Lumbrineridae (Annelida: Polychaeta) from the Grand ...

J. Mar. Biol. Ass. U.K. (2001), 81, 599^621 Printed in the United Kingdom

Lumbrineridae (Annelida: Polychaeta) from the Grand Caribbean region with the description of six new species Luis F. Carrera-Parra El Colegio de la Frontera Sur, Unidad Chetumal. Apartado Postal 424, Chetumal QR. 77000, Mexico. E-mail: [email protected]

Twenty-seven species of Lumbrineridae are characterized; six are described as new: Lumbrinerides uebelackerae sp. nov., Lumbrineris perkinsi sp. nov., L. salazari sp. nov., Ninoe vargasi sp. nov., N. wardae sp. nov. and Lysarete raquelae sp. nov. Twenty-one species from the Grand Caribbean region are recognized herein and characterized based upon type and non-type materials: Augeneria bidens, Eranno bilabiata comb. nov., Lumbrinerides crassicephala, L. dayi, L. jonesi, Lumbrineriopsis cf. paradoxa, Lumbrineris £oridana, L. cf. latreilli, L. nonatoi, L. paucidentata, Ninoe nigripes, Scoletoma branchiata comb. nov., S. candida comb. nov., S. ernesti, S. robusta comb. nov., S. tenuis, S. testudinum comb. nov., S. treadwelli comb. nov., S. verrili, S. parvapedata comb. nov. and S. elongata comb. nov. A key to identify all lumbrinerid species recorded from the Grand Caribbean region is included.

INTRODUCTION The former idea about uniformity in external characters in lumbrinerids (simplicity and reduction of many morphological parts) resulted in a simpli¢ed generic system for the family, lumping in only three or four genera all of the known species (Carrera-Parra, 2001). From Hartman (1944) and Fauchald (1970) some genera included in Lumbrineridae (Augeneria Monro, Cenogenus Chamberlin, Eranno Kinberg and Scoletoma de Blainville) were put under synonymy of Lumbrineris de Blainville. In the 1970s, Orensanz (1973) employing mainly characters of the mandibles and maxillary apparatus began a new taxonomic approach in lumbrinerids and emended Augeneria and erected Lumbrineriopsis and Lumbrinerides. This approach was suggested previously by Crossland (1924), but was not followed by early taxonomists. Mohammad (1973) erected Kuwaita and Fauchald (1977) suggested that Kuwaita could be placed into a separate family. Levenstein (1977) erected Paraninoe, which has been recently been shown to be a junior synonym of Cenogenus (Carrera-Parra, 2001). Since 1979, three genera have been established (Arabellonereis Hartmann-Schro«der, Abyssoninoe Orensanz and Lumbricalus Frame) and two others have been emended (Eranno Kinberg and Scoletoma de Blainville). Currently 13 genera of Lumbrineridae are recognized. Nonetheless, the taxonomy of lumbrinerid polychaetes is confusing, a large part of the described species have been regarded as being of cosmopolitan distribution, which cannot be explained by the reduced natural dispersal abilities of the species; all studied species being brooders (Wilson, 1991). Rather, this wide spread distribution is a result of the indiscriminate use of faunas or guides from di¡erent regions, resulting in the incorporation of names to the local fauna of species that are not necessarily present in that locality. Journal of the Marine Biological Association of the United Kingdom (2001)

In this contribution, the taxonomic status and characterization of species described from the Grand Caribbean region were analysed based upon type and additional material from the region. Six new species are described, seven species are transferred to di¡erent genera, and all 27 species are characterized and illustrated. A key to identify lumbrinerid species recorded from the Grand Caribbean region is also included.

MATERIALS AND METHODS Type materials were borrowed from the collections of the National Museum of Natural History (USNM), Smithsonian Institution, Washington, DC; the Museum of Comparative Zoology (MCZ), Harvard University, Cambridge; the Peabody Museum of Natural History (YPM), Yale University, New Haven; the American Museum of Natural History (AMNH), New York; Los Angeles County Museum of Natural History, Allan Hancock Foundation Polychaete Collection (LACMAHF); the Zoologisches Museum (ZMB), Berlin and the Natural History Museum (BMNH), London. The nontype material were obtained from the collections of the USNM; the Museum of Marine Invertebrates (MMI), University of Miami, Miami; Centro de Colecciones Naturales del Instituto de Oceanolog|¨ a (IOH), La Habana, Coleccion de poliquetos, Instituto de Ciencias del Mar y Limnologia, UNAM (CP-ICML-UNAM) and Coleccion de Referencia de ECOSUR-Chetumal (ECOSUR), Chetumal. An antero-dorsal dissection was made to extract the maxillary apparatus, it was mounted dorsally and then ventrally on a slide to analyse details of both the maxillae and mandible under a compound microscope; the maxillary apparatus was later returned to its original position.

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Lumbrineridae from the Grand Caribbean region

The nomenclature of maxillary apparatus is based on Orensanz (1990); a true tooth is herein considered to exit only if a pulp cavity is present. The designation for type localities of syntypes described here follow the recommendations of article 73.2.3 of the International Code of Zoological Nomenclature (2000). The measurements were standardized to setiger 10; they are abbreviated as L10 for length through setiger 10, and W10 for width at setiger 10 (excluding parapodia). Maxillae as M followed by a Roman numeral indicates the position in the maxillary apparatus from dorsal to ventral side. A correlation analysis between some characters and body size (standardized to the length up to setiger 10) were made to clarify their relationship. Illustrations were made using camera lucida. Records of lumbrinerids from the Grand Caribbean follow the lists provided by Perkins & Savage (1975) and Salazar-Vallejo (1996). The original spelling of Lumbrineris de Blainville was modi¢ed to Lumbriconereis or Lumbrinereis and several species were described in each of them (Frame, 1992); however, both are regarded herein as spelling variants and not as di¡erent genera. Thus, following the article 51.3.1 (International Code of Zoological Nomenclature, 2000) the author names are not modi¢ed since they are being retained in Lumbrineris.

RESULTS AND DISCUSSION Family LUMBRINERIDAE Schmarda, 1861 Augeneria Monro, 1930 emended Orensanz, 1973 Augeneria bidens (Ehlers, 1887) Figure 1A^ D Lumbriconereis bidens Ehlers, 1887:103, plate 31, ¢gures 7^17. Augeneria bidensöUebelacker 1984:41.17^41.18, ¢gures 14A^H. Material examined

Syntypes of Lumbriconereis bidens Ehlers: MCZ1239 (1) south of Dry Tortugas, Florida (24808'N 82851'W) Blake 1877^1878, station 43, 620 m; MCZ760 (1) o¡ Coral Reef, Florida, BIBB Gulf Stream Exploration 1868^1870, station 7, 642 m; MCZ750 (1) o¡ Carysfort Reef, Florida, BIBB Gulf Stream Exploration 1868^1870, station 2, 214 m; MCZ756 (1) o¡ Bah|¨ a Honda, Cuba (23801'N 83814'W) RV `Blake' 1877^1878, station 23, 348 m. Non-type materials: MMI G823 (1) RV `Gerda', cruise 6709, o¡ Florida (25835'N 80800'W), 07 July 1967, 174 m; MMI G847 (1) RV `Gerda', cruise 6713, o¡ Florida (25851'N 80803'W), 02 August 1967, 169 m; MMI G848 (1) RV `Gerda', cruise 6713, o¡ Florida, (25853'N 80803'W), 02 August 1967, 201m; MMI G854 (4) RV `Gerda', cruise 6713, o¡ Florida (25827'N 80802'W), 25 August 1967, 221m; CP-ICMLUNAM IMCA 2, station 33 (1), 03 October 1988, o¡ Tabasco, Mexico; CP-ICML-UNAM IMCA 2, station 75 (1), 10 February 1988, o¡ Campeche, Mexico. Description

Syntypes all incomplete, MCZ1239 with 83 setigers (L102.9 mm, W101.2 mm), MCZ760 with 73 setigers (L106 mm, W102.3 mm), MCZ750 and MCZ756 in poor condition. Prostomium conical, longer than wide, with a pair of nuchal organs and three small antennae in the nuchal fold. Peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, Journal of the Marine Biological Association of the United Kingdom (2001)

ventrally with a shallow lip; anterior ring 0.75 of total peristomial length. First four parapodia smaller and slightly ventral. Anterior parapodia with short presetal lobe, rounded and digitiform postsetal lobe with in£ated base; in posterior parapodia presetal lobe slightly longer and postsetal lobe digitiform. Postsetal lobe always longer than presetal one. Setae limbate, composite multidentate hooded hooks from setiger 1 to 7^15 and simple multidentate hooded hooks from setigers 8^16 (Figure 1A,B); acicula yellow. Mandibles with short and divergent shaft (Figure 1C); short and stout maxillary apparatus, with four pairs of maxillae (Figure 1D). Maxillary carriers shorter than MI slightly rounded anteriorly; MI forcepslike with bridles well developed; MII with three rounded teeth; MIII and MIV with whitish central and dark peripheral areas. Remarks

The ending of composite multidentate hooded hooks and the beginning of the simple multidentate hooded hooks are size dependent (r0.974, P50.0001).

Discussion

Sol|¨ s-Weiss et al. (1995) recorded this species from Campeche, Mexico, but this record is doubtful.Their specimens lack antennae and the authors reported structures similar to branchial scars on the dorsal parapodial surface, more perceptible in anterior parapodia.

Distribution

North Carolina, Florida, Alabama, Texas, Tabasco and Campeche. Eranno Kinberg, 1865 emended Orensanz, 1990 Eranno bilabiata (Treadwell, 1901) comb. nov. Figure 1E ^ I

Lumbriconereis bilabiata Treadwell, 1901:199, ¢gures 45^46. Lumbrineris bilabiata ö Hartman, 1942a:120. Material examined

Holotype of Lumbriconereis bilabiata Treadwell USNM16015, Mayaguez Harbor, Puerto Rico, 20 January 1899, station (135)6063, Fish Hawk, 135 m. Non-type material: USNM16746 (1), Caribbean Sea, RV Àlbatross', station 2145, 02 April 1884, US Bureau of Fisheries.

Description

Holotype incomplete, with 59 setigers (L102.9 mm, W101.2 mm); USNM16746 with 168 setigers (L10 5.5 mm, W102 mm). Prostomium conical, as long as wide, with a pair of divergent dorsal longitudinal black bands, with a pair of nuchal organs, and an antenna in the nuchal fold (Figure 1E). Peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a shallow lip; anterior ring 0.75 of total peristomial length. All parapodia well developed, but ¢rst three smaller. Anterior parapodia with inconspicuous presetal lobe and postsetal lobe digitiform with in£ated base (Figure 1F); from setiger 20^25 presetal lobe well developed, gradually lengthening on posterior parapodia, becoming digitiform. In median and


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Figure 1. Augeneria bidens (Ehlers) (Syntype MCZ1239). (A) Composite multidentate hooded hook; (B) simple multidentate hooded hook; (C) mandible; (D) maxillary apparatus. Eranno bilabiata (Treadwell) (E^G USNM16746, HÎ Holotype USNM16015). (E) Anterior end; (F) anterior parapodium; (G) posterior parapodium; (H) mandible; (I) maxillary carriers and MI^MII. Lumbrinerides crassicephala (Hartman) (Paratype LACM-AHF0723). (J) Anterior end; (K) anterior parapodium; (L) posterior parapodium; (M) simple bidentate hooded hook; (N) mandible; (O) maxillary apparatus. Lumbrinerides acuta (Verrill) (Syntype USNM13392). (P) Simple bidentate hooded hook. Scale bars: A,B,M&P, 0.01 mm; C&D, 2 mm; E&J, 1 mm; F&G 0.1 mm; H&I 0.2 mm; K,L,N&O, 0.05 mm.

posterior parapodia postsetal lobe digitiform, longer than presetal lobe (Figure 1G). In both specimens, almost all setae broken; limbate setae with black core from the base to near to the tip where they become translucent. By the condition of the material, it was impossible to determine the shape and kind of hooded hooks; acicula black. Mandibles as long as maxillary apparatus, divided about half total length (Figure 1H), maxillary apparatus with ¢ve pairs of maxillae. Maxillary carriers shorter than MI, constricted anteriorly; MI forceps-like with bridles well developed; MII with ¢ve teeth, about half the length of MI, base with a narrow connecting plate (Figure 1I); MIII and MIV each have a prominent single tooth; MV partially fused with MIV. Remarks

The species is herein assigned to Eranno Kinberg based on the maxillary apparatus; the knowledge of setae is Journal of the Marine Biological Association of the United Kingdom (2001)

incomplete and the presence of nuchal antennae must be reviewed in the genus. Distribution

Puerto Rico, Caribbean Sea. Lumbrinerides Orensanz, 1973 Lumbrinerides crassicephala (Hartman, 1965) Figure 1J ^ O

Lumbrineris crassicephala Hartman,1965:117, plate 20, ¢gure C^F. Lumbrinerides crassicephalaöPerkins, 1979:419^420. Lumbrinerides ?acutaöUebelacker,1984:41.11^41.13 ¢gure 8A^G. Lumbrinerides aberrans ö Solis-Weiss et al., 1995:65 (in part). Material examined

Paratype of Lumbrineris crassicephala Hartman LACMAHF0723, o¡ Bermuda slope, 1000 m. Non-type materials:

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USNM75336 (4) o¡ Florida (26816'43''N 83847'40''W), So£a station 12A, 90 m; USNM75337 (1) o¡ Florida (26816'43''N 83847'40''W), So£a station 12D, 90 m. CP-ICML-UNAM IMCA 2, Station 2 (1), 28 October 1990, o¡ Tamaulipas, Mexico. Description

Paratype incomplete with 36 setigers (L103.2 mm, W100.5 mm). Prostomium conical, as long as wide, with a pair of nuchal organs. Peristomium shorter than prostomium, with two rings, anterior ring 0.75 of total peristomial length, separation between rings distinct dorsally and laterally, ventrally with a shallow lip (Figure 1J). First six parapodia smaller; anterior parapodia with presetal lobe short, rounded and postsetal lobe digitiform; both lengthening on posterior parapodia, postsetal lobe always longer than presetal one (Figure 1K,L). Setae limbate, simple bidentate hooded hooks beginning in setiger 6, with proximal tooth directed laterally (Figure 1M); acicula yellow. Mandibles completely fused (Figure 1N), stout maxillary apparatus, with four pairs of maxillae (Figure 1O); maxillary carriers longer than MI, rounded anteriorly and constricted posteriorly; MI forceps-like with bridles well developed, with an accessory tooth near middle; MII with three blunt teeth; MIII and MIV like edentate plates.

Discussion

Hartman (1965) and Ramos (1976) noted only one peristomial ring, but two peristomial rings are present. The species resembles L. acuta (Verrill, 1875) from New England in the maxillary formula; however, a review of the type material of L. acuta (USNM13392, USNM12895) suggested that the species can be separated by the beginning of simple bidentate hooded hooks and by the shape of hooks. Lumbrinerides acuta has both teeth directed distally (Figure 1P) while in L. crassicephala the proximal tooth is directed laterally and the other one is directed distally. Distribution

Bermuda, Florida and Tamaulipas. Lumbrinerides dayi Perkins, 1979 Figure 2A^ F

Lumbrinerides dayi Perkins, 1979:421^423, ¢gure 1C ^ E.; Uebelacker, 1984:41.13, ¢gure 10A^ G. Lumbrinerides aberrans ö Solis-Weiss et al., 1995:65 (in part). Material examined

Holotype of Lumbrinerides dayi Perkins USNM51145, o¡ Beaufort, North Carolina (34834'N 78825'W), 19 April 1965, 20 m; paratypes USNM 55875 (1) o¡ Panama City, Florida, November 1977 (29848'N 86809'W), RV `Columbus Iselin', 47 m; USNM52209 (8), Barceloneta, Puerto Rico (18829'N 66833'W), August 1974, 23^28 m; USNM55598 (16), o¡ Beaufort, North Carolina (34834'N 78825'W), 19 April 1965, 20 m; CP-ICML-UNAM DINAMO 2, station 2 (1), 28 October 1990, o¡ Tamaulipas; CP-ICML-UNAM DINAMO 2, station 4 (1), 29 October 1990, o¡ Tamiahua Lagoon, Veracruz; CP-ICML-UNAM DINAMO 2, station 33 (2), 31 October 1988, o¡ Tabasco; CP-ICML-UNAM IMCA 2, station 8 (3), 21 September 1988, o¡ Tamiahua Lagoon, Veracruz; ECOSUR LUMB1 Journal of the Marine Biological Association of the United Kingdom (2001)

E33COLT-I (1), o¡ Tamiahua Lagoon, Veracruz, Mexico, October 1994, 52 m. Description

Holotype incomplete with 41 setigers (L103 mm, W100.6 mm). Prostomium elongated, longer than wide, with a pair of nuchal organs. Peristomium shorter than prostomium, both rings of similar size, separation between rings distinct dorsally and laterally, ventrally with a shallow lip (Figure 2A). All parapodia short, the ¢rst seven inconspicuous; parapodia with presetal lobe inconspicuous and postsetal one digitiform, longest in posterior parapodia (Figure 2B,C). Setae limbate, simple bidentate hooded hooks from setigers 1^5 with proximal tooth directed laterally (Figure 2D); acicula yellow. Pygidium without anal cirri. Mandibles completely fused (Figure 2E); short maxillary apparatus, with four pairs of maxillae (Figure 2F). Maxillary carriers longer than MI, rounded anteriorly and constricted in the middle; MI forceps-like with bridles well developed, with two teeth in the middle; MII stout with three teeth; MIII and MIV as edentate plates.

Distribution

North Carolina, Florida, Alabama, Veracruz and Puerto Rico. Lumbrinerides jonesi Perkins, 1979 Figure 2G ^ J Lumbrinerides jonesi Perkins, 1979:423^425, ¢gure 2A ^ F. Material examined

Holotype of Lumbrinerides jonesi Perkins USNM57435, Hutchinson Island, Florida (27821'N 80813'W), 03 January 1972, 10.3 m. Paratypes USNM57437 (1), Hutchinson Island, Florida (27821'N 80813'W), September 1972, 11.2 m; USNM57436 (2), Hutchinson Island, Florida (27820.7'N 80812.8'W), 14 March 1972, 10.3 m. Description

Holotype complete with 118 setigers (L104.2 mm, W100.8 mm). Prostomium twice longer than wide, with a pair of nuchal organs. Peristomium shorter, as long as ¢rst setiger; both rings of similar size; separation between rings distinct dorsally and laterally, ventrally with a shallow lip. First ten parapodia smaller; with presetal lobe inconspicuous and postsetal one short, digitiform; longest in median and posterior parapodia (Figure 2G,H). Setae limbate, simple bidentate hooded hooks beginning not before of setiger 23 (Figure 2I); acicula yellow. Pygidium without anal cirri. Mandibles translucent, completely fused; stout maxillary apparatus, with four pairs of maxillae (Figure 2J). Maxillary carriers longer than MI, broad anteriorly and constricted in the posterior end; MI forceps-like with bridles well developed, without accessory tooth; MII short, stout, with edentate distal end and three teeth, the ¢rst one in the middle and the others smaller, in parallel line; MIII and MIV as edentate plates. Distribution

Hutchinson Island, Florida.


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Figure 2. Lumbrinerides dayi Perkins (A^D Holotype USNM511455, E^F Paratype USNM52209). (A) Anterior end; (B) anterior parapodium; (C) posterior parapodium; (D) simple bidentate hooded hook; (E) mandible; (F) maxillary apparatus. Lumbrinerides jonesi Perkins (G^H Holotype USNM57435, I^J Paratype USNM57437). (G) Anterior parapodium; (H) posterior parapodium; (I) simple bidentate hooded hook; (J) maxillary apparatus. Lumbrinerisdes uebelackerae sp. nov. (Holotype USNM67774). (K) Anterior end; (L) anterior end; (M) posterior end; (N) composite multidentate hooded hook; (O) simple bidentate hooded hook; (P) mandible; (Q) maxillary apparatus. Scale bars: A&K, 0.5 mm; B,G,H,L&M, 0.02 mm; D,I&N,O, 0.01 mm; E,F,J,P&Q, 0.1 mm.

Lumbrinerides uebelackerae sp. nov. Figure 2K ^ Q Lumbrinerides sp. A Uebelacker 1984:41.9^41.11, ¢gure 6A ^ H. Material examined

Holotype USNM67774, o¡ Mobile Bay, Alabama, USA, 106 m, MAFLA stationVI-2645 (29835'00''N 87820'02''W), February 1978.

Description

Holotype incomplete with 33 setigers (L102 mm W100.5 mm). Prostomium conical, more than twice longer than wide, with a pair of nuchal organs. Peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a shallow lip; anterior Journal of the Marine Biological Association of the United Kingdom (2001)

ring 3/4 of total peristomial length (Figure 2K). All parapodia well developed, but anterior parapodia shorter than posterior parapodia; postsetal lobe rounded inconspicuous and postsetal digitiform (Figure 2L,M). Setae limbate; hooks of two kinds, composite multidentate hooded hooks beginning from setiger 1^5, with short appendage and with up to four teeth (Figure 2N); simple multidentate hooded hooks from setiger 6 with proximal tooth directed laterally (Figure 2O); acicula black. Mandibles completely fused (Figure 2P); short maxillary apparatus, with four pairs of maxillae (Figure 2Q). Maxillary carriers longer than MI, with broad and rounded anterior end, constricted in the middle; MI forceps-like with bridles well developed, without accessory tooth; MII short and stout, with three blunt teeth; MIII and MIV as edentate plates.

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Discussion

Lumbrinerides uebelackerae sp. nov. resembles L. jonesi Perkins, 1979 (Florida), L. laubieri Miura, 1980 (Mediterranean Sea), L. amoureuxi Miura, 1980 (Mediterranean Sea) and L. acuta sensu Ramos 1976 (Mediterranean Sea) by the presence of MI without accessory tooth. However, Lumbrinerides uebelackerae sp. nov. is the only species in the genus that has composite multidentate hooded hooks and black acicula.

Type locality

Alabama, Gulf of Mexico, USA.

Etymology

The species is named in honour of Joan M. Uebelacker in recognition of her amazing e¡ort to edit the Atlas of Polychaetes from the Gulf of Mexico and her study of lumbrinerid polychaetes from the Gulf of Mexico.

Gender

Feminine.

Lumbrineriopsis Orensanz, 1973 Lumbrineriopsis cf. paradoxa (de Saint-Joseph, 1888) Figure 3A ^ B Lumbrineriopsis paradoxa ö Ramos,1976: 127^129, ¢gures 22^23; Uebelacker, 1984:41.15, ¢gure 12A ^ H. Material examined

ECOSUR LUMB2, E39COLT-I (1) o¡ Lobos Island, Veracruz, Mexico, October 1994, 39 m; M1E11 (1) o¡ Tonala, Veracruz, Mexico (18825'N 94813'W). Description

Both specimens in poor condition. Prostomium conical, longer than wide, with a pair of nuchal organs. Peristomium shorter than prostomium; only one ring is visible, ventrally with a shallow lip (Figure 3A). All parapodia small; with presetal lobe, inconspicuous and postsetal lobe digitiform; both lobes lengthening in median and posterior setigers. Setae limbate and simple bidentate hooded hooks from setiger one (Figure 3B); acicula yellow. Maxillary apparatus

Figure 3. Lumbrineriopsis cf. paradoxa (de Saint-Joseph) (ECOSUR LUMB2). (A) Anterior end; (B) simple bidentate hooded hook. Lumbrineris £oridana Ehlers (Syntype MCZ838). (C) Anterior end in ventral view; (D) anterior parapodium; (E) composite multidentate hooded hook; (F) simple multidentate hooded hook; (G) mandible; (H) maxillary apparatus. Lumbrineris cf. latreilli Audouin & Milne-Edwards (ECOSUR LUMB4). (I) Anterior parapodium; (J) posterior parapodium; (K) composite multidentate hooded hooks with long hood; (L) composite multidentate hook with short hood; (M) simple multidentate hooded hook; (N) mandibles; (O) maxillary apparatus. Scale bars: A&C, 0.5 mm; B, 0.02 mm; D,G^J&N, 0.2 mm; E,F&K^M, 0.01 mm; O, 0.1 mm. Journal of the Marine Biological Association of the United Kingdom (2001)

Lumbrineridae from the Grand Caribbean region with four pairs of maxillae, maxillary carriers longer than MI; MI forceps-like with bridles well developed; MII with 5^6 teeth; MIII unidentate and MIV with up ten teeth.

Remarks

Discussion

Discussion

Lumbrineriopsis paradoxa (de Saint-Joseph, 1888) was described with biannulated peristomium. The material here examined have only one peristomial ring, but because of the poor condition of the material, these specimens cannot be assigned to any species, but they are herein considered close to L. paradoxa.

Distribution

Gulf of Mexico. Lumbrineris de Blainville, 1828 Lumbrineris £oridana Ehlers, 1887 Figure 3C ^ H

Lumbriconereis £oridana Ehlers, 1887:103, plate 30, ¢gures 10^15. Lumbrinereis£oridana ö Treadwell,1921:100^101, ¢gures 365^370. Material examined

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The ending of composite multidentate hooded hooks are size dependent (r0.737, P0.0062). Pettibone (1963) regarded this species as a junior synonym of L. coccinea Renier, 1804 from the Mediterranean Sea. Both species are similar in having composite multidentate hooded hooks in the anterior parapodia and MIII bidentate. However, they can be separated based on the peristomium shape, L. £oridana has muscular lip, transversally separated and by the shape of MIII, compared with Ramos's ¢gure (1976:110, Figure 4). Both species are here considered to be distinct; the records of L. coccinea in the Grand Caribbean should be assigned to L. £oridana.

Distribution

Grand Caribbean region, from Florida to Venezuela. Lumbrineris cf. latreilli Audouin & Milne-Edwards, 1834 Figure 3I ^ O

Syntypes of Lumbriconereis £oridana Ehlers, MCZ838 (2), MCZ841 (1), Key West, Florida, Blake 1877^1878, 2^13 m, Coll. A. Agassiz. Non-type materials, IOH 181 (1) Laguna de Cayo Juan Garc|¨ a, Cuba, 28 January 1986, 2 m, IOH245 (1), Cayo Juan Garc|¨ a, Cuba, July 1988, 2 m, IOH, Me¨dano de la Vela, Archipelago Jardines de la Reina, Cuba, 30 April 1997, 0.5 m, MMI P-709 (1) RV `Pillsbury', cruise 6806, o¡ Isla los Testigos, Venezuela (11825'N 62840'W), 19 July 1968, 71m; MMI P-965 (1) RV `Pillsbury', cruise 6907, Saba Bank, Lesser Antilles (17826'N 63823'W), 20 July 1969, 24 m; MMI P-975 (1) RV `Pillsbury', cruise 6907, o¡ Barbuda, Lesser Antilles (17829'N 61855'W), 21 July 1969, 29 m; ECOSUR LUMB3 E42COLT-I (1), south of Lobos Island, Veracruz, October 1994, 33 m; M1E11 (1) o¡ Tonala, Veracruz, Mexico (18825'N 94813'W); LAG-1 (2) Ria Lagartos, Yucatan, Mexico, 3 m; BP1(1) Bajo Pepito, Isla Mujeres, Mexico, in algae, 4 m; AVE (2) Aventuras, Quintana Roo, Mexico, 2 m; MAJ-1 (1) Majahual, Quintana Roo, 2 m; A70 (1) Xahuaychol, Quintana Roo, 1m.

IOH56 (1) Batabano¨, Cuba, 8 m; ECOSUR LUMB4 E33COLT-I (1) o¡ Tamiahua Lagoon, Veracruz, Mexico, 52 m; CA (1) Caleta, Quintana Roo; BV1(1) Buena Vista, Quintana Roo, Mexico; EL-2772 (1) RV Èdwin Link', north of Isla de la Pasion, Cozumel Island (20839.59'N 86849.64'W); EL-2774 (2) RV Èdwin Link', Cayo Norte, Chinchorro, Quintana Roo, Mexico (18845.63'N 87815.84'W), 60 m; EL-2777 (1) RV Èdwin Link', south of Chinchorro, Quintana Roo, Mexico (18826.02'N 87818.82'W), 66 m; EL-2783 (1) RV Èdwin Link', o¡ Majahual, Quintana Roo, Mexico (18841.85'N 87841.58'W), 58 m; CHI-1 (2) Cayo Norte, Chinchorro, Mexico, 2 m; B24 (1) Majagual, Quintana Roo, Mexico, 0.70 m; C19 (2) Buena Vista, Quintana Roo, Mexico, 04 June 1998, 1.35 m; T19 (1) Buena Vista, Quintana Roo, Mexico, 27 September 1997, 1.10 m.

Description

Description

Syntype complete (MCZ841) with 109 setiger (L103.7 mmW101.4 mm). Prostomium globular, as long as wide, with a pair of nuchal organs. Peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a muscular lip, transversally separated (Figure 3C). All parapodia well developed, but the ¢rst four smaller; parapodia with presetal lobe short, rounded and postsetal lobe digitiform, elongated (Figure 3D). Setae limbate; with two kind of hooks: composite multidentate hooded hooks from setiger 1 to 7^12, all appendages short (Figure 3E); simple multidentate hooded hooks beginning in setiger 8^13 (Figure 3F); acicula yellow. Pygidium without anal cirri. Mandibles fused for about two thirds of its total length (Figure 3G); maxillary apparatus with ¢ve pairs of maxillae (Figure 3H). Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles well developed; MII with four teeth of similar size; MIII bidentate, distal tooth longer; MIV with prominent pointed tooth; MV free.

Journal of the Marine Biological Association of the United Kingdom (2001)

Lumbrineris latreilliöUebelacker, 1984:(41) 39^41, ¢gure 36A^H. Material examined

Specimens incomplete, (BV1 with 50 setigers, L106 mm W102.5 mm). Prostomium conical, longer than wide, with a pair of nuchal organs. Peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a muscular lip; anterior ring 0.75 of total peristomial length. All parapodia well developed, with short dorsal cirrus, represented by several notoaciculae; presetal lobe short, rounded and postsetal lobe digitiform, elongated. Both lobes longest in anterior parapodia (Figure 3I,J). Setae limbate; two kinds of hooks, composite multidentate hooded hooks from setiger 1 continued to setigers 12^24, with appendages longest in more anterior setigers (Figure 3K), gradually becoming shorter in posterior setigers (Figure 3L); simple multidentate hooded hooks beginning in setigers 13^25, all with short hood (Figure 3M); acicula yellow. Mandibles slightly divided posteriorly (Figure 3N); maxillary apparatus with ¢ve pairs of maxillae (Figure 3O). Maxillary carriers shorter than MI, anterior end constricted; MI

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forceps-like with bridles well developed; MII with ¢ve teeth, as long as MI; MIII bidentate with distal tooth longer; MIV unidentate, with pointed tooth; MV free. Discussion

Lumbrineris latreilli has been considered to be a cosmopolitan eurybathic species recorded from intertidal to 2360 m. Because of the reproductive mechanisms of lumbrinerids and the lack of pelagic larvae (Sato et al., 1982; Wilson, 1991; Rouse, 2000), it is di¤cult to understand this widespread distribution; rather L. latreilli should be regarded as a species complex. There are two names employed for similar species in the western Atlantic, L. nasuta Verrill, 1900 and L. nuchalis Treadwell, 1921. If they are di¡erent from the European L. latreilli, then we should use the former because the second might be the same.

Distribution

Questionably cosmopolitan. Lumbrineris nonatoi Ramos, 1976 Figure 4A^ F

Lumbrineris nonatoi Ramos, 1976:124^127, ¢gures 19^21. Lumbrineris sp. D Uebelacker, 1984:41.44^41.45, ¢gures 42A ^ H.

(Figure 4A,B). Setae limbate; two kinds of hooks, composite multidentate hooded hooks from setiger 1 to 8^19, appendages short, all of similar length (Figure 4C); simple multidentate hooded hooks from setigers 9^20, all with short hood (Figure 4D); acicula yellow. Pygidium with two anal cirri. Mandibles translucent and fused for about two thirds its length (Figure 4E); maxillary apparatus short, stout, with ¢ve pairs of maxillae (Figure 4F). Maxillary carriers as long as MI, distally constricted; MI forceps-like with bridles; MII with three robust teeth; MIII and MIV both with one tooth; MV free. Remarks

The ending of composite multidentate hooded hooks are size dependent (r0.864, P50.01).

Discussion

The description of Lumbrineris sp. D Uebelacker, 1984 was based on incomplete specimens; then, she could not notice the increase of presetal lobe in posterior parapodia. Her specimens belong to L. nonatoi based mainly in the peculiar maxillary apparatus. Distribution

Mediterranean and Gulf of Mexico.

Material examined

USNM90998 (2) o¡ Port Isabel, Texas (26810'N 97808'W), Stocs station IV-4, 15 m; USNM75350 (1) o¡ Florida (26845'50''N 82845'11''W), So£a station 2B, 24 m; CP-ICML-UNAM IMCA 2, station 18 (1), September 1988, o¡ Veracruz; CP-ICML-UNAM IMCA 2, station 76 (2), 27 September 1988; o¡ Campeche. ECOSUR LUMB5 E3COLT-I (1) o¡ Tamaulipas, Mexico 34 m; E33COLT-I (1) o¡ Tamiahua Lagoon, Veracruz, Mexico, 52 m; E19COLT-III (1) o¡ Tamaulipas, Mexico; E31COLT-III (1) o¡ Tamaulipas, Mexico; E43COLT43 (1) Tamaulipas, Mexico; E5EMOAPP-II (3) o¡ north Veracruz, Mexico, 23 m; E6EMOAPP-II (2) o¡ Tamiahua Lagoon, Veracruz, Mexico, 23 m; E8EMOAPP-II (1) o¡ the north of Veracruz, 56 m; E11COSTA (1) Tamaulipas, Mexico, 27 m; M1E1 (2) o¡ Sontecomapan, Veracruz, Mexico (18835'N 94852'W); M1E5 (1) o¡ Punta San Juan, Veracruz, Mexico (18820'N 94838'W); M1E6 (1) o¡ Coatzacoalcos, Veracruz, Mexico (18811'N 94825'W); M1E7 (3) o¡ Coatzacoalcos, Veracruz, Mexico (18820'N 94825'W); M1E14 (1) (18823'N 93848'W), o¡ Carmen Lagoon, Tabasco, Mexico; E7PROMEX-III(1) Gulf of Mexico; E12PROMEX-III (1) o¡ Punta Zapotitlan, Veracruz, Mexico (18831'N 94836'W), E30PROMEX-III (1) o¡ Campeche, Mexico (18858'N 92802'W), 20 m. Description

Specimen complete (E31COLT-III) with 149 setigers (L101.9 mm W100.5 mm). Prostomium globular, as long as wide, with a pair of nuchal organs. Peristomium shorter than prostomium, both rings of similar length, separation between rings distinct dorsally and laterally, ventrally with a shallow lip. Parapodia well developed; anterior parapodia with presetal lobe short, rounded and postsetal lobe digitiform, elongated; median parapodia with both lobes of similar length and posterior parapodia with presetal lobe longer than postsetal lobe, digitiform Journal of the Marine Biological Association of the United Kingdom (2001)

Lumbrineris paucidentata Treadwell, 1921 Figure 4G ^ J Lumbrinereis paucidentata Treadwell, 1921:99^100, plate 9, ¢gures 1^4, text-¢gures 357^364. Lumbrineris paucidentata ö Hartman, 1956:288. Material examined

Holotype of Lumbrinereis paucidentata Treadwell AMNH1199, Dry Tortugas, Florida. Non-type material: ECOSUR LUMB6 XC-1 (1) Xcacel, Quintana Roo, Mexico, intertidal. Description

Holotype a mature female, incomplete with 199 setigers (L105.5 mm W102 mm), LUMB6 incomplete with 28 setigers (L104.1mm W101.7 mm). Prostomium conical, as long as wide, with a pair of nuchal organs. Peristomium shorter than prostomium, both rings of similar length; separation between rings distinct dorsally and laterally, ventrally with a shallow lip. All parapodia well developed, of similar length throughout; presetal lobe short, rounded and postsetal lobe digitiform, elongated. Setae limbate; two kinds of hooks, composite multidentate hooded hooks from setiger 1 to 11, with long appendages (Figure 4G); simple multidentate hooded hooks from setiger 12, with short hood (Figure 4H); acicula yellow. Mandibles with short and slightly divergent shaft (Figure 4I); maxillary apparatus with ¢ve pairs of maxillae (Figure 4J). Maxillary carriers shorter than MI, constricted anteriorly and rounded posteriorly; MI forceps-like with bridles; MII with four teeth; MIII unidentate; MIV with short tooth; MV tiny, free. Discussion

The maxillary apparatus was removed from the holotype, as noted by Hartman (1965). The maxillary apparatus

Lumbrineridae from the Grand Caribbean region of LUMB6 is identical to the one described and illustrated by Treadwell (1921). Distribution

Dry Tortugas, Florida; Xcacel, Quintana Roo, Mexico. Lumbrineris perkinsi sp. nov. Figure 4K ^ P

Material examined

Syntypes, USNM (10) San Juan de Ulua, Veracruz, Mexico, 3 m; BM(NH) (10) San Juan de Ulua, Veracruz, Mexico, 3 m; MNHN (10) San Juan de Ulua, Veracruz, Mexico, 3 m; ECOSUR LUMB8 E12 EMOAPP-II (1) o¡ Laguna San Andres, Tamaulipas, Mexico, 81m; SJU-I (16) San Juan de Ulua, Veracruz, Mexico, 3 m; IE-I (5) Isla de Enmedio Reef, in sponge, 6 m; E7 PROMEX-III (1) Gulf of Mexico; Cha13 (2) Champoton, Campeche, Mexico, in coralline-rock, 6 m; Cha9 (1) 12 km south of Champton, Campeche, Mexico, in coralline-rocks, 3 m; Cha14 (2)

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Champoton, Campeche, Mexico, in coralline-rock, 6.5 m; LAG-1 (3) Ria Lagartos, Yucatan, in coralline-rock, 3 m; LAG-2 (1) San Felipe, Yucatan, Mexico, in coralline-rock, 2 m; CEL-1 (2) Ria Celestun, Yucatan, Mexico, in coralline-rock, 0.6 m; IMU-1 (2) Isla Mujeres, Mexico, intertidal; PM (12) Puerto Morelos, Quintana Roo, Mexico, in sponge; COZ-3 (1) Playa Azul, Cozumel Island, Mexico, 1.5 m; XC-I (2) Xcacel, Quintana Roo, Mexico, in coralline-rock, intertidal; CV9 (27) Cayo Valencia, Ascension Bay, Quintana Roo, Mexico, in mangrove roots; CCE-1 (4), Cedros, Ascension Bay, Quintana Roo, Mexico, in mangrove roots; EL-2771 (1) RV Èdwin Link', east coast of Cozumel Island (20825'N 87802'W); E24 (1) BuenaVista, Quintana Roo, Mexico, 1.2 m; IOH (1) archipelago Jardines de la Reina, Cuba, 0.5 m; MMI P968 (2) RV `Pillsbury', cruise 6907, north of Antigua, Lesser Antilles (17818'N 61852'W), 20 July 1969, 18 m; MMI P1284 (1) RV `Pillsbury', cruise 7006, south of Cabo Beata, Dominican Republic (17835'N 71825'W), 19 July 1970, 20 m.

Figure 4. Lumbrineris nonatoi Ramos (ECOSUR LUMB5). (A) Anterior parapodium; (B) posterior parapodium; (C) composite multidentate hooded hook; (D) simple multidentate hooded hook; (E) mandible; (F) maxillary apparatus. Lumbrineris paucidentata Treadwell (G^H Holotype AMNH1199, I^J ECOSUR LUMB6). (G) Composite multidentate hooded hook; (H) simple multidentate hooded hook; (I) mandible; (J) maxillary apparatus. Lumbrineris perkinsi sp. nov., (Syntype USNM). (K) Anterior end in ventral view; (L) anterior parapodium; (M) composite multidentate hooded hook; (N) simple multidentate hooded hook; (O) mandibles; (P) maxillary apparatus. Lumbrineris in£ata Moore (Syntype USNM16840). (Q) Maxilla III; (R) anterior end in ventral view. Scale bars: A&B, 0.05 mm; C,D,G,H,M&N, 0.01 mm; E,F,J,Q&P, 0.1 mm; I,K&R, 0.2 mm. Journal of the Marine Biological Association of the United Kingdom (2001)

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Description

Prostomium globular, as long as wide; with a pair of nuchal organs. Peristomium shorter than prostomium, separation between rings distinct dorsally and laterally, ventrally with a complete shallow lip; anterior ring 0.75 of total peristomial length (Figure 4K). All parapodia well developed but anterior parapodia longer than posterior ones; presetal lobe short, rounded and postsetal lobe digitiform elongated (Figure 4L). Pygidium with two pairs of anal cirri of similar length. Setae limbate; two kinds of hooks, composite multidentate hooded hooks from setiger 1 to 5^16, with short appendage (Figure 4M); simple multidentate hooded hooks from setiger 6^17, with short hood (Figure 4N); acicula yellow. Mandibles as long as maxillary complex, fused for about two thirds its total length (Figure 4O); maxillary apparatus with ¢ve pairs of maxillae (Figure 4P). Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles well developed; MII with ¢ve teeth, of similar length to MI; MIII with four teeth, distal one longest; MIV bidentate with distal tooth longest; MV free.

Remarks

The ending of composite multidentate hooded hooks are size dependent (r0.864, P50.01). The smallest specimen (L100.8 mm W100.3 mm) does not have any composite multidentate hooded hooks. Discussion

Lumbrineris perkinsi sp. nov. resembles L. in£ata Moore, 1911 (California) by having MIII multidentate and MIV bidentate but can be separated by the di¡erent number of teeth in MIII and by the shape of peristomial ventral side. Lumbrineris in£ata has MIII with only three teeth (material examined: syntypes USNM16840 (2), USNM17205 (1); non-type material USNM40599 (2), USNM40597 (1), USNM40598 (2), USNM40596 (8), USNM40595 (5), USNM40594 (45)) and a peristomium transversally separated (Figure 5Q,R); while L. perkinsi sp. nov. always has four teeth in MIII and a peristomium ventrally complete. In L. perkinsi sp. nov. the basal tooth of MIII is smaller and sometimes worn, but a pulp cavity is always present. Type localities

Gulf of Mexico, Mexican Caribbean, Cuba, Dominican Republic and Lesser Antilles.

Etymology

The species is named in honour of Thomas H. Perkins in recognition of his many publications on polychaetes and especially for his work with lumbrinerids.

Gender

Florida (25816'54''N 83843'11''W), So£a station 24F, 88 m. ECOSUR LUMB7 E1PROMEX-III (1), o¡ Tuxpan, Veracruz (20831.5'N 96858.1'W), 39 m. Description

Prostomium conical, as long as wide, with a pair of nuchal organs; peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a shallow lip; anterior ring 0.75 of total peristomial length (Figure 5A). All parapodia well developed, but ¢rst 3^4 smaller. Anterior parapodia with presetal lobe short, slightly conical and postsetal lobe conical, longer than presetal one; posterior parapodia with both lobes digitiform, but presetal longer than postsetal (Figure 5B,C). Pygidium with two pairs of anal cirri, dorsal pair longer than ventral one. Setae limbate; two kinds of hooks, composite multidentate hooded hooks from setiger 1 to 8^13, with short appendage (Figure 5D); simple multidentate hooded hooks from setigers 9^14, with short hood (Figure 5E); acicula yellow. Mandibles distally rounded, fused for about one third its total length, with two internal blacks bands from distal end to middle (Figure 5F); maxillary apparatus with ¢ve pairs of maxillae (Figure 5G). Maxillary carriers as long as MI, anterior end slightly notched; MI slender, forceps-like with bridles well developed; MII with four teeth, as long as MI, with a small portion of the posterior membrane partially sclerotized; MIII with two teeth, distal one longest; MIV with two teeth, the basal tiny; MV free.

Remarks

The ending of composite multidentate hooded hooks is size dependent (r0.958, P50.01).

Discussion

Lumbrineris salazari sp. nov. is similar to Lumbrineris obtusa Kinberg, 1865 from Chile; in both, MIII and MIV are bidentate, but they di¡er in the relative size of setal lobe in posterior parapodia; L. salazari sp. nov. has longer presetal lobe and L. obtusa has longer postsetal lobe. Type localities

Florida and Veracruz, Gulf of Mexico.

Etymology

This species is named in honour of my mentor and good friend Sergio I. Salazar-Vallejo in recognition of his invaluable labour in the improvement of the taxonomy of polychaetes.

Gender

Masculine.

Masculine. Lumbrineris salazari sp. nov. Figure 5A ^ G

Lysarete Kinberg, 1865 Lysarete raquelae sp. nov. Figure 5H ^ O

Lumbrineris sp. E Uebelacker, 1984:41.41, ¢gure 38A ^ H.

Lysarete brasiliensisöGilbert, 1984:43.6^43.7, ¢gure 4A^ G (non Kinberg).

Material examined

Material examined

Syntypes USNM75353 (10) o¡ Florida (26816'43''N 83812'49''W), So£a station 8C, 48 m; USNM75355 (1) o¡


Syntypes USNM31521 (2) Virginia Beach, Virginia, taken on beach after tidal storm; USNM49190 (1) Gulf of


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Figure 5. Lumbrineris salazari sp. nov. (Syntype USNM75353). (A) Anterior end; (B) anterior parapodium; (C) posterior parapodium; (D) composite multidentate hooded hook; (E) simple multidentate hooded hook; (F) mandible; (G) maxillary apparatus. Lysarete raquelae sp. nov. (H^L Syntype USNM31521, M Syntype USNM30964, N Syntype USNM53040, O Syntype USNM30968). (H) Anterior end; (I) anterior parapodium; (J) posterior parapodium; (K) mandible; (L) maxillary apparatus; (M) maxillary apparatus; (N) maxillae IÎV; (O) maxillae IV, VI. Scale bars: A&I^L, 0.5 mm; B&C, 0.2 mm; D&E, 0.01 mm; F,G&MÔ, 0.1 mm; H, 2.5 mm.

Mexico (29835'N 88850'W), south of Chandeleur Island, 9 m. USNM55884 (1) Gulf of Mexico (29854'58.8''N 86804'58.5''W), RV `Columbus Iselin' station 2528, June 1976, 37 m; USNM60380 (1) o¡ the east coast of Florida (29831'N 80840'W), 18 m; USNM53040 (1) Bogue Sound, Morchead City, North Carolina, shallow deep on pilings; USNM98179 (1) o¡ southern Florida; USNM15812 (1) Beaufort, North Carolina; USNM30968 (4) Alligator Harbor, Florida; USNM30964 (1) o¡ Georgia (31859.5'N 80829.5'W), from body cavity of tunicate, 15 m. Description

Prostomium short, rounded, wider than long, with three occipital antennae, tapering, the lateral antennae reaches second peristomial ring, the median antennae reach ¢rst setiger; with two pairs of eyes. Peristomium with a dorsal groove from ¢rst ring to ¢rst setiger, ventrally with a muscular lip (Figure 5H). All parapodia well developed, but ¢rst 6^7 slightly smaller; dorsal cirri from ¢rst setiger, become longer in posterior parapodia, in anterior parapodia are digitiform and foliaceous in posterior segments, they are vascularized; both setal lobes well


developed from ¢rst setiger, presetal lobe conical and postsetal digitiform, are longer and vascularized in posterior parapodia (Figure 5I,J). Only limbate setae; acicula black. Pygidium with two pairs of anal cirri of similar length. Mandibles fused for about half its length (Figure 5K); maxillary apparatus with six pairs of maxillae (in adults L10 10^16.1mm, W10 6^8.8 mm) (Figure 5L). Maxillary carriers twice longer than MI, anterior end constricted; MI forceps-like with bridles well developed, with two internal accessories teeth in basal position (in adults, see remarks); MII with 6^7 teeth; MIII with four teeth and MIV with four teeth, both maxillae with distal tooth bigger; MV free, lateral to MIV; MVI uppermost MIV with a long conical tooth. The shape and number of teeth are variables during ontogenetic development (see remarks). Remarks

Small specimens (L10 2.2^5.4 mm, W10 0.5^1.5 mm) have a di¡erent shape and numbers of teeth in maxillary apparatus; becoming similar to maxillae of Oenonidae. Specimen with L10 53 mm (USNM30964) has maxillary apparatus with only three maxillae. Maxillary carriers

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posteriorly incomplete, fused to MI; MI with internal border denticulate, with ten teeth and without bridles; MII asymmetrical, left plate smaller than right plate and MIII with ¢ve teeth, basal tooth smaller (Figure 5M). Specimen with L105 mm (USNM53040) has maxillary apparatus with four pairs of maxillae. Maxillary carriers well de¢ned, anterior end constricted; MI with distal end forceps-like and basal end denticulate, with bridles, (Figure 5N), MII with both plates of similar size with nine teeth; MIII and MIV multidentate with distal tooth bigger. Specimen with L105.4 mm (USNM30968) has maxillary apparatus with ¢ve pair of maxillae; MI ^ MIVas previous specimen; MV tiny; MVI asymmetrical (Figure 5O), left tooth smaller. The size of antennae and dorsal groove in the peristomium are size dependent, but all are present. Discussion

There are only two species in Lysarete: L. brasiliensis Kinberg (Rio de Janeiro, Brazil) and L. australiensis Benham (southern Australia).The former has been recorded in two discontinuous regions; in North America, from North Carolina to Florida and Gulf of Mexico and in South America from Rio de Janeiro, Brazil to Golfo San Mat|¨ as, Argentina. The specimens from North America are di¡erent species by the di¡ering mainly in the number of teeth in the maxillary apparatus: L. raquelae sp. nov. has MI with two accessories teeth in a basal position, the distal accessory tooth blunt, bigger than the basal one while L. brasiliensis has a MI with only one prominent pointed tooth. Furthermore, L. raquelae sp. nov. has more teeth in MIII and MIV (four teeth) than L. brasiliensis which has both maxillae tridentate. Lysarete australensis has a di¡erent maxillary formula being MI with one pointed tooth and few teeth in MIII and MIV (three and two respectively). Type localities

North Carolina, Florida and Gulf of Mexico.

Etymology

This species is named in honour of my mother Raquel Parra, in recognition of her support and encouragement.

Gender

Feminine.

Ehlers MCZ1238 (8) south of Dry Tortugas (24808'N 82851'W), Blake 1877^1878, station 43, 620 m; MCZ881 (1) o¡ the Mississippi delta (28851'30''N, 89801'30''W), Blake 1877^1878, station 49, 216 m. Non-type materials: YPM2190 (5), YPM2171 (15) o¡ Martha's Vineyard, Vineyard Sound, Massachusetts, 29^31.5 m. MMI G848 (2) RV `Gerda', cruise 6713, Florida (25853'N 80803'W), 02 August 1967, 201m; MMI G850 (1) RV `Gerda', cruise 6713, Florida (25848'N 80803'W), 02 August 1967, 214 m; CP-ICML-UNAM IMCA 2, station 32 (1), 23 September 1988, o¡ Tabasco; CP-ICML-UNAM IMCA 2, station 52 (1), 24 September 1988, o¡ Campeche; CP-ICML-UNAM IMCA 3, station 63 (1), 09 March 1988, o¡ Campeche. Description

Prostomium conical, as long as wide, with a pair of nuchal organs; peristomium shorter than prostomium, both rings of similar size; separation between rings distinct dorsally and laterally, ventrally with a shallow lip (Figure 6A). All parapodia well developed; presetal lobe rounded short and postsetal lobe digitiform, longer in the branchial region (Figure 6B). Branchiae from setigers 3 to 20^31, with up to ¢ve branchial ¢laments. Pygidium with a pair of anal cirri. Setae limbate; simple multidentate hooded hooks with long hood from setigers 1 to 20^31 (Figure 6C), simple multidentate hooded hooks with short hood from setigers 21^31 always present after branchial region (Figure 6D); all setae with black core from the base to near to the tip where they become translucent; acicula black. Mandibles fused for about half the total length, with two internal black bands from distal end to mid-mandible (Figure 6E); maxillary apparatus with ¢ve pairs of maxillae (Figure 6F). Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles; MII with six teeth, subdistal tooth slightly smaller; MIII with one prominent tooth; MIV multidentate with up to 17 teeth, distal tooth longest; MV free.

Remarks

The ending of both branchial region and simple multidentate hooded hooks with large hood, and the beginning of the simple multidentate hooded hooks with short hood are size dependent (r827, P50.001).

Discussion Ninoe Kinberg, 1865 Ninoe nigripes Verrill, 1873 Figure 6A ^ F

Ninoe nigripes Verrill, 1873:595; Hartman, 1942b:53^54, ¢gures 94^97; Pettibone, 1963:266^268, ¢gure 68A ^ G (include N. kinbergi and N. nigripes gracilis). Ninoe kinbergi Ehlers, 1887: 105^106, plate 32, ¢gures 1^9. Ninoe nigripes gracilis Hartman, 1951:61, 63, plate 16, ¢gures 1^2.

Hartman (1951) erected the subspecies N. nigripes gracilis on the basis of having MIII multidentate. The maxillary apparatus of the type is in poor condition, but it has MIII with only one tooth. The synonymy of Pettibone (1963) is herein considered to be valid.

Distribution

Massachusetts to Florida, Louisiana, Tabasco and Campeche.

Material examined

Syntypes of Ninoe nigripes Verrill YPM2735 (13) o¡ Martha's Vineyard, Vineyard Sound, Massachusetts, US Fish Commission, 45 m;YPM2736 (1) US Fish Commission, no other locality data; YPM2737 (13) New Brunswick, Bay of Fundy, Grand Manan Island, 108 m. Holotype of Ninoe nigripes gracilis Hartman LACM4289, ten miles south of Grand Isle, Louisiana, 6 m. Syntypes of Ninoe kinbergi


Ninoe vargasi sp. nov. Figure 6G ^ K Ninoe sp. B Uebelacker, 1984:41.7; ¢gure 4A ^ G. Material examined

Syntypes USNM90979 (1) Louisiana, South Timbalier Lease Area (28834'09''N 90824'32''W). Coll. SWRI for


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Figure 6. Ninoe nigripes (Verrill) (A, E^F Syntype YPM2735, B^D Syntype MCZ1238). (A) Anterior end; (B) parapodium with branchiae; (C) simple multidentate hooded hook from anterior setiger; (D) simple multidentate hooded hook from posterior setiger; (E) mandible; (F) maxillary apparatus. Ninoe vargasi sp. nov. (Syntype ECOSUR LUMB9). (G) Anterior end; (H) parapodium with branchiae; (I) simple multidentate hooded hook; (J) mandible; (K) maxillary apparatus. Scale bars: A&G, 0.5 mm; B&C, 0.2 mm; C,D&I, 0.01 mm; E,F,J&K, 0.1 mm.

BLM, May 1978; station 04P, 45 m; USNM55881 (1) Alabama (29853'30''N 88812'27.5''W), RV `Columbus Iselin', station 2639, June 1975, 32 m; USNM90980 (1) Texas, o¡ Port Aransas (27840'N 096859'W); coll. for BLM, 1976; Stocs station 11-1, 22 m; MMI G812 (1) RV `Gerda', cruise 6617, south of Florida (25833'N 80804'W), 22 November 1966; MMI G854 (1) RV `Gerda', cruise 25827'N 80802'W, 25 August 1967, 221m; BM(NH) (2) o¡ Coatzacoalcos, Veracruz, Mexico (18821'N 94825'W); MNHN (2) o¡ Carmen Lagoon, Tabasco, Mexico (18823'N 93848'W); ECOSUR LUMB9 E4-N Panuco (1) o¡ Tampico, Tamaulipas, Mexico; E43COLT-I (1) south of Tamiahua Lagoon, Veracruz, Mexico; M1E1 (1) o¡ Sontecomapan, Veracruz, Mexico (18835'N 94852'W); M1E5 (1) o¡ Punta San Juan, Veracruz, Mexico (18820'N 94838'W); M1E6 (2) o¡ Coatzacoalcos, Veracruz, Mexico (18821'N 94825'W); M1E8 (1) o¡ Coatzacoalcos, Veracruz, Mexico (18831'N 94825'W); M1E10 (1), o¡ Tonala, Veracruz, Mexico (18841'N 94813'W); MIE14 (2) o¡ Carmen Lagoon, Tabasco, Mexico (18823'N 93848'W); E6 PROMEX-III (1) o¡ Punta Zempoala, Veracruz (19830'N 95859'W), 188 m; E12 PROMEX-III (1) north of Punta San Juan, Veracruz, Mexico (18831'N 94836'W), 63 m; E38 PROMEX-III (2) o¡ Terminos Lagoon, Campeche, Mexico (19827'N 91856'W), 47 m; CP-ICML-UNAM DINAMO 2, station 52 (3), 02 November 1990, o¡ Campeche; CP-ICML-UNAM Journal of the Marine Biological Association of the United Kingdom (2001)

DINAMO 2, station 60 (1), 09 November 1990, o¡ Campeche; CP-ICML-UNAM IMCA 2, station 21 (1), 22 January 1989, o¡ Tabasco; CP-ICML-UNAM IMCA 1, station 80 (1), 10 March 1988, o¡ Campeche; CP-ICMLUNAM IMCA 1, station 55 (2), 08 February 1988, o¡ Campeche. Description

Syntype (ECOSUR LUMB9-E12) complete with 35 setigers (L101.2 mm W100.4 mm). Prostomium conical, more than twice as long as wide, with a pair of divergent dorsal longitudinal black bands, with a pair of nuchal organs. Peristomium shorter than prostomium, both rings of similar size; separation between rings distinct dorsally and laterally, ventrally with a shallow lip (Figure 6G). All parapodia well developed with inconspicuous rounded presetal lobe and digitiform postsetal lobe, longest in branchial region (Figure 6H). Branchiae from setigers 4^5 extending to setigers 12^38, with up to six branchial ¢laments. Setae limbate, simple multidentate hooded hooks always after branchial region, with short hood (Figure 6I); acicula black. Pygidium with a par of anal cirri. Mandibles divided for 0.75 length, with two internal black bands from distal end to middle (Figure 6J); maxillary apparatus with ¢ve pairs of maxillae (Figure 6K). Maxillary carriers as long as MI, anterior end constricted; MI forceps-like with

612

L.F. Carrera-Parra


bridles well developed; MII with seven teeth; MIII multidentate with six teeth, the distal one bigger; MIV multidentate with up to eight teeth, distal one longest; MV free. Remarks

The ending of the branchial region and the beginning of simple multidentate hooded hooks are size dependent (r0.89, P0.001).

Discussion

Ninoe vargasi sp. nov. is similar to N. armoricana Gle¨marec (Mediterranean Sea), N. gemmea Moore (California) and N. wardae sp. nov. (Gulf of Mexico) in having MIII multidentate. Ninoe vargasi sp. nov. di¡ers from the others by having simple hooded hooks restricted to setigers in the postbranchial region.

Type locality

Gulf of Mexico.

Discussion

Ninoe wardae sp. nov. resembles N. armoricana Gle¨marec, 1968 (Mediterranean Sea), N. gemmea Moore, 1911 (California) and N. vargasi sp. nov. (Gulf of Mexico) by having MIII multidentate. Ninoe armoricana and N. vargasi sp. nov. can be separated from the other two because the hooks begin more posteriorly, while N. gemmea and N. wardae sp. nov. have hooks beginning at the ¢rst setiger. Ninoe wardae sp. nov. is the only species with black mandibles. Type locality

Apalachicola River, Florida, Gulf of Mexico.

Etymology

This species is named in honour of Linda Ward in recognition of her extraordinary e¡ort in the collections of the National Museum of Natural History, Smithsonian Institution, and by her kind support during part of this study.

Gender

Etymology

This species is named in honour of Juan Manuel VargasHerna¨ndez, a former colleague and friend, and in recognition of his studies of reef fauna from the Gulf of Mexico, mainly from Veracruz.

Gender

Masculine. Ninoe wardae sp. nov. Figure 7A^ G

Ninoe sp. A Uebelacker, 1984:(41) 5, 7, ¢gure 2A ^ H. Material examined

Holotype USNM67768, o¡ Apalachicola River, Florida, USA, 175 m, Ma£a station IV-2427 (29849'59''N 85837'02''W), July 1976.

Description

Holotype incomplete, broken into two fragments, with 50 setigers (L102.6 mm W100.8 mm). Prostomium longer than wide, with a pair of divergent dorsal longitudinal black bands; with a pair of nuchal organs. Peristomium shorter than prostomium, both rings of similar size; separation between rings distinct dorsally and laterally, ventrally with a shallow lip (Figure 7A). All parapodia well developed, parapodia in prebranchial region with short rounded presetal lobe and rounded postsetal lobe; in branchial region parapodia with inconspicuous presetal lobe and long digitiform postsetal lobe; in postbranchial region both lobes short (Figure 7B,C). Branchiae from setigers 11 to 30, with only one branchial ¢lament. Setae limbate, simple multidentate hooded hooks from ¢rst setiger; in anterior setigers with long hood, becoming shorter in posterior ones (Figure 7D,E); acicula black. Mandibles black, fused at the anterior end (Figure 7F); maxillary apparatus with ¢ve pairs of maxillae (Figure 7G). Maxillary carriers longer than MI, anterior end constricted; MI forceps-like with bridles well developed; MII with eight teeth, distal tooth longest; MIII with 13 teeth, distal one longest; MIV with 11 teeth, the distal prominent; MV free. Journal of the Marine Biological Association of the United Kingdom (2001)

Feminine. Scoletoma de Blainville, 1828 Scoletoma branchiata (Treadwell, 1921) comb. nov. Figure 7H ^ M

Lumbrinereis branchiata Treadwell, 1921:94^95, plate 8, ¢gures 5^6, text-¢gures 333^343. Lumbrineris branchiate ö Hartman, 1956:287. Lumbrineris alata Hartman, 1951:59^61, plate 15, ¢gures 1^4. Material examined

Holotype of Lumbrinereis branchiata Treadwell AMNH1338, Bucco Bay, Tobago, April 1918. Holotype of Lumbrineris alata Hartman LACM-AHF0710, Port Aransas, Texas. Non-type material, ECOSUR LUMB10 (2) Mochima Bay, Venezuela, 1.5 m.

Description

Holotype incomplete with 436 setigers (L106.5 mm W102 mm). Prostomium conical, longer than wide; with a pair of nuchal organs. Peristomium shorter than prostomium, both rings of similar size; separation between rings distinct dorsally and laterally, ventrally with a shallow lip. All parapodia well developed, but ¢rst seven smaller; with notopodia button-like with internal notoaciculae. Presetal lobe inconspicuous, rounded, and postsetal lobe digitiform, elongated; both lobes more developed in posterior parapodia; nephridial papillae poorly developed in mid-anterior parapodia, then developing gradually towards posterior end (Figure 7H,I). Pygidium with two pairs of anal cirri, dorsal longer than ventral. Setae limbate, simple multidentate hooded hooks from ¢rst setiger, in anterior setigers with long hood becoming shorter in posterior ones (Figure 7J,K); acicula yellow. Mandibles fused for about two thirds its total length (Figure 7L); maxillary apparatus with ¢ve pairs of maxillae (Figure 7M). Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles well developed; MII as long as MI, with 4^5 teeth, with a portion of the posterior membrane partially sclerotized; MIII with two teeth of similar size; MIV with pointed tooth; MV free.

Lumbrineridae from the Grand Caribbean region Discussion

Treadwell (1921) inverted the parapodium and confused the nephridial papillae with dorsal branchiae. Based on this feature Hartman (1951) described Lumbrineris alata as a di¡erent species. Hartman (1956) reviewed the type of L. branchiata, emended Treadwell's description and regarded L. alata as a probable synonym.

Distribution

Port Aransas, Texas; Bucco Bay, Tobago and Mochima Bay, Venezuela. Scoletoma candida (Treadwell, 1921) comb. nov. Figure 8A ^ E

Lumbrinereis candida Treadwell, 1921:96^97, plates 7^9, text¢gures 344^350. Lumbrineris candida ö Hartman, 1956:288; Uebelacker, 1984:(41) 21,24, ¢gure 18A ^ H. Material examined

Holotype of Lumbrinereis candida Treadwell AMNH1277, Bucco Bay, Tobago in ¢ne sand, April 1918. Non-type

L.F. Carrera-Parra

613

material, ECOSUR LUMB11 PC1 (1) Paraiso, Cozumel Island, Quintana Roo, Mexico, in sand; CHI-1 (1) Cayo Norte, Chinchorro, Mexico, in sand, 2 m. Description

Holotype incomplete, broken into three parts, Treadwell (1921) wrote ``. . . The bottle labelled type contains an anterior and a posterior end from this lot, but I can not be certain that both came from the same individual . . .''. The anterior end has 200 setigers (L107.3 W10 1.5 mm). Prostomium conical, twice longer than wide, with a pair of nuchal organs; peristomium shorter than prostomium, rings of similar size; separation between rings distinct dorsally and laterally, ventrally with a shallow lip. All parapodia well developed; presetal lobe short rounded and postsetal lobe digitiform elongated; both lobes longer in posterior parapodia (Figure 8A,B). Setae limbate, simple multidentate hooded hooks from ¢rst parapodia, in anterior setigers, with long hood becoming shorter in posterior (Figure 8C,D); acicula yellow. Mandibles with short and slight divergent shaft; maxillary apparatus short, stout, with ¢ve pairs of maxillae (Figure 8E). Maxillary carriers as long as MI,

Figure 7. Ninoe wardae sp. nov. (Holotype USNM67768). (A) Anterior end; (B) parapodium with branchia; (C) posterior parapodium; (D) simple multidentate hooded hook from anterior setiger; (E) simple multidentate hooded hook from posterior setiger; (F) mandible; (G) maxillary apparatus. Scoletoma branchiata (Treadwell) (Holotype AMNH1338); (H) anterior parapodium; (I) posterior parapodium in ventral view; (J) simple multidentate hooded hook from anterior setiger; (K) simple multidentate hooded hook from posterior setiger; (L) mandible; (M) maxillary apparatus. Scale bars: A, 0.4 mm; B,C,G&F, 0.1 mm; D,E,J&K, 0.01 mm; L&M, 0.2 mm. Journal of the Marine Biological Association of the United Kingdom (2001)

614

L.F. Carrera-Parra


distal end constricted and rounded posterior end; MI forceps-like with bridles; MII with four teeth; MIII edentate; MIV with one tooth; MV tiny. Remarks

Hartman (1956) redescribed MIII as being bidentate; however, the MIII is edentate as was originally described by Treadwell (1921).

Distribution

Quintana Roo, Mexico and Bucco Bay, Tobago. Scoletoma elongata (Treadwell, 1931) comb. nov. Figure 8F ^ I

Lumbriconereis elongata Treadwell, 1931:3, ¢gure 2A ^ D. Material examined

Holotype of Lumbriconereis elongataTreadwell USNM19622, Grand Island, Louisiana. Description

Specimen(s) broken in several parts, with 1064 setigers (L103.5 mm W101.2 mm): the assignation of all fragments to the same specimen is doubtful (Treadwell, 1931). Prostomium conical, as long as wide, with a pair of nuchal organs. Peristomium short, separation between rings distinct dorsally and laterally, ventrally with a muscular lip; anterior ring 0.75 of total peristomial

length. First three parapodia smaller, all parapodia lateral (Figure 8F); presetal lobe short rounded and postsetal lobe digitiform, more elongated in posterior parapodia (Figure 8G,H). Setae limbate, simple multidentate hooded hooks; almost all setae in anterior setigers broken, in posterior setiger hooks with short hood, distally truncated (Figure 8I); acicula yellow. Holotype lacks maxillary apparatus, but it was described by Treadwell ``. . . the forceps have a prominent carrier and a slender terminal portion. The left paired plate has a slender terminal tooth followed by three much heavier ones and a basal hump that may be the remnant of tooth. The second paired plate has two teeth, the terminal plate only one . . .'' Discussion

Hartman (1956) regarded this species as a junior synonym of S. parvapedata (Treadwell, 1901) with the only di¡erence being in the number of teeth of MII. The specimens of S. parvapedata are bigger than the holotype of S. elongata (L104.5 mm W102.2 mm and L103.5 mm W101.2 mm respectively). The former have a peristomium distinctly shorter without ventral lip; furthermore, the ¢rst three parapodia of the specimens are in a more dorsal position. On the basis of these di¡erences, both species are considered to be valid.

Distribution

Louisiana, Gulf of Mexico.

Figure 8. Scoletoma candida (Treadwell) (Holotype AMNH1277). (A) Anterior parapodium; (B) posterior parapodium; (C) simple multidentate hooded hook from anterior setiger; (D) simple multidentate hooded hook from posterior setiger; (E) maxillary apparatus. Scoletoma elongata (Treadwell) (Holotype USNM19622). (F) Anterior end in lateral view; (G) anterior parapodium; (H) posterior parapodium; (I) simple multidentate hooded hook. Scoletoma ernesti (Perkins) (J^K Paratype USNM55878, L^M Paratype USNM45598). (J) Anterior parapodium; (K) simple multidentate hooded hook; (L) mandible; (M) maxillary apparatus. Scale bars: A&B, 0.05 mm; C,D,I&K, 0.01 mm; E,M,L&J, 0.2 mm; F, 0.5 mm; G&H, 0.1 mm. Journal of the Marine Biological Association of the United Kingdom (2001)


615

Tabasco, Mexico (18823'N 93848'W); IDP (1) Veracruz, Mexico.

Scoletoma ernesti (Perkins, 1979) Figure 8J ^ M Lumbrineris ernesti Perkins, 1979:429^433, Uebelacker, 1984:(41) 28^30, ¢gure 26A ^ G. Scoletoma ernersti ö Frame, 1992:203.

L.F. Carrera-Parra

¢gures

5^6;

Material examined

Holotype of Lumbrineris ernesti Perkins USNM57438, Anclote Anchorage Tarpon Springs, Florida (28812.6'N 82847.6'W), 3.5 m; paratypes USNM55878 (1), Gulf of Mexico (29855'N 88843.5'W), RV `Columbus Iselin'; USNM54580 (1), o¡ Beaufort, North Carolina (34829'N 76813'W), 33 m; USNM45598 (7), Tampa Bay, Florida. Non-type materials, MMI G1032 (1) RV `Gerda', cruise 6905, Straits of Florida (24836'N 81864'W), 26 February 1969, 46 m; MMI P699 (1) RV `Pillsbury', cruise 6806, Venezuela (09830'N 60815'W), 16 July 1968, 64 m; MMI P756 (1) RV `Pillsbury', cruise 6806 o¡ Puerto Cumarebo, Venezuela (11834'N 69812'W), 27 m, 27 July 1968; ECOSUR LUMB12 E1COLT-III (1) o¡ Tamaulipas, Mexico; M1E5 (1) o¡ Punta San Juan, Veracruz, Mexico (18820'N 94838'W); M1E6 (1) o¡ Coatzacoalcos, Veracruz, Mexico (18811'N 94825'W); M1E14 (1) o¡ Carmen Lagoon,

Description

Holotype complete with 259 setigers (L105.3 mm W102.5 mm). Prostomium conical, as long as wide, with a pair of nuchal organs. Peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a shallow lip; anterior ring 0.75 of total peristomial length. All parapodia well developed, with notopodia button-like with internal notoaciculae; anterior parapodia with presetal lobe rounded and postsetal lobe digitiform basally in£ated; in median and posterior parapodia both lobes become more developed (Figure 8J). Setae limbate, simple multidentate hooded hooks from setigers 21^50 (Figure 8K); acicula yellow. Pygidium with two pairs of anal cirri. Mandibles fused for about two thirds its total length (Figure 8L); maxillary apparatus with ¢ve pairs of maxillae (Figure 8M). Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles well developed; MII as long as MI, left plate with four teeth and right plate

Figure 9. Scoletoma parvapedata (Treadwell) (Syntype USNM16019). (A) Anterior end in lateral view; (B) anterior parapodium; (C) posterior parapodium; (D) simple multidentate hooded hook; (E) mandible; (F) maxillary apparatus. Scoletoma robusta (Ehlers) (Syntype MCZ747). (G) Simple multidentate hooded hook; (H) mandible; (I) apparatus maxillary. Scoletoma tenuis (Verrill) (Syntype USNM57439). (J) Anterior parapodium; (K) posterior parapodium; (L) simple multidentate hooded hook; (M) mandible; (N) maxillae IÎI with carriers. Scale bars: A, 1 mm; B,C,J&K, 0.1 mm; D,G&L, 0.01 mm; E,F,M,N,H&I, 0.2 mm. Journal of the Marine Biological Association of the United Kingdom (2001)

616

L.F. Carrera-Parra Lumbrineridae from the Grand Caribbean region

with ¢ve teeth, with a portion of the posterior membrane partially sclerotized; MIII with two teeth, the distal bigger; MIV with pointed tooth; MV free. Remarks

Mature specimens with small dark swelling in ventral position close to the base of median and posterior parapodia. The beginning of simple multidentate hooded hooks is size dependent (r0.825 P0.0001).

Distribution

Florida, Gulf of Mexico and Venezuela. Scoletoma parvapedata (Treadwell, 1901) comb. nov. Figure 9A ^ F

Lumbriconereis parvapedata Treadwell, 1901:198, ¢gures 38^40. Lumbrineris paravapedata ö Hartman, 1942a:118^119, ¢gure 10 I,J & M. Material examined

Syntypes of Lumbriconereis paravapedata Treadwell USNM16019 (2), Ensenada Honda, Culebra. Fish Hawk 1898^1899. Description

The syntypes are two anterior and one posterior fragments, the longest with 415 setigers (L104.5 mm, W102.2 mm). Prostomium conical, short, as long as wide, with a pair of nuchal organs. Peristomium distinctly shorter than prostomium, as long as ¢rst setigers, shallow separation between rings, without ventral lip. First ¢ve parapodia smaller, the ¢rst three in a more dorsal position (Figure 9A); presetal lobe short and postsetal lobe digitiform; parapodia more developed in median and posterior segments (Figure 9B,C). Pygidium with two pairs of anal cirri. Setae limbate; simple multidentate hooded hooks, almost all setae broken in anterior setigers, in posterior setigers, hooks with short hood (Figure 9D); acicula yellow. Mandibles divided for a short distance posteriorly (Figure 9E); maxillary apparatus with ¢ve pair of maxillae (Figure 9F). Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles; MII with six teeth, as long as MI, with a small portion of the posterior membrane partially sclerotized; MIII with two teeth; MIV with short blunt tooth; MV free.

Discussion

Treadwell (1901) described hooks from second setiger; this information can not be con¢rmed because of the poor condition of the material.

Distribution

Puerto Rico. Scoletoma robusta (Ehlers, 1887) comb. nov. Figure 9G ^ I

Lumbriconereis robusta Ehlers, 1887:104^105, plate 31, ¢gures 1^6. Material examined

Syntypes of Lumbriconereis robusta Ehlers MCZ742 (1), o¡ Havana, Cuba, 320 m; MCZ747 (1); Non-type material, MMI G812 (1) RV `Gerda', cruise 6617, o¡ Miami, Florida Journal of the Marine Biological Association of the United Kingdom (2001)

(25833'N 80804'W), 22 November 1966; MMI G823 (5) RV `Gerda', cruise 6709, o¡ Miami, Florida (25835'N 80800'W), 7 July 1967, 174 m, MMI G850 (1) RV `Gerda', cruise 6713, o¡ Miami, Florida (25848'N 80803'W), 2 August 1967, 214 m; MMI G854 (1) RV `Gerda', cruise 6713, o¡ Florida (25827'N 80802'W), 25 August 1967, 221m; MMI P919 (1) RV `Pillsbury', cruise 6907, o¡ Guadeloupe, Lesser Antilles (16805'N 61819'W), 12 July 1969, 695 m. Description

Syntype (MCZ742) incomplete with 115 setigers (L106.4 mm W103.2 mm). Prostomium conical, as long as wide, with a pair of nuchal organs. Peristomium shorter than prostomium, separation between rings distinct dorsally and laterally, ventrally with a shallow lip. All parapodia well developed, but ¢rst six smaller; presetal lobe short, rounded and postsetal lobe digitiform with base in£ated in anterior parapodia, digitiform in posterior ones. Setae limbate, simple multidentate hooded hooks from setigers 33^55 (Figure 9G); acicula black. Mandibles fused for about half its length (Figure 9H); maxillary apparatus with ¢ve pairs of maxillae (Figure 9I). Maxillary carriers shorter than MI, anterior end constricted; MI forcepslike with bridles well developed; MII as long as MI, with ¢ve teeth, with a small portion of the posterior membrane partially sclerotized; MIII with one tooth and a basal hump; MIV with one pointed tooth; MV free.

Remarks

Mature specimens with dark swelling in ventral position close to the base of parapodia in the median and posterior segments. The beginning of simple multidentate hooded hooks is size dependent (r0.98 P50.001).

Distribution

Florida, Cuba and Guadeloupe, Lesser Antilles. Scoletoma tenuis (Verrill, 1873) Figure 9J ^ N

Lumbrineris tenuis ö Perkins, 1979:433^436, ¢gure 7. Scoletoma tenuis ö Frame, 1992:203, 215. Material examined

Syntype of Lumbrineris tenuis Verrill YPM2733 (1), Connecticut, New Haven Co. Stony, Creek. Non-type material, USNM57439 (10), Boca Ciega Bay, Tampa Bay, Florida (24843'26''N 82843'37''W), 1m; USNM67772 (1), o¡ Mobile Bay, Alabama, (29853'30''N 88812'27''W), 32 m; USNM75349 (2) o¡ Florida (26816'43''N 83847'40''W), 90 m; ECOSUR LUMB13 E4 EMOAPII (2) o¡ north of Veracruz; E7 PROMEX-III (1) Gulf of Mexico; E38 PROMEX-III (3) o¡ Terminos Lagoon, Campeche, Mexico, (19827'N 91856'W), 47 m; MIE5 (3) o¡ Punta San Juan, Veracruz, Mexico (18820'N 94838'W); MIE6 (3) o¡ Coatzacoalcos, Veracruz, Mexico (18811'N 94825'W); MIE14 (2) (18823'N 93848'W), o¡ Carmen Lagoon, Tabasco, Mexico. Description

Syntype with 67 setigers (L102.9 mm W100.4 mm) broken into three fragments. Prostomium slightly conical, as long as wide, with a pair of nuchal organs. Peristomium

Lumbrineridae from the Grand Caribbean region shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a shallow lip, both rings of similar size. All parapodia well developed; presetal lobe rounded short and postsetal lobe conical, more elongated in posterior parapodia (Figure 9J,K). Pygidium with two pairs of similar anal cirri. Setae limbate, simple multidentate hooded hooks from setigers 10^18, with short hood (Figure 9L); acicula yellow. Mandibles fused for about half its length (Figure 9M); maxillary apparatus with ¢ve pairs of maxillae. Maxillary carriers shorter than MI, anterior end slightly constricted; MI forceps-like with bridles well developed, with rough internal border (Figure 9N); MII with 5^6 teeth, as long as MI, with a small portion of the posterior membrane partially sclerotized; MIII unidentate; MIV with one pointed tooth; MV present. Remarks

Uebelacker (1984) described Lumbrineris sp. B and regarded it as similar to S. tenuis because of the maxillary apparatus, but di¡ering by having an early beginning of the hooks, postsetal lobe more elongated in anterior parapodia and mandibles fused in two thirds of its total

L.F. Carrera-Parra

617

length. The beginning of the hooks is a size dependent character; all specimens reviewed by her were incomplete with no more than 47 setigers, the postsetal lobe is longer in more posterior parapodia, and the mandibles of her specimens are fused about half its total length as in the type material. Lumbrineris sp. B (partim USNM57439 and USNM67772) is herein considered as conspeci¢c with S. tenuis. Distribution

Massachusetts to Florida and Gulf of Mexico. Scoletoma testudinum (Augener, 1922) comb. nov. Figure 10A ^ D

Lumbriconereis testudinum Augener, 1922:46 Lumbrineris testudinum ö Perkins, 1979:439^441, ¢gure 9D ^ H. Material examined

Holotype of Lumbriconereis testudinum ZMB6399, Tortugas, south-west Channel, Florida. Non-type materials, ECOSUR LUMB14 MIE6 (1) o¡ Coatzacoalcos,Veracruz, Mexico (18811'N 94825'W); M1E11 (1) o¡ Tonala,Veracruz, Mexico (18825'N 94813'W); HU1 (1) Hualaxtoc, Quintana Roo, Mexico.

Figure 10. Scoletoma testudinum (Augener) (Holotype ZMB6399). (A) Anterior parapodium; (B) posterior parapodium; (C) simple multidentate hooded hook from anterior setiger; (D) simple multidentate hooded hook from posterior setiger. Scoletoma treadwelli (Hartman) (Syntype USNM16018); (E) anterior end; (F) anterior parapodium; (G) posterior parapodium; (H) simple multidentate hooded hook from anterior setiger; (I) simple multidentate hooded hook from posterior setiger; (J) mandible; (K) maxillary apparatus. Scoletoma verrilli (Perkins) (Paratype USNM57442); (L) simple multidentate hooded hook from anterior setiger; (M) large simple multidentate hooded hook from posterior setiger; (N) small simple multidentate hooded hook from posterior setiger; (O) maxillary apparatus. Scale bars: A,B,F&G, 0.2 mm; C,D,H,I,L,M&N, 0.01 mm; E, 0.5 mm; K, 0.2 mm. Journal of the Marine Biological Association of the United Kingdom (2001)

618

L.F. Carrera-Parra Lumbrineridae from the Grand Caribbean region

Description

Holotype mature, complete with 93 setigers (L101.5 mm W100.5 mm), broken into two fragments. Prostomium globular, as long as wide; with a pair of nuchal organs. Peristomium shorter than prostomium; separation between rings distinct dorsally and laterally, ventrally with a shallow lip; both rings of similar size. All parapodia well developed, becoming shorter in posterior segments; anterior parapodia with presetal lobe rounded short and postsetal lobe digitiform with base in£ated; in posterior parapodia both lobes become conical (Figure 10A,B). Pygidium with two pairs of anal cirri, the dorsal pair longest. Setae limbate; simple multidentate hooded hooks from ¢rst setiger, in anterior setigers hooks with teeth of similar size, in posterior setigers hooks with a proximal tooth bigger and directed laterally (Figure 10C,D); acicula yellow. Mandibles translucent, anterior end rounded, fused for about half its length; maxillary apparatus with ¢ve pairs of maxillae. Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles well developed; MII with ¢ve teeth, as long as MI; MIII with one tooth; MIV with a short rounded tooth; MV present. Distribution

Florida, USA; southern Veracruz and Quintana Roo, Mexico.

Scoletoma treadwelli (Hartman, 1956) comb. nov. Figure 10E ^ K Lumbriconereis maculataTreadwell, 1901:198^199, ¢gures 42^44. Lumbrinereis maculataöTreadwell, 1921:103^104, plate 8, ¢gure 10, text-¢gures 378^385. Lumbrineris treadwelli Hartman, 1956:288. Material examined

Syntype of Lumbriconereis maculataTreadwell USNM16018 (2), Puerto Real, Puerto Rico, Fish Hawk 1898^1899, US Fish Commission Non-type material, ECOSUR LUMB15 E1L1T1 (1) Veracruz, Gulf of Mexico.

Description

Syntypes incomplete, large specimen with 112 setigers (L104.4 mm W102 mm), short one with 38 setigers (L103 mm W101.2 mm). Prostomium conical, slightly longer than wide; with a pair of nuchal organs. Peristomium shorter than prostomium, separation between rings distinct dorsally and laterally, ventrally with a shallow lip (Figure 10E). All parapodia well developed, but ¢rst ¢ve parapodia smaller; parapodia with presetal lobe short rounded and postsetal lobe digitiform with in£ated base in anterior parapodia; both lobes more developed in anterior parapodia (Figure 10F,G). Setae limbate; simple multidentate hooded hooks, in anterior setigers with long hood becoming short in posterior setigers (Figure 10H,I). Mandibles divided for a short distance posteriorly (Figure 10J); maxillary apparatus with ¢ve pairs of maxillae (Figure 10K). Maxillary carriers shorter than MI, anterior end constricted; MI forceps-like with bridles well developed; MII with four teeth, as long as MI, with a small portion of the posterior membrane partially sclerotized;


MIII with two teeth, the distal tooth longest; MIV with a pointed tooth; MV free. Discussion

Hartman (1956) replaced the name L. maculata with L. treadwelli because it was preoccupied.

Distribution

Florida, USA; Veracruz, Mexico; Cuba; Puerto Real, Puerto Rico and Guadeloupe, Lesser Antilles. Scoletoma verrilli (Perkins, 1979) Figure 10L ^ O

Lumbrineris verrilli Perkins, 1979:441^444, ¢gures 10^11; Uebelacker, 1984:(41) 24, 26, ¢gure 20A ^ H. Scoletoma verrilli ö Frame, 1992:203. Material examined

Holotype of Lumbrineris verrilli Perkins USNM57440, lower Tampa Bay, Florida (27833'27''N 82842'36''W), 5 m; paratypes USNM57442 (39) Biscayne Bay, Miami, Florida; USNM45701 (10) Tampa Bay, Florida; USNM57441 (28) lower Tampa Bay, Florida (27833'27''N 82842'36''W), 5 m; USNM57443 (3) Hutchinson Island, Florida (27822'08''N 80813'46''W), 10 m. Non-type material, ECOSUR LUMB16 E11COLT-I (1) o¡ Tamaulipas, Mexico, 18 m; E41COLT-I (3) o¡ north Veracruz, Mexico, 16 m; E24COLT-II (1) o¡ Altamira Tamaulipas, Mexico; E19-COLT-III (1) o¡ Tamaulipas, Mexico; E43COLT43 (1) Tamaulipas, Mexico; E11-COSTA (1) Tamaulipas, 27 m; E4N-PANUCO (1) o¡ northVeracruz; E3EMOAP-II (4) o¡ AltamiraTamaulipas, Mexico; E18EMOAPP-II (3) o¡ north Veracruz, 56 m; MIE1 (3) o¡ Sontecomapan, Veracruz, Mexico (18835'N 94852'W); MIE4 (1) o¡ Punta Zapotitlan, Veracruz, Mexico (18831'N 94838'W); MIE5 (7) o¡ Punta San Juan, Veracruz, Mexico (18820'N 94838'W); MIE6 (8) o¡ Coatzacoalcos, Veracruz, Mexico (18811'N 94825'W); MIE8 (1) o¡ Coatzacoalcos, Veracruz, Mexico (18831'N 94825'W); MIE11 (1) o¡ Tonala, Veracruz, Mexico (18825'N 94813'W); MIE12 (2) o¡ Tonala, Veracruz, Mexico (18814'N 94813'W); MIE14 (5) o¡ Carmen Lagoon, Tabasco, Mexico (18823'N 93848'W); IDP (1) Veracruz, Mexico; E1PROMEX-III (1), o¡ Nautla, Veracruz (20831.5'N 96858.1'W), 39 m; E7 PROMEX-III (2) Gulf of Mexico; E30PROMEX-III (1) o¡ Campeche, Mexico (18858'N 92802'W), 20 m; E48 PROMEX-III (1) Gulf of Mexico (19845'N 91808'W), 20 m; A70 (1) Xahuaychol, Quintana Roo, Mexico, 1m; MMI P368 (1) RV `Pillsbury', cruise 6607, o¡ Golfo Morrosquillo, Colombia (09831'N 75840'W), 13 July 1966, 36 m; MMI P619 (1) RV `Pillsbury', cruise 6802, o¡ Punta Sal, Honduras (15858'N 87834'W), 20 March 1967, 40 m. Description

Holotype complete with 160 setigers (L101.2 mm W100.3 mm). Prostomium slightly conical, as long as wide, with a pair of nuchal organs; Peristomium shorter than prostomium, separation between rings distinct dorsally and laterally, ventrally with a shallow lip. All parapodia well developed in anterior and posterior segments, but smaller in median segment. Presetal lobe short rounded and postsetal lobe rounded in anterior parapodia becoming digitiform elongated in posterior

Lumbrineridae from the Grand Caribbean region parapodia. Setae limbate; simple multidentate hooded hooks from ¢rst setiger, in anterior setigers with long hood, in median and posterior setigers with hooks of two di¡erent size, the uppermost bigger (Figure 10L ^ N); acicula yellow. Pygidium with two anal cirri. Mandibles translucent fused for about 0.75 its total length; maxillary apparatus with ¢ve pairs of maxillae (Figure 10O). Maxillary carriers as long as MI, anterior end constricted; MI forceps-like with bridles poorly developed; MII with 3^4 teeth, as long as MI, with a small portion of the posterior membrane partially sclerotized; MIII and MIV with one tooth; MV tiny. Distribution

Florida, Gulf of Mexico, Honduras and Colombia. Key to lumbrinerids recorded from the Grand Caribbean region

1. Maxillary apparatus with six pairs of maxillae; MI with at least one prominent internal tooth . . . . . Lysarete MI with two accessory teeth, MIII with 5 teeth, MIV with 4 teeth . . . . . . . . . . . . . . . . . Lysarete raquelae sp. nov. ö Maxillary apparatus with up to 5 pairs of maxillae; MI forceps-like (it could have small blunt teeth) . . . . . . . . 2 2 (1) MII shorter than MI (about half its size length), with connecting plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 ö MII as long as MI, without connecting plates . . . . . . . 4 3 (2) Connecting plates broad; with dorsal branchiae and nephridial papillae in median and posterior parapodia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kuwaita* ö Connecting plates narrow; without branchiae nor nephridial papillae . . . . . . . . . . . . . . . . . . . . . . . . . . Eranno1 ö MIII and MIV unidentate; median and posterior parapodia with presetal lobe longer than postsetal; acicula black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eranno bilabiata 4 (2) Mandibles fused over their total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lumbrinerides2 ................................................. 5 ö Mandibles divided at least for a short distance posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 5 (4) MI with accessory teeth . . . . . . . . . . . . . . . . . . . . . . . 6 ö MI without accessory teeth . . . . . . . . . . . . . . . . . . . . . . . 7 6 (5) MI with one accessory tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lumbrinerides crassicephala ö MI with two accessory teeth . . . . . . . Lumbrinerides dayi 7 (5) MII with teeth on the internal border, with composite and simple hooded hooks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lumbrinerides uebelackerae sp. nov. ö MII with posterior teeth in parallel, with simple hooded hooks . . . . . . . . . . . . . . . . . . . . Lumbrinerides jonesi 8 (4) MIV multidentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 ö MIV with up to two teeth . . . . . . . . . . . . . . . . . . . . . . . 12 9 (8) With parapodial branchiae . . . . . . . . . . . . . . . . Ninoe3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 ö Without branchiae . . . . . . . . . . . . . . . . . . . Lumbrineriopsis . . . . . . . . Peristomium with one ring; MIII unidentate; simple bidentate hooded hooks from setiger 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lumbrineriopsis cf. paradoxa 10 (9) MIII unidentate . . . . . . . . . . . . . . . . . . Ninoe nigripes ö MIII multidentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11 (10) Simple multidentate hooded hooks from setiger 1, mandibles black . . . . . . . . . . . . . . . . Ninoe wardae sp. nov. Journal of the Marine Biological Association of the United Kingdom (2001)

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ö Simple multidentate hooded hooks from median setigers (always after branchial region); mandibles only with black cores . . . . . . . . . . . . . . . . . . . . . Ninoe vargasi sp. nov. 12 (8) Maxillary apparatus with four maxillae. . . . . . . 13 ö Maxillary apparatus with ¢ve maxillae. . . . . . . . . . . 15 13 (12) MIV with whitish central area; mandibles with short divergent shaft . . . . . . . . . . . . . . . . . . . . . . Augeneria With three antennae; MIII and MIV unidentate; posterior parapodia with lobules not elongated; composite and simple multidentate hooded hooks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Augeneria bidens ö MIV without whitish central area; mandibles with long shaft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14 (13) MIV as edentate plate, with parapodial branchiae and one nuchal antennae . . . . . . . . . . . . . . . . . Cenogenus4 ö MIV as broad rectangular plate with tooth prominent protruding from middle of its border; without branchiae or nuchal antennae . . . . . . . . . . . . . . . . . . . . . Abyssoninoe* 15 (12) With hooded hooks . . . . . . . . . . . . . . . . . . . . . . . . . 16 ö Without hooded hooks . . . . . . . . . . . . . . . . . Arabellonereis MIII bidentate; postsetal lobe well developed, elevated dorsally. . . . . . . . . . . . . . . . . . . . . . . Arabellonereis janerensis 16 (15) With composite spinigers. . . . . . . . . . . Lumbricalus5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 ö Without composite spinigers . . . . . . . . . . . . . . . . . . . . . 18 17 (16) MIII bidentate . . . . . . . . . . . . . . Lumbricalus januarii ö MIII multidentate, with up to 5 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . Lumbricalus januarii sensu Hartman 1944 18 (17) With composite and simple multidentate hooded hooks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lumbrineris6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 ö Only simple multidentate hooded hooks . . . . Scoletoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 19 (18) MIII with one tooth . . . . . . . . . . . . . . . . . . . . . . . . 20 ö MIII with two teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 ö MIII with four teeth. . . . . . Lumbrineris perkinsi sp. nov. 20 (19) Posterior parapodia with presetal lobe longer than postsetal . . . . . . . . . . . . . . . . . . . . . . . . . Lumbrineris nonatoi ö Posterior parapodia with presetal lobe shorter than postsetal . . . . . . . . . . . . . . . . . . . . Lumbrineris paucidentata 21 (20) MIV bidentate . . . . . . Lumbrineris salazari sp. nov. ö MIV unidentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22 (21) Composite multidentate hooded hooks with short appendage; peristomium with ventral lip transversally separated . . . . . . . . . . . . . . . . . . . . . . . Lumbrineris £oridana ö Composite multidentate hooded hooks with long appendage in anterior setigers, becoming shorter in posterior setigers; peristomium with complete ventral lip . . . . . . . . . . . . . . . . . . . . . . . . . . . Lumbrineris cf. latreilli 23 (18) MIII edentate. . . . . . . . . . . . . . . . . . Scoletoma candida ö MIII unidentate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 ö MIII bidentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 24 (23) Acicula black . . . . . . . . . . . . . . . . . . Scoletoma robusta ö Acicula yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25 (24) Posterior parapodia with more developed postsetal lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 ö Posterior parapodia with shorter postsetal lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scoletoma testudinum 26 (25) MI with bridle poorly developed; middle and posterior setigers with one simple multidentate hooded hook stouter than others . . . . . . . . . . . . Scoletoma verrilli ö MI with bridles well developed; all hooks similar . 27

620

L.F. Carrera-Parra


27 (26) Simple multidentate hooded hooks from ¢rst setiger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 ö Simple multidentate hooded hooks never anterior to setiger 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scoletoma tenuis 28 (27) First three parapodia dorsal to the lateral alignment; peristomium without muscular lip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scoletoma parvapedata ö All parapodia lateral; peristomium with muscular lip ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . Scoletoma elongata 29 (28) With nephridial papillae in median and posterior parapodia . . . . . . . . . . . . . . . . . . . . . . . Scoletoma branchiata ö Without nephridial papillae . . . . . . . . . . . . . . . . . . . . . 30 30 (29) Simple multidentate hooded hooks from ¢rst setiger, with long hood in anterior setiger becoming shorter in posterior one . . . . . . . . . . . Scoletoma treadwelli ö Simple multidentate hooded hooks from median setigers (21^50), all with short hood . . . . . . . . . . Scoletoma ernesti

General remarks. *, not recorded in the Grand Caribbean region. 1, Parker (1956 cit. Perkins & Savage, 1975) recorded E. bi¢liaris (Ehlers, 1901) in the Mississippi Delta; this record is doubtful because E. bi¢liaris was described from Chile. 2, Sol|¨ s-Weiss et al. (1995) recorded from Campeche, Mexico Lumbrinerides aberrans (Day, 1963), but this species was described from South Africa. Their specimens were apparently misinterpreted (number of accessory teeth of MI). Their specimens are L. crassicephala and L. dayi. 3, Ninoe brasiliensis Kinberg, 1865 and N. leptognatha Ehlers, 1900 were recorded by Granados-Barba & Sol|¨ s-Weiss (1994) and Sol|¨ s-Weiss et al. (1995) from the Gulf of Mexico. However, both species have a restricted distribution in South America (Orensanz, 1973, 1990); records on the Gulf of Mexico are doubtful. Other specimens collected from nearby localities from those in the cited works, belong to N. nigripes and N. vargasi sp. nov. 4, the only species of Cenogenus recorded from the Gulf of Mexico is C. brevipes (McIntosh, 1903, Cape Finisterre); it was recorded by Uebelacker (1984), Granados-Barba & Sol|¨ s-Weiss (1994) and Sol|¨ s-Weiss et al. (1995). I reviewed the type material of Lumbriconereis brevipes McIntosh (BMNH1921.5.1.1612) and the specimen recorded by Uebelacker (USNM90993). The latter is a di¡erent species but unassignable because of its poor condition, without parapodial branchia in anterior parapodia, and with a di¡erent number of teeth in the MIII. The other two records lack a description of the material examined. The species is not included in the key. 5 , Frame (1992), using Hartman's description of MIII of Lumbricalus januarii, (MIII with up to ¢ve teeth), considered the specimens recorded by Day (1973) and Uebelacker (1984), having MIII bidentate to be a distinct species, and named it L. dayi. However, examination of the original description of L. januarii, and additional material from the type locality, indicate that MIII is bidentate. The species is not included in the key; it should be re-evaluated from additional material from the region. 6, Lumbrineris cingulata has been reported by Frame (1992) and Sol|¨ s-Weiss et al. (1995) from the North Atlantic and Gulf of Mexico; however, the distribution of the species was restricted by Orensanz (1990) to the Magellanic biogeographic province. The records of L. cingulata in the Grand Caribbean region are doubtful. Journal of the Marine Biological Association of the United Kingdom (2001)

I wish to thank Kristian Fauchald (USNM), Eric Lazo-Wasem (YPM), Birger Neuhaus (ZMB), Ardis Baker Johnston (MCZ), James Cordeiro (AMNH), Leslie H. Harris (LACM-AHF); Miranda Lowe (BMNH) and Alejandro Granados-Barba (CPICML-UNAM) for making available the type and non type materials that made this study possible. I also wish to thank Diana Ibarza¨bal (IOH), NancyVoss (MMI), and K. Fauchald for making laboratory facilities available to work in their collections. I am grateful to Victor H. Delgado-Blas (Universidad de Quintana Roo) andJuan M.Vargas-Herna¨ndez (UniversidadVearacruzana) for the specimens donated for this study. Special thanks to Sergio I. Salazar-Vallejo (ECOSUR-Chetumal) for encouraging me in the development of this work and for his critical reading of the manuscript. I also thank Scott Monks and Angel de Leo¨n-Gonza¨lez for their help in the preparation of the ¢nal version of this manuscript. This study was partially ¢nanced by a research fellowship from El Colegio de la Frontera Sur and by CONACyT (32529-T).

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