A catalogue of the scaleworm genus Lepidonotus (Polynoidae, Polychaeta) from South America, with two new records for Brazilian waters.

The genus Lepidonotus is the largest in number of species within the Polynoidae, with more than 70 described species and subspecies. A catalogue of 18 nominal species and subspecies of Lepidonotus from South America is provided, with valid names, synonyms and original citations. Redescriptions and illustrations of two species based on new specimens collected along the littoral of the State of Paraíba, northeastern Brazil are included. Lepidonotus carinulatus and Lepidonotus natalensis are reported for the first time for Brazilian waters. A comparative table of characters for all reported species and subspecies of Lepidonotus from South America is provided.

Introduction

The scaleworm belongs to the family , and contains more than 70 described species (Read and Fauchald 2015). They have been found from the intertidal to the bathyal zones, in diverse marine environments (Day 1967, Fauchald 1977, Salazar-Silva 2006, Wehe 2006).

Leach (1816) established the genus for the species Montagu (1808), which had been described earlier. This author did not provide identifying characters of the genus, which led to subsequent confusion in the literature, causing many synonyms. Later, Seidler (1923) made a very extensive review of the , presenting descriptions and keys to more than 50 species of , but there are almost no illustrations to supplement the descriptions, nor is it clear from the text which specimens or types were examined (Wehe 2006). However, Wehe (2006) clarified that this paper is invaluable in providing base-line data and access to the literature on lepidonotid genera.

has a short body with 26 segments, is dorsoventrally flattened, and subrectangular in the cross-section. The prostomium is bilobed, extending anteriorly into the ceratophores of the terminally-attached lateral antennae. The antennae and cirri are smooth. A facial tubercle is present; the buccal segment is with or without nuchal fold. Twelve pairs of elytra are present on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21 and 23; elytra are with or without tubercles and papillae. The notopodia small or vestigial, unidentate notochaetae are short. The neuropodia are large, with or without an acicular lobe; the neurochaetae are stout, long, with subdistal spines and unidentate or occasionally bidentate tips (Fauchald 1977, Amaral and Nonato 1982, Ruff 1995, Wehe 2006).

In this paper, a catalogue of the genus from South America is provided, and and are redescribed, collected in the intertidal region of the State of Paraíba, northeastern Brazil. These two species are reported for the first time for Brazilian waters.

Material and methods

The species accounts in the catalogue are given alphabetically. Each account contains author, publication year, number of pages, figures, types, and deposition numbers, together with the abbreviation of the museum or institution in which the type material is deposited, type locality with coordinatesand geographical distribution, when available. In some cases, remarks on taxonomic status of somes species are included. Synonyms are listed chronologically. A comparative table for all reported species and subspecies from South American is provided (Table 1).

Table 1.

Comparative table for all species and subspecies of the genus from South America.

	Characters
Species	Prostomium	Palps	Nuchal nodules	Posterior part of the elytra	Microtubercule	Macrotubercule	Notochaetae	Neurochaetae
Lepidonotus arenosus	Rounded	Smooth	Present	Numerous posterior papillae	Small papillae egg-shaped	Bigger central papillae egg-shaped	Thin capillaries in all chaetigers	Stout, curved, with spinous row, tip bidentate
Lepidonotus brasiliensis	Rounded	Smooth	Absent	Numerous thin papillae	Small papillae lunar crate-shaped	Big papillae mushroom-shaped	Spinous ciliated capillaries	Stout, curved, with spinous rows, tip unidentate
Lepidonotus brasiliensis laevis	Rounded	Smooth	Absent	Numerous papillae in posterior elytra	Small papillae lunar crate-shaped	Big papillae mushroom-shaped	Thin capillaries, with 3 ciliated filaments	Stout, straight, smooth blade, unidentate
Lepidonotus caeruleus	Longer than wide	Papillate	Absent	Numerous short papillae on posterior portion	Conical	Absent	Longer capillaries shorter, curved and dentate; shorter capillaries long and barbed	Long, strongly curved,with spinous row, tip bidentate
Lepidonotus carinulatus	Rounded to hexagonal, buccal segment with two nuchal nodules	With 8 rows of longitudi nal papillae	Present	Papillae along external edge	Small warty tubercles	Carinate tubercles scattered	Spinous, superior row slender, serrate on convex side	Stout, falcate, subdistally thickened, few rows of spines, tip bidentate
Lepidonotus crosslandi	Rounded, with posterior nuchal fold	Smooth	Absent	Numerous short posterior papillae	Small and thin tubercles	Absent	Fairly numerous capillaries	Stout, slightly curved tip, with a row of spines, bidentate in two chaetigers
Lepidonotus crosslandi peruana	Rounded	Smooth	Present	Numerous short posterior papillae	Small and thin tubercles	Absent	Fairly numerous capillaries	Stout, tip slightly curved, with rows of strong spines, bidentate in all chaetigers
Lepidonotus furcillatus	Rounded	With numerous papillae	Absent	Digitiform papillae with central dark spot	Small and conical	Small, nodular	Curved, short, smooth	Stout, weakly ornamented, bidentate
Lepidonotus hupferi	Rounded to hexagonal	Smooth	Absent	Fringe covering posterior part of elytra	Rounded, surface rough	Warty and pointed	Capillaries spinous in all chetigers	Stout, spinous rows with fine teeth, bidentate
Lepidonotus margaritaceus	Convex, posterior rounded	Smooth	Absent	Slender papillae on posterior border	Small granules	Absent	Short, numerous rows of small spines	Stout, numerous rows of distal spines, unidentate
Lepidonotus natalensis	Slightly hexagonal	With 8 rows of Longitud-inal papillae		Long digitiform external papillae and group of infero-central papillae	Crown-like tip or without tip	Crown-like tip in central region	Spinous, superior row slender, tips fine, pointed	Stout, falcate, subdistally slightly thickened, few rows of spines, tip unidentate
Lepidonotus nesophilus	Convex, posterior rounded	smooth	Absent	Absent	Hemisferical, spinous	Long spines	Spinous capillaries in all chaetigers	Stout, with spinous rows, bidentate
Lepidonotus panamensis	Largely hidden, convex lobe projecting over posterior half	A few papillae		Numerous posterior papillae and tulf of long central papillae	Numerous small spines on centre	Excavate disk, cap covered by many spines	Chaetae shorter, falcate	Long, stout, with curved tip, unidentate
Lepidonotus savygni	Rounded	Smooth	Absent	Slender short posterior papillae	Numerous small warty-shaped spines	Warty-shaped	Spinous capillaries in all chaetigers	Long, stout, tip curved, unidentate
Lepidonotus sublevis	Slightly longer than wide	Papillate	Absent	Additional posterior papillae	Conical, without areola	Absent	Spinous capillaries in all chaetigers	Stout, few spinous rows, tip slightly hooked
Lepidonotus teinuetosus	Oblong, no cephalic peaks	Smooth	Absent	Slender posterior papillae	Blunt or somewhat warty tips	Very small, conical or globular	Slender, numerous rows of small spines and capillary tips tapering	Stouter, falcate, numerous distal spine rows, d mostly unidentate, few spines minutely bidentate
Lepidonotus tomentosus	Rounded	Smooth	Absent	Numerous large posterior papillae	Small warty tips	Numerous warts on central elytra	Spinous capillaries in all chaetigers;	Stout, with spinous row, bidentate
Lepidonotus variabilis	Rounded	Smooth	Absent	Numerous large posterior papillae	Small, cylindrical	Absent	Spinous capillaries in all chaetigers	Stout, with spinous rows, secondary teeth small, bidentate

Specimens were collected by handpicking during low tides from the intertidal region (0.0–0.2 m) and by snorkeling to a depth of up to 5 meters along the coast of the state of Paraíba. Specimens were fixed in formaldehyde (10% in seawater), and PageBreakPageBreakPageBreaklater rinsed with fresh water and transferred to 70% ethanol. General structures were observed with Stereomicroscope Olympus Nikon SMZ800. Chaetae and elytrae were observed with an Olympus BX41 compound microscope. All illustrations were drawn using a camera lucida, and photographs were edited in Photoshop, PhotoScape and CorelDraw X7. Specimens are deposited in the ‘Coleção de Invertebrados Paulo Young’, Departamento de Sistemática e Ecologia da Universidade Federal da Paraíba’, Brazil.

The nomenclature of appendages and other characteristics of polynoids mentioned in this paper follow Tebble and Chambers (1982), Hanley and Burke (1991), Ruff (1995), Imajima (1997), and Wehe (2006).

The following abbreviations are used in the text:

AMNH

American Museum of Natural History, New York

BMNH

The Natural History Museum, London, Great Britain (formerly British Museum of Natural History)

LIPY

Laboratório de Invertebrados Paulo Young, Paraíba, Brazil

MNHN Poly Type

type collection, Museum National d’Histoire Naturelle, Paris

NCB

Naturalis, The Netherlands Centre for Biodiversity, Leiden

PMNH

Peabody Museum of Natural History, Yale University

POLY-UFPB

Coleção de do Laboratório de Invertebrados Paulo Young

SSM, Naturhistoriska

Rijsmuseet, Stockholm

USNM

National Museum of Natural History, Smithsonian Institution, Washington D.C.

ZUEC-POL

Polychaete Collection, Zoological Museum of the State University of Campinas, São Paulo

ZMB

Naturhistorisches Forschungsinstitut, Museum für Naturkunde, Zentralinstitutder Humboldt-Universität zu Berlin, Germany

ZMH

Zoologisches Institut und Museum der Universität Hamburg, Germany

Results

Family Malmgren, 1867

Leach, 1816

Type species.

Montagu, 1808

Leach 1816, Hanley and Burke 1991, Ruff 1995, Chambers and Muir 1997, Barnich and Fiege 2003.

Diagnosis.

Body short, arched, with 26 segments. Bilobed prostomium extending anteriorly into ceratophores of terminally-attached lateral antennae. Antennae and cirri smooth. Facial tubercle present; buccal segment with or without nuchal fold. Twelve PageBreakpairs of elytra on segments 2, 4, 5, 7.... 21 and 23. Notopodia small or vestigial; unidentate notochaetae short, slender, spinose, or notochaetae capillaries sometimes present. Neuropodia large, with or without acicular lobe; neurochaetae stout, long, with subdistal spines and unidentate or occasionally bidentate tips.

Remarks.

The genus contains more than 70 species distributed worldwide (Ruff 1995). However, only 18 species and subspecies have been reported for South America, including the two new records described here.

Ehlers, 1901b

Ehlers, 1901b: 253–254 (Calbuco, Chile),

Holotype.

NCB Verm. 3643.

Type locality.

Calbuco, Chile.

Distribution.

Chile and New Caledonia.

(Quatrefages, 1866)

Quatrefages, 1866: 246–247 (Bahia, Brazil).-

-

Syntype of .

Poly Type 78.

Bahia, Brazil.

Known only from the type-locality in Bahia.

Rullier & Amoureux, 1979

Rullier & Amoureux, 1979: 150, fig. d. (Brazil).-

Syntype.

Poly Type: 1304.

Bahia, Brazil.

This species occurs along the Brazilian littoral.

Solís-Weiss et al. (2006) considered only the species, however, in Read and Fauchald (2015), the status as subspecies is considered valid.

Kinberg, 1856

Kinberg, 1856: 384 (off Rio de Janeiro-Brazil),

.-

.-

.-

.-

.-

.-

ZUEC-POL 2919.

off Rio de Janeiro-Brazil.

Western Pacific from Japan. Eastern Pacific from Alaska to California. Southwestern Atlantic from Paraíba to São Paulo. 7.3–932 m (Figure 1).

Figure 1.

Map showing the distribution of genus in South America: 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 .

Biology.

The species occurs in colonies of the sponge (Bowerbank, 1875), in colonies of the bryozoan (Waters, 1879), on tubes of sabellariid (Moore, 1906), in tubes of cirratulid , with the barnacle (Darwin, 1854), and the mussel . In rhodolith beds.

presents a wide distribution. Futher studies are required to enable us to understand if it is a cryptogenic species, because there are no studies to show that it represents a species complex, and its origin was not determined.

Grube, 1869

Figures 2 , 3

Figure 2.

A Anterior end of showing the dark ring in the antennae and tentacular cirrus (anterodorsal view of the prostomium) B Anterior end showing the dorsal anus in the last chaetigerous segment. Scale bars: 1 mm (A, B). (p, palp; ma, median antennae; la, lateral antennae; tc, tentacular cirri; pr, prostomium; nn, nuchal nodules; dc. Dorsal cirrus; e, elytra; pl, papillae; a, anus; ac, anal cirri.

Figure 3.

A pair of nuchal nodules on 2nd segment in dorsal anterior view B right parapodia of 4th segment, dorsal view C notochaetae, dorsal view D neurochaetae, ventral view E first right elytra, dorsal view F–G macrotubercle H microtubercles I right elytra of 5th segment J macrotubercle. Scale bars: 1 mm (A); 0,5 mm (B–D), 0.05 mm (E–J).

—

.—

. —

With two nuchal nodules and without nuchal folds; black pigmentation on antennae, cirrophores and tentaculophores; elytra with dark pigmentation; elytral surface reticulate, with oval to rounded macro- and microtubercles, anterior ones flattened, smooth or carinate, central and posterolateral ones warty; margin with fringing papillae; neurochaetae bidentate.

Description.

Body elongated, flattened dorsoventrally, subrectangular in cross-section; 2 mm in length, including palps and pygidial cirri; 26 chaetigerous segments, and pygidium (Figure 2a–b). Prostomium bilobed, rounded to hexagonal, lateral antenna with terminal insertion (Figure 3a). Two pairs of eyes; anterior pair dorsolateral, near widest portion of prostomium, posterior pair near posterior end of prostomium, converging towards midline, buccal segment without nuchal fold, but with pair of nuchal nodules (Figure 3a–b). Median and lateral antennae, tentacular and dorsal cirri with two dark rings (Figure 2a), both having subdistal swelling, culminating abruptly in sharp point; ceratophores cylindrical, median antenna larger than lateral antennae. Pair of palps, slightly smaller than median antenna, culminating in thin point, with 8 longitudinal rows of papillae.

Tentacular segment with two pairs of cylindrical tentaculophores, with three prostomial chaetae on anterodorsal bases. Buccal cirri larger than ventral cirri, with cylindrical cirrophores. Pharynx with nine pairs of papillae and two pairs of maxillae. Facial tubercle present. Dorsal cirri with same coloration as median antenna, larger than ventral cirri, with cylindrical cirrophores.

Parapodia biramous (Figure 3b), prechaetal lobe quadrate or subtriangular, postchaetal lobe short and subtriangular, both with acicula; short notopodia on anterodorsal side of neuropodia. Notochaetae spinous, superior row slender, serrated on convex side (Figure 3c). Neuropodia large, truncate, distally cleft with prechaetal lobe slightly longer than poschaetal lobe. Neurochaetae stouter than notochaetae, falcate, subdistally thickened, with several rows of spines, distal spines usually much larger than basal ones, and bidentate tips with small secondary tooth (Figure 3d).

Twelve pairs of elytra, covering dorsum entirely, with dark-brown pigmentation; pairs until last chaetiger segment following order: 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21 and 23; long papillae along external edge. First three pairs of elytrae more ornate, with tubercles (Figure 3e); on 11th and 12th pair with ornamentation similar to that in first pair. Most carinate macrotubercles in central region (Figure 3f–g), surrounded by microtubercles (Figure 3h); after 4th or 5th pair (Figure 3i), small warty tubercles give impression of smooth elytra (Figure 3j).

Nephridial papillae starting from chaetiger 7, with peduncular aspect. Short ventral cirri with thin tip; pair of long anal cirri with same coloration as median antenna; dorsal anus in last chaetiger segment (Figure 2b).

Habitat.

Recorded between the intertidal and 60 m; elsewhere reported down to 200 m (Hartmann-Schröder and Hartmann 1991, Wehe 2006).

ZMB 1071.

Bohol, Philippines.

Red Sea, Arabian Sea: Socotra Archipelago, Gulf of Oman, Arabian Gulf. Elsewhere: Indo-West Pacific: Madagascar, Sri Lanka, Indonesia, Philippines, Hong Kong, South China Sea, Australia, New Caledonia, Japan (Willey 1905, Fauvel 1953, Hanley and Burke 1991, Hanley 1992, Imajima 1997, Barnich et al. 2004).

New records: Coast of Paraíba, Brazil: Pedra da Galé, Pitimbú (07°28'17"S, 34°47'26"W), POLY–UFPB 1501; Rio Mamanguape (06°48'44"S, 34°54'48"W), POLY–UFPB 1502, 1503.

Zenetos et al. (2010) assigned as an exotic species with an origin in the Indo-Pacific/Red Sea. Its establishment success in the Mediterranean is questionable, because its description, based on local specimens, was insufficient. It is an exotic species in Brazilian waters with casual establishment success; because only the present records are known, it is presumed to be non-established in the Mediterranean area (Zenetos et al. 2010).

Monro, 1928

Monro, 1928: 553–555, figs 1–4 (Callao and Bahia Independencia, Peru).-

USNM 54378.

Balboa and Taboga, Panama.

El Salvador to Peru.

Hartmann-Schröder, 1962b

.-

ZMH.

Callao and Bahia de Independencia, Peru

Presently known only from Peru. 0–112.8 m.

Ehlers, 1901b

Ehlers, 1901b: 254–255 (Tumbes and Cavancha, Chile),

Syntypes.

NCB 3682, 3701.

Tumbes and Cavancha, Chile.

Western Pacific from Australia and New Zealand. East Pacific from Colombia to Chile and Galapagos Islands.

Augener, 1918

Augener, 1918: 133–136, pl. 2: figs 7–11 (Gold Coast, western Africa, Gana).-

Holotype

. Fauvel, 1950: 345 (Senegal). ZMH V-530

Gold Coast, western Africa, Ghana.

Eastern Atlantic from Senegal and Cape Verde. Western Pacific from New Caledonia and Australia. Eastern Pacific from Mexico to Ecuador. 10–30 m.

presents a wide distribution. Futher studies are required to enable us to understand if it is a cryptogenic species, because there are no studies to show that it is a species complex, and its origin was not determined

Kinberg, 1856

Kinberg, 1856: 383 (Guayaquil, Ecuador);

.-

.

Guayaquil, Ecuador.

Known only from the type material from Ecuador.

Day, 1951

Figures 4 , 5

Figure 4.

A Anterior end of showing the pigmentation of antennae and tentacular cirri B dorsal anus in the 23rd segment chaetigerous, dorsal posterior view. Scale bars: 1 mm (A–B). (p, palp; ma, median antennae; la, lateral antennae; tc, tentacular cirri; pr, prostomium; nn, nuchal nodules; dc, dorsal cirrus; e, elytra; pl, papillae; a, anus; ac, anal cirri.

Figure 5.

A anterodorsal view of prostomium B second segment showing the anterior peak C right parapodia of 4th segment, dorsal view D notochaetae, dorsal view E neurochaetae, ventral view F first right elytra, dorsal view G macrotubercle H microtubercles I right elytra of 6th segment J macrotubercle K group of papillae. Scale bars: 1 mm (A); 0,5 mm (B–D); 0.05 mm (B–D).

Day, 1951; fig. 1 e–l [removed from synonymy with

Fauvel, 1927: 414 [not Gravier, 1902].

Without nuchal fold; some elytrae with group of papillae in center, dark pigmentation and small tubercles after 4th or 5th pair, giving impression of smooth elytra; elytra margin and surface with long slender, digitiform papillae; notochaetae partially threadlike; neurochaetae unidentate.

Body elongate, flattened dorsoventrally; with 26 chaetigerous segments, 2 mm in length, including palps and pygidial cirrus, 2 mm in width, including chaetae (Figure 4a–b). Dorsum and sides of parapodia pigmented black. Prostomium bilobed, rounded, slightly hexagonal, black pigmentation at base of ceratophores (Figure 5a). Lateral antenna with terminal insertion; two pairs of eyes, anterior pair dorsolateral, near widest portion of prostomium, posterior pair near base of prostomium, converging towards midline (Figure 5b). Median and lateral antennae, tentacular and dorsal cirri with two dark rings; first more elongated than second, both having subdistal swelling, culminating abruptly in sharp point; ceratophores cylindrical, median antenna larger than lateral antennae. Palps paired, same length as median antenna, culminating almost abruptly into thin point, with 8 longitudinal rows of papillae.

Tentacular segment with two pairs of cylindrical tentaculophores, with three prostomial chaetae on anterodorsal bases. Buccal cirri larger than ventral cirri, with cylindrical cirrophores. Pharynx with 9 pairs of papillae and 2 pairs of maxillae. Facial tubercle present. Dorsal cirri with same coloration as median antenna, larger than ventral cirri, with cylindrical cirrophores.

Parapodia biramous (Figure 5c), prechaetal lobe quadrate or subtriangular, postchaetal lobe short and subtriangular, both with acicula; short notopodia on anterodorsal side of neuropodia. Notochaetae spinous, superior row slender, partially thread-like, with fine, pointed tips (Figure 5d). Neuropodia large, truncate, distally cleft, with prechaetal lobe slightly longer than poschaetal lobe. Neurochaetae stouter than notochaetae, falcate, subdistally slightly thickened, with few rows of spines below slightly bent, blunt, unidentate tips (Figure 5e).

Twelve pairs of elytra covering dorsum entirely, with dark or dark-brown pigmentation; pairs until last chaetiger segment following order: 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21 and 23. First three pairs more ornate, with macro and microtubercles (Figure 5f-h); these tubercles located more posteriorly on elytra after 4th or 5th pair, giving impression of smooth elytra; on 11th and 12th pairs ornamentation similar to that in first pairs. Most of macrotubercles on central region, surrounded by microtubercles (Figure 5i-j). Very long digitiform papillae along external edge, and group of papillae near inferior base of elytra or sometimes next to center (Figure 5k).

Nephridial papillae starting from chaetiger 7, with peduncular aspect. Short ventral cirri with thin tip; one pair of short anal cirri with same coloration as median antenna; anus dorsal in 23rd chaetigerous segment (Figure 4b).

Intertidal zone (algae, rhodoliths) to 5 m, from the coast of Paraíba.

BMNH 1961.16.1

Natal, South Africa.

Suez Canal, Arabian Sea: Karachi; Port Edward, Natal, South Africa, Indian Ocean (Day 1951; Wehe 2006).

New records.

Barra de Camaratuba, Mataraca, Paraíba, Brazil (06°36'14.17"S, 34°57'48.79"W).

(POLY-UFPB 1504), adult from Barra de Camaratuba, Mataraca (06°36'14.17"S, 34°57'48.79"W), Paratypes (POLY-UFPB 1505, 1506), Prainha, Baía da Traição (06°41'23.77"S, 34°55'48.47"W), Paraíba, Brazil.

presents a wide distribution. Futher studies are required to enable us to understand if it is a cryptogenic species, because there are no studies to show that it is a species complex, and its origin was not determined.

Chamberlin, 1919

Chamberlin, 1919: 75–78, pl. 4: figs 1–7; pl. 5: fig. 13 (Chatham Island, Galapagos Archipelago).-

USNM 19400.

Chatham Island, Galapagos Archipelago

Eastern Pacific from Mexico to Galapagos.

Hartman, 1939

Hartman, 1939: 44–46, pl. 6: figs 70–77 (Pacific Panama, 27–91m).

.-

USNM 47981.

Pacific, Panama.

Eastern Pacific: Panama Bay. Southwestern Atlantic from Bahia and Espírito Santo. Brazil, 27–91 m.

Grube, 1856

Grube, 1856: 45 (Callao, Peru);

.-

.-

Callao, Peru.

Eastern Pacific from Peru. Western Atlantic from the West Indies.

Verrill, 1873

-

.-

Treadwell, 1939a: 3, figs 10–12 (Texas).

.-

[not (Linnaeus, 1758); [=

3-5.

New England (Northeastern USA).

Western Atlantic from Massachusetts to Colombia, with one record for Antarctica. Eastern Pacific from Mexico. From 2 to10 m.

This species is commensal with the hermit crabs (Bosc, 1802), (Stimpson, 1860), (Benedict, 1892) Say, 1818, and Say, 1818. It was found on the lower surface of the sea pansy, Kölliker, 1872 (Martin and Britayev 1998).

(Gravier, 1902)

Gravier, 1902: 222–226, figs 228–231, pl. 8: figs 123–125 (Djibouti, Gulf of Tadjoura, Gulf of Aden).-

.-

Day, 1951: 9, fig. 1e–l (Port Edward, Natal, South Africa).-

.-

Holotype of .

Poly type 263.

Djibouti, Gulf of Tadjoura, Gulf of Aden.

Southwestern Atlantic from Bahia to São Paulo. Eastern Atlantic from Mediterranean to South Africa. Indian Ocean, Madagascar, Persian Gulf, and Red Sea. Western Pacific from Russia to Australia. 0–0.5 m.

Zenetos et al. (2010) and Çinar (2013) assigned as an exotic species for the Mediterranean Sea, with an origin in the Indo-Pacific/Red Sea. Its establishment success is uncertain, because it was recorded only once. We consider this species as an exotic species for the Brazilian coast.

(Grube, 1856)

Grube, 1856 (Punta Arenas, Chile).-

Grube, 1878.

.-

Punta Arenas, Chile.

Eastern Pacific from Costa Rica and Chile. Western Atlantic from French Guyana.

There is material in the USNM from the Galapagos Islands.

Webster, 1879

Webster, 1879: 205–208, pl. 1: figs 6–11; pl. 2: figs 12–14 (Virginia).-

not .-

Virginia Coast, North American.

Western Atlantic from Massachusetts to Colombia; 0.2–18 m.

Associated with the oyster (Sowerby, 1871), and with the sponge Schmidt, 1862. Food of the winter flounder, (Walbaum, 1792). Lives among hydroids and sponges.

Discussion

Herein, all information on members of the genus found around South American coasts in the literature have been gathered, and additional data on two species collected in northeastern Brazil is provided. Eighteen species and subspecies are catalogued from South America, and three of them represent endemic taxa: and the subspecies are endemic for Bahia, Brazil, while is endemic from Ecuador. The subspecies is very similar to the species . However, only a detailed review can confirm if the two taxa are synonyms.

The species , , , , and have a broad distribution, and have been reported from several countries. According to Zenetos et al. (2010) and Çinar (2013), the species and are exotic species in the Mediterranean and have possibly originated in the Indo-Pacific region/Red Sea. Their introduction area was through the Mediterranean and Sea of Marmara. Despite the broad distributions of , and , more studies are needed to indicate if they may possibly represent exotic species, their possible areas of introduction, and into which ecological category they belong accordPageBreaking to the classification scheme of Çinar (2013). The possible origin of is Natal, South Africa, and it was possibly reported from the Suez Canal, Arabian Sea, and Karachi, Pakistan. It is herein reported from the southwest Atlantic, in the state of Paraíba. was first described off Rio de Janeiro, and was reported from the Pacific coast of North America and Japan. was first described from the Eastern Atlantic from Senegal and Cape Verde, and later reported for the Pacific from New Caledonia, Australia, Mexico, and Ecuador. The remaining species present narrow distributions spanning few countries.

For some records essential features are not presented clearly, such as the ornamentation of the elytra, or the shape of the prostomium and chaetae. Some characters, such as form of nuchal folds, pigmentation of the antennae, and dorsal cirri, are not mentioned for the species and . We are left with the view that species are very similar and difficult to distinguish. Therefore, revisionary studies of are needed to establish whether cryptic species occur.