Annelida: Polychaeta: Polynoidae

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Revision of Halosydna Kinberg, (Annelida: Polychaeta: Polynoidae) from the Tropical Eastern Pacific and Grand Caribbean with descriptions of new species Patricia Salazar-Silva

a b

a

Departamento de Ecología Acuática, El Colegio de la Frontera Sur, Unidad Chetumal, Quintana Roo, Mexico b

Departamento de Ingenierias, Instituto Tecnológico de Bahía de Banderas, Nayarit Mexico Published online: 22 Mar 2013.

To cite this article: Patricia Salazar-Silva (2013): Revision of Halosydna Kinberg, (Annelida: Polychaeta: Polynoidae) from the Tropical Eastern Pacific and Grand Caribbean with descriptions of new species, Journal of Natural History, 47:17-18, 1177-1242 To link to this article: http://dx.doi.org/10.1080/00222933.2012.752934

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Journal of Natural History, 2013 Vol. 47, Nos. 17–18, 1177–1242, http://dx.doi.org/10.1080/00222933.2012.752934

Revision of Halosydna Kinberg, 1856 (Annelida: Polychaeta: Polynoidae) from the Tropical Eastern Pacific and Grand Caribbean with descriptions of new species Patricia Salazar-Silva* Departamento de Ecología Acuática, El Colegio de la Frontera Sur, Unidad Chetumal, Quintana Roo, Mexico; and Departamento de Ingenierias, Instituto Tecnológico de Bahía de Banderas, Nayarit Mexico

Downloaded by [189.164.77.232] at 16:59 11 May 2013

(Received 8 January 2012; final version received 19 November 2012; first published online 22 March 2013)

The genus Halosydna is a member of the family Polynoidae, comprising a group of polychaete annelids commonly known as scale worms. The main results in this study are the redescriptions of 14 species. Four new species are described from the Mexican Pacific. Halosydna leucohyba is newly recorded for the Mexican Caribbean, and H. leius and H. tuberculifer from the Mexican Pacific. The identities of some widely recorded species are clarified, among them H. brevisetosa, H. glabra, H. fuscomarmorata and H. parva. Finally, H. nebulosa and H. virgini are re-established as valid species and a dichotomous key for Halosydna specimens from the tropical eastern Pacific, Grand Caribbean, and other worldwide localities is included. http://www.zoobank.org/urn:lsid:zoobank.org:pub:AE52ECA9-090C-469B-9503557B3DBB24BC Keywords: Polynoids; Gulf of California; Mexican Pacific; Mexican Caribbean; systematics

Introduction The generic name Halosydna was first proposed by Kinberg (1856: 384) for Halosydna patagonica, as the type species for the genus. The genus belongs to the Polynoidae, a family distributed from the intertidal zone to the deep sea, the members are commonly known as scale worms because they are characterized by the presence of pairs of elytra on the dorsum. There are about 21 currently accepted Halosydna species, which are widely distributed in both hemispheres. Ten species are known from the tropical eastern Pacific whereas only one occurs in the Grand Caribbean (Salazar-Vallejo 1996; Salazar-Vallejo and Londoño-Mesa 2004). Individuals of this genus have been described living as commensals, as epibionts of bivalve molluscs, or among rhizomes of sea grasses. Hartman (1938) redescribed the genus, with members having 36 segments instead of 37 as mentioned by Kinberg (1856, 1858), and 18 pairs of elytra. Hartman (1938) also reassigned several species to Halosydnella Hartman, 1938, and recognized 15 Halosydna species as valid. Hartman (1949) redescribed and synonymized some of Kinberg’s species, while Hartman (1939a) reported the known species from the *Email: [email protected] © 2013 Taylor & Francis

1178 P. Salazar-Silva


American Pacific and provided most of the known records for the tropical eastern Pacific: some of the species described from the South Pacific were recorded in the Gulf of California, whereas others described in the North Pacific were recorded in the south-eastern Pacific. Like many Polynoidae, members of several Halosydna species are either reported as having wide distributions along the eastern Pacific or as having an amphi-American distribution. In turn, members of many species are poorly described, with numerous synonymies, a lack of recent regional faunistic studies, and few redescriptions. Hence, the purpose of this contribution is to review the available type and non-type material of Halosydna species recorded from the tropical eastern Pacific and in the Grand Caribbean region. Redescriptions of species considered valid were made in a standardized manner, detailing the ornamentation of elytra, a feature which has proved reliable for establishing species (Salazar-Silva 2010). Material and methods Type and non-type material of Halosydna species recorded in the Grand Caribbean and tropical eastern Pacific were examined. In addition, samples were collected at localities in the Mexican Pacific and Mexican Caribbean. Specimens were obtained from beneath rocks, as epibionts of oysters, and from among sea grass rhizomes. Specimens were fixed with 10% seawater–formalin, preserved in 70% ethanol and subsequently dissected and examined using a stereomicroscope. Parapodia, chaetae and elytra were examined using a compound microscope. Drawings were made using a camera lucida and photographs were taken with a digital camera. Scanning electron micrographs were taken by Electron Services of ECOSUR-Tapachula, Mexico. Specimens examined are deposited in the following collections and institutions: Museum acronyms used are as follows: AMNH, American Museum of Natural History, New York City, USA; ECOSUR, Reference Collection, El Colegio de la Frontera Sur-Chetumal, Mexico; LACM-AHF, Allan Hancock Foundation Polychaete Collection, Natural History Museum of Los Angeles County, Los Angeles, CA, USA; MCZ, Museum of Comparative Zoology, Harvard University, USA; SMNH, Swedish Museum of Natural History, Department of Invertebrate Zoology, Sweden; USNM, Smithsonian Institution, National Museum of Natural History, USA; YPM, Peabody Museum of Natural History, Yale University, USA; ZMB, Zoologisches Museum, Museum für Naturkunde der Humboldt Universitat, Berlin, Germany; ZMH, Zoologisches Institut and Museum der Universitat Hamburg, Germany. Abbreviations used: coll., collected; cr, cirrophore; dc, dorsal cirrus; el, elytron; fr, fringe; id., identified; ma, macrotubercles; mi, microtubercles; mp, micropapillae; ne, neurochaetae; no, notochaetae; nr, neuropodium; nt, notopodium; pr, prostomium; pa, parapodium; pt, principal tooth; st, subdistal tooth; vc, ventral cirrus. Taxonomy Class POLYCHAETA Grube, 1850 Family POLYNOIDAE Kinberg, 1856 Subfamily LEPIDONOTINAE Willey, 1902 Genus Halosydna Kinberg, 1856


Journal of Natural History 1179 Halosydna brevisetosa Kinberg, 1856 Halosydna fuscomarmorata (Grube, 1876) Halosydna glabra Hartman, 1939a Halosydna hartmanae (Kudenov, 1975) Halosydna johnsoni (Darboux, 1899) Halosydna latior Chamberlin, 1919a Halosydna leius (Chamberlin, 1919a) Halosydna leucohyba (Schmarda, 1861) Halosydna nebulosa (Grube, 1876) Halosydna nesiotes (Chamberlin, 1919b) Halosydna parva Kinberg, 1856 Halosydna patagonica Kinberg, 1856 Halosydna tuberculifer Chamberlin, 1919a Halosydna virgini Kinberg, 1856 Halosydna olgae sp. nov. Halosydna salazarvallejoi sp. nov. Halosydna silvamariae sp. nov. Halosydna riojaenriquei sp. nov. Class POLYCHAETA Grube, 1850 Family POLYNOIDAE Kinberg, 1856 Subfamily LEPIDONOTINAE Wiley, 1902 Genus Halosydna Kinberg, 1856

Type species Halosydna patagonica Kinberg, 1856: 385.

Diagnosis Body thin, sub-rectangular, with 36 segments. Prostomium bilobed, without cephalic peaks; two pairs of small eyes, both pairs on posterior half of prostomium; three antennae, median antenna with ceratophore inserted frontally, with long, thick, style, subdistally expanded and tip filiform, surface not papillose; lateral antennae with ceratophores inserted terminally as prolongations of prostomium, at same level of ceratophore of median antenna, style of similar shape to median antenna. Two palps. Pharynx with nine pairs of marginal papillae and two pairs of hard jaws. Tentacular segment not visible dorsally, tentaculophores lateral to prostomium, tentacular cirri similar to antennae. Body with 18 pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 25, 27, 28, 30, 31, 33. Posteriormost three segments with dorsal cirri. Elytra imbricated; margins with or without papillae; surface with papillae and/or tubercles (sclerotized or vesicular). Elytrophores rounded. Parapodia biramous. Notopodia shorter than neuropodia. Neuropodia distally truncated, with small rounded lobe near acicula tip. Dorsal cirri subdistally expanded and distally with filiform tip; cirrophore basally expanded. Dorsal tubercle inconspicuous. Ventral cirri shorter than neuropodia, tapering to fine tip. Notochaetae shorter than neurochaetae; with rows of spines; less abundant than neurochaetae; the smaller curved, with blunt tip, remaining ones slender tapering

1180 P. Salazar-Silva to capillary tips. Neurochaetae with rows of spines on upper region, tips entire or bidentate. Anus dorsal, between segments 35–36. Pygidium with one pair of anal cirri.


Remarks Members of Halosydna were described by Kinberg (1858) as having 18–21 pairs of elytra. However, all specimens examined in this study have 18 pairs, as defined by Hartman (1938). In addition, members of Halosydna species are recognized by eytral insertion: elytra of segments 27 and 28 are contiguous, also elytra of segments 30 and 31. Elytral arrangement is constant and important to confirm generic placement. Members of Halosydna have robust bodies, which neither autotomize nor lose elytra when collected or fixed. Members of all species consistently bear a small rounded lobe near the neuroacicular tip. Halosydna brevisetosa Kinberg, 1856 (Figure 1) Halosydna brevisetosa Kinberg, 1856: 385; Kinberg, 1858:18. Halosydna johnsoni (Darboux, 1899): Darboux, 1899:246 (partim). Halosydna brevisetosa: Hartman, 1949: 17–18; Salazar-Silva, 2006: 146–147.

Type material Two syntypes (SMNH 400) of Halosydna brevisetosa, Sausalito, San Francisco Bay, California, USA, Pacific North America, Eugenia Exped., 1851–53. One syntype (LACM-AHF POLY 1583) of Polynoe reticulata, San Pedro, CA, USA, rocky shores near low water mark, probable commensal with “huge amphitrite”, 2 August 1895, coll. & id. as P. reticulata by H. P. Johnson. One syntype (LACM-AHF POLY 1584) of P. reticulata, San Pedro, CA, USA, rocky shores near low water mark, probable commensal with “huge amphitrite”, 6 January 1896, coll. & id. as P. reticulata by H.P. Johnson.

Description Two syntypes in poor condition, but complete; description based on best preserved specimen. Body with 36 segments, 2.2 cm long, 0.3 cm wide, pigmentation absent. Prostomium bilobed, as wide as long, without cephalic peaks; facial tubercle pointed; two pairs of eyes, anterior pair dorsolateral on widest part of prostomium, posterior pair dorsal, near posterior margin of prostomium; median antenna with ceratophore inserted frontally, style subdistally expanded, tapering to filiform tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, at same level as ceratophore of median antenna, styles of similar shape to median antennae but shorter. Palps robust, tapering to filiform tips, surfaces with tiny papillae. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores ventral to prostomium, with slender chaetae; tentacular cirri similar in shape to antennae. Body with 18 pairs of elytra on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments cirrigerous; elytra fleshy,


Journal of Natural History 1181 overlapping, covering dorsum (Figure 1A). First pair of elytra circular (Figure 1B); with marginal papillae (Figure 1C); surface covered with tiny digitiform papillae; macrotubercles conical, sclerotized, some thin, distally acute and slightly curved, others conical-thick and prominent over elytrophore scar (Figure 1D,E). Second and third pairs of elytra with similar ornamentation (Figure 1A) but sclerotized tubercles shorter, distally rounded. Elytra in posterior half of body oval (Figure 1F), with short marginal papillae; surfaces with granular appearance (Figure 1F,G) with numerous microtubercles and micropapillae; microtubercles conical-truncate, sclerotized, with tiny cusps or distally rounded. Parapodia biramous, with small oocytes inside. Notopodia shorter than neuropodia. Neuropodia distally truncate, without projecting prechaetal lobe, with small rounded lobe near acicular tip. Dorsal cirri reaching neurochaetae tips, similar to antennae. Cirrophores cylindrical, basally expanded. Ventral cirri tapering to filifom tips. Nephridial papillae shorter on anterior segments. Anus dorsal, on segment 35. Pygidium wider than long, anal cirri absent. Notochaetae shorter than neurochaetae with rows of spines; the smaller thick, curved, with blunt tips; remaining ones tapering to capillary tips. Neurochaetae with rows of spines on upper region, tips entire. First pair of parapodia with spinous neurochaetae, thiner than remaining, with tips entire.

Remarks Halosydna brevisetosa can be distinguished by having all elytra margins fringed, with minute conical, truncate microtubercles, and conical macrotubercles on first pair of elytra; neurochaetae mainly unidentate, with only a tiny subdistal tooth in some midbody neuroparapodia. The second syntype of SMNH 400 with a damaged prostomium, 3.7 cm long, 36 segments, 18 pairs of elytra. Pharynx everted, with nine pairs of papillae, two pairs of jaws with smooth margins. This specimen appears to correspond to Kinberg’s (1858) description. The sytypes LACM-AHF POLY 1583 and LACM-AHF POLY 1584 of Halosydna johnsoni (Darboux, 1899) agree with those of H. brevisetosa. However, the other syntypes clearly differ so that both species are herein considered as valid and H. johnsoni is redescribed below. Members of this species have been extensively recorded from the tropical eastern Pacific, as well as worldwide. Here, the distribution is limited to California, USA. All other records are considered questionable. This species was synonymized with Halosydna nebulosa Marenzeller, 1902, and so later recorded in Japon (Imajima and Gamó 1970). The type materials of both species were examined and they are different species. Halosydna nebulosa is redescribed below.

Type locality Sausalito, San Francisco California, USA.

Distribution Known from California, USA: San Pedro, San Francisco.



Figure 1. Halosydna brevisetosa Kinberg, 1856. Syntype SMNH 400. (A) Anterior end, dorsal view; (B) first elytron; (C) same, fringe of papillae; (D, E) same, macrotubercles; (F) left elytron from middle segment; (G) same, fringe and microtubercles.

Journal of Natural History 1183 Halosydna fuscomarmorata (Grube, 1876) (Figures 2, 3) Polynoe fuscomarmorata Grube, 1876: 62–63. Halosydna fuscomarmorata: Augener, 1906: 117–119, plate 3, figures 41–44; Hartman, 1938: 109; Hartman, 1939a: 32, pl. 9, figures 119, 120 (partim); Seidler, 1924: 120-122; Salazar-Silva, 2006: 147.

Type material Holotype ZMB 1171, Payta, Perú, coll. Grube, Jelski leg.


Aditional material One specimen (LACM-AHF POLY 2433), Independencia Bay, Perú, 14◦ 14 08 S, 76◦ 08 30 W R/V Velero III, sta 380-35, shore, rock, 14 January 1935, coll. Allan Hancock Foundation, id. Hartman (1939).

Remarks The type material of Halosydna fuscomarmorata consists of two elytra and one midbody parapodium, dried, clearly belonging to the genus because of the shape of the neuropodium (prechaetal lobe with a small lateral lobe near the acicular tip), notochaetae and neurochaetae. Elytra without fringe of marginal papillae; with abundant microtubercles, sclerotized, rounded distally, scattered over the surface, some larger and more scattered. Notopodia shorter than neuropodia. Notochaetae shorter than neurochaetae, the smaller slightly curved with blunt tips, larger ones tapering to capillary tips. Neurochaetae thicker than notochaetae, with bidentate tips. Augener (1906) and Seidler (1924), described H. fuscomarmorata has having 18 pairs of elytra, with microtubercles along margins and some larger ones over the surface, as were illustrated by Augener (1906) and coincide with observations of available type material. Halosydna fuscomarmorata was recorded by Hartman (1939a) from Independencia Bay, Perú; Port Utria, Colombia and from Piñas Bay, Panamá. However, the features of these specimens agree with those of Halosydna elegans Monro (1928: 567) not Kinberg (1858) from Limon Bay (Panamá) in having small, soft, vesicular tubercles in addition to sclerotized on the three first pairs of elytra and posterior elytra. The specimen from Independencia Bay, Perú (LACM-AHF POLY 2433), examined by Hartman (1939a), agrees with type material of H. fuscomarmorata. The specimen LACM-AHF POLY 2433 lacks some posteriormost segments, but is in good condition. It is 1.6 cm long, 0.4 cm wide. Prostomiun bilobed; facial tubercle rounded, dark; two pairs of eyes; ceratophore of median antenna inserted frontally, style subdistally expanded and tip filiform; ceratophores of lateral antennae thinner, inserted terminally as prolongations of prostomium lobes. Palps robust abruptly tapering to small tips; segment two does not project over prostomium; tentaculophores with chaetae. The elytra are greyish (Figure 2A), fleshy, with smooth surfaces, without fringe of marginal papillae. The first pair of elytra are circular (Figure 2B), with abundant



Figure 2. Halosydna fuscomarmorata (Grube, 1876). LACM-AHF POLY 2433. (A) Anterior end, dorsal view; (B) first elytron; (C–E) same, macrotubercles; (F) right elytron from middle segment; (G, H) same, marginal microtubercles.

sclerotized microtubercles, conical, scattered on the surface, the largest over the elytrophore scar (Figure 2C) and the smaller around the margin (Figure 2D,E). Second pair of elytra and posterior elytra oval (Figure 2F), with papillate surfaces

Journal of Natural History 1185


and microtubercles short, sclerotized, conical-truncated, abundant around margins (Figure 2G,H). Parapodia biramous. Notopodia shorter than neuropodia (Figure 3A). Neuropodia distally truncated, prechaetal lobe with small rounded lobe over acicular tip. Notochaetae shorter than neurochaetae, not reaching distal neuropodial margin, the smaller curved and blunt tips, remaining slender tapering to capillary tips. Neurochaetae with rows of spines on upper region, bidentate tips (Figure 3B,C). The record of H. fuscomarmorata from Bahía Magdalena and Cabo San Lucas, Baja California, Mexico, includes a short description which does not mention the elytral shape (Hartman, 1939b: 4) so that the presence of this species in Baja California is uncertain.

Type locality Payta, Perú.

Distribution Known from Perú: Payta and Independencia Bay. Halosydna glabra Hartman, 1939 (Figure 4) Halosydna glabra Hartman, 1939a: 35-36, plate 4, figures 43–50 (partim); Salazar-Silva, 2006:147. Lepidonotus hupferi: Hartman, 1939a: 43 (partim) (non L. hupferi Augener, 1918). Halosydna johnsoni (Darboux, 1899): Darboux, 1899:246 (partim).

Type material Holotype LACM-AHF POLY 13, Bahía Concepción, Baja California Sur, Mexico, 26◦ 41 40 N, 111◦ 51 05 W, R/V Velero, station 688-37, 21.9 m, mud sand, 16 March 1937, coll. Allan Hancock Foundation. One paratype LACM-AHF POLY 14, Bahía Concepción, Baja California Sur, Mexico, 26◦ 41 40 N, 111◦ 51 05 W, R/V Velero III, station 688-37, 21.9 m, mud sand, coll. Hancock Foundation 16 March 1937. One syntype LACM-AHF POLY 1591 of Polynoe reticulata, Pacific Grove, Monterey, California, USA, in tube of Amphitrite or Thelephus sp. 11 July 1896 coll. H.P. Johnson.

Additional material Mexico: one specimen (LACM-AHF POLY 2379) off Bahía Concepción, Baja California Sur, 26◦ 53 50 N, 111◦ 52 25 W, 12 fm, corallines, R/V Velero III, station 683-37, 21.9 m, 15 March 1937, coll. Allan Hancock Foundation. One specimen (LACM-AHF POLY 2380), Bahía Concepción, Baja California Sur, 26◦ 53 50 N, 111◦ 52 25 W, 21.9 m, corallines, R/V Velero III, station 683-37, Id. as Lepidonotus hupferi in Hartman (1939a), 15 March 1937, coll. Allan Hancock Foundation. Three



Figure 3. Halosydna fuscomarmorata (Grube, 1876). LACM-AHF POLY 2433. (A) Parapodium; (B) bidentate neurochaeta; (C) same, details of the neurochaetae tips.



specimens (LACM-AHF POLY 2383) from Bahía San Carlos, off Guaymas, Sonora, coll. R. Brusca, 24 February 1972. One specimen (LACM-AHF POLY 2384) from Puerto Lobos, Sonora, Intertidal Rocky, M. E., Niemil, coll. J. Kudenov, No. 251, 20 March 1969. One specimen (LACM-AHF POLY 2385) Puerto Peñasco, Sonora, Station Beach, MTZ, coll. J. Kudenov, sandy tide pool, under basalt boulder in sand or on rock, 19 January 1973. One specimen (LACM-AHF POLY 2386) Topolobampo, Sinaloa, on small rocks, No. 12, coll. Univ. Arizona and R. Brusca, in mid-intertidal 26 February 1971.

Description Holotype incomplete, with 25 segments, lacking posterior segments. Prostomium bilobed, elongate, without cephalic peaks; facial tubercle rounded, long; two pairs of eyes, anterior pair dorsolateral on widest part of prostomium, posterior pair near posterior prostomium margin; median antenna with ceratophore inserted frontally between prostomial lobes, style subdistally expanded with ring of pigmentation, distally tapering to fine tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, styles similar in shape to median antenna but shorter. Palps stout, surface annulate, with short papillae. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores ventral to prostomium; tentacular cirri similar in shape to median antenna. Segment 2 projecting anteriorly on to prostomium as short nuchal lobe with two small tubercles. Thirteen pairs of elytra, on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 25; surfaces with diffuse pigmentation, margins without fringe. First pair of elytra circular, surface with granular appearance, abundant tubercles (Figure 4A,B) and micropapillae. Macrotubercles sclerotized of two kinds: conical, truncated and hemispherical with wrinkled surface (Figure 4C–F). Elytra in median and posterior segments oval, surfaces with few micropapillae and microtubercles sclerotized (Figure 4G,H) over anterior half. Parapodia biramous. Notopodia shorter than neuropodia. Neuropodia distally truncate, with protruding acicular tip, prechaetal lobe with small rounded lobe near acicular tip. Dorsal cirri reach neurochaetal tips, similar to antennae and filiform tips. Cirrophores basally expanded. Elytrophores more prominent than dorsal tubercles. Nephridial papillae visible from segment 8, stouter and longer at mid-body. Notochaetae shorter than neurochaetae, the smaller curved, with blunt tips; remaining notochaetae slender tapering to capillary tips. Neurochaetae with rows of spines on upper region, bidentate tips.

Remarks The paratype (LACM-AHF 14) is in two pieces, but agrees with the holotype. Halosydna glabra can be distinguished by the absence of fringe papillae on elytra, and the two kinds of sclerotized macrotubercles: conical, truncate, with wrinkled surfaces, and hemispherical, almost ovoid, and bidentate neurochaetae. Halosydna glabra was described from Concepción, Bay, Mexico, and recorded also in Panama (Hartman 1939a). The features of Panama specimens from Station 444–35 do not agree with H. glabra, so its presence in Panama is doubtful.



Figure 4. Halosydna glabra Hartman, 1939a. Holotype LACM-AHF POLY13. (A) First elytron; (B–D) same, macrotubercles; (E, F) same details of the macrotubercles; (G, H) Same, microtubercles.

The specimens of Lepidonotus hupferi from Station 683-37 in Bahía Concepción (Hartman, 1939a) agree with H. glabra. This record, together with the new ones from the Mexican Pacific, suggest a distribution restricted to this region.

Journal of Natural History 1189 Type locality Bahía Concepción, Baja California, Mexico, muddy sand, 26◦ 41 40 N, 111◦ 51 05 W.

Distribution Known from California, USA (Monterrey); Sonora, Mexico (Bahía San Carlos, Puerto Peñasco, Puerto Lobos); Sinaloa (Topolobampo). Halosydna hartmanae (Kudenov, 1975) (Figures 5, 6)


Malmgrenia hartmanae Kudenov, 1975: 77–79, figure 2. Halosydna hartmanae: Hanley, 1987: 160; Salazar-Silva, 2006: 147.

Type material Holotype LACM-AHF POLY 1118, as Malmgrenia hartmanae, off Puerto Peñasco, Sonora, Mexico, 31◦ 10 N, 113◦ 50 W, among lateral chaetae of Aphrodita mexicana, brought up by shrimp trawler, 28 February 1971. coll. R. Brusca, id. J. Kudenov. One syntype (LACM-AHF POLY 1590) Monterrey County, USA, in tube of Amphitrite or Thelepus, coll. & id. by H.P. Johnson as Polynoe reticulata.

Additional material One specimen (LACM–AHF POLY 2462), Punta Cholla Sonora, Mexico, low intertidal, commensal in tubes of Thelepus setosus, coll. S.A. Glassell, 9 May 1941, id. by O. Hartman as Lepidametria gigas.

Description Holotype complete, with 30 segments (Figure 6A), 0.5 cm long, 0.2 cm wide. Prostomium bilobed, as long as wide; lateral and frontal lobes rounded, slightly separated anteriorly (Figure 5A); facial tubercle stout, long; two pairs of round eyes, dorsolateral anterior pair on widest part of prostomium, posterior pair partially covered by segment 2. Median antenna ceratophore inserted frontally between prostomial lobes, style subdistally expanded, distally tapering to fine tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, styles similar to to median antenna. Palps robust tapering to filiform tips, surfaces with small papillae. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores lateral to prostomium, chaetae present; tentacular cirri similar to median antenna. Segment 2 projecting on to prostomium as short nuchal lobe. First pair of elytrophores lateral to prostomium. Body with 16 pairs of elytrophores on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 25, 27, 28, 30; some elytra missing. Elytra overlapping, covering dorsum, thin, translucent (Figure 6A), surfaces with small dots of greyish pigment (Figure 6B). First pair of elytra missing. Elytra of anterior and posterior segments without fringe of papillae on margin; surfaces smooth; anterior half of elytra with patch



Figure 5. Halosydna hartmanae (Kudenov, 1975). Holotype LACM-AHF POLY 1118. (A) Anterior end, dorsal view; (B) elytron from middle segment; (C) same, microtubercles; (D) parapodium, posterior view; (E) notochaetae; (F) neurochaetae.



Figure 6. Halosydna hartmanae (Kudenov, 1975). Holotype LACM-AHF POLY 1118. (A) Body; (B) elytron from middle segment; (C) notochaeta; (D) neurochaeta; (E) neurochaeta tip.



of microtubercles (Figures 5B, 6B); posterior half smooth. Microtubercles truncate (Figure 5C). Notopodia shorter than neuropodia (Figure 5D). Neuropodia distally truncate with protruding acicular tip, prechaetal lobe with small rounded lobe over acicular tip. Dorsal cirri subdistally expanded and tapering to filiform tips; cirrophores basally expanded; elytrophores wider than dorsal tubercles. Ventral cirri taper to fine tips. Nephridial papilla not visible. Anus dorsal. Pygidium with one pair of anal cirri (one missing in holotype), similar to dorsal cirri. Notochaetae shorter than neurochaetae; the smaller curved with blunt tips; remaining notochaetae slender, tapering to capillary tips (Figures 5E, 6C). Neurochaetae with rows of spines on upper region, bidentate tips, main tooth straight, subdistal tooth shorter than main tooth (Figures 5F, 6D,E).

Remarks The holotype of Halosydna hartmanae may be a juvenile based on the low number of segments. However, the parapodia, chaetae and insertion of elytra correspond to Halosydna. The species was assigned to Malmgrenia and later referred to Halosydna (Hanley 1987). I agree with the aforementioned paper because the lateral antennae are inserted terminally (ventrally in Malmgrenia) and the elytra posterior to pair 12 alternate with one, instead of two dorsal cirri, and the segments 14 and 15 pairs do not alternate with one dorsal cirrus. However, neuropodia are truncate, without prechaetal and postchaetal lobes as in Malmgrenia. Halosydna hartmanae is distinguished by having elytra with smooth margins and smooth surfaces except for microtubercles forming a patch on the anterior half. The non-type material is a 4.5 cm long, 1.0 cm wide specimen, whose elytra from mid-body and posterior segments agree with the species description. Also, it has bidentate neurochaetae and the first and second pairs of elytra with conical, sclerotized microtubercles.

Type locality Off Puerto Peñasco, Sonora, Mexico (31◦ 10 N, 113◦ 50 W).

Distribution Known from Sonora, Mexico (Puerto Peñasco and Punta Cholla). Halosydna johnsoni (Darboux, 1899) (Figures 7, 8) Polynoe reticulata Johnson, 1897: 170–172, plate 7, figures 32, 41, plate 8, figure 47 (partim). Lepidonotus johnsoni Darboux, 1899: 246 footnote (replacement name). Polynoe californica Johnson, 1901: 387 footnote (replacement name). Halosydna californica: Treadwell, 1914: 180–181. Halosydna johnsoni: Hartman, 1939a: 34–35 synonymy (partim); Salazar-Silva, 2006:147.

Journal of Natural History 1193 Type material Syntype (LACM-AHF POLY 1585) of Polynoe reticulata Johnson, 1897, Pacific Grove, Monterey, County, California, USA, in tube of Amphitrite or Thelepus sp., coll. & id. H.P. Johnson. Two syntypes (LACM-AHF POLY 1582) of P. reticulata, San Pedro, Los Angeles, County, California, USA, rocky shore near low water mark, probably commensal with “huge Amphitrite”, 25 June 1895, coll. & id. H.P. Johnson. One syntype (LACM-AHF POLY 1581) of P. reticulata, San Pedro, California, USA, rocky shore, near low water mark, probably commensal with “huge amphitrite”, 25 June 1895, coll. & id. H.P. Johnson. One syntype (LACM-AHF POLY 1588) of P. reticulata, Avalon, Santa Catalina Island, California, USA, 31 May 1896, coll. H.B. Torrey, id. H.P. Johnson.


Description Syntype (LACM-AHF POLY 1585) complete, in good condition, subrectangular in cross-section, 36 segments, 2.6 cm long, 0.4 cm wide, some elytra detached (Figure 7A). Prostomium bilobed, cephalic peaks absent (Figure 7B); facial tubercle rounded, long; two pairs of eyes, anterior pair on widest part of prostomium, second segment partially obscuring posterior pair; median antenna with ceratophore inserted frontally on prostomial lobes, style slightly expanded subdistally, tapering to filiform tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, styles similar to median antenna. Palps robust with papillate surfaces. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores with some chaetae; tentacular cirri similar to median antenna. Segment 2 projecting on prostomium as short, rounded lobe with two small tubercles dorsally (Figure 7B). Body with 18 pairs of elytra on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. First three pairs of elytra fringed. First pair of elytra round (Figure 7C), with short fringe of marginal papillae (Figure 7D,E); surfaces with abundant microtubercles and macrotubercles sclerotized, conical-truncate, larger macrotubercles over elytrophore (Figure 7F,G). Second pair of elytra ovoid (Figure 8A); short fringe of marginal papillae (Figure 8B); surfaces granulate, with micropapillae and microtubercles; microtubercles shorter than on first pair. Elytra in posterior half of body without fringe of papillae (Figure 8C), surfaces smooth, with microtubercles mainly along posterior margin. Notopodia shorter than neuropodia (Figure 8D). Neuropodia distally truncate; prechaetal lobe with small, rounded lobe over acicular tip. Dorsal cirri similar to antennae, subdistally expanded, with pigmented band, distally tapering to filiform tips; cirrophore basally expanded. Elytrophores wider than dorsal tubercles. Ventral cirri long. Nephridial papillae from segment 2, thicker in median segments. Anus dorsal, near segment 36. Pygidium with two anal cirri, similar to dorsal cirri. Notochaetae shorter than neurochaetae, not reaching distal neuropodial ends, with rows of spines; the smaller slightly curved; blunt tips; remaining notochaetae slender, tapering to capillary tips. Neurochaetae with rows of spines on upper region; tips bidentate, subdistal tooth short (Figure 8E–G); some subacicular neurochaetae with subdistal tooth tiny (Figure 8H).



Figure 7. Halosydna johnsoni (Darboux, 1899). Syntype LACM-AHF POLY 1585. (A, B) Anterior end, dorsal view; (C) first elytron; (D, E) same, fringe of papillae; (F, G) same, microtubercles.

Remarks Polynoe reticulata Johnson, 1897, was a pre-occupied name, which was renamed to Lepidonotus johnsoni (Darboux, 1899), then to Polynoe californica (Johnson, 1901), and finally to Halosydna johnsoni (Hartman, 1939a).



Figure 8. Halosydna johnsoni (Darboux, 1899). Syntype LACM-AHF POLY 1585. (A) Left elytron from middle segment, (B) same, fringe of papillae; (C) elytron from posterior segments; (D) parapodium; (E) bidentate neurochaetae; (F–H) neurochaetae tips.

Halosydna johnsoni has been confused and synonymized with H. brevisetosa. However, according to Johnson (1897), the latter has all elytra with a fringe of papillae on margins (absent in mid-body and posterior elytra in H. johnsoni), those of anterior segments with prominent tubercles (smooth with only microtubercles along the posterior margin except for the first pair, which has abundant conical macrotubercles, in H. johnsoni) and neurochaetae with entire tips (bidentate in H. johnsoni).

1196 P. Salazar-Silva Other syntypes of H. johnsoni also examined were LACM-AHF POLY 1583; LACM-AHF POLY 1584; LACM-AHF POLY 46; LACM-AHF POLY 1589; LACMAHF POLY1587; LACM-AHF POLY 1591; LACM-AHF POLY 1586; LACM-AHF POLY 1590; LACM-AHF POLY1592. All of them have labels indicating that Johnson did not designate types. Hence, the syntypes represent his original material. However, some specimens do not agree with the description of H. johnsoni and are herein reassigned to other species.

Type locality


San Pedro, California, USA, San Diego, California, USA and Santa Catalina Island, USA.

Distribution California, USA (Monterey, San Pedro, Santa Catalina Island). Halosydna latior Chamberlin, 1919 (Figures 9, 10) Halosydna latior Chamberlin, 1919a: 1–2. Halosydna obtusa-cirrata Treadwell, 1937: 143–144, plate 1, figures 8–11.5. Halosydna latior: Hartman, 1939a: 32; Salazar-Silva, 2006: 148. Halosydna brevisetosa: Skobsberg, 1942: 482–489 (non Kinberg, 1856).

Type material Holotype (MCZ 2138) of Halosydna latior, Mussel Point, Laguna Beach, California, USA, coll. Hamilton, Field # PO3. One paratype (AMNH 2379) of Halosydna obtusacirrata Treadwell, 1937, East of Cedros Island, Baja California Sur, Mexico, 73.15 m, dried, 22 May 1936.

Additional material Three specimen (LACM-AHF POLY 2387) of Puerto San Carlos, Sonora, Mexico, 27◦ 57 15 N, 111◦ 04 45 W, rock shingle, shore, R/V Velero III, station 1091-40, 8 February 1940, coll. Allan Hancock Foundation. Commensal worm, found in shells with large hermit, commensal in shell apex inhabited by Petrochiros californiensis, Bouvier, id. as H. latior.

Description Holotype with 36 segments, 4.1 cm long, 1.2 cm wide, subrectangular in cross-section, pigmentation absent. Prostomium bilobed, as wide as long, slightly retracted into segment 2; lobes slightly separated, rounded, without peaks; facial tubercle short and rounded; two pairs of eyes, anterior pair on widest part of prostomium; posterior pair near posterior margin; median antenna with ceratophore inserted frontally, style


Journal of Natural History 1197 subdistally expanded and ring of pigmentation, distally tapering to fine tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, styles similar to median antenna. Palps smooth, papillae absent. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores lateral to prostomium, chaetae absent; tentacular cirri similar to antennae. Segment 2 with two small tubercles dorsally. First pair of elytrophores lateral to prostomium, prominent. Body with 18 pairs of elytra, on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. First pair of elytra missing. Second pair reniform (Figure 10A). Elytra in median and posterior segments oval. All elytra with fringe of short and thick marginal papillae; surfaces granular, with microtubercles and macrotubercles. Microtubercles conicalblunt (Figure 9A), scattered over entire surface. Macrotubercles over elytrophore scar, larger, circular as flattened disc-shape (Figure 9B). Notopodia shorter than neuropodia (Figure 9C). Neuropodia distally truncate, prechaetal lobe with small rounded lobe near acicular tip. Dorsal cirri reaching neurochaetae tips, with ring of pigment subdistally. Cirrophores basally expanded. Elytrophores wider than dorsal tubercles. Ventral cirri short, not reaching distal neuropodial margin. Nephridial papillae long in median segments. Anus dorsal. Pygidium with anal cirri missing. Notochaetae shorter than neurochaetae; with rows of spines;

Figure 9. Halosydna latior Chamberlin, 1919a. Holotype MCZ 2138. (A) Microtubercles from elytra of middle segment; (B) same, macrotubercles; (C) parapodium, posterior view; (D) notochaeta, detail; (E) unidentate neurochaetae.



Figure 10. Halosydna latior Chamberlin, 1919a. (A, B) Holotype MCZ 2138. (A) Right elytron of middle segment; (B) same, details. (C–G) paratype AMNH 2379 of Halosydna obtusa-cirrata, (C) first elytron; (D) left elytron from middle segment; (E) right elytron from middle segment; (F) same, macrotubercles; (G) same, microtubercles.



tapering to capillary tips (Figure 9D), the smaller curved with blunt tips. Neurochaetae with rows of spines on upper region, tips entire (Figure 9E).

Remarks The other specimens of H. latior measure from 7.9 cm long and 1.6 cm wide to 1.8 cm long and 0.8 cm wide. However, all coincide with regard to elytra, parapodia and chaetae. The holotype lacks the anterior elytra, which are present in non-type specimens, where the first three pairs show a fringe of marginal papillae (Figure 10C), granular surfaces with abundant microtubercles, macrotubercles over the elytrophore scar. The features of the remaining elytra agree with those of the holotype (Figure 10D–G). Although the paratype of H. obtusa-cirrata Treadwell, 1937, is dried, it is possible to distinguish the fringe and microtubercles on all elytra (Figure 10C–E) and the macrotubercles on median elytra, like those present in H. latior (Figure 10E–G) Hence H. obtusa-cirrata is considered a synonym of H. latior. Halosydna latior was described from Laguna Beach, California, and the records of Hartman (1939a) and the paratype of H. obtusa-cirrata are from Baja California, Mexico. Rioja (1963) recorded the species from Topolobampo, Mexico. Skobsberg (1942) reported H. brevisetosa from Monterey and Carmel Bays, but the features described agree with H. latior.

Type locality Laguna Beach at Mussel Point, California, USA.

Distribution Laguna Beach, California, USA; Cedros Island, Baja California Sur, Mexico; Puerto San Carlos, Sonora, Mexico. Halosydna leius (Chamberlin, 1919) (Figures 11, 12) Lepidonotus leius Chamberlin, 1919a: 4. Halosydna johnsoni (Darboux, 1899): 246 (partim). Halosydna leius: Salazar-Silva, 2006: 148.

Material examined Holotype (MCZ 2142) Lepidonotus leius Chamberlin 1919, Laguna Beach, California, USA, coll. W.A, Hilbaun, dredge, Original locality label says “Laguna Beach Po. 8”. One syntype (LACM-AHF-POLY 1587), San Pedro, California, USA, rocky shore near low water, July 1895, coll. & id. as Polynoe reticulata by H.P. Johnson. One syntype (LACM-AHF POLY 46), San Diego, California, USA, coll. S.J. Holmes, 25 October 1896, id. as Polynoe reticulata by H.P. Johnson. One syntype (LACM-AHF POLY 1586), San Pedro, California, USA, rocky shore near low water mark, 9 January 1895, coll. & id. as Polynoe reticulata by H.P. Johnson. One syntype (LACM-AHF



POLY 1592), San Diego County, California, USA, on pilings in bay, 12 July 1895, coll. and id. as Polynoe reticulata by H.P. Johnson.

Additional materials Pacific, Mexico: one specimen (LACM-AHF POLY 2388), Santa Rosalía Bay, Baja California, Sur, 29◦ 54 20 N, 115◦ 48 20 W, R/V Velero III, station 610-37, 27.43 m, sand and kelp, 28 February 1937, coll. Allan Hancock Foundation, id. as H. parva in Hartman (1939a). One specimen (LACM-AHF POLY 2389), South Bay, Cedros Island, Baja California, 28◦ 04 45 N, 115◦ 21 05 W, 18.2–27.43 m, R/V Velero III, station 287-34, rock along margin of kelp bed, 10 March 1934, coll. Allan Hancock Foundation. One specimen (LACM-AHF POLY 2443) coll. in oil platform Hidalgo, 10 km southwest of Point Arguello, California, USA, 34◦ 29.70 N, 120◦ 42.13 W, subtidal, from Mytilus sp. and Corynactis california on support beams, station. HIHF008, L. Harris 24 March 2003. One specimen (ECOSUR P02577), off Villas las Rosas, Ensenada, Baja California, 31◦ 52 19.24 N, 116◦ 40 43.24 W, on sea grass roots, 6 March 2004. One specimen in two fragments (ECOSUR P02578), Villa las Rosas, Ensenada, Baja California, 31◦ 52 19.24 N, 116◦ 40 43.24 W, on sea grass roots, 5 March 2004. One specimen (ECOSUR P02579), Villas las Rosas, Ensenada, Baja California, 31◦ 52 19.24 N, 116◦ 40 43.24 W, on sea grass roots, 6 March 2004.

Description Holotype in good condition, complete; 36 segments, 1.2 cm long, 0.3 cm wide, posteriormost segments regenerating. Prostomium bilobed, wider than long; lobes without cephalic peaks; facial tubercle not examined; two pairs of dark, circular eyes (Figures 11A, 12A), anterior pair dorsolateral on widest part of prostomium, posterior pair near posterior margin of prostomium; median antenna with ceratophore inserted frontally on prostomium lobes, style missing; lateral antennae ceratophores inserted terminally as prolongations of prostomium lobes, styles subdistally expanded with ring of pigment, surfaces without papillae. Palps robusts, tapering to filiform tips. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores with chaetae; tentacular cirri similar to antennae. Segment 2 slightly projects over prostomium as short rectangular lobe with two small tubercles dorsally (Figures 11A, 12A). First pair of elytrophores directed forward, lateral to the tentaculophores. Body with 18 pairs of elytrophores on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33; three posteriormost segments with dorsal cirri; most elytra detached. First pair of elytra circular (Figure 11B) with sparse fringe of short marginal papillae (Figure 12B), surfaces with micropapillae, microtubercles and macrotubercles. Microtubercles sclerotized, amber-coloured, conical, abundant over all surfaces. Macrotubercles larger, sclerotized, conical-truncate or distally rounded with rough surface (Figures 11C, 12C,D). Elytra in median and posterior segments oval (Figure 11D–F), with fringe of short marginal papillae; surfaces granular, with some microtubercles sclerotized, conical, scattered on entire surface, other microtubercles not sclerotized, as vesicles (Figure 11D–G); some microtubercles truncate, distally rounded, with rough surfaces.



Figure 11. Halosydna leius (Chamberlin, 1919a). Holotype MCZ 2142. (A) Anterior end, dorsal view; (B) first elytron; (C) same, macrotubercles; (D) right elytron from middle segments; (E) left elytron from middle segments; (F) posterior elytron; (G) soft-macrotubercle; (H) parapodium posterior view.



Figure 12. Halosydna leius (Chamberlin, 1919a). Holotype MCZ 2142. (A) Anterior end, dorsal view; (B) elytra fringe; (C, D) first elytron, macrotubercles; (E) parapodia, posterior view; (F) bidentate neurochaetae.

Notopodia shorter than neuropodia (Figures 11H, 12E). Neuropodia distally truncate; protruding acicular tip; prechaetal lobe with small rounded lobe near acicular tip (Figure 11H). Dorsal cirri subdistally expanded, tapering to filiform tip. Cirrophores



expanded basally (Figure 12E). Elytrophores wider than dorsal tubercles. Anus dorsal. Pygidium thick, anal cirri missing. Notochaetae shorter than neurochaetae; with rows of spines; the shorter curved with blunt tips; remaining notochaetae slender tapering to capillary tips. Neurochaetae with rows of spines on upper region; bidentate tips, the main tooth slightly curved, subdistal tooth smaller (Figure 12F). First pair of parapodia with spinous neurochaetae with entire tips.

Remarks The distribution of H. leius is only known from the type locality. The additional materials examined here are from Ensenada, Baja California, and represent new records of H. leius in the Mexican Pacific. The non-type material shows the following: brownish pigmentation on ceratophores and midventrum; most specimens with elytra having green pigmentation and a black spot near the elytrophores; one specimen with elytra of orange pigment and a small facial tubercle; everted pharynx with nine pairs of papillae and two pairs of maxillae not fused to each other; anus dorsal; pygidium with two long cirri. The syntypes LACM-AHF POLY 46, LACM-AHF POLY 1586, LACM-AHF POLY 1587 and LACM-AHF POLY 1592 of H. johnsoni correspond to H. leius (Chamberlin 1919a).

Type locality Laguna Beach, California, USA.

Distribution Known from California, USA (Laguna Beach, San Pedro, San Diego); Baja California, Mexico (Ensenada); Baja California Sur, Mexico (Cedros Island and Santa Rosalia). Halosydna leucohyba (Schmarda, 1861) (Figure 13) Polynoe leucohyba Schmarda, 1861: 153–154, figure 308. Halosydna leucohyba: Webster, 1884: 309–310, plate 7, figures 16–18, plate 8, figures 19–20; Seidler, 1924: 122–124 (Synonym); Hartman, 1944: 9; Liñero-Arana, 1993: 25, plate 6, figures 1–8.

Additional material Caribbean: two specimens (ZMH V563), la Guayra, coll. Nepperschmidt, donor Nepperschmidt 26.07.1894, id. Hermann Augener in 1919. One specimen (YPM 1369), Castle Harbor, Bermuda, on coral rock, coll. A. Verrill, Expedition of 1898. One specimen (YPM 1155), Bermuda, coll. W.R Coe, 1903, id. Pettibone (1962). One specimen (LACM-AHF 2390), Punta Arenas, Puerto Rico, coll. W.G. Hewatt, 11 June 1946. One specimen (LACM-AHF POLY 2429), Dry Tortugas, Florida, USA, coll.


1204 P. Salazar-Silva A.L. Treadwell, June–July 1909. One specimen (LACM AHF 2436), Guana Island, British Virgin Islands, station BVI-99-11, 18.482◦ N, 64.575◦ W, North Beach, in front of beach house, on coral rubble overgrown by compact morph of Caulerpa racemosa, Amphiroa fragilisima tufts, and Cladophora sp., depth 2.74 m, coll. L. Harris, 25 July 1999, LH99-088. One specimen (LACM-AHF POLY 2437), station BVI-99-95, 18.476◦ N, 64.578◦ W, White bay, sample consists of Halimeda opuntia and associated epifauna, depth 5.48–12.8 m, 10 August 1999, coll. G. Hendler, LH99-772. One specimen (LACM-AHF POLY 2438), station BVI-00-62, 18.486◦ N, 64.583◦ W, Atlantic side of Long Point, 13 July 2000, among barnacles on vertical rock wall overgrown with calcareous and non-calcareous red algae, tunicates and sponges, on vertical rock wall, intertidal and shallow subtidal, coll. T. Zimmerman, G. Hendler, J. Martin, R. Ware, Vc0690. One specimen (LACM AHF POLY 2439), station BVI-00-62, 18.486◦ N, 64.583◦ W, Atlantic side of Long Point, among large barnacles on vertical rock wall overgrown with calcareous and non-calcareous red algae, tunicates and sponges, intertidal and shallow subtidal, 13 July 2000, coll. T. Zimmerman, G. Hendler, J. Martin, R. Ware, Vc0698. One specimen (LACM-AHF POLY 2440), station BVI00-6218.486◦ N, Atlantic side of Long Point, among large barnacles on vertical rock wall overgrown with calcareous and non-calcareous red algae, tunicates and sponges, intertidal and shallow subtidal, 13 July 2000, coll. T. Zimmerman, G. Hendler, J. Martin, R. Ware, Vc00700. One specimen (LACM-AHF POLY 2441), station BVI00-62, 18.486◦ N, 64.583◦ W, Atlantic side of Long Point, among large barnacles on vertical rock wall overgrown with calcareous and non-calcareous red algae, tunicates and sponges, intertidal and shallow subtidal, 13 July 2000, coll. T. Zimmerman, G. Hendler, J. Martin, R. Ware, Vc1356. One specimen (LACM-AHF POLY 2442), station BVI-00-62, 18.486◦ N, 64.583◦ W, Atlantic side of Long Point, among large barnacles on vertical rock wall overgrown with calcareous and non-calcareous red algae, tunicates and sponges, intertidal and shallow subtidal, 13 July 2000, coll. T. Zimmerman, G. Hendler, J. Martin, R. Ware, Vc1406. Venezuela: 10 specimens (LACM-AHF POLY 2391), Tortugas Island, 10◦ 58 00 N, 65◦ 23 75 W, coral shore, R/V Velero III Station A20-39, 13April 1939, coll. Allan Hancock Foundation. One specimen (LACM-AHF POLY 2430), Cubagua, 10◦ 48 48 N, 64◦ 13 13 W, station A30-39, Rocky Shore, R/V Velero III, 15 April 1939, coll. Allan Hancock Foundation. Mexican Caribbean, Quintana Roo: one specimen (ECOSUR P01328) of Rancho Buenavista, 18◦ 30 42 N, 87◦ 45 30 W, 3 m, coll. S. Salazar, on coral rock, 4 June 1998. One specimen (ECOSUR P01329), Rancho Buenavista, 18◦ 30 42 N, 87◦ 45 30 W, 2 m, on coral rock, coll. L. Carrera, 27 September 1996. One specimen (ECOSUR P01331), Xcalak, 18◦ 15 33 N, 87◦ 50 21.4 W, 20 m, 28 August 2002. One specimen (ECOSUR P01332), Xahuayxol, 18◦ 30 15 N, 87◦ 45 32 W, 2 August 2002. Four specimens (ECOSUR P01333), Punta Nizuc, 20◦ 02 N, 86◦ 44 W, 4 m, 10 February 2002. One specimen (ECOSUR P01334), Playa Aventuras, 20◦ 20 15 N, 87◦ 20 31 W, station DIF Aventuras, 24 May 1998. Five specimens (ECOSUR P01335), Contoy Island, 18◦ 30 8.4 N, 86◦ 47 45 W, 2 March 2001. Two specimens (ECOSUR P01336), Mahahual, 18◦ 40 9.6 N, 87◦ 43 1.4 W, 4 m, coll. R. Bastida, 21 March 1998. One specimen (ECOSUR P01337), Mahahual, 18◦ 40 9.6 N, 87◦ 43 1.4 W, st North side, 25 February 20001. Two specimens (ECOSUR P01338), Cozumel, 20◦ 23 45 N, 86◦ 51 53 W, station Playa Azul, coll. J. Ruiz, 25 March 2001. Two specimens (ECOSUR P01339), Punta Herradura, 18◦ 32 23 N, 87◦ 32 W, on coral rock, coll. L. Carrera and S. Salazar, 28 October 1997. One specimen (ECOSUR P01340), Punta



Nizuc, 21◦ 2 5.4 N, 86◦ 47 41.9 W, 4 m, 10 February 2001. One specimen (ECOSUR P01341), Contoy Sur, 21◦ 30 8.4 N, 86◦ 47 45 W, 2 m, 2 March 2001. One specimen (ECOSUR P01345), Xcacel, 2 m, 25 October 2002. One specimen (ECOSUR P0 1355), Punta Allen, 19◦ 46 46 N, 87◦ 28 33 W, coll. E. Donath, 24 February 1986. One specimen (ECOSUR OH-P0259), Islache reef, 21◦ 26 2.5 N, 86◦ 46 56.1 W, South of Contoy Island, 2 m, in coralline rock, coll. Salazar-Vallejo and Carrera-Parra, 25 February 2008.

Description One specimen (ECOSUR OH-P0259) entire. Body subrectangular, 36 segments, 2.3 cm long, 0.4 cm wide. Dorsum and elytra with brown pigment. Prostomium bilobed, partially retracted in second segment, without cephalic peaks; two pairs of faded eyes; facial tubercle small, rounded; median antenna with ceratophore inserted frontally, style without papillae, subdistally expanded, tapering to filiform tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, at same level as median antenna, with diffuse pigment, styles shorter than that of median antenna, papillae absent, subdistally clavate, abruptly tapering to filifom tip. Palps robust without pigmentation, surface with short papillae. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores with small, thin chaetae; tentacular cirri similar to antennae. Segment 2 projecting anteriorly as short, rectangular lobe on prostomium, with two small dorsal tubercles. Body with 18 pairs of elytra, inserted on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. Elytra fleshy, imbricate, covering dorsal midline (Figure 13A). First pair of elytra circular, with smooth margins (Figure 13C), surfaces with abundant microtubercles (Figure 13D). Microtubercles sclerotized, colour amber, conical, larger over elytrophore scar, others vesicular, distally blunt. Second and third pairs of elytra without papillae on margins; surfaces with sclerotized microtubercles, amber colour on anterior half, posterior half with smooth macrotubercles, not sclerotized (Figure 13E,F). Elytra in median and posterior segments without microtubercles (Figure 13G), with uncolored, vesicular macrotubercles (Figure 13H,I). Notopodia shorter than neuropodia. Neuropodia distally truncate with protruding acicular tip and small rounded lobe near acicular tip. Elytrophores wider than dorsal tubercles. Dorsal cirri subdistally expanded with ring of pigmentation, distally tapering to filiform tip; cirrophores basally expanded; ventral cirri tapering to fine tip, not reaching distal neuropodial margin; nephridial papillae elongate on posterior parapodia. Anus dorsal on last three segments. Pygidium with anal cirri missing. Notochaetae with rows of spines; the shorter, curved, with blunt tips; remaining notochaetae slender, tapering to capillary tips; neurochaetae with rows of spines on upper region; bidentate tips, main tooth thick, curved, subdistal tooth shorter than main tooth (Figure 13J).

Remarks Halosydna leucohyba (Schmarda, 1861) is widely recorded from the Grand Caribbean. Although Schmarda’s (1861) original description is brief, the features of the present



Figure 13. Halsydna leucohyba (Schmarda, 1861). ECOSUR OH-P0259. (A) Anterior end, dorsal view; (B) posterior end, dorsal view; (C) first elytron; (D) same microtubercles; (E) left elytron from posterior segment; (F) same macrotubercles; (G) right elytron from posterior segment; (H, I) same, macrotubercles; (J) bidentate neurochaeta.

Journal of Natural History 1207 specimens agree with later characterizations (e.g. Webster 1884; Hartman 1944). The morphology of elytra and neurochaetae are consistent in specimens of different size. The chaetae on the tentaculophores are very slender, few in number but present (although might be broken). The type material was not available in the Zoologisches Institut and Museum Hamburg.

Type locality Jamaica.


Distribution Bermuda; Florida; Puerto Rico; Venezuela; Virgin Islands; Mexican Caribbean, multiple records in this paper. Halosydna nebulosa (Grube, 1876) (Figures 14, 15) Polynoe (Halosydna) nebulosa Grube, 1876: 49. Halosydna nebulosa: Marenzeller, 1902: 567–569, plate 1, figure 1; Hartman, 1938: 109.

Type material One syntype (ZMB 1170) of Chefoo, China, Leg. Reimann, id. Grube.

Description Syntype complete, with 36 segments. Prostomium bilobed, wider than long, without cephalic peaks; facial tubercle present, without pigmentation; two pairs of eyes; median antenna with ceratophore inserted at same level as prostomium, style subdistally expanded, tapering to filiform tip, lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, at same level as ceratophore of median antenna, styles similar to median antenna. Palps robust taper to filiform tips, surfaces with small papillae. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores with slender chaetae, acicula tip emergent. Segment 2 with two small dorsal tubercles. Body with 18 pairs of elytra inserted on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. First pair of elytra rounded (Figure 14A) with short papillae on margins (Figure 14B,D), surfaces with microtubercles, papillae and macrotubercles. Microtubercles conicalslender, sclerotized, abundant, scattered over entire surface. Macrotubercles conicalslender, sclerotized, abundant (Figure 14C–F). Second pair of elytra with thick papillae on margins; surfaces with microtubercles and macrotubercles conical, sclerotized, amber, the microtubercles on anterior half, macrotubercles on posterior half. Elytra in median and posterior segments oval (Figure 14G,H), with papillae on margins and surfaces with microtubercles short, conical, distally blunt, scarce.



Figure 14. Halosydna nebulosa (Grube, 1876). Syntype ZMB 1170. (A, B) First elytron; (C) same, macrotubercles; (D) same, fringe; (E, F) same, macrotubercles; (G) left elytron from posterior segment; (H) same, microtubercles.

Notopodia shorter than neuropodia. Neuropodia distally truncate with protruding acicular tip, prechaetal lobe with small rounded lobe near acicula tip. Dorsal cirri similar to antennae. Cirrophores basally expanded. Elytrophores wider than dorsal



Figure 15. Halosydna nebulosa (Grube, 1876). Syntype ZMB 1170. (A) Neurochaetae; (B) lower notochaetae; (C) upper notochaetae; (D–F) unidentate neurochaetae.

tubercles. Nephridial papillae elongated in median and posterior segments. Ventral cirri taper to filiform tips (Figure 15A). Pygidium with two anal cirri similar to dorsal cirri. Notochaetae with rows of spines, the smaller curved, with blunt tips (Figure 15B); the larger slender, taper to capillary tips (Figure 15C). Neurochaetae with rows of spines on upper region (Figure 15F), tips entire, lightly curved (Figure 15D–E).

Remarks Halosydna nebulosa was redescribed by Marenzeller (1879, 1902). The syntype herein examined is from Chefoo (now Yantai city), China, This species has not been recorded

1210 P. Salazar-Silva in the Grand Caribbean or the tropical eastern Pacific, but is included to clarify their identity because this species is not considered valid and has been listed as a synonym of H. brevisetosa for Imajima and Gamó (1970). However, the latter has prominent macrotubercles on the anterior elytra, principally over the elytrophore mark, and microtubercles mainly around the margin, while the former has abundant macrotubercles and microtubercles scattered over all the elytral surface. Therefore, both species are herein redescribed separately, and the respective distributions are clarified.


Type locality Chefoo (now Yantai city), China.

Distribution Same as type locality. Halosydna nesiotes (Chamberlin, 1919) (Figure 16) Polynoe nesiotes Chamberlin, 1919b: 72–74, plate 8, figure 8, plate 9, figures 1–5. Malmgrenia nesiotes: Hartman, 1938: 122. Halosydna nesiotes: Hanley, 1987: 160; Salazar-Silva, 2006: 148.

Type material Holotype (USNM 19460) of Polynoe nesiotes Chamberlin, 1919b, Isla Santa Margarita, Baja California Sur, Mexico, Albatross expedition 1891, id. R.V Chamberlin.

Description Holotype in two fragments, not complete, anterior fragment with 14 segments, posterior fragment with 21 segments. Body dried, but the following features can be distinguished. Body with 18 pairs of elytrophores, seven on anterior fragment, 11 on posterior fragment, posteriormost segments with dorsal cirri. Elytra attached on anterior fragment absent on posterior fragment. Elytra without marginal papillae; surfaces granular due to numerous microtubercles (Figure 16A). The microtubercles conical-truncate, short, scattered over posterior half of surface (Figure 16B). Some elytra with tubercles hemispherical, large, sclerotized. Notopodia shorter than neuropodia. Neuropodia distally truncate, small rounded lobe near acicula tip. Dorsal cirri and cirrophore dehydrated (Figure 16C). Anus dosal. Pygidium with anal cirri missing. Notochaetae with rows of spines, the smaller curved, blunt tips, remaining ones slender tapering to capillary tips. Neurochaetae with rows of spines on upper region, tips bidentate, main tooth slightly curved, subdistal tooth shorter (Figure 16D–F).



Figure 16. Halosydna nesiotes (Chamberlin, 1919b). Holotype USNM 19460. (A) elytron from middle segment; (B) same, microtubercles; (C) parapodium, posterior view; (D) bidentate neurochaetae; (E, F) neurochaetae tips.

Remarks Although the holotype is dried, the parapodia, neurochaetae, number of elytra and segments agree with those of Halosydna as stated by Salazar-Silva (2006).



Halosydna nesiotes is distinguished by having elytra without marginal papillae, but with microtubercles conical-truncate (abundant and short on posterior half) and longer hemispherical tubercles in some elytra. Chamberlin (1919b) decribed the species as having a “shrunken” prostomium (but prostomial appendages were not described), 34 segments and 15 pairs of elytra. However, it was described as having “granular appearance due to the presence of papillae and tubercles subconical, all of the same type excepting for variation in size, those toward the periphery of the papillose area becoming smaller”. The second pair were described as similar to the first, and those of median and posterior segments as almost smooth, with minute microtubercles. The elytra margins were not described, but were illustrated as smooth, which agrees with present observations.

Type locality Santa Margarita Island, Baja California, Mexico.

Distribution Baja California, Mexico (Santa Margarita Island). Halosydna parva Kinberg, 1856 (Figures 17, 18) Halosydna parva Kinberg, 1856, 385; 1858: 17–18; 1910: 17–18, plate 5 figure 24; Seidler, 1924: 116; Hartman, 1939a, 33, plate 21, figure 265–267 (partim); Hartman 1949, 18–19, plate 1, figures 5–9 (partim); Salazar-Silva, 2006: 148.

Type material Lectotype (SMNH 402), Chincha Island, Peru, Leg Eugenie Expedition 1851, station 531, February to March, 1852, id. J. Kinberg.

Additional material Ecuador: one specimen (LACM-AHF POLY 2431), La Plata Island, Galapagos, 1◦ 16 S, 81◦ 05 10 W, R/V Velero III, station 22-33, shore, 22 January 1933, coll. Allan Hancock Foundation. id. O. Hartman. One specimen (LACM-AHF POLY 2434), Tagus Cove, Albemarle Island Galapagos, 0◦ 16 08 S, 91◦ 22 44 W, shallow water, coral, R/V Velero III, station 152-34, 14 January 1934, id. O. Hartman. Peru: One specimen (LACM-AHF POLY 2432), off Viejas Island, Independencia Bay, 14◦ 15 05 S, 76◦ 12 W, R/V Velero III, st. 374-35, 12 fm, sand and mud, 12 January 1935, coll. Allan Hancock Foundation, id. O. Hartman. One specimen (LACMAHF POLY 2433), Independencia Bay, 14◦ 14 08 S, 76◦ 08 30 W, R/V Velero III, station 380-35, shore, rock, 14 January 1935, coll. Allan Hancock Foundation, id. O. Hartman.



Description Lectotype complete but in poor condition. Body with 36 segments, pigmentation absent. Prostomium bilobed, without cephalic peaks, laterally rounded, wider than long, slightly retracted into second segment (Figures 17A, 18A); facial tubercle not examined; two pairs of eyes, both on posterior half of prostomium; median antenna with ceratophore inserted frontally on prostomial lobes, style missing; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, at same level of median antenna ceratophore, style missing. Palps missing. Pharynx partially everted with stout jaws. Tentacular segment not visible dorsally; tentaculophores with chaetae; tentacular cirri subdistally expanded, with short, filiform tips. Segment 2 not projecting over prostomium. First pair of elytrophores lateral to prostomium. Body with 18 pairs of elytra on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. First pair of elytra with thick marginal papillae (Figure 17B); surfaces with conical microtubercles sclerotized (Figure 17C,D). Second and third pairs of elytra

Figure 17. Halosydna parva Kinberg, 1856. Holotype SMNH 402. (A) Anterior end, dorsal view; (B) first elytron fringe; (C, D) first elytron, microtubercles; (E) notochaetae; (F) bidentate neurochaeta.



Figure 18. Halosydna parva Kinberg, 1856. Holotype SMNH 402. (A) Anterior end, dorsal view; (B) left elytron from middle segment; (C) right elytron from middle segment; (D) same, fringe; (E) same, macrotubercles; (F, G) microtubercles.

with microtubercles sclerotized and some macrotubercles distally rounded. Elytra in mid-body oval (Figure 18B,C), marginal papillae scarce (Figure 18D), surfaces with macrotubercles sclerotized, distally rounded (Figure 18E), microtubercles abundant



(Figure 18F,G). Elytra in posterior segments without marginal papillae, surfaces with microtubercles and macrotubercles of similar shape to those in anterior elytra. Notopodia shorter than neuropodia. Neuropodia distally truncate, prechaetal lobe with small rounded lobe near acicular tip. Dorsal cirri missing; cirrophores basally expanded. Elytrophores wider than dorsal tubercles. Ventral cirri taper to filiform tips. Anus dorsal. Pygidium thick with anal cirri missing. Notochaetae with rows of spines; the smaller curved with blunt tips (Figure 17E); remaining ones slender tapering to capillary tips. Neurochaetae, with lateral rows of spines on upper region; bidentate tips, subdistal tooth smaller than main tooth (Figure 17F).

Remarks The type material is labelled as holotype, but the species was originally described from several specimens. This specimen should be considered a syntype. However, it was redescribed by Hartman (1949) so herein it is being designated as Lectotype. The species was reported from Bahía Santa Rosalia, Baja California Sur, Mexico (Hartman 1939a). However, Salazar-Silva (2006) examined the specimen and corresponds to H. leius. Hence, H. parva does not occur in the Mexican Pacific. The non-type material ranged from 1.2 cm long and 0.3 cm wide, to 3.1 cm long and 0.55 cm wide, and shows a short rounded lobe projecting over the prostomium on segment 2, eyes round dark, antennae similar in shape to tentacular cirri and papillate palps. The type material of Halosydna virgini Kinberg, 1856, has been re-examined. Contrary to Hartman (1949), the species is considered as valid and so it is redescribed below.

Type locality Peru, Chincha Island.

Distribution Ecuador (la Plata Island; Tagus cove Island, Galapagos), Peru (Chicha Island, Independencia Bay). Halosydna patagonica Kinberg, 1856 (Figures 19, 20) Halosydna patagonica Kinberg, 1856: 385; 1858: 17, plate 5, figure 23; Hartman, 1949: 16–17, plate 2, figures 15–18.

Type material Holotype (SMNH 408), York Bay, Magellan Sound, Chile, 53◦ 34 S, 072◦ 20 W, Expedition Eugenie 1851-53.



Description Holotype in poor condition, in two fragments, lacking some median segments. Anterior fragment with 15 segments, posterior fragment with 11 segments (0.65 + 0.7 cm length, 0.55 cm width), body subrectangular in cross-section, pale-yellow. Prostomium bilobed (Figure 19A), without cephalic peaks; facial tubercle absent; two pairs of eyes, anterior pair on widest part of prostomium, posterior pair near posterior margin; median antenna with ceratophore, inserted frontally, style without papillae expanded subdistally, tip filiform; lateral antennae ceratophores inserted terminally as prolongations of prostomium lobes, at same level as median antenna ceratophore, styles subdistally tapering to filiform tips, surfaces smooth. Palps robust, surfaces smooth. Pharynx dissected and missing. Tentacular segment not visible dorsally; tentaculophores with slender chaetae; tentacular cirri similar to antennae. Segment 2 projects slightly over prostomium. Body with 15 pairs of elytrophores. Anterior fragment with eight pairs of elytrophores inserted on segments 2, 4, 5, 7, 9, 11, 13, 15. Posterior fragment with five pairs of elytrophores inserted on segments 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. First pair of elytra circular (Figure 19B), marginal papillae short, scarce; surfaces with macrotubercles, conical, sclerotized, scattered, smaller along margins, longer and curved over elytrophore scar (Figure 19B,C). Elytra of third pair with marginal papillae short and microtubercles scarce on surfaces. Microtubercles distally blunt, surfaces rough, with areola basally thicker and blunt on elytrophore scar. Elytra of median and posterior segments oval, with marginal papillae (Figure 19D–F), microtubercles thick, shorter. Notopodia shorter than neuropodia. Neuropodia distally truncate (Figure 20A,B), protruding acicular tips, prechaetal lobe with small rounded lobes near acicular tips. Elytrophores wider than dorsal tubercles. Dorsal cirri with surfaces smooth, similar to antennae. Cirrophores basally expanded. Ventral cirri taper to filiform tips. Nephridial papillae elongated in median and posterior segments. Anus dorsal. Pygidium with two anal cirri similar to dorsal cirri. Pharynx not everted. Notochaetae shorter than neurochaetae, with rows of spines; the smaller curved (Figure 20F), tips blunt; remaining ones tapering to capillary tips (Figure 20G). Neurochaetae with rows of spines on upper region (Figure 20C), tips bidentate, main tooth curved, subdistal tooth shorter (Figure 20D,E).

Remarks The holotype was originally described as having 36 segments and 18 pairs of elytra. It was labelled as syntype, but the description was based in a single specimen, which must be considered as the holotype. Halosydna patagonica has not been recorded in the Grand Caribbean or tropical eastern Pacific.

Type locality York Bay, Magellan Sound, Chile, 53◦ 34 S, 072◦ 20 W.

Distribution Same as type locality.



Figure 19. Halosydna patagonica Kinberg, 1856. Syntype SMNH 408. (A) Anterior end, dorsal view; (B) first elytron; (C) same, macrotubercles; (D, E) right elytron from middle segment; (F) same, microtubercles and margin with papillae.



Figure 20. Halosydna patagonica Kinberg, 1856. Syntype SMNH 408. (A) Parapodium, anterior view; (B) parapodium, posterior view; (C, D) bidentate neurochaeta; (E) neurochaeta tip; (F) notochaetae; (G) upper notochaetae.

Halosydna tuberculifer Chamberlin, 1919 (Figures 21, 22) Halosydna tuberculifer Chamberlin, 1919a: 2; Rioja, 1963: 148–153, figures 49–57. Halosydna species B Hartman, 1939a: 38, plate 22, figures 273–279; Salazar-Silva, 2006: 149. Material examined Pacific Ocean, Mexico: one specimen (LACM-AHF POLY 2435), Bahía San Juánico, Baja California Sur, 26◦ 11 50 N, 112◦ 29 05 W, R/V Velero III, station 617-37,



43.9 m, Velero, sand and kelp, 2 March 1937, coll. Allan Hancock Foundation, id. as Halosydna species B by O. Hartman. Two specimens (ECOSUR P02574), Villa las Rosas, Ensenada, Baja California, 31◦ 52 19.24 N, 116◦ 40 43.24 W, on sea grass roots, 6 March 2004. Two specimens (ECOSUR P02575), Villas las Rosas, Ensenada, Baja California, 31◦ 52 19.24 N, 116◦ 40 43.24 W, on sea grass roots, 6 March 2004. Two specimens (ECOSUR P02576), Villas las Rosas, Ensenada, Baja California, M31◦ 52 19.24 N, 116◦ 40 43.24 W, on sea grass roots, 6 March 2004. Description Specimen (ECOSUR P02574), complete, in good condition. Body with 36 segments, 1.6 cm long and 0.4 cm wide. Dorsum with brown pigment along dordal midline, ceratophores, and palps. Prostomium bilobed, wider than long, without prostomial peaks (Figure 22A); facial tubercle small, pigmented; two pairs of round, dark eyes, anterior pair dorsolateral on widest part of prostomium, posterior pair smaller, near posterior border; median antenna with ceratophore inserted frontally, dark pigmented, style missing; lateral antennae with ceratophores, inserted terminally as prolongations of prostomium, darkly pigmented, styles subdistally expanded, almost clavate, filiform distal tip. Palps robust, diffuse pigment and micropapillae, abruptly tapering to short tips. Pharynx everted, nine pairs of papillae distally, two pairs of amber-coloured jaws, not fused to each other. Tentacular segment not visible dorsally; tentaculophores with chaetae present; tentacular cirri similar to lateral antennae. Segment 2 projects over prostomium as short lobe with two small, lateral tubercles. Body with 18 pairs of elytra on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31. Three posteriormost segments with dorsal cirri. Elytra with small spots of greyish pigment, becoming diffuse towards posterior margin, unpigmented medially and over elytrophore scar. First pair of elytra circular (Figure 21A), short, numerous papillae on margins (Figures 21C, 22B); surfaces with blunt microtubercles, some macrotubercles conical, distally pointed (Figures 21B,C, 22C,D) and others distally blunt (Figures 22F). Elytra in median and posterior segments oval (Figure 21D), with marginal papillae (Figure 21D,E), surfaces granulate due to abundant microtubercles (Figure 21E), macrotubercles, and cylindrical micropapillae (Figure 21F); microtubercles and macrotubercles sclerotized, distally blunt. Notopodia shorter than neuropodia (Figure 22G). Neuropodia distally truncate, without distinct prechaetal and postchaetal lobes, small rounded lobe near acicula tip. Dorsal cirri similar to antennae, with ring of subdistal pigment. Cirrophores basally expanded. Elytrophores wider than dorsal tubercles. Nephridial papillae elongated on median and posterior segments. Ventral cirri tapering to filiform tip. Anus dorsal. Pygidium with anal cirri missing. Notochaetae shorter than neurochaetae; with rows of spines; the smaller curved, blunt tips (Figure 22H); remaining notochaetae slender taper to capillary tips. Neurochaetae with rows of spines on upper region, tips entire (Figure 22I). Neurochaetae are more spinous in first two chaetigers, distal ends capillary, entire. Remarks The specimen LACM-AHF POLY 2435 [reported by Hartman (1939a) as Halosydna species B] is in poor condition and incomplete, with 32 segments, 0.45 cm wide and



Figure 21. Halosydna tuberculifer Chamberlin, 1919a. ECOSUR P02574. (A) First elytron; (B) same, macrotubercle; (C) same, fringe; (D) right elytron from posterior segment; (E) same, fringe; (F) same, microtubercles.

1.6 cm long. However, it agrees with the specimen herein described, as well as with the features described by Chamberlin (1919a) for the specimens found in Laguna Beach, California. The species was not illustrated by Chamberlin but the original description refers to the presence of abundant conical tubercles on elytra, especially larger in the



Figure 22. Halosydna tuberculifer Chamberlin, 1919a. ECOSUR P02575. (A) Anterior end, dorsal view; (B) elytral fringe; (C, D) first elytron, macrotubercles; (E) same, conical macrotubercles; (F) same, rounded macrotubercles; (G) parapodium, posterior view; (H) notochaeta detail; (I) unidentate neurochaetae.

1222 P. Salazar-Silva first three chaetigers, which agrees with material examined in this study. The specimens reported by Rioja (1963) from Isla Cedros also agree with the original description. The type material was not examined because it cannot be found in the MCZ. New records of H. tuberculifer are included herein for Baja California, Mexico. The larger specimen being 2.3 cm long and 0.4 cm wide. All specimens show brownish spots on elytra, abundant blunt microtubercles, and some macrotubercles. Halosydna tuberculifer resembles H. brevisetosa in having unidentate neurochaetae and elytra with prominent macrotubercles, but the latter has median and posteriormost elytra only with microtubercles.

Type locality


Laguna Beach, California.

Distribution California, USA; Baja California, Mexico; Baja California Sur, Mexico. Halosydna virgini Kinberg, 1856 (Figure 23) Halosydna virgini Kinberg, 1856: 384; Kinberg, 1858: 15–16; Seidler, 1924: 120; Hartman, 1938: 109.

Type material Holotype (SMNH-Type 410), Honolulu, Hawaii Island, 21◦ 19 N, 157◦ 52 W, leg Eugenie, Expedition 1851–53.

Description Holotype incomplete; an anterior fragment with 19 segments, yellow-pale. Prostomium in poor condition. Seven pairs of elytra attached, overlapping. Elytra margins with thick, long papillae (Figure 23A,B), surfaces covered with conical microtubercles (Figure 23C), some truncate and others pointed. Notopopodia shorter than neuropodia (Figure 23D). Neuropodia distally truncate, prechaetal and postchaetal lobes indistinct, prechaetal lobe with small rounded lobe near acicular tip. Dorsal cirri detached. Cirrophores basally expanded. Elytrophores wider than dorsal tubercles. Ventral cirri tapering to filiform tip. Notochaetae shorter than neurochaetae, with rows of spines; the smaller chaetae shorter, curved, blunt tips; the remaining notochaetae slender tapering to capillary tips (Figure 23F). Neurochaetae with rows of spines on upper region; bidentate tips, main tooth curved, subdistal tooth short (Figure 23E).

Remarks The species, known from Hawaii, was originally described as having 35 segments, 18 pairs of elytra and bidentate neurochaetae. However, it was synonymized with



Figure 23. Halosydna virgini Kinberg, 1856. Holotype SMNH-Type 410. (A) First elytron; (B) same, fringe; (C) microtubercles; (D) parapodium, posterior view; (E) bidentate neurochaetae; (F) notochaetae.

Halosydna parva from Peru (Hartman 1949). Despite the present condition of the holotype, its features are clear: elytra margins with long papillae thicker than in H. parva and elytra lack the thick macrotubercles present in this species. Therefore, it must be considered a valid species.

1224 P. Salazar-Silva Type locality Honolulu, Hawaii Island, 21◦ 19 N, 157◦ 52 W.

Distribution Same as type locality. Halosydna olgae sp. nov. (Figures 24–26)


Halosydna species A Hartman, 1939a: 37, plate 4: figures 51–55; Salazar-Silva, 2006: 148–149.

Type material Holotype (ECOSUR 0132), Villa las Rosas, Ensenada, Baja California, Mexico, 31◦ 52 19.24 N, 116◦ 40 43.24 W, on seagrass roots, 6 March 2004. Paratypes: One specimen (ECOSUR 0133), Villas las Rosas, Ensenada, Baja California, Mexico, 31◦ 52 19.24 N, 116◦ 40 43.24 W, 6 March 2004. One specimen (LACM-AHF POLY 2467), Consag Rock, Gulf of California, Mexico, station 719-37, R/V Velero III, coll. Allan Hancock Foundation, 24 March 1937, id. as Halosydna, species A of Hartman (1939a). One specimen (LACM-AHF POLY 3608), Scammon’s Lagoon, Baja California Sur, 27◦ 50 N, 114◦ 13 W, 3.66 m, among shell, 14 September 1953, coll. J.W. Knudsen and D.S. Gorsline, R/V Horizon, station KG-8, id. as Halosydna species A of Hartman, 1939a. One specimen (LACM-AHF POLY 3609), Port San Bartolomé, Baja California Sur, Mexico, 1.1 miles from Kelp Point, 27◦ 41 06 N, 114◦ 53 38 W, R/V Velero IV, station 2603-54, from kelp in rocky intertidal, 11 February 1954, id. as Halosydna species A of Hartman (1939a). One specimen (LACM-AHF POLY 3610), off Redondo, California, USA, Flat Rock Point, 33◦ 47 50 N, 118◦ 24 25 W, rocky intertidal with Kelp and boulders, 18 October 1941.

Additional material Mexico, Pacific Ocean: One specimen (LACM-AHF POLY 3611), San Eugenio Point, Dewey channel, Baja California Sur, 27◦ 49 50 N, 115◦ 06 05 W to 27◦ 49 35 N, 115◦ 06 00 W, R/V Velero III, station 1260-41, 43.9 m, on coralline rock, 27 February 1941, id. as Halosydna species A of Hartman (1939a). Two specimens (LACM-AHF POLY 3612), Scammon’s Lagoon, Laguna ojo de liebre, Baja California Sur, 27◦ 50 N, 114◦ 13 W, 5.4–9.1 m, rock, coll. J.W. Knudsen and D.W. Gorsline R/V Horizon, station No. KG4, 14 September 1953, id. as Halosydna species A of Hartman (1939a). One specimen (LACM-AHF POLY 3613), off Pto. Malarrino, Baja California Sur, 27◦ 49 00 N, 114◦ 42 00 W, station 2025-51, R/V velero IV, shore rock, 18 April 1951, id. as Halosydna species A of Hartman (1939a).

Etymology This species is named in honour of Olga Hartman in recognition of her many contributions to the knowledge of polychaetes in the Pacific Coast.



Description Holotype complete, with 36 segments, 2.3 cm long and 0.3 cm wide. Dorsum with two transverse bands of dark pigment on each segment, cirrophores pigmented, two spots on notopodia (Figure 24A,B). Ventrum with transverse bands of dark pigment along middle line and laterally on each segment. Elytra with small spots of pale pigment, except over the elytrophores. Prostomium bilobed (Figure 24A), wider than long, laterally rounded with diffuse pigment; facial tubercle small; two pairs of eyes, anterior pair on widest part of prostomium, posterior pair dorsolateral, round and dark; median antenna with dark pigmented ceratophore, inserted frontally, style with diffuse brown pigment, subdistally expanded with ring of pigment, abruptly tapering to filiform tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, with brown pigment, style as long as prostomium, pigmented basally, similar to median antenna. Palps robust, surfaces with diffuse pigment and papillae, tapering to filiform tips. Pharynx everted, with nine pairs of papillae and two pairs of jaws. Tentacular segment not visible dorsally; tentaculophores unpigmented, with short chaetae; tentacular cirri of similar shape as antennae. Segment 2 projecting over prostomium as short lobe, with two small dorsal tubercles. Body with 18 pairs of elytra inserted on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. First pair of elytra circular, with short marginal papillae (Figure 24C–E), surfaces with microtubercles and macrotubercles. Microtubercles abundant, conical, sclerotized, distally blunt. Macrotubercles larger, vesicular, not sclerotized, hemispherical (Figure 24C,D). Elytra in median segments (Figure 24F) with few marginal papillae, surfaces with microtubercles and macrotubercles. Microtubercles conical short, sclerotized, numerous on anterior half. Macrotubercles hemispherical, vesicular, not sclerotized (Figure 24G), numerous on posterior half. Elytra in posterior segments without margin papillae, macrotubercles not sclerotized. Notopodia shorter than neuropodia. Neuropodia robust, prechaetal and postchaetal lobes indistinct, distally truncate; prechaetal lobe with a small rounded lobe near acicular tip. Dorsal cirri similar to antennae. Cirrophores basally expanded. Elytrophores wider than dorsal tubercles. Ventral cirri taper to fine tips. Nephridial papillae long, from segment 8. Anus dorsal; pygidium with two anal cirri similar to dorsal cirri. Notochaetae shorter than neurochaetae, with rows of spines, the smaller curved, with blunt tips; remaining ones slender, tapering to capillary tips. Neurochaetae with rows of spines, bidentate tips, subdistal tooth short. Chaetiger 1 neurochaetae thinner than remaining parapodia and sharp tip.

Remarks Paratype LACM-AHF POLY 2467 herein assigned was previously identified as a possible new species by Hartman (1939a), Halosydna species A. However, its features are not comparable with any other known species, which combined with examination of other material, shows the same features (Figures 25 and 26) and allows the formal description of this new species.



Figure 24. Halosydna olgae sp. nov. Holotype ECOSUR 0132. (A) Anterior end, dorsal view; (B) middle segments, dorsal view; (C) first elytron; (D) same, microtubercles; (E) same, fringe of papillae; (F) left elytron from middle segment; (G) same, microtubercles not sclerotized on posterior half.



Figure 25. Halosydna olgae sp. nov. Paratype LACM-AHF POLY 3610. (A) Left elytron from middle segment; (B) same, macrotubercles; (C) left elytron from posterior segment; (D–G) same, macrotubercles.



Figure 26. Halosydna olgae sp. nov. Paratype LACM-AHF POLY 3609. (A) First elytron; (B) same, macrotubercles; (C) right elytron from middle segment; (D) right elytron from posterior segment; (E) same, macrotubercles; (F) notochaetae, chorter; (G) bidentate neurochaeta; (H) neurochaeta tip.

Journal of Natural History 1229 Type locality Villa las Rosas, Ensenada, Baja California, Mexico.

Distribution California, USA; Gulf of California and Baja California Sur, Pacific Side. Halosydna salazarvallejoi sp. nov. (Figure 27) Halosydna sp 1 Salazar-Silva, 2006: 154


Type material Holotype (LACM-AHF POLY 3614), Scammon’s Lagoon, Laguna ojo de liebre, Baja California Sur, Mexico, R/V Hori-zon, station KG-8, 3.66 m, 14 September 1953, coll. J.W. Knudsen and Gorsline, id. as Halosydna species A of Hartman (1939a).

Etymology This species is named in honour of Sergio Salazar-Vallejo for his dedication to the taxonomy of the polychaetes along Mexico’s coast.

Description Holotype incomplete, an anterior fragment with 31 segments, 1.5 cm long, 0.3 cm wide. Dorsum and ventrum with dark pigment, also on ceratophores. Prostomium bilobed, without cephalic peaks; facial tubercle acute; two pairs of dark eyes, anterior pair on widest part of prostomium, posterior pair near posterior margin and smaller; median antenna with ceratophore, inserted frontally, style slightly subdistally expanded, tapering to filiform tip; lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, at same level as median antenna ceratophore, styles similar to median antenna. Palps missing. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores with slender chaetae; tentacular cirri similar to antennae. Segment 2 projecting over prostomium as short nuchal lobe, with two small tubercles. Body with 17 pairs of elytra inserted on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, all with diffuse pigmentation. First pair of elytra circular (Figure 27A) with few, short marginal papillae; surfaces with abundant micropapillae and macrotubercles. Macrotubercles prominent, vesicular, not sclerotized, almost ovoid, elongate, scattered over elytra surfaces (Figure 27A–C). Second elytra pair and median elytra with few short marginal papillae; vesicular macrotubercles, ovoid, not sclerotized (Figure 27D). Elytra in posterior segments with few short marginal papillae; vesicular macrotubercles ovoid, not sclerotized, shorter than in more anterior elytra (Figure 27E). Notopodia shorter than neuropodia. Neuropodia distally truncate with protruding acicular tip, prechaetal lobe with small, rounded lobe near acicula tip. Dorsal cirri



Figure 27. Halosydna salazarvallejoi sp. nov. Holotype LACM-AHF POLY 3614. (A) First elytron; (B, C) same, macrotubercles; (D) elytron from middle segment; (E) elytron from posterior segment; (F, G) notochaetae; (H) neurochaeta.

short, similar to antennae. Cirrophores short, basally expanded. Ventral cirri tapering to filoform tips. Nephridial papillae elongate on median and posterior segments. Anus dorsal; pygidium with two anal cirri missing.

Journal of Natural History 1231 Notochaetae shorter than neurochaetae with rows of spines; the smaller curved, blunt tips; remaining ones slender tapering to capillary tip (Figure 27F,G). Neurochaetae with lateral rows of long spines on upper region, tips bidentate, subdistal tooth short and thinner (Figure 27H).

Remarks


The holotype was previously labelled as Halosydna species A sensu Hartman (1939a). However, the features of these species differ. They coincide in having elytra with soft tubercles (i.e. not sclerotized), but H. salazarvallejoi sp. nov. lacks sclerotized microtubercles on the elytra, the marginal papillae are scarce, and the soft tubercles are prominent and abundant on elytra.

Type locality Scammon Lagoon, Baja California Sur, Mexico.

Distribution Same as type locality. Halosydna silvamariae sp. nov. (Figures 28, 29) Halosydna sp 2 Salazar-Silva, 2006: 154.

Type material Holotype (LACM-AHF POLY 3615), Scammon’s Lagoon, Baja California Sur, Mexico, R/V Hori-zon, station KG-8, 3.66 m, 14 September 1953, coll J.W. Knudsen and Gorsline id. as Halosydna species A of Hartman (1939a). One paratype (LACMAHF POLY 3616), Scammon’s Lagoon, Baja California Sur, Mexico, station KG-3, 5.5–8.2 m, R/V Hori-zon, 13 September 1953, coll. J.W. Knudsen and Gorsline, id. as Halosydna species A of Hartman (1939a).

Etymology The species name is derived from the name of the author’s mother.

Description Holotype complete, with 36 segments, 2.0 cm long and 0.4 cm wide. Prostomium bilobed, without cephalic peaks; facial tubercle small, rounded, with dark pigment; two pairs of eyes, anterior pair on widest part of prostomium, posterior pair near posterior margin; median antenna with ceratophore inserted frontally on prostomial lobes, style slightly expanded subdistally, tapering to short tip; lateral antennae with


1232 P. Salazar-Silva ceratophores inserted terminally as prolongations of prostomium lobes, styles similar to median antenna. Palps papillate, taper abruptly to small tips. Pharynx not everted. Tentacular segment not visible dorsally; tentaculophores short; tentacular cirri similar to antennae. Segment 2 projecting over prostomium as short nuchal lobe, with two small dorsal tubercles. Body with 18 pairs of elytra on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. Elytra with green pigment on midline, first pair less pigmented. First pair of elytra circular, without marginal papillae; surfaces with vesicular macrotubercles not sclerotized, ovoid, abundant, more prominent near margins (Figure 28A,B), microtubercles absent. Elytra of median and posterior segments without marginal papillae (Figure 28C,D); surfaces with micropapillae and vesicular macrotubercles. Macrotubercles less prominent and scattered (Figure 28E,F). Notopodia shorter than neuropodia. Neuropodia distally truncate, acicula tip protruding, prechaetal lobe with small rounded lobe near acicular tip. Dorsal cirri subdistally expanded and tapering to short tips. Cirrophores basally expanded. Nephridial papillae, present from segment 16. Ventral cirri tapering to filiform tip. Anus dorsal, pygidium with two anal cirri similar to dorsal cirri. Notochaetae shorter than the neurochaetae with rows of spines; the smaller curved, with blunt tips; the remaining ones slender tapering to capillary tips. Neurochaetae, with rows of spines on upper region, tips bidentate.

Remarks The features of the paratype (Figure 29A–G) are consistent with those of the holotype. The material examined was previously labelled as Halosydna sp. A sensu Hartman (1939a) but the elytra of Halosydna silvamariae sp. nov. lack marginal papillae and sclerotized microtubercles, and the vesicular macrotubercles are around the margin. Halosydna salazarvallejoi sp. nov. resembles H. silvamariae sp. nov. but differs in having a fringe of marginal papillae on the anterior elytra and soft macrotubercles less prominent and scattered on the elytral surface.

Type locality Scammon Lagoon, Baja California, Mexico.

Distribution Same as type locality. Halosydna riojaenriquei sp. nov. (Figures 30, 31)

Type material Holotype (ECOSUR 0134). Playa los pinos, Mazatlán, Mexico, in front of Facimar, depth 4 m, on rock oyster, coll. M. Tovar and P. Salazar, 27 February 2004. Paratypes:



Figure 28. Halosydna silvamariae sp. nov. Holotype LACM-AHF POLY 3615. (A, B) Elytra of first pair; (C) right elytron from middle segment; (D) right elytron from posterior segment; (E, F) same, macrotubercles.

Four specimens (ECOSUR 0135) of Playa los Pinos, Mazatlan, Mexico, station in front of Facimar, depth 4 m, on rock oyster, coll. M. Tovar and P. Salazar, 27 February 2004. One specimen (ECOSUR 0136), Playa los pinos, Mazatlán, Mexico, in front



Figure 29. Halosydna silvamariae sp. nov. Paratype LACM-AHF POLY 3616 (A) First elytron; (B) same, macrotubercles; (C) right elytron from posterior segment; (D) same, micropapilla; (E) notochaetae; (F) shorter notochaetae; (H) bidentate neurochaetae.

of Facimar, depth 4 m, on rock oyster, coll. M. Tovar and P. Salazar, 27 February 2004. One specimen (ECOSUR 0137), Playa los Pinos, Mazatlán, Mexico, in front of Facimar, depth 4 m, on rock oyster, coll. M. Tovar and P. Salazar, 27 February 2004.



Additional material Mazatlan, Mexico, Pacific Ocean: Two specimens (ECOSUR P02580), off Playa los Pinos, Mazatlan, station Cañon, coll. M. Tovar and P. Salazar, 22 February 2004. Two specimens (ECOSUR P02581), Playa los Pinos, Mazatlan, Station Cañon, coll. M. Tovar and P. Salazar, 22 February 2004. One specimen (ECOSUR P02582), Playa los Pinos, Mazatlán, near Marine Biology School, on rock oyster, coll. M. Tovar and P. Salazar, 23 February 2004. One specimen (ECOSUR P02583), Playa los Pinos, Mazatlan, near Marine Biology School, coll. M. Tovar and P. Salazar, 27 February 2004, Two specimens (ECOSUR P02584), playa los Pinos, Mazatlan, near Marine Biology School, depth 4 m, on rock oyster, coll. M. Tovar and P. Salazar, 27 February 2004. One specimen (ECOSUR P02585) of Playa los Pinos, Mazatlan, near Marine Biology School, depth 4 m, on rock oyster, coll. M. Tovar and P. Salazar, 27 February 2004. One specimen (ECOSUR P02586) of Playa los Pinos, Mazatlan, in front Marine Biology School, depth 4 m, on rock oyster, coll. M. Tovar and P. Salazar, 27 February 2004. Guerrero, Mexico Pacific Side. One specimen (ECOSUR P02587) of La Quebrada, Acapulco, on Spondylus calcifer, coll. M. Tovar and P. Salazar, 25 May 2000. One specimen (ECOSUR P02588) of La Quebrada, Acapulco, on Spondylus, 15 m, coll. A. Medina, 25 May 2000. One specimen (ECOSUR P02589) of La Quebrada, Acapulco, 25 May 2000, on Pinctada mazatlanica. coll. M. Medina. One specimen (ECOSUR P0 2590) of Los Cantiles, Acapulco, coll. A. Medina, on Spondylus, 26 May 2000. One specimen (ECOSUR P02591) of La Quebrada, Acapulco, on Muricanthus spondylus, 25 May 2000, coll. A Medina. One specimens (ECOSUR P02592) of La Condesa, Acapulco, coll. A. Medina, 27 November 1999, Three specimens (ECOSUR P02593) of La Quebrada, Acapulco, 6 m depth, coll. A. Medina, on Spondylu calcifer, 25 May 2000. One specimen (ECOSUR P02594) of La Quebrada, Acapulco, coll. A. Medina, 25 May 2000. One specimen (ECOSUR P02595) of La Roqueta, Acapulco, on sponge, coll. A. Medina, 26 May 2000. Five specimens (ECOSUR P02596) of Los Cantiles, la Quebrada, Acapulco, on Spondylus coll. A. Medina, 26 May 2000. One specimen (ECOSUR P02597) of Los Cantiles, la Quebrada, Acapulco, on Spondylus, coll. A. Medina, depth 8 m, 26 May 2000. One specimen (ECOSUR P02598) of Los Cantiles, la Quebrada, Acapulco, on Spondylus, coll. A. Medina, 26 May 2000. One specimen (ECOSUR P02599) of Los Cantiles, Acapulco, on Spondylus, depth 8 m, coll. A. Medina, 26 May 2000. One specimen (ECOSUR P02600) of La Quebrada, Acapulco, 6 m, coll. A. Medina, on Pinctada, 25 May 2000. Oaxaca, Mexico Pacific Side. One specimen (ECOSUR P02601) of Puerto de Abrigo, Huatulco, on Spondylus, 22 May 2000. One specimen (ECOSUR P02602) of Puerto de Abrigo, Huatulco, 22 May 2000. Four specimens (ECOSUR P02603) la Entrega, Huatulco, 3 m, on coral rock, 23 May 2000. Two specimens (ECOSUR P02604) of La Entrega, Huatulco, 3 m, on stromatolite, 23 May 2000. One specimen (ECOSUR P02605) of La Entrega, Huatulco, Transect 1, coll. P. Gomez 1991. One specimen (ECOSUR P0 2606) of Puerto de Abrigo, Huatulco, 22 May 2000.

Etymology This species is named for Enrique Rioja in recognition of his studies of polychaetes from Mexico.



Description Holotype with 36 segments, 1.05 cm long and 0.25 cm wide. Body pale yellow, dorsum with a band of green pigment on each segment. Brownish pigment on ceratophores and on antennae. Prostomium elongate with dots of pigment dorsally; facial tubercle long, rounded, with dark pigment; two pairs of dark eyes, anterior pair dorsolateral on widest part of prostomium, posterior pair near posterior margin; median antenna with ceratophore inserted frontally on prostomial lobes, style subdistally expanded with dark pigment, tip filiform, lateral antennae with ceratophores inserted terminally as prolongations of prostomium lobes, styles similar to median antenna. Palps robust, surfaces with papillae, distally tapering to filiform tips. Pharynx everted, with nine pairs of papillae, two pairs of jaws not fused to each other medially. Tentacular segment not visible dorsally. Tentaculophores with chaetae. Tentacular cirri similar to median antenna. Segment 2 projecting slightly over prostomium as small nuchal lobe, two small tubercles dorsally. Body with 18 pairs of elytra on segments 2, 4, 5, 7, 9, 11, then alternate to 21, 23, 25, 27, 28, 30, 31, 33. Three posteriormost segments with dorsal cirri. Elytra overlapping mid-dorsally, with spots of olive green towards the dorsal midline, without pigmentation over elytrophores. First pair of elytra circular, without marginal papillae (Figure 30A); surfaces smooth except for small micropapillae and sclerotized, conical microtubercles (Figure 30B), scattered, larger near elytrophore (Figure 30A). Second pair of elytra with granular surfaces (Figure 30C) due to presence of abundant microtubercles, conical-truncate, short (Figure 30D,E). Elytra in median and posterior segments (Figure 30F) without marginal papillae, surfaces with tiny micropapillae (Figure 30G) and microtubercles near margins. Notopodia shorter than neuropodia (Figure 30H). Neuropodia distally truncate, prechaetal lobe with small rounded lobe near acicula tip. Elytrophores wider than dorsal tubercles. Dorsal cirri subdistally expanded, pigmented, tapering to filiform tips. Cirrophores basally, expanded. Ventral cirri tapering to filiform tips. Nephridial papillae visible from segment 11, increasing in size toward posterior segments. Anus dorsal. Pygidium with anal cirri missing. Notochaetae shorter than neurochaetae, with rows of spines, the smaller curved, blunt tips, the remaining ones slender, tapering to long capillary tip. Neurochaetae with rows of spines on upper region, tip bidentate, subdistal tooth well developed. On first parapodia the neurochaetae have entire tips.

Remarks Most specimens of Halosydna riojaenriquei sp. nov. have olive-green elytra, but some have the pigment concentrated medially or covering the entire surface. Other specimens have yellow-brown pigment and all elytrophore scars are unpigmented. The prostomium can be unpigmented or with small black spots. The features of the paratypes (Figure 31A–D) are consistent with those of the holotype, the largest specimen is 2 cm long. Halosydna riojaenriquei sp. nov. resembles H. hartmanae (Kudenov, 1975) from Sonora, in lacking marginal papillae, but the former has anteriormost elytra with abundant microtubercles scattered over the entire surface and median and posterior



Figure 30. Halosydna riojaenriquei sp. nov. Holotype ECOSUR 0134. (A) First elytron; (B) same, microtubercles; (C) right elytron from second pair; (D) same, microtubercles of anterior border; (E) microtubercles; (F) left elytron from posterior segment; (G) same, micropapillae; (H) parapodium.



Figure 31. Halosydna riojaenriquei sp. nov. Paratype ECOSUR 0136 (A, B) Microtubercles; (C) notochaetae; (D) bidentate neurochaetae.

elytra with microtubercles around posterior borders, while H. hartmanae has median and posterior elytra with microtubercles forming patchs on the anterior half. Type locality Playa los Pinos, Mazatlan, Mexico. Distribution Mexico, Pacific Ocean: Mazatlan, Guerrero, Oaxaca. Key to species of Halosydna Kinberg, 1856, of eastern Pacific, Grand Caribbean and other worldwide localities (species marked with an asterisk) 1. Neurochaetae with tips entire (Figure 9E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Neurochaetae with tips bidentate (Figure 3C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2. Elytra thick, fleshy; elytra of anterior segments without macrotubercles; elytra of the mid-body and posterior segments with macrotubercles disc-shape short (Figure 10F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. latior Chambelin, 1919a Elytra fragile; elytra of anterior segments with macrotubercles; elytra of middle and posterior segments without macrotubercles . . . . . . . . . . . . . . . . . . . . 3

Journal of Natural History 1239 3. Macrotubercles of anterior elytra of two types, some blunt and others conical thick (Figure 21B) . . . . . . . . . . . . . . . . . . . . . . . H. tuberculifer Chamberlin, 1919a Macrotubercles of anterior elytra of one type, conical . . . . . . . . . . . . . . . . . . . . . 4 4. Macrotubercles of anterior elytra conical-slender, abundant, scattered on all elytral surface (Figure 14C) . . . . . . . . . . . . . . . . . . . . . . H. nebulosa (Grube, 176)∗ Macrotubercles of anterior elytra conical-thick, most prominent over the scar of the elytrophore (Figure 1D) . . . . . . . . . . . . . . . . . H. brevisetosa Kinberg, 1856 5. Elytra with fringe of marginal papillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Elytra without fringe of marginal papillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12


6. Elytra with vesicular tubercles, not sclerotized . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Elytra without vesicular tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7. Macrotubercles from first pair of elytra vesicular, not sclerotized, ovoid, prominent and long (Figure 27A); sclerotized tubercles absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. salazarvallejoi sp. nov. Macrotubercles from first pair of elytra vesicular, hemispherical, no prominent, short; tubercles sclerotized present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Fringe of marginal papillae on all elytra; first pair of elytra with macrotubercles long, sclerotized conical-truncated (Figure 11B); on posterior elytra vesicular macrotubercles scarce . . . . . . . . . . H. leius (Chamberlin, 1919a) Fringe of marginal papillae only on anterior elytra; first pair of elytra with vesicular macrotubercles long, not sclerotized (Figure 24D), on posterior elytra the vesicular macrotubercles are abundant . . . . . . . . . . . H. olgae sp. nov. 9. Elytra of anterior segments with tubercles sclerotized short and long . . . . . . 10 Elytra of anterior segments with tubercles sclerotized short . . . . . . . . . . . . . . . 11 10. Elytra of anterior segments with macrotubercles distally rounded, short (Figure 18B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. parva Kinberg, 1856 Elytra of anterior segments with macrotubercles conical, long (Figure 19B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. patagonica Kinberg, 1856∗ 11. Elytra of anterior segments with macrotubercles conical (Figure 7C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. johnsoni (Darboux, 1899) Elytra of anterior segments with microtubercles conical-truncated (Figure 23C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. virgini Virgini Kinberg, 1856∗ 12. Elytra with vesicular tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Elytra without vesicular tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13. First pair of elytra without microtubercles sclerotized; with vesicular macrotubercles ovoid (Figure 28A), abundant; elytra of middle and posterior segments with macrotubercles soft, ovoids long . . . . . . . H. silvamariae sp. nov. First pair of elytra with microtubercles sclerotized (Figure 13D), macrotubercles hemispheric, short, scarce; elytra of middle and posterior segments with vesicular microtubercles hemispherical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. leucohyba (Schmarda, 1861) 14. Elytra with macrotubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Elytra without macrotubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

1240 P. Salazar-Silva 15. Macrotubercles of elytra of anterior segments abundant, conical long and ovoid, surface wrinkled (Figure 4E) . . . . . . . . . . . . . . H. glabra Hartman, 1939a Macrotubercles on elytra of anterior segments scattered conical short blunt, surface smooth (Figure 2C) . . . . . . . . . . . . . . . H. fuscomarmorata (Grube, 1876) 16. Elytra of middle and posterior segments with microtubercles forming a patch on anterior part of the elytra (Figure 6B) . . . . . H. hartmanae (Kudenov, 1975) Elytra of middle and posterior segments with microtubercles over the half posterior elytral surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17 17. Microtubercles abundant, scattered over the half posterior elytral surface (Figure 16B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. nesiotes (Chamberlin, 1919b) Microtubercles scarce, around of posterior margin . . . H. riojaenriquei sp. nov.


Conclusion The genus Halosydna belongs to the group of polynoids with a constant number of segments and elytra, and a relatively short but not fragile body. Halosydna leucohyba is the only species occurring in the Grand Caribbean region but has also been reported for the Mexican Caribbean. The genus is clearly more diverse in the tropical eastern Pacific than in the Grand Caribbean. This pattern of diversity might be caused by a higher diversity of habitats in the tropical eastern Pacific, as reported for other groups (Muss et al. 2001). However, both regions require more taxonomic studies to increase records of Halosydna species in less-studied localities. Halosydna species are geographically distributed both in cold and warm waters. The species of the tropical eastern Pacific and Grand Caribbean species have neurochaetae with bidentate tips, and those of north and south Pacific coasts have neurochaetae with entire tips. This pattern might be related to the predominant modes of life as epibionts in the former and commensals in the latter. Martin and Britayev (1998) noted that members of commensal species tend to have less elaborate chaetae. Acknowledgements I thank Ardis B. Johnston (MCZ); Eric Lazo-Wasen (YPM); Mark Sidall (AMNH); Elin Sigvaldadottir, Karin Sindemark Kronestedt and Stefan Lundber (SMNH); Birger Neuhaus (ZMB); Angelika Brandt, Kathrin Philipps-Bassau, Petra Wagner (ZMH), Emilia GozalezVallejo (ECOSUR) for the loan of type specimens and other material. Special thanks go to Sergio I. Salazar-Vallejo (ECOSUR) for the opportunity to participate in the polychaete project; Guadalupe Nieto (ECOSUR-Tapachula) for the scanning electron micrographs; Nancy Voos (UMML) for arranging loans of type specimens; Leslie Harris (LACM-AHF) and Kristian Fauchald (USNM) for the attention received during my stay at the museums; Luis F. CarreraParra (ECOSUR), anonymous reviewers and Kirk Fitzhugh (LACM-AHF) made suggestions and improved the English. Part of the results of this study were supported by Semarnat-Conacyt 2002-C01-0182, other were supported by CONACYT (6169).

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