Echinoidea

Journal of the Marine Biological Association of the United Kingdom, page 1 of 35.
# Marine Biological Association of the United Kingdom, 2016

doi:10.1017/S0025315416001569
Echinoidea and Holothuroidea

(Echinodermata) of the Trindade and Martin
Vaz Archipelago, off Brazil, with new
records and remarks on taxonomy and
species composition
luciana martins1, camilla souto2, joel braga1 and marcos tavares1
1
Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42494, São Paulo–SP, CEP: 04218– 970, Brazil, 2Department of
Integrative Biology, Museum of Paleontology, University of California, Berkeley, 1101 Valley Life Science Building, Berkeley –CA
94720, USA
The degree of isolation of oceanic islands makes these environments an excellent model for evolutionary studies. Proper
knowledge of the species composition of oceanic islands, however, is required to better understand evolutionary processes
(e.g. speciation events). A 3-year survey in the shallow waters (up to 30 m) of the Trindade and Martin Vaz oceanic
insular complex, and a literature review on the data published for these islands and for Fernando de Noronha and São
Pedro and São Paulo oceanic archipelagos have been conducted to document the biodiversity of echinoids and holothuroids
from these isolated Brazilian oceanic islands. Sixteen species were collected and characterized morphologically, including two
first records for the South Atlantic and one for Brazil. Comparison with conspecific specimens from the Brazilian coast and
congeners was also done. Species richness increased from six to 18; the richness in Trindade Island being the highest among the
South Atlantic oceanic islands. However, these islands remain undersampled beyond 30-m depth. Endemism was very low,
suggesting the potential role of oceanic currents and seamounts as stepping-stones in transoceanic dispersal of species to
remote islands. The Brazilian oceanic islands are impoverished oceanic outposts of the Brazilian Province; nevertheless,
endemic species and intraspecific morphological variations compared with the mainland suggest they may also be regions
of speciation. Documenting their biodiversity is critical for effective management and conservation of their marine ecosystems.
Keywords: Oceanic islands, Saint Peter and Paul Rocks, biodiversity, species richness, new records
Submitted 12 June 2016; accepted 18 October 2016
INTRODUCTION shallow-water species and is directly reflected in the taxo-

nomic composition of the TMV invertebrate fauna (Leal &
The Trindade and Martin Vaz Oceanic Archipelago (TMV) is Bouchet, 1991; Gasparini & Floeter, 2001; Anker et al., 2016).
the above-water portion of a submarine volcanic chain of The levels of endemism and species richness in the TMV
about 3–3.5 my rising 5500 m above the seafloor (Marques vary among taxa (Carvalho, 1950; Oliveira, 1951; Vannucci,
et al., 1999). Trindade Island and the much smaller Martin 1951; Albuquerque & Guille, 1991; Tavares, 1999; Leal,
Vaz islands are only 49 km apart from each other, but 2000; Floeter & Gasparini, 2000; Paiva, 2006; Coimbra et al.,
almost 1200 km away from the Brazilian coast and some 2009; Moraes, 2011; Coimbra & Carreño, 2012; Anker et al.,
4200 km from the nearest African shore. The archipelago is 2016). However, to a certain extent, this may be only a reflec-
mainly under the influence of the southward flow of the tion of sample size (McClain, 2007), as not only the Brazilian
Brazil Current, which runs along the Brazilian coast from oceanic islands and seamount chains are yet to be intensively
about 108S to 388S (Stramma et al., 1990; Peterson & sampled, but also many surveys in the area have been and con-
Stramma, 1991; Podesta et al., 1991; Stramma, 2001). The tinue to be taxon-oriented.
influence of the oceanic main current patterns in this zone The current knowledge about the echinoderm diversity of
and the existence of a seamount chain oriented east-west the TMV is restricted to a few studies (Oliveira, 1951; Krau,
between TMV (whose tops can be as shallow as 50 m 1952; Bernasconi, 1955b, 1957; Brito, 1971; Lima-Verde,
[Motoki & Motoki, 2013]) and the Brazilian continental 1969; Guille & Albuquerque, 1987, 1990; Vadon, 1988;
shelf, can potentially serve as stepping stones for spillover of Albuquerque & Guille, 1991). Ophiuroids are the most
studied echinoderms, while holothuroids have never been
reported from the archipelago. During a 4-year project
Corresponding author:
(ProTrindade/CNPq), five campaigns to the TMV oceanic
L. Martins archipelago were conducted between 2012 and 2015, which
Email: [email protected] resulted in about 605 lots of shallow-water echinoderms,
Downloaded from http:/www.cambridge.org/core. Instituto De Biociencias, on 24 Nov 2016 at 10:24:50, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms.
http://dx.doi.org/10.1017/S0025315416001569
2 luciana martins et al.
collected mainly by scuba diving and thus forming a solid base (see Taxonomic List). Seven species of echinoids were found
for taxonomic assessment of the shallow-water echinoderm in the Trindade Island (three new records) and three in the
fauna of the archipelago. Here we report on 16 echinozoan Martin Vaz archipelago (all new records). Nine species of
species, 11 of which represent new records for TMV. Three holothuroids were found in the Trindade Island and two in
species are recorded from the south-western Atlantic and the Martin Vaz archipelago (all new records); three species
Brazilian territorial waters for the first time. Additionally, were recorded for the SPP for the first time: Eucidaris tribu-
we studied a few unreported samples from the São Pedro loides, Holothuria (Halodeima) manningi and Isostichopus
and São Paulo Oceanic Archipelago, a group of small islets badionotus.
rising from the mid-Atlantic ridge located about 1146 km None of the species sampled are endemic to the SPP and
away from Brazil and some 1960 km from the nearest TMV. Holothuria (Halodeima) manningi, previously regarded
African coast. We take the opportunity to elaborate on the as an Ascension endemic is herein recorded for Trindade
taxonomy of the seven echinoid and the nine holothuroid Island and the SPP, and is apparently an insular species
species reported herein, providing morphological descriptions restricted to the South Atlantic Ocean. Actinopyga agassizii
and illustrations for all of them. We also present morpho- and Pseudoboletia maculata have their first record from the
logical comparisons between the studied species and their south-western Atlantic, and Euapta lappa from Brazil. The
relevant closest congeners and between insular and coastal endemic species from Trindade Island, Centrostephanus bes-
specimens, and provide some notes on the echinozoan nardi Bernasconi and Clypeaster oliveirai Krau as well as den-
species composition of the Trindade and Martin Vaz drochirotids, which are very abundant on the Brazilian coast,
Oceanic Archipelago. were not re-collected there. Known depth range was extended
for Holothuria (Halodeima) manningi and Pseudoboletia
maculata (see Systematics).
MATERIALS AND METHODS
TAXONOMIC LIST
Most of the material reported herein was collected during 168 SUBPHYLUM ECHINOZOA
scuba diving operations between 4–30 m, at numerous sta- CLASS ECHINOIDEA
tions around Trindade Island (20830′ S 29820′ W) and two sta-
tions off Martin Vaz Islands (20830′ S 28851′ W) (see Anker Order Cidaroida
et al., 2016: figure 1). Also included in this study are a few Family Cidaridae Gray
unreported samples from the São Pedro and São Paulo Eucidaris tribuloides (Lamarck, 1816)
Archipelago (20830′ S 29818′ W, also known as Saint Peter
and Saint Paul Rocks). All specimens were preserved in 75% Order Diadematoida
ethanol; some were photographed alive in situ and some Family Diadematidae Gray
photographed after preservation. Diadema ascensionis Mortensen, 1909
Holothuroid morphological techniques and terminology
are after Rowe & Doty (1977) and Samyn et al. (2006). Order Camarodonta
Echinoid pedicellariae were classified according to Coppard Family Echinometridae Gray
et al. (2010). Ossicles, pedicellariae and spines were dried Echinometra lucunter (Linnaeus, 1758)
and mounted on metal stubs with double-sided tape, coated Family Toxopneustidae Troschel
with gold and observed with a LEO 440 Scanning Electron Pseudoboletia maculata Troschel
Microscope (SEM). Measurements: maximum lengths of Tripneustes ventricosus (Lamarck, 1841)
holothuroids were obtained from ethanol fixed specimens;
maximum sizes of echinoids were accessed from test diameter Order Clyperasteroida
and size of table ossicles from disc diameter. Family Mellitidae
Abbreviations: TMV, Trindade and Martin Vaz Leodia sexiesperforata (Leske, 1778)
Archipelago; SPP, São Pedro and São Paulo Archipelago
(Saint Peter and Saint Paul Rocks); BOI, Brazilian oceanic Order Spatangoida
islands; ASC, Ascension Island; STH, Saint Helena Island; Family Brissidae Gray
AST, Ascension and Santa Helena Archipelago; my, million Plagiobrissus grandis (Gmelin, 1791)
years; spm, specimen.
Acronyms: California Academy of Sciences, San Francisco, CLASS HOLOTHUROIDEA
USA (CAS-IZ), Museu de Zoologia da Universidade de São
Paulo, Brazil (MZUSP), Natural History Museum, London, Order Apodida
UK (NHM-UK), National Museum of Natural History, Family Chiridotidae Östergren
Washington DC, USA (USNM), University of California, Chiridota rotifera (Pourtalès, 1851)
Museum of Paleontology, Berkeley, USA (UCMP). Family Synaptidae Burmeister
Euapta lappa (J. Muller, 1850)
RESULTS Order Aspidochirotida

Family Holothuriidae Eschscholtz
Actinopyga agassizii (Selenka, 1867)
Echinoids and holothuroids from Trindade Holothuria (Cystipus) pseudofossor Deichmann, 1930
A total of 10 families of Echinozoa were sampled, including Holothuria (Halodeima) manningi Pawson, 1978
seven species of echinoids and nine species of holothurians Holothuria (Halodeima) grisea Selenka, 1867
http://dx.doi.org/10.1017/S0025315416001569
echinozoans from the brazilian oceanic islands 3
Holothuria (Semperothuria) surinamensis Ludwig, 1875 13 m, 23.8.2013, 4 spms 15 –30 mm long (MZUSP 664).
Holothuria (Thymiosycia) arenicola Semper, 1868 Trindade Island, ES, Brazil (20829′ – 20831′ S 29817′ –
Family Stichopodidae Haeckel 20820′ W) – Andrada beach, 10 m, 17.7.2013, 2 spms 9–
Isostichopus badionotus (Selenka, 1867) 24 mm long (MZUSP 666); intertidal, 21.7.2013, 2 spms
15 –30 mm long (MZUSP 654, 671). Cabritas beach, 9 m,
14.6.2012, 3 spms 20– 40 mm long (MZUSP 690). Calheta
systematics
beach, 23.7.2011, 1 spm 35 mm long (MZUSP 737); 4 m,
Class ECHINOIDEA
24.6.2012, 1 spm 30 mm long (MZUSP 677); 14 m,
Order CIDAROIDA Claus, 1880
26.6.2012, 3 spm 15– 40 mm long (MZUSP 669); 17 m,
Family CIDARIDAE Gray, 1825
8.5.2014, 1 spm 25 mm long (MZUSP 983). Calheta Island,
Genus Eucidaris Pomel, 1883
12 m, 18.6.2012, 2 spms 11 –25 mm long (MZUSP 675,
Eucidaris tribuloides (Lamarck, 1816)
676); 4 m, 14.7.2013, 3 spms 8–25 mm long (MZUSP 658).
(Figures 1 –3)
Crista do Galo beach, 15 m, 21.5.2014, 3 spms 5–10 mm
Cidarites tribuloides Lamarck, 1816, p. 56. long (MZUSP 990). Enseada da Cachoeira, 12 m, 16–
Eucidaris clavata Mortensen, 1928, pp. 408 –411, pls. 42, 67, 20.6.2012, 3 spms 6– 35 mm long (MZUSP 685, 689); 18 m,
73, 86. 4.7.2012, 1 spm 5 mm long (MZUSP 647); 14 m, 9.7.2012, 1
Cidaris tribuloides – Rathbun, 1879, p. 143; Agassiz, 1881, spm 30 mm long (MZUSP 687); 21 m, 12.7.2012, 1 spm
pp. 36– 37, pl. 1; Brito, 1968, pp. 17 –18, pl. 8. 35 mm long (MZUSP 683); 10 m, 9.7.2013, 2 spms 20–
Eucidaris tribuloides – Döderlein, 1887, p. 42, pl. 9; 30 mm long (MZUSP 667). Enseada de Orelhas, 8 m,
Mortensen, 1928, pp. 400 – 408, pls. 41, 48, 73, 86; 24.1.2012, 1 spm 30 mm long (MZUSP 665); 10 m,
Bernasconi, 1955a, pp. 52– 54, pl. 1. 15.6.2012, 5 spms 10– 25 mm long (MZUSP 682); 14 m,
Eucidaris tribuloides var. africana Mortensen, 1909, pp. 30.6.2012, 2 spms 31.5– 36 mm long (MZUSP 679); 15 m,
40 –43, pl. 6, 10, 13 –16. 25.6.2012, 1 spm 20 mm long (MZUSP 656); 14 m, 6.7.2013,
Eucidaris tribuloides tribuloides – Serafy, 1979, pp. 15 –19, 6 spms 15 –35 mm long (MZUSP 673); 12 m, 4.5.2014, 1
figure 3. spm 26 mm long (MZUSP 1013); 10 m, 16.5.2014, 2 spms
30 –40 mm long (MZUSP 989); 11 m, 21.5.2014, 2 spms
material examined 25 –30 mm long (MZUSP 986). Enseada Portuguesa, 15 m,
Martin Vaz Island, ES, Brazil (20828′ –20831′ S 29818′ – 10 –15.7.2012, 9 spms 5–35 mm long (MZUSP 650, 663,
20851′ W) – 19 m, 22.1.2012, 1 spm 15 mm long (MZUSP 688). Farol beach, 13 m, 17.4.2014, 7 spms 10 –35 mm long
659); 13 m, 23.7.2013, 3 spms 4.9 – 16.3 mm long (MZUSP (MZUSP 987, 988); 14 m, 22.4.2014, 1 spm 10 mm long
655); 12 m, 24.7.2013, 2 spms 25 mm long (MZUSP 668); (MZUSP 1016); 13 m, 20.5.2014, 6 spms 10 –40 mm long
Fig. 1. Eucidaris tribuloides (Lamarck, 1816): A– E, G – H (MZUSP 679); F (MZUSP 651). (A) Specimen from Trindade Island photographed in situ, at 14 m
depth; (B) aboral view; (C) oral view; (D) ambulacrum; (E) interambulacrum; (F) detail of the apical disc; (G) Aristotle’s lantern; (H) apophysis. Scale bars:
B– E, 10 mm; F, 5 mm.
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Fig. 2. Eucidaris tribuloides (Lamarck, 1816): A –E, I– L, O (MZUSP 669); F –H (MZUSP 1028); M– N (MZUSP 679). SEM images – (A – B) valves of narrow
fistulate globiferous pedicellariae, and (C) detail of the head; valve of broad fistulate globiferous pedicellariae from a specimen (D) from the BOI and (F) from
Boipeba island (Brazilian coast); (E, G) detail of the pedicellariae head, respectively; (H) detail of the edge of the pedicellaria depicted in (F); (I) slender
tridentate pedicellariae from a specimen from Boipeba island; (J) plate from peristomial membrane; (K – M) tube feet ossicles; and (N – O) detail of a spine.
Scale bars: A – B, D, F, I, 150 mm; J, 400 mm; K – O, 50 mm.
(MZUSP 984). Lixo beach, 23.5 m, 24.6.2012, 4 spms 8– Castelhanos beach, Boipeba, 17.9.2012, 2 spms 38 –45 mm
17.5 mm long (MZUSP 653); 25 m, 2.7.2012, 2 spms 25– long (MZUSP 1028, 1030); Itapuã Beach, Salvador,
30 mm long (MZUSP 678). Ponta do Monumento, 12 m, 29.7.2007, 1 spm 22 mm long (CAS-IZ 175395). Colombia
16.6.2012, 1 spm 25 mm long (MZUSP 651); 19.5 m, (108N 758W) – Cartagena Bay, 3 spms 22– 26 mm long
30.6.2013, 1 spm 32 mm long (MZUSP 670). Ponta do (CAS-IZ 91365); La Cieba, 1931, 1 spm 30– 34 mm long
Paredão, 19 m, 19.6.2012, 3 spms 6– 25 mm long (MZUSP (UCMP 123406). Gulf of Mexico (24826′ N 81848′ W) –
652); 20 m, 20.6.2012, 4 spms 10– 40 mm long (MZUSP 68 m, 15.1.1885, 3 spms 31– 51 mm long (CAS-IZ 91340).
686). Ponta Noroeste, 12 m, 4.7.2012, 1 spm 10 mm long Jamaica (188N 77′ W) – 3 spms 30– 34 mm long (UCMP
(MZUSP 657). Ponta Norte, 14 m, 18.7.2013, 1 spm 20 mm 123429). Panama (Atlantic coast) – 6.1913, 1 spm 52 mm
long (MZUSP 662). Prı́ncipe beach, 19.5 m, 9.7.2013, 1 spm long (CAS-IZ 91372). San Pedro Macoris beach, Hispaniola
35 mm long (MZUSP 672). Racha Island, 27 m, 22.6.2012, 5 Island, Dominic Republic (18842′ N 69812′ W) – 1.5 m,
spms 10– 30 mm long (MZUSP 684); 30 m, 12.7.2012, 1 4.3.1965, 1 spm 14 mm long (UCMP 123408). St. John
spm 30 mm long (MZUSP 681); 24 m, 13.7.2012, 2 spms Island, Virgin Islands (188N 648W) – 4.1958, 1 spm 28 mm
15 –35 mm long (MZUSP 691); 25 m, 16.7.2012, 1 spms long (UCMP 123407). Eucidaris galapagensis Döderlein,
15 mm long (MZUSP 661). Secon, 11 m, 23.4.2014, 3 spms 1887: Galapagos Islands, Ecuador (0815′ S –0815′ N 89845′ –
5– 8 mm long (MZUSP 1011); 9 m, 12.5.2014, 6 spms 15– 90815′ W) – Academy Bay, Santa Cruz Island, 0– 3 m, 24–
30 mm long (MZUSP 985). Tartarugas beach, 9.5 m, 25.1.1964, 2 spms 46– 49 mm long (UCMP 123409). Baltra
26.4.2012, 1 spm 30 mm long (MZUSP 680). Island, 2– 4.5 m, 12.2.1964, 2 spms 39– 51 mm long (UCMP
123410, 123436). Darwin Bay, Genovesa Island, 0–3 m, 1–
additional material 16.2.1964, 3 spms 25 –34 mm long (UCMP 123411,
São Pedro and São Paulo Archipelago, PE, Brazil (20830′ S 123437 – 123438). Plaza Island, 8.2.1964, 1 spm 20 mm long
29818′ W) – 21.2 –5.3.2009, 2 spm 30– 35 mm long (MZUSP (UCMP 123412). Venedig Lagoon, Santa Cruz Island, 3–
735, 736). Ilha Belmonte, intertidal, 27.10.2007, 1 spm 4.5 m, 19.2.1964, 1 spm 48 mm long (UCMP 123413).
28 mm long (MZUSP 1029). Eucidaris metularia (Lamarck, 1816): Marianas Islands,
Guam, U.S.A. (12 – 218N 1458W) – 0.5 – 1.5 m, 10.8.1992, 1
comparative material examined spm 23 mm long (CAS-IZ 108829). Oahu, Hawaii, USA
Eucidaris tribuloides: Alagoas, Brazil (8840′ –10829′ S 3589′ – (218N 1578W) – 21.7.1939, 1 spm 23 mm long (CAS-IZ
36823′ W) – 21.11.1998, 4 spms 28 –36 mm long (CAS-IZ 96779). Eucidaris thouarsii (L. Agassiz & Desor, 1846):
15913, 98043). The Bahamas – 1 spm 50 mm long (CAS-IZ Brasilito Bay, Costa Rica (10824′ N 85850′ W) – 0–12 m,
91346); Abaco Island, 8.8.1954, 2 spms 35– 42 mm long 13.5.1968, 1 spm 31 mm long (UCMP 123428). Colombia
(CAS-IZ 91363); Gibson Bay – 3 spms 32 –48 mm long (2857′ –6816′ N 77826′ –79811′ W) – Punta Cotudo, 2 –27 m,
(UCMP 123405). Bahia, Brazil (12857′ –13839′ S 388W) – 1– 2.5.1968, 1 spm 25 mm long (UCMP 123417). Punta
http://dx.doi.org/10.1017/S0025315416001569
Fig. 3. Ratios of structures of Eucidaris specimens plotted against their test diameter. Upper graph: apical disc diameter/peristome diameter ratio; bottom graph:
(Int) median interambulacral width/(Are) ambital areole diameter. Measurements taken from E. clavata from AST (data from Pawson (1978)); E. tribuloides from
the Caribbean, West Africa (includes data from Pawson (1978), and North-east Brazil (Alagoas and Bahia states (NE BR)) (in white), and SPSP and TMV (in
black); and E. galapagensis, E. metularia and E. thouarsii from the eastern Pacific (in grey).
Mono, Cauca, 3–33.5 m, 27.4.1968, 1 spm 21 mm long Perradial zone about 1.5 wider than poriferous zone, two
(UCMP 123418). Culebra Island, Canal Zone, Panama series of secondary tubercles in each half ambulacrum
(8855′ N 79832′ W) – 0– 4.5 m, 5.4.1968, 2 spm 29 –33 mm (outer series larger than inner). Interambulacra (Figure 1E)
long (UCMP 123419, 123445). Mexico – Islas Revillagigedo, more than twice as wide as ambulacra (Figure 1D); primary
Clarion Island, 2 spms 44.5 – 59 mm long (CAS-IZ 91360). tubercle areola at ambitus almost twice width of middle inter-
Jalisco, Tenacatita Bay, 4.5 –12 m, 27.5.1968, 1 spm 22 mm ambulacral region. Peristomial membrane covered with
long (UCMP 123420). Los Angeles, Sonora, intertidal, plates. Cidaroid lantern (Figure 1G). Ear-shaped apophyses
7.4.1959, 1 spm 32 mm long (UCMP 123440). Puerto (Figure 1H). Tubercles perforate, non-crenulate.
Vallarta, intertidal, 5.4.1968, 1 spm 22 mm long Primary spines pencil-like (mean diameter 3.5 mm). Spine
(UCMP 123421). Punta Sur, Esmeraldas, Ecuador (180′ N shaft with short projections pointing outwards (resembling
79854′ W) – 7.5 –12 m, 23.4.1968, 1 spm 30 mm long warts in optical microscope), ending in crown; scrobicular
(UCMP 123416). spines short, broad flattened; marginal ambulacral spines
short, narrow, flattened. Two types of fistulate globiferous
description pedicellariae. Narrow type slender, blade margins with
Test thick, globular in outline, diameter 5–40 mm. Apical disc numerous small teeth throughout, glandular cavity with
(Figure 1F) monocyclic or hemicyclic, ocular plates II and III large teeth, and terminal tooth often bifurcated or completely
exsert (ocular plates I, IV and V in contact with elongated split into two (370 –520 mm; Figure 2A –C). Broad type
periproctal plates); periproctal plates numerous, tubercles slightly curved, broad base, top half of blade with long mar-
tiny, numerous. Peristome about 1.5 larger than apical disc. ginal teeth, else smooth, glandular cavity with large teeth, no
http://dx.doi.org/10.1017/S0025315416001569
terminal tooth (570– 690 mm; Figure 2D). Tridentate pedicel- Variation within E. tribuloides: the shape of the terminal
lariae not found. Ossicles showing as thick plates with small tooth of the small globiferous pedicellariae from the
perforations (1200 –1320 mm; Figure 2J) in peristomial mem- Trindade specimens is slightly different from the terminal
brane; thorny rods (115 –225 mm; Figure 2L, M) and crowns tooth of E. tribuloides from the Caribbean; however, data
(220 –260 mm; Figure 2K) in the tube feet. from specimens from the other oceanic islands are needed
Colour light brown, denuded test green with white tuber- for further comparison.
cles; apical disc plates with transversal dark green stripe Specimens from Boipeba island (Bahia, north-eastern
across middle of plate forming circle around anus; primary Brazil): two specimens from Bahia analysed for comparison
spines brownish, warts dark, crown pale, scrobicular spines have some morphological differences when compared with
creamy with dark brown tips, marginal ambulacral spines other Eucidaris tribuloides specimens: the DA/WI ratio is
apical brown or greenish with dark tips. very high (ratios are 1.25 and above; see Figure 3); slender tri-
dentate pedicellariae are abundant (815 –890 mm; Figure 2I)
distribution (vs. rare or lacking); and the broad fistulate globiferous pedi-
Known from North Carolina (USA) to Rio de Janeiro (Brazil), cellariae have a straight edge (vs. curved), the blade has mar-
and Gulf of Guinea; oceanic island distribution: Bermuda, ginal teeth throughout (vs. only up to the top half), and the
Cape Verde, São Tomé, Fernando de Noronha, ASC, STH, glandular cavity has fewer large teeth (1– 2 vs.3–4 in each
TMV (Clark, 1898; Mortensen, 1928; Bernasconi, 1955a; side), and a terminal tooth (300 –330 mm; Figure 2F) (vs. ter-
Brito, 1968; Lima-Verde, 1969; Pawson, 1978; Serafy, 1979; minal tooth lacking). The blades of the broad fistulate globi-
Brown, 2015; present paper). Bathymetric range: 0–800 m berous pedicellariae from Eucidaris depicted in Mortensen
(Serafy, 1979). (1909, 1928) are curved in Eucidaris tribuloides and only den-
ticulated up to their top half in all Eucidaris species; if those
drawings are accurate, the pedicellariae from the Boipeba
biological notes island specimens are unique within the genus. Narrow fistu-
In TMV, specimens were often found in association with bra- late globiferous pedicellariae is very similar (445– 520 mm).
chyuran crabs (Majidae), sponges and algae. Intraspecific variation within E. tribuloides could be related
to little geographic isolation between the oceanic islands and/
remarks or local environmental differences; however, the specimens
The specimens from the BOI differ from Eucidaris australiae from Boipeba Island could be a new species and need add-
Mortensen, 1950 in having the primary spines ending in a itional observations. Further molecular studies with more
crown and narrower globiferous pedicellariae, from E. metu- genes and additional specimens from the oceanic and contin-
laria by the presence of tubercles widespread in the genital ental islands are necessary to study their connectivity.
plates, from E. thouarsii by colouration of the scrobicular
spines (light with dark tip vs. all dark in E. thouarsii); and Order DIADEMATOIDA Duncan, 1889
from E. galapagensis by the absence of thick spines (5 – Family DIADEMATIDAE Gray, 1855
7 mm). Pawson (1978) showed that the thickness of the Genus Diadema Gray, 1825
spines in Eucidaris from the Ascension Island increases with Diadema ascensionis Mortensen, 1909
the depth, but all E. galapagensis analysed herein, from (Figures 4 & 5)
various depths, had thick spines compared with the other Diadema ascensionis Mortensen, 1909, pp. 55– 58, pls. 7, 16.
Eucidaris species. Diadema setosum – Rathbun, 1879, p. 143.
Eucidaris tribuloides vs. Eucidaris clavata Mortensen, 1928: Diadema ascensionis – Mortensen, 1940, pp. 279 –281, pls.
according to Pawson (1978), there are two valid Eucidaris 48, 54, 61, 73; Tommasi, 1966, p. 11, figures 3–12; Brito, 1968,
species in the Atlantic Ocean, E. tribuloides and E. clavata, pp. 19 –20, pl. 9; Brito, 1971, p. 264; Gondim et al., 2013, p.
both differentiated by the ratio between the apical disc and 432, figures 1 & 2.
the peristome width (AW/PW), and by the ratio between Diadema antillarum ascensionis – Pawson, 1978, p. 17.
the diameter of the primary tubercle’s areola and the width
of the denuded area of the interambulacrum (DA/WI; both examined material
measurements taken at the ambitus). Figure 3 shows the Martin Vaz Island, ES, Brazil (20830′ S 29818′ W) – 13 m,
ratios calculated herein combined with data extracted from 23.7.2013, 3 spms 15– 25 mm long (MZUSP 641, 644).
Pawson (1978). Only specimens over 20 mm were plotted. Trindade Island, ES, Brazil (20829′ – 20831′ S 29817′ –
The AW/PW ratio (Figure 3) shows that the specimens 20820′ W) – Andrada beach, 10 m, 17.8.2013, 3 spms 25–
from the BOI are more similar to E. clavata and to E. thouarsii 60 mm long (MZUSP 639, 643). Cabritas beach, 9.5 m,
than to the Caribbean E. tribuloides. Specimens from the 14.6.2012, 1 spm 40.8 mm long (MZUSP 610). Calheta
Brazilian coast were nested within both groups. Conversely, beach, 16 m, 20.5.2014, 3 spms 10– 21 mm long (MZUSP
the DA/WI ratio (Figure 3) shows that the specimens from 1018). Calheta Island, 12.6.2012, 1 spm 30 mm long
the BOI are more similar to E. tribuloides than to the other (MZUSP 600). Crista do Galo beach, intertidal, 12.6.2012, 1
Eucidaris species. However, this similarity is only present in spm 40 mm long (MZUSP 640); 12.7.2012, 1 spm 35 mm
specimens over 35 mm in test diameter because specimens long (MZUSP 601); 15 m, 21.5.2014, 3 spms 40– 50 mm
from the BOI did not change the ratio with ontogeny. long (MZUSP 1003). Enseada da Cachoeira, 10.5 m,
Because these differences between E. tribuloides and 9.7.2013, 1 spm 31.8 mm long (MZUSP 646). Enseada
E. clavata are very subtle, we recommend accepting only the Portuguesa, 9.5 m, 14.6.2012, 1 spm 16 mm long (MZUSP
species E. tribuloides in the Atlantic Ocean, which is in agree- 638); 15.7.2013, 1 spm 50 mm long (MZUSP 599); 10 m,
ment with the molecular analysis with the COI gene per- 18.4.2014, 3 spms 20 –25 mm long (MZUSP 1005). Farol
formed by Lessios et al. (1999). beach, 12 m, 15.7.2013, 1 spm 25 mm long (MZUSP 642);
http://dx.doi.org/10.1017/S0025315416001569
Fig. 4. Diadema ascensionis Mortensen, 1909: A– C, E –F (MZUSP 599); D, I (MZUSP 603); G (MZUSP 600); H (MZUSP 643). (A) Specimen from Trindade
Island photographed in situ, at 9.5 m depth; images of the carapace showing (B) aboral view, (C) ambulacrum, interambulacrum of a specimen (D) 39.2 mm
long and (E) 50 mm long; detail of the (F) apical disc, (G) Aristotle’s lantern, (H) buccal notches, and (I) auricle. Scale bars: B– E, 10 mm; F, 5 mm.
Fig. 5. Diadema ascensionis Mortensen, 1909: A (MZUSP 603); B – E (MZUSP 1003); F, H – K (MZUSP 599); G (MZUSP 600); L – N (MZUSP 590). SEM images –
(A) valve of triphyllous pedicellariae; (B – D) valves of narrow rostrate pedicellariae, and (E) detail of the head; (F) plate from peristomial membrane; tube feet
ossicles are (G) crown, (H– I) perforated plate, and (J– K) ‘doubled-sided hair-comb’; (L – M) longitudinal view and (N) cross section of the distal portion of a
spine. Scale bars: A– D, 100 mm; F, 500 mm; G – K, 100 mm; M, 500 mm.
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13 m, 17.4.2014, 3 spms 15 –40 mm long (MZUSP 1004). (perforated plate (230 –350 mm; Figure 5H, I), ‘doubled-
Farrilhões beach, 10.5 m, 9.7.2013, 1 spm 35 mm long sided hair-comb’ (90 – 200 mm; Figure 5J, K), and crown
(MZUSP 646). Secon, 10 m, 29.4.2014, 2 spms 25 –65 mm (270 –350 mm; Figure 5G)).
long (MZUSP 1002). Tartarugas beach, 9.5 m, 26.4.2012, 1 Colour black, denuded test and tubercles white; porous
spm 15 mm long (MZUSP 739); 2 m, 29.6.2013, 6 spms 15– region of madreporite and anal cone black; peristomial mem-
40 mm long (MZUSP 602, 603, 645). brane dirt white with purple-brownish ring around the
mouth; primary spines dark, varying from brown to purple
additional material (banded white/brownish spines in juveniles), oral spines black.
Fernando de Noronha archipelago, PE, Brazil (3848′ S
32823′ W) – Rata Island, 13.4.1999, 1 spm 73 mm long distribution
(MZUSP 604). Occurs alongside the Brazilian coast, Alagoas probably to Rio
de Janeiro (Brazil) (considering that all Brazilian specimens
comparative material examined were mistaken for D. antillarum); oceanic island distribution:
Diadema antillarum Philippi, 1845: Discovery Bay, Jamaica – Rocas Atoll, Fernando de Noronha, ASC, STH, TMV
1 spm 58 mm long (private collection, in Smith & Kroh, (Mortensen, 1940; Tommasi, 1966; Lima-Verde, 1969; Brito,
2011). Colon, Honduras (16800′ N 86830′ W) – 25.5.1975, 1 1971; Pawson, 1978; Gondim et al., 2013; present paper).
spm 80 mm long (UCMP 123422). New Providence Island, Bathymetric range: 0 –30 m (Mortensen, 1940).
The Bahamas (25815′ N 77815′ W) – 8.1956, 1 spm 80 mm
long (UCMP 123423). Diadema mexicanum A. Agassiz, biological notes
1863: Clipperton Island, France (Pacific territory) (10817′ N In TMV, specimens were gregarious, and usually found in reef
109813′ W) – 24.10.1956, 1 spm 55 mm long (UCMP crevices during the day and also moving around areas with
123424). Isla Montuosa, Panama (7829′ N 82815′ W) – 3 – high percentage of green algae cover.
7.5 m, 9.4.1968; 1 spm 55 mm long (UCMP 123425). Santa
Cruz Island, Galapagos Islands, Ecuador (0815′ S –0815′ N remarks
89845′ –90815′ W) – 0– 3 m, 24 –25.1.1964, 1 spm 75 mm Morphological differences among the Diadema species rival
long (UCMP 123426). Santa Elena Bay, Ecuador (2811′ S intraspecific differences. Although molecular data support
80856′ W) – 6 m, 17.4.1968, 1 spm 65 mm long (UCMP most of the species (e.g. Lessios et al., 2001), identifying speci-
123427). mens when DNA sequences are not available is very hard. We
assigned the specimens from TMV to D. ascensionis based on
description the absence of broad tridentate pedicellariae, rostrate pedicel-
Test circular in outline, low, diameter 10– 65 mm. Apical disc lariae (narrow tridentate) with broad tip, the number of solid
(Figure 4F) small, hemicyclic, ocular plates II and III exsert, wedges on the spines, and the interambulacra tuberculation
1– 2 tubercles may occur in ocular plates only. Hydropores pattern.
of madreporic plate enclosed in crescent moon-shaped area, The broad and narrow types of tridentate pedicellariae are
apparently constrained by arch-shaped depression. probably not homologous since D. mexicanum has both types.
Gonopores large, located on distal tip of triangular, wide Diadema antillarum has mainly the broad type, while
genital plates; periproct large with tiny plates peripherally. D. ascensionis apparently has only the narrow type. In add-
Ambulacra (Figure 4C) slightly elevated, narrow, trigeminate ition, the number of solid wedges in the spines was higher
plates composed of pore-pairs in single series, in distinct than the average for D. antillarum (18 solid wedges;
arcs of three and widening to four pore-pairs adorally. Coppard & Campbell, 2004). Regarding the tuberculation
Primary tubercles small, arranged into two parallel series pattern in the interambulacra, D. antillarum develops the
between columns of pore-pairs; secondary tubercles in third column of tubercles early on; specimens 60 mm in test
zigzag series along perradial suture. Interambulacra diameter already have three columns in each interambula-
(Figure D) wide, three series of primary tubercles in specimens crum and the inner column is not contiguous. We also ana-
over 70 mm in carapace diameter (two series in smaller speci- lysed specimens from Honduras, and they had the same
mens): outer series parallel to midline of plates; tubercles in pattern observed in our specimens. Molecular data have
middle series smaller, series shorter (only nearby ambitus); shown that D. ascensionis is nested within D. antillarum,
tubercles in inner series arranged in single columns. however, the analysis was based on only one gene and it is
Secondary tubercles scattered, although most concentrated not possible to infer if both species are indeed separated
on lateral edges of plates. Peristomial membrane with spine- without a proper analysis to the population level.
less ambulacral plates, thick and undulating buccal podia. The specimens from the BOI differ from D. africanum
Buccal notches pronounced (Figure 4H). Aulodont lantern Rodrı́guez, Hernández, Clemente & Coppard, 2013 by the
(Figure 4G). Auricles fused perradially, lobed edge shape of the rostrate pedicellariae and the interambulacra
(Figure 4I). Tubercles perforate, crenulate. tuberculation pattern (three columns in D. africanum), from
Primary spines long, brittle, with hollow lumen, 22 –25 D. mexicanum by the absence of broad tridentate pedicellar-
solid wedges; secondary spines smaller, thinner than iae, from D. palmeri Baker, 1967 by having a hemicyclic
primary spines. Spine shaft verticillate. Rostrate pedicellariae apical system (vs. monocyclic), from D. paucispinum
slender, blade smoothly serrated (double columns), narrow A. Agassiz, 1863 in having arch-shaped depression and only
blade tip (335– 870 mm; Figure 5B –E); tryphylous pedicellar- 0– 2 tubercles on genital plates (vs. 2– 6 tubercles), from
iae with long, broad blade (165– 265 mm; Figure 5A); clavi- D. savignyi (Audouin, 1829) in having 22– 25 solid wedges
form pedicellariae not found. Peristomial membrane ossicles on spines (vs. mean of 18), and from D. setosum (Leske,
showing as perforated plates of varied shapes (590– 1778) by the absence of long and slender tridentate
1850 mm; Figure 5F); tube feet ossicles of various shapes pedicellariae.
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Order CAMARODONTA Jackson, 1912 long (MZUSP 606). Tunel beach, 11.6.2012, 2 spms 13–
Family ECHINOMETRIDAE Gray, 1855 35 mm long (MZUSP 624).
Genus Echinometra Gray, 1825
Echinometra lucunter (Linnaeus, 1758) comparative material examined
(Figures 6 & 7, Table 1) Echinometra lucunter lucunter: Bolivar Province, Colombia
(108N 758W) – 4 spms 50 –54 mm long (CAS-IZ 91207).
Echinus lucunter Linnaeus, 1758, p. 665. ‘Gayanca stone reef’, Brazil – 3 spms 30 mm long (CAS-IZ
Echinometra lucunter – Mortensen, 1943b, pp. 357 –368, 15938). Jorge Grego Island, Angra dos Reis, RJ, Brazil
pls. 41 –44; Bernasconi, 1955a, pp. 62– 63, pl. 2; Brito, 1968, (coord) – 9.8.2009, 2 spms 44.5 –58 mm long (MZUSP
pp. 21– 22, pl. 10. 1290). Echinometra insularis H.L. Clark, 1912: Ecuador – 3
Echinometra subangularis – Rathbun, 1879, pp. 143 –144. spms 40 –45 mm long (CAS-IZ 91093). Echinometra
mathaei (Blainville, 1825): Midway Island, Pacific Ocean
(288N 1778W) – 2 spms 58 mm long (CAS-IZ 91179). The
examined material Philippines – 2 spms 45 mm long (CAS-IZ 189015). Upolu
Martin Vaz Island, ES, Brazil (20830′ S 29818′ W) – 13 m, Island, Western Samoa (138S 1718W) – 2 spms 39 mm long
26.6.2013, 1 spm 30 mm long (MZUSP 611); 13 m, (CAS-IZ 91184). Echinometra oblonga (Blainville, 1825):
23.7.2013, 3 spm 5– 39 mm long (MZUSP 618, 648). Islas Revillagigedo, Socorro Island, Mexico (188N 1108W) –
Trindade Island, ES, Brazil (20829′ – 20832′ S 29817′ – 2– 12.5.1925, 2 spms 32– 42 mm long (CAS-IZ 91202).
20820′ W) – Andrada beach, intertidal, 5– 21.7.2013, 6 spms Echinometra vanbrunti A. Agassiz1863: Gulf of California,
5– 30 mm long (MZUSP 605, 625); 10 m, 17.7.2013, 5 spms Mexico (228–248N 1098– 1128W) – Bahia Magdalena,
15 –25 mm long (MZUSP 620). Calheta Island, intertidal, 27.4.1916, 1 spm 45 mm long (CAS-IZ 91232); Cape San
11.6.2012, 4 spms 25– 30 mm long (MZUSP 614). Enseada Lucas, 2 spms 60 mm long (CAS-IZ 17104).
da Cachoeira, 10.5 m, 9.7.2012, 2 spms 35 –42 mm long
(MZUSP 622); 14 m, 9.7.2014, 1 spm 47 mm long (MZUSP description
621). Enseada de Orelhas, 15 m, 25.6.2012, 1 spm 17 mm Test subpentabonal or oval in outline, longer axis along ocular
long (MZUSP 660). Enseada Portuguesa, 12 m, 15 – plate I and genital plate 3, diameter 5– 47 mm. Apical disc
18.7.2013, 4 spms 25– 30 mm long (MZUSP 616, 617). (Figure 6F) monocyclic or hemicyclic, ocular plate V exsert
Ponta do Monumento, 12 m, 16.6.2012, 3 spms 10 –25 mm (rarely, ocular plate I also exsert). Plates bearing three or
long (MZUSP 649); 8 m, 2.4.2014, 3 spms 24 –37 mm long more tubercles, madreporic plates almost entirely covered
(MZUSP 1001). Farrilhões beach, 12.5 m, 31.1.2012, 1 spm by hydropores, periproctal plates large, few. Ambulacral
10 mm long (MZUSP 608). Secon, 9 m, 12.5.2014, 9 spms plating (Figure 6D) polygeminate, five to six (rarely seven)
15 –35 mm long (MZUSP 1000). Tartarugas beach, 11 m, 1 pore-pairs to a compound plate, arranged in arcs. Poriferous
spm 5 mm long (MZUSP 607); 9.5 m, 26.6.2012, 4 spms zone a bit larger than perradial zone. Primary tubercles
15 –25 mm long (MZUSP 609, 740); 14 m, 28.6.2012, 4 small, arranged into two parallel series between columns of
spms 10– 25 mm long (MZUSP 613); intertidal, 15.7.2012, 2 pore-pairs; secondary tubercles in zigzag series along perradial
spms 28.5– 37 mm long (MZUSP 619); 2 m, 29.6.2013, 7 suture. Interambulacra (Figure 6E) with two parallel series of
spms 20 –25 mm long (MZUSP 623); 12 m, 3.7.2013, 1 spm large primary tubercles arranged in midline of plates, one
35 mm long (MZUSP 612); intertidal, 5.7.2013, 4 spms 30 – series of smaller primary tubercles near each ambulacrum,
40 mm long (MZUSP 615); 14.5 m, 11.7.2013, 1 spm 10 mm one in zigzag along the interradial suture. Peristomial
Fig. 6. Echinometra lucunter (Linnaeus, 1758): A –C, F (MZUSP 619); D– E, G –H (MZUSP 616); I (MZUSP 1290). (A) Specimen from Trindade Island
photographed in situ, between 0– 1 m depth; images of the carapace showing (B) aboral view, (C) oral view, (D) ambulacrum, and (E) interambulacrum;
detail of the (F) apical disc, (G) Aristotle’s lantern, and auricle of (H) specimen from the BOI, and (I) from the mainland. Scale bars: B – E, 10 mm; F, 5 mm.
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Fig. 7. Echinometra lucunter (Linnaeus, 1758): A, E– L (MZUSP 623); B – D (MZUSP 616). SEM images – (A) valve of triphyllous pedicellaria; (B –C) valves of
fanged globiferous pedicellariae, and (D) detail of the head; (E – H) valves of ophicephalous pedicellariae; (I) plate from peristomial membrane; (J) crown from tube
feet; and (K – L) detail of a spine. Scale bars: A, 50 mm; B, 200 mm; C, 150 mm; E –H, 100 mm; I, 200 mm; J – K, 100 mm.
membrane with few dispersed plates. Buccal notches shallow. with irregularly denticulate margins (375– 600 mm;
Camarodont lantern (Figure 6G). Auricles fused perradially, Figure 7E– H); triphyllous pedicellariae broad, flat, margin
distal portion broad, lobed edge (Figure 6H). Tubercles imper- smooth (100 –110 mm; Figure 7A); tridentate pedicellariae
forate, non-crenulate. not found. Ossicles showing as perforated plates in peristomial
Primary spines all cylindrical, medium-sized. Spine shaft membrane (800– 830 mm; Figure 7I), crowns in tube feet
with tiny denticulations arranged into transversal groups of (330 –585 mm; Figure 7J), C-shaped ossicles in both (17 –
four teeth, about 15 mm apart from the next group. Fanged 43 mm).
globiferous pedicellariae slender, margins smooth, short Peristomial membrane brownish with dark green rim
lateral unpaired tooth on blade, long terminal tooth (730– around the mouth, tube feet green-brownish with pale tips,
1050 mm; Figure 7B –D); ophicephalous pedicellariae robust denuded test beige; primary spines dark purple orally and
Table 1. Morphological traits of the test used to distinguish the Echinometra species. Numbers in parentheses indicate rare amounts of pores.
E. insularis E. lucunter E. mathaei E. oblonga E. vanbrunti E. viridis

H.L. Clark (Linnaeus) (Blainville) (Blainville) A. Agassiz A. Agassiz
Plates in peristomial membrane Scattered Scattered No/rare No/rare Scattered Scattered

Pores in ambulacral arc 5–6 5–6(7) 4–5 4 –5 (6)–7–(8) 5(6)
Strong concavity orally No No Yes No No No
Tag in auricle Inconspicuous Conspicuous Mild Conspicuous Inconspicuous Mild
Test shape Oval Oval Oval Oval Circular Varied
Geographic distribution Pacific islands Atlantic Indo-West Pacific Indian/Pacific East Pacific Atlantic
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black aborally, secondary spines greenish to purple (from base Pseudoboletia maculata Troschel, 1869
to tip); apical plates beige with brownish spots. (Figures 8 & 9)
Pseudoboletia maculata Troschel, 1869, p. 96.
distribution Pseudoboletia occidentalis H.L. Clark, 1921, pp. 115 – 118,
From North Carolina (USA) to Santa Catarina (Brazil) pl. 2.
(western Atlantic); from Senegal to Angola (eastern Pseudoboletia occidentalis – Mortensen, 1943a, pp.
Atlantic); oceanic island distribution: Bermuda, Cape Verde, 538 –540.
São Tomé, Fernando de Noronha, ASC, STH, Abrolhos, Pseudoboletia maculata maculata – Pawson, 1978, pp.
TMV (Rathbun, 1879; Agassiz, 1881; Clark, 1898; 17 –20.
Mortensen, 1943b; Pawson, 1978; Hendler et al., 1995;
present paper). Bathymetric range: 0– 45 m (Serafy, 1979). examined material
Trindade Island, ES, Brazil (20829′ – 20830′ S 29818′ –
biological notes 29820′ W) – Enseada de Orelhas, 15 m, 24.10.2014, 2 spms
In TMV, specimens were gregarious and found in rocky 16 –23 mm long (MZUSP 1123). Enseada Portuguesa, 15 m,
bottoms. 10.7.2012, 1 spm 35 mm long (MZUSP 1026). Secon, 9 m,
12.5.2014, 1 spm 55 mm long (MZUSP 1027).
remarks
The specimens reported herein are very similar to the coastal
comparative material examined
Pseudoboletia atlantica: Prosperous Bay, STH – 27 m,
populations, but the subpentagonal test shape is unusual and
17.12.1945 (NHM-UK 1945.12.17.8).
the shape of the auricle is different (Figure 6I). The subspecies
Pseudoboletia maculata: Guam, USA – 2 m, 3.1992, 1 spm
Echinometra lucunter polypora Pawson, 1978 was described
73 mm long (CAS-IZ 81252). Lubang Islands, The Philippines
from AST populations and differs from Echinometra lucunter
(13850′ N 120805′ E) – 1 –18 m, 23.5.2014, 1 spm 60 mm long
lucunter by the presence of seven pore-pairs in the arcs (vs.
(CAS-IZ 203488). Pseudoboletia indiana (Michelin, 1862):
six/five) and the greenish or white colour of the adapical
Hawaii, USA – 2 spms 65 mm long (CAS-IZ 91174, 92324);
region of the test. However, the number or pore-pairs in the
Oahu, 7.5 m, 2 spm 50– 60 mm long (CAS-IZ 91339).
arcs are very variable in the genus Echinometra and molecular
analysis with the COI gene does not support the separation of
E. lucunter into subspecies (McCartney et al., 2000). description
The differences between E. lucunter and the other Test circular in outline, diameter 35– 55 mm. Apical disc
Echinometra species are summarized in Table 1. We consid- small, hemicyclic (Figure 8F), ocular plates II, III and IV
ered E. mathaei (Blainville, 1825) and E. oblonga (Blainville, exsert. Madreporic plate twice wider than other genital
1825) as separate species because of morphological differences plates; 2 –3 tubercles on proximal edge of plates; periproctal
regarding their auricle and test shape. This separation has also plates covering periproct, sometimes bearing tubercle.
been supported by molecular data (see McCartney et al., 2000; Ambulacra polygeminate plates composed of arc with four
Landry et al., 2003). pore-pairs (Figure 8D). Primary tubercles arranged into two
parallel series among pore-pairs; tubercles from outer series
Family TOXOPNEUSTIDAE Troschel, 1872 absent in aboral region. Interambulacra (Figure 8E) with
Genus Pseudoboletia Troschel, 1869 four parallel series of primary tubercles, innermost series
Fig. 8. Pseudoboletia maculata Troschel, 1869: A (MZUSP 1027); B – H (MZUSP 1026). (A) Specimen fixed in ethanol; images of the carapace showing (B) aboral
view, (C) oral view, (D) ambulacrum, and (E) interambulacrum; detail of the (F) apical disc, (G) Aristotle’s lantern, and (H) auricle. Scale bars: B – E, 10 mm; F,
5 mm.
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Fig. 9. Pseudoboletia maculata Troschel, 1869 (MZUSP 1027). SEM images – (A– B) valves of triphyllous pedicellariae; (C – D) valves of ophicephalous
pedicellariae; (E –G) valves of fanged globiferous pedicellariae; (H) C-shaped ossicle; (I) rosette-like plate from peristomial membrane; (J) crown, (K) plate,
and (L) C-shaped ossicle from tube feet ([M] detail of the tip of the C-shaped ossicles depicted in (L)), and (K –L) detail of a spine. Scale bars: A– B, 50 mm;
C– D, 100 mm; E, 200 mm; F, 100 mm; G, 50 mm; H, 5 mm; I– K, 100 mm; L, 20 mm; O, 250 mm.
smaller, present only at ambitus; secondary tubercles scat- paper). Bathymetric range: 9 –183 m (Mortensen, 1943a;
tered. Peristomial membrane with dispersed small plates, present paper).
five pairs of buccal plates with spines and pedicellariae. Tip
of oral tube feet crenulated. Buccal notches pronounced.
Camarodont lantern (Figure 8G). Auricles widened at edges
and fused perradially (Figure 8H). Tubercles imperforate, remarks
non-crenulate. Pawson (1978) did an extensive revision of this genus and
All spines cylindrical, small. Fanged globiferous pedicellar- in this paper we adopt his suggestion that Pseudoboletia
iae slender, margins smooth, perforations for insertion of occidentalis Clark, 1921, is a junior synonym of P. maculata.
adductor muscles widespread on proximal region of blade, Pseudoboletia maculata differs from Pseudoboletia atlantica
terminal tooth present (300– 910 mm; Figure 9E –G); ophice- H.L. Clark, 1912 by the presence of four pore-pairs in
phalous pedicellariae robust, broad, margins denticulate, each arc (vs. five) and by the colouration of the test (sparse
slightly serrate (430– 670 mm; Figure 9C, D); triphyllous flat, dark blotches vs. dark stripes from apical disc to peristome),
margin apparently smooth, narrow neck (130– 240 mm; and from P. indiana in having dark spots on the test (vs.
Figure 9A, B); tridentate pedicellariae not found. Rosette- uniform colour). Although known worldwide, this is the
like oval ossicles in the peristomial membrane (300– first record of P. maculata in the South Atlantic, and the
475 mm; Figure 9I); tube feet with crowns (205– 555 mm; first record of the species for Brazil. This species can easily
Figure 9J), perforated plates (265 –390 mm; Figure 9K) and be mistaken for other toxopneustids, such as Tripneustes
thin C-shaped ossicles with bifurcated tips (40 mm; only one ventricosus and Lytechinus variegatus; therefore, we would
ossicle found; Figure 9H); C-shaped ossicles on all pedicellar- not be surprised if it occurs elsewhere along the Brazilian
iae (11 –30 mm). coast.
Denuded test creamy, brown blotches on aboral region;
primary spines with brown/green base, white tips; apical Genus Tripneustes L. Agassiz, 1841
disc plates with brown ring; tube feet and pedicellariae light Tripneustes ventricosus (Lamarck, 1816)
pink. (Figures 10 & 11)
Echinus ventricosus Lamarck, 1816, p. 44.
distribution Tripneustes ventricosus – Agassiz & Desor, 1846, p. 363;
Venezuela, Barbados, Trindade Islands and Indo-West Pacific Mortensen, 1943a, pp. 490 – 498, pls. 33, 34, 37, 38, 56;
(Smith & Kroh, 2011; Alvarado & Solı́s-Marı́n, 2013; present Tommasi, 1962, p. 59, figures 1 & 2.
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Fig. 10. Tripneustes ventricosus (Lamarck, 1816): A– F, H (MZUSP 632); G (MZUSP 633). (A) Specimen from Trindade Island photographed in situ, between 0–
1 m depth; images of the carapace showing (B) aboral view, (C) oral view, (D) ambulacrum, and (E) interambulacrum; detail of the (F) apical disc, (G) Aristotle’s
lantern, and (H) auricle. Scale bars: B – E, 10 mm; F, 5 mm.
Fig. 11. Tripneustes ventricosus (Lamarck, 1816): A, H – I (MZUSP 1020); B – G, J – L (MZUSP 632). SEM images – (A) valve of triphyllous pedicellaria; (B – D)
valves of ophicephalous pedicellariae; (E – F) valves of fanged globiferous pedicellariae; (G –H) valves of tridentate pedicellariae; (I) rosette-like plate from
peristomial membrane; (J) crown from tube feet; and (K – L) detail of a spine. Scale bars: A, 100 mm; B –H, 200 mm; I– J, 150 mm, L, 100 mm.
http://dx.doi.org/10.1017/S0025315416001569
examined material long, widened at edges, fused perradially (Figure 10H).

Trindade Island, ES, Brazil (20829′ – 20831′ S 29818′ – Tubercles imperforate, non-crenulate.
29820′ W) – Andrada beach, intertidal, 21.7.2013, 1 spm All spines cylindrical, small; spine shaft with tiny vertical
40 mm long (MZUSP 632). Cabritas beach, 9 m, 14.6.2012, projections arranged in horizontal groups of 2– 4, about
2 spms 75 –80 mm long (MZUSP 630); 7.5 m, 28.4.2014, 1 20 mm apart from next group. Fanged globiferous pedicellar-
spm 75 mm long (MZUSP 998). Calheta beach, 16 m, iae slender, margins smooth, perforations for insertion of
29.4.2014, 2 spms 40– 55 mm long (MZUSP 991); 17 m, adductor muscles only in half innermost part of proximal
8.5.2014, 2 spms 30 mm long (MZUSP 995); 16 m, region, a terminal tooth present (430 –465 mm; Figure 11E,
20.5.2014, 1 spm 20 mm long (MZUSP 1017). Calheta F); ophicephalous pedicellariae robust, margins serrate, den-
Island, 10 m, 24.7.2011, 1 spm 34 mm long (MZUSP 628); ticulate (240 –655 mm; Figure 11B –D); tridentate slender,
12 m, 18.6.2012, 1 spm 60 mm long (MZUSP 629); 4 m, distal cap denticulate, some spines widely separated in
14.7.2013, 1 spm 60 mm long (MZUSP 633). Enseada da middle of valve (370 –750 mm; Figure 11G, H); triphyllous
Cachoeira, 14 m, 9.7.2012, 1 spm 23.5 mm long (MZUSP pedicellariae broad, flat, upper margin finely serrate (160–
738). Enseada de Orelhas, 14 m, 30.6.2012, 2 spms 35– 205 mm; Figure 11A). Dumbbell-shaped ossicles on all pedi-
51 mm long (MZUSP 637). 14 m, 6.7.2013, 5 spms 15– cellariae, larger ones curved (9 – 20 mm), tube feet with
80 mm long (MZUSP 635, 674); 12 m, 18.4.2014, 5 spms crown (340 –425 mm; Figure 11J), peristomial membrane
30 –70 mm long (MZUSP 999); 14 m, 9.5.2014, 1 spm with rosette-like round ossicles (255– 315 mm; Figure 11I).
77 mm long (MZUSP 1014); 10 m, 21.5.2014, 1 spm 50 mm Colour dark grey, denuded test white-brownish, tubercles
long (MZUSP 993). Enseada Portuguesa, 14 m, 10.7.2012, 3 pale; primary spines light grey or pale green with brownish
spms 30– 40 mm long (MZUSP 627); 12 m, 11.7.2012, 4 tips, secondary spines grey-purplish; apical plates light grey
spms 35– 55 mm long (MZUSP 631); 12 m, 15.7.2013, 5 with dark grey-purplish spots around the gonopores; tube
spms 30 –70 mm long (MZUSP 636). Farol beach, 13 m, feet and tridentate pedicellariae yellowish, ophicephalous
22.4.2014, 5 spms 50 –60 mm long (MZUSP 996); 13 m, 8 – pedicellariae dark brown.
20.5.2014, 12 spms 30 –60 mm long (MZUSP 994, 1020,
1021). Farrilhões beach, 10 m, 9.7.2013, 2 spms 20 –90 mm
distribution
long (MZUSP 626, 634); 14 m, 5.5.2014, 2 spms 10 –15 mm
Occurs from Florida (USA) to São Paulo (Brazil); oceanic
long (MZUSP 1015). Lixo beach, 14 m, 22.4.2014, 1 spm
island distribution: Fernando de Noronha, ASC (?), STH (?),
60 mm long (MZUSP 1012). Ponta do Monumento, 8 m,
and Trindade Islands (Mortensen, 1943a; Tommasi, 1962;
2.4.2014, 1 spm 55 mm long (MZUSP 997).
Lima-Verde, 1969; Pawson, 1978). Bathymetric range: 0–
55 m (Serafy, 1979).
comparative material examined
Tripneustes ventricosus: Arraial do Cabo, RJ, Brazil (228S biological notes
428W) – 1– 2 m, 15.10.1998, 1 spm 112.39 mm long (CAS- In TMV, specimens were usually solitary and found in rocky
IZ 116108). Puerto Rico – 1934, 1 spm 71 mm long (CAS- bottoms.
IZ 91159). Tripneustes depressus A. Agassiz, 1863: Isla
Danzante, Baja California, Mexico (25847′ N 111815′ W) –
9 m, 2 spms 57 –78 mm long (CAS-IZ 101402). Islas remarks
Revillagigedo, Clarion Island, Mexico (188N 1148W) – The specimens described herein have no apparent morpho-
19.3.1993, 1 spm 89 mm long (CAS-IZ 103105). Tripneustes logical variations from specimens from mainland or other
gratilla (Linnaeus, 1758): Hawaii, USA – Hawaii, 2 spms islands. Lessios et al. (2003) found three clades within
60 –62 mm long (CAS-IZ 108120); Oahu, 1 spm 111 mm Tripneustes ventricosus (Caribbean, Brazil and São Tomé)
long (CAS-IZ 91164). and it would be interesting to sequence molecular data from
mid-Atlantic specimens to access the connectivity between
these regions.
description Only three extant Tripneustes species are recognized as
Test circular in outline, diameter 10– 90 mm. Apical disc valid worldwide, although the molecular data presented by
(Figure 10F) small, hemicyclic, ocular plates II and III Lessios et al. (2003) did not support the Indo-Pacific
exsert; ocular plate IV may be exsert or not, touching peri- species. Morphologically, T. ventricosus can be distinguished
proct only slightly through an extension between the genital from T. gratilla in being densely covered with tubercles, espe-
plates. Madreporic plate enlarged, 2– 3 tubercles on proximal cially in the interambulacra; and from T. gratilla and
edge; periproctal plates covering periproct, sometimes bearing T. depressus in the reduced/absence of plates in the peristomial
a tubercle. Genital plates usually bearing two tubercles, gono- membrane.
pores large. Ambulacra (Figure 10D) with trigeminate plates Order CLYPEASTEROIDA A. Agassiz, 1872
composed of three columns of pore-pairs; inner column Family MELLITIDAE Stefanini, 1912
almost straight, pores in middle column arranged diagonally. Genus Leodia Gray, 1851
Primary tubercles arranged in parallel series between columns Leodia sexiesperforata (Leske, 1778)
of pore-pairs; tubercles in inner series larger than in outer (Figures 12 & 13)
series. Secondary tubercles scattered. Interambulacra
(Figure 10E) with two parallel series of primary tubercles Echinodiscus fexiesperforatus Leske, 1778, p. 199.
arranged in midline of plates and scattered tubercles of Mellita sexforis – A. Agassiz, 1972 –1974, p. 141; Rathbun,
various sizes. Peristomial membrane naked. Buccal notches 1879, p. 144.
pronounced. Camarodont lantern (Figure 10G). Auricles Mellita sexiesperforata – Bernasconi, 1947, p. 104.
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Fig. 12. Leodia sexiesperforata (Leske, 1778) (MZUSP 1070). (A) Specimen from Trindade Island photographed in situ, between 16 –17 m depth; images of the
carapace showing (B) aboral view and (C) oral view; detail of the (D) apical disc, (E) peristome, and (F) periproct; Aristotle’s lantern (G) oral and (H) aboral side,
and (I – J) detail of individual pyramids. Scale bars: B– C, 20 mm; D – F, 2 mm; G – H, 5 mm; I –J, 1 mm.
Leodia sexiesperforata – Mortensen, 1948b, pp. 429 – 432, comparative material examined
pls. 58, 61, 72; Serafy, 1979, pp. 74 –75, figure 32. Leodia sexiesperforata: Boipeba, BA, Brazil (13836′ S 38853′ W)
– Bainema beach, 17 –31.1.2015, 17 spms 55 –83.5 mm long
(MZUSP 1198).
examined material
Trindade Island, ES, Brazil (20829′ – 20830′ S 29818′ – description
29820′ W) – Calheta beach, 16– 17 m, 20– 24.10.2014, 31 Test pentagonal in outline, posterior edge sharp, flattened,
spms 39 –60 mm long (MZUSP 1070, 1126); 17 m, 25 –70 mm long. Six long, narrow lunules; anal lunule’s prox-
4.11.2014, 48 spms 25– 70 mm long (MZUSP 1124, 1125). imal margin right after end of petals. Apical disc monobasal
Enseada de Orelhas, 13.9 m, 6.I.2014, 5 spms 47 –65 mm (Figure 12D), four gonopores (G5 missing), hydropores
long (MZUSP 1190). Racha Island, 25 m, 12.11.2014, 4 spread throughout apical disc, ocular plates small. Petals
spms 60 –65 mm long (MZUSP 1191). broad, short (1/3 of the ambulacrum), petal III longest. Both
Fig. 13. Leodia sexiesperforata (Leske, 1778) (MZUSP 1070). Optical microscope images of the (A –D) bidentate pedicellariae, of the spines from the (E) edge of
the test, and from the (F) aboral region; and (G) SEM image of the tip of a spine depicted in (F). Scale bars: A– D, 10 mm; G, 100 mm.
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columns of pore-pairs bowed, closing at end; poriferous zone biological notes

broad, fully tuberculated, up to seven primary tubercles per In TMV, specimens were found in soft bottoms (medium-
row. One to two extra pore-pairs beyond petals. Peristome grained sand).
small, circular (Figure 12E). Clypeasteroid lantern with prom-
inent, strongly lamellar pyramid wings, supra-alveolar pro-
cesses located far from tip (Figure 12G – J). Periproct remarks
(Figure 12F) small, longer than wide, on first interambulacral Leodia sexiesperforata is the only extant species of the genus.
plate, slightly closer to peristome than to anal lunule. Food Some intraspecific variations to coastal Brazilian specimens
grooves bifurcate near peristome, broadens, and narrow include the shape of the test (subpentagonal to round in
down again towards posterior margin of test, bifurcating coastal specimens) and the location of the anal lunule
again. Oral tube feet extend into interambulacra. Test fully (placed more proximally, usually between posterior petals,
tuberculated, tubercles larger on oral ambulacrum than on in coastal specimens).
interambulacrum. Tubercles perforate, crenulate.
Order SPATANGOIDA L. Agassiz, 1840
Spines thin, small, mainly of three types: straight spines
Family BRISSIDAE Gray, 1855
of varying sizes orally (largest between peristome and peri-
Genus Plagiobrissus Pomel, 1883
proct), curved spines on peristome and at ambitus, swab-
Plagiobrissus grandis (Gmelin, 1791)
shaped spines aborally. Bidentate pedicellariae with broad
(Figures 14 & 15)
serrate blade, short base on aboral region (32 –38 mm;
Figure 13A –D). Echinus grandis Gmelin, 1791, p. 3200.
Specimen blackish in vivo, test pale in ethanol; spines pale. Metalia pectoralis – A. Agassiz, 1881, p. 199.
Plagiobrissus grandis – Mortensen, 1951, pp. 496 – 368, pls.
40 –41, 63; Serafy, 1979, pp. 98 –99, figure 43.
distribution
From South Carolina (USA) to Rio La Plata (Argentina); examined material
oceanic island distribution: Bermuda Island and Trindade Trindade Island, ES, Brazil (20830′ S 29818′ W) – Calheta
Islands (H.L. Clark, 1898; Bernasconi, 1947; Mortensen, beach, 16 m, 24.10.2014, 9 spms 58 –76 mm long (MZUSP
1948b; present paper). Bathymetric range: 0– 490 m (A. 1084); 17 m, 4.11.2014, 7 spms 60 –100 mm long (MZUSP
Agassiz, 1972 –1974; Serafy, 1979). 1117, 1289).
Fig. 14. Plagiobrissus grandis (Gmelin, 1791): A (MZUSP 1084); B, G (MZUSP 1289); C –F (MZUSP 1117). (A) Specimen from Trindade Island photographed in
situ, at 16 m depth; images of the carapace showing (B) aboral view, (C) oral view and (D) lateral view of the body; and detail of the (E) apical disc, (F) peristome
and (G) periproct. Scale bars: B – D, 20 mm; E– G, 5 mm.
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Fig. 15. Plagiobrissus grandis (Gmelin, 1791) (MZUSP 1289). SEM images – (A – B) valves of triphyllous pedicellariae; (C) valve of rostrate pedicellaria; (D– E)
valves of tridentate pedicellariae; (F– G) valves of long globiferous pedicellariae; (H – I) rods from tube feet; and (K – L) detail of a spine. Scale bars: A– B, 30 mm; C–
E, 50 mm; F– G, 100 mm; H – J, 20 mm; K, 200 mm.
comparative material examined narrow, serrate, base broad (165– 270 mm; Figure 16D, E).
Plagiobrissus grandis: The Bahamas (24802′ –25801′ N 74829′ – Triphyllous pedicellariae near peristome, blade long, serrate
77824′ W) – Nassau, 1 spm 195 mm long (NHM-UK margin, narrow neck, base short (100– 150 mm; Figure 15A,
87.4.25.6); San Salvador Island, 3 m, 23.10.1983, 1 spm B). Regular globiferous and ophicephalous pedicellariae not
87 mm long (USNM E32634). Florida, USA – 1 spm found. Phyllopodia ossicles showing as thin rods, straight or
148 mm long (USNM E13737); 5–11 m, 1964, 1 spm slightly curved, sometimes bifurcated at middle or at tips
87 mm long (USNM E36776). Panama (9824′ N 79851′ W) – (35 – 100 mm); flat perforated plates (50 –95 mm).
Galeta Island, 10.7 m, 30.9.1980, 1 spm 97 mm long (USNM Respiratory tube feet with rods only (40 – 140 mm;
E36447). Figure 15H, I).
Colour in vivo: aboral surface of specimen dark brown with
description white patches, oral surface with pale spines, phyllopodia red,
Test oval in outline, flattened, 58 –90 mm in length. Frontal long globiferous pedicellariae dark brown. Test white in
groove slight but present, more developed in larger specimens. ethanol.
Apical disc ethmolytic (Figure 14E), four gonopores and
genital plate 2 separates oculars III and IV. Petals almost
180 degrees in younger specimens, angle diminishing in
largest specimen. Paired petals deeply sunk; anterior petal
slightly sunk. First half of inner columns of paired petals
less developed than outer columns; distal plates in petals
occluded. Interporiferous zone much narrower than porifer-
ous zones. Peripetalous fasciole indented in interambulacral
2 and 3, subanal fasciole shield-shaped, anal and subanal fas-
cioles confluent. Primary tubercles inside peripetalous fasciole
developed. Periproct large, tear-shaped (Figure 14G), trun-
cate. Peristome broad, kidney shaped (Figure 14F); labrum
short, in contact with both sternal plates. Sternal plates paral-
lel, very long, narrow, with an elevated midline, completely
tuberculated. Phyllodes well developed, with four to five
pores in each half ambulacrum. Tubercles perforate, crenulate.
Spines thin; large inside peripetalous fasciole, around peri-
proct and on oral region (except on plastron); small elsewhere. Fig. 16. Chiridota rotifera (Pourtalès, 1851): A, C (MZUSP 944); B (MZUSP
Long globiferous pedicellariae triangular, robust, with dense 946); D (MZUSP 973). Specimen fixed in ethanol: (A) lateral view of the
stereom (510– 700 mm; Figure 15F, G), located near periproct. body, (B) miliar granules (indicated by the arrow) on the posterior region of
the body, (C) detail of the anterior region (arrow indicates warts
Rostrate pedicellariae with short denticulate blade, long and (aggregations of wheel) in tentacles and body wall), and (D) detail of
narrow neck, smooth margins, broad base (350– 400 mm; tentacles and calcareous ring (indicated by the arrow). Scale bars: A– B,
Figure 15C). Tridentate pedicellariae near petals, blade 5 mm; D, 1 mm.
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distribution Class HOLOTHUROIDEA

Occurs from Florida (USA) to São Paulo (Brazil); oceanic Order APODIDA Brandt, 1835
island distribution: Trindade Island (Mortensen, 1951; Brito, Family CHIRIDOTIDAE Östergren, 1898
1962; present paper). Bathymetric range: 1 –210 m (Serafy, Genus Chiridota Eschscholtz, 1829
1979). Chiridota rotifera (Pourtalès, 1851)
(Figures 16 & 17)
biological notes Synapta rotifera Pourtalès, 1851, p. 15.

In TMV, specimens were found in soft bottoms, usually com- Chiridota rotifera – Ludwig, 1881, pp. 116 –117.
pletely buried in medium-grained sand. Toxodora ferruginea – Verrill, 1882, p. 220; Clark, 1907,
p. 126; Deichmann, 1930, p. 213.
Chiridota ferruginea – Deichmann, 1940, pp. 228 –229.
remarks
The specimens from the BOI differ from the Caribbean speci- examined material
mens in having plates with similar width in each column of Trindade Island, ES, Brazil (20829′ – 20831′ S 29818′ –
the anterior interambulacral (when comparing same-sized 20820′ W) – Enseada de Orelhas, 14 m, 9.5.2014, 2 spms 1–
specimens from both regions). The inner column of the 2 cm long (MZUSP 945). Farol beach, 13 m, 8.5.2014, 2
Caribbean specimens is much narrower than the outer spms 0.5 –2 cm long (MZUSP 942); 12 m, 8.4.2014, 3 spms
column. Also, the tubercles are slightly larger on the speci- 1– 2 cm long (MZUSP 937); 13 m, 8–22.4.2014, 4 spms 2–
mens from the Caribbean. This variation is very slight and 3 cm long (MZUSP 938 – 941). Farrilhões beach, 12 m,
does not justify description of a new species. 4.4.2014, 1 spm 1 cm long (MZUSP 946). Secon, 12 m,
Plagiobrissus grandis differs from P. africanus (Verrill, 12.4.2014, 1 spm 3.5 cm long (MZUSP 944).
1871) by the absence of large primary tubercles close to the
frontal ambulacrum (there are medium-sized tubercles in description
the specimens from the BOI), and by the shape of the peri- Body vermiform (Figure 16A), maximum length 3.5 cm. Body
proct, which is obliquely truncate and not visible in oral wall transparent, miliar granules posteriorly (Figure 16B).
view (vs. vertically truncate and visible in oral view); it Twelve white, digitate (fingerlike) tentacles subdivided into
differs from Plagiobrissus (Rhabdobrissus) costae (Gasco, five pairs of digits (Figure 16C). Calcareous ring with 12
1876), P. (R.) jullieni (Cotteau, 1889) and P. (R.) pacificus pieces, radial pieces perforated on top for insertion of radial
(Clark, 1940) in having sunk petals (vs. flush petals). nerves (Figure 16D). Ten to 20 Polian vesicles.
Fig. 17. Chiridota rotifera (Pourtalès, 1851): (MZUSP 944). SEM and optical microscope images of the ossicles – wheels from body wall in (A) ventral, (B) dorsal
and (C) lateral view; (D) early developmental stage of a wheel; (E) C-shaped ossicles from body wall; (F – H) rods from tentacles. Scale bars: A –D, 20 mm; E, 10 mm;
F– H, 50 mm.
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Body wall wheels (30 –100 mm; Figure 17A – D) with six differences were observed between coastal and oceanic
spokes each, inner rim serrated, margins smooth; C-shaped insular specimens of C. rotifera.
ossicles (20–60 mm; Figure 17E) with few small projections
mainly at tips. Tentacles with wheels as in body wall (30 – Family SYNAPTIDAE Burmeister, 1837
70 mm); slightly curved rods (30–90 mm; Figure 17F) with Genus Euapta Östergren, 1898
some projections at tips (rarely along the body); bi or tri- Euapta lappa (Müller, 1850)
radiated, curved rods, branched at ends rods (40 –60 mm; (Figures 18 & 19)
Figure 17G, H). Synapta lappa Müller, 1850, pp. 134 – 135.
Synapta polii Ludwig, 1875, p. 80.
distribution Euapta lappa – Clark, 1907, p. 73, pl. 4; Clark, 1924, pp.
Occurs from Florida to Brazil (to São Paulo); oceanic island 464 –465, pl. 1; Heding, 1928, pp. 136 – 137, pl. 8.
distribution: Trindade Island (Deichmann, 1930; Tommasi, Euapta tobagoensis – Heding, 1928, pp. 133 –136, pl. 8.
1969; Hendler et al., 1995; present paper). Bathymetric
range: from the intertidal down to 360 m (Alvarado & Solı́s- examined material
Marı́n, 2013). Trindade Island, ES, Brazil (20829′ S 29819′ W) – Enseada
Portuguesa, 14 m, 10.7.2012, 12 spms 3 –15 cm long
biological notes (MZUSP 289, 982); 12 m, 15.7.2013, 10 spms 3– 12 cm long
In TMV specimens found in association with calcareous algae. (MZUSP 317, 318). Farol beach, 13 m, 8– 17.4.2014, 26
spms 1.5–25 cm long (MZUSP 976, 980, 981); 14 m, 22–
remarks 27.4.2014, 2 spms 13 –20 cm long (MZUSP 977, 979); 13 m,
Eight species of Chiridota are known from the Atlantic Ocean, 8.5.2014, 3 spms 6–11 cm long (MZUSP 978).
of which only C. rotifera, C. marenzelleri Perrier, 1904 and C.
pisanii Ludwig, 1887 have been recorded from the South
Atlantic so far.The tentacle morphology is the main character comparative material examined
to distinguish C. rotifera from its Atlantic congeners: C. rotif- Euapta lappa: Bacia do Espı́rito Santo, ES, Brazil, (20826′ S
era differs from C. peloria Deichmann, 1930, C. ferruginea 39844′ W) – 1350 m, 9.2010, 2 spms 0.5 – 1 cm long
(Verrill, 1882), C. laevis (Fabricius, 1780) and C. heheva (MZUSP 290). Great Sound, Long Island, Bermudas – 2
(Pawson & Vance, 2004) in the numbers of digits (10 vs. 14, spms 5– 6 cm long (USNM 25001). Euapta godeffroyi
10 –12, 4– 6, 20, respectively), and can be separated from (Semper, 1868): Baja California, Mexico – intertidal,
C. hydrothermica (Smirnov & Gebruk, 2000) in having 11.1.1959, 1 spm 8 long (CAS 192451); 17.12.1960, 3 spms
finger-like tentacles (vs. lobe-like). Regarding the South 10 cm long (CAS 105518); 3.1889, 1 spm 7 cm long (USNM
Atlantic species, C. rotifera differs from C. pisanii in the E 2437); 15 m, 30.4.1888, 1 spm 6 cm long (USNM E 2431).
number of Polian vesicles (7 vs. 10 –12), and differs from C.
marenzelleri by the presence of stellate ossicles. Clark (1907) description
recommended a thorough comparison between C. rotifera Body vermiform (Figure 18A, B) with brown, beige stripes,
and C. ferruginea as both species share similar ossicles and maximum length 25 cm. 15 pale pinnate tentacles
overlap in the number of tentacles digits. No morphological (Figure 18C). Calcareous ring (Figure 18D) with 15 pieces
Fig. 18. Euapta lappa (Müller, 1850) (MZUSP 289). (A) Specimen from Trindade Island photographed in situ, at 14 m depth, and (B) fixed in ethanol; (C) detail of
pinnate tentacles; (D) calcareous ring, arrow indicates place for insertion of radial nerves. Scale bars: B, 5 cm; E, 2 mm.
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Fig. 19. Euapta lappa (Müller, 1850) (MZUSP 289). SEM images of the ossicles – anchors from body wall in (A) dorsal and (B) ventral view, (C) detail of minute
tubercles on anchor vertex, and (D) detail of branched anchor stock; anchor plates from body wall in (E – F) dorsal and (G– H) ventral view; detail of (I) dentate
holes and of (J) anchor handle; (K) rods and (L) rosettes from tentacles. Scale bars: A– C, 100 mm; D, 20 mm; E – H, 100 mm; I –L, 20 mm.
and radial pieces, perforated at top for insertion of radial remarks

nerves. Many Polian vesicles, madreporite single. There are four valid species of Euapta worldwide: E. lappa
Body wall anchors (200 –360 mm; Figure 19A – C) with differs from E. magna Heding, 1928 in the presence of
smooth arms, vertex with minute knobs (about 12), stock webbed digits on the tentacles, and from E. tahitiensis
branched (8 –9), minute knobs at ends (60 – 100 mm; Cherbonnier, 1955 in the shape of the anchor plate. Euapta
Figure 19D). Anchor plates oval (200 –300 mm long, 130 – lappa and E. godeffroyi were historically separated by their
180 mm wide; Figure 19E –J), margins smooth, undulating; pattern of geographic distribution (West-Indian and Indo-
anterior region broad, seven dentate holes, three in midline, Pacific, respectively). Clark (1924) distinguished them by the
two in each side (denticles forming circles in both ventral size of their anchors and anchor plates (larger in E. lappa),
and dorsal sides); bridge smooth, well developed; posterior while Heding (1928) found differences in the shape of their
region narrow with two large and 5– 6 marginal holes. anchors.
Tentacles with spiniform rods (100– 300 mm; Figure 19K), This is the first record of E. lappa from the Brazilian coast.
slightly branched ends, and rosettes (10 – 40 mm; Figure 19L). The specimens from the BOI differ from the specimens from
Ascension islands in having larger anchors and anchor plates.
No other morphological difference was found between them.
distribution
Occurs from the Gulf of Mexico to Brazil (Espı́rito Santo); Order ASPIDOCHIROTIDA Grube, 1840
oceanic island distribution: ASC, STH and Trindade Islands Family HOLOTHURIIDAE Burmeister, 1837
(Pawson, 1978; Hendler et al., 1995; present paper). Genus Actinopyga Bronn, 1860
Bathymetric range: intertidal to 1350 m (Hendler et al., Actinopyga agassizii (Selenka, 1867)
1995; Alvarado & Solı́s-Marı́n, 2013). (Figures 20 & 21)
Mülleria agassizii Selenka, 1867, pp. 311.

biological notes Actinopyga agassizii – Hendler et al., 1995, pp. 282 – 284,
In TMV this species has been found near rhodolites. figures 157, 180.
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Fig. 20. Actinopyga agassizii (Selenka, 1867) (MZUSP 293). (A) Specimen from Trindade Island photographed in situ, depth not recorded; (B) calcareous ring; (C)
top view of mouth showing numerous tentacular ampullae (indicated by a yellow arrow in (D)) and a mottled brown Polian vesicle (indicated by a white arrow in
(D)); and (E) detail of anal teeth. Scale bars: B, 5 mm; C, 1 mm; D, 5 mm; E, 3 cm.
examined material 294); 12.5 m, 31.1.2012, 1 spm 17 cm long (MZUSP 297);

Martin Vaz archipelago, ES, Brazil (20830′ S 29818′ W) – 16 m, 2.7.2012, 2 spms 15 cm long (MZUSP 303). Lixo
Martin Vaz Island, 13 m, 23.7.2013, 13 spms 9 –15 cm long beach, 14 m, 22.4.2014, 2 spms 15 –20 cm long (MZUSP
(MZUSP 325, 327). Trindade Island, ES, Brazil (20829′ – 964). Pedra da Garoupa, 10 m, 1 spm 3 cm long (MZUSP
20831′ S 29817′ –20820′ W) – Andrada beach, 10 m, 316). Rachada Island, 27 m, 2.6.2012, 1 spm 12 cm long
17.7.2013, 1 spm 14 cm long (MZUSP 326). Calheta beach, (MZUSP 302). Tartarugas beach, 12.7.2013, 1 spm 12 cm
12 m, 16.6.2012, 4 spms 15 –17 cm long (MZUSP 292); long (MZUSP 415).
18.6.2012, 1 spm 14 cm long (MZUSP 293); 14 m,
26.6.2012, 1 spm 12 cm long (MZUSP 420); 12 m, description
18.7.2012, 2 spms 13– 15 cm long (MZUSP 723); 3.7.2012, 4 Body thick (Figure 20A), without distinct warts dorsally or lat-
spms 9 cm long (MZUSP 305); 16 m, 28.4.2014, 6 spms 15– erally, maximum length 28 cm. Body wall mottled brown,
28 cm long (MZUSP 957, 960); 15 m, 29.4.2014, 1 spm orange or yellow. 20 –30 large yellow peltate tentacles.
15 cm long (MZUSP 966); 12 m, 14.5.2014, 1 spm 11 cm Calcareous ring (Figure 20B) without posterior projections;
long (MZUSP 954); 16 m, 20.5.2014, 1 spm 11 cm long radial and interradial plates square-shaped, united along all
(MZUSP 953). Enseada da Cachoeira, 9 m, 8.8.2013, 1 spm length. Radial plates with two notches, three short projections
14 cm long (MZUSP 418). Enseada de Orelhas, 10 m, in anterior region, base slightly curved. Interradial plates with
15.6.2012, 4 spms 12 –15 cm long (MZUSP 304); 10 m, three anterior projections (middle one bifurcated), straight
6.7.2013, 1 spm 14 cm long (MZUSP 414). Farol beach, base. Tentacular ampullae large, numerous (20 –30;
12 m, 17.7.2013, 1 spm 3 cm long (MZUSP 320); 13 m, Figure 20C), mottled brown Polian vesicle (Figure 20D).
22.4.2014, 4 spms 4–28 cm long (MZUSP 957, 963). Five conspicuous, white, calcareous teeth surrounding anus
Farrilhões beach, 31.1.2012, 1 spm 12 cm long (MZUSP (Figure 20E).
Fig. 21. Actinopyga agassizii (Selenka, 1867) (MZUSP 293). SEM images of the ossicles – rosette-like elements from body wall: (A) ‘dog biscuit’-shaped and (B)
branched rods; (C– D) rods from tentacles; and (E) end plate and (F) rod from tube feet. Scale bars: A– B, 20 mm; C– E, 100 mm; F, 20 mm.
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Body wall rosette-like ossicles varying from simple ‘dog examined material
biscuit’ shapes (20 – 60 mm; Figure 21A) to complex dichot- Trindade Island, ES, Brazil (29830′ S 29820′ W) – Racha Island,
omously branched rods (20 – 40 mm; Figure 21B). Tentacles 27 m, 29.7.2015, 1 spm 6 cm long (MZUSP 1305).
with two types of rods: curved and spiniform rods with perfo-
rated ends (150– 200 mm; Figure 21C) and straight rods,
smooth along their length and spiniform tips (40 –70 mm; description
Figure 21D) (40–800 mm). Tube feet with perforated end Body elongate (Figure 22A). Tube feet throughout whole
plate (200 –400 mm; Figure 21E); rods minutely branched, body, maximum length 10 cm. Colour white in ethanol. Ten
smooth along length, tips branched (30 – 100 mm; Figure 21F). peltate tentacles. Calcareous ring simple, delicate, without
posterior projections (Figure 22B, C). Radial plates rectangu-
distribution lar-shaped, anterior region with one large notch, posterior
Occurs from South Carolina (USA) to the Caribbean; oceanic region curved. Interradial plates with triangular anterior
island distribution: Bermuda, TMV (Pawson et al., 2010; region, curved base. Internal organs were degraded and
present paper). Bathymetric range: intertidal up to 54 m body was filled with sediment.
(Pawson et al., 2010). Body wall tables four-pillared (50–70 mm; Figure 23A –C),
disc with one large central hole, eight marginal oval perfora-
tions, disc margin undulated, knobs bifurcated, adjacent to
biological notes marginal perforations; spire knobbed, ending in 15 –30 teeth
Found in association with the pearlfish Carapus sp.
projected laterally and upwards; buttons elongated, knobbed
(100 –160 mm; Figure 23D), margins undulated, 12 –14 per-
remarks forations of unequal size arranged into two parallel rows;
According to Panning (1944), the genus Actinopyga has about middle bar knobbed, usually projecting beyond one extremity
18 valid species subdivided into three main groups: ‘Act. leca- (rarely beyond both extremities, or not projecting at all).
nora’, characterized by having simple and smooth rosettes, Ventral region of body with four-pillared tables, disc with
which occasionally have lateral extensions and swollen ends; one central hole, 6– 8 marginal perforations, disc margin
‘Act. echinites’, characterized by the narrowing and the elong- undulated, spire tall, several projections at tip (spire height:
ation of the rosettes from the ventral body wall; and the 40 –70 mm, disc diameter: 60 –70 mm; Figure 23E, F).
‘obesa-gruppe’, characterized by the presence of rough and Tentacles with two types of spiniform rods: large curved
often branched rods. Actinopyga agassizii belongs to the rods with pointed projections along outer side and at tips,
‘Act. echinites’ group, and is the only species of the genus inner side smooth, tips sometimes with tiny perforations
that occurs in the western Atlantic. (100 –180 mm; Figure 23G –I); small straight rods with
Actinopyga agassizii has been mentioned from Porto coarse projections along whole ossicle (40–70 mm;
Seguro (Brazil) in a checklist without any characterization Figure 23J). Tube feet end plate (100 –240 mm; Figure 23K)
or illustration (see Magalhães et al., 2005). This is, therefore, with large perforations, smaller perforations on edges; rods
the first formal record of this species from the South with 4– 10 central perforations, 2 –3 perforations at tips
Atlantic. SEM images of ossicles, images of specimens in (100 –200 mm; Figure 23L, M).
situ and of internal morphology are provided.
Genus Holothuria Linnaeus, 1767 distribution
Holothuria (Cystipus) pseudofossor Deichmann, 1930 Occurs from the Gulf of Mexico to São Paulo (Brazil); oceanic
(Figures 22 & 23) island distribution: Trindade Island (Tommasi, 1974;
Holothuria pseudofossor Deichmann, 1930, pp. 57 –58, figures Alvarado & Solı́s-Marı́n, 2013; present paper). Bathymetric
9– 14. range: 3– 370 m (Alvarado & Solı́s-Marı́n, 2013).
Holothuria pseudofossor — Deichmann, 1954, pp. 389, 394,
figure 66, 29– 34. remarks
Holothuria (Cystipus) pseudofossor — Rowe, 1969, pp. 157; There are three species of Holothuria (Cystipus) in the Atlantic
Tommasi, 1974, pp. 2, figure 1; Borrero-Pérez et al., 2012, pp. Ocean: H. (C.) pseudofossor differs from H. (C.) cubana
182 –183; Prata et al., 2014, pp. 138 –140, Figure 5. Ludwig, 1875 by the absence of ‘large supporting plates’
(sensu Deichmann, 1930) in the tube feet and the simple
morphology of the table spire (vs. heavily knobbed spires
sometimes ending in a sphere with blunt teeth; note that
simpler tables are also found in this species, and these are
probably an earlier developmental stage of the tables), and
from H. (C.) occidentalis Ludwig, 1875 in having simple flat-
tened knobbed buttons (vs. complex ovoid knobbed buttons
sometimes folded and/or with protuberances).
No morphological differences were observed between spe-
cimens from the BOI and the Brazilian coast. However, onto-
genetic variation, which has already been documented for
Caribbean specimens (Deichmann, 1930; Ortiz-Gómez et al.,
Fig. 22. Holothuria (Cystipus) pseudofossor Deichmann, 1930 (MZUSP 1305).
(A) Specimen fixed in ethanol, (B) photograph and (C) drawing of the
2006), was found. The specimens described by Tommasi
calcareous ring (‘R’, radial plate, ‘I’, interradial plate). Scale bars: A, 1 cm; B, (1974; length ¼ 2.4 cm) and Prata et al. (2014; length ¼
2 mm. 4 cm) from Paraı́ba (North-east of Brazil) have buttons and
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Fig. 23. Holothuria (Cystipus) pseudofossor Deichmann, 1930 (MZUSP 1305). SEM images of the ossicles – tables from body wall in (A) bottom, (B) lateral and
(C) top view; (D) button from body wall; tables from ventral region in (E) lateral and (F) top view; (G– H) large curved rods ((I) detail of the tip of the rod depicted
in (H) showing tiny perforation), and (J) small straight rod from tentacles; (K) end plate and (L –M) rods from tube feet. Scale bars: A– C, 20 mm; D, 50 mm; E– F,
20 mm; G – H, 50 mm; J, 10 mm; K– M, 100 mm.
tables with undeveloped knobs, and the teeth at the top of

their tables are also undeveloped.
Holothuria (Halodeima) grisea Selenka, 1867

(Figures 24 & 25)
Holothuria grisea Selenka, 1867, p. 328, pl. 18.

Holothuria unicolor Selenka, 1867, p. 329, pl. 18.
Holothuria (Holothuria) grisea – Panning, 1935, p. 31,
Fig. 24. Holothuria (Halodeima) grisea Selenka, 1867 (MZUSP 1057). (A)
figure. 23; Cutress, 1996, pp. 51 –55, figures 5 & 6; Pawson
Specimen fixed in ethanol; detail of calcareous ring (B) before and (C) after et al., 2010, p. 36, figure 29; Prata et al., 2014, pp. 132 –134,
bleach treatment. Scale bars: A, 6 cm; B –C, 5 mm. figure 3.
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Fig. 25. Holothuria (Halodeima) grisea Selenka, 1867 (MZUSP 1057). SEM images of the ossicles: body wall tables in (A) lateral, (B) top and (C) bottom view; (D–
F) different developmental stages of body wall buttons; rods from (G – M) tentacles, and (N) the tube feet. Scale bars: A, 20 mm; B – C, 10 mm; D – F, 20 mm; G – N,
100 mm.
examined material from grey to brownish; tube feet yellow, covering body,
Trindade Island, ES, Brazil (20830′ S 29818′ W) – Calheta more abundant in ventral region forming a sole. 20 –25
beach, 5 m, 10.4.2014, 1 spm 12 cm long (MZUSP 1057). black peltate tentacles. Calcareous ring (Figure 24B, C)
short, simple, without projections. Radial plates square-
comparative material examined shaped, anterior region with single-notched, posterior region
Holothuria (Halodeima) grisea: North-eastern Brazilian coast curved. Interradial plates with triangular anterior region,
– Cairu, BA, 17.9.2012, 1 spm 11 cm long; Maceió, AL, inter- base curved.
tidal, 31.5.2011, 1 spm 10 cm long (MZUSP 540). Natal, RN, Body wall tables four-pillared (30 –80 mm; Figure 25A –
intertidal, 16.11.2009, 1 spm 14 cm long (MZUSP 534). Porto C), one central, 9–10 marginal perforations; disc with 12 –
Seguro, BA, 1 spm 14 cm long (MZUSP 563). South-eastern 14 marginal spines, spire with 16 spines (4 groups of 4) at
Brazilian coast – Paraty, RJ, 19.9.1960, 1 spm 12 cm long tip; buttons oval (30 – 60 mm; Figure 25D – F), 2–4 central
(MZUSP 531). São Sebastião, SP, 2 m, 31.10.2013, 1 spm and 2– 6 marginal perforations. Tentacles with two types
14 cm long (MZUSP 422). South Brazilian coast – Balneário of rods: curved rods, smooth along length, spiniform at
da Penha, SC, 1 spm 13 cm long (MZUSP 552). tips (70 – 120 mm; Figure 25G); complex rods with many
perforations, sometimes highly ramified (140– 600 mm;
description Figure 2H– M). Tube feet with end plate (400 –600 mm),
Body cylindrical, (Figure 24A) warts distinct, flattened ven- rods smooth (90–300 mm; Figure 25N), 5– 6 perforations
trally; maximum length 12 cm. Body wall colour varying at tips.
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distribution description
Western Atlantic: from Florida to Brazil; oceanic island distri- Body cylindrical (Figure 26A –B), brown maximum length
bution: ASC and Trindade Islands (Pawson, 1978; Hendler 22 cm. 20 –24 olive green peltate tentacles. Calcareous ring
et al., 1995; present paper). Bathymetric range: intertidal (Figure 26C) robust, square-shaped plates lacking posterior
down to 25 m (Prata et al., 2014). projections, united along length. Radial plates three-notched,
four thick anterior projections, slightly curved base.
biological notes Interradial plates with one notch and two thick anterior pro-
In TMV was found in rocky bottoms, near Actinopyga jections. Tentacular ampullae numerous, one Polian vesicle,
agassizii. dark-spotted (Figure 26D). Tube feet sparsely scattered dor-
sally, more numerous ventrally, but not forming conspicuous
sole. Anus with five thick papillae (Figure 26E).
remarks Body wall tables (40 – 70 mm; Figure 27A) with reduced
There are four Holothuria (H.) species in the western Atlantic: smooth discs (20 – 30 mm; Figure 27B), long spire surrounded
H. (H.) grisea differs from H. (H.) manningi Pawson, 1978 by by 20 projections at top; two small perforations at tips of
the morphology of the body wall tables (Knobbed in H. (H.) buttons (15 – 30 mm; Figure 27C), two large middle perfora-
manningi); and from H. (H.) floridana (Pourtalès, 1851) and tions. Tentacles with minute spiniform rods (50–70 mm;
H. (H.) mexicana Ludwig, 1875 by the number of Polian vesi- Figure 27D – F), often curved and very scarce. Tube feet with
cles (1 –3 vs. 2–3 and 1, respectively) and in having free stone flat perforated plates (80 – 200 mm; Figure 27G, H), and end
canals (vs. stone canals forming lateral tufts). plates (100– 200 mm; Figure 27I).
Holothuria (H.) grisea is an amphi-Atlantic species, occur-
ring along the entire Brazilian coast. Probably because of its
wide geographic distribution, this species is thought to be a
‘species-complex’. However, thorough morphological com- distribution
parisons are required to properly address this issue. The speci- Ascension (type locality), SPP and Trindade Islands (Pawson,
mens from the BOI differ from the specimens from the 1978; present paper). Bathymetric range: up to 12 m (Pawson,
Brazilian coast only in the shape of the tentacle rods, which 1978; present paper).
are actually more similar to the rods from Bahamas specimens
(see Pawson, 1978); and from the specimens from Ascension
by the larger size of their tables and buttons.
remarks
Holothuria (Halodeima) manningi Pawson, 1978 Holothuria (H.) manningi can be distinguished from all other
(Figures 26 & 27) Holothuria (Halodeima) species in having tables with reduced
discs, buttons with four perforations, and flat perforated
Holothuria (Halodeima) manningi Pawson, 1978, p. 26, pl. 11. plates. No morphological differences were found between
the specimens from the BOI and the specimens from
examined material Ascension Island. Surprisingly, the ossicles (i.e. tables and
Trindade Island, ES, Brazil (20831′ S 29817′ W) – Tartarugas flat perforated plates) of Holothuria (H.) manningi show
beach, 12 m, 1 spm 17 cm long (MZUSP 463). greater similarity to the Indo-Pacific species H. (H.) nigralutea
O’Loughlin, 2007 and H. (H.) edulis Lesson, 1830 (see
additional material examined O’Loughlin et al., 2007).
São Pedro e São Paulo archipelago, PE, Brazil (20830′ S This is the first record of this species in the western Atlantic
29818′ W) – Ilha Belmonte, 22.9.1979, 1 spm 22 cm long Ocean.
(MZUSP 725); 27.10.2007, 1 spm 19 cm long (MZUSP 306); Holothuria (Semperothuria) surinamensis Ludwig, 1875
05.3.2009, 1 spm 19 cm long (MZUSP 301). (Figures 28 & 29)
Holothuria surinamensis Ludwig, 1875, p. 111, figure 27.
Holothuria surinamensis – Deichmann, 1930, pp. 63 –64,
figures 12 –15, 19.
Holothuria (Semperothuria) surinamensis – Rowe, 1969,
pp. 135 –136, figure 6.
Semperothuria surinamensis – Deichmann, 1958, p. 303.
examined material
Trindade Island, ES, Brazil (20829′ – 20831′ S 29818′ –
20820′ W) – Calheta beach, 16 m, 28.4.2014, 1 spm 30 cm
long (MZUSP 958). Enseada de Orelhas, 14 m, 5.4.2014, 1
spm 10 cm long (MZUSP 951). Principe beach, 14 m,
23.4.2014, 2 spms 18 –24 cm long (MZUSP 959). Secon,
10 m, 29.4.2014, 2 spms 2.5 – 3 cm long (MZUSP 947, 948).
Fig. 26. Holothuria (Halodeima) manningi Pawson, 1978 (MZUSP 301). (A)
Dorsal and (B) ventral view of specimen fixed in ethanol; (C) calcareous ring;
Secon, 9 m, 12.5.2014, 1 spm 12 cm long (MZUSP 1009).
(D); arrows indicate the Polian vesicles; and (E) detail of anal papillae. Scale Farrilhões beach, 12 m, 04.4.2014, 2 spms 16– 26 cm long
bars: A –B, 10 cm; C– D, 2 cm; E, 1 cm. (MZUSP 1058).
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Fig. 27. Holothuria (Halodeima) manningi Pawson, 1978 (MZUSP 301). SEM images of the ossicles – body wall tables in (A) lateral view and (B) detail of reduced
disc; (C) buttons from body wall; (D – F) rods from tentacles; and (G– H) flattened plates and (I) end plate from tube feet. Scale bars: A– B, 20 mm; C, 10 mm; D– F,
20 mm; G – I, 50 mm.
comparative material examined Body wall tables four-pillared (30–70 mm; Figure 29A –C),
Holothuria (Semperothuria) surinamensis: Angra dos Reis, RJ, usually cylindrical, with reduced or absent discs, 10– 16 pro-
Brazil – 2.1956, 4 spms 6– 8 cm long (MZUSP 518). Vila jections in double layers on top of spire. Tentacles with two
Velha beach, 21.7.1996, 1 spm 8 cm long (MZUSP 521). types of rods: minute curved rods, often branched at tips,
some projections along length (40 – 70 mm; Figure 29D, E);
large curved rods smooth along length, tips spiniform, some-
description times perforated with tiny holes (100 –600 mm; Figure 29F,
Body cylindrical (Figure 28A), maximum length 30 cm. Body G). Tube feet with flattened plates, undulating margins, two
wall light brown, 20 pale peltate tentacles (Figure 28B). large middle perforations, small perforations elsewhere
Calcareous ring (Figure 28C) simple, short, without projec- (100 –350 mm; Figure 29H); straight rods with solid midline,
tions. Radial plates square-shaped, one central notch, base perforations at tips and/or along margins (100– 200 mm;
curved. Interradial plates small, triangular, slightly curved at Figure 29I, J); end plate with several small perforations
base, united to radial plates at base. One stone canal, head (400 –600 mm; Figure 29K).
oblong, Polian vesicle very long, madreporite long, filament-
ous (Figure 28D). Tube feet scarce dorsally, abundant distribution
ventrally. Occurs from Bermuda to Southern Brazil; oceanic island dis-
tribution: Trindade Island (Clark, 1898; Tommasi, 1969;
Pawson et al., 2010; present paper). Bathymetric range: inter-
tidal down to 42 m (Pawson et al., 2010).
remarks
Seven species are recognized in the subgenus Holothuria
(Semperothuria): Holothuria (S.) surinamensis differs from
H. (S.) imitans Ludwig, 1875 (circumtropical) and H. (S.) lan-
guens Selenka, 1867 (eastern Pacific) in the number of spines
at top of table (10 – 16 vs. 8 and 8), in the morphology of the
perforated plate-like rods from the tube feet (undulating vs.
dentate outline); from H. (S.) flavomaculata Semper, 1868
(Indo-Pacific) and H. (S.) roseomaculata Kerr, 2013
(western Pacific) in the morphology of the rods from the
tube feet (smooth vs. spiniform); and from H. (S.) cinerascens
(Brandt, 1835) and H. (S.) granosa Cherbonnier, 1988 (Indo-
Pacific) in the presence of perforated plates in the tube feet.
No difference was observed between species from the BOI
and from the Brazilian coast. According to Cutress (1996), the
Fig. 28. Holothuria (Semperothuria) surinamensis Ludwig, 1875 (MZUSP
body wall tables vary over ontogeny in H. (S.) surinamensis: in
1058). (A) Specimen fixed in ethanol; (B) peltate tentacles; (C) calcareous young specimens (up to 3 cm), tables have large discs with
ring; (D) madreporite. Scale bars: A, 5 cm; C, 2 mm; D, 1 mm. several perforations (7 – 8) while in adults tables have
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Fig. 29. Holothuria (Semperothuria) surinamensis Ludwig, 1875 (MZUSP 1058). SEM images of the ossicles – body wall tables in (A) lateral, (B) bottom, and (C)
top view; (D– G) rods from tentacles; (H) plate, (I– J) rods, and (K) end plate from tube feet. Scale bars: A– C, 20 mm; D, 10 mm; E, 20 mm; F– G, 100 mm; H,
50 mm; I – K, 100 mm.
reduced discs with a single perforation. This same pattern was Bradtothuria arenicola – Deichmann, 1958, p. 290, pl. 1.
observed in the young specimens of H. (S.) surinamensis Holothuria arenicola – Deichmann, 1930, p. 66, pl. 4.
studied herein. Holothuria (Holothuria) arenicola – Panning, 1935, pp.
88 –89, figure 73.
Holothuria (Thymiosycia) arenicola Semper, 1868 Holothuria (Thymiosycia) arenicola – Rowe, 1969, p. 147;
(Figures 30 & 31) Cutress, 1996, pp. 84 –88, figures 24 & 25.
Holothuria arenicola Semper, 1868, p. 61, pls. 20, 30, 35.

examined material
Martin Vaz archipelago, ES, Brazil (20830′ S 29818′ W) –
Martin Vaz Island, 13 m, 26.6.2013, 1 spm 12 cm long
(MZUSP 611); 20 m, 23.7.2013, 1 spm 9 cm long (MZUSP
648). Trindade Island, ES, Brazil (20829′ – 20832′ S 29817′ –
20820′ W) – Andrada beach, intertidal, 5– 21.7.2013, 2 spms
11 –15 cm long (MZUSP 323, 416). Calheta beach,
26.6.2012, 2 spms 8– 12 cm long (MZUSP 296, 421); 14 m,
16.9.2013, 12 spms 8– 13 cm long (MZUSP 413); 4 m,
18.5.2014, 4 spms 4–7 cm long (MZUSP 973, 974). Enseada
da Cachoeira, 10 m, 9.7.2013, 1 spm 11 cm long (MZUSP
417). Enseada de Orelhas, 15 m, 20.6.2012, 1 spm 11 cm
long (MZUSP 313). Enseada Portuguesa, 14 m, 10.7.2012, 4
spms 9 –12 cm long (MZUSP 312); 10 m, 18– 23.4.2014, 28
spms 3–12 cm long (MZUSP 972, 975). Farilhões beach,
Fig. 30. Holothuria (Thymiosycia) arenicola Semper, 1868 (MZUSP 949). (A) 11 m, 4.7.2012, 1 spm 9 cm long (MZUSP 308). Farol beach,
Specimen from Trindade Island photographed in situ, at 10 m depth (the 13 m, 22.4.2012, 1 spm 8 cm long (MZUSP 950); 12 m,
colour may have been affected by light gradients), and (B) fixed in ethanol;
(C) peltate tentacles; (D) calcareous ring; (E) white arrow indicates
17.7.2013, 1 spm 8 cm long (MZUSP 321); 13 m, 22.4.2014,
tentacular ampullae attached to the calcareous ring (red arrow). Scale bars: 1 spm 5 cm long (MZUSP 962). M beach, 15 m, 8.7.2013, 1
B, 5 cm; C, 2 cm; D, 2 mm; E, 1 mm. spm 16 cm long (MZUSP 322). Pedra da Garoupa, 12 m,
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Fig. 31. Holothuria (Thymiosycia) arenicola Semper, 1868 (MZUSP 949). SEM images of the ossicles – tables from body wall in (A) top and (B) lateral view; (C)
button from body wall; (D –F) rods from tentacles; (G– H) rods and (I) end plate from tube feet. Scale bars: A –C, 20 mm; D – H, 50 mm; I, 100 mm.
16.7.2013, 1 spm 13 cm long (MZUSP 324). Ponta Noroeste, distribution

14 m, 27.1.2012, 1 spm 9 cm long (MZUSP 309). Ponta Occurs from the Gulf of Mexico to Brazil (Western
Norte, 12 m, 17.7.2013, 1 spm 13 cm long (MZUSP 313); Atlantic) and in the Gulf of California (eastern Pacific);
11 m, 1.4.2014, 2 spms 6– 16 cm long (MZUSP 956). Secon, oceanic island distribution: ASC, STH and TMV (Pawson,
10 m, 29.4.2014, 1 spm 10 cm long (MZUSP 949). 1978; Hendler et al., 1995; Solı́s-Marı́n et al., 2009; present
Tartarugas beach, 14 m, 06.7.2012, 1 spm 9 cm long paper). Bathymetric range: intertidal to 121 m (Solı́s-Marı́n
(MZUSP 311). et al., 2009).
remarks
description Holothuria (T.) arenicola is considered a circumtropical
Body cylindrical (Figure 30A, B), maximum length 20 cm. 20 species and surprisingly, there are only few differences in
small peltate tentacles (Figure 30C). Body brown or beige with the size and shape of the ossicles throughout its geographic
brown spots, tentacles white. Calcareous ring (Figure 30D) range (Deichmann, 1930). There are three species of
short, delicate, tentacular ampullae numerous (Figure 30E), Holothuria (Thymiosycia) in the Atlantic Ocean: H. (T.) are-
two Polian vesicles. Tube feet scattered throughout body. nicola differs from H. (T.) impatiens Forsskål, 1775 (and
Body wall tables four-pillared (50 – 100 mm; Figure 31A, B), Holothuria (T.) thomasi Pawson & Caycedo, 1980 by the
subcircular, smooth disc perforated by one central, 4–8 mar- morphology of the body wall table discs (circular vs. square-
ginal hole, spire ending in 8– 12 spines, buttons elongated shaped and irregular outline, respectively).
(60 – 80 mm; Figure 31C), three pairs of abundant elongated The specimens from the BOI differ from the specimens
perforations. Tentacles with straight, curved rods, smooth from Ascension Island in having larger buttons and tables.
along length, some spines at tips (80 –120 mm; Figure 31D– Body wall ossicles composition in aspidochirotids changes
F). Tube feet rods (130– 180 mm; Figure 31G, H) with solid over ontogenic; buttons of H. (T.) arenicola, for instance,
midline, 4–6 marginal perforations; end plate (200– are not present in younger specimens (Deichmann, 1930;
300 mm; Figure 31I) with large perforations. Pawson, 1978; Cutress, 1996). In the specimens studied
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955); 15 m, 29.4.2014, 1 spm 20 cm long (MZUSP 965);

16 m, 12.5.2014, 1 spm 15 cm long (MZUSP 968); 17 m,
28.4.2014, 4 spms 15 –26 cm long (MZUSP 935, 967, 969,
971). Enseada de Orelhas, 10 m, 16.5.2014, 1 spm 14 cm
long (MZUSP 970). Enseada Portuguesa, 11 m, 23.4.2014, 1
spm 19 cm long (MZUSP 933). Farol beach, 13 m,
22.4.2014, 1 spm 4 cm long (MZUSP 961); 14 m, 22.4.2014,
2 spms 5 –8 cm long (MZUSP 932). Farrilhões beach, 17 m,
20.6.2012, 1 spm 13 cm long (MZUSP 307). Ponta Norte,
11.5 m, 1.4.2014, 1 spm 12 cm long (MZUSP 936).
additional material examined

São Pedro and São Paulo Archipelago, PE, Brazil (20830′ S
Fig. 32. Isostichopus badionotus (Selenka, 1867) (MZUSP 936). (A) Specimen 29818′ W) – Ilha Belmonte, 1980, 1 spm 14 long (MZUSP
from Trindade Island photographed in situ, at 11.5 m depth, and (B) fixed in 727).
ethanol; (C) calcareous ring; (D) tentacular ampullae indicated by arrow; and
(E) detail of the madreporite. Scale bars: B, 5 cm; C– E, 5 mm.
description
herein, body wall buttons were present only in specimens Body thick (Figure 32A, B), warts distinct dorsally and lat-
larger than 3 cm. erally, maximum length 26 cm. 20 brown peltate tentacles.
Body wall molted dark brown, mustard. Calcareous ring
Family STICHOPODIDAE Haeckel (Figure 32C) robust, plates are united throughout. Radial
Genus Isostichopus Deichmann plate square-shaped, strongly dipping posteriorly, projections
Isostichopus badionotus (Selenka, 1867) short. Interradial plates with triangular apex, curved base. One
(Figures 32 & 33) Polian vesicles; one stone canal; large tentacular ampullae
(Figure 32D); madreporite long, filamentous (Figure 32E).
Stichopus badionotus Selenka, 1867, p. 316, pl. 18. Dorsal and ventral surfaces sharply defined by lateral rim of
Isostichopus badionotus – Deichmann, 1958, pp. 279 –280; conspicuous papillae. Ventral surface flat, covered with
Hendler et al., 1995, p. 280, figures156, 187; Cutress, 1996, pp. numerous cylindrical tube feet organized into three rows.
105 –113, figures 37– 40. Body wall tables four-pillared (30 – 45 mm; Figure 33A, B),
Stichopus badionotus – Deichmann, 1930, p. 80, pl. 5. disc with one central hole, 8–9 marginal perforations, margin
smooth, undulating, spire ending in about 24 teeth; C-shaped
examined material and S-shaped ossicles (50 – 60 mm). Tentacles with two types
Trindade Island, ES, Brazil (20829′ – 20831′ S 29818′ – of rods: curved and thick rods with minute spines (300–
20820′ W) – Calheta beach, 12 m, 18.6.2012, 1 spm 15 cm 800 mm; Figure 33C, D), nearly straight and thin rods with
long (MZUSP 291); 14 m, 26.6.2012, 1 spm 14 cm long large spines (50 – 120 mm; Figure 33E). Tube feet rods (200–
(MZUSP 295); 12 m, 14.5.2014, 1 spm 12 cm long (MZUSP 400 mm; Figure 33F, G) with perforations in central region,
Fig. 33. Isostichopus badionotus (Selenka, 1867) (MZUSP 936). SEM images of the ossicles – tables from body wall in (A) lateral and (B) top view; (C– E) rods
from tentacles; (F– G) rods and (H) end plate from tube feet. Scale bars: A, 20 mm; B, 5 mm; C– D, 200 mm; E, 50 mm; F– G, 100 mm; H, 200 mm.
http://dx.doi.org/10.1017/S0025315416001569
sometimes at the tips; end plates (300– 600 mm; Figure 33H) of larval pressure and of the role of the oceanic islands as step-
large, with small perforations. ping stones in the dispersal of the shallow-water benthic fauna
(Table 2).
distribution Dendrochirotids and crinoids were not found in the TMV
Gulf of Mexico to Brazil (western Atlantic), and Gulf of and ASP, nor in the AST (Pawson, 1978, present paper) even
Guinea (eastern Atlantic); oceanic island distribution: SPP, though they are the most diverse shallow-water holothuroids
ASC and Trindade Islands (Pawson, 1978; Hendler et al., in the Brazilian coast (Alvarado & Solı́s-Marı́n, 2013).
1995; present paper). Bathymetric range: intertidal up to Different life history strategies (e.g. reproductive mode,
65 m (Hendler et al., 1995). larval duration, ecology and dispersal potential) will affect dis-
persal and endemicity of species (Floeter & Gasparini, 2000;
remarks Leal, 2000). The absence of dendrochirotids and crinoids in
Isostichopus is composed by three species: I. badionotus differs the BOI are probably due to the lack of a planktonic larval
from I. macroparentheses (Clark, 1922a) in having smaller stage (Pawson, 1978). An additional explanation is the lack
C-shaped ossicles and from I. fuscus (Ludwig, 1875) in of suitable habitat. Many dendrochirotid species (e.g.
having tables with circular outline (vs. squarish). According Coronatum baiensis Martins & Souto and Ocnus braziliensis
to Clark (1922b) and Deichmann (1930), the three species (Verrill)) live preferably under rocks; however, loose rocks
cannot be distinguished from one another using external were scarce in the TMV.
morphology. Unexpectedly, despite the prevalence of the trans-Atlantic
No differences were observed among the specimens from South Equatorial Current (Colling, 2001), which brings
the BOI, Ascension Island and the Brazilian coast. warm waters from West Africa towards Brazil, therefore facili-
According to Pawson (1978) and Cutress (1996), the ossicles tating the spillover of eastern Atlantic species westward, the
change throughout ontogeny in Isostichopus. Similarly, in echinozoan fauna of the TMV has a strong western Atlantic
the Brazilian specimens of I. badionotus tables decreased in component (78.9%), whereas the effect of the eastern
width and height with increasing size and C-shaped ossicles, Atlantic component on the insular echinozoan fauna of
absent in juveniles, were present in larger specimens. TMV is too low (21%). This is consistent with the results
found by Anker et al. (2016) from a dataset of 120 alpheid
shrimp species. Indeed, about 20% of the echinoid and holo-
DISCUSSION thuroid species found along the Brazilian coast are also found
in Trindade and Martin Vaz, supporting the suggestion that
TMV (and other Brazilian oceanic islands) is to a great
Zoogeographic notes extent a subset of the Brazilian shallow-water fauna in
A total of 16 echinozoan species are reported herein from the oceanic domain (Edwards & Lubbock, 1983; Floeter &
TMV, and SPP. Of these, 15 (78.9%) are also known from the Gasparini, 2000; Anker et al., 2016).
western Atlantic coast, 4 (21%) are eastern Atlantic species, 1 Based on our morphological observations and from the
(5%) is a so far restricted insular species and 16 (84%) are number of subspecies described so far, there is little morpho-
amphi-Atlantic in distribution. The Trindade Island is home logical variation between the conspecific apodid and aspido-
to two endemic echinoids, Centrostephanus besnardi chirotid holothuroids throughout the Atlantic. Their
Bernasconi, 1955 and Clypeaster oliveirai Krau, 1952, none similarity could be either the result of the relative simplicity
of them found in the present survey. Centrostephanus besnardi of body organization and/or their great effectiveness as disper-
and C. oliveirai are said to inhabit depths greater than 30 m sers. Actually, we found slight morphological differences
(Krau, 1952; Bernasconi, 1955b) and have not been reported between conspecifics from the TMV and the coast; however,
again since they were described, probably because they live we preferred to be conservative and not describe new
in depths less accessible to scuba divers. Although the depth species based on slight differences that could be a result of
range for C. besnardi is acually unknown (Bernasconi, environmental plasticity. Whether molecular data can
1955b), it supposedly inhabits deep waters as do its congeners provide further resolution and help in clarifying the separ-
known from depths beyond 40 m (Pawson & Miller, 1983). ation of TMVs and mainland populations is unclear. Indeed,
Clypeaster oliveirai was caught at 50 m. two echinoids have been described from Ascension Island
The holothuroid Actinopyga agassizii and the echinoid based on slight morphological variation from coastal speci-
Pseudoboletia maculata are new records from the south- mens (Echinometra lucunter polyplora and Eucidaris
western Atlantic, whereas the holothuroid Euapta lappa is clavata). In both cases, molecular data (although restricted
recorded from the Brazilian territorial waters for the first to the COI gene) have not fully supported the proposed sep-
time. The echinoid Eucidaris tribuloides tribuloides is first aration (Lessios et al., 1999; McCartney et al., 2000).
recorded from the SPP. The echinoids Leodia sexiesperforata, Although quite far from the coast, the TMV are connected
Plagiobrissus grandis and Pseudoboletia maculata are first to the mainland by seamounts, which probably contribute to
records from Trindade, whereas Diadema ascensionais, the species richness on these islands. At about 250 km apart
Eucidaris tribuloides and Echinometra lucunter are recorded and with peaks at 50 –120 m of depth, the six major sea-
from Martin Vaz for the first time. The nine holothuroid mounts act as stepping stones between the mainland and
species reported herein are all first records from Trindade. the islands for shallow-water species (Leal & Bouchet, 1991;
That all 16 echinozoan species known from the TMV also Peterson & Stramma, 1991; Floeter & Gasparini, 2000;
occur in at least one of the oceanic islands of the south Pinheiro et al., 2015; Anker et al., 2016). Even though these
Atlantic (SPP, Atol das Rocas, Fernando de Noronha, seamounts cannot support species restricted to very shallow
Ascension Island, Saint Helena Island, Cape Verde habitats, the connectivity between the coast and TMV was
Archipelago, and islands of the Gulf of Guinea) is an indicator possibly facilitated by the low sea level during the
http://dx.doi.org/10.1017/S0025315416001569
http://dx.doi.org/10.1017/S0025315416001569
Table 2. Geographic distribution of echinozoan species so far known from the tropical southern Atlantic oceanic islands. Abbreviations and symbols as following: ASC, Ascension Island; BR, continental coast and shelf
of Brazil, including Abrolhos Archipelago; FN, Fernando de Noronha Archipelago; MV, Martin Vaz Archipelago; NEA, North-east Atlantic (continental coast and shelf of West Africa, from Morocco to Equatorial
Guinea, Canary Islands, Cape Verde Archipelago, São Tomé and Prı́ncipe Island, and West European coast); NWA, North-west Atlantic (North Carolina to Florida and Texas, Gulf of Mexico, Caribbean Sea,
Bahamas and Bermuda); SPP, São Pedro and São Paulo Archipelago; SEA, South-east Atlantic (continental coast and shelf of West Africa, from Gabon to South Africa); STH, Saint Helena Island; TI, Trindade
Island; UA, continental coast and shelf of Uruguay and Argentina (including Falkland/Malvinas Islands). X, Previously known record; ∗ , New record; ?, Record to be confirmed.
Taxa TI MV FN SPP BR UA NWA ASC STH SEA NEA References
Echinoidea Brissus unicolor (Leske) X X ? X Mortensen (1951), Pawson (1978)

Centrostephanus besnardi Bernasconi X Bernasconi (1955b)
Cidaris rugosa (H.L. Clark) X X Mortensen (1928)
Clypeaster oliveirai Krau X Krau (1952); Brito (1968)
Coelopleurus floridanus A. Agassiz X ? Pawson (1978)
∗
Diadema ascensionis Mortensen X X X X X Pawson (1978), Gondim et al. (2013), present paper
Eucidaris tribuloides clavata Mortensen ? X X Pawson (1978); Edwards & Lubbock (1983)
∗ ∗
Eucidaris tribuloides tribuloides (Lamarck) X X X X X X Rathbun (1879), Mortensen (1928), present paper
∗
Echinometra lucunter lucunter (Linnaeus) X X X X X X Rathbun (1879), Mortensen (1943a), Pawson (1978), present paper
Echinometra lucunter polypora Pawson X X Pawson (1978)
Echinoneus cyclostomus Leske X X Mortensen (1948a), Pawson (1978)
Heliophora orbiculus (Linnaeus) X X X Mortensen (1948b), Pawson (1978)
∗
Leodia sexiesperforata (Leske) X X X Bernasconi (1947), Mortensen (1948b), present paper
Lytechinus variegatus (Lamarck) X X X Rathbun (1879), Mortensen (1943b)
echinozoans from the brazilian oceanic islands

∗
Plagiobrissus grandis (Gmelin) X X Mortensen (1951), present paper
Pseudoboletia atlantica H.L. Clark X X Mortensen (1943b), Pawson (1978)
∗
Pseudoboletia maculata Troschel X Mortensen (1943b), present paper
Tretocidaris spinosa Mortensen X X Mortensen (1928)
Tripneustes ventricosus (Lamarck) X X X X ? ? X X Mortensen (1943b), Lima-Verde (1969), Pawson (1978), present paper
Taxa from South Atlantic oceanic islands
∗ ∗
Holothuroidea Actinopyga agassizii (Selenka) X Pawson et al. (2010), present paper
∗
Chiridota rotifera (Pourtalès) X X Hendler et al. (1995), Gondim et al. (2008), present paper
∗
Euapta lappa (Muller) X X X ? Pawson (1978), Hendler et al. (1995), present paper
∗ ∗
Isostichopus badionotus Selenka X X X X Hendler et al. (1995), present paper
∗
Holothuria (C.) pseudofossor Deichmann X X Deichmann (1930), Tommasi (1974), present paper
∗
Holothuria (H.) grisea Selenka X X X ? Pawson (1978); Hendler et al. (1995); present paper
∗ ∗
Holothuria (H.) manningi Pawson X Pawson (1978), present paper
∗
Holothuria (S.) surinamensis Ludwig X X Hendler et al. (1995), present paper
Holothuria (P.) sanctori Delle Chiaje X X Pawson (1978)
∗ ∗
Holothuria (T.) arenicola Semper X X X X ? Pawson (1978), Hendler et al. (1995), present paper
31
Pleistocene glaciations (Andrade et al., 2003; Coimbra & with new records, description of a new species of Synalpheus and
Carreño, 2012). This connectivity has most probably influ- remarks on zoogeographical patterns in the oceanic islands of the trop-
enced species spillover and, not surprisingly, resulted in the ical southern Atlantic. Zootaxa 4138, 1–58.
low level of endemism of echinoids and holothuroids in the Bernasconi I. (1947) Distribucion geografica de Los Equinoideos
TMV. True, a moderate to high level of endemism has been Argentinos. Anales de la Sociedad Argentica de Estudos Geográficos
documented for other marine organisms (e.g. prosobranch 8, 97–114.
gastropods, reef fishes, sponges; Leal, 2000; Moraes &
Bernasconi I. (1955a) Equinoideos y asteroideos de la collección del
Muricy, 2007; Floeter et al., 2008; Pinheiro et al., 2015 – Instituto Oceanográfico de la Universidad de San Pablo. Boletim do
but see Coimbra & Carreño, 2012 (ostracods)), but whether Instituto Oceanográfico 6, 51–77.
this is only a reflection of sample size (e.g. McClain, 2007) is
yet to be clarified. Bernasconi I. (1955b) Una nueva especie de diadematideo tropical.
Neotropica 6, 92.
The Trindade and Martin Vaz Oceanic Archipelago has
rather harmonic faunal elements (Holdgate, 1960; Carlquist, Bernasconi I. (1957) Otra nueva especie de Asteroideo Brasileiro.
1974; Whittaker et al., 2014), harbouring only two species Neotropica 111, 33–34.
with disharmonic geographic distribution patterns:
Borrero-Pérez G.H., Benavides-Serrato M. and Diaz-Sanchez C.M.
Holothuria (Thymiosycia) arenicola (also known from the (2012) Equinodermos del Caribe colombiano II: Echinoidea
Eastern Pacific) and Pseudoboletia maculata (also occurring y Holothuroidea. Serie de Publicaciones Especiales de Invemar 30,
in the Indo-West Pacific). 250 pp.
Brito I.M. (1962) Ensaio de catálogo dos Equinodermas do Brasil.
Universidade do Brasil Faculdade Nacional de Filosofia. Centro de
ACKNOWLEDGEMENTS
Estudos Zoológicos 13, 1 –10.
We are grateful to the anonymous reviewers. We thank Erica Brito I.M. (1968) Asteróides e equinóides do estado da Guanabara e adja-
Clites (UCMP), Jon Norenburg (USNM) and Rich Mooi cências. Boletim do Museu Nacional, Rio de Janeiro 260, 1 –51.
(CAS) for granting us access to the echinoderm collections, Brito I.M. (1971) Contribuição ao conhecimento dos Equinodermas da
and Lara Guimarães (MZUSP) for technical support with Ilha da Trinidade, Brasil. Arquivos do Museu Nacional 54, 261–265.
SEM images.
Brown J. (2015) Marine life of St. Helena. Newbury: Pisces Publications,
220 pp.
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LM thanks FAPESP for providing financial support by way of Carvalho J. (1950) Resultados cientı́ficos do cruzeiro do “Baependı́” e do
a doctoral fellowship (201318202-9). MT thanks CNPq “Vega” à I. da Trindade. Peixes. Boletim do Instituto Paulista
(403940/2012-5) for financial support to the ProTrindade Oceanografico 1, 97–133.
Project and a technical grant given to JBM, and CNPq
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Journal of the Marine Biological Association of the United Kingdom, page 1 of 35.

# Marine Biological Association of the United Kingdom, 2016

Echinoidea and Holothuroidea

Submitted 12 June 2016; accepted 18 October 2016

INTRODUCTION shallow-water species and is directly reﬂected in the taxo-

RESULTS Order Aspidochirotida

E. insularis E. lucunter E. mathaei E. oblonga E. vanbrunti E. viridis

Plates in peristomial membrane Scattered Scattered No/rare No/rare Scattered Scattered

examined material long, widened at edges, fused perradially (Figure 10H).

columns of pore-pairs bowed, closing at end; poriferous zone biological notes

distribution Class HOLOTHUROIDEA

biological notes Synapta rotifera Pourtalès, 1851, p. 15.

and radial pieces, perforated at top for insertion of radial remarks

Mülleria agassizii Selenka, 1867, pp. 311.

examined material 294); 12.5 m, 31.1.2012, 1 spm 17 cm long (MZUSP 297);

tables with undeveloped knobs, and the teeth at the top of

Holothuria (Halodeima) grisea Selenka, 1867

Holothuria grisea Selenka, 1867, p. 328, pl. 18.

Holothuria arenicola Semper, 1868, p. 61, pls. 20, 30, 35.

16.7.2013, 1 spm 13 cm long (MZUSP 324). Ponta Noroeste, distribution

955); 15 m, 29.4.2014, 1 spm 20 cm long (MZUSP 965);

additional material examined

Taxa TI MV FN SPP BR UA NWA ASC STH SEA NEA References

Echinoidea Brissus unicolor (Leske) X X ? X Mortensen (1951), Pawson (1978)

echinozoans from the brazilian oceanic islands

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