Catalogue
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Catalogue
A catalogue of the scaleworm genus Lepidonotus (Polynoidae, Polychaeta) from South America, with two new records for Brazilian waters
expand article infoJosé Eriberto De Assis, Rafael Justino de Brito§, Martin Lindsey Christoffersen§, José Roberto Botelho de Souza
‡ Universidade Federal da Paraíba, Recife, Brazil
§ Centro de Ciências Exatas e da Natureza - UFPB, João Pessoa, Brazil
Open Access

Abstract

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.

Keywords

Benthos, Annelida, catalogue, new record, distribution

Introduction

The scaleworm Lepidonotus belongs to the family Polynoidae, 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 Lepidonotus for the species Aphrodita clava 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 Lepidonotinae, presenting descriptions and keys to more than 50 species of Lepidonotus, 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.

Lepidonotus 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 Lepidonotus from South America is provided, and L. carinulatus and L. natalensis 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).

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

Characters
Species Prostomium Palps Nuchal nodules Posterior part of the elytra Microtubercule Macrotubercule Notochaetae Neurochaetae
L. 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
L. 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
L. 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
L. 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
L. 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
L. 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
L. 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
L. furcillatus Rounded With numerous papillae Absent Digitiform papillae with central dark spot Small and conical Small, nodular Curved, short, smooth Stout, weakly ornamented, bidentate
L. 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
L. 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
L. 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
L. nesophilus Convex, posterior rounded smooth Absent Absent Hemisferical, spinous Long spines Spinous capillaries in all chaetigers Stout, with spinous rows, bidentate
L. 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
L. 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
L. 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
L. 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
L. 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
L. 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 later 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 Polychaetatype 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 Polychaeta 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 Polynoidae Malmgren, 1867

Lepidonotus Leach, 1816

Type species

Aphrodita clava 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 pairs 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 Lepidonotus 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.

Lepidonotus arenosus Ehlers, 1901b

Lepidonotus arenosus Ehlers, 1901b: 253–254 (Calbuco, Chile), 1901a: 49, pl. 2: figs 9–12 (Chile).-Wesenberg-Lund 1962: 15.-Hartwich 1993: 80.-Pleijel 2007: 179 (New Caledonia).

Holotype

NCB Verm. 3643.

Type locality

Calbuco, Chile.

Distribution

Chile and New Caledonia.

Lepidonotus brasiliensis (Quatrefages, 1866)

Polynoe brasiliensis Quatrefages, 1866: 246–247 (Bahia, Brazil).-Solís-Weiss et al. 2004: 358.

Lepidonotus brasiliensis-Seidler 1924: 37.-Amaral and Nonato 1982: 25.-Salazar-Vallejo 1996: 15.-Amaral et al. 2013.

Syntype of Polynoe brasilienis

MNHN Poly Type 78.

Type locality

Bahia, Brazil.

Distribution

Known only from the type-locality in Bahia.

Lepidonotus brasiliensis laevis Rullier & Amoureux, 1979

Lepidonotus braziliensis laevis Rullier & Amoureux, 1979: 150, fig. d. (Brazil).-Solís-Weiss et al. 2004: 358.

Syntype

MNHN Poly Type: 1304.

Type locality

Bahia, Brazil.

Distribution

This species occurs along the Brazilian littoral.

Remarks

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

Lepidonotus caeruleus Kinberg, 1856

Lepidonotus caeruleus Kinberg, 1856: 384 (off Rio de Janeiro-Brazil), 1858: 13–14, pl. 4: fig. 16, pl. 10, fig. 51.-Baird 1865: 183.-Grube 1876: 61.-Seidler 1924: 69.-Hartman 1939: 108–109.-Nonato and Luna 1970a: 63 (Alagoas, 19–35 m); 1970b: 66–67, pl. 1: figs 8–14 (Alagoas, 19–35 m).-Rullier and Amoureux 1979: 152 (Alagoas and Bahia).-Morgado and Amaral 1981: 93 (São Paulo, in bryozoan colonies).-Amaral and Nonato 1982: 25.-Salazar-Vallejo 1996: 15.-Duarte and Nalesso 1996: 142 (São Paulo, in colonies of Zygomycale parishii).-Morgado and Tanaka 2001: 178 (São Paulo; in colonies of Schizoporella errata).-Berlandi et al. 2012: 282 (off Espírito Santo State, rhodolith beds).-De Assis et al. 2012: 17 (Paraíba).-Cunha et al. 2013: 146 (off Bahia).

Polynoe caerulea.-Quatrefages 1866: 224.

Lepidonotus caeloris.-Moore 1903: 412–414, pl. 23: fig. 12 (Japan, 115–280 m), 1906: 546–547, pl. 36: figs 36–37 (Alaska); 1908: 331 (Alaska and Pacific Canada), 1910: 333–334 (California).-Essenberg 1918: 184 (Alaska to California, 53–932 m).-Hartman and Reish 1950: 5 (Oregon).-Díaz-Castaneda and Rodríguez-Villanueva 1998: 12 (Pacific Mexico).

Polynoë caelora.-Izuka 1912: 23–25, fig. (Japan).

Lepidonotus caelorus.-Treadwell 1914: 182 (California).-Chamberlin 1918: 174 (California).-Berkeley 1923: 213 (Pacific Canada).-Hartman 1939: 44, 1944: 244 (California).-Rioja 1941: 680 (Pacific Mexico), 1947: 199 (Pacific Mexico).-Berkeley and Berkeley 1942 (Alaska).-Pettibone 1953: 15–16, pl. 1: figs 1–8; pl. 2: figs 9–19 (Washington and Oregon 7.3–256 m, with Volsella modiolus, Balanus nubilis, on tube of Neosabella [as Sabellaria] cementarium, in calcareous tubes of Dodecaceriapacifica).-Reish 1968: 100 (California).

Lepidonotus coelorus.-Treadwell 1937: 141 (California).-Berkeley and Berkeley 1942: 187 (Pacific Canada), 1948: 9–10, figs 6–7 (Pacific Canada).-Pequegnat 1964: 278 (California).

Lepidonotus caerulus.-Berkeley and Berkeley 1941: 20 (California).

Holotype

ZUEC-POL 2919.

Type locality

off Rio de Janeiro-Brazil.

Distribution

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 Lepidonotus in South America: 1 L. arenosus 2 L. brasiliensis 3 L. brasiliensis laevis 4 L. caeruleus 5 L. carinulatus 6 L. crosslandi 7 L. crosslandi peruana 8 L. furcillatus 9 L. hupferi 10 L. margaritaceus 11 L. natalensis 12 L. nesophilus 13 L. panamensis 14 L. savignyi 15 L. sublevis 16 L. tenuisetosus 17 L. tomentosus 18 L. viriabilis.

Biology

The species occurs in colonies of the sponge Zygomycale parishii (Bowerbank, 1875), in colonies of the bryozoan Schizoporella errata (Waters, 1879), on tubes of sabellariid Neosabellaria cementarium (Moore, 1906), in tubes of cirratulid Dodecaceriapacifica, with the barnacle Balanus nubilis (Darwin, 1854), and the mussel Volsella modiolus. In rhodolith beds.

Remarks

Lepidonotus caeruleus 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.

Lepidonotus carinulatus Grube, 1869

Figures 2, 3

Polynoe (Lepidonotus) carinulataGrube 1869; Grube 1878: 26–27, pl. 3: figs 2–2 b.

Lepidonotus carinulatus.—Willey 1905: 248–249, pl. 1: figs 7–11, Fauvel 1911: fig. 1, 1918, 1919a, Augener 1922: figures 3–3b, Seidler 1923, Fauvel 1933, Wesenberg-Lund 1949, Fauvel 1955, Mohammad 1971, Day 1975: figs 2 g–k, Amoureux et al. 1978, Hanley and Burke 1991: fig 20, Imajima 1997: figs 45–46, Rasheed and Mustaquim 2003: figs 7–8, Barnich et al. 2004.

Lepidonotus tenuisetosus. — Mohammad 1971: 288, Gravier 1902.

Diagnosis

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.

Figure 2.

A Anterior end of Lepidonotus carinulatus 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.

Lepidonotus carinulatus 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).

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).

Syntype

ZMB 1071.

Type locality

Bohol, Philippines.

Distribution

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.

Remarks

Zenetos et al. (2010) assigned L. carinulatus 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).

Lepidonotus crosslandi Monro, 1928

Lepidonotus crosslandi Monro, 1928: 553–555, figs 1–4 (Callao and Bahia Independencia, Peru).-Hartman 1939: 109, pl. 5, figs 62–69, 1959.-Fauchald and Reimer 1975: 80 (Panama).-Reimer 1976: 242 (Pacific Panama).-Fauchald 1977: 6 (Pacific Panama).-Von Prahl et al. 1979 (Pacific Colombia).-Cruz et al. 1980: 92 (Ecuador).-Laverde-Castillo 1986 (Colombian Pacific).-Rivera 2008: 23 (El Salvador).

Holotype

USNM 54378.

Type locality

Balboa and Taboga, Panama.

Distribution

El Salvador to Peru.

Lepidonotus crosslandi peruana Hartmann-Schröder, 1962b

Lepidonotus crosslandi.-Hartman 1939: 42–43, pl. 5: figs 62–69 (Peru, 0–112.8m). [not Lepidonotus crosslandi Monro 1928]

Lepidonotus crosslandi peruana Hartmann-Schröder 1962b: 109–112, pl. 1: fig. 3; pl. 2: figs 1–2, 4 (Peru).-Hartman 1965: 9.-Romero et al. 1988: 138 (Peru).

Holotype

ZMH.

Type locality

Callao and Bahia de Independencia, Peru

Distribution

Presently known only from Peru. 0–112.8 m.

Lepidonotus furcillatus Ehlers, 1901b

Lepidonotus furcillatus Ehlers, 1901b: 254–255 (Tumbes and Cavancha, Chile), 1901a: 52–53, pl. 2, figs 1–8 (Chile).-Augener 1913: 102–103 (Australia).-Seidler 1924: 67–69.-Hartman 1939: 42, pl. 5: figs 57–58 (Colombian Pacific, Ecuador, Pacific Panama, and Chatham Island, New Zealand, 55–101m).-Wesenberg-Lund 1962: 15.-Day 1975: 171 (Australia).-Laverde-Castillo 1986.-Blake 1991 (Galapagos).-Hartwich 1993: 101.-Salazar-Vallejo and Londoño-Mesa 2004: 49.

Syntypes

NCB 3682, 3701.

Type locality

Tumbes and Cavancha, Chile.

Distribution

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

Lepidonotus hupferi Augener, 1918

Lepidonotus hupferi Augener, 1918: 133–136, pl. 2: figs 7–11 (Gold Coast, western Africa, Gana).-Seidler 1924: 69–70.-Day 1934: 20 (Angola).-Hartman 1939: 43, pl. 6: figs 78–82 (Ecuador, Pacific Panama, and Pacific Mexico, 3.7–22 m).-Steinbeck and Rickets 1941 (Pacific Mexico).-Rioja 1947: 198–199, figs 1–8 (Pacific Mexico), 1962: 141 (Pacific Mexico).-Tebble 1955: 80 (Australia).-Fauvel and Rullier 1957: 48 (Senegal), 1959a: 146 (Gulf of Guinea), 1959b: 500 (Senegal and Cape Verde).-Day 1967: 37 (southern Africa).-Intes and Le Loeuff 1975: 275 (Ivory Coast, 10–30 m).-Cruz et al. 1980: 92 (Ecuador).-Batisda-Zavala 1993 (Pacific Mexico).-Hernández-Alcantara et al. 2003: 6 (Pacific Mexico).-Salazar-Vallejo and Londoño-Mesa 2004: 49.-Pleijel 2007: 179 (New Caledonia).

Holotype

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

Type locality

Gold Coast, western Africa, Ghana.

Distribution

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

Remarks

Lepidonotus hupferi 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

Lepidonotus margaritaceus Kinberg, 1856

Lepidonotus margaritaceus Kinberg, 1856: 383 (Guayaquil, Ecuador); 1858: 11–12, pl. 3: fig. 12; pl. 10: fig. 49.-Baird 1865: 182.-Grube 1876: 62.-Seidler 1924: 33–34.-Hartman 1948: 23, pl. 3: figs 1–3.

Polynoe margaritacea.-Quatrefages 1866: 223–224.

Holotype

SSM.

Type locality

Guayaquil, Ecuador.

Distribution

Known only from the type material from Ecuador.

Lepidonotus natalensis Day, 1951

Figures 4, 5

Lepidonotus natalensis Day, 1951; fig. 1 e–l [removed from synonymy with L. tenuisetosus (sensu Day 1967)].

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

Diagnosis

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.

Description

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.

Figure 4.

A Anterior end of Lepidonotus natalensis 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.

Lepidonotus natalensis 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).

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).

Habitat

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

Holotype

BMNH 1961.16.1

Type locality

Natal, South Africa.

Distribution

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.

Remarks

Lepidonotus natalensis 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.

Lepidonotus nesophilus Chamberlin, 1919

Lepidonotus nesophilus Chamberlin, 1919: 75–78, pl. 4: figs 1–7; pl. 5: fig. 13 (Chatham Island, Galapagos Archipelago).-Hartman 1939: 38–39, pl. 7: figs 83–95 (Galapagos Islands, 22–73 m) [not Tenacatitla Bay, Mexico = undescribed species].-Rioja 1941: 680 (Mexican Pacific), 1962: 141 (Mexican Pacific).-Fauchald and Reimer 1975: 80 (Panama).-Fauchald 1977: 6 (Pacific Panama).-Van der Heiden and Hendrickx 1982 (Mexican Pacific).-Blake 1991 (Galapagos).-Hernández-Alcantara et al. 2003: 6.-Salazar-Vallejo and Londoño-Mesa 2004: 49.-Salazar-Silva 2006: 151.

Holotype

USNM 19400.

Type locality

Chatham Island, Galapagos Archipelago

Distribution

Eastern Pacific from Mexico to Galapagos.

Lepidonotus panamensis Hartman, 1939

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

Lepidonotus panamensis.-Hartman 1959: 90.-Rullier and Amoureux 1979: 152 (Bahia).-Amaral and Nonato 1982: 25 (Espírito Santo).

Holotype

USNM 47981.

Type locality

Pacific, Panama.

Distribution

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

Lepidonotus savignyi Grube, 1856

Polynoe Savignyi Grube, 1856: 45 (Callao, Peru); 1876: 61.-Seidler 1924: 88.

Lepidonotus savignyi.-Baird 1865: 183.-Augener 1925 (West Indies).-Salazar-Vallejo and Eibye-Jacobsen 2012: 1398.

Polynoe Savignyi.-Quatrefages 1866: 225.

Type locality

Callao, Peru.

Distribution

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

Lepidonotus sublevis Verrill, 1873

Lepidonotus sublevis-Verrill 1973.-Verrill and Smith 1874: 581, pl. 10: fig. 42 (Savin Rock, near New Haven, and Vineyard Sound, Massachusetts); Verrill 1882: 300, pl. 4: fig. 2; pl. 6: fig. 3 (New England).-Dorner 1877: 66 (Massachusetts).-Wood 1885: 532 (New England).-Heilprin 1888 [1906]: 6, pl. 8: fig. 8 (New Jersey).-Andrews 1891: 278 (North Carolina).-Drowne 1896: 74 (Massachusetts).-Summer et al. 1913: 618 (Massachusetts).-Pratt 1923: 285.-Hartman 1942: 22–23, figs 7–12, 1944; 1945: 10 (North Carolina); 1951: 17–18 (Gulf of Mexico), 1954: 413 (Gulf of Mexico).-Ferguson and Jones 1949 (Virginia).-Behre 1950: 11 (Louisiana).-Hedgpeth 1950: 75 (Texas; on lower surface of Renilla muelleri [as R. mülleri]).-Pratt 1951: 328.-Wells 1961: 247 (North Carolina).-Pettibone 1963: 18–19, fig. 3e (Massachusetts to Florida, in gastropod shells with Pagurus pollicaris).-Wells and Gray 1964 (North Carolina).-Smith 1964: 76 (Massachusetts).-Yentsch et al. 1966: 117 (Massachusetts).-Calder and Brehmer 1967 (Virginia).-Conner et al. 1972: 1503.-Boesch 1973: 230 (Virginia).-Day 1973: 6 (Beaufort, North Carolina, 0–100 m).-Fotheringham 1976: 574 (Texas; comensal with Clibanarius vittatus and Pagurus pollicaris).-Gardiner 1976: 86, fig. 1f–g (North Carolina; associated with Clibanarius vittatus, Pagurus impressus, P. longicarpus, and P. pollicaris).-Kinner and Maurer 1978: 217 (Delaware Bay).-Dueñas 1981 (Caribbean Colombia), 1999 (Caribbean Colombia).-Stainken 1984: 100 (New York).-Weston 1984: 27 (Gulf of Mexico).-Ismail 1985: 384 (New Jersey).-Ibarzabal 1986: 2 (Cuba, 2 m); 1989b: 2–3, figs. 1a–f (Cuba).-Steimle and Caracciolo-Ward 1989: 148 (New York).-Dauer 1991 (comensal of Clibanarius vittatus, Pagurus annulipes, P. longicarpus, and P. pollicaris).-Bastida-Zavala 1995: 14 (Pacific Mexico).-Pollock 1998: 197 (northeast North America).-Posey et al. 1998: 151 (Gulf of Mexico).-Sagasti et al. 2000: 481 (Chesapeake Bay).-McDermott 2001: 624 (New Jersey; obligate symbiont of Pagurus longicarpus).-Baéz and Ardilla 2003: 102 (Caribbean Colombia, to 10 m).-Williams 2003: 17 (commensal of Pagurus longicarpus).-Williams and McDermott 2004: 28.-Struck et al. 2008: 631.-Fauchald et al. 2009: 768 (Gulf of Mexico).-Gobin 2010: 151 (Trinidad and Tobago).-Struck and Halanych 2010: 271.-Piquet et al. 2011: 417 (Antarctica).-Golombek et al. 2013: 315.

Lepidonotus sublaevis.-Seidler 1924: 41.-Gambi et al. 1997: 1055.

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

Lepidonotus squamatus.-Warren 1942: 45 (Louisiana).-Cowles 1930: 341.

[not Lepidonotus squamatus (Linnaeus, 1758); [= Lepidonopsis humilis (Augener 1922)].

Holotype

PMNH 3-5.

Type locality

New England (Northeastern USA).

Distribution

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

Biology

This species is commensal with the hermit crabs Clibanarius vittatus (Bosc, 1802), Pagurus annulipes (Stimpson, 1860), P. impressus (Benedict, 1892) P. longicarpus Say, 1818, and P. pollicaris Say, 1818. It was found on the lower surface of the sea pansy, Renilla muelleri Kölliker, 1872 (Martin and Britayev 1998).

Lepidonotus tenuisetosus (Gravier, 1902)

Euphione tenuisetosa Gravier, 1902: 222–226, figs 228–231, pl. 8: figs 123–125 (Djibouti, Gulf of Tadjoura, Gulf of Aden).-Fauvel 1911: 368 (Persian Gulf).-Solís-Weiss et al. 2004: 13.

Lepidonotus tenuisetosus.-Fauvel 1919a: 330–332 (Madagascar), 1927: 411 (Suez Canal); 1933: 15 (India); 1953: 36–37, fig. 14c–f (India).-Seidler 1924: 25–27 (Red Sea).-Day 1934: 20 (Madagascar); 1953: 400 (South Africa); 1962: 632 (Madagascar); 1967: 82, fig. 1.14a–e (South Africa, Mozambique, Madagascar, and Red Sea); 1974 (Mozambique).-Monro 1934: 358 (China).-Aziz 1938 (Pakistan).-Okuda 1940: 4–6, fig. 2 (Japan).-Day and Morgans 1956 (South Africa).-Kalk 1958: 232 (Mozambique).-Macnae and Kalk 1958 (Mozambique).-Imajima and Hartman 1964: 27 (Japan).-Tampi and Rangarajan 1964: 100 (Andaman Islands).-Wu 1968: 27–28 (China).-Achari 1969: 31 (Andaman Islands).-Mohammad 1971: 288 (Kuwait).-Ben-Eliahu 1972: 190, 195 (Suez Canal).-Santhakumari 1973: 179 (India).-Hartman 1974 (Indian Ocean).-Peyrot-Clausade 1974 (Australia).-Sarma 1974: 158 (India).-Rullier and Amoureux 1979: 152 (Bahia).-Buzhinskaya et al. 1980: 228 (Indo-Pacific).-Soota et al. 1980 (Andaman and Nicobar Islands), 1981 (India).-Amaral and Nonato 1982: 25 (Espírito Santo).-Shin 1982 (China), 1998 (China), 2000 (China).-Uschakov 1982: 106, 107, pl. 29: figs 1–8 (Russia).-Kirkegaard 1983: 194, fig. 1.14a–e (Sierra Leone and French Guinea, 15–65 m).-Ansari et al. 1986: 363 (India).-Cantone 1987: 75, 80 (Somalia).-Gil et al. 1987: 1 (Korea).-Srikrishnadhas et al. 1987 (India).-Palpal-Latoc 1990 (Philippines); 1994: 67 (Philippines).-Hanley 1992: 366–367 (China, 0–0.5 m).-Hong and San 1993 (Vietnam).-Wang and Huang 1994: 4 (China).-Misra 1995: 93 (India).-Mustaquim 1997: 221 (Pakistan).-Wu et al. 1997 (China).-Che et al. 1999 (China).-Kumar 2000: 441 (India).-Misra and Chakraborty 2000: 219 (India).-Paxton and Chou 2000: 210 (China).-Bellan 2001: 224 (Europe).-Pillai 2001: 122 (India).-Sato 2001 (Japan).-Wehe and Fiege 2002: 113 (Arabian Peninsula).-Barnich and Fiege 2003: 86, fig. 44 (Mediterranean Sea).-Rasheed and Mustaquim 2003: 70–72, fig. 12 (Pakistan).-Barnich et al. 2004: 300–301 (China).-Khan and Murugesan 2005: 116 (India).-Zenetos et al. 2005: 73 (casual alien in Mediterranean), 2010 (introduced into Mediterranean).-Galil 2006 (Suez Canal), 2007: 301 (alien in Israel).-Kato et al. 2006: 30 (Japan).-Paiva 2006: 268 (Central Brazilian Plataform).-Wehe 2006: 107–109, fig. 24a–l (Arabian Peninsula).-Pleijel 2007: 179 (New Caledonia).-Li and Ping 2008 (China).-Çinar 2009: 2286, fig. 2a (Turkish Mediterranean), 2013: 1259 (introduced from Red Sea to Mediterranean).-Naeini and Rahimian 2009: 59–60 (Persian Gulf and Gulf of Oman).-Sukumaran and Devi 2009: 1443 (India).-Li et al. 2010: 110 (China).-Amaral and Nallin 2011: 556 (São Paulo).-Çinar 2013: 264 (alien species in Turkey).-Rizzo et al. 2011: 133 (São Paulo).-Wang 2011: 746 (China).-Amaral et al. 2013: 453 (São Paulo).-Rajasekaran and Fernando 2012: 3 (Andaman and Nicobar Islands).-Katsanevakis et al. 2012: (alien species in European waters).-Kazmi and Naushaba 2013: 253 (Pakistan).

Lepidonotus natalensis Day, 1951: 9, fig. 1e–l (Port Edward, Natal, South Africa).-Wehe 2006: 101–103, fig. 21a–j (Arabian Peninsula).-Naeini and Rahimian 2009: 55–59 (Gulf of Oman).

Lepidonotus cf. tenuisetosus.-Yan and Huang 1993: 133 (China).

Holotype of Euphione tenuisetosa

MNHN Poly type 263.

Type locality

Djibouti, Gulf of Tadjoura, Gulf of Aden.

Distribution

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.

Remarks

Zenetos et al. (2010) and Çinar (2013) assigned L. tenuisetosus 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.

Lepidonotus tomentosus (Grube, 1856)

Polynoe tomentosa Grube, 1856 (Punta Arenas, Chile).-Quatrefages 1866: 225–226.

Polynoe (Lepidonotus) pilosella Grube, 1878.

Lepidonotus tomentosus.-Fauvel 1919b: 472–473, fig. 1a–d (French Guyana), 1923 (French Guyana).-Hartman 1959.-Perkins and Savage 1975: 21.-Dean 2004 (Costa Rica).

Type locality

Punta Arenas, Chile.

Distribution

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

Remarks

There is material in the USNM from the Galapagos Islands.

Lepidonotus variabilis Webster, 1879

Lepidonotus variabilis Webster, 1879: 205–208, pl. 1: figs 6–11; pl. 2: figs 12–14 (Virginia).-Andrews 1891: 278 (North Carolina; among hydroids and sponges).-Hoagland 1919: 572 (Puerto Rico, 18 m).-Seidler 1924: 70–72.-Treadwell 1939b: 185 (Puerto Rico).-Warren 1942: 45 (Louisiana).-Hartman 1945: 10 (Florida), 1951: 18 (Gulf of Mexico), 1954: 413–414 (Gulf of Mexico).-Behre 1950: 11 (Louisiana).-Pearse and Williams 1951 (North and South Carolina).-Renaud 1956: 3, fig. 2 (Bahamas).-Rioja 1958: 221 (eastern Mexico).-Wells 1961: 247 (North Carolina).-Wells and Gray 1964 (North Carolina).-Ebbs 1966: 493–496, fig. 2a–h (Florida).-Forbes 1966: 278 (Florida; associated with Cryptostrea [as Ostrea] permollis and Stelletta grubii).-Dauer 1973: 193 (Gulf of Mexico; in sponge).-Day 1973: 6 (North Carolina, to 18 m).-Rullier 1974: 20 (Cuba; in sponges).-Gardiner 1976: 86, fig. 1k–n (North Carolina).-MacPhee 1978: 15 (Massachusetts; food of Pseudopleuronectes americanus).-Rodríguez-Gómes 1979: 27 (Caribbean Colombia); 1988 (Caribbean Colombia).-Tagatz et al. 1982: 134 (Alabama).-Ibarzábal 1986: 2 (Cuba, 3 m).-San Martín et al. 1986: 6–7 (Cuba).-Perkins 1998: 85 (Florida).-Dueñas 1999 (Caribbean Colombian).-Baéz and Ardilla 2003: 102 (Caribbean Colombian, 0.2–2 m).-Fauchald et al. 2009: 768 (Gulf of Mexico).-Gobin 2010: 5 (Trinidad and Tobago).

not Lepidonotus variabilis.-Treadwell 1939b: 341. [=Lepidonotus subleavis Verrill, 1874]

Type locality

Virginia Coast, North American.

Distribution

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

Biology

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

Discussion

Herein, all information on members of the genus Lepidonotus 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: L. brasiliensis and the subspecies L. brasiliensis brevis are endemic for Bahia, Brazil, while L. margaritaceus is endemic from Ecuador. The subspecies L. brasiliensis brevis is very similar to the species L. brasiliensis. However, only a detailed review can confirm if the two taxa are synonyms.

The species L. caeruleus, L. carinulatus, L. hupferi, L. natalensis, and L. tenuisetosus have a broad distribution, and have been reported from several countries. According to Zenetos et al. (2010) and Çinar (2013), the species L. carinulatus and L. tenuisetosus 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 L. caeruleus, L. hupferi and L. natalensis, 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 according to the classification scheme of Çinar (2013). The possible origin of L. natalensis 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. Lepidonotus caeruleus was first described off Rio de Janeiro, and was reported from the Pacific coast of North America and Japan. Lepidonotus hupferi 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 Lepidonotus brasiliensis and L. panamensis. We are left with the view that species are very similar and difficult to distinguish. Therefore, revisionary studies of Lepidonotus are needed to establish whether cryptic species occur.

Acknowledgments

We heartily thank Dr Mathan Magesh (Project Associate Conservation of Coastal and the Marine Resources Division National Centre for Sustainable Coastal Management, Chennai, India) for the corrections of the English grammar, and for contributions to the manuscript. To all members of the Laboratory of Invertebrates Paulo Young (LIPY) for their work in assembling the collection of marine invertebrates, and in particular to Dr. Carmen Alonso Samiguel for her dedication in the preservation of this collection. We thank Dr Christopher Glasby (MAGNT), and the anonymous referees for suggestions to improve this paper. We acknowledge CNPq and FACEPE for pos-doctoral scholarship to J.E. De Assis and the productivity grant to M.L. Christoffersen.

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