs (Maury 2002). e most common formations
along the coast are arenitic beach rocks (Mabesoone and Coutinho 1970), which har-
bour scleractinian corals and calcarious algae. Reefs composed exclusively of calcarious
algae and corals also occur commonly. At the South of the State of Bahia, singular
mushroom-shaped coral-reef formations, denominated locally as “chapeirões”, are pre-
sent oshore (Abrolhos region).
e northeastern continental shelf is narrow and shallow. Due to the reduced con-
tinental inuence 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) (Coutinho 2006). e platform ends very abruptly
around depths of 60–80 m. e isobath of 20 m coincides mostly with the extent of
the platform. e continental shelf may reach an extent of 60 miles at Cape São Roque
(RN), and less than 5 miles in front of Recife (PE) (Kempf et al. 1970).
e biogenic carbonates form sand and gravel, consisting mainly of incrusting and
ramied calcareous algae, with local occurrences of Halimeda spp. ese sediments may
contain up to 5% of carbonatic mud derived from the desintegration of larger organ-
isms. In certain areas, these sediments undergo a process of litication, also aecting the
quartsoze sands of the inner platform, originating the extensive coastal reefs known as
beach rocks. ese then become overgrown by algae and corals (Lana et al. 1996).
Figure 1. Map of the Americas showing the northeast region of Brazil divided in the nine states. MAMa-
ranhão PI Piauí CE Ceará RN Rio Grande do Norte PB Paraíba PE Pernambuco AL Alagoas SESergipe,
and BA Bahia.

Taxonomic guide and historical review of starshes in northeastern Brazil... 5
Figure 2. Some littoral environmnets and shallow-water biotopes used as habitats by sea stars in north-
eastern Brazil. A Fringing reef B Beach rocks C Algae banks D Rhodolith bed E Seagrass bed F Patch
reef, and G Mangroves. Photos: elma LP Dias.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
6
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.
Results
A total of 21 species, belonging to ve orders, 10 families and 12 genera were identi-
ed. ese are listed and described below, following the taxonomic organization of
Clark and Downey (1992).
Checklist of Starshes from northeastern Brazil
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

Taxonomic guide and historical review of starshes in northeastern Brazil... 7
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)
Systematics
Order Paxillosida Perrier, 1884
Family Luidiidae Sladen, 1889
Luidia alternata alternata (Say, 1825)
Figure 3a–d
Asterias alternata Say, 1825: 144–145.
Luidia alternata Lütken, 1859: 42–43. Brito 1968: 12–13, pl. 3, g. 4. Tommasi 1970:
8, g. 24. Tommasi and Aron 1987: 5. Tommasi et al. 1988: 6. Ventura et al. 2007:
236. Miranda et al. 2012: 9.
Luidia granulosa Perrier, 1869: 109–110, pl. 2, g. 18.
Luidia variegata Perrier, 1875: 337.
Luidia numidica Koehler, 1911: 3, pl. 1, gs 8–11.
Luidia quequenensis Bernasconi, 1942: 253. Tommasi 1970: 8, g. 23. Carrera-Rodri-
guez and Tommasi 1977: 62, 65.
Luidia bernasconiae A.H.Clark, 1945: 19–21.
Luidia alternata var. numidica Madsen, 1950: 206–209, g. 9.
Luidia alternata numidica A.M.Clark, 1953: 388–389, pl. 41, g. 1.
Luidia alternata alternata Clark & Downey, 1992: 8–9; Magalhães et al. 2005: 63.
Material examined. 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.
Type locality. Dry Tortugas, Florida Keys, Florida (Clark and Downey 1992)–
Neotype.
Description. Body pentagonal (Fig. 3a, b). Five elongate and thin arms. Abactinal
surface with paxillae (Fig. 3a). Carinal paxillae smaller than adradial paxillae, with 1–4

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
8
Figure 3. 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.

Taxonomic guide and historical review of starshes in northeastern Brazil... 9
blunt central spinelets and with one or two series of marginal spinelets, numbering
12–20 per series. One of the central spines is slightly longer than remaining spines.
Paxillae rounded and arranged into regular transversal rows. Some adradial paxillae
with one long, conical and pointed central spine (~1.90 mm) (Fig. 3c). Inferomarginal
plate with 1 or 2 long, thin spines (~1.76 mm), placed vertically and forming a well
dened marginal row. Actinal surface with inferomarginal plates densely covered with
spines of diverse shapes and sizes, there being 1–3 longer spines located centrally (Fig.
3c, d). Adambulacral plate with 4 spines placed vertically. e adambulacral spine is
the smallest, being slightly curved and compressed. e two subambulacral spines are
subequal in shape and size. A short spine is located laterally to the most external spine.
Oral spines narrow and elongate. With bi, tri and tetravalvular pedicellariae on actinal
surface (Fig. 3b).
Colour. Abactinal surface white or cream-coloured, with dark transversal bands.
e colour of these bands varies, and may be brown, black, green, or purple. Actinal
surface whitish or beige.
Distribution. North Carolina, the Bahamas, Florida, Gulf of Mexico, Cuba,
Jamaica, Honduras, Puerto Rico, Panama, Colombia, Venezuela, Brazil, Uruguay,
and Argentina (Bernasconi 1943, Tommasi 1958, Downey 1973, Walenkamp 1976,
Clark and Downey 1992, Hendler et al. 1995, Ventura et al. 2007, Alvarado et al.
2008). In Brazil from AL, BA, RJ, and SP (Verrill 1915, Brito 1962, 1968, Tommasi
1970, Magalhães et al. 2005, Miranda et al. 2012). is is the rst record of the species
for the coast of Paraíba. From 1 to 200 m in depth (Clark and Downey 2002), most
common between 10 and 30 m.
Remarks. e 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. e abactinal colouration
is distinctive. e nominal subspecies diers from its congeneric form Luidia alternata
numidica Koehler, 1911, from West Africa, for attaining a larger size and for having
longer paxillar spines. Clark (1982) provided a good discussion on the synonymy of
the nominal subspecies and designates a neotype. Specimens analysed in this study
are young individuals that may attain 200 mm along its larger ray. Furthermore, our
specimens presented no morphological variations, agreeing with the descriptions of
this species. Despite not receiving much attention in recent taxonomic surveys, pedi-
cellariae are an important taxonomic character to distinguish species of asteroids. Clark
(1982) and Clark and Downey (1992) furnished excellent illustrations of the species of
Luidia known from the Atlantic.
Ecological notes. is 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 (Clark and Downey 1992, Benavides-Serrato et al. 2011). According
to Hendler et al. (1995), the subspecies does not occur in large numbers, but is often
found among the accompanying fauna in trawling nets. L. alternata alterna is carnivo-
rous, feeding on epifaunal organisms, especially other echinoderms.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
10
Luidia clathrata (Say, 1825)
Figure 3e–h
Asterias clathrata Say, 1825: 142.
Luidia clathratta Lütken, 1859: 37–39. Rathbun 1879: 150. Bernasconi 1943: 6–7.
Tommasi 1970: 8. Magalhães et al. 2005: 63.
Luidia clathrata Lütken, 1859: 37. Tommasi 1958: 9, pl. 2, g. 1; 1970: 8, g. 22.
Brito 1962: 4; 1968: 11–12, pl. 2, g. 2. Carrera-Rodriguez and Tommasi 1977:
63–64. Tommasi and Aron 1987: 3. Tommasi et al. 1988: 6. Magalhães et al.
2005: 63. Ventura et al. 2007: 237. Manso et al. 2008: 185, g. 7a–e. Lima and
Fernandes 2009: 58. Xavier 2010: 75.
Material examined. Paraíba: 6°57'S; 34°41'W, 2 spec., UFPB/ECH.875, 12.II.1981, 26m.
Type locality. Probably no longer existant (Clark and Downey 1992).
Description. Five long and narrow arms (Fig. 3e, f). Abactinal surface paxillar.
Lateral paxillae quadrangular, forming three regular rows (Fig. 3g). Carinal paxillae
small, smaller than lateral paxillae. Dorsal paxillae rounded, small, with 1–6 short,
blunt, central spinelets and 6–18 slender, marginal spinelets. Inferomarginal plates
with two long, pointed, conical spines (~2.58 mm), positioned vertically, the inferior
one the largest. Actinal surface with inferomarginal plates densely covered with at-
tened, lanceolate spines (Fig. 3f). A row of short, actinolateral plates, with 1–3 short,
lanceolate, divergent spines. Ambulacral plates with three spines placed vertically (Fig.
3h). e adambulacral spine is the smallest, being slightly curved and compressed. Of
the two subambulacral spines, the inner one is longer and thicker than the outer spine.
Oral spines long (~1.32 mm), thin and pointed, forming dense tufts on the inner angle
of the jaw (Fig. 3f). Ocular plate granulose and elongate.
Colour. Abactinal surface bluish-gray, frequently with a darker line occupying
the carinal region of the arm and disk. Hendler et al. (1995) cite other colour patters
for the abactinal surface: brown, rose and salmon. Actinal surface white or cream-
coloured.
Distribution. Bermuda, Gulf of Mexico, Belize, Honduras, Nicaragua, Panama,
Colombia, Venezuela, and Brazil (Downey 1973, Clark and Downey 1992, Hendler
et al. 1995, Alvarado et al. 2008, Benavides-Serrato et al. 2005). In Brazil: PE, BA,
RJ, SP, and SC (Rathbun 1879, Bernasconi 1943, Tommasi 1958, 1970, Brito 1960,
1962, Walenkamp 1976, Magalhães et al. 2005, Lima and Fernandes 2009, Xavier
2010). is study records the species for the rst time in the State of Paraíba. From
intertidal to 175 m in depth (Ventura et al. 2007), being most common in depths
under 40 m (Hendler et al. 1995).
Remarks. Luidia clathrata diers from L. alternata alternata for not having the
abactinal surface spinulose. It diers from L. senegalensis for having only 5 arms and
from L. ludwigi scotti for not having pedicellariae. According to Walenkamp (1976),
the number of central spinelets on the paxillae and of inferomarginal spines increases
with ontological development, the maximum numbers being, respectively, 7 and 3. In

Taxonomic guide and historical review of starshes in northeastern Brazil... 11
this study the examined specimen had only two inferomarginal spines, and 1–6 central
spinelets on the paxillae. ese characters indicate a juvenile individual. According
to Hendler et al. (1995), adult individuals may attain 20 to 30 cm in disk diameter.
Knott and Hopkins (1998) recognized two morphotypes of L. clathrata for the Co-
lombian Caribbean, one with a gray colour and the other with three colours, which
were separated by Hopkins and Knott (2010) into L. clathrata and L. lawrencei. ose
authors also established and described a neotype for L. clathrata.
Ecological notes. is species lives in sandy or muddy areas with low hydrody-
namism 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 (Machado et al. 2008, Benavides-Serrato et al. 2011). It feeds on a
great variety of prey, including molluscs, crustaceans, and ophiuroids (Hendler et al.
1995). Like L. alternata and L. senegalensis, Ludia clathrata is host for the small poly-
chaete Podarke obscura Verrill, 1873, that lives in the interior of its ambulacral groove.
L. clathrata may form dense populations, and is thus frequently captured in trawling
nets used in shrimp sheries (McClintock and Lawrence 1985, Hendler et al. 1995).
Presently it is considered to be a species vulnerable to extinction along the Brazilian
coast (Machado et al. 2008).
Luidia ludwigi scotti Bell, 1917
Figure 4a–d
Luidia scotti Bell, 1917: 8–9. Tommasi 1970: 8, g. 25. Carrera-Rodriguez and
Tommasi 1977: 62, 65–66.
Luidia doello-juradoi Bernasconi, 1941: 117; 1943: 8–11. Brito 1962: 3.
Luidia rosaurae Jonh & Clark, 1954: 142–145.
Luidia doello-juradol Brito, 1968: 12, pl. 3, g. 5.
Luidia ludwigi Walenkamp, 1976: 32–37, g. 9, pl. 2, gs 1–3, pl.4, g. 3. Machado
et al. 2008: 179–180. Xavier 2010: 75.
Luidia ludwigi scotti A.M.Clark, 1982: 171–173. Tommasi 1985: 3. Tommasi et al.
1988: 6. Manso 1989: 357.
Luidia rosaurae John & Clark, 1954: 142–145, pl. 6, g. 1.
Material examined. Paraíba: 6°39'05"S; 34°49'W, 1 spec., UFPB/ECH.878,
28.V.1981, 20m.
Type locality. Rio de Janeiro, Brazil (Clark and Downey 1992).
Description. Body attened. Five arms that taper gradually towards their extremi-
ties (Fig. 4a, b). Abactinal surface with paxillae (Fig. 4a). Dorsal paxillae small, ordered,
with 1–6 central, short, blunt spinelets, and 18 marginal, slender, and denticulate
spinelets (Fig. 4c). Adradial paxillae rectangular or quadrangular, slightly larger than
the carinal paxillae. Inferomarginal plates with one pointed, elongate spine (~2.04
mm) forming the marginal fringe (Fig. 4a). e remaining part of the plate is covered

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
12
Figure 4. 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.

Taxonomic guide and historical review of starshes in northeastern Brazil... 13
by short, hyaline and denticulate spinelets. Actinal surface with inferomarginal plates,
densely covered by attened and lanceolate spines (Fig. b, d). Actinolateral plate with
three short, slender, divergent spines. e median of these is the largest. e ambu-
lacral spine long, attened and slender (Fig. 4d). Bivalved pedicellariae present only
on the actinal surface, mainly on the ventro-lateral plates and in the areas close to the
arms, never occurring on the abactinal surface. Oral spines slender, long (~1.18 mm)
with blunt tip, forming a bundle of spines at apex of jaw (Fig. 4c).
Colour. e 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 (Walenkamp 1976). Benavides-Serrato et al. (2011) and Clark and
Downey (1992) observed specimens with dark pink spots in the center of the dorsal
surface of the disc and transversal bands of the same tone on the arms. Oral surface
with colour varying between white and cream. When observed in alcohol they are
uniformly white.
Distribution. Florida, Gulf of Mexico, Colombia, Venezuela, Guyana, French
Guyana, Brazil, and Argentina (Mar del Plata) (Bernasconi 1943, Tommasi 1970,
Clark and Downey 1992, Benavides-Serrato et al. 2005). In Brazil: RJ, SP, and SC
(Brito 1962, 1968, Xavier 2010). is is the rst record of the species for the littoral
of northeast Brazil. From 20 to 126 m in depth. Clark and Downey (1992) considered
the previous record at 5 m for the State of São Paulo to be doubtful. us the present
study expands the bathymetric distribution to 20 m deep, previously established at
33–126 m.
Remarks. is 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. e synonymy between Luidia rosaurae John & Clark, 1954 and L.
scotti was proposed by Clark (1982), who observed that the dierences in the infero-
marginal spines previously established between the two species were insignicant and
unable to support the independence of the two species. Furthermore, with the obser-
vations of Walenkamp (1976) that L. rosaurae is conspecic with L. ludwigi Fisher,
1906, Clark and Downey (1992) recognized the subspecies Luidia ludwigi scotti for
specimens from the Atlantic, as these have a paxillar arrangement that is distinct from
the remaining species previously cited. According to Clark and Downey (1992), the
relationships between L. patriae Bernasconi, 1941 and this subspecies still need to be
investigated. e specimen analised in this study, even though representing a juvenile
individual, did not present signicant morphological variations when compared with
the characters described for adult specimens, indicating that the morphological char-
acters of L. ludwigi scotti do not vary signicantly during ontogeny.
Ecological notes. is subspecies occurs in non-consolidated sediments containing
ne or coarse sand (Machado et al. 2008). In contrast to other taxa of Luidia, which

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
14
do not present prey selectivity, in Luidia ludwigi scotti only eight dierent food types
have been recorded, of which bivalves, foraminiferans and ophiuroids are their main
prey (Benavides-Serrato et al. 2011). e presence of dierent prey species of distinct
sizes in the stomachs of L. ludwigi scotti probably reects a response to competition and
coexistence with other species of sea-stars (Brögger and Penchaszadeh 2008).
Luidia senegalensis (Lamarck, 1816)
Figures 4e–h, 12a
Asterias senegalensis Lamarck, 1816: 567.
Luidia senegalensis Müller & Troschel, 1842: 78. Tommasi 1958: 9, pl. 2, g. 1; 1970:
8, g. 21; 1985: 3. Brito 1968: 10–11, pl. 3, g. 1. Lima-Verde 1969: 10. Nomura
and Fausto Filho 1966: 19. Fernandes et al. 2002: 422. Magalhães et al. 2005: 63.
Manso et al. 2008: 185, g 8c–e. Lima and Fernandes 2009: 58. Xavier 2010: 75.
Luidia marcgravii Steenstrup in Lütken 1859: 43–46.
Material examined. 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. Para-
íba: Lucena, Costinha Beach, 5 spec., UFPB/ECH.1673, 08.XI.2003; Cabedelo, Mi-
ramar 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.
Type locality. Supposedly Senegal (‘L’ océan d’Afrique, les côtes du Senegal’), but
probably West Indies (Clark and Downey 1992).
Description. Body attened. Disk rounded. Nine long and narrow arms (rarely 7)
(Fig. 4e, f). Paxillae on abactinal surface. Paxillae small, arranged irregularly, occupy-
ing center of disk and of arms. Paxillae of carinal regions rounded, with 1–4 central,
short, rounded spinelets and 12–16 marginal spinelets disposed into two rows. Adradi-
al paxillae quadrangular, disposed in regular longitudinal and tranversal rows, bearing
4–10 central, short, rounded spines (Fig. 4g). Inferomarginal plates with two short,
conical and slightly compressed spines (~2.20 mm), the upper one being the smallest.
Towards the mouth the inferomarginal plates are covered by short, lanceolate spines
(Fig. 4f). Between these and the lateral margins slender spinelets occur. Adambulacral
plates with four spines, two elongate, compressed and slightly curved adambulacral
spines, and two elongate, attened and lanceolate subambulacral spines (Fig. 4h). Oral
spines long (~2.59 mm) and thin, forming a dense tuft of spines on the apex of jaw.
Ocular plates well developed and grunuliform. Pedicellariae absent.
Colour. Dorsally bluish-grey or greenish-gray, with a strong dark line along the
central region of the disk and of the arms. e actinal surface is white to cream-coloured.
Distribution. Florida, Cuba, Jamaica, Honduras, Nicaragua, Costa Rica, Pan-
ama, Venezuela and Brazil (Tommasi 1958, Clark 1982, Abreu-Pérez et al. 2005,
Alvarado et al. 2008, del Valle García et al. 2008). In Brazil: PI, CE, PB, PE, AL, BA,

Taxonomic guide and historical review of starshes in northeastern Brazil... 15
RJ, SP, and SC (Rathbun 1879, Verrill 1915, Bernasconi 1943, Krau 1950, Tommasi
1958, 1970, Brito 1962, 1968, Lima-Verde 1969, Fernandes et al. 2002, Magalhães
et al. 2005, Gondim et al. 2008, 2013, Gondim and Giacometti 2010, Xavier 2010,
Miranda et al. 2012). is study provides the rst record for the coast of Rio Grande
do Norte. From 1 to 64 m in depth, being rare below 40 m (Clark and Downey 1992).
Remarks. 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 Mex-
ico and south Brazil is the closest species, but it may be distinguished from L. senegalensis
for having 6 arms, inframarginal plates with two long and narrow spines, and the usual
presence of pedicellariae. Walenkamp (1979) identied one specimen with 6 arms from
Guyana as L. senegalensis. His identication was questioned for some time, but the pres-
ence of dark bands on the carinal region of the arms and disk and the depth of collection
(32m, while the minimum known depth of L. barbadensis is 73m) conrm the validity
of this identication (Clark and Downey 1992). Our observation of both juvenile (dd ≤
15 cm) and adult individuals permitted the conclusion that morphological characters do
not vary during ontogeny. us forms with less than 7 arms or more than 9 arms (this
later condition never having been observed in nature) must be the result of failures dur-
ing metamorphosis (Hotchkiss 2000). As for the remaining Paxillosida, the structure of
the paxillae represent a key taxonomic character for the identication of species.
Ecological notes. e species lives in environments of low hydrodynamism, in
sediment containing sand, mud, or a combination of both of these (Hendler et al.
1995). It feeds mainly on molluscs, but also of other echinoderms such as sea-stars of
the genus Astropecten and irregular echinoids, as well as ophiuroids, copepods, deca-
pods, scaphopods, and polychaetes (Penchaszadeh and Lera 1983). Sometimes a small
commensal crustacean (Minyoceras angustus Dana, 1852) is found in the interior of its
ambulacrum. is crustacean was also observed in L. clathrata (Brito 1960). According
to Hendler et al. (1995), individulas may attain 30–40 cm in disk diameter (dd), and
attain sexual maturity when at 15 cm in diameter. Presently it is considered vulnerable
to extinction along the Brazilian littoral. e main causes of population decline are its
high succeptibility to the eects of pollution of the water column and the frequent ac-
cidental capture in trawling nets (Machado et al. 2008).
Family Astropectinidae Gray, 1840
Astropecten alligator Perrier, 1881
Figure 5a–d
Astropecten alligator Perrier, 1881: 30.
Astropecten nuttingi Verrill, 1915: 181, pl. 12, g. 2e, pl. 21, gs 1–2.
Material examined. Paraíba: 7°01'S; 34°41'05"W, 1 spec., UFPB/ECH.881,
13.II.1981, 24m.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
16
Figure 5. 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.

Taxonomic guide and historical review of starshes in northeastern Brazil... 17
Type locality. Alligator Reef, Florida Keys, Florida (Clark and Downey 1992).
Description. Body attened dorso-ventrally. Disk small, with ve long and nar-
row arms (~4.93 mm) (Fig. 5a, b). Epiproctal cone pronounced (this region of the disk
and the surrounding areas are inated) (Fig. 5a). Abactinal surface at and covered by
small paxillae disposed in regular rows (Fig. 5a). Paxillae with a central spinelet and
8–10 (usually 9) thick, marginal spinelets with blunts extremities. Superomarginal
plates short (~1.14 mm) and narrow (~1.59 mm), covered by short and blunt spine-
lets, giving a granulose appearance to animals. Each of these plates has an elongate,
conical and pointed spine (Fig. 5a, c), which has the same length from the interbra-
chial area to the extremity of each arm. Inferomarginal plates with two long, attened
and pointed spines, placed in parallel to each other. ese plates are also covered by
short and narrow spinelets. Towards the mouth the inferomarginal plates are covered
by many attened and pointed spines, which become elongated at the margins. Adam-
bulacral plates with three long, attened adambulacral spines, placed in parallel to each
other, the median one being slightly longer than the other two (Fig. 5d). ere are
three to four long and attened subambulacral spines, the median of which is longer
and wider than the other two.
Colour. Reddish-brown dorsally, with conspicuous dark bands along the center of
each arm (Clark and Downey 1992), or uniformly orange-red.
Distribution. North Carolina, the Bahamas, Florida, Honduras, Nicaragua Co-
lombia, and Brazil (Verrill 1915, Gray et al. 1968, Clark and Downey 1992, Bena-
vides-Serrato et al. 2005, 2011, Alvarado et al. 2008). In Brazil: Benavides-Serrato et
al. (2005) cite its occurrence in the north of Brazil. is study expands the distribution
southward to the State of Paraíba, being the rst reference for the northeast region
of Brazil. From 22 to 576 m in depth (Benavides-Serrato et al. 2011), most frequent
between 22 and 114 m (Clark and Downey 1992).
Remarks. Astropecten alligator difers from the remaining species in the genus re-
corded for Brazil for presenting a single, elongate, erect conical spine on all the supero-
marginal plates. is character is present in both juvenile and adult individuals (dd =
41.0 mm). For a long time it was believed that Perrier (1881) had described A. alligator
based on a juvenile specimen and the recognition of this species remained uncertain.
Clark and Downey (1992) analised the type material of A. alligator. ey were able to
validate the species and consider Astropecten nuttingi Verrill, 1915 a junior synonym
of this taxon. In a molecular phylogenetic analysis of Astropecten, Zulliger and Lessios
(2010) concluded that A. alligator and A. americanus probably belong to the same spe-
cies and that the species needs to be extensively revised.
Ecological notes. Usually inhabits muddy environments, but may also be found
associated to communities of azooxanthelate corals (Benavides-Serrato et al. 2011). It
is common along the coast of Florida (Clark and Downey 1992), but for a long time
it has been erroneously cited there as Astropecten nuttingi. In other areas A. alligator is
a rare species, being little cited in the literature.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
18
Astropecten articulatus (Say, 1825)
Asterias articulata Say, 1825: 144.
Astropecten dubius Gray, 1840: 182.
Asterias aranciaca Gould, 1841: 349 (non Asterias aranciaca Linnaeus, 1758).
Astropecten articulates Müller & Troschel, 1842: 72. Tommasi 1970: 6, g. 17. Tommasi
et al. 1988: 5. Manso 1989: 357.
Astropecten buschi Müller & Troschel, 1843 (a variety of Astropecten articulatus (Say,
1825) according to Doderlein (1917)).
Astropecten articulatus dubius Verrill, 1915: 165.
Astropecten articulatus var. valenciennesi A.H.Clark, 1939: 442.
Astropecten articulatus duplicatus Zoppi de Roa, 1967: 277, g. 6. (non Astropecten
duplicatus Gray, 1840).
Material examined. Caraguatatuba, São Paulo, 1 spec., MZUSP484, 16.X.2001, 19m.
Type locality. Florida (Clark and Downey 1992).
Description. Body pentagonal, attened dorso-ventrally. Five long and narrow
arms (length of arm corresponds to approximately four times its width), which be-
come 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 plates granulose, with a large spine (~1.54
mm) positioned internally on the interbrachial plates. Some distal supermarginal plates
have a short spine positioned more externally. Inferomarginal plates with two elongate,
attened and pointed marginal spines, positioned horizontally. Small oral spine. ree
adambulacral spines divergent, the median one being the largest.
Colour. Dorsally dark blue or purple in paxillar region. Superomarginal plates
white or orange. Oral surface white of beige (Hendler et al. 1995; Benavides-Serrato
et al. 2011).
Distribution. North Carolina, Florida, the Bahamas, Mexico, Cuba, Puerto Rico,
Dominican Republic, Panama, Colombia, Venezuela, Brazil, and Uruguay (Tommasi
1970, Clark and Downey 1992, Hendler et al. 1995, Alvarado 2011). In Brazil: PI, CE,
RJ, SP, and RS (Lima-Verde 1969, Netto 2006). From 0 to 550 m in depth, being most
common between 5 and 20 m (Clark and Downey 1992, Hendler et al. 1995).
Remarks. Astropecten articulatus diers from the congeneric A. cingulatus for hav-
ing a spine or tubercule on the distal supermarginal plates, two at spines on each
inferomarginal plate and subambulacral spines larger that the adambulacral spines.
e original description of A. articulatus Say (1825) is excelent, but the emphasis given
by this author on the presence of spines and tubercules resulted in some uncertainty
regarding the validity of A. duplicatus (Gray, 1840) (Clark and Downey 1992). Ac-
cording to Clark and Courtman-Stock (1976) and Clark and Downey (1992) the con-
dence in the presence of spines on the superomarginal plates as a specic criterium is
doubtful. As in other species of the genus, A. articulatus presents a series of morpho-
logical variations. Verrill (1915) provides a good discussion of these variations, and

Taxonomic guide and historical review of starshes in northeastern Brazil... 19
stresses the dierences found between juvenile and adult individuals. We have only
been able to examine a single juvenile specimen, which nevertheless presented all the
diagnostic characters of the adults.
Ecological notes. Inhabits sandy environments. According to Benavides-Serrato
et al. (2011) and Hendler et al. (1995), this species is common oshore over conti-
nental shelf, being particularly abundant in North Carolina. Astropecten articulatus
is a voracious and non-selective predator (Hendler et al. 1995). Wells et al. (1961)
recorded 91 food items for 124 specimens collected in North Carolina, gastropods, bi-
valves, and scaphopods being the most importante items. Small crustaceans, juveniles
of Mellita sp. and A. articulatus itself were also recorded as food items for the species,
although in a lower level of importance (Hendler et al. 1995).
Astropecten brasiliensis Müller & Troschel, 1842
Astropecten brasiliensis Müller & Troschel, 1842: 68. Tommasi et al. 1988: 5. Manso
1989: 357. Fernandes et al. 2002: 422. Netto 2006: 25–26, pl. 2a, g. 16a. Ven-
tura et al. 2007: 230. Machado et al. 2008: 350. Lima and Fernandes 2009: 58.
Xavier 2010: 75. Miranda et al. 2012: 143, 144.
Astropecten braziliensis Rathbun, 1879: 150. Tommasi 1970: 6.
Astropecten brasiliensis riensis Döderlein, 1917: 84.
Astropecten brasiliensis armatus Jonh, 1948: 503.
Astropecten armatus brasiliensis Tortonese, 1956: 329. Tommasi 1958: 12–13, pl. 2, g.
3; 1970: 7, g. 19. Brito 1962: 3; 1968: 7–8, pl. 4, g. 3. Lima-Verde 1969: 10.
Carrera-Rodrigues and Tommasi 1977: 81–83. Magalhães et al. 2005: 63.
Astropecten brasiliensis brasiliensis Döderlein, 1917: 83.
Astropecten armatus riensis Tommasi, 1958: 13–14, pr. 2, g. 4. Brito 1962: 3; 1968:
8, pl. 4, g. 2.
Astropecten riensis Tommasi, 1970: 7. Carrera-Rodrigues and Tommasi 1977: 89. Tom-
masi and Aron 1988: 3. Tommasi et al. 1988: 5.
Material examined. Ceará: Fortaleza, Mucuripe, 1 spec., MNRJ285, 1945. Rio Gran-
de do Norte: Areia Branca, Ponta do Mel, 1 spec., UFPB/ECH.1919, 23.VI.182.
Bahia: Salvador, 2 spec., UFBA00132, 01.III.2000.
Type locality. São Sebastião Island, São Paulo, Brazil (Clark and Downey 1992).
Description. Disk small with long, slender, dorsally attened 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. Sup-
eromarginal plates 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 plates with two long and attened spines positioned vertically
and forming a marginal fringe, the most ventral one being larger than the dorsal one.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
20
Adambulacral plates with three ambulacral spines, the inner one being slightly longer
than the other two. Without pedicellariae.
Colour. According to Bernasconi (1957), live animals have an intense violet col-
our with the spines of the marginal fringe yellowish or pinkish. Actinal surface light
salmon colour. When dry specimens may be light pink or whitish.
Distribution. Honduras, Panama, Suriname, Brazil, Uruguay, and Argentina
(Mar del Plata) (Tommasi 1958, 1970, Clark and Downey 1992, Alvarado et al. 2008,
Ventura et al. 2007). In Brazil: CE, PE, AL, BA, RJ, SP, SC, and RS, incluing the
islands of Fernando de Noronha and Trindade (Rathbun 1879, Verrill 1915, Tom-
masi 1958, 1970, Walenkamp 1976, Brito 1962, Lima-Verde 1969, Fernandes et al.
2002, Miranda et al. 2012). In this paper we establish the rst record for Rio Grande
do Norte. From 7 to 45 m in depth (Ventura et al. 2007).
Remarks. is species diers 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 plates. e vast
synonymy presented by Astropecten brasiliensis evidences the plasticity of some of its
characters, such as the number of spines on the superomarginal plates. is character,
together with the shape of the marginal spines and paxillae spinelets formed the basis
for the establishment of the ve known subspecies. Döderlein (1917) considered A.
armatus Gray, 1840 and A. erinaceus Gray, 1840 subspecies of A. brasiliensis on the
basis of dierences in the adambulacral and superomarginal spines. Boone (1933),
Clark (1940) and John (1948) disagreed with Döderlein (1917) and considered the
three species not to be dierent, stating that the characters proposed to diagnose the
species were not signicant. Walenkamp (1976) listed a series of variations observed in
specimens from Surinam and established A. brasiliensis and A. riensis as distinct species.
Bernasconi (1957), Brito (1968) and Tommasi (1958, 1970) adopted the subspecies
proposed by Müller and Troschel (1842) and Döderlein (1917) for Brazilian material.
Clark and Downey (1992), analysing the neotype of the species, concluded that A.
brasiliensis and A. armatus are distinct, while A. riensis is a synonym of the rst. Fur-
thermore, he considered all the described subspecies to be synonyms. We agree with
the proposal of Clark and Downey (1992) and include the subspecies (A. brasiliensis
riensis, A. brasiliensis armatus, A. brasiliensis brasiliensis and A. armatus riensis) and A.
riensis as synonyms of A. brasiliensis. e specimen examined in this study was broken,
but its taxonomic characters were observable, except for the loss of the superomarginal
spines. Scars of these spines remained on the plates, however, and we were able to es-
tablish that two spines occurred per plate, characterizing an adult individual.
Ecological notes. 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 bi-
valves, gastropods, crustaceans, echinoderms, and polychaetes (Ventura et al. 2007).
As all species inhabiting soft sediments, Astropecten brasiliensis is suering the impact
of excessive collecting, being captured in bottom trawling nets and frequently do not
resist the damage inicted by these shing eorts (Machado et al. 2008). Presently the
species is considered to be vulnerable to extinction along the Brazilian coast.

Taxonomic guide and historical review of starshes in northeastern Brazil... 21
Astropecten cingulatus Sladen, 1833
Figure 5e–h
Astropecten cingulatus Sladen, 1883: 266. Brito 1962: 3; 1968: 9, pl. 4, g. 4. Tommasi
1970: 5, g. 16; 1985: 3. Carrera-Rodrigues and Tommasi 1977: 84–86. Tommasi
and Aron 1987: 3. Manso 1989: 357. Tommasi et al. 1988: 5. Ventura et al. 2007:
231. Xavier 2010: 75.
Astropecten mesactus Studer, 1884: 46.
Astropecten jarli Madsen, 1950: 181.
Material examined. Rio de Janeiro: Cabo Frio, 1 spec., MNRJ1853, 18.VI.1997.
Type locality. Pernambuco, Brazil (Clark and Downey 1992).
Description. Body pentagonal, attened dorso-ventrally. Five long (~37.70 mm)
and narrow (~9.56 mm) arms (length of arm corresponds to approximately four times
its width) (Fig. 5e). Madreporite oval (~1.52 mm) and marginal. Epiproctal cone pro-
nounced. Paxillae small and granulose, with 1–4 central spinelets and 9–12 marginal
spinelets. Paxillar spinelets granulose (Fig. 5e). Superomarginal plates granulose (Fig.
5g). Two fringes of marginal spines aligned horizontally. First row of spines of infero-
marginal plates with three marginal spines, disposed in parallel to each other, two being
of the same length and one shorter. Second row with four elongate and slightly attened
spines, three of which are subequal in length and one much shorter. Oral spines short,
forming a bundle at the apex of the jaw (Fig. 5f). Adambulacral plates with 3–4 diver-
gent spines, the median one being the largest (Fig. 5h). Subambulacral spines forming a
bundle of elongate and slightly attened (lanceolate) spines. Pedicellariae rarely present.
Colour. According to Benavides-Serrato et al. (2011) and Bernasconi (1957),
the species has the abactinal surface red or orange-red and the actinal surface white.
Ventura et al. (2007) recorded a cream-colour on the dorsal surface and white on the
oral surface in Brazilian specimens.
Distribution. North Carolina, the Bahamas, Gulf of Mexico, Mexico, Nicaragua,
Costa Rica, Panama, Colombia, Brazil, Uruguay, Argentina, and Africa (Tommasi
1970, Carrera-Rodríguez and Tommasi 1977, Clark and Downey 1992, Ventura et
al. 2007, Alvarado et al. 2008). In Brazil: PE, RJ, SP, and SC, including the submarine
banks and mountain ranges Vitória-Trindade and Vitória Island (SP) (Brito 1962,
Tommasi 1970, 1985, Tommasi and Aron 1987, Manso 1989, Xavier 2010). Inter-
tidal to 1350 m in depth (Clark and Downey 1992), being most frequent between 51
and 129 m (Carrera-Rodríguez and Tommasi 1977).
Remarks. Astropecten cingulatus diers from its closest species, A. articulatus, for
having three rounded spines on each inferomarginal plate, and for having subambu-
lacral spines that are smaller than the adambulacral spines and rounded. Bernasconi
(1957) stresses a few morphological variations observed in specimens from Uruguay
and Argentina. Among these are the aspect of the superomarginal plates, which do not
have large spines, or then there are only a few small, granuliform spines on the rst
plates. According to Ventura et al. (2007) the specimens collected along the Brazilian

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
22
coast have short arms, in contrast to the indication in the original diagnosis. e single
individual we examined, on the other hand, corresponds to that indicated in the litera-
ture (lenght about four time its width).
Ecological notes. is species lives on sandy or muddy bottoms of the littoral re-
gion up to depths of 50 m (Tommasi 1970, Machado et al. 2008). It feeds mainly on
gastropods, bivalves, crustaceans, and cirripeds (Ventura et al. 2007). Astropecten cin-
gulatus, and other species of the genus, are frequently captured in trawling nets and the
species is presently included among those vulnerable to extinction in Brazil (Machado
et al. 2008). According to Brito (1962), this species is abundant along the coast of
Pernambuco, but no other works conducted in this state conrm this observation.
Astropecten marginatus Gray, 1840
Figures 6a–e, 12b
Astropecten marginatus Gray, 1840: 181. Tommasi 1958: 14, pl. 2, g. 5; 1970: 5, g. 15.
Brito 1962: 3; 1968: 7, pl. 4, g. 1. Lima-Verde 1969: 11. Carrera-Rodrigues and
Tommasi 1977: 88–89. Tommasi et al. 1988: 5. Nomura and Fausto Filho 1966:
19. Gondim et al. 2008: 155. Lima and Fernandes 2009: 58. Xavier 2010: 75.
Astropecten ciliatus Grube, 1857: 340.
Astropecten richardi Gary, 1840: 181.
Astropecten orans Sluiter, 1895: 54.
Material examined. 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.
Type locality. Unknown (Clark and Downey 1992).
Description. Body attened. Five broad, triangular arms (Fig. 6a, b). Abactinal
surface covered by paxillae arranged in regular transversal rows. Circular madrepo-
rite positioned marginally in one of the abactinal intermediate areas. Carinal paxillae
slightly smaller than the adradials (Fig. 6d). Paxillae small with 6–8 central spinel-
ets and 12 marginal spinelets, all short and blunts. Superomarginal plates granulose,
broader (~3.89 mm) than long (~2.00 mm) (Fig. 6c, d). Inferomarginal plates granu-
lose, with two thick, blunt, parallel spines of similar length (~3.92 mm) and one small
spine positioned laterally in relation to the other two. ese spines form a well dened
marginal bundle. Actinal surface with inferomarginal plates partially naked, having
two series of marginal spines positioned laterally and one other marginal series with
four thin, elongate, and attened spines (~1.66 mm), located behind the aboral mar-
ginal row of spines (Fig. 6e). Adambulacral plates with a series of small spines on the
proximal face and three long, narrow, and attened adambulacral spines, the median

Taxonomic guide and historical review of starshes in northeastern Brazil... 23
Figure 6. Some species of the family Astropectinidae recorded in northeastern Brazil. Astropecten marginatus
(A–E) A Abactinal view B Actinal view C Abactinal view of the arm D Detail of the superomarginal plate,
and E Actinal view of the arm.
of which is longest and widest. Six elongate, narrow, and pointed oral spines (~1.67
mm). Ocular plate small and bilobed.
Colour. Dorsally either blue with white margins (Clark and Downey 1992) or uni-
formly orange-coloured (Benavides-Serrato et al. 2011). Most specimens collected in
northeastern Brazil vary from cream-coloured to grayish. Dried specimens become white.
Distribution. Costa Rica, Puerto Rico, Colombia, Venezuela, Guyana, and Brazil
(Tommasi 1958, Clark and Downey 1992, Hendler et al. 1995, Alvarado et al. 2008,
Benavides-Serrato et al. 2005, 2011). In Brazil: CE, PB, PE, RJ, SP, SC, and RS
(Bernasconi 1955, Brito 1962, Lima-Verde 1969, Netto 2006, Gondim et al. 2008,
Lima and Fernandes 2009, Xavier 2010). is is the rst record for the littoral of Rio
Grande do Norte. From 1 to 130 m in depth (Clark and Downey 1992).
Remarks. Astropecten marginatus diers 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 plate. Unlike other species of Astropecten, A. marginatus

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
24
shows little morphological variation in characters considered of taxonomic interest
(Walenkamp 1976). We observed both juveniles (R = 11.73 mm) and adults (R =
81.41 mm).
Ecological notes. is species lives in substrates containing sand or sand with
mud (Ortega et al. 2010). As most species of the genus, it is an active predator with
a generalist food diet (Ortega et al. 2010). is is one of the most common species
in northeastern Brazil, being abundant below 2–4 m (Benavides-Serrato et al. 2011).
Together with other species in the genus, populations of A. marginatus suer great col-
lecting pressures from bottom trawling shing nets. Presently it is included among the
Brazilian species vulnerable to extinction (Machado et al. 2008).
Order Valvatida Perrier, 1884
Family Asterinidae Gray, 1840
Asterinides folium (Lütken, 1860)
Figure 7a–e
Asterina minuta Gray, 1840: 289.
Asteriscus folium Lütken, 1860: 60–61.
Asterina folium A. Agassiz, 1877: 106, pl. 14, gs 7–9.
Asterinides folium Verrill, 1913: 479; Brito 1962: 3; 1968: 17, pl. 7, gs 2–3; 1971:
262. Tommasi 1970: 15, g. 38. Oliveira et al. 2010: 3, g. 2a.
Material examined. 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.
Type locality. Saint omas, Virgin Islands (O’Loughlin 2002).
Description. Body inated, pentagonal (Fig. 7a, b). Five short arms (some speci-
mens may have 4–6 arms). Abactinal plates imbricated, decreasing in size towards
the margin of the body, with 2–4 short, hyaline, divergent spines, which have the
extremity trifurcate (Fig. 7c). Between each of these plates there is a papula (Fig. 7c).
Anus located in the center of the abactinal surface. Superomarginal plates similar to
the remaining abactinal plates, but with more numerous and slightly longer spines,
forming a dense bundle (Fig. 7a). Papulae restricted to the abactinal surface. Actinal
plates similar to the abactinal plates, but a little bigger and longer, having 1–3 diver-
gent spines, also similar to the dorsal ones, but slightly longer (Fig. 7b). Inferomarginal
plates similar to the remaining actinal plates. Adambulacral plates with three thin,
vitreous, elongate spines (~0.035 mm), having the tips trifurcate (Fig. 7d). Six thin,
vitreous, elongate oral spines (~0.43 mm) (Fig. 7e).

Taxonomic guide and historical review of starshes in northeastern Brazil... 25
Figure 7. Some species of the order Valvatida recorded in northeastern Brazil. Asterinides folium (A–E)
A Abactinal view, in detail the superomarginal plates 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 clavi-
gera (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.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
26
Colour. Juvenile specimens vary from white to cream-coloured, larger juveniles
are yellow to reddish, while adults are blue or greenish-blue (Hendler et al. 1995). Bri-
to (1968) recorded a dark grayish-blue for 25 specimens from Trindade Island (ES).
Distribution. Bermudas, Florida, Bahamas, Belize, Panama, and Brazil (Verrill
1915, Clark and Downey 1992, Hendler et al. 1995, Alvarado et al. 2008, Benavides-
Serrato et al. 2011). In Brazil: BA, RJ, and Trindade Island (Brito 1962, 1968, 1971,
Oliveira et al. 2010). In the presente study we record for the rst time its presence in
the State of Paraíba. Intertidal to 15 m in depth (Hendler et al. 1995).
Remarks. Only two species of the genus Asterinides are known from the Atlantic
Ocean, A. folium and A. hartmeyeri (Döderlein, 1910). e rst is recorded for Ber-
mudas and southern Brazil and the second only for the Caribbean region. According
to Clark and Downey (1992) these species are partially sympatric, and for this reason
have previously easily been confused with each other. In a revision of family Aster-
inidae based on molecular and morphological data, O’Loughlin and Waters (2004)
transferred A. folium and A. hartmeyeri from the genus Asterina to the genus Asterin-
ides, remarking on the morphological similarities between these two species. Asterinides
folium diers from A. hartmeyeri for having bigger papular pores, 6 to 7 series of actinal
plates and abactinal plates arranged into two rows. A broad discussion of the main dif-
ferences between these two species is given by Clark and Downey (1992). According
to Hendler et al. (1995), this species rarely reaches 2.5 cm in diameter. Specimens
examined in this study had a larger ray (R, maximum of 7.76 mm) and diered from
those described by Hendler (op cit.) for not having narrow radial areas inated, having
instead the abactinal surface completely inated.
Ecological notes. is species lives in association with coral reefs, being found par-
ticularly under rocks or corals of the reef at (Hendler et al. 1995). e specimen from
Paraíba recorded in this study was found associated with rhodolite beds at 10 m depth.
Although the species is reported from several localities, is has never been found in large
numbers (Hendler et al. 1995, Benavides-Serrato et al. 2011). Only Brito (1971) ob-
served this species to be relatively abundant mainly under rocks at Trindade Island.
Family Mithrodiidae Viguier, 1878
Mithrodia clavigera (Lamarck, 1816)
Figure 7f–i
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.

Taxonomic guide and historical review of starshes in northeastern Brazil... 27
Mithrodia clavigera Perrier, 1875: 378.
Mithrodia victoriae Bell, 1882: 123, pl. 6, g. 2. Brito 1962: 3; 1968: 16. Tommasi
1970: 19, g. 55.
Material examined. Paraíba: 7°04'S; 34°41'W, 1 spec., UFPB.ECH.880, 17.II.1981, 26m.
Type locality. Unknown (Clark and Downey 1992).
Description. Disk small (Fig. 7f). Five cylindrical and narrow arms (~2.52 mm)
(Fig. 7f, g). Abactinal and actinal surfaces granulose (Fig. 7h). Skeleton formed by
polygonal primary plates (usually hexagonal) that are widely spaced and united by
secondary plates of rectangular shape, forming a reticulum. Some carinal and adradial
plates with a long, narrow, and blunt spine (~0.77 mm). Papula large and single, found
between the abactinal plates. Papulae restricted to abactinal surface. Granules covering
body and spines small and with spinous tip. Granules from base of spines larger than
at other localities. One conical and elongate subambulacral spine (~0.32 mm), form-
ing a well dened row at base of ambulacral groove (Fig. 7i). Four slightly attened
adambulacral spines, the median ones being the largest. Eight short and rectangular
oral spines, of which the median ones are largest (Fig. 7g).
Colour. Arms banded with dark brown or red lines, unusually green (Clark and
Downey 1992). Specimens preserved in alcohol become uniformly white.
Distribution. Mexico, Cuba, Nicaragua, Brazil, Indo-Pacic (except Hawaii)
(Hayashi 1940, Abreu-Pérez et al. 2005, Alvarado et al. 2008). In Brazil: ES (Vitória
Banks) (Brito 1968, Clark and Downey 1992). In this study we provide the rst record
for northeastern Brazil. From 24 to 71m in depth (Clark and Downey 1992).
Remarks. Mithrodia clavigera is the only species of the family Mithrodiidae re-
corded for the Western Atlantic. For some time, two species were considered present:
M. clavigera and M. victoriae. e later species was described by Bell (1882), based
on two small specimens from submerged banks of Vitória (Victoria Bank) (Espírito
Santo, Brazil). Since its description the validity of M. victoriae was questioned, and the
distinction between these two species was discussed by Engel et al. (1948) and Pope
and Rowe (1977). ese authors concluded that the species should be synonymized
but, due to the lack of material, this action was not formally carried out. Clark and
Downey (1992) analised a large number of individuals of several sizes and agreed with
the observations of Engel et al. (op. cit.) and Pope and Rowe (op. cit.), considering
the two species to be synonyms. e individual we analised is juvenile (R = 9.80 mm)
and corresponds to the characterization of juvenile specimens by Engel et al. (1948).
ese authors provide details on the aspect of the pedicellariae of Mithrodia clavigera,
but pedicellariae were not found in our specimen.
Ecological notes. is species lives on hard substrates covered by incrusting or-
ganisms and in reef gravel (Abreu-Pérez et al. 2005). For Paraíba it was recorded as-
sociated with rhodolith banks. According to Guille et al. (1986) this species is more
active during the night.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
28
Family Oreasteridae Fisher, 1911
Oreaster reticulatus (Linnaeus, 1758)
Figure 8a–g, 12c
Asterias gigas Linnaeus, 1753: 114.
Asterias reticulata Linnaeus, 1758: 661.
Pentaceros reticulatus Gray, 1840: 276.
Oreaster reticulatus Linnaeus, 1758. Tommasi 1958: 16–17, pl. 3, g. 2; 1970: 10–11, g.
31. Brito 1962: 3; 1968: 5–6, pl. 2, gs 1–3. Lima-Verde 1969: 11. Fernandes et al.
2002: 422. Magalhães et al. 2005: 63. Ventura et al. 2007: 238. Manso et al. 2008: 185,
g. 8c, d, e. Xavier 2010: 75. Alves et al. 2010: 757. Miranda et al. 2012: 143, 144.
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, gs 1–2.
Material examined. Ceará: o Fortaleza, 1spec., UFPB/ECH.1255, Geomar XXIV,
V.1985. Paraíba: 1spec., UFPB/ECH.1579, 26.X.1980; Cabedelo, Farol de Cabede-
lo 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.
Type locality. ‘Las Antillas Occidentales’ or ‘Spanish West Indies’ (Clark and
Downey 1992).
Description. Body pentagonal. Disk high, inated, concave on actinal side (Fig. 8a,
b). Five short arms (Fig. 8a). Abactinal plates with one thick, short, blunt spine (~2.43
mm). Among these plates there occur papular areas, which are covered by small granules
and bivalve pedicellariae (Fig. 8c). ese granules cover the entire body, including the
bases of the spines. Superomarginal plates with one short, thick and blunt spine (~3.38
mm), determining the margin of the body. Inferomarginal plates similar to superomar-
ginal plates, with one or two small and thick spines. Papular areas restricted to abactinal

Taxonomic guide and historical review of starshes in northeastern Brazil... 29
Figure 8. 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.
surface. Actinal surface granulose, with a great number of pedicellariae, especially in
areas near the mouth and abulacral groove. Actinal plates with 1–2 short, conical, and
blunt spines (~1.53 mm). Ambulacral plates with 5–6 short and attened spines, of
which the median spines are the largest (Fig. 8e). Short, conical, spines (~3.16 mm)
form a well dened row of spines on the margins of the ambulacral grooves. Four short,
thick, blunt oral spines (Fig. 8d). Sessile bivalve pedicellariae distributed over entire
body of animal (Fig. 8f). Skeleton formed by conical, abactinal plates interconnected by
secondarily elongated and widened plates, arranged into a reticulum (Fig. 8g).
Colour. According to Hendler et al. (1995) and Verrill (1915) the colour pattern
of this species is very variable, even among individuals from a same population. Along
the Brazilian coast the most common colour is orange or brownish red. Yet juvenile in-
dividuals dier signicantly from adults. According to Benavides-Serrato et al. (2011)
and Hendler et al. (1995) the aboral surface of juveniles is frequently olive-green and
usually presents green-grey or coe-coloured spots. In the adults, on the other hand,
this colour is orange with lighter or darker tubercles on the disk and arms. e oral
surface in both stages is beige or cream-coloured.
Distribution. North and South Carolina, the Bermudas, the Bahamas, Belize, Gua-
temala, Honduras, Cuba, Nicaragua, Costa Rica, Venezuela, Brazil, and Cabo Verde
(Tommasi 1970, Walenkamp 1976, Hendler et al. 1995, Ventura et al. 2007, Alvarado
et al. 2008, del Valle García et al. 2008). In Brazil: MA, CE, PB, PE, AL, BA, RJ, SP,
and SC, incluinding Abrolhos and Trindade Island (Rathbun 1879, Verrill 1915, Clark
1942, Tommasi 1958, 1970, Brito 1962, 1968, Lima-Verde 1969, Fernandes et al.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
30
2002, Magalhães et al. 2005, Xavier 2010, Miranda et al. 2012). In this study we record
the species for the rst time in the States of Rio Grande do Norte, and Alagoas. From 0
to 800 m in depth, being most abundant up to 50m.
Remarks. Only two species of the genus Oreaster are known for the Atlantic Ocean,
Oreaster clavatus and O. reticulatus. e rst is known only from the Island of Cape
Verde, São omé and the Gulf of Guinea. e second, occurs widely throughout the
West Atlantic, from North Carolina to the south of Brazil (Clark and Downey 1992),
its known southern limit being located in the State of Santa Catarina. Oreaster reticula-
tus diers from its congeneric O. clavatus for presenting an inated body and abactinal
plates with tubercules or spines. Clark (1942) described the variety O. reticulatus var.
bermudensis on the basis of the irregular placement of spines and papulae on the abacti-
nal surface and of the presence of only one spine on the actinal plates. However, these
characters also occur in other specimens from the Atlantic and thus do not sustain the
name. According to Hendler et al. (1995) the species may attain a disk diameter of up to
500 mm. We observed morphological variations in both juveniles and adults, but were
not able to correlate these with colour patterns in this preserved material.
Ecological notes. e species lives in shallow reef environments with calm water,
coastal lagoons, seagrass beds (alassia, Halodule and Syringodium), and mangrove
channels (Benavides-Serrato et al. 2011). In this study the species was recorded in
rhodolite beds and coastal reefs below 6 m. According to Verrill (1915) Oreaster reticu-
latus was the most abundant species in the States of Bahia and Pernambuco. Presently
it is dicult to nd along the northeastern coast of Brazil, and is listed as vulnerable
to extinction (Machado et al. 2008). is is an omnivorous species, feeding mainly
on microorganisms from organic matter associated with the sand of seagrass beds and
algal substrates. However, it is also an opportunistic predator of echinoderms, such
as Tripneustes ventricosus (Lamarck, 1816) and Meoma ventricosa ventricosa (Lamark,
1816), as well as of individuals belonging to its own species and to a large variety of
sponges (Hendler et al. 1995). e only known predator of adults belonging to this
species is the gastropod Charonia variegata (Lamarck, 1816), while young individuals
are known to be eaten by a great variety of shes (Scheibling 1980).
Order Velatida Perrier, 1884
Family Pterasteridae Perrier, 1875
Calyptraster coa Sladen, 1882
Figure 9a–h
Calyptraster coa Sladen, 1882: 207. Tommasi 1970: 13.
Calyptraster personatus Madsen, 1947: 3–7, gs 1–2.
Material examined. MZUSP (without registration number), 1spec., W Besnardi, dredging
5142. MZUSP (without registration number), 2spec., W Besnardi, dredging 5363.

Taxonomic guide and historical review of starshes in northeastern Brazil... 31
Figure 9. 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 pedi-
callariae (optical microscopic image), and M Lateral view of the arms, in detail the bivalve pedicellariae
(optical microscopic image).

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
32
Type locality. Recife, Pernambuco, Brazil (Clark and Downey 1992).
Description. Body pentagonal (Fig. 9a, b, g, h). Five short arms. Supradorsal
membrane thin and transparent (Fig. 9a). Spiracules moderately large, numerous, ir-
regularly distributed. Oscule large, surrounded by long oscular valves with an enlarged
extremity (Fig. 9g). Abactinal surface with paxillae. Paxillae with long peduncles and a
crown of 5 to 6 long and vitreous spinelets (Fig. 9c). Skeletal plates narrow, long and
vitreous, forming a reticulum. Actinal surface slightly concave. Oral spines long and
vitreous, the lateral ones being longer and thicker (Fig. 9e). Adambulacral plates with
three or four spines (Fig. 9d).
Colour. From light brown to hey-coloured (Sladen 1889).
Distribution. e Bahamas, Florida, and Brazil (Clark and Downey 1992). In
Brazil: PE (Tommasi 1970; Clark and Downey 1992). From 260 to 933 m in depth
(Clark and Downey 1992).
Remarks. e genus Calyptraster presently contains ve species (Mah 2013),
four of which occur in the Western Atlantic: C. coa, C. personatus (Perrier, 1885),
C. tenuissimus Bernasconi, 1966 and C. vitreus Bernasconi, 1972. e rst two
have a similar geographical distribution, occurring from the Bahamas to Brazil and
Colombia, respectively, while the last two are restricted to the coast of Argentina.
According to Clark and Downey (1992), this genus is problematic, and the distinc-
tions between C. coa and C. personatus remain to be better established. Walenkamp
(1979) provides good descriptions and discusses the main morphological diferences
between these species. In the phylogenetic analysis of the family Pterasteridae by
Villier et al. (2004), it is concluded that Calyptraster representes a monophyletic
genus. Calyptraster coa diers from C. personatus for having conical suboral spines,
an osculum surrounded by valves with enlarged extremities, and for being distrib-
uted no deeper than 1000 m. Specimens observed in this study were all juveniles.
Smaller specimens tend to have longer arms and a more attened body. Our mate-
rial was badly preserved in general, not permitting a detailed description of their
morphological characters.
Ecological notes. is species is quite rare, from deep waters, with little known
of its biology and ecology. e species was collected in bottoms containing red mud
(Sladen 1889).
Family Ophidiasteridae Verril, 1870
Linckia guildingi Gray, 1840
Figure 10a–e, 12d
Linckia guildingii Gray, 1840: 285. Tommasi 1958: 17. Brito 1962: 3; 1968: 4–5, pl.
1, g. 3; 1971: 262. Lima-Verde 1969: 11. Tommasi and Aron 1988: 3. Tommasi
et al. 1988: 6. Fernandes et al. 2002: 422. Gondim et al. 2008: 154.
Linckia pacica Gray, 1840: 285.

Taxonomic guide and historical review of starshes in northeastern Brazil... 33
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. Miranda et al. 2012: 144.
Material examined. 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, Pa-
ripueira 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 Ver-
melha, 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.
Type-locality. Saint Vincent, West Indies (Clark and Downey 1992).
Description. Disk small. From four to six long, thin, cylindrical arms (Fig. 10a,
b). Abactinal and actinal surfaces granuliform. Two or more madreporites. Abactinal
plates rounded, inated, irregularly arranged. Among these plates there are papular
areas with 5–23 pores (Fig. 10c). Papular areas restricted to abactinal surface. Supero-
marginal and inferomarginal plates similar and uniform in size, separated by a row of
papular areas. Actinal plates forming 2–3 series, which extend to the tip of the arms
and are covered by granules a little larger than those on abactinal surface. Adambu-
lacral plates with two short, blunt, parallel spines, one being much larger than the
other (Fig. 10d). Behind these there is one wide, thick, blunt, ambulacral spine. Oral
spines slightly longer (~0.68 mm) than remaining spines, having their tips rounded
(Fig.10e). Ocular plates also granulose.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
34
Figure 10. 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.

Taxonomic guide and historical review of starshes in northeastern Brazil... 35
Colour. Extremely variable, usually juveniles and adults having dierent colours
(Hendler et al. 1995). Juvenile individuals have brown, red or violet spots, while adults
are uniformly reddish-brown, yellowish-brown, violet, or olive-green (Brito 1960,
Hendler et al. 1995, Benavides-Serrato et al. 2011). In the Brazilian material the most
common colour observed is brown and yellow.
Distribution. Tropicopolitan (Tommasi 1970, Alvarado et al. 2008). It also oc-
curs throghout the tropical Indo-West Pacic. In Brazil: PB, PE, AL, BA, RJ, and SP,
including Abrolhos and Trindade Island (Rathbun 1879, Verrill 1915, Bernasconi
1955, Tommasi 1958, Brito 1960, 1962, 1968, 1971, Lima-Verde 1969, Tommasi
and Aron 1988, Fernandes et al. 2002, Magalhães et al. 2005, Gondim et al. 2008,
Miranda et al. 2012). From 0 to 298 m in depth (Clark and Downey 1992).
Remarks. 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 south-
eastern Brazil. Tommasi and Aron (1988) cite Ophidiaster guildingi Gray, 1840 for
southeast Bahia, a locality we were not able to conrm. Linckia guildingi diers from
L. nodosa for having small triangular abactinal plates, 18–30 pores per papular area,
and two subambulacral spines. Juvenile individuals of L. guildingi may be confused
with Ophidiaster guildingi Gray, 1840, which have the same colour and occupy the
same habitat (Hendler et al. 1995). Ophidiaster guildingi diers from L. guildingi for
having less than 15 pores per papular area. Although we observed both juvenile and
adult individuals, no morphological variations were noted.
Ecological notes. is species lives in environments with consolidated sub-
strates or sand banks among reefs (Machado et al. 2008). It has cryptic habits,
being found mostly under rocks. Possibly L. guildingi uses the lm of microorgan-
isms adhered to the substrate as food (Hendler et al. 1995; Machado et al. 2008).
According to Brito (1971) this species is abundant along the northeastern coast of
Brazil. However, its populations are becoming reduced, especially in southeastern
Brazil. e species is presently included among the species vulnerable to extinction
(Machado et al. 2008). L. guildingi is known for its strong propensity to autotomize
and its capacity to regenerate. Specimens with four, six or seven arms are common
(Tommasi 1958).
Narcissia trigonaria Sladen, 1889
Figure 10f–j
Narcissia trigonaria Sladen, 1889: 414, pl. 65, gs 5–8. Tommasi 1966: 244; 1970: 9,
g. 26. Brito 1960: 5, pl. 1, gs 4–5; 1962: 3; 1968: 5. Tommasi and Aron 1988: 3.
Tommasi et al. 1988: 6. Miranda et al. 2012: 144.
Narcissia trigonaria var. helenae Mortensen, 1933: 429.
Material examined. Alagoas: Lagoa Azeda, Jequidá da Praia, 1spec., MNRJ (no reg-
istration number), 22.VI.2002. Bahia: Salvador, north coast, 1spec., UFBA00570,

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
36
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.
Type-locality. Bahia, Brazil (Clark and Downey 1992).
Description. Disk high and pyramidal (Fig. 10f, h). Five long and triangular
arms in transversal section (Fig. 10f, h). Abactinal and actinal surfaces granulose
(Fig. 10i). Abactinal plates rounded, placed in irregular rows, covered by attened
and polygonal granules. Among these plates are papular areas with up to three papu-
lae. Superomarginal plates short, wide (~2.69 mm), granulose and visible only later-
ally. Papular areas restricted to abactinal surface. Inferomarginal and superomarginal
plates similar. Actinal plates slightly rectangular and granulose, these granules being
bigger and taller than the dorsal granules. Actino-lateral plates with two rows of
spines, the outer series with 4–5 short and blunt spines. e inner row is formed by
3–4 large, attened spines, being longer than the outer row. Adambulacral plates
with 3–4 series of attened and prismatic spines, of which the innermost are the
largest (Fig. 10j).
Colour. Live specimens are cream-coloured with red spots (Benavides-Serrato
et al. 2011).
Distribution. North Carolina, Florida, Panama, Colombia, and Brazil (Tommasi
1970; Clark and Downey 1992; Alvarado et al. 2008; Benavides-Serrato et al. 2011).
In Brazil: AL, BA, and RJ (Verrill 1915, Brito 1960, 1962, Tommasi 1970, Tommasi
and Aron 1988, Miranda et al. 2012). From 5 to 91 m in depth (Tommasi 1970,
Clark and Downey 1992).
Remarks. Narcissia trigonaria is a well established species, with a small list of syno-
nyms and little morphological variation. It diers from N. canariensis (d’Orbigny,
1839) for having subambulacral spines arranged into three series and paired papulae.
Downey (1973) records the sugar-tongs type of pedicellariae among the carinal plates
of N. trigonaria. However, we did not observe this type of pedicellaria. Walenkamp
(1976) gives an excellent discussion on the presence or absence of pedicellariae and on
small morphological variations found in his material. He emphasizes the great mor-
phological dierences existing between juvenile and adult specimens. In general, very
juvenile individuals have shorter and wider arms. ese tend to become thinner and
longer during ontogenetic development.
Ecological notes. e species lives in consolidated substrates, either rocks or
coral (Machado et al. 2008). Presently it is considered vulnerable to extinction along
the Brazilian littoral. e main causes of its populational decline are the eects of pol-
lutants and its illegal and indiscriminate collecting for aquarium rearing (Machado
et al. 2008).

Taxonomic guide and historical review of starshes in northeastern Brazil... 37
Order Spinulosida Perrier, 1884
Family Echinasteridae Verril, 1867
Echinaster (Othilia) brasiliensis Müller & Troschel, 1842
Figure 11a–e
Echinaster brasiliensis Müller & Troschel, 1842: 22. Tommasi 1958: 22–23, pl. 4, g.
3; 1970: 17, gs 44–45. Brito 1962: 3; 1968: 13–14, pl. 6, g. 6. Carrera-Rodri-
guez and Tommasi 1977: 103–104. Tommasi and Aron 1987: 3. Tommasi et al.
1988: 6. Ávila-Pires 1983: 440–442, gs 8–9. Fernandes et al. 2002: 422. Netto
2006: 30–32, pl. 5a. Alves et al. 2010: 758. Miranda et al. 2012: 144.
Echinaster braziliensis Verrill, 1915: 41–42, pl. 26, g. 1.
Echinaster antonioensis Bernasconi, 1955: 72–73, pl. 6, gs 1–2. Tommasi 1958: 22,
pl. 4, g. 2. Brito 1968: 15.
Echinaster sentus Bernasconi, 1956: 136–137, pl. 4, g. 3. Tommasi 1958: 23–24, pl. 4,
g. 4; 1970: 17–18, g. 46 a 48. Brito 1968: 14, pl. 6, gs 3–4.
Echinaster spinulosus Bernasconi, 1956: 138–139, pl. 4, g.4. Tommasi 1958: 21–22,
pl. 4, g. 1. Brito 1968: 14, pl. 6, g. 1–2.
Echinaster densispinulosus Tommasi, 1970: 18–19, gs 49–51.
Echinaster nudus Tommasi, 1970: 18–19, gs 52–54. Gondim et al. 2008: 154.
Echinaster (Othilia) brasiliensis Clark & Downey, 1992: 21–22, pl. 4a. Hopkins et al.
2003: 98–100. Machado et al. 2008: 182–183. Lima and Fernandes 2009: 59.
Xavier 2010: 75.
Material examined. 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; Cabe-
delo, 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 Be-
ach, 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.
Type-locality. Ubatuta, São Paulo, Brazil (Walenkamp 1976).
Description. Disk small (Fig. 11a). Body concave on abactinal surface, plane
on actinal surface. Five long-narrow to short-thick arms (usually decreasing rapidly
in thickness towards tips) (Fig. 11a, b). Abactinal plates attened or slightly mam-
miform (Fig. 11e), forming a reticulum and bearing a short, conical spine (Fig. 11c).
Abactinal plates separated by small, elongated, secondary plates. Anus normally
placed within the central pentagon of the disk and surrounded by 4–6 spines. Ma-
dreporite circular, with numerous small granules and irregular sulci. Superomarginal
plates more granulose than inferomarginal plates. Papular areas more numerous on
dorsal and lateral regions of arms (Fig. 11c). Adambulacral plates with three spines,

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
38
Figure 11. 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.

Taxonomic guide and historical review of starshes in northeastern Brazil... 39
the inner one being rudimentary (Fig. 11d). e two outer spines are subequal and
larger than the remaining spine.
Colour. Quite variable, being light brown, reddish-brown, dark red or even yel-
lowish red (Gray et al. 1968; Benavides-Serrato et al. 2011).
Distribution. Florida, Cuba, Honduras, Panama, Colombia, Brazil, and Argentina
(Tommasi 1958, Alvarado et al. 2008, Benavides-Serrato et al. 2011). In Brazil: PB,
PE, AL, BA, ES, SP, RJ, SC, and RS (Verrill 1915, Tommasi 1958, Carrera-Rodriguez
and Tommasi 1977, Fernandes et al. 2002, Magalhães et al. 2005, Gondim et al. 2008,
Xavier 2010, Miranda et al. 2012). is paper provides the rst record for the State of
Rio Grande do Norte. From 1 to 360 m in depth (Benavides-Serrato et al. 2011).
Remarks. 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 diers from E. (O.) echinophorus
for presenting a larger number of rows of dorsal longitudinal spines (9–15) and actinal
plates not mammiform. E. (O.) sentus diers for having more numerous and shorter
dorsal spines and for having mammiform abactinal plates. Walenkamp (1976) pointed
out some variations observed in specimens from Suriname, such as: number of adam-
bulacral spines (from 2 to 5), number of spines per dorsal longitudinal row (from 1 to
15), and size of the dorsal papular areas. In our study, as well as the large plasticity in
the number of dorsal longitudinal spines (from 7 to 13), a character used as diagnotic
for the genus, the number of spines surrounding the anus and the shape of the arms
also proved to be quite variable. However, these morphological variations do not seem
to be related to ontogenetic stages. Despite both adults and juveniles being present in
our material, these variations occurred among specimens of a same size class. According
to Machado et al. (2008), the most common shape of E. (O.) brasiliensis presents nar-
row and elongate arms, while a smaller proportion of individuals have short and thick
arms and less numerous spines. Tommasi (1970) synonymized E. (O.) antonioensis De
Loriol, 1904 with E. (O.) brasiliensis, though without providing further details. Ac-
cording to him (Tommasi op. cit.) the characters used by De Loriol to distinguish the
two species are all dependent on xation mode or represent highly variable characters.
An excelent discussion on the synonyms of the several species of E. (O.) brasiliensis is
found in Clark and Downey (1992). Avila-Pires (1983) proposed the presence of only
two species of Echinaster for the Brazilian littoral, E. (O.) echinophorus being restricted
to the northeastern coast and E. (O.) brasiliensis to the south and southeastern coast.
We disagree with this opinion, suggesting that E. (O.) brasiliensis also occurs in north-
eastern Brazil. More taxonomic studies are clearly needed in order to better establish
the interspecic limits between these two species.
Ecological notes. is species lives in sand, substrates of sand with mud, and
consolidated substrates, often associated with the bivalves Mytillus sp. and Lithophaga
sp. (Penchaszadeh 1973), having also been observed in banks of alassia sp. (Bena-
vides-Serrato et al. 2011). Echinaster (O.) brasiliensis is frequently found in intertidal
regions or shallow waters, and may be strongly inuenced by water salinity (Machado
et al. 2008). In this study, the species was found mainly in reef environments and

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
40
hypersaline mangrove areas, always together with E. (O.) echinophorus. According to
Machado et al. (2008), this species is common along the coast of Rio de Janeiro, being
intensively collected by aquarists without futher control on their extraction and com-
mercialization. Alves and Dias (2010) recorded its use for medical purposes (treatment
of asthma). E. (O.) brasiliensis is listed among the species vulnerable to extinction
(Machado et al. 2008).
Echinaster (Othilia) echinophorus (Lamarck, 1816)
Figures 11e–j, 12e–f
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. Brito 1962: 3. Lima-Verde 1969: 11.
Avila-Pires 1983: 436–440, gs 6–7. Tommasi 1970: 16–17, gs 41–43. Tom-
masi and Aron 1988: 3. Fernandes et al. 2002: 422. Gondim et al. 2008: 155, g.
3a. Alves et al. 2010: 757. Miranda et al. 2012: 144.
Othilia echinophora Fisher, 1919: 432.
Echinaster (Othilia) echinophorus Clark & Downey, 1992: 367–371. Magalhães et al.
2005: 63. Machado et al. 2008: 183–184. Lima and Fernandes 2009: 59. Gondim
et al. 2011: 6, g. 3e.
Material examined. 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,

Taxonomic guide and historical review of starshes in northeastern Brazil... 41
Figure 12. Some common starshes in their natural habitat. A Luidia senegalensis in a sand beach B As-
tropecten marginatus in a hypersaline mangrove C Oreaster reticulatus in a sand beach D Linckia guildingi
on coral reefs E Echinaster (O.) echinophorus in seagrass beds and F E. (O.) echinophorus over the muddy
bottom of a hypersaline mangrove. Photos: elma LP Dias.
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.,

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
42
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 Camaruxa-
tiba 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.
Type-locality. ‘Amérique du Nord’ (lectotype) (Clark and Downey 1992).
Description. Disk small (Fig. 11f). Body convex dorsally and attened ventrally.
Five (rarely three to six) short and thick arms, with a robust skeleton (Fig. 11f, g).
Abactinal plates mammiform (Fig. 11j), with one large spine. Seven to 11 series of dor-
sal longitudinal spines. Anus normally positioned within central pentagone of disk and
surrounded by 4–6 spines. Madreporite circular with numerous small granules and ir-
regular grooves. Papular areas more numerous on abactinal and lateral regions of arms
(Fig. 11h). Papular areas on actinal surface small and widely dispersed. Adambulacral
plates with 3 to 4 spines arranged transversally (Fig. 11i).
Colour. Red, orange-red, becoming brownish-red when conserved in alcohol and
dark brown when dry (Verril 1915, Hendler et al. 1995).
Distribution. Florida, the Bahamas, Gulf of Mexico, Puerto Rico, Jamaica, Ni-
caragua, Colombia, Venezuela, and Brazil (Alvarado 2011, Benavides-Serrato et al.
2011). In Brazil: CE, RN, PB, PE, AL, BA, ES, and RJ, including Abrolhos (Verrill
1915, Krau 1950, Brito 1962, Lima-Verde 1969, Tommasi 1970, Avila-Pirez 1983,
Fernandes et al. 2002, Magalhães et al. 2005, Miranda et al. 2012). From 0 to 55 m
in depth (Hendler et al. 1995).
Remarks. Echinaster (O.) echinophorus diers from E. (O.) sentus for having few
large and conspicuous spines on arms and a uniform colour (Hendler et al. 1995). It dif-
fers from E. (O.) brasiliensis for having thicker arms and mammiform abactinal plates.
According to Atwood (1973), E. (O.) echinophorus appears to contain several morpho-
logically distinct forms or closely related species. For Walenkamp (1979) the number of
series of dorsal longitudinal spines, which Perrier considered to be the main character
for separating species in the genus, is quite variable. e examined specimens have a
broad morphological variation, among which: number of rows of dorsal longitudinal
spines (from 7 to 9), number of spines surrounding the anus (from 4 to 6) and number
of granules on the madreporite (from 8 to 15). Notwithstanding, these morphological
variations do not seem to be related to ontogenetic development, because even though
specimens examined included both juvenile and adult individuals, the dierences also
aected specimens of the same size. An excellent discussion of the synonymies proposed
for the dierent species of E. (O.) echinophorus and of the taxonomic history of the spe-
cies is found in Clark and Downey (1992) and Walenkamp (1979).
Ecological notes. Lives usually in environments containing consolidated sub-
strates (Hendler et al. 1995), and may be found in estuarine regions (Nobre and

Taxonomic guide and historical review of starshes in northeastern Brazil... 43
Campos-Creasey 2000). It feeds preferably on incrustating organisms belonging to
the epifauna and on organic detritus deposited in the substrate (Jangoux and Lawrence
1982). Kempf (1966) found that E. (O.) echinophorus may occur in salinities up to
47. We found it in salinities of 52 (Tubarão River/Rio Grande do Norte), forming
dense populations. Alves and Dias (2010) commented on the use of this species for
medicinal purposes and Machado et al. (2008) remarked that one of the main threats
relates to its collecting for aquarists. Presently it is included among the Brazilian spe-
cies vulnerable to extinction (Machado et al. 2008).
Order Forcipulatida Perrier, 1884
Family Asteriidae Gray, 1840
Coscinasterias tenuispina (Lamarck, 1816)
Figure 9i–m
Asterias tenuispina Lamarck, 1816: 561–562.
Asteracanthion tenuispinus Müller & Troschel, 1842: 16.
Asterias atlantica Verrill, 1868: 368. Rathbun 1879: 145.
Asterias (Stolasterias) tenuispina Sladen, 1889: 565, 583.
Polyasterias tenuispina Perrier, 1894: 108.
Lytaster inaequalis Perrier, 1894: 98–99.
Coscinasterias tenuispina Verrill, 1914: 45. Brito 1960: 4; 1962: 2. Netto 2006: 34, g.
16c, pl. 5b. Ventura et al. 2007: 228.
Coscinasterias tenuispina var. atlantica Verrill, 1915: 20–21. Tommasi 1966: 24–244.
Stolasterias tenuispina Verrill, 1907: 324.
Coscinasterias (Stolasterias) tenuispina Fisher, 1926: 197.
Material examined. 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.
Type locality. ‘I’ ocean eropéen’ (Clark and Downey 1992).
Description. Disk small, with 1–3 madreporites (in some cases up to 5 were ob-
served). Six to nine (rarely 5 and unusually 7) thin, elongate (Fig. 9i, j), cylindrical
arms, usually of dierent sizes, the larger ones grouped to one side and the smaller
ones to the opposing side. Abactinal plates with one long, conical and pointed spine
(~2.17 mm), with base densely surrounded by bivalve pedicallariae with overlapping
valves (Fig. 9l). Carinal plates arranged in a regular series. Dorso-lateral plates forming
a reticulum. Papular regions occuring on the abactinal and actinal surfaces. Among
the abactinal plates, mainly in the intermediate areas, large bivalve pedicellariae are
found. Pedicellariae of dorsal spines sessil and with overlapping bivalves, with denteate
margins. Oral pedicallariae bivalve (Fig. 9m), consisting of a basal piece into which
two valves with smooth margins t in.

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
44
Colour. Specimens from Brazil vary from brown to orange colour (Ventura et al.
2007). According to Clark and Downey (1992), specimens from the Mediterranean
are usually yellow or whitish-yellow with black or brown spots on abactinal surface and
crowns of rusty-red pedicellariae. Individuals from the Bermudas, on the other hand,
with dorsal surface purple and the oral surface yellow, blue or violet (Verrill 1915).
Distribution. North Carolina, Gulf of Mexico, Antilles, Bermudas, Cuba, Brazil,
Portugal, Spain, France and Montenegro (Tommasi 1970, Downey 1973, Clark and
Downey 1992, Alves et al. 2002, Waters and Roy 2003, Kascelan and Mandic 2007).
In Brazil: BA, ES, and SP, including Abrolhos (Rathbun 1879, Verrill 1915, Brito
1960, Tommasi 1970, Ventura et al. 2007). Intertidal to 165 m in depth (Clark and
Downey 1992).
Remarks. Two species of the genus Coscinasterias Verrill, 1870 are known for the
Atlantic, C. tenuispina and C. calamaria (Gray, 1840). e rst is widely distributed
through the Atlantic and Mediterranean, while the second is restricted to South Africa,
Angola, and Madagascar (Mah 2013). C. tenuispina diers from C. calamaria for hav-
ing intercrossing pedicellariae with a well developped terminal tooth (except in some
specimens from Brazil). Clark and Downey (1992) suggested a subspecic distinc-
tion for these species on the basis of morphological similarities observed in Brazilian
and South African specimens. However, according to Waters and Roy (2003) these
observations were made on the basis of juvenile specimens and it is thus necessary to
undertake new morphological analyses to clarify the close relationships between C.
calamaria and C. tenuispina (Waters and Roy 2003). In the phylogeographic analysis
of Waters and Roy (op. cit.) for species of Coscinasterias, morphological variations were
observed between populations from Brazil on the one side and from the Bermudas
and the Mediterranean on the other. No morphological variations are observed in the
specimens examined by us.
Ecological notes. Lives in consolidated substrates, including areas with strong
hydrodynamism (Machado et al. 2008). Coscinasterias tenuispina has extra-oral diges-
tion and feeds on epifaunal organisms, mainly mussels (Ventura et al. 2007). It is
a ssiparous species, which presents an annual gonadal cycle and a long period of
spawning (Alves et al. 2002). According to these authors, the preponderance of males
in the population of Itaipu Beach (Niterói/Rio de Janeiro) suggests that assexual re-
production by ssion is predominant and, consequently, that the number of clones
must be signicant. According to Brito (1962), this species is very common in Cabo
Frio (Rio de Janeiro). However, since the rst record by Rathbun (1879) of C. ten-
uispina for Abrolhos (Bahia), the species has not been cited again o the northeast
region of Brazil. As pointed out by Machado et al. (2008), although this species has a
wide geographical distribution, its range is discontinuous, probably due to its assexual
reproduction, that limits dispersal ability. Presently the species is listed among those
vulnerable to extinction, having among the main causes of population decline the
constant destruction of its habitat, the erosion of the substrate, the eects of pollut-
ants, the precarious sanitation and the excess of tourists and divers within their range
of occurrence (Machado et al. 2008).

Taxonomic guide and historical review of starshes in northeastern Brazil... 45
Discussion
e 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 (Tommasi 1970). One exception is Mithrodia clavigera, which
represents a new record for the northeast and is typically a species of deeper waters.
Another two species represent new records for northeast Brazil: Astropecten alligator
and Luidia ludwigi scotti.
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). ese 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 (Zulliger et al. 2010). Although common in
marine communities, the taxonomy of the species composing this order and the phylo-
genetic relationships of the Paxillosida are still uncertain and contradictory (Matsubara
et al. 2005). During many years this order was considered to be the most primitive in
the class, due mainly to the absence of an anus and of ventosae on the ambulacral feet
(Jangoux 1982). However, a reexamination of characters evidenced that these charac-
ters represent adaptations to life in sandy environments that produced character losses
instead of being primitive absences (Matsubara et al. 2005).
e genus Astropecten represents one of the most complex taxa within the class As-
teroidea, in which species exibit great morphological plasticity, making identication
of species dicult. According to Zulliger et al. (2010), the high phenotypic variability
of this genus resulted in the naming of several subspecies. Six of these were recognized
in Brazil (Tommasi 1970). Presently they are all synonymized (Clark and Downey
1992; Mah 2013) and the records of Astropecten armatus for the Brazilian coast repre-
sent synonyms of A. brasiliensis (Tommasi 1999, unpublished data). Among the sev-
eral taxonomic characters used for the identication of species, the appearance of the
paxillae and of the superomarginal plates, together with the number and shape of the
spines of the marginal fringe are the characters that contribute most for the identica-
tion of species. On the other hand, the number and shape of the adambulacral spines,
characters that were much used by authros such as Bernasconi (1955, 1957), Tommasi
(1970) and Clark and Downey (1992), proved to be very similar among specimens
and were thus not considered to be good characters for the taxonomy of the genus.
Another taxonomicly complex genus is Echinaster that, similarly to Astropecten, pre-
sents large morphological variability, making species identication dicult. According
to Clark and Downey (1992), the species of this genus occurring in Brazil are polymor-
phic, with possible hybridization among them. Presently three occurrences of Echinaster
(Othilia) are considered valid along the Brazilian coast: E. (O.) brasiliensis, E. (O.) echi-
nophorus, and E. (O.) guyanensis. According to Avila-Pires (1983), the rst occurs only
in the south and southeastern regions, below Espírito Santo, and the second is restricted
to the northeastern region. at author does not conrm the presence of Echinaster (O.)
guyanensis, but Clark and Downey (1992) indicate its distribution as being from Cen-

Anne Isabelley Gondim et al. / ZooKeys 449: 1–56 (2014)
46
tral America to Belém (Pará). Magalhães et al. (2005) recorded E. (O.) guyanensis for
the littoral of Bahia, but this occorrence could not be conrmed in the present study.
On the basis of the material we examined we cannot agree with Avila-Pires (1983) that
E. (O.) brasiliensis does not occur in the northeastern region. We conrm its presence
in northeastern Brazil, although it is rarer there than its congener E. (O.) echinophorus.
Among the several characters used in the taxonomy of the genus, the aspect of the en-
doskeleton plates represent one of the most important criteria for species recognition.
e number of spine rows on the arms was used by Avila-Pires (1983) and Bernasconi
(1957) to separate species, but it turned out be be quite variable in this study.
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). ere
are still no records of asteroids for the coasts of Maranhão and Sergipe. ese numbers
indicate the scarce knowledge available on the Asteroidea from northeast Brazil, which
represents one of the least known areas regarding the benthic macrofauna (Ventura et
al. 2007). e species refered for northeast Brazil (n = 21 spp) are mostly shallow water
species (up to 30 meters in depth), with only 4 species representing deep-water forms.
It remains crucial to conduct inventories in little explored areas such as northeast Bra-
zil, particularly in deep waters.
Presently 374 species of sea-stars are known for the Atlantic Ocean (Clark and
Downey 1992), the Gulf of Mexico being the most speciose area, with 126 recorded
species (Pawson et al. 2009), followed by the Caribbean, with 116 spp (Alvarado et
al. 2011). e fauna from Brazil represents only 20.5% of the known species from the
Western Atlantic, while those from northeast Brazil represent 4.82%. Considering the
similarity of the Brazilian fauna with the Caribbean region, and taking into account
the extent of the Brazilian coastline, the necessity to make an inventory and to describe
the Brazilian fauna becomes urgent. According to Marques and Lamas (2006), the
degree of knowledge of the marine fauna from Brazil is far from ideal and the most no-
table gap in our knowledge refers to the the invertebrates mainly from northeast Brazil.
Another three species recorded previously for northeast Brazil were not conrmed
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 (Brito 1962); Asterina stel-
lifera (Möbius, 1859) (as Enoplopatiria marginata (Hupe, 1857)) was cited by Bernas-
coni (1955) for the Abrolhos Archipelago and Echinaster (O.) guyanensis was recorded
by Magalhães et al. (2005) for the coast of Bahia. Ventura et al. (2007) cited A. acuti-
radiatus, N. arenatus and P. dentatus for the continental shelf of Bahia. Unfortunately,
we have been unable to retrieve these species in the present study.
Due to the negligible previous knowledge on the Asteroidea from the littoral of north-
eastern 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
scientic collections turned out to be of fundamental importance, permitting a historical
panorama of the Asteroidea from northeast Brazil. Deep-water sampling in the studied

Taxonomic guide and historical review of starshes in northeastern Brazil... 47
region, restricted to sporadic expeditions, as accounted for in Clark and Downey (1992),
has still not provided many published records on asteroids. e diversity of the deep-water
and abyssal marine fauna of the South West Atlantic perhaps remains the least known in
the world, and clearly represents the next marine frontier to be systematically sampled and
studied, both for asteroids and for other marine groups.
Acknowledgements
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 col-
leagues 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 schol-
arship and Martin L. Christoersen 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.
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