Sabellids (Polychaeta: Sabellidae) from the Grand Caribbean María Ana Tovar-Hernández* and Sergio I

Home , Bispira, Sabellastarte, Sabellidae

Zoological Studies 45(1): 24-66 (2006)

Sabellids (Polychaeta: Sabellidae) from the Grand Caribbean María Ana Tovar-Hernández* and Sergio I. Salazar-Vallejo Laboratorio de poliquetos, El Colegio de la Frontera Sur, Av. Centenario Km. 5.5, C. P. 77900, Chetumal, Quintana Roo, Méxi co

(Accepted September 3, 2005)

María Ana Tovar-Hernández and Sergio I. Salazar-Vallejo (2006) Sabellids (Polychaeta: Sabellidae) from the Grand Caribbean. Zoological Studies 45(1): 24-66. A taxonomic key for the 40 valid species of sabellids (Polychaeta: Sabellidae) occurring in the Grand Caribbean is provided. Eighteen species are herein recorded from the Grand Caribbean, and 4 species new to science are described: Anamobaea phyllisae sp. nov. (Guana Island), Bispira paraporifera sp. nov. (Mexican Caribbean), Megalomma perkinsi sp. nov. (Florida), and Pseudopotamilla fitzhughi sp. nov. (Mexican Caribbean). Identifications were corroborated by comparisons with type and non-type material loaned from several museums. An annotated checklist of the sabellid polychaetes from the Grand Caribbean is provided, including type localities, museums where material is deposited, and taxonomic remarks whenever necessary. The checklist comprises 56 species, of which 13 remain as questionable records, either because there is no type material, their records are isolated, or their type localities are far away from the Grand Caribbean. By presenting a complete overview of all records for sabellids in the area, this work summarizes our current knowledge of the diversity of this polychaete family in the Grand Caribbean, providing baseline data for future research. http://zoolstud.sinica.edu.tw/Journals/45.1/24.pdf

Key words: Fan worms, Key, Checklist, New species.

The Sabellidae Latreille 1825 is a family of b), (Hartman 1942 1951), (Rioja 1946), (Carpenter polychaete annelids (Polychaeta Grube 1850) 1956), (Renaud 1956), (Marsden 1960), (Jones commonly known as“fan worms”“, feather-duster 1962), and (Uebelacker and Johnson 1984), worms”, or“sea flowers”. They are easily recog- among others. However, none of these studies nized because living specimens have a colorful focused exclusively on sabellids. Salazar-Vallejo crown frequently projecting from the mouth of their (1996) listed 50 species of sabellids, belonging to tubes. According to Giangrande (in Rouse and 22 genera, occurring in the Grand Caribbean Pleijel 2001), the family consists of 490 species, of region. which 75 belong to the subfamily Fabriciinae (13 The state of knowledge of the species of genera), and over 400 correspond to the sabellids occurring in the Grand Caribbean, how- Sabellinae (30 genera) (Fitzhugh and Rouse ever, is still very poor and doubtful due to several 1999). reasons: for many years, identifications of The Grand Caribbean region comprises the Caribbean material were made using literature and Gulf of Mexico, Caribbean Sea, Bermuda, and the taxonomic keys from other regions of the world. northern littoral of Brazil (Salazar-Vallejo 2000). In Moreover, the identification of species rested the entire area, a number of important studies on mainly on drawings (which were seldom original), the polychaetes have been carried out: (Schmarda without comparison to the type material. Also, 1861), (McIntosh 1885), (Ehlers 1887), (Treadwell inadequate preservation of the material may 1901 1917 1921 1924a b 1928 1936 1939), sometimes have led to misinterpretations of body (Augener 1906 1922 1927), (Monro 1928 1933a structures or proportions. Finally, many of the

*To whom correspondence and reprint requests should be addressed. Tel: 52-983-8350440 ext. 4329. E-mail: [email protected]

24 Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 25 original descriptions used to identify local fauna or more times longer than the distance between are incomplete and not illustrated. All of the above the breast and crest. Further, uncini were separat- reasons led to misidentifications and records of ed into 2 different types with respect to the relative species from localities far away from the Grand development of the breast; thus, breasts were con- Caribbean. sidered well developed if they had a rounded pro- Thus, the main objectives of this research jected lobe, or breasts were reduced, if this lobe were to determine the composition of species of was missing, having a sligthly rounded lobe in lat- sabellids occurring in the Grand Caribbean region, eral view. describe 4 new species, and provide an updated The checklist for the sabellids from the Grand checklist and taxonomic key for the species occur- Caribbean region is arranged in alphabetic order, ring in the area. including information as follows: author(s) and date of publication, type locality (TL), abbreviations of museums where the type material is deposited; MATERIALS AND METHODS other letters in parentheses are HT for holotype, PT for paratype, ST for syntype, LT for lectotype, Type and non-type materials were loaned NT for neotype, SST for schizosyntype, and in from the following institutions: BMNH, ECOSUR, some cases, remarks. In this contribution, the FSBC, LACM-AHF, UMML, USNM, and ZMA (see species of Branchiomma and fabriciin sabellids are explanation of acronyms below). Samples from only included in the checklist. The revisions of the different localities in the Mexican Caribbean were tropical American species of Branchiomma and the collected at different depths from the intertidal Grand Caribbean species of Chone are going to zone to 10 m, by skin or scuba diving. Poly- be published elsewhere (Tovar-Hernández and chaetes were obtanined from dead coral blocks, Knight-Jones, in press; Tovar-Hernández, in algae, sponges, sea grasses, or sediments. press). Fabriciin sabellids are documented in a Worms were placed in plastic bags, transferred to series of contributions by Fitzhugh (1983 1990 fresh water for up to 15 min in the shade, causing 1996). an osmotic shock which enhanced the relaxation of organisms and avoided contraction at fixation; Abbreviations samples were then fixed with a 10% formaline solution, washed in the lab with tap water for 24 h, The following abbreviations are used in the and transferred to 70% ethanol for long-term text or in the checklist: AMNH, American Museum preservation. of Natural History, New York, NY, USA; BMNH, Specimens were generally examined with the The Natural History Museum, London, UK; ECO- aid of a stereomicroscope; characteristics of the SUR, El Colegio de la Frontera Sur, Chetumal, chaetae, eyes, internal structure of the dorsal lips, Mexico; FSBC, Florida Marine Research Institute, and number of thoracic segments were observed St. Petersburg, FL, USA; LACM-AHF, Los Angeles using a compound microscope. Every specimen County Museum of Natural History, Allan Hancock was analyzed to obtain the following measure- Foundation, Los Angeles, CA, USA; MCZ, ments: width of the posterior thorax, body length Museum of Comparative Zoology, Harvard Univ., (from the peristomium to the pygidium), length of Cambridge, MA, USA; NHMW, Naturhistorisches the radiolar crown, and number of thoracic seg- Museum Wien, Wien, Austria; NMI, National ments. Drawings were made with a camera luci- Museum of Ireland, Dublin, Ireland; NMW, National da, while some illustrations of diagnostic structures Museum and Galleries of Wales, Wales, UK; were obtained using scanning electron microscopy UMML, Marine Invertebrates Museum, Rosenstiel (SEM). School of Marine and Atmospheric Science, Univ. In order to standardize the relative size of the of Miami, Miami, FL, USA; USNM, The Natural manubria and facilitate comparison between History Museum, Smithsonian Institution, species, the following clasification was used: a) Washington DC, USA; UZIU, Museum of manubria were considered short if they were short- Evolution, Zoology Section, Uppsala Univ., er than the distance between the breast and crest; Uppsala, Sweden; ZMA, Zoological Museum, Univ. b) medium, if they were as long as the distance of Amsterdam, Amsterdam, The Netherlands; between the breast and crest; c) long, if they were ZMH, Zoologisches Institut und Museum, Univ. 1.5 times longer than the distance between the Hamburg, Hamburg, Germany; and ZMUC, breast and crest; and d) extra long, if they were 2 Zoologisk Museum, Univ. København, Copen- 26 Zoological Studies 45(1): 24-66 (2006) hagen, Denmark. - Radioles without stylods ...... Sabellastarte; Base of the crown involuted, forming spirals, thorax with 7 Key for sabellids from the Grand Caribbean or 8 chaetigers ...... S. magnifica 15. (14) Stylodes poorly developed (Fig. 18B); compound radiolar eyes present or absent ...... Pseudobranchiomma; 1. Breast of thoracic uncini narrow, poorly developed, giving Radioles with serrations, radiolar tips long, digitiform...... uncini an acicular appearance (Fig. 7L) ...... 2 ...... P. emersoni - Breast of thoracic uncini well developed, giving uncini a Z- - Stylodes well developed; compound radiolar eyes pre- shaped or avicular appearance (Fig. 1M) ...... 10 sent, set in pairs ...... Branchiomma 2. (1) Abdominal uncini with an elongate manubrium below 16. (11) Abdomen with paleate neurochaetae (Fig. 1N, O)...17 dentate region...... 3 - Abdomen without paleate neurochaetae ...... 19 - Abdominal uncini without or with reduced manubrium (Fig. 17. (16) Crown with dorsal and ventral basal flanges (Figs. 7K) ...... 7 1C, 3C) ...... Anamobaea 3. (2) Branchial crown with 2 pairs of radioles.....Manayunkia - Basal flanges absent ...... 18 - Branchial crown with 3 pairs of radioles ...... 4 18. (17) Chaetiger 1 with chaetae arranged in a row (Fig. 4. (3) Entire anterior peristomial ring collar membranous, or 12A) ...... Notaulax reduced to a low ridge ...... 5 - Chaetiger 1 with chaetae arranged in a bundle...... - Anterior peristomial ring collar as a low ridge dorsally and ...... Hypsicomus laterally, ventrally as a well-developed lobe ...... 6 19. (16) Inferior thoracic notochaetae broadly hooded (Fig. 5. (4) Anterior peristomial ring collar complete middorsally; 10J)...... 20 ventral filamentous appendages, if present, vascularized.. - Inferior thoracic notochaetae paleate ...... 21 ...... Pseudofabriciola 20. (19) Radioles with subdistal compound eyes (Fig. 9C, D).. - Entire anterior peristomial ring collar reduced to a low ...... Megalomma ridge, except for conical to elongate lobes on either side - Radioles without compound eyes ...... Demonax of dorsal midline; inferior thoracic neurochaetae in 21. (19) Radioles with compound eyes, except the dorsalmost chaetigers 3-8, pseudospatulate ...... Fabricinuda pair (Fig. 19A) ...... Pseudopotamilla; 6. (4) Ventral filamentous appendages branched; dorsal lips 3 or 4 compound eyes per radiole; dorsolateral collar mar- well developed, erect ...... Augeneriella; gins entire (Fig. 19C)....Pseudopotamilla fitzhughi sp. nov. Peristomial eyes rounded, well developed...... - Radioles without compound eyes ...... Perkinsiana; ...... A. hummelincki long ventral lappets (Fig. 17A); up to 20 thoracic - Ventral filamentous appendages unbranched; dorsal lips chaetigers ...... P. fonticula reduced to low ridges ...... Novafabricia Inferior thoracic pseudospatulate notochaetae limited to chaetigers 3-5; manubrium of abdominal uncini twice Amphicorina longer than dentate region ...... N. infratorquata 7. (2) Anterior peristomial ring with narrow, triangular, ventral 1. Posterior peristomial ring collar crenulate laterally, with lobe; abdominal uncini with equal-sized teeth above main 10-12 abdominal chaetigers ...... A. anneae fang...... Amphicorina - Posterior peristomial ring collar with smooth margin; with - Anterior peristomial ring without triangular ventral lobe; 5 abdominal chaetigers ...... A. androgyne abdominal uncini with unequal-sized teeth above main fang...... 8 Anamobaea 8. (7) Palmate membrane absent...... Jasmineira; Collar well developed; dorsal lobes triangular, ventral 1. Two dorsal kidney-shaped shields over anterior margin of lobes rounded...... J. bilobata base of crown; flanges smooth (without papillae) (Fig. 3C) - Palmate membrane present ...... 9 ...... Anamobaea phyllisae sp. nov. 9. (8) Last several abdominal chaetigers modified as a ven- - Dorsal kidney-shaped shields absent; flanges wrinkled tral anal depression ...... Euchone; (with papillae) (Fig. 1C) ...... Anamobaea orstedi Anal depression in last 3 chaetigers; collar ventrally entire; 8 abdominal chaetigers...... E. incolor - Anal depression absent ...... Chone Bispira 10. (1) Manubrium of thoracic uncini extra long....Potamethus; Collar ventral lobes very prolonged; thorax with 8 1. Radioles with eyes arranged in pairs ...... B. melanostigma chaetigers ...... P. spathiferus - Radioles without eyes...... 2 - Manubrium of thoracic uncini short, medium-sized, or 2. With dorsal spongy cushions (Fig. 6A); collar with short absent ...... 11 rounded ventral lobes (Fig. 6B) .... B. paraporifera sp. nov. 11. (10) Abdomen with neuropodial chaetae in a group ...... 12 - Without spongy cushions; collar with long triangular ven- - Abdomen with neuropodial chaetae in a row ...... 16 tral lobes (Fig. 4B)...... B. brunnea 12. (11) Companion chaetae present (Figs. 1K, L, 2C, 5G, 11E, F, 13G, 21E) ...... 13 Branchiomma - Companion chaetae absent ...... 14 13. (12) Abdominal neurochaetae arranged in an incomplete 1. Macrostylodes strap-like, tongue-like at midlength of radi- spiral or C-shaped pattern ...... Bispira ole; collar with rounded ventral lobes; uncini with 3 rows - Abdominal neurochaetae arranged in a tight spiral pattern of teeth above main fang ...... B. bairdi ...... Sabella - Microstylodes at midlength of radiole twice as long as the 14. (12) Radioles with stylods...... 15 others; uncini with 1 row of teeth above main fang...... Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 27

...... B. nigromaculatum for small part of ocular region (Fig. 12D); about 30 ocelli per group ...... N. bahamensis Chone - Collar segment wider than long, about as long as following 2 segments (Fig. 14B); radioles not flanged between palmate membrane and distal part of ocular region (Fig. 1. Pygidial cirrus present; crown with 10 pairs of radioles; 43 14C); 30~70 ocelli per group...... N. nudicollis C americana abdominal chaetigers ...... 3. Dorsal margin of collar with a deep incision (Fig. 16A); - Pygidial cirrus absent (Fig. 7A); crown with 5 pair of radi- short incision in ventral margin of collar (Fig. 16B); radi- Chone oles; 17 abdominal chaetigers...... sp. 1 oles with 4 or 5 ocelli (Fig. 16C), in small groups...... N. paucoculata Demonax - Dorsal margin of collar without deep incision; long incision in ventral margin of collar ...... 4 1. Ventral shield of collar segment longer than wider...... 4. Thorax with 13 chaetigers; radioles with many ocelli per ...... D. flecatus row (> 50) ...... N. circumspiciens - Ventral shield of collar segment at least twice as wide as - Thorax with 8 chaetigers; radioles with fewer ocelli per long ...... 2 row (up to 30) ...... 5 2. Radioles with numerous pairs of minute ocelli; branchial 5. Numerous ocelli (30 per row) ...... N. occidentalis crown as long as 1/4 of body; radioles with 4~6 dark - Few ocelli (7 per row) ...... N. midoculi bands ...... D. microphthalmus - Radioles without ocelli; branchial crown as long as 1/2 of Pseudofabriciola body ...... 3 3. Ventral shield of collar segment as long as wide; thoracic 1. Ventral filamentous appendages present, vascularized; D lacunosus tori not reaching ventral shields ...... dorsal lips well developed ...... P. quasiincisura - Ventral shield of collar segment twice wider than long; - Ventral filamentous appendages absent...... 2 thoracic tori indenting ventral shields (Fig. 8B)...... 2. Anterior margin of anterior peristomial ring with a large D jamaicensis ...... conical structure (dorsal to mouth, separated from collar); abdominal uncini with manubrium 1.5-times longer than Fabricinuda dentate region...... P. sofla - Anterior margin of anterior peristomial ring with no promi- 1. Branchial crown dorsally displaced; anterior peristomial nent structure; abdominal uncini with manubrium as long ring dorsally entire ...... F. trilobata as dentate region...... P. longa - Branchial crown not dorsally displaced; anterior peristomial ring ventrally U-shaped ...... F. pseudocollaris Anamobaea Krøyer, 1856

Manayunkia Anamobaea is known from a single Caribbean species, A. orstedi, for 150 years, which 1. Dorsalmost radiole with unpaired pinnules; ventral fila- is a very attractive species due to the colors of the mentous appendage wrinkled ...... M. speciosa - Dorsalmost radiole with paired pinnules; ventral filamen- branchial crown. In the phylogenetic analysis of tous appendage smooth ...... M. aestuarina Fitzhugh (1989), Anamobaea is defined by 1 rever- sal of the chaetal state: the posterior rows of the Megalomma anterior abdominal neurochaetae are modified, elongate, narrowly hooded. In this revision, we 1. Dorsal margins of collar fused to fecal groove; caruncle redescribe A. orstedi and describe a new species on anterior margin of shelf between dorsal lips, above from Guana I. (Anamobaea phyllisae). mouth (Fig. 10B) ...... M. lobiferum - Dorsal margins of collar not fused to fecal groove; caruncle absent ...... 2 Anamobaea orstedi Krøyer, 1856 2. Eyes present on most radioles ...... 3 (Figs. 1, 2) - Eyes present only on dorsalmost pair of radioles, large, with visible ommatidia ...... M. bioculatum Anamöbaea Orstedii Krøyer 1856: 32. 3. Eyes projected; segment 1 with quadrangular ventral Anamobaea orstedi.- San Martín et al. 1994: 556, fig. 1A-I.- shield (Fig. 9B)...... M. heterops Humann 1992: 135. - Eyes flat; segment 1 with rectangular ventral shield (wider than long) ...... M. perkinsi sp. nov. Material: [ECOSUR] Mexican Caribbean: I. Cozumel, SEDENA, Coll. A. Medina, 24 Mar. 2001 Notaulax (9). I. Contoy, Punta Sur, Coll. S. I. Salazar, 2 Mar. 2001 (5); Campamento de pescadores, Coll. M. A. 1. Ventral margin of collar entire ...... 2 Tovar, 1 Mar. 2001 (7). Cancún, Punta Nizuc, Coll. - Ventral margin of collar incised...... 3 2. Collar segment as long as wide, longer than 3 following S. I. Salazar and L. F. Carrera, 1 Apr. 1997 (1). segments (Fig. 12B); radioles completely flanged except Buenavista, E-24, Coll. S. I. Salazar and L. F. 28 Zoological Studies 45(1): 24-66 (2006)

Carrera, 27 Sept. 1996 (3). Xahuayxol, CR8, RC1, Vc 0556, airport (15). [LACM-AHF] LH 668, Sta. Coll. S. I. Salazar and L. F. Carrera, 2 June 1998 110, Coll. T. Zimmerman, 25 July 2000, between (2). Veracruz, I. Verde, Coll. S. I. Salazar, 15 June White Beach and Harris Ghut (1). [UMML] 1985 (3). [LACM-AHF] British Virgin Is.: Vc 0564, Jamaica: 22.754, Pedro Bank, S Dominican AF00-59, Guana, Beef I., Trellis Bay, Coll. G. Republic, R/V Pillsbury, Sta. 1254, 17 11'N, 78 Hendler, J. Martin, K. Fitzhugh, and R. Wear, 12 31'W, Cruise 7006, 14 July 1970, °20 m (1).° July 2000, tubes taken from coral rubble (1). [UMML] Lesser Antilles: 22.753, R/V Pillsbury, [LACM-AHF] BVI 2000, Sta. 59 (1). [LACM-AHF] 852, 11°53'N, 61°53'W, 3 July 1969, 13 m (3).

(A) (B) (C)

vbf

(D) (G) (H) (K) pi

(I) (J) (L)

(E)

rs m

(F) dra (M) (N) (O) dpa

dl ma

Fig. 1. Anamobaea orstedi. (A) Anterior end, dorsal view; (B) same, ventral view; (C) same, lateral view; (D) radiole, midlength; (E) radiolar skeleton in cross-section; (F) dorsal lip, lateral view; (G, H) short, superior thoracic spinelike notochaetae; (I, J) inferior thoracic paleate notochaetae, superior row; (K, L) companion chaetae; (M) thoracic uncinus, (N, O) abdominal paleate chaetae, posterior row. (A-O) ECOSUR, Isla Contoy, Mexico. Scale bars (A-C, F) 1 mm; (D) 0.1 mm; (E) 0.06 mm; (G-O) 0.05 mm. dl, dorsal lip; dpa, dorsal pinnular appendage; dra, dorsal radiolar appendage; ma, manubrium; m, mucro; pa, papillae; pi, pinnule; rs, radiolar skeleton; se, simple eyespot; vbf, ventral basal flange; vl, ventral lip. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 29

[UMML] Puerto Rico: 22. 763, 1423, 21 41'N, 71 thoracic notochaetae paleate, symmetrical (Figs. 23'W, 19 July 1971, 18 m (3). ° ° 1I, J, 2A), arranged in 2 transverse rows of 5 or 6 Description: Body 11-124 mm long. Crown chaetae each. Superior thoracic notochaetae long (19-23 mm), rigid with red and white bands. spinelike, short, 5 or 6 per group (Figs. 1G, H, 2A). Radioles 19-27 pairs. Base of crown red (in live Thoracic tori not reaching ventral shields, long for material), 2-2.5 mm long. Two pairs of erect and 20 anteriormost chaetigers, then gradually rigid flanges on basal lamina, 1 pair dorsal, the decreasing in length. Thoracic uncini avicular, other ventral, terminating on posterior peristomial with 22-26 rows of equal-sized teeth above main ring, with rounded papillae on surface (Fig. 1A-C). fang in frontal view (seen under SEM), covering Palmate membrane present above basal lamina 1/2 of extension of main fang (Fig. 2B), breast well (1.5-times length of basal lamina). Simple black developed, manubrium medium (Fig. 1M). radiolar eyespots in a small group above palmate Companion chaetae with roughly symmetrical tips, membrane, in a narrow band slightly above radio- as teardrop-shaped membranes (Figs. 1K, L, 2C). lar midlength (Fig. 1D). Radiolar skeleton with 6 Abdominal chaetigers 56-150. Abdominal neu- or 7 cells in cross-section (Fig. 1E). Dorsal lips rochaetal fascicles in 2 transverse rows; anterior short, with dorsal pinnular appendage (Fig. 1F). and posterior rows on all chaetigers with asym- Ventral lips kidney-shaped (Fig. 1B), parallel metrical paleate chaetae, mucronate with small lamellae present. Dorsal margin of collar slightly teeth at base of mucro (Figs. 1N, O, 2D), and indented by fecal groove. Ventral lappets triangu- modified, elongate, narrowly hooded chaetae. lar, not overlapping (Fig. 1B). Ventral shield of Abdominal uncini avicular, with 30-37 rows of segment 1 rectangular, longer than following equal-sized teeth above main fang in frontal view shields (Fig. 1B). Thorax 13-23 mm long, 1.5-4 (seen under SEM), covering 3/4 of extension of mm wide with many chaetigers (32-55). Inferior main fang, breast well developed, manubrium

(A) (B)

66.6 µm 7.69 µm

10.0 µm 50.0 µm

Fig. 2. Anamobaea orstedi. (A) Thoracic chaetigers; (B) thoracic uncini; (C) companion chaetae; (D) abdominal paleate chaetae. (A- D) ECOSUR, Isla Cozumel, Mexico. 30 Zoological Studies 45(1): 24-66 (2006) long. Tube distally thinner, semi-transparent, cov- Material: Holotype [LACM-AHF POLY 2148] ered by fine sand and algae; thicker at midlength, British Virgin Is.: Off Guana Head, ARM, Coll. G. leathery, with incrusting calcareous algae basally. Hendler and R. Wear, 10 July 2000, 21.6 m, sand bottom, fine sand mixed with silt. [LACM-AHF] Anamobaea phyllisae sp. nov. AF0056, VC0519, White Bay, 18.2 m directly off (Fig. 3) beach house, Coll. K. Fitzhugh, 11 July 2000, 2 m, crevice on side of large brain coral with sabellid

(A) (B) (C)

vbf

(G) (H) (I)

(D) (F)

(P) se (N) (K) (J) m (O)

(E) ps

(L) (M) rs

Fig. 3. Anamobaea phyllisae sp. nov. (A) Anterior end, dorsal view; (B) same, ventral view; (C) same, lateral view; (D) radiole, midlength; (E) short, superior thoracic spinelike notochaeta; (F, G) inferior thoracic paleate notochaeta; superior row; (H) thoracic uncinus; (I) radiolar skeleton in cross-section; (J-M) companion chaetae; (N) abdominal paleate chaetae, posterior row; (O) modified, elongate, narrowly hooded chaeta; (P) abdominal uncinus, oblique view. (A-P) LACM-AHF POLY 2148, Guana Island, holotype. Scale bars (A-C) 1 mm; (D, I) 0.12 mm; (E-H, J-P) 0.05 mm. fg, fecal groove; m, mucro; pi, pinnules; ps, pinnular skeleton; rs, radiolar skeleton; se, simple eyespots; ss, sheath tissue; vbf, ventral basal flange. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 31 tubes extending out (1). Remarks: Anamobaea phyllisae sp. nov. is Etymology : This species is named in honor of unique by the presence of smooth basal flanges, Dr. Phyllis Knight-Jones (Univ. of Swansea, UK) in without rounded papillae and 2 dorsal kidney- recognition of her many years of dedication to the shaped shields located on the anterior margin of taxonomy of sabellid polychaetes. the dorsal pair of the basal flanges, and a dorsal Description: Holotype complete, 37 mm long. collar margin divided by a deep fecal groove. In Crown long (21 mm), rigid, with 26 pairs of radi- the Caribbean, there is another undescribed oles. Basal flanges and basal 1/3 of radioles pale species of Anamobaea (Anamobaea sp. 1: brown, next 1/3 pale, distal 1/3 yellowish-brown (in [LACM-AHF] BVI 99, Sta. 123, LH 1005, Guana: preserved material). Two pairs of erect, rigid, and Bigelow Beach, Coll. T. Zimmerman, 15 Aug. smooth flanges (not papillated) on basal lamina, 1 1999, 9-11 m (14). AF0042, VC0489, Off Guana pair dorsal, the other ventral, terminating on poste- Head, Coll. G. Hendler and R. Wear, 10 July 2000, rior peristomial ring (Fig. 3C). Two dorsal kidney- 21.6 m, sand bottom, fine sand mixed with silt). It shaped shields on anterior margin of dorsal pair of differs from A. phyllisae by having a different radio- basal flanges (Fig. 3A, C). Palmate membrane lar color pattern in preserved material, as follows: present above basal lamina (1.5 times length of white-purple-transparent-orange-transparent with basal lamina). Simple radiolar eyespots, orange, the transparent flanges, and for lacking the kidney- in a small group above palmate membrane (Fig. shaped shields (see Humann 1992: 131). 3D), in a narrow band slightly above radiolar Anamobaea orstedi and Anamobaea sp. 1 have midlength. Radiolar skeleton with 8 cells in cross- different radiolar color patterns in live specimens section (Fig. 3I). Dorsal lips short, with dorsal pin- (red-white in A. oerstedi and white-purple-transpar- nular appendages. Ventral lips kidney-shaped, ent-orange-transparent in Anamobaea sp. 1), but with parallel lamellae. Collar with dorsal margin both have bright white patches dorsally and ven- indented by fecal groove (Fig. 3A). Ventral lappets trally on the basal flanges. rounded, overlapping, (Fig. 3B), brownish-red. Ventral shield of segment 1 divided into 2 triangles, Bispira Krøyer, 1856 longer than the following shields. Thorax 20 mm long, 5 mm wide, with 74 chaetigers. Inferior tho- Krøyer (1956) defined Bispira without includ- racic notochaetae paleate, symmetrical (Fig. 3F, ing any species. Knight-Jones and Perkins (1998) G), arranged in 2 transverse rows of 5 or 6 designated Amphitrite volutacornis Montagu, 1804, chaetae each. Superior thoracic notochaetae as the type species. Bispira is recognized by the spinelike, short, 5 or 6 per group (Fig. 3E). independent adquisition of radiolar flanges and Thoracic tori long, diminishing in length towards dorsal pinnular appendages (Fitzhugh 1989). posterior end, not indenting ventral shields. Knight-Jones and Perkins (1998) recognized 19 Thoracic uncini avicular, with 18 equal-sized teeth valid species; only B. brunnea (Treadwell, 1917) above main fang in lateral view, covering 1/2 of and B. melanostigma (Schmarda, 1861) are found extension of main fang (seen under light in the Grand Caribbean. microscopy), breast well developed, manubrium medium (Fig. 3H). Companion chaetae with Bispira brunnea (Treadwell, 1917) roughly symmetrical tips, as teardrop-shaped (Fig. 4) membranes (Fig. 3J-M). Abdominal chaetigers 70. Abdominal neurochaetal fascicles in 2 transverse Metalonome (sic) brunnea Treadwell 1917: 268, pl. 3, figs. 24- rows; anterior and posterior rows on all chaetigers 27. Sabella bahamensis Augener 1922: 48 fide Knight-Jones and with asymmetric paleate chaetae, mucroate, with Perkins 1998. small teeth at base of mucro (Fig. 3N) and modi- Bispira brunnea.- Knight-Jones and Perkins 1998: 433, figs. 19, fied, elongate, narrowly hooded chaetae (Fig. 3O). 20. Abdominal uncini avicular, with 27 equal-sized teeth above main fang in lateral view, covering 3/4 Material: [ECOSUR] Mexican Caribbean: of extension of main fang (seen under light Punta Allen, Coll. P. Gómez, 23 Apr. 1992 (34). microscopy), breast well developed (Fig. 3P), Puerto Morelos, Coll. M. S. Jimenez, 7 Sept. 1986 manubrium long. Tube distally thinner, semi-trans- (22). Majahual, Coll. J. R. Bastida and P. Salazar, parent, covered by fine sand and algae; thicker at 21 Mar. 2000 (73). Buenavista, Coll. S. I. Salazar midlength, leathery, with incrusted calcareous and L. F. Carrera, 27 Sept. 1996 (52). I. Cozumel, algae; very hard basally. Chankanaab, Coll. S. I. Salazar, 2 Apr. 1992 (78); 32 Zoological Studies 45(1): 24-66 (2006)

SEDENA, Coll. M. A. Tovar, 5 Mar. 2001 (89). Blanche Bay, Coll. P. Wagenaar Hummelinck, Sta. Laguna de Términos, July 1984, 30.27a (3). 1125A, 26 Apr. 1949, conglomeratic rocks with bal- [LACM-AHF] British Virgin Is.: Vc 0627, Sta. 59 anids, some sand, 0.5-1.5 m (88). Guana, Beef I., Trellis Bay, Coll. G. Hendler, J. Description: Gregarious species. Body long Martin, K. Fitzhugh, and R. Wear, 12 July 2000 (5.5-36 mm). Crown brown at base with white (30). [LACM-AHF] Grand Ghut, Coll. G. Hendler, 6 radioles (in preserved materials). Crown longer July 2000, algal clumps near Graham Forrester than body, 6-38 mm (Fig. 4A), radioles arranged in (15). [LACM-AHF] AF00-16A, Vc 0236, Grand 2 semicircles with slightly involuted ventral edges, Ghut, near Graham Forrester's transect, reef slope with 13 or 14 pairs of radioles. Radioles with bare beyond spur and groove zone, algal clumps, 3-4 tips as long as equivalent space of 16 pinnules cm high, mainly Laurencia and Dictyota, 19.8 m, 7 (Fig. 4C), without eyes. Radiolar skeleton with 8 June 2000. [LACM-AHF] Vc 0237. [LACM-AHF] cells in cross-section disposed in a reniform Vc 0238. [ZMA] St. Martin: V. Pol. Great Bay, Pt. arrangement at base of radioles; narrow flanges

(A) (B) (C)

(F)

(D) (D)

Fig. 4. Bispira brunnea. (A) Whole worm, dorsal view; (B) anterior end, ventral view; (C) radiolar tip; (D) thoracic uncinus, oblique view; (E) abdominal uncinus, non-lateral oblique; (F) inferior thoracic spinelike notochaeta. (A-F) ECOSUR, Punta Allen, Mexico. Scale bars (A) 2 mm; (B) 1 mm; (C) 0.2 mm; (D, E) 0.01 mm; (F) 0.02 mm. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 33 along edges of outer surface. Dorsal lips with radi- Sabella bipunctata Baird 1865: 158-159 fide Knight-Jones and olar appendages extending beyond palmate mem- Perkins 1998: 415. brane; pinnular appendages present, surrounding sheath tissue absent, radiolar appendage skeleton Material: [BMNH] St. Vincent: [1839.12.27./27]. composed of a single row of cells, and connective 16-20 m, Rev. Landstone Guilding Coll. (3 speci- tissue at dorsal lip lamellae. Ventral lips and paral- mens labeled as syntypes of Dasychone nigro- lel lamellae present. Collar with wide dorsal gap, maculata, correspond to Bispira melanostigma). lateral margins extending well beyond origin of [ECOSUR] Mexican Caribbean: Akumal, 12 Apr. crown. Long, triangular ventral lappets (Fig. 4B), 1986 (2). Banco Chinchorro, 28 July 1990, 2.5 m with small lobular extensions overlapping at mid- (1). Canal Zaragoza, Coll. J. Espinosa, 9 Oct. line. Anterior margin of shield of segment 1 W- 1997, in mangrove roots (1). I. Cozumel, shaped, following thoracic shields rectangular with Chankanaab, 2 Apr. 1992 (3). Desembocadura de wide gap to adjacent tori (Fig. 4B). Spinelike Bacalar Chico, Coll. J. Espinosa, 9 Oct. 1997, in chaetae on segment 1. Thoracic and abdominal Rhizophora mangle (15). DIF Puerto Aventuras, inter-ramal eyespots present. Thorax 2-6.5 mm Coll. S. I. Salazar, 21 Mar. 1992 (5). Hualalpich, long, 1.5-2 mm wide, with 10 or 11 chaetigers. 17 Apr. 1986 (1), 19 Apr. 1986 (5). I. Contoy, Superior and inferior thoracic notochaetae spine- Punta Sur, Coll. M. A. Tovar and S. Frontana, 28 like (Fig. 4F) arranged in bundles, with irregular, Feb. 2001 (3). Majahual Norte, Coll. M. A. Tovar, longitudinal rows of chaetae. Thoracic uncini avic- 19 Jan. 2001 (21); muelle, Coll. H. ten Hove, 18 ular, with 4 or 5 equal-sized teeth above main fang Mar. 2001 (various dozens). Cancún, Laguna in lateral view, covering 1/2 of extension of main Nichupté, 20 Apr. 1988, E7M1+1 (2); Punta Nizuc, fang (seen under light microscopy), breast well Coll. S. I. Salazar and L. F. Carrera, 1 Sept. 1997 developed (Fig. 4D), manubrium medium. (3). Punta Gavilán, 9 Apr. 1992, Coll. S. I. Salazar Companion chaetae with distinctly asymmetrical (5); Coll. M. Londoño, 10 Feb. 2001, 4 m (7). membranes. Abdomen with 22-84 chaetigers. Punta Río Indio, Coll. M. A Tovar, 17 Mar. 2001 Abdominal neuropodia as conical lobes. (3). Río Lagartos, Coll. J. R. Bastida and S. I. Abdominal neurochaetal fascicles partially spiraled Salazar, 18 Feb. 1999 (1). Santa Cecilia, bolsa or C-shaped. Anterior abdominal neurochaetae 101, 4 Nov. 1990 (1). Xahuayxol, Coll. S. I. spinelike on anterior and posterior rows; posterior Salazar and L. F. Carrera, 27 Sept. 1996 (1). abdominal neurochaetae spinelike on anterior Xamach, Coll. E. Donath, 28 Feb. 1986 (2). rows, and modified, elongate, narrowly hooded on Xcacel, Coll. S. I. Salazar, 3 Apr. 1992 (3). Xcalay, posterior rows. Abdominal uncini avicular, with 5 B 115, 4 Nov. 1990 (10). Yalahau, 8 Sept. 1993 or 6 equal-sized teeth above main fang in lateral (1). Panama: Fuerte Sherman, Colón, Coll. S. I. view, covering 1/2 of extension of main fang (seen Salazar, 2 June 2002 (3). [LACM-AHF] British under light microscopy), breast well developed, Virgin Is.: BVI-99-12F, LH 0143, Guana, North Bay, manubrium short (Fig. 4E). Tube built with fine Coll. L. Harris, 25 July 1999, west rocky intertidal, white sand. clumps of zoanthids Zoanthus pulchellus (3). Remarks: Bispira brunnea is similar to B. [LACM-AHF] BVI-99-25F, LH 142, Tortola, Toll porifera (Grube, 1878), as both species lack radio- bridge, Coll. L. Harris, 29 July 1999, mangroves lar eyes; however, the collar segment in that (6). [LACM-AHF] BVI-99-53, LH 0361, Pelican species is very elongate and has a dorsal spongy Ghut, Coll. T. Zimmerman, 3 Aug. 1999 (1). structure, both characters of which are absent from [LACM-AHF] BVI-99-81, LH 0754, Tortola, Coll. L. B. brunnea. This is a gregarious species, very Harris, 7 Aug. 1999 (8). [LACM-AHF] AF00-04, Vc common in the Caribbean, easily recognized alive 0050, White Bay, Coll. K. Fitzhugh, 4 July 2000, by the white tubes and bicolored crown. 0.5 m, tubes attached to coral boulder (1). [LACM- AHF] AF00-04, Vc 0051 (1). [LACM-AHF] AF00- Bispira melanostigma (Schmarda, 1861) 04, Vc 0052F (6). [LACM-AHF] AF00-04, Vc 0054 (Fig. 5) (1). [LACM-AHF] AF00-54, Vc 0555, Muskmelon Bay, Long Point, near ARM site, Coll. G. Hendler Sabella melanostigma Schmarda 1861: 36, pl. 22, fig. 190.- and R. Wear, 11 July 2000, 9.1-15.2 m, gorgonians Uebelacker 1984: 54-42, 54-43. and sponges, sediment extremely silty, covered by Sabella variegata Krøyer 1856: 29-30 fide Knight-Jones and red, green, and brown algae (2). [LACM-AHF] Perkins 1998: 415. AF00-57, Vc 0480, White Bay, Coll. K. Fitzhugh, Sabella thoracica Krøyer 1856: 31-32 fide Knight-Jones and Perkins 1998: 415. 11 July 2000, 2 m, coral rock (1). [LACM-AHF] 34 Zoological Studies 45(1): 24-66 (2006)

AF00-59, Beef I., Trellis Bay, Coll. G. Hendler, J. Wagenaar Hummelinck, Sta. 1540B, 26 July 1967, Martin, K. Fitzhugh, and R. Wear, 12 July 2000 eroded Thalassia flat, 0.5-1 m (7). [ZMA] St. (15). [LACM-AHF] AF00-59, Vc 0628 (1). [LACM- Martin: V. Pol. Great Bay, Pt. Blanche Bay, Coll. P. AHF] AF00-60, Vc 0673, Muskmelon Bay off Crab Wagenaar Hummelinck, Sta. 1125A, 26 June Cove, Coll. G. Hendler, T. Zimmerman, J. Martin, 1949, rocks, sand, low tide and lower zone (20). and R. Wear, 13 July 2000, 21.3 m (1). [LACM- Description: Medium-sized specimens, 21-68 AHF] Bahamas: LH03-29, Sta. 15; LH03-30, Sta. mm long. Thorax inflated (Fig. 5B), with anterior 15; LH03-31, Sta. 15; LH03-34, Sta. 010 (5); purple spots. Crown purple (in live and preserved LH03-72, Sta. 21; LH03-203, Sta. 045 (2); LH03- material), 13-30 mm long, with 16-21 pairs of radi- 204, Sta. 045 (1); LH03-215, Sta. 55; LH03-225, oles in semicircular arrangement. Radioles with 2 Sta. 55. [UMML] Mexican Caribbean: 22. 764, or 3 pairs of dark-brown compound eyes (Fig. 5C). Majahual, muelle, Coll. H. ten Hove, 18 Mar. 2001 Radiolar skeleton with 8-10 cells in cross-section, (20); 22.765, Xcalay, B115, Nov. 4, 1990 (6). obtuse epithelial corners on outer surface (Fig. [ZMA] Jamaica: V. Pol. Kingston Harbour, Small 5D). Dorsal lips with radiolar appendages extend- Boat Channel, E of Port Royal, Coll. P. Wagenaar ing beyond palmate membrane; pinnular Hummelinck, Sta. 1678, 7 May 1973, Rhizophora appendages present. Ventral lips and parallel timber, 0-1 m (50). V. Pol. flood gate of The lamellae present. Dorsal margins of collar promi- Falsees Great Salt pond, Coll. P. Wagenaar nent and separated by fairly wide gap, lateral mar- Hummelinck, Sta. 024, 8 May 1975 (10). [ZMA] gins just covering origin of crown (Fig. 5A). Ventral Antigua: V. Pol. Dickinson Bay N, Coll. P. lappets prominent and medially involuted, forming

(A) (B)

re ss

Fig. 5. Bispira melanostigma. (A) Anterior end, dorsal view; (B) same, ventral view; (C) radiolar tip; (D) radiolar skeleton in cross-section; (E) thoracic uncinus; (F) abdominal uncinus; (G) companion chaeta. (A-G) ECOSUR, Majahual, Mexico. Scale bars (A, B) 0.5 mm; (C) 0.2 mm; (D) 0.12 mm; (E-G) 0.02 mm. ce, compound eye; rs, radiolar skeleton; ss, sheath tissue. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 35 small pockets flanking midline cleft (Fig. 5B). larity between this species and B. porifera. Ventral thoracic shields trapezoidal on segment 1. Description: Holotype complete, 16 mm long. Spinelike chaetae on segment 1. Purple patches Crown pale (in preserved material), 8 mm long on ventral sacs and dorsally on thorax, dark patch- with unflanged bases and 14 pairs of radioles, es close to dorsal sides of all parapodia, and small arranged in 2 semicircles with slightly involuted marks at ventral end of each thoracic torus and ventral edges. Radioles with 4 pairs of eyes, long ventral to each abdominal fascicle. Thorax with 8- bare tips, as long as equivalent space of 8 pin- 14 chaetigers, 4.5-9 mm long, 4.5-7 mm wide. nules, radiolar flanges present in area between Thoracic ventral shields trapezoidal, not indented eyes and radiolar tip. Radiolar skeleton with 8 by tori. Inferior thoracic notochaetal arranged in cells in cross-section. Dorsal lips purple, with radi- bundles, with irregular, longitudinal rows of spine- olar appendages extending beyond palmate mem- like chaetae. Thoracic uncini avicular, with 3 or 4 brane; pinnular appendages present. Ventral lips equal-sized teeth above main fang in lateral view, and parallel lamellae present. Collar dorsal mar- covering 1/2 of extension of main faing (seen gins with orange spongy, cushion-like mass under light microscopy), breast well developed, extending to 6th thoracic chaetiger, superior border manubrium medium (Fig. 5E). Companion heart-shaped, inferior border V-shaped (Fig. 6A); chaetae with distinctly symmetrical distal mem- ventral lappets small, rounded with purple spots, branes (Fig. 5G). Abdomen with 64-72 chaetigers. overlapping (Fig. 6B). Thorax 5 mm long, 3 mm Abdominal neurochaetal fascicles in partially spi- wide with 8 chaetigers. Ventral shield of segment raled or C-shaped arrangement formed by posteri- 1 W-shaped (Fig. 6B); following shields rectangu- or chaetal row, posterior abdominal neurochaeta lar, not indented by tori. Thoracic and abdominal spinelike on anterior rows, and modified, elongate, inter-ramal eyespots present (Fig. 6B). Inferior narrowly hooded on posterior rows. Abdominal thoracic notochaetal fascicles arranged in bundles, uncini avicular, with 3 or 4 equal-sized teeth above with irregular, longitudinal rows of spinelike main fang in lateral view, covering 1/2 of extension chaetae (Fig. 6F, G). Thoracic uncini avicular, with of main faing (seen under light microscopy), breast 5 equal-sized teeth in lateral view, covering 1/2 of well developed, manubrium short (Fig. 5F). Tubes extension of main fang (seen under light made of fine white sand. microscopy), breast well developed, manubrium Remarks: Bispira melanostigma is a common medium (Fig. 6C). Companion chaetae with dis- species in the Caribbean, being very attractive tinctly asymmetrical distal membranes (Fig. 6E). because of the purple color of the branchial crown. Abdomen with 38 chaetigers. Abdominal neuropo- Knight-Jones and Perkins (1998) considered dia as conical lobes, in partially spiraled or C- Sabella thoracica Krøyer, 1856, B. variegata shaped arrangement. Abdominal neurochaetae (Krøyer, 1856), and Bispira melanostigma spinelike on anterior and posterior rows (Fig. 6H), (Schmarda, 1861) as synonyms. Although S. tho- and modified, elongate, narrowly hooded on poste- racica and B. variegata have priority over B. rior rows (Fig. 6I, J). Abdominal uncini avicular, melanostigma, those names have not been used with 3 equal-sized teeth in lateral view, covering since 1856, except B. variegata by Perkins (1984). 1/2 of extension of main fang (seen under light Conversely the name melanostigma has been in microscopy), breast well developed, manubrium frequent use since its inception and is the name short (Fig. 6D). Gametes visible on 4th abdominal used by the curators of most museums. chaetiger. Therefore, to conserve the name melanostigma, Remarks: The spongy cushion of Bispira the precedence of S. thoracica and B. variegata paraporifera sp. nov. is similar to that of B. porifera should be waived (Art. 23, Sect. 9, International (Grube, 1878), described from the Philippines. Code of Zoological Nomenclature, 2000). However, B. paraporifera has the spongy cushion- like mass restricted to the dorsal side of the tho- Bispira paraporifera sp. nov. racic segments, its inferior border is V-shaped, and (Fig. 6) it extends to the 6th thoracic chaetiger, while in B. porifera, the spongy cushion-like mass extends Material: Holotype [LACM-AHF POLY 2149] around the peristomium to the ventral sacs into Mexican Caribbean: Holotype, DIF Puerto smaller cushions; its inferior border is rounded and Aventuras, Coll. S. I. Salazar, 22 Mar. 1992, on less developed, and it extends to the 8th thoracic dead coral. chaetiger. Further, B. porifera has radioles without Etymology: Specific name refers to the simi- eyes, with incipient paired flanges along the outer 36 Zoological Studies 45(1): 24-66 (2006) surface becoming more distinct distally, radiolar embryos. tips very short, and ventral lappets prominent (see Knight-Jones and Perkins 1998, fig. 16A-P). Chone Krøyer, 1856 Willey (1905) suggested that the spongy cushions of B. porifera are glandular, while Knight-Jones Banse (1972) emended Chone, stressing the and Perkins (1998) suggested that variability in girdle of glands on the 2nd chaetiger, the ventral texture and degree of coverage in spongy cush- shields, the“bayonet-like chaetae”, and the ions in B. porifera may be due to a developmental “subespatulate chaetae”. He included Mega- series and may be involved in the incubation of chone Johnson, 1901, redescribed or improved the

(A) (B) sc

ire

(D)

(E)

Fig. 6. Bispira paraporifera sp. nov. (A) Anterior end, dorsal view; (B) same, ventral view; (C) thoracic uncinus, non-lateral oblique; (D) abdominal uncinus; (E) companion chaetae; (F, G) inferior thoracic spinelike notochaetae; (H) abdominal spinelike neurochaetae; (I, J) abdominal modified, elongate, narrowly hooded chaetae. (A-J) LACM-AHF POLY 2149, Puerto Aventuras, Mexico, holotype. Scale bars (A, B) 1 mm, (C-E) 0.02 mm, (F-J) 0.04 mm. ire, interamal eyespot; sc, spongy cushions. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 37 knowledge of some species, described 3 new ly. In addition, Fitzhugh (1989) suggested that species, and recognized that the taxonomy of this Chone possibly be divided into several mono- genus was confusing because some species pre- phyletic genera based on the retention or loss of sent variations in the morphology of the abdominal dorsal radiolar appendages and the morphology of uncini. the abdominal uncini. Fitzhugh (1989) placed Chone within the subfamily Sabellinae, giving emphasis to the presence Chone sp. 1 of the branchial skeleton and to the origin of the (Fig. 7) collar. He pointed out that Chone is not defined by any synapomorphy; also, he emended the genus Material: [ECOSUR] Panama: Club de Yates, because some specimens previously identified as Colón, Coll. S. I. Salazar, 1 June 2002 (1). [ZMA] C. infundibuliformis have dorsal lips with dorsal Grenada: V. Pol. Hog I. near Point Salines, radiolar appendages, whereas other specimens do Thalassia in muddy sand near Rhizophora, 0.5-1 not have such structures. Fitzhugh (1989) named m, 8 July 1967, Coll. P. Wagenaar Hummelinck, those specimens Chone1 and Chone2, respective- Sta. 1551A (1).

(A) (B) (C)

(J)

(D) (E) (F) (G) (H) (I)

(L)

(K)

Fig. 7. Chone sp. 1. (A) Whole worm, lateral view; (B) radiolar tip; (C) anterior end, ventral view; (D-G) thoracic paleate chaeta; (H, I) thoracic elongate narrowly hooded chaetae; (J) thoracic bayonet chaeta; (K) abdominal uncinus; (L) thoracic uncinus. (A-L) ECOSUR, Colon, Panama. Scale bars (A, C) 1 mm; (B) 0.1 mm; (D-L) 0.01 mm. 38 Zoological Studies 45(1): 24-66 (2006)

Description: Specimen from Panama com- Jones (1983), Knight-Jones and Walker (1985), plete, small, cylindrical body 5.5 mm long (Fig. Perkins (1984), and Fitzhugh (1989). The primary 7A). Crown 1.5 mm long, palmate membrane character distinguishing Demonax is that compan- along 3/4 of radiolar length, 5 pairs of radioles with ion chaetae are distally dentate (Fitzhugh 1989). broad radiolar flanges (Fig. 7C) and short bare Only 3 species of Demonax have been described tips, as long as equivalent space of 6 pinnules from the Grand Caribbean: D. flecatus (Hoagland, (Fig. 7B). Pinnules of same length along radioles. 1919) from Puerto Rico, D. jamaicencis (Augener, Dorsal margins of collar fused to fecal groove; lat- 1922) from Jamaica, and D. lacunosus Perkins, eral and ventral margins of collar entire. Radiolar 1984 from Hutchinson I., FL. skeleton with 2 rows of cells in cross-section. Three pairs of ventral radiolar appendages. Dorsal Demonax jamaicensis (Augener, 1922) lips erect, ventral lips rounded. Anterior margin of (Fig. 8) anterior peristomial ring with narrow-ventral lobe. In lateral view, ventral margin slightly elevated Parasabella jamaicensis Augener 1922: 48. (Fig. 7A). Segment 1 with broad, dark ventral Demonax jamaicensis.- Johansson 1927: 136.- Perkins 1984: 315-317, fig. 17. shield, with rounded anterior margin. Thorax with 8 chaetigers, each with 2 pigmented bands (Fig. Material: [ZMA] St. Thomas: V. Pol. Benner 7C). Glandular girdle present on chaetiger 2. Bay lagoon, 0-1 m, 30 Apr. 1973, Coll. P. Inferior thoracic notochaetae in 2 transverse Wagenaar Hummelinck, Sta. 1674, Rhizophora groups: anterior rows with 3 or 4 bayonet chaetae (3). (Fig. 7J), posterior rows with 3 or 4 paleate Description: Medium-sized body, 13-23 mm chaetae with small mucro (Fig. 7D-G). Nine acicu- long, 1.5 mm wide. Branchial crown 5 mm long, lar uncini per tori, with 5 equal-sized teeth above with 10 pairs of radioles, with 5 yellow crossbands main fang in lateral view (seen under light on pinnules (in preserved material). Radioles in microscopy) (Fig. 7L). Abdomen with 17 semicircles, without flanges or stylodes. Dorsal chaetigers. Abdominal neurochaetae arranged in margin of dorsal lip extending up along inner mar- 2 transverse rows per fascicle; anterior row on gin of dorsalmost radiole and fused with 1-3 of the anterior chaetigers with 4 or 5 elongate, narrowly more-proximal pinnules. Ventral lips triangular, hooded chaetae; posterior row with modified, elon- well developed, beginning between ventral lappets gate, narrowly hooded chaetae; posterior abdomi- extending dorsally to near dorsal lips. Collar short, nal fascicles with modified, elongate, narrowly widely separated dorsally by fecal groove, begin- hooded chaetae in both rows (Fig. 7H, I). ning slightly anterior and medial to chaetae from Abdominal uncini with main fang surmounted by 3 segment 1 (Fig. 8A). Thorax with 8 chaetigers. or 4 small-sized teeth in lateral view (seen under Ventral lappets rounded (Fig. 8B). Ventral shield light microscopy), breast well developed, manubri- of segment 1 rectangular, about twice as wide as um absent (Fig. 7K). Pygidium rounded, cirrus long, broader than following segments, anteriorly and eyespots absent. Sperm with rectangular indented at midlength; following shields laterally heads, and 2 small mitochondrial lobes. indented by tori (Fig. 8B). Spinelike chaetae on Remarks: In the Grand Caribbean Region no segment 1 in 2 short rows. Inferior thoracic species of Chone has been described. Chone notochaetae broadly hooded (Fig. 8E), arranged in americana Day, 1973 (described from Beaufort, several transverse rows. Thoracic uncini avicular, NC) has a pygidial cirrus, while our specimens with equal-sized teeth above main fang, breast have 5 pairs of radioles, broad flanges, short radio- well developed, manubrium medium (Fig. 8C). lar tips, 17 abdominal segments, all paleate chaeta Companion chaetae with very stout shaft and long with well-developed mucro, sperm with rectangular mucro. Abdomen with 55 chaetigers. Abdominal heads, and no pygidial cirrus. Material of Chone neurochaetae in 2 transverse rows of elongate, sp. 1 is available at ECOSUR and ZMA; we includ- narrowly hooded chaetae (Fig. 8F). Abdominal ed this species in the present paper in order to uncini avicular, with main fang surmounted by 7 facilitate and encourage further research. equal-sized teeth in lateral view, covering 1/2 of main fang length (seen under light microscopy), Demonax Kinberg, 1867 breast well developed, manubrium short (Fig. 8D). Generic diagnosis was provided by Knight- Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 39

Megalomma Johansson, 1927 latter author divided 23 species of Megalomma into 5 groups based on (1) whether the middorsal Megalomma was studied by Perkins (1984), collar margins are fused to the fecal groove or not; Fitzhugh (1989), and Knight-Jones (1997). The (2) whether the dorsolateral margins of the collar

(A) (B)

(E) (F)

(C)

(D)

Fig. 8. Demonax jamaicensis. (A) Anterior end, dorsal view; (B) same, ventral view; (C) thoracic uncinus; (D) abdominal uncinus; (E) thoracic broadly hooded chaeta; (F) abdominal narrowly hooded chaeta. (A-F) ZMA, St. Thomas, 1674. Scale bars (A, B) 1 mm; (C-F) 0.01 mm. 40 Zoological Studies 45(1): 24-66 (2006) form pockets or not; and (3) the extent to which Megalomma heterops Perkins, 1984 eyes occur in the radioles. Nishi (1998) described (Fig. 9) a new species from Thailand, and added a new group (2D) to the description proposed by Knight- Megalomma heterops Perkins 1984: 359-363, figs. 42-43. Jones. Fitzhugh (2002) described a new species from Thailand included in group 1A, and later Material: [FSBC] Florida: Paratypes, 27696, (Fitzhugh 2003) described a new species from St. Lucie Co., Hutchinson I., 27°20'24''N, 83°13' Taiwan (group 1B) and emended the diagnosis of 04''W, Sta. 4, 12.1 m, Coll. R. Gallagher and M. the genus to point out that the radiolar Hollinger, 15 Sept. 1971, Id. T. H. Perkins (1). appendages of the dorsal lips do not have an inter- [FSBC] I 27698, Hutchinson I., St. Lucie Co., 27° nal skeleton, and that dorsal pinnular appendages 21'23''N, 80°13'24''W, 11.2 m, Coll. R. Gallagher can be either present or absent. Furthermore, and M. Hollinger, 15 Sept. 1971, Id. T. H. Perkins Fitzhugh emphasized the implications of those (1). [USNM] Florida: Paratypes, 067953, Sta. J, findings for the determination of cladistic relation- 20 miles W of Sanibel I., 26°24'N, 82°28'W, R/V ships among Sabellinae genera. Hernán Cortés, Coll. Presley, 11 May 1966, 18 m, Three species of Megalomma have been Id. T. H. Perkins (5). [ECOSUR] Mexican described from the Grand Caribbean: M. biocula- Caribbean: I. Cozumel, Coll. S. I. Salazar, 24 Apr. tum (Ehlers, 1887), M. lobiferum (Ehlers, 1887), 2001, coral (2). Cayo Valencia, Coll. M. S. and M. heterops Perkins, 1984, all of them from Jiménez, 29 Apr. 1983 (1). DIF Puerto Aventuras, Florida. Coll. S. I. Salazar, 21 Mar. 1992 (1). Xcacel, Coll. S. I. Salazar, 3 Apr. 1992 (1). Ana y José, Coll. S.

(A) (B) (C) (D) p (E)

(I) (J)

(F)

ss (G) (H) (K)

Fig. 9. Megalomma heterops. (A) Anterior end, dorsal view; (B) same, ventral view; (C) radiolar eye from dorsalmost pair; (D, E) radiolar eyes from lateral radioles; (F) radiolar skeleton in cross-section; (G, H) inferior thoracic paleate notochaetae; (I) thoracic uncinus; (J) abdominal uncinus; (K) pygidium. (A, B) UMML 22.250 Pennekamp Site 5, Florida. (C) FSBC I 27696, paratype, St. Lucie Co., Hutchingson I. FL. (D, G, H) LACM-AHF, Guana I. (E, F) ECOSUR, Isla Cozumel, Mexico. (I, J) USNM 067953, paratype. Scale bars (A, B, K) 0.5 mm; (C-F) 0.12 mm; (G-J) 0.02 mm. ce, compound eye; pe, pygidial eye; p, pocket; rs, radiolar skeleton; ss, sheath tissue. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 41

I. Salazar, J. R. Bastida and J. Ruíz, 11 Feb. 2001 lamellae, beginning between ventral lappets of col- (1). San Felipe, Coll. J. R. Bastida, 19 Feb. 1999 lar on anterior margin of collar, extending anteriorly (3). Xahuayxol, Coll. S. I. Salazar, L. F. Carrera, 1 and dorsally to dorsal lips. Collar not fused to June 1997 (3). [LACM-AHF] British Virgin Is.: Vc fecal groove (Fig. 9A), pockets present. Collar 0566, Guana (1). [UMML] Florida: 22.250, with 1 pair of distally rounded, overlapping, ventral Pennekamp Site 5, Florida Reef Tract, Grecian lappets, which extend beyond beginning of radi- Rocks, July 1980 (1). [USNM] 067955, Off W oles (Fig. 9B). Thorax with 8 chaetigers, 1-5 (3.5- coast, middle ground Florida, Sta. 151, 28 32' 9) mm long, and 0.5-2.5 (0.5-6.5) mm wide. 02''N, 84 18'36''W, 05 Oct. 1978, 25-27 m,° on Ventral shield of segment 1 entire, with rounded Madracis ° decactis, Id. T. H. Perkins (3). [USNM] anterolateral corners and slight anteromedial Puerto Rico: 42791, J2 1963, 7.6 m, Coll. Hulings indentation, twice as long as other shields and and Feray, muddy sand, Id. N. Foster. [USNM] about the same width, ventral shields rectangular, 53233, Guayanilla Bay, Coll. T. Morgan, 28 Oct. not indented by tori (Fig. 9B). Tori gradually short- 1971, Thalassia bed, 1.2 m, Id. T. H. Perkins. ening posteriorly. Superior thoracic notochaetae, Description (variations observed in paratypes elongate narrowly hooded. Inferior thoracic given in parentheses): Body pale (in preserved notochaetae broadly hooded (Fig. 9G, H), material), 4-47 (4.5-27) mm long, anterior margin arranged in 2 or more transverse rows. Thoracic of segment 1 with paired purple spots dorsally uncini avicular, with 16 equal-sized teeth in lateral below collar, paired spots on chaetigers 1-3 dorsal view, covering 1/2 of extension of main fang (seen to notopodia. Branchial crown long, with a row of under light microscopy), breast well developed, pigmented spots on palmate membrane, narrow manubrium long (Fig. 9I). Companion chaetae purple bands on outer and lateral surfaces of radi- with roughly symmetrical tips as teardrop-shaped oles, not extending to pinnules, and 13-15 pairs of membranes. Abdomen with 36-98 (26-92) radioles alternating 5 or 6 purplish-yellow bands (in chaetigers. Abdominal neurochaetae in 2 trans- preserved material). Radioles with subterminal verse rows: anterior and posterior rows on anterior compound eyes, 1st pair on dorsalmost radioles chaetigers with elongate, narrowly hooded and enlarged with distinct ommatidia; other eyes much modified, elongate, narrowly hooded chaetae, smaller, gradually decreasing in size ventrally, with respectively. Abdominal uncini avicular, with 13 smooth surface and ommatidia visible only under equal-sized teeth in lateral view, covering 1/2 of high magnification of light microscope; dorsalmost extension of main fang (seen under light radioles subdistally enlarged, with short tips microscopy), breast well developed, manubrium extending beyond eyes (Fig. 9C-E). Radiolar short (Fig. 9J). Pygidium rounded with irregularly skeleton with 8 cells in cross-section surrounded arranged eyespots (Fig. 9K). Tubes with pieces of by moderately thick sheath and columnar epitheli- shells and coral fragments of different sizes, poste- um (Fig. 9F). Dorsal lips with dorsal radiolar rior end translucent amber. appendages and dorsal pinnular appendages, pig- Remarks: Perkins (1984) described M. het- mented on upper surface. Ventral lips with parallel erops from Florida, and it corresponds to the group

Table 1. Species of Megalomma recorded or discussed in this study and their classification according to Knight-Jones (1997)

Group Dorsal margin of collar Subterminal eyes Caruncle Species Type locality

1A Fused to fecal groove, on most radioles present M. lobiferum Florida pockets present 2A Not fused to fecal groove, on most radioles absent M. heterops Florida pockets present absent M. neapolitanum Italy absent M. perkinsi sp. n. Florida 2B Not fused to fecal groove, dorsalmost pair of absent M. bioculatum Florida pockets absent radioles present M. pigmentum Baja California present Megalomma sp. 1 Venezuela absent Megalomma sp. 2 Florida 42 Zoological Studies 45(1): 24-66 (2006)

2A (Table 1) designated by Knight-Jones (1997). ommatidia in the eyes of the 2nd and 3rd dor- Perkins indicated that there are 2 similar forms of salmost pairs. Megalomma sp. A is formally M. heterops from North Carolina, Florida, and described as M. perkinsi sp. nov. (see below). Venezuela (sp. A), and from Puerto Rico (sp. C), both with reduced flat eyes. Specimens of Megalomma lobiferum (Ehlers, 1887) Megalomma sp. C., are in poor condition [USNM (Fig. 10) 42791 (1 incomplete) and USNM 53233 (1 complete)]; they have some rounded pigmented areas Branchiomma lobiferum Ehlers 1887: 254-259, pl. 53, figs. 10- in the 3rd dorsal pair of the radioles, and distinct 15.- Hoagland 1919: 577.- Treadwell 1924a: 18; 1939:

(A) (B)

(J)

(G) (H) (I)

(M)

(F) (E) (D) (N)

Fig. 10. Megalomma lobiferum. (A) Anterior end, dorsal view; (B) same, ventral view; (C) thoracic shields; (D-F) companion chaetae; (G-I) spinelike chaetae from segment 1; (J) inferior thoracic paleate notochaetae; (K, L) abdominal spinelike chaetae; (M) abdominal uncinus; (N) thoracic uncinus. (A-N) ECOSUR, Isla Contoy, Mexico. Scale bars (A-C) 0.5 mm; (D-L) 0.02 mm. ca, caruncle. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 43

291-292, fig. 105.- Fauvel 1953: 17. elongate, narrowly hooded chaetae, respectively Megalomma lobiferum.- Johansson 1927: 132.- Perkins 1984: (Fig. 10K, L). Abdominal uncini avicular, with 13 354-357, figs. 39, 40. equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (visible Material: [ECOSUR] Mexican Caribbean: I. under light microscopy), breast well developed, Contoy, Punta Sur, Coll. S. I. Salazar, 2 Mar. 2001 manubrium short (Fig. 10M). (1). Veracruz, I. Verde, Coll. S. I. Salazar, 5 Aug. Remarks: The most-distinctive character in M. 1985 (1). Panama: Fuerte Sherman, Colón, Coll. lobiferum is the“caruncle”; this organ also occurs S. I. Salazar, 2 June 2002, T=30, S=30/00 (1). in M. pigmentum and Megalomma sp. 1 (Table 1). [LACM-AHF] British Virgin Is.: AF00-59, Vc 0634, Megalomma lobiferum differs of M. pigmentum and Guana, Beef I., Trellis Bay, Coll. K. Fitzhugh, 12 Megalomma sp. 1 in the occurrence of subterminal July 2000, tubes taken from coral rubble (2). eyes on most radioles, dorsal collar margins fused [LACM-AHF] AF00-59, Vc 0565, Beef I., Trellis to the fecal groove, and the presence of pockets. Bay, directly off airport, Coll. K. Fitzhugh, 12 July 2000 (1). [USNM] Panama: 073019, Galeta reef, Megalomma perkinsi sp. nov. Coll. A. A. Reimer, Id. T. H. Perkins, Apr. 1982 (6). (Fig. 11) Description: Body 15-36 mm long. Branchial crown purple, 4-14 mm long with 15-27 pairs of Megalomma sp. A Perkins 1984. radioles, cross-banded with yellowish-orange pigment extending to pinnules (in preserved material). Etymology : This species is named in honor of Subterminal compound eyes on all radioles, mod- Thomas Perkins (Florida Marine Research erately large on dorsalmost pair, gradually Institute, St. Petersburg, FL) in recognition of his decreasing in size laterally and ventrally. Except important contribution to the knowledge of poly- for size, all eyes similar, well defined, spherical, chaetes in general, and especially of sabellids. with distinct ommatidia. Radiolar skeleton with 4 Material: [USNM] Florida: Holotype 53976, or more rows of cells in cross-section. Branchial Cape Lookout, NC, Coll. S. L. Gardiner, 16 Apr. crown with long fused region. Dorsal lips long, 1976, intertidal sand mixed with gravel and shell with dorsal radiolar appendages, with margin of fragments 129, Id. S. L. Gardiner. [USNM] upper lamellae fused proximally to base of basal Paratype 53977, Cape Lookout, NC, 28 July 1976, pinnules of dorsalmost pair of radioles. Ventral lips Coll. C. Jenner, intertidal sand mixed with gravel short, rounded, with parallel lamellae. Caruncle and shell fragments 146, Id. S. L. Gardiner as M. above mouth, anterior to and in between dorsal lobiferum. lips (Fig. 10B). Dorsal margin of collar fused to Description (variations observed in paratype fecal groove (Fig. 10A) forming deep pockets. given in parentheses): Specimen large, 107 (92) Ventral lappets short, rounded, overlapping (Fig. mm long. Branchial crown 12 mm long, with 22 10B). Ventral shield of segment 1 up to twice as (23) pairs of radioles with 3 purplish-brown bands broad as long, convex laterally, straight to convex at 1st 1/2 of crown, dorsalmost pair longer and anteriorly (Fig. 10C); following shields concave, pale (in preserved material). Radioles with subter- rounded laterally around tori. Thorax 3-6 mm minal compound eyes on most radioles, flat, those wide, with 8 chaetigers. Collar with long, narrowly from dorsalmost pair bigger than others (Fig. 11H), hooded chaetae (Fig. 10G-I). Superior thoracic surrounding almost covering tip dorsally, small notochaetae, elongate narrowly hooded. Inferior ventrally. Radiolar skeleton with 4 or more rows of thoracic notochaetae broadly hooded, arranged in cells in cross-section (Fig. 11G). Dorsal lips long 2 or more transverse rows (Fig. 10J). Thoracic with broad base (Fig. 11C), with orange pigment uncini avicular with several rows of equal-sized laterally and radiolar pinnular appendages. Ventral teeth in lateral view, covering 1/2 of extension of lips with parallel lamellae. Dorsal margins of collar main fang a number of teeth not visible under light not fused to fecal groove, with 2 concave lobes microscopy), breast well developed, manubrium forming deep pockets (Fig. 11B); collar with a long (Fig. 10N). Companion chaetae with roughly small lateral indentation (Fig. 11A). Ventral shield symmetrical tips, as teardrop-shaped membranes of segment 1 as wide as long (Fig. 11C), following (Fig. 10D-F). Abdomen with 43-84 chaetigers. shields rectangular. Thorax 11 (10) mm long, 4.5 Abdominal neurochaetae in 2 transverse rows: (5) mm wide, with 8 chaetigers. Superior thoracic anterior and posterior rows on anterior chaetigers notochaetae, elongate narrowly hooded. Inferior with elongate, narrowly hooded and modified, thoracic notochaetae broadly hooded (Fig. 11K), 44 Zoological Studies 45(1): 24-66 (2006) arranged in 2 or more transverse rows. Thoracic manubrium short (Fig. 11I). Maturating oocytes in uncini avicular, with 22 equal-sized teeth above abdomen (chaetiger 11). Tube with large shells main fang in lateral view, covering 1/2 of extension and stone fragments. of main fang, breast well developed, manubrium Remarks: Megalomma perkinsi sp. nov. long (Fig. 11D). Companion chaetae with roughly belongs to group 2A proposed by Knight-Jones symmetrical tips, as teardrop-shaped membranes (1997), which is defined by the dorsal margins of (Fig. 11E-F). Abdomen with 66 (104) chaetigers. collar not being fused to the fecal groove, the pres- Abdominal neurochaetae in 2 transverse rows: ence of dorsolateral collar pockets, and the occur- anterior and posterior rows on anterior chaetigers rence of eyes on most radioles. The other species with elongate, narrowly hooded and modified, belonging to this group include M. heterops elongate, narrowly hooded chaetae, respectively Perkins 1984 from Florida and M. neapolitanum (Fig. 11J). Abdominal uncini avicular, with 18 (Claparède, 1868) from Italy (Table 1). equal-sized teeth in lateral view, covering 1/2 of Based on the description and illustration pro- extension of main fang, breast well developed, vided by Claparède (1868: pl. XXII, fig. 5), the

(A) (B) (C) dl

(D)

(E) (F)

(J) (K) (G) (I) ce (H)

Fig. 11. Megalomma perkinsi sp. nov. (A) Anterior end, lateral view; (B) same, dorsal view; (C) dorsal lips; (D) thoracic uncinus; (E, F) companion chaetae; (G) radiolar skeleton in cross section; (H) radiolar eye; (I) abdominal uncinus; (J) abdominal elongate narrowly hooded chaeta; (K) inferior thoracic broadly hooded chaeta. (A-K) USNM-53976, holotype. Florida. Scale bars (A-C) 1 mm; (D-F, J, K) 0.06 mm; (G, H) 0.1 mm; (I) 0.05 mm. bv, blood vessel; ce, compound eye; dl, dorsal lips; rs, radiolar skeleton. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 45 anterior dorsal margin of the collar in M. neapoli- proposed by Knight-Jones (1997) (Table 1). tanum is of even height, forming 2 small pockets, In Megalomma sp. 1 the 1st shields are simi- compound radiolar eyes are hemispherical with big lar to those of M. pigmentum Reish; however, ommatidia (more than in M. heterops), covering all Megalomma sp. 1 has a caruncle, a structure pre- radiolar tip. In M. perkinsi, the anterior dorsal mar- viously recorded only for M. lobiferum, and in this gin of the collar is high; forming 2 big, deep pock- study for M. pigmentum Reish. Megalomma bioc- ets, compound radiolar eyes are rounded with ulatum has a collar with triangular ventral lappets, small ommatidia, not covering the radiolar tip. and a quadrangular ventral shield on segment 1 Megalomma heterops differs from M. perkinsi with an anterior M-shaped margin, and lacks the because in the latter species, the anterior margin caruncle, while M. pigmentum Reish has a collar of the collar has a lateral depression, reduced and with lanceolated ventral lappets, the ventral shield flattened radiolar compound eyes, with distinct of segment 1 is divided into 2 asymmetrical parts, ommatidia only in the 2nd and 3rd dorsalmost and there is a caruncle, as corroborated by topo- pairs, and rounded pigmented areas in 3rd dorsal typical specimens [ECOSUR: Bahía San Quintín, pair of radioles. Coll. L. E. Calderón, 8 Dec. 1981]. We included this species in the present paper Megalomma sp. 1 in order to facilitate and encourage further research, because the presence of a caruncle in Megalomma pigmentum.- Perkins 1984: 357-359, fig. 41C (non species of Megalomma should be more frequently Reish). included in taxonomic studies since it provides additional information for investigation of phyloge- Material: [USNM] Venezuela: 57945, Bahía netic relationships. de Mochima, Cumaná (7), Coll. Edwards, 22 June 1971, 6-10 m, in calcareous sand, Id. T. H. Megalomma sp. 2 Perkins, 1981. Description: Five complete, pale (in preserved Megalomma pigmentum Perkins 1984: 358, fig. 41A, B (non material), small specimens, 4-23 mm long. Crown Reish). 3-7 mm long, with 12 pairs of radioles, base and interradiolar spots orange on preserved material. Material: [USNM] Florida: 45691 (7), Tampa Dorsal lips with radiolar and pinnular appendages. Bay, Coll. J. L. Taylor, 1963, Id. T. H. Perkins, Ventral lips with parallel lamellae. Ventral lappets 1981. lanceolated. Dorsal margins of collar not fused to Description (data of best specimen given in fecal groove, pockets absent, eyes only present on parentheses): Two complete specimens and 7 dorsalmost pair of radioles. Caruncle thin, above fragments. Body pink (in preserved material), 3-19 the mouth, anterior to and in between dorsal lips. (24) mm long. Crown 4-6 (5.5) mm long, with 10 First ventral shield divided into 2 parts, basal part or 11 (10) pairs of radioles. Dorsal lips with dorsal larger, remaining thoracic shields rectangular. radiolar appendages and dorsal pinnular Thorax 2.5-5.5 mm long, 1-1.5 mm wide, with 8 appendages. Ventral lips with parallel lamellae. chaetigers. Inferior thoracic notochaetae broadly Ventral lappets lanceolated. Dorsal margins of col- hooded, arranged in transverse rows. Companion lar not fused to fecal groove, pockets absent, eyes chaetae with roughly symmetrical tips. Abdomen only present on dorsalmost pair of radioles. with 14-43 chaetigers. Abdominal neurochaetae Caruncle absent. First ventral shield divided into 2 with elongate, narrowly hooded chaetae. parts, basal part larger, remaining thoracic shields Remarks: Perkins (1984) recorded M. pig- rectangular. Thorax 2-5.5 (3.5) mm long, 1-2 mm mentum Reish from Florida and Venezuela; he wide, with 7-9 (8) chaetigers. Inferior thoracic assumed that the main difference between Atlantic notochaetae broadly hooded, arranged in trans- specimens and the type material was the color. verse rows. Companion chaetae with roughly Nevertheless, the material from the Grand symmetrical tips. Abdomen with 3-21 (60) Caribbean region (USNM), identified by Perkins as chaetigers. Tube covered with transparent crys- M. pigmentum (non Reish), corresponds to 2 tals. undescribed species, one from Venezuela Remarks: Megalomma sp. 2 is close to (Megalomma sp. 1), the other from Florida Megalomma sp. 1, the 1st ventral shield of both is (Megalomma sp. 2 see below). The 3 species cor- divided into 2 pairs; nevertheless, they only differ respond to group 2B according to the classifictaion because Megalomma sp. 2 lacks a caruncle (Table 46 Zoological Studies 45(1): 24-66 (2006)

1). Drawings of Perkins (1984: 358, fig. 41A, B) Hypsicomus to Notaulax, described 2 new species, illustrate the anterior region of Megalomma sp. 2 and provided a key for the identification of species very clearly. Descriptions of Megalomma sp. 1 of Notaulax. The genus is characterized by the and Megalomma sp. 2 are brief and without draw- arrangement of collar chaetae, the presence of ings; however, the material is available at the superior thoracic spinelike notochaetae, and the USNM. long branchial base flanges (Fitzhugh, 1989). Six species have been described from the Grand Notaulax Tauber, 1879 Caribbean: N. bahamensis Perkins, 1984 and N. paucoculata Perkins, 1984 from the Bahamas; N. Perkins (1984) made a preliminary revision of circumspiciens (Ehlers, 1887), and N. midoculi Hypsicomus and Notaulax, in which he (Hoagland, 1919) from Florida; N. nudicollis redescribed some species, moved most species of (Krøyer, 1856) from St. Thomas, and N. occiden-

(D) (C) rf

(A) (B) se

(G) (E) (F) (H)

(O) (P)

m (N) (I) (J) (K) (L)

(M)

Fig. 12. Notaulax bahamensis. (A) Anterior end, dorsal view; (B) same, ventral view; (C) flanged radiolar tip; (D) radiole, area of eyes; (E, F) collar chaetae; (G) thoracic uncinus; (H) abdominal uncinus, non-lateral oblique; (I, J) thoracic paleate inferior notochaeta; (K, L) thoracic superior spinelike notochaetae; (M) companion chaeta; (N) abdominal, modified, elongate, narrowly hooded chaetae; (O, P) anterior abdominal paleate chaetae. (A-P) ECOSUR, Isla Cozumel, Mexico. Scale bars (A, B) 0.5 mm; (C) 0.06 mm; (D) 0.1 mm; (E, F, I, J, N-P) 0.01 mm; (G, H, K-M) 0.02 mm. cc, collar chaetiger; m, mucro; rf, radiolar flange; se, simple eyespot. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 47 talis (Baird, 1856) from St. Vincent. verse rows of chaetae: anterior row on anterior and posterior abdominal chaetigers paleate, Notaulax bahamensis Perkins, 1984 mucroate with small teeth at base of mucro (Fig. (Fig. 12) 12O, P); posterior row on anterior abdominal chaetigers elongate, narrowly hooded; posterior Notaulax bahamensis Perkins 1984: 348, figs. 35, 36. row in posterior abdominal chaetigers modified, elongate, narrowly hooded (Fig. 12N). Abdominal Material: [ECOSUR] Mexican Caribbean: I. uncini avicular, with 14 equal-sized teeth above Cozumel, Chankanaab, Coll. S. I. Salazar, 2 Apr. main fang in lateral view, covering 3/4 of extension 1992 (1). [USNM] Holotype 062082, Lucaya, of main fang (seen under light microscopy), breast Grand Bahama I., Hydrolab, off Bell Channel, 26 well developed, manubrium short (Fig. 12H). 33'00''N, 78°34'00''W, Coll. B. A. Vittor and T. S. Pygidial eyespots present. Hopkins, 28 Jan. 1974, on coral. ° Remarks: Notaulax bahamensis and N. nudi- Description: Body pale in preserved materials, collis share a roughly similar arrangement of eye- 34 mm long. Crown with 4 brown bands (in pre- spots, but in the 1st species, the radioles are served material), 2 in palmate membrane, another flanged except in a small portion on the eye band, immediately above it, and the last at midlength of while in the latter species, the radioles are com- free parts of radioles. Crown 15 mm long, with 11 pletely flanged. Further, N. bahamensis has a very or 12 pairs of radioles united proximally by palmate long collar (longer than the next 3 segments), membrane for about 40% of their length above while the collar of N. nudicollis is as long as the basal lamina, flanged from palmate membrane to next 2 segments. The holotype is damaged, the tips (Fig. 12C) except for a small part of region of crown is almost separated from the body, and simple eyespots (Fig. 12D). Basal lamina very some radioles were already removed. long, as long as palmate membrane, eyes in oval groups (30), separated from palmate membrane Notaulax midoculi (Hoagland, 1919) for about 1/3 of its length; dorsal and ventral mar- (Fig. 13) gins of basal lamina flanged, from bases of lobes to origins of dorsalmost and ventralmost radioles. Parasabella midoculi Hoagland 1919: 579, pl. 31, figs. 10-14, Radiolar skeleton with 4 or more rows of cells in pl. 32, figs. 1, 2. cross-section. Simple radiolar eyes present above Parasabella sulfurea.- Treadwell 1924a: 18 fide Perkins 1984: 343. palmate membrane (Fig. 12D). Dorsal lips with Hypsicomus midoculi.- Johansson 1927: 141. radiolar appendages; pinnular appendages absent. Ventral lips and parallel lamellae present. Collar Material: [ECOSUR] Mexican Caribbean: I. as a single lobe, beginning near dorsal midline at Contoy, Boca Norte, Pueblo Viejo, Coll. M. A. margin of posterior peristomial ring and extending Tovar, 1 Mar. 2001, in sponge on dead coral (1); anteriorly, almost straight laterally, with convex Punta Sur, Coll. M. A. Tovar, 2 Mar. 2001 (4). I. ventral margin (Fig. 12A, B); segment 1 much Cozumel, SEDENA, Coll. A. Medina, 24 Mar. 2001 longer than following segments (Fig. 12A, B), ven- (4); Playa Azul, Coll. L. E. González, 25 Mar. 2001 tral shield subtriangular. Chaetae on segment 1, (1); Chankanaab, Coll. S. I. Salazar, 2 Apr. 1992, as single, elongate row of spinelike chaetae (Fig. QR-7 (1); Muelle Puerta Maya, Coll. A. Medina, 26 12E, F) on posterior 1/3 of collar, fascicles longitu- Mar. 2001 (2). Majahual Norte, Coll. S. I. Salazar, dinal to oblique (Fig. 12A). Thorax with 8 18 Mar. 2001, on dead coral (1). [UMML] Florida: chaetigers, 5 mm long, 2.8 mm wide. Thoracic 22.166, Reef Tract, Margot Fish Shoal, 16 Feb. superior notochaetae spinelike (Fig. 12K, L); inferi- 1962, MFS (18 l), Coll. Jones, Work, Bayer, and or notochaetae paleate, arranged in 2 transverse Ebbs (1). [USNM] Barbados: 020304, Bathsheba, rows, symmetrical (Fig. 12I, J). Thoracic uncini Coll. Univ. of Iowa, Summer, 1918, Barbados- avicular, with 16 equal-sized teeth above main Antigua Expedition, Id. T. H. Perkins, 1980 (1). fang in lateral view, covering 1/2 of extension of Description: Body 18-28 mm long. Crown 8- main fang (seen under light microscopy), hood 14 mm long, with 10-16 pairs of radioles united absent, breast well developed, manubrium medi- above basal lamina by a palmate membrane, um (Fig. 12G). Companion chaetae with roughly extending for 1/6 of their length. Radioles with symmetrical tips, as teardrop-shaped membranes purple (in preserved material) very long basal lami- (Fig. 12M). Abdomen with 108 chaetigers. na (Fig. 13A, B), followed by a white band (10 pin- Abdominal neurochaetal fascicles with 2 trans- nules), at midlegth an orange band (13 pinnules), 48 Zoological Studies 45(1): 24-66 (2006) another white band (5 pinnules), and a pale- 13B). Shield of segment 1 not well defined; follow- orange band (6 pinnules), the latter with 2 groups ing segments with trapezoidal ventral thoracic of 9-14 small simple eyespots at both sides of radi- shields, lateral sides strongly indented by tori (Fig. ole (Fig. 13C). Radioles slightly flanged near tips. 13B). Thorax 3-4 mm long, 2-3 mm wide, with 8 Radiolar skeleton with 6 rows of cells in cross-sec- chaetigers. Single, elongate, longitudinal to tion (Fig. 13D). Dorsal lips with radiolar oblique row of spinelike chaetae on segment 1. appendages but without dorsal pinnular Superior thoracic notochaetae spinelike (Fig. appendages. Ventral lips and parallel lamellae 13M), inferior notochaeta paleate, arranged in 2 present. Collar with rounded ventral lappets (Fig. transverse rows, with symmetrical and asymmetri-

(A) (B) (C)

(D)

(J) (E) (F) (G) (H)

(I)

(K) (L) (M) (N) (O) (P)

Fig. 13. Notaulax midoculi. (A) Anterior end, dorsal view; (B) same, ventral view; (C) radiole, area of eyes; (D) radiolar skeleton, transverse view; (E) thoracic uncinus; (F) abdominal uncinus; (G-J) companion chaetae; (K, L) thoracic inferior paleate notochaeta; (M) thoracic broadly hooded chaeta; (N, O) anterior abdominal paleaete chaetae; (P) abdominal, modified, elongate, narrowly hooded chaetae. A-P: ECOSUR, Isla Contoy, Mexico. Scale bars (A, B) 0.5 mm; (C, D) 0.02 mm; (E-P) 0.01 mm. m, mucro; rs, radiolar skeleton; se, simple eye; bv, blood vessel. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 49 cal chaetae (Fig. 13K, L). Thoracic uncini avicular, Punta Nizuc, Coll. S. I. Salazar and L. F. Carrera, with 22 equal-sized teeth above main fang in later- 30 Aug. 1997 (2). I. Cozumel, Paraiso, Coll. S. I. al view, covering 1/2 of extension of main fang Salazar, 5 June 1996 (1). Buenavista, E-25, Coll. (seen under light microscopy), hood absent, breast S. I. Salazar and L. F. Carrera, 27 Sept. 1996 (3); well developed, manubrium medium (Fig. 13E). R-5 (1); R-4 (1); E-24 (1). Majahual, Coll. M. A. Companion chaetae with roughly symmetrical tips, Tovar, 10 Jan. 2001 (1). Rio Indio, Coll. S. as teardrop-shaped membranes (Fig. 13G-J). Frontana, 17 Mar. 2001 (1). San Felipe, Coll. J. R. Abdomen with 22-41 chaetigers. Abdominal neu- Bastida and S. I. Salazar, 19 Nov. 1999 (1). Sol y rochaetae fascicles in 2 transverse rows of Mar (2). Xahuayxol, Coll. S. I. Salazar and L. F. chaetae: anterior row on anterior and posterior Carrera, 27 Sept. 1996, E-4 (2), 3 June 1998, C28 abdominal chaetigers paleate, with long mucro RC1 (2); 1 June 1997, A-46, in Strombus gigas (3); (Fig. 13N, O) with small teeth at base of mucro; 27 June 1997, E2 (1); 19 Oct. 1997, B-45, Coll. S. posterior row on anterior abdominal chaetigers I. Salazar and L. F. Carrera (1). [LACM-AHF] elongate, narrowly hooded; posterior row on poste- British Virgin Is.: BVI-99, LH0995, Guana, Sta. 035 rior abdominal chaetigers modified, elongate, nar- (3). [UMML] Florida: 22.87, Straits off Florida, R/V rowly hooded (Fig. 13P). Abdominal uncini avicu- Gerda, Sta. 1033, 24 36'N, 81 06'W, 42 m, 26 lar, with 19 equal-sized teeth above main fang in Feb. 1969, Coll. G-1033,° Id. T.° H. Perkins (6). lateral view, covering 3/4 of extension of main fang [USNM] 52042, Big Pine Key, FL, Coll. D. (seen under light microscopy); breast well devel- Maynahan, 14 Aug. 1973, from rock wall of canal, oped, manubrium short (Fig. 13F). 0.6-1.8 m, Id. T. H. Perkins (3), lot contains a Remarks: Perkins (1984) reviewed a speci- brown crown, 1 thorax, 2 fragments and rest of men from Barbados in which the ventral margin of tranlucid tubes. [USNM] 57933, Safe Harbor, Stock the collar had a slight incision, and the ventral I., near Key West, FL, Sta. 7A, 2.4 m, 7 July 1970, lobes were not very well defined; in his figures, the Coll. R. Chester, Id. T. H. Perkins (2), lot contains 1 shield of segment 1 is distinct and rectangular, and complete specimen, 1 incomplete (missing posteri- the other shields are rectangular with concave lat- or segments), and 2 crowns. [ZMA] North eral sides. Specimens identified in this study as N. Carolina: V. Pol. Bogue Sound, wooden piling at midoculi have a collar that is well incised ventrally, Fisheries Institute, Coll. P. Wagenaar Hummelinck, forming 2 rounded lobes, and the ventral shield of 28 Aug. 1963, 0-1 m (7). segment 1 is indistinct, as originally illustrated by Description: Body 12-24 mm long. Branchial Hoagland (1919 fig. 10) and Perkins (1984 fig. crown 5-5.5 mm long with 5 brown to purple cross 32H) from the holotype. The USNM specimen was bands (in preserved material) and 10-14 pairs of collected in 1918, and has been reviewed by radioles; radiolar eyes in elongate-oval groups of Treadwell (as Parasabella sulfurea), Hartman (as 30-70 eyes of each side, diminishing in number Hypsicomus circumspiciens), and Perkins. It is distally (Fig. 14C). Radiolar flanges beginning broken into 3 parts, has a transverse incision above eyes and extending to tips. Radiolar skele- showing the coelom full of oocytes, and another ton with 4 rows of cells in cross-section (Fig. 14D). dorsal incision from chaetiger 5 (thorax) towards Dorsal lips extending laterally in almost straight the midbody. line, with radiolar appendages but without pinnular appendages. Ventral lips and parallel lamellae Notaulax nudicollis (Krøyer, 1856) present. Dorsal and ventral margins with flanges. (Fig. 14) Dorsal margin of collar incised (Fig. 14A), ventral margin entire (Fig. 14B). Thorax with 8 chaetigers, Sabella nudicollis Krøyer 1856: 30-31. 2.5 mm long and 1-1.5 mm wide. Ventral shield of Protulides elegans Webster 1884: 325-326, pl. 11, figs. 63-74 segment 1 rectangular to trapezoidal (Fig. 14B). fide Perkins 1984: 332. Notaulax nudicollis.- Perkins 1984: 331, figs. 25-28. Chaetae of segment 1 in a longitudinal to oblique single elongate row of spinelike chaetae. Superior Material: [ECOSUR] Mexican Caribbean: DIF thoracic notochaetae spinelike. Inferior thoracic Puerto Aventuras, QR-4, Coll. S. I. Salazar, 21 notochaeta paleate, nearly symmetrical (Fig. 14H- Mar. 1992 (3). Contoy, Coll. L. F. Carrera and S. I. J), arranged in 2 rows. Thoracic uncini avicular Salazar, 10 June 1999 in Ircinia (3); Faro, Coll. P. with 17 or 18 equal-sized teeth above main fang in Salazar, 1 Mar. 2001 (1); Camping, 22 Feb. 1999 lateral view (seen under light microscopy), cover- (1); Coll. S. I. Salazar, 1 Mar. 2001, in Strombus ing 1/2 of extension of main fang, hood absent, gigas (3); Punta Sur, 28 Feb. 2001 (2). Cancún, breast well developed, manubrium medium (Fig. 50 Zoological Studies 45(1): 24-66 (2006)

14E). Companion chaetae with roughly symmetri- microscopy), covering 3/4 of extension of main cal tips, as teardrop-shaped membranes (Fig. fang, breast well developed (Fig. 14F), manubrium 14G). Abdomen with 29-115 chaetigers. short. Abdominal neurochaetae in 2 transverse rows of Remarks: Perkins (1984) recorded N. nudicol- chaetae: anterior row paleate with short mucro on lis from North Carolina, Florida, Puerto Rico, the anterior and posterior abdominal chaetigers (Fig. Virgin Is. Brazil, Western Africa, and the Mexican 14L); posterior row on anterior abdominal Pacific; he stated:“Minor differences between chaetigers elongate, narrowly hooded; posterior specimens from the Caribbean Sea and adjacent row on posterior abdominal chaetigers modified, areas and those from West Africa, Brazil, and elongate, narrowly hooded (Fig. 14K). Abdominal Western Mexico are attributed to geographic sepa- uncini avicular, with 13 equal-sized teeth above ration but are not considered to be sufficient to main fang in lateral view (seen under light indicate speciation”. Nevertheless, the morpholo-

(A) (B) (C)

bv (D)

ss (E)

m (H) (I) (J) (K) (L)

(G)

(F)

Fig. 14. Notaulax nudicollis. (A) Anterior end, dorsal view; (B) same, ventral view; (C) radiole, area of eyes; (D) radiolar skeleton in cross-section; (E) thoracic uncinus; (F) abdominal uncinus, non-lateral oblique; (G) companion chaetae; (H-J) inferior thoracic paleate notochaeta; (K) abdominal modified, elongate, narrowly hooded chaeta; (L) abdominal paleate chaeta. (A-L) ECOSUR, Isla Contoy, Mexico. Scale bars (A, B) 0.05 mm; (C-G, K) 0.01 mm; (H-J, L) 0.02 mm. bv, blood vessel; m, mucro; pi, pinnule; rs, radiolar skeleton; se, simple eyespots; ss, sheath tissue. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 51 gy of the collar and the distribution of the eyes on fide Perkins 1984: 339. the radioles were variable in his material; further, Notaulax occidentalis.- Perkins 1984: 339-342, figs. 29, 30. Perkins (1984: 332) synonymized Sabella torquata Grube, 1877, described from Western Africa, with Material: [ECOSUR] Mexican Caribbean: I. N. nudicollis (Krøyer, 1856) described from St. Cozumel, Paraíso, Coll. S. I. Salazar, 5 June 1995 Thomas. However, these species have remark- (1); Chankanaab, Coll. S. I. Salazar, 2 Apr. 1992 able differences: the ventral, anterior collar margin (2). I. Contoy, Punta Sur, Barlovento, Coll. S. I. of the syntypes of S. torquata is triangular and Salazar, 2 Mar. 2001 (3). DIF Puerto Aventuras, medially indented (Perkins, Fig. 25F), and there QR-4, Coll. S. I. Salazar, 21 Mar. 1992 (2). are groups of more than 40 eyes, separated from Majahual Norte, Coll. M. A. Tovar, 19 Jan. 2001 the palmate membrane by a distance shorter than (2). Cancún, Punta Nizuc, Coll. S. I. Salazar and that in N. nudicollis (Perkins 1984, fig. 25D). L. F. Carrera, 1 Sept. 1997 (2); 31 Aug. 1997, Perkins mentioned that these characters were sim- rocks 4 (2). Buenavista, R4, Coll. S. I. Salazar and ilar in 1 specimen from the Mexican Pacific L. F. Carrera, 27 Sept. 1996 (1). Venezuela: (Zihuatanejo), but with up to 30 eyes. However, Turpialito, Cumaná, Coll. M. Liñero, 22 Feb. 2002, this record is questionable, since in Perkins’ in Millepora (3). [LACM-AHF] British Virgin Is.: Vc drawings (Perkins 1984, fig. 26C, D), the ventral 0675, Guana (1); AF00-49, Vc 0517 (1); AF00-65, collar is complete, of the same width all along its Vc 0791 (1); AF01-67, Vc 0592 (1). [UMML] extension, and the dorsal margin of collar is more Florida: 22.761, Reef Tract, Margot Fish Shoal, incised than in the holotype of N. nudicollis. Red Head, 16 Feb. 1962 (1), Coll. Jones, Work, Further, the material of Zihuatanejo (USNM 41528, Bayer, and Ebbs (1). [UMML] British Virgin Is.: Coll. Klawe, 6 Sept. 1968, on dead coral, Id. T. H. 22.98, St. John, Great Cruz Bay, Virgin Is., Coll. Perkins) consists only of a trunk fragment in a very Thomas, 15 Dec. 1958 (2). [USNM] Puerto Rico: poor condiction missing the crown, collar, and 16218, Reef at Ponce, 1898-1899, Coll. Str. Fish pygidium. Hawk, Donor US Fish. Comm., Id. T. H. Perkins Two specimens from Bahía de los Ángeles, (2). [USNM] Bahamas: 062083, Lucaya, Grand Baja California (ECOSUR: 23 Aug. 1987, Coll. Bahama I., Hydrolab, off Bell Channel, 26 33' , ° SISV) are identical to Perkins drawings for the 00''N, 78°34'00''W, Coll. B. A. Vittor and T. S. Zihuatanejo specimen; but we considered them as Hopkins, 28 Jan. 1974, 15 m, Id. T. H. Perkins (1). belonging to a different species. The number of [USNM] Panama: 073020, Galeta Reef, Coll. A. A. eyes is a character which changes with age, but Reimer, 5 Oct. 1970, Id. T. H. Perkins (2). the pattern of radiolar bands is constant and useful Description: Body brown with a brown band to separate species. Thus, some species have along dorsum, dark brown ventrally, 22-57 mm bands near the basal membrane, others at radiolar long. Base of crown purple to brown (in preserved midlength, and still others near the distal part of material). Palmate membrane darker than body, , the radioles. Perkins (1984) drawings of N. nudi- with 4-8 cross bands of purple to brown pigmenta- collis from Florida (Perkins 1984, fig. 27A-G), show tion, pigment extending into pinnules. Branchial at least 2 different forms, differentiated by the ven- crown 9-18 mm long with 17-30 pairs of radioles; tral morphology of the collar: one with a wide, palmate membrane extending for about 1/3 of total entire, distally rounded anterior margin (Perkins length of branchial crown, with basal lamina about 1984, fig. 27D), and the other with the collar seg- 2/3 as long as palmate membrane; radiolar flanges ment wide, incised, with a heart-shaped anterior extending until radiolar tips (Fig. 15C). Flanges margin (Perkins 1984, fig. 27F). Thus, they may beginning at about midlength of free parts, pro- be different species, and some additional differ- gressively widening distal; bare tips, flat, long, ences can be found by studying their chaetae. about the length of 2 or 3 pinnules, tongue-shaped (Fig. 15C); eyes in single rows, 10-30, beginning Notaulax occidentalis (Baird, 1865) just above palmate membrane (Fig. 15J). Radiolar (Fig. 15) skeleton with 4 cells in cross-section. Dorsal lips longer than palmate membrane (1-2 mm), with Sabella occidentalis Baird 1865: 159, pl. 5, figs. 7, 8. dorsal radiolar appendages but without pinnular Sabella alba Treadwell 1917: 266-267, pl. 3, figs. 10-15; 1939: appendages. Ventral lips and parallel lamellae 294, fig. 108 fide Perkins 1984: 339. present. Branchial lobes very long, dorsal and Parasabella sulfurea Treadwell 1917: 267, pl. 3, figs. 16-23 fide ventral margins with flanges extending from base Perkins 1984: 339. Hypsicomus purpureus Treadwell 1924: 20-21, pl. 2, figs. 30-33 of lobes to origin of dorsal and ventralmost radi- 52 Zoological Studies 45(1): 24-66 (2006) oles. Collar bilobed (Fig. 15A), with triangular and (Fig. 15N). Inferior thoracic notochaetae paleate, distally rounded ventral lobes. Thorax 3.5-5.5 mm asymmetrical (Fig. 15L, M), arranged in 2 trans- long with 8 chaetigers, 2.5-4 mm wide. Ventral verse rows. Thoracic uncini avicular, with 18 shield of segment 1 rectangular, divided into 2 equal-sized teeth above main fang in lateral view asymmetrical parts (Fig. 15B), following ventral (seen under light microscopy), covering 1/2 of shields trapezoidal, with broader anterior margin extension of main fang (Fig. 15F), hood absent, and lateral margins indented by tori. Segment 1 breast well developed, manubrium medium (Fig. with fascicles modified as a single, elongate, longi- 15D). Companion chaetae with roughly symmetri- tudinal to oblique row of spinelike chaetae (Fig. cal tips, as teardrop-shaped membranes (Fig. 15K). Superior thoracic notochaetae spinelike 15G-I). Abdomen with 78-162 chaetigers.

(A) (B) (C) rf

(J) se

(H) (I) (D) (E) (F) (G)

m (K) (L) (M) (N) (O) (P) (Q)

Fig. 15. Notaulax occidentalis. (A) Anterior end, dorsal view; (B) same, ventral view; (C) radiolar tip; (D) thoracic uncinus; (E) abdominal uncinus; (F) thoracic uncinus, frontal view; (G-I) companion chaetae; (J) radiole, area of eyes; (K) chaeta from segment 1; (L, M) inferior thoracic paleate notochaeta; (N) superior thoracic spinelike notochaetae; (O, P) abdominal paleate neurochaeta; (Q) abdominal modified, elongate, narrowly hooded chaeta. (A-Q) ECOSUR, Isla Cozumel, Mexico. Scale bars (A, B) 0.5 mm; (C, J) 0.1 mm; (D-I, K) 0.01 mm; (L-R) 0.05 mm. m, mucro; rf, radiolar flange; se, simple eyespot. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 53

Abdominal neurochaetae in 2 transverse rows of Posterior abdominal segments containing coelomic- chaetae: anterior row on anterior and posterior madurating oocytes. Pygidial eyes present. abdominal chaetigers paleate, with long mucro Remarks: Both N. occidentalis and N. circum- (Fig. 15O-P); posterior row on anterior abdominal spiciens (Ehlers, 1887) have 2 rows of eyes per chaetigers elongate, narrowly hooded; posterior radiole; the 1st species has up to 30 eyes per radi- row on posterior abdominal chaetigers modified, ole, while the 2nd has up to 50. In N. circum- elongate, narrowly hooded (Fig. 15Q). Abdominal spiciens (described from Carysfort Reef, FL), the uncini avicular, with 24 equal-sized teeth above collar is divided into 4 lobes, and the ventral lobes main fang in lateral view, covering 1/2 of extension are rounded and longer than those in N. occiden- of main fang (seen under light microscopy); breast talis; in lateral view; the collar of N. circumspiciens well developed, manubrium short (Fig. 15E). has a shorter indentation than that in N. occiden-

(A) (B) (C)

(E) (F) (D)

(G) (H) (L) (M) (N) (K)

m (I) (J)

Fig. 16. Notaulax paucoculata. (A) Anterior end, dorsal view; (B) same, ventral view; (C) radiole, area of eyes; (D, E) chaetae from segment 1; (F) companion chaetae; (G) thoracic uncinus; (H) abdominal uncinus, non-lateral oblique; (I, J) inferior thoracic paleate notochaeta; (K) superior thoracic notochaeta; (L, M) abdominal paleate neurochaetae, anterior row, (N) abdominal modified, elongate, narrowly hooded chaeta. (A-N) USNM 062077, holotype. Scale bars (A, B) 0.5 mm; (C) 0.05 mm; (D-N) 0.01 mm. m, mucro; se, simple eyespot. 54 Zoological Studies 45(1): 24-66 (2006) talis. Abdominal neurochaetal fascicles in 2 transverse rows of chaetae: anterior row throughout abdomen Notaulax paucoculata Perkins, 1984 with symmetrical paleate chaetae with long dentic- (Fig. 16) ulate mucro (Fig. 16L, M); posterior row on anterior abdominal chaetigers with elongate, narrowly Notaulax paucoculata Perkins 1984: 345, figs. 33, 34. hooded chaetae; posterior row on posterior abdominal chaetigers with modified, elongate, nar- Material: [ECOSUR] Mexican Caribbean: I. rowly hooded chaetae (Fig. 16N). Abdominal unci- Cozumel, Ixchen, Coll. S. I. Salazar, 4 June 1995 ni avicular, with 17 equal-sized teeth above main (1); SEDENA, Coll. L. F. Carrera, 24 Mar. 2001, in fang in lateral view, covering 3/4 of extension of Ircinia strobilina (1); Coll. A. Medina (3). DIF main fang (seen under light microscopy), breast Puerto Aventuras, Coll. J. R. Bastida and S. I. well developed, manubrium short (Fig. 16H). Salazar, 28 Nov. 1999 (1). Punta Herradura, Coll. Pygidium with eyespots. L. F. Carrera and S. I. Salazar, 28 Oct. 1997, Com Remarks: Notaulax paucoculata has few radi- B90 (1). San Felipe, Coll. J. R. Bastida and S. I. olar eyes, and the dorsal margin of the collar is Salazar (1). [UMML] Mexican Caribbean: 22.766, deeply incised. The holotype is incomplete; the I. Cozumel, SEDENA, Coll. A. Medina, March 24, crown and posterior end of the body are in poor 2001 (3). [USNM] Holotype 062077, Lucaya, condition. Grand Bahama I., Hydrolab, off Bell Channel, 26 33'00''N, 78 34'00''W, Coll. S. A. Earle and A. Perkinsiana Knight-Jones, 1983 Hurley 8 July° 1974, Sta. 8-1, 41.1 m. ° Description: Body medium-sized, 12 mm long. Perkinsiana was erected to accommodate 11 Branchial crown very long at 11 mm with 8 pairs of species which did not reasonably fit into Potamilla radioles, 3 cross-banded with purplish-brown pig- Malmgren,1866, Potamethus Chamberlin, 1919, or ment (in preserved material). Radioles united Demonax Kinberg, 1867 (Knight-Jones 1983). proximally above basal lamina by palmate mem- Fitzhugh (1989) found that Perkinsiana could not brane for 2/5 of their length, flanged above be diagnosed in terms of apomorphy. Only 1 palmate membrane for their its length. Simple species of Perkinsiana has been described from radiolar eyes present above palmate membrane, 4 the Grand Caribbean: P. fonticula from Puerto or 5 eyes in a small group on lateral borders of Rico, recorded below. radioles (Fig. 16C), at distal end of middle color band. Radiolar skeleton with 4 rows of cells in Perkinsiana fonticula (Hoagland, 1919) cross-section. Dorsal lips with radiolar append- (Fig. 17) ages, but without pinnular appendages. Ventral lips and parallel lamellae present. Collar long, Parasabella fonticula Hoagland 1919: 579, pl. 31, figs. 3-9. covering origin of branchial crown, deeply incised Potamilla fonticula.- Fauchald 1977: 62, fig. 13c-k. dorsomedially (Fig. 16A). Thorax with 8 Perkinsiana fonticula.- Knight-Jones 1983: 282, fig. 17A-L. chaetigers, 3 mm long, 1.5 mm wide. Chaetae from segment 1 in a single, elongate, longitudinal Material: [ECOSUR] Mexican Caribbean: to oblique row of spinelike chaetae (Fig. 16D, E). Majahual, dead coral, Coll. S. I. Salazar and B. Ventral shield of segment 1 rectangular, divided Trujillo, 7 June 2003 (1). DIF Puerto Aventuras, into 2 asymmetrical regions, basal one well Coll. S. I. Salazar, 21 Mar. 1992 (5). Tulum, Punta defined (Fig. 16B), following shields trapezoidal, Piedra, Coll. LEGE, 11 Feb. 2001 (3). I. Contoy, indented by tori. Superior thoracic notochaetae Punta Sur, Coll. S. I. Salazar, 2 Mar. 2001 (1). I. spinelike (Fig. 16K). Inferior thoracic notochaeta Cozumel, Puerta Maya, Coll. A. Medina, 26 Mar. paleate, asymmetrical and symmetrical, arranged 2001 (1). in 2 transverse rows (Fig. 16I, J). Thoracic uncini Description: Body medium-sized, 22-30 mm avicular, with 18 equal-sized teeth above main long, 1 mm wide. Branchial crown supported by a fang in lateral view (seen under light microscopy), deep cartilaginous base that joins 2 halves dorsally covering 1/2 of extension of main fang, hood (Fig. 17B), 5 mm long, with 6-8 pairs of radioles absent, breast well developed, manubrium medi- fused at bases. Anterior margin of anterior peristo- um (Fig. 16G). Companion chaetae with roughly mial ring low, posterior peristomial ring collar pre- symmetrical tips, as teardrop-shaped membranes sent (Fig. 17A, B). Dorsal lips tapered with outer (Fig. 16F). Abdomen with 31-42 chaetigers. lamella fused to slightly enlarged pinnules. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 55

Elongated triangular collar lappets (Fig. 17C), par- verse rows of chaetae: anterior and posterior rows allel lamellae present. Anterior margin of shield of with elongate, broadly hooded chaetae throughout. segment 1 rounded, M-shaped (Fig. 17C). Thorax Abdominal uncini avicular, with 19 equal-sized with 20 chaetigers. Thoracic fascicles very small teeth above main fang in lateral view, covering 1/3 with only 1 or 2 slender, narrowly hooded superior of extension of main fang (seen under light chaetae (Fig. 17F, G), and about 5 inferior paleate microscopy), breast well developed, manubrium notochaetae with short mucro (Fig. 17D, E), long. arranged in 2 or more transverse rows. Thoracic Remarks: The holotype of P. fonticula as tori with 6-9 avicular uncini, with 16 equal-sized specimens recorded in this study has 20 thoracic teeth above main fang in lateral view, covering 1/2 chaetigers. Fauchald (1977) recorded specimens of extension of main fang (seen under light from Panama with 14-22 chaetigers, but probably microscopy), hood absent, breast well developed, this variation is due to regeneration of the anterior manubrium long. Companion chaetae with roughly end of the body or may depend on the age of the symmetrical tips, as teardrop-shaped membranes organisms. (Fig. 17I, J). Abdominal neurochaetae in 2 trans-

(A) (B) (C)

(D)

(E) (F) (G) (H) (I) (J)

Fig. 17. Perkinsiana fonticula. (A) Anterior end, lateral view; (B) same, dorsolateral view; (C) same, ventral view; (D, E) inferior thoracic paleate notochaetae; (F, G) superior thoracic chaeta spinelike; (H) abdominal elongate broadly hooded chaeta; (I, J) companion chaetae. (A-J) ECOSUR, Isla Cozumel, Mexico. Scale bars (A-C) 1 mm; (D-J) 0.01 mm. 56 Zoological Studies 45(1): 24-66 (2006)

Pseudobranchiomma Jones, 1962 49'34''N, 80 08'20''W, 1.2 m, 10 Mar. 1982, Id. H. D. Rudolph° (1). [FSBC] I 43730, near Miami Fitzhugh (1989) used 3 characters to sepa- Beach shore, 25 48'49''N, 80 08'48''W (2). rate Pseudobranchiomma from Branchiomma: the [FSBC] I 43731, Sta.° 40, 25 47'47''N,° 80 11' presence of radiolar flanges, dorsal lips without 06''W, 1.7 m, 11 Mar. 1982, Id. °H. D. Rudolph° (2). pinnular support, and abdominal fascicles with [FSBC] I 43732, Sta. 38, 25 46'40''N, 80 11'57''W, long chaetae. Knight-Jones (1994) indicated, how- 2.7 m, 11 Mar. 1982, Id.° H. D. Rudolph° (1). ever, that the last character is also present in [FSBC] I 43733, South Biscayne Bay, 25 32'01''N, Branchiomma; the presence or absence of the pin- 80 10'17''W, 2.4 m, 21 Nov. 1982, °Id. H. D. nular support in the dorsal lip is not very robust, Rudolph° (2). [UMML] 22.762, Broad River Delta, since the holotype of P. emersoni has 1 lip with Chatham River Delta, Coot Bay Delta, Coll. W. L. this appendage, while the other has a free end. In Rouse, 29 Dec. 1965. the same work, she indicated that P. odhneri and Description: Body pale or pink (in preserved P. longa (Kinberg, 1867) have radiolar eyes (this material), small, 1-14 mm long. Base of crown feature was sufficient to emend the genus), and involuted midventrally, each side forming a partial that radiolar serrations extend through a variable circle with dark inter-radiolar spots (Fig. 18C). range, since they can run along the radiole, are Radioles without eyes, with paired serrated stylods restricted to the distal region, or are missing com- for most their length (5-9 serrations) (Fig. 18K). pletely. Palmate membrane extending for 1/8 length of Nogueira and Knight-Jones (2002) suggested branchial crown. Radioles with bare tips, as long that a thorax with few chaetigers can be attributed as equivalent space of 9 pinnules (Fig. 18K). to an imperfect regeneration after scissiparity; Dorsal lips long (1/2 lengths of radioles) with dor- some records of Pseudobranchiomma species sal radiolar appendages. Ventral lips rounded. show variations of 4, 6, and 8 chaetigers. Also, Midline fecal groove deep on 1st segment forming they indicated that the external radiolar mounds at each side, collar well separated dorsal- appendages of Pseudobranchiomma are only reg- ly. Ventral lappets rounded, overlapping (Fig. ular serrations of the radiolar flange, which differs 18B). Anterior margin of 1st ventral shield M- from the paired stylodes of Branchiomma. shaped (Fig. 18A). Interramal eyespots present Knight-Jones and Giangrande (2003) divided (Fig. 18C). Thorax, 0.5-2 mm long and 0.3-1.5 13 species of Pseudobranchiomma into 3 groups: mm wide with 4-7 chaetigers. Spinelike chaetae (A) those with paired serrated flanges along all or on segment 1 arranged in compact fascicles. most of the extensions of radioles, as in the type Thoracic notochaetae arranged in irregular, species P. emersoni; (B) those with such flanges oblique rows of superior and inferior chaetae; only on the distal parts of the radioles; and (C) superior chaetae spinelike with knee slightly those with reduced or absent flanges (i.e., without broader than shaft (Fig. 18H); inferior chaetae serrations). Only 2 species of Pseudobranch- spinelike with knee up to twice as broad as shaft iomma have been described from the Grand (Fig. 18I). Thoracic uncini avicular, with 4 rows of Caribbean: P. emersoni Jones, 1962 (Jamaica) equal-sized teeth above main fang in frontal view and P. perkinsi Knight-Jones and Giangrande, (Fig. 18E), occupying 1/2 of extension of main fang 2003 from Cape Canaveral. (seen under light microscopy), breast well developed, manubrium short (Fig. 18F). Abdomen with Pseudobranchiomma emersoni Jones, 1962 18-23 chaetigers. Abdominal neuropodia as coni- (Fig. 18) cal lobes; anterior abdominal row with short, spine- Pseudobranchiomma emersoni Jones 1962: 198-201, figs. 115- like chaetae, posterior row with modified, elongate, 124. narrowly hooded chaetae (Fig. 18J). Abdominal uncini avicular, with 3 equal-sized teeth above Material: [FSBC] Florida: I 23705, North main fang in lateral view (Fig. 18G), covering 1/2 Biscayne Bay, Sta. 28040349, 25 49'34''N, 80 08' of extension of main fang (seen under light 20''W, 1 m, 22 Feb. 1979, Id. T. ° H. Perkins, Coll.° microscopy), breast well developed, manubria Biosystems Research, Miami (1). [FSBC] I 43728, short (Fig. 18G). Pygidium bilobed (Fig. 18D). Sta. 32, 25°44'03''N, 80°12'05''W, 11 Oct. 1981, Tubes flexible with fine sand. Id. H. D. Rudolph (3). [FSBC] I 43729, Sta. 4a, 25° Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 57

Pseudopotamilla Bruguiére, 1789 numerous questionable records in many areas far apart from its type locality (Iceland): Fauchald and Müller (1771: 194) described“Die nieren- Reimer (1975) and Fauchald (1977) recorded it formige Amphitrite”from Iceland. Bruguière from Panama; Perkins and Savage (1975) listed it , (1789: 57) gave a binomial name to Müller s from Florida, the Gulf of Mexico, and the species: Amphitrite reniformis, and considered it to Caribbean; Laverde-Castillo and Rojas-García be the type species for Pseudopotamilla. (1983) from Bahía de Cartagena (Colombia); and Unfortunatelly, the type material is missing; Knight- Uebelacker (1984) from Florida. We herein state Jones et al. are working in a revision of species that the specimens studied by Uebelacker (1984), from the North Sea, and describing the neotype of from the northern Gulf of Mexico and identified as P. reniformis. P. cf. reniformis correspond to a new species Pseudopotamilla reniformis (Müller, 1771) has described below as P. fitzhughi sp. nov.

(A) (B) (C) (D)

(E)

sty (K)

(H) (I) (J) (F)

(G)

Fig. 18. Pseudobranchiomma emersoni. (A) Anterior end, ventral view; (B) ventral lappets and thoracic shields; (C) anterior end, lateral view; (D) posterior end; (E) thoracic uncinus, frontal view; (F) same, lateral view; (G) abdominal uncinus; (H, I) superior thoracic spinelike notochaetae; (J) abdominal posterior modified, elongate, narrowly hooded chaetae; (K) radiole. (A-K): UMML 22.762, Florida. Scale bars (A-D) 1 mm; (E-G) 0.02 mm; (H-J) 0.3 mm; (K) 0.1 mm. sty, stylode. 58 Zoological Studies 45(1): 24-66 (2006)

Natural History) for his significant contribution to Pseudopotamilla fitzhughi sp. nov. the knowledge and taxonomy of the Sabellidae, (Figs. 19-21) and for his sustained advice and recommendations Potamilla cf. reniformis Uebelacker 1984: 53-36, 54-38, figs. throughout this study. 54-29. Material: Holotype [LACM-AHF POLY 2150] Mexican Caribbean: I. Contoy, camping, on dead Etymology: This species is named in honor of coral, 1.5 m, Coll. S. I. Salazar, 1 Mar. 2001. Dr. Kirk Fitzhugh (Los Angeles County Museum of [LACM-AHF POLY 2151] One paratype. [ECO-

(A) (B) (C)

(D) (E) (F)

itnp

ce (G) (H)

Fig. 19. Pseudopotamilla fitzhughi sp. nov. (A) Anterior end, dorsal view; (B) same, ventral view; (C) same, lateral view; (D) compound eyes; (E) dorsal pinnular appendage; (F) thoracic chaetiger; (G) thoracic uncinus; (H) abdominal uncinus. (A-H) LACM-AHF POLY 2150, Isla Contoy, Mexico, holotype. Scale bars (A-C) 0.5 m; (D) 0.15 mm; (E, G, H) 0.01 mm; (F) 0.2 mm. dl, dorsal lip; ce, compound eye; itnp, inferior thoracic notochaetae paleate. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 59

SUR 0050] Three paratypes (one mounted in gold 2645, 106 m, Id. J. M. Uebelacker (3). for SEM). [UMML 22.767] One paratype. [ECO- Description (variations observed in paratypes SUR] I. Contoy, Coll. L. F. Carrera and S. I. given in parentheses): Holotype complete, with Salazar, June 1999 (5). San Felipe, Coll. J. R. medium-sized body, 8 (1.5-11.5) mm long, 1 (0.75- Bastida and S. I. Salazar, 19 Feb. 1999 (5). 1.5 mm) mm wide with purple crown and thorax (in Cancún, Punta Nizuc, Coll. S. I. Salazar and L. F. live and preserved materials). Radiolar crown Carrera, 30 Mar. 1997, rock 2 (2). I. Cozumel, long, measuring about 1/2 of body length, 4 (1.5- SEDENA, Coll. S. I. Salazar, 24 Mar. 2001 (1). 5.5) mm long. Base of crown with erect rigid dor- Majahual, Coll. J. R. Bastida and P. Salazar, 22 sal, basal flanges. Crown with unpaired com- Mar. 2000 (1). [FSBC] Florida: I 23695, east edge pound eyes only in dorsal radioles, except on dor- , of Jeff s Reef, east coast of Florida, 27 32.8'N, 79 salmost pair, limited to proximal 1/2 of radioles, 3 58.8'W, Sta. JSL I-438 B, 23 Aug. 1977,° Id. T. H.° or 4 teardrop-shaped eyes per radiole (Figs. 19D, Perkins as P. reniformis, Coll. L. E. Edmiston, 81.4 21A). Nine pairs of radioles, diminishing in length m (20). [FSBC] Curacao: I 60311, Netherlands ventrally. Radioles with bare tips, as long as the Antilles, 9.1 m, Coll. Y. Steward-Van Es, Oct. 1977, space equivalent to 8 pinnules. Outer radiolar Id. T. Perkins as P. reniformis (3). [UMML] margins quadrangular in cross-section. Radiolar Mexican Caribbean: 22.768, San Felipe, Coll. J. R. skeleton with 4 rows of cells in cross-section. Bastida and S. I. Salazar, 19 Feb. 1999 (4). Dorsal lips long, measuring about 1/2 of length of [USNM] Texas: 090424, Hospital rock, 27 32' radioles, triangular and with distinct longitudinal 0.5''N, 96 28'19''W, Coll. BLM, Sta. HR1, 1976,° 75 ridge (mid-rib), with radiolar and pinnular m. Id. J. °M. Uebelacker as P. cf reniformis (1). appendages (Fig. 19E). Ventral lips and parallel [USNM] 090425, Southern Bank, 27 26'06''N, 96 lamellae present. Anterior margin of anterior peris- 31'47''W, BLM-OCS, Spring 1976, Sta.° SB3, 82 m,° tomial ring low, of even height in lateral view (Fig. Id. J. M. Uebelacker (1). [USNM] Alabama: 19C), forming 2 deep pockets dorsally (Fig. 19A); 090426, 29 40'30''N, 87 37'00''W, BLM-OCS, posterior peristomial ring collar present. Ventrally, May 1974, Sta.° 16 F, 36 m,° Id. J. M. Uebelacker collar with a pair of triangular lappets, not overlap- (1). [USNM] 090429, off Mobile Bay, 29°35'00''N, ping, nearly reaching proximal margin of 1st ven- 87°20'02''W, Coll. BLM-OCS, Feb. 1978, Sta. VI- tral shield, when reflexed (Fig. 19B). Dorsal mar-

(B) (C)

(A)

(D) (E)

Fig. 20. Pseudopotamilla fitzhughi sp. nov. (A) Upper row of inferior thoracic paleate notochaetae; (B, C) inferior thoracic paleate notochaetae from lower row; (D) thoracic uncinus and companion chaetae; (E) elongate, broadly hooded abdominal chaeta. (A-E) LACM-AHF POLY 2150, Isla Contoy, Mexico, holotype. Scale bars (A-F) 0.01 mm. 60 Zoological Studies 45(1): 24-66 (2006) gin of collar fused to fecal groove (Fig. 19A). described from Singapore and later synonymized Ventral shield of segment 1 divided into 2 asym- by Johansson (1927) with P. lacioniosa Ehlers, metrical regions, anterior one with distal margin 1904 from New Zealand. Pseudopotamilla rounded (Fig. 19B). Thorax 1.5 mm long, 0.3-1 oligophthalmus and P. fitzhughi sp. nov. have mm wide, with 9 chaetigers. Each ventral thoracic entire dorsolateral collar margins, with rounded shield divided into 2 regions by a dark band. dorsal margins of the collar near the fecal groove Segment 1 with spinelike chaetae. Following tho- (Table 2). They differ because (A) P. oligophthal- racic chaetigers with 3 spinelike superior mus has rounded ventral lappets, while in P. notochaeta (Fig. 19F), and 5-7 paleate inferior fitzhughi sp. nov., they are triangular. notochaetae (Fig. 19F), arranged in 2 transverse Other similar species are P. aspersa (Krøyer, rows, upper row with short and medium length 1856), P. reniformis (Bruguière, 1879), and P. mucro (Fig. 20A), lower row with short mucro (Fig. saxicava (de Quatrefages, 1866); P. fitzhughi sp. 20B, C). Thoracic tori longer than abdominal tori, nov. differs from these species because (A) in P. not reaching ventral shields. Thoracic avicular reniformis, the dorsal margins of the collar are uncini18 or 19 per torus, with main fang surmount- quadrangular near the fecal groove (rounded in P. ed by 19-22 rows of equal-sized teeth in frontal fitzhughi sp. nov.); (B) P. aspersa and P. reniformis view (seen under SEM), occupying 1/2 of exten- have indistinct ventral lappets (distinct and triangu- sion of main fang, breast well developed, manubri- lar in P. fitzhughi sp. nov.); (C) the manubrium of um long (Figs. 19G, 20D). Thoracic uncini reach- the thoracic uncini in P. saxicava are as long as ing 3/4 of length of manubria of companion the shaft of the companion chaetae (3/4 in P. chaetae (Fig. 20D). Companion chaetae with fitzhughi sp. nov.); and (D) P. aspersa and P. reni- roughly symmetrical tips, as teardrop-shaped formis are not found in rocks (P. fitzhughi sp. nov. membranes (Figs. 20D, 21E). Abdomen with 11- and P. saxicava live in rocks) (Table 2). 32 chaetigers. Tori smaller than those on thorax, with 10-13 uncini. Abdominal neurochaetae in 2 Sabellastarte Krøyer, 1856 transverse rows: anterior and posterior rows with 3 or 4 elongate, broadly hooded chaetae on all Fitzhugh (1989) provided a complete diagno- chaetigers (Fig. 20E), those of anterior rows short- sis for the genus; he noted that Sabellastarte does er (Fig. 21F). Abdominal uncini avicular, with 22- not appear to be an apomorphic definable group. 24 rows of equal-sized teeth above main fang in Knight-Jones and Mackie (2003) recognized 8 frontal view (seen under SEM), occupying 3/4 of valid species in the genus, and included typical but extension of main fang, breast well developed, not unique characters in their generic diagnosis. manubrium short (Fig. 19H). Tubes made of Only 1 species has been described from the Grand mucus and fine sand. Caribbean: S. magnifica (Shaw, 1800) from Remarks: Pseudopotamilla fitzhughi sp. nov. Jamaica. resembles P. oligophthalmus (Grube, 1878),

Table 2. Comparison of selected features between the species of Pseudopotamilla

Collar margin Ventral Thoracic uncini Substrate Type locality lappets length vs. shaft Dorsally, near of companion fecal grooveDorsolaterally chaeta

P. aspersa rounded V-shaped notches indistinct 1/2 ? Greenland P. fitzhughi sp. nov. rounded entire distinct, triangular 3/4 dead coral Mexican Caribbean P. oligophthalmus rounded entire distinct, rounded 3/4 ? Singapore P. reniformis quadrangular V-shaped notches indistinct 3/4 kelp holdfasts, Iceland among encrusting fauna on surfaces of sheltered, hard substrata P. saxicava rounded entire distinct, triangular 1 in rocks SW France Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 61

Sabellastarte magnifica (Shaw, 1800) 99, Sta. 42, LH-316, Coll. T. Zimmerman, 1 Aug. (Fig. 22) 1999 (1). [UMML] Virgin Is.: 22.121, St. John, Greater Lameshur Bay, Coll. Voss, Kumpf, and Tubularia magnifica Shaw 1800: 228, pl. IX, figs. 1-6. Adams, 22 Jan. 1959 (2); [UMML] 22.752, Coll. Sabella magnifica.- Savigny 1822: 78.- de Quatrefages 1866: Manning and Moore, 5 Jan. 1961 (1); [UMML] 443.- Fitzsimons 1965: 643, figs. 1-13. Sabella lingva Krøyer 1856: 27 fide Knight-Jones and Mackie 22.127, Coll. and Id. Thomas, 14 Dec. 1958 (1); 2003. [UMML] 22.120, St. John, Kiddle Bay, Coll. G. Sabella melania Schmarda 1861: 35, fig. 188 fide Knight-Jones Voss, Id. J. Evald, 23 Jan. 1959 (1). Venezuela: I. and Mackie 2003. Larga, Coll. S. Tovar, 10 Nov. 1962 (1). Sabella splendida Kinberg 1867: 353 fide Knight-Jones and Description: Large-bodied species, 70-142 Mackie 2003. Sabellastarte magnifica.- Augener 1922: 48.- Rioja 1958: 287.- mm long. Branchial crown 53-69 mm long, each Knight-Jones and Mackie 2003: 2278-2280, fig. 3A-P. side of crown base involuted ventrally to almost form a circle on each side, outer surface of radi- Material: [ECOSUR] Mexican Caribbean: oles dark with pale longitudinal lines. Radioles 69- Veracruz, La Galleguilla, Coll. M. A. Tovar and S. 82 pairs; radiolar tips short and blunt. Radiolar Frontana, 12 May 2001 (4); I. Verde, Coll. SISV, 15 skeleton with 8-10 rows of cells in cross-section June 1983 (1). Panama: Fuerte Sherman, Colón (Fig. 22D). Dorsal lips long (3/4 of length of (1), Coll. S. I. Salazar, 2 June 2002 (1); Coll. L. crown), with radiolar appendages, but without pin- Harris (8). [LACM-AHF] British Virgin Is.: BVI-00, nular appendages. Ventral lips with parallel lamel- Sta. 28 Guana, Coll. K. Fitzhugh (1); BVI-00, Sta. lae. Lateral margins of collar transverse to axis of 59 (1); BVI-99, Sta. 13, LH128, 27 Aug. (1); BVI- body, well above junction between crown and tho-

(A) (B) (C)

25.0 µm 20.0 µm 7.69 µm

(D) (E) (F)

10 µm 10.1 µm 28.5 µm

Fig. 21. Pseudopotamilla fitzhughi sp. nov. (A) Crown, right part; (B) inferior thoracic notochaetae paleate; (C) thoracic uncini; (D) spinelike chaeta from segment 1; (E) companion chaetae; (F) elongate, broadly hooded abdominal chaetae. (A-F) ECOSUR 0050, Isla Contoy, Mexico, paratype mounted in gold for SEM. Scale bars (D) 10 µm. 62 Zoological Studies 45(1): 24-66 (2006) rax (Fig. 22C); dorsal margins equally high, with ly enclosed by anterior row. Anterior abdominal notches above collar pockets, defining well-devel- neurochaetae spinelike on anterior rows (Fig. oped dorsal lappets, with rounded margins (Fig. 22K); elongate, narrowly hooded on posterior 22A). Ventral lappets triangular, overlapping at rows; posterior abdominal neurochaetae spinelike midline (Fig. 22B). Thorax twice as wide as long, in anterior rows; modified, elongate, narrowly 5-12 mm long, 7-17 mm wide with 8-10 chaetigers. hooded on posterior rows (Fig. 22J). Abdominal Thoracic inter-ramal eyespots present. First seg- uncini avicular, with 13 equal-sized teeth above ment somewhat longer than following segments. main fang, covering 1/3 of main fang length (seen Thoracic tori long, extending towards ventral under light microscopy), breast well developed, shields. Inferior thoracic notochaetae arranged in manubrium short (Fig. 22G). Pygidium rounded. bundles with irregular, longitudinal rows of spinelike chaetae (Fig. 22H, I). Thoracic uncini avicular, Checklist of sabellids from the Grand Caribbean with 13 equal-sized teeth above main fang in lateral view (seen under light microscopy), covering 1/3 Species with an asterisk " * " are not included of length of main fang (Fig. 22E), hood absent, in the key until any revision sustains their distribu- breast well developed, manubrium medium (Fig. tion, because those records from the Grand 22F). Abdomen with 188-204 chaetigers. Caribbean area are questionable, either because Abdominal neurochaetae in partially spiraled material reviewed by some authors was unavail- arrangement, formed by anterior chaetal row; origi- able, or their identifications have not been corrobo- nal posterior chaetal row as a small bundle partial- rated with the type material. In Salazar-Vallejo

(C)

(A) (B)

rs (E) (D) (H) (I)(J) (K) ss

(F) (G)

Fig. 22. Sabellastarte magnifica. (A) Anterior end, dorsal view; (B) same, ventral view; (C) same, lateral view; (D) radiolar skeleton in cross-section; (E) main fang from thoracic uncinus, lateral view; (F) thoracic uncinus; (G) abdominal uncinus, non-lateral oblique; (H, I) inferior thoracic spinelike notochaetae; (J) abdominal spinelike neurochaetae; (K) abdominal elongate narrowly hooded chaeta. (A-K) LACM-AHF, Guana I. Scale bars (A-C) 2 mm; (D) 0.1 mm; (E-K) 0.01 mm. rs, radiolar skeleton; ss, sheath tissue; bv, blood vessel. Tovar-Hernández and Salazar-Vallejo -- Sabellids from the Grand Caribbean 63

(1996), references of all these records are includ- TL: Belize. USNM 74679 (HT, PT). ed. Jasmineira bilobata (Day, 1973) Amphicorina androgyne (Rouse, 1994) TL: Beaufort, NC. USNM 43134 (HT). TL: Carrie Bow, Belize. USNM 157629 (HT). *Jasmineira pacifica Annenkova, 1937 Amphicorina anneae (Rouse, 1994) TL: Peter the Great Bay, Asia. Type material not found. TL: Florida. USNM 157620 (HT). Manayunkia aestuarina (Bourne, 1883) *Amphicorina rivularis (Annenkova, 1929) TL: Western Europe. Species broadly distributed in TL: Schantar Is., Sea of Okhotsk, Russia. Type material European waters and in the North American coasts: Bell not found. (1982) recorded it from South Carolina, Bishop (1984) *Amphiglena mediterranea (Leydig, 1851) from the Gulf of Mexico, and Rouse (1995) from TL: Nice, France. USNM 5095 (HT). Chesapeake Bay (Maryland). Anamobaea orstedi (Krøyer, 1856) Manayunkia speciosa Leidy, 1855 TL: West Indies. ZMUC 18.9.1845 (HT). TL: Schuylkill River, Philadelphia, PA. Type material not Anamobaea phyllisae sp. nov. found. TL: Guana I., British Virgin Is. LACM-AHF (HT, PT). Megalomma bioculatum (Ehlers, 1887) Augeneriella hummelincki Banse, 1957 TL: Straits off Florida. MCZ 669 (PT), MCZ 824 (PT). TL: Salinja Plenchi, Bonaire. ZMH V-11917a (HT). Megalomma heterops Perkins, 1984 Bispira brunnea (Treadwell, 1917) TL: Florida. USNM 54721 (HT), FSBC I 27696, 27698 TL: Nassau Harbour, Bahamas. AMHN 982 (HT). (PT). Synonyms: Sabella bahamensis Augener, 1922 fide Megalomma lobiferum (Ehlers, 1887) Knight-Jones and Perkins (1998: 433). TL: Florida. MCZ (HT). Bispira melanostigma (Schmarda, 1861) *Megalomma pacifica Johansson, 1927 TL: Jamaica. NHMW (ST). Synonyms: Sabella thoracica TL: Gilbert Is., Melanesia. UZIU (HT). Krøyer, 1856 and Sabella variegata Krøyer, 1856 fide Megalomma perkinsi sp. nov. Knight-Jones and Perkins (1998: 415). TL: Florida. USNM 53976 (HT). Bispira paraporifera sp. nov. *Megalomma pigmentum (Reish, 1963) TL: Puerto Aventuras, Mexico. LACM-AHF (HT). TL: San Quintín, Mexico. USNM 57945 (HT). Records of Branchiomma bairdi (McIntosh, 1885) M. pigmentum from Perkins (1984) correspond to 2 unde- TL: Bermuda. BMNH 1885.12.1.391 (LT). scribed species (informally described herein as Branchiomma conspersum (Ehlers, 1887) Megalomma sp. 1 and Megalomma sp. 2 from Venezuela TL: Key West, FL. MCZ 848 (HT). and Florida, respectively). Branchiomma nigromaculatum (Baird, 1865) *Megalomma vesiculosum (Montagu, 1815) TL: St. Vincent, West Indies. BMNH 1839.12.27.16.20 TL: Devonshire, UK. NMW.1995.024 (NT). (LT, SST). Synonyms: Sabella crispa Krøyer, 1856, Notaulax bahamensis Perkins, 1984 Dasychone ponce Treadwell, 1901. Syntypes of TL: Bahamas. USNM 62082 (HT). Dasychonopsis arenosa Treadwell, 1924b correspond in Notaulax circumspiciens (Ehlers, 1887) part to young B. nigromaculatum and B. conspersum. TL: Florida. MCZ (HT). *Branchiomma wyvillei (McIntosh, 1885) Notaulax midoculi (Hoagland, 1919) TL: St. Thomas, West Indies. Type material is missing, TL: Dry Tortugas, FL. AMNH 1186 (HT). but the original drawings of thoracic uncini do not have Notaulax nudicollis (Krøyer, 1856) the typical shape of Branchiomma. It probably corre- TL: St. Thomas, West Indies. USNM 62084 (HT). sponds to Sabellastarte magnifica (Shaw, 1800). Synonyms: Sabella brevicollaris Grube, 1859, S. torquata Chone americana Day, 1973 Grube, 1877, and Protulides elegans Webster, 1884 fide TL: Beaufort, NC. USNM 43134 (HT). Perkins (1984: 331). *Chone duneri Malmgren, 1867 Notaulax occidentalis (Baird, 1865) TL: Spitsbergen, Norway. Type material not found. TL: St. Vincent, West Indies. BMNH (HT). Demonax flecatus (Hoagland, 1919) Notaulax paucoculata Perkins, 1984 TL: Puerto Rico. AMHN (HT). TL: Bahamas. USNM 62077 (HT). Demonax jamaicensis (Augener, 1924) *Notaulax phaeotenia (Schmarda, 1861) TL: Jamaica. ZMH V-6791 (HT). TL: Sri Lanka. NHMW (HT). Demonax lacunosus Perkins, 1984 Novafabricia infratorquata (Fitzhugh, 1983) TL: Hutchinson I., FL. USNM 54725 (HT). TL: Twin Cays, Belize. USNM 74644 (HT). *Demonax leucaspis Kinberg, 1867 Perkinsiana fonticula (Hoagland, 1919) TL: San Lorenzo, Peru. NRS 575 (HT). TL: Guanica, Puerto Rico. AMNH (HT). Demonax microphthalmus (Verrill, 1873) Potamilla floridana Augener, 1922 TL: Vineyard Sound, MA. USNM 13079 (ST). TL: Tortugas, SW Channel, Bird Key Reef, FL. MCZ *Euchone incolor Hartman, 1965 (HT). TL: Continental shelf off New England. LAMNH-261 (HT), Potamethus spathiferus (Ehlers, 1887) -262 (PT). TL: Florida. MCZ (HT). *Euchone southerni Banse, 1970 Pseudobranchiomma emersoni Jones, 1962 TL: Ballynakill Harbor, Ireland. NMI 77.1908 (HT). TL: Jamaica. AMNH 3612 (HT). Fabricinuda pseudocollaris Fitzhugh, 1990 Pseudobranchiomma perkinsi Knight-Jones and Giangrande, TL: Florida. USNM 122023 (HT). 2003 Fabricinuda trilobata (Fitzhugh, 1983) TL: Indian River at Cape Canaveral, FL. NMW CP94003 (HT), LACM-AHF, USNM, FSBC (PT). 64 Zoological Studies 45(1): 24-66 (2006)

Pseudofabriciola longa Fitzhugh, 1990 Augener H. 1906. Reports on the results of dredging, under TL: Highland Beach, FL. USNM 122056 (HT). the supervision of Alexander Agassiz in the Gulf of Pseudofabriciola quassiincisura Fitzhugh, 1996 Mexico and the Caribbean Sea, and on the east coast of TL: Coco Plum I., Belize. USNM 169974 (HT). the United States, 1877 to 1880, by the USS Coast Pseudofabriciola sofla Fitzhugh, 1996 Survey Steamer Blake, 42. Westindische Polychaeten. TL: Florida. USNM 169980 (HT). Synonym: Fabricia sp. Bull. Mus. Comp. Zool. 43: 91-196. A Uebelacker, 1984. Augener H. 1922. Über litorale Polychaeten von Westindien. *Pseudopotamilla reniformis (Müller, 1771) Gesells. Naturforsch. Freunde Berlin Schrifte 3-5: 38-53. TL: Iceland, Finland, Norway. Type material missing. Augener H. 1927. Polychaeten von Curaçao , 9 figs. Bijdr. Knight-Jones et al. are currently describing a neotype Dierk. 25: 39-82 (Knight-Jones pers. comm.). Baird W. 1865. On new tubiculous annelids, in the collection of Pseudopotamilla fitzhughi sp. nov. the British Museum, 2. J. Linn. Soc. 8: 157-160. TL: I. Contoy, Mexico. LACM-AHF (HT), ECOSUR (PT). Banse K. 1957. Die Gattungen Oriopsis, Desdemona und Sabellastarte magnifica (Shaw, 1800) Augeneriella (Sabellidae, Polychaeta). Medd. Dansk TL: Jamaica. BMNH 1998.932 (NT). Naturh. Foren. 119: 67-105. Banse K. 1970. The small species of Euchone Malmgren Acknowledgments: This work was possible (Sabellidae: Polychaeta). Proc. Biol. Soc. Wash. 35: 387- 408. thanks to the support provided by several col- Banse K. 1972. Redescription of some species of Chone leagues. Kirk Fitzhugh and Eduardo Suárez pro- Kröyer and Euchone Malmgren, and three new species vided much useful advice throughout this study. (Sabellidae, Polychaeta). Fish. Bull. US Fish Wildl. Serv. João Miguel de Matos Nogueira revised the manu- 70: 459-495. script and provided valuable suggestions. Leslie Bell SS. 1982. On the population biology and meiofaunal characteristics of Manayunkia aesturiana (Polychaeta: Harris and Andy Cohen made possible the inclu- Sabellidae: Fabriciinae) from a South Carolina salt marsh. sion of one of the authors (SISV) in the Pew Exotic Estuar. Coast. Shelf Sci. 14: 215-221. Species Expedition to Panama. Todd L. Bishop TD. 1984. A range extension for Manayunkia aestuari- Zimmerman and J. W. Martin made possible a na (Bourne, 1883) (Polychaeta: Sabellidae) to the Gulf series of expeditions to Guana I. (British Virgin Coast of the United States with a review of previous habi- tat information. Gulf Res. Rep. 7: 389-392. Islands) under a grant from the Biotic Surveys and Bourne AG. 1883. On Haplobranchus, a new genus of Inventories Program of the US National Science Capitobranchiate annelids. Q. J. Microsc. Sci. 23: 168- Foundation (DEB 9972100), during which L. Harris 176. collected important material. Harry ten Hove Bruguière LG. 1789. Encyclopedie methodique. Hist. nat. (ZMA) made an intense search of sabellids in his vers Paris Panchoucke 1: 1-344. Carpenter DG. 1956. Distribution of polychaete annelids in the collections from the Antilles and provided some Alligator Harbor Area, Franklin County, Florida. FL State useful material, and also made important sugges- Univ. Stud. 22: 89-110. tions to the manuscript. Andrew Cabrinovic Chamberlin RV. 1919. The annelida polychaeta of the (BMNH), Sandra Farrington (FSBC), Nancy Voss Albatross tropical Pacific expedition, 1891-1905. Mem. (UMML), and Kristian Fauchald (USNM) loaned Bull. Mus. Comp. Zool. Harv. Univ. 48: 1-514, 80 pls. Claparède É. 1868. Les Annelides Chetopodes du Golfe de material revised in this study. Leslie Harris provid- Naples. Mém. Soc. Phys. Hist. Nat. Gen. 19: 313-584, 16 ed motivation and hospitality, both extremely pls. important to culminate this work. We would like to Day JH. 1973. 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